Nis Me TEN Leecrere € : etn ‘ ve uy Sa Seekoes pyc tye She prey bara hk abed aR SIT RE ARIS: . corse rettees Cees "Se ee vt Pay tier’ ied 3 ae Seka ree HARVARD UNIVERSITY Ernst Mayr Library of the Museum of Comparative Zoology = 4 y M4 | Ai 7 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 118, Number 1 January-March 2004 125" Anniversary of the Ottawa Field-Naturalists Club 1879-2004 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 information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining 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 2004 Council President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor Francis R. Cook Cendrine Huemer To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, 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 IRO; (613) 269-3211; e-mail: fecook @achilles.net Copy Editor: Elizabeth Morton Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john @pwgsc.gc.ca 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 except Book Reviews should be addressed to the Editor and sent by postal mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, 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 members (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: January—March 2004 (December 2004). Cover: A male Northern True Katydid, Prerophylla camellifolia Ottawa, from location 3 Figure 1 in Darbyshire pages 124-126. This is the same individual as in Figure 2, page 125. Photographed by Stephen J. Darbyshire. MCZ LIBRARY JAN § 2005 HARVARD THERCANAIDEAIN (nt tae © FIELD-NATURALIST Volume 118 2004 Volume 120 The Ottawa Field-Naturalists’ Club Transactions THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA ate The Canadian Field-Naturalist Volume 118, Number | January—March 2004 Origins and History of The Ottawa Field-Naturalists’ Club DANIEL F. BRUNTON 216 Lincoln Heights Road, Ottawa, Ontario K2B 8A8 Canada; e-mail: dbruntonn211@rogers.com Brunton, Daniel F. 2004. Origins and history of The Ottawa Field-Naturalists’ Club. Canadian Field-Naturalist 118(1): 1-38. The Ottawa Field-Naturalists’ Club (OFNC) represents an unbroken chain of organized, non-governmental natural history investigation and education dating back to the early days of the city of Ottawa itself. The Club originated in 1863 with the formation of the Ottawa Natural History Society which became the Natural History branch of the Ottawa Literary and Scientific Society in 1870, from which the OFNC formally separated in March 1879. Since that time, it has grown into Canada’s oldest and largest regional natural history organization and has produced a diverse and internationally recognized publication program. Since 1880 The Canadian Field-Naturalist and its predecessors have constituted the scientific core of the OFNC’s publication program, with Trail & Landscape being an important Ottawa Valley publication since the late 1960s. The importance of both publications to the growth and health of the organization is reflected in the major surges in Club membership experienced when each of these publications was established. The focus of membership activities has changed over the history of the OFNC, with enlightened natural resource management, then original scientific research and local exploration directing energies in the early decades. By the early years of the 20" century the publications program become the raison d’etre of the Club, almost to the exclusion of local field activities. A renewed interest in field discovery and the growth of conservation awareness in the 1960s, however, rekindled local activities and re-established the balance which has sustained the organization throughout its history. Natural environment education has remained a critical theme within OFNC programs and activities. Over and above inspiring the professional careers and private interests of thousands of individuals for more than a century, the OFNC has had an important and lasting impact on the conservation of natural environment features and landscapes in Canada and North America. Key Words: Ottawa Field-Naturalists’ Club; Ottawa Natural History Society; The Canadian Field-Naturalist; Trail & Landscape. Canada was a mere 12 years old in 1879 when 34 members of the Ottawa Literary and Scientific Society (OLSS) gathered together on an early spring evening to discuss a growing problem within Ottawa’s fledg- ling naturalist community. Most were full of Victorian enthusiasm for discovery and intellectual advance- ment and were convinced of the limitless potential of their new country. These young men — and they were all men, mostly in their 20s or 30s — were frustrated by what they saw as an ineffective, moribund OLSS Natural History Branch which did not serve their needs. These impatient “young Turks” wanted to actually do things, to get out into the countryside of the Ottawa Valley to explore and discover its natural wonders. And with true Victorian missionary zeal, they wanted to share these revelations amongst themselves and with the larger Canadian — even international — commu- nity. All of this in the name of personal intellectual development as well as the chance to advance the scientific and applied benefits of such knowledge. That was heady, revolutionary stuff in the staid, conserva- tive Ottawa of March 1879 when how close one was to Rideau Hall — the literal and figurative operational base of the Governor General and his politically and socially powerful entourage — dictated more about one’s status and options than personal wealth or political position (Gwyn 1984). The events which unfolded that cool spring evening in the OLSS rooms perched above the muddy streets of Ottawa launched careers, changed government policy, protected tens of thousands of hectares of Canadian natural landscape, produced internationally recognized and significant scientific publications, made huge con- tributions to our understanding and appreciation of North American natural sciences, and enriched thou- sands upon thousands of lives. Oh yes ... and initiated what has become the largest and oldest regional natu- ralist organization in Canada’s history, The Ottawa Field-Naturalists’ Club (OFNC). The Pioneers (1840s-1863) Although the formal beginning of The Ottawa Field- Naturalists’ Club in 1879 was 125 years ago, natural- ists’ organizations in the Capital actually pre-date Can- ada itself (Brault 1946; Dore 1968; Taylor 1986). Prior to Confederation, Ottawa (and Bytown before it) was D THE CANADIAN FIELD-NATURALIST a rude little lumber town characterized more by saw- dust, beer and brawls than by intellectual achievement. The only adult education or research institution pre- sent in those early days was the Mechanics Institute, a charitable organization initiated in 1847 as something akin to a continuing education facility and library for working men. There were no such things as “night school” or public libraries, let alone publicly accessi- ble research organizations. An informal group known as the Silurian Society interested in geological (and mining?) issues was also reported to be active in the 1850s. “Active” may be a misnomer, since they left little reference of their doings, other than to suggest that their meetings were held “in the City of Ottawa” (Anonymous 1854). The only natural environment research being under- taken in the Ottawa Valley before the 1860s was by three highly active individuals. Edward Van Cortlandt (1805-1875) was the most socially prominent of these. He was one of the first doctors in Bytown, arriving in 1832 to attend to the military personnel stationed on Barracks Hill (now Parliament Hill). He developed an extensive private museum of curiosities and artifacts in the 1840s, liberally mixing archaeological specimens found at aboriginal sites along the Ottawa River with natural items dug up, collected, and/or shot in the vicinity of the town. He was, by all accounts, a remark- ably energetic man who used his high social standing to influence local business leaders in natural resource- oriented concerns (Moffatt 1986). Another dynamo was Elkanah Billings (1820-1876) (Figure 1), second son of one of Ottawa’s first pio- neer families. Billings was passionately interested in natural history in general and geology/paleontology in particular. At various times he was a newspaper pub- lisher (The Bytown Citizen, forerunner of The Ottawa Citizen), a lawyer and finally, Canada’s first profes- sional paleontologist. Indeed, this latter science was his true calling and he came to be known as “the father of Canadian paleontology” (Whiteaves 1876; Clarke 1971). He presumably was involved in the aforemen- tioned Silurian Society, but no direct evidence of that was found. The Bytown Citizen was full of natural his- tory items during Billings’s tenure (1852-1856), mostly representing accounts of his own observations or text reprinted from European or American publications. These publication activities led directly to his produc- tion in Ottawa of Ontario’s first natural science journal, The Canadian Naturalist and Geologist, in February 1856 (Figure 2). Delightful and insightful articles in that first volume such as “On the species of woodpeck- ers observed in the vicinity of Ottawa” (Billings 1856) were based on his extensive travels in the Ottawa Valley. They demonstrated both excellent powers of observation and a keen appreciation of the importance of documenting the appearance and constitution of original landscape conditions. His move to Montreal later that year to join Sir William Logan at the Cana- Vol. 118 FiGure |. Elkanah Billings. Billings was born in 1820, became one of the earliest naturalists in Ottawa, and pub- lished the first journal on natural history in Ontario in 1856 (see Figure 2). He moved to Montreal later that year to become the first palaeontologist of the Geo- logical Survey of Canada, and the initial curator of its museum. (reproduction of the OFNC-commissioned portrait, from The Ottawa Naturalist, February 1901). dian Geological Survey, however, permanently ended his Ottawa connection (Zaslow 1975). Elkanah Billings’ older brother, Braddish Billings Jr. (1819-1871), completes the trio of Ottawa’s pioneer resident naturalists. Braddish was a keen botanist and used his position as chief clerk on the Prescott & Otta- wa Railway to gain access to a wide variety of habitats across eastern Ontario. Although he published nothing during this time and relatively little even later, in 1868 he did produce the first list of vascular plants for the city of Ottawa. It was a superb effort for its day, chron- icling over 400 species that he found within close proximity to the City in 1866 (Dore 1968). Billings was seen as a distinguished figure in natural history investigations in the Ottawa Valley in the 1860s; like his younger brother Elkanah, Braddish was widely consulted by natural science researchers elsewhere in Canada (Dore 1968). The days of exclusively private investigation of the Ottawa Valley natural environment ended in the early ————s S i, ai Niclan i ae epee ; ES mn F EN_AS AN Vise AD OAS CPS ON BESO Oh _ B i 4 Gentiemen to whom this Number is sent, and who do wot wish to i (O Sabserive, arc respectfully requesied to return the same. RG eee re ey s ee «68H CANADIAN Waturalist «Grnlogist, & By E. BILLINGS, Barrister at Law, OTTAWA, CANADA WEST. Pay lURee b, of — oe Eee Corama Fosstts, (Glyptocrizus rainulosus) x new species of Lily Enerivite, fromthe — ~ Trenion Linestose, Bighanys Lake, Coumy of Gilawa.— See quge 54, : : UBLISHED AT THE OFFICE or tue OTTAWA CITIZEN, OTTAWA. FIGURE 2. The Canadian Naturalist and Geologist, first pub- lished in 1856 at the office of The Ottawa Citizen by Elkanah Billings, and later continued in Montreal. 1860s with the transfer of the seat of government of the colony of Canada to Ottawa, heralded by the start of construction of the new Parliament Buildings in 1860 (Eggleston 1961). This represented a mega-project with huge economic benefits to local businesses. Similarly, the 1863 transfer of some 300 bright, educated, and rel- atively financially secure members of the civil service from the old capital represented both economic and social opportunities for the city and the Ottawa Valley. Confederation-era city business and social leaders quickly embraced the idea of broadening and deepen- ing the intellectual resources of the community. In furtherance of this, they encouraged the development of various clubs and societies to enhance the prestige and intellectual capacity befitting the new capital (Brault 1946; Gwyn 1984). It is useful to consider how really rough and ready Ottawa and the larger world were in 1863. The Arctic was still largely unknown to Europeans and North Americans alike but was being charted with great speed BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 3 by the continuing search for the ill-fated Third Frank- lin Expedition. The United States Civil War and the accompanying ferocious slavery debate raged uncom- fortably close to the south, as did intense arguments in the Old World regarding the newly published (1859) “heresies” of Charles Darwin’s On the origin of spe- cies. Closer to home, Ottawa was a bustling, rapidly growing city of approximately 15 000 people which boasted but a single operating sewer line along Welling- ton Street in front of the new Parliament Buildings. The first railway train had puffed into town fewer than ten years earlier and a municipal drinking-water system was still 12 years off (Brault 1946; Eggleston 1961). Ottawa was very much straddling the line between pioneer lumber town and developing political centre. The Ottawa Natural History Society (1863- 1869) It is against this backdrop that a veritable “Who’s- who” of the new business and professional elite of Confederation-era Ottawa (notably including both the aforementioned Edward Van Cortlandt and Braddish Billings Jr.) assembled on 3 October 1863 to form the Ottawa Natural History Society (ONHS). These gentle- men — and the ONHS constitution made it clear that membership was open only to gentlemen — were very economically oriented. The original minute books maintained in the OFNC collection at Library and Archives Canada (LAC) records that they were “ ... desireous to develope [sic] the Natural History of the Ottawa and general resources of the surrounding coun- try” (LAC OFNC Collection, 3 October 1863). Explor- ing and researching the natural sciences of the Ottawa Valley were all well and good but these gentlemen — at least initially — wanted to see a profit result from it. An important aspect of the growth of such endeay- ours was the potential participation of the relatively large body of newly arrived civil servants (Brault 1946). Activity within quasi-professional associations (it’s called “networking” today) was a very important un- official avenue for professional advancement within the small professional community of public servants in the Capital at this time (Gwyn 1984). The ONHS offered such an outlet and avenue for civil servants interested in natural resources and natural environment issues. A definite pecking order mirroring that of their profes- sional relationships was soon evident amongst found- ing ONHS members who were also senior civil ser- vants. When powerful Finance Department Deputy Minister John Langton (1808-1894) was ONHS Presi- dent in the late 1860s, for example, his ambitious sub- ordinate, Interior Department Deputy Minister Edmund Meredith (1817-1898), was conspicuously involved in the organization, but was careful not to hold higher office nor to publicly disagree with Langton (Gwyn 1984). The social/political significance of the organi- zation, over and above the fact that 77 men paid the $1.00 membership fee for 1863/1864, can also be meas- 4 THE CANADIAN FIELD-NATURALIST Vol. 118 ured by the prominent involvement of individuals such as Sir James A. Grant (1831-1920), the Governor Gen- eral’s personal physician (Travill 1988) (LAC OFNC Sls = = Collection, 3 October 1863). iD = oben re ois The ONHS members chose widely appreciated and £ g 2 a 2 z si cs 6 politically-neutral Braddish Billings Jr. as their first 2) 5 E ee = mm O68 S president, although he was not very active in subse- S A lai = = = a Ss ice quent Society affairs (LAC OFNC Collection, 3 Oc- tober 1863 — 28 October 1864). The Council (Board of Directors) soon established a schedule for regular = meetings and made arrangements for the development s = of “The Cabinet’, a series of cases displaying natural ARE Aer mes = E : 3 E 3 ‘oO oo av 2 >) history specimens which were to be held in the rooms eg Weipa) ds se 2 they rented at the Mechanics Institute building on ea Bg Bi = & mm Ss Sparks Street. Van Cortlandt was elected museum coho areas Re) ead Teal curator for the Society and remained in that position throughout the life of the organization. s Although precise records are sketchy, records at a os = Library and Archives Canada indicate that the Society eS ees Saas : Xe) os oY & o met regularly for lectures and field trips. The last C2 Milfsseehy ssi Sho 2 Saturday of each month was fixed for the latter (LAC Ss 8 ae S = = OS OFNC Collection, 28 April 1865), during which mem- chr een ee areal i age bad ifs teal bers actively searched for natural history specimens for The Cabinet. Indeed, this seemed to be a major focus s of the organization (Dore 1968). In October 1869, only wee & months before the organization’s amalgamation with ur 2 ge gel es 5 the Mechanics Institute, Braddish Billings offered his ca) = = Se Ss a herbarium to the ONHS for $60 (LAC OFNC Collec- Ra ES Es & = & A s tion, 1 October 1869). Although no supporting motion Co Zagtesie tes te ee} (e\ al for such a purchase is noted, Billings’s collection did end up there (it may have been deposited after his 3 death), and was examined in the OLSS museum in the 5 oa thie & 1880s (Fletcher 1888). Unfortunately, the OLSS herbar- e [2+ Bee Fs 5 ium, including the Billings’s specimens, disappeared 2 bere seee 2 sometime in the early 20" century (Dore 1968). g Ali olepese Haya daeiply The ONHS seems to have been most active between oe ea PA (as esis ea eh teal 1865 and 1867 and to have achieved considerable ic} standing in the community. In 1866 a group met with S D es federal Minister Thomas Darcy McGee to discuss a oe in 2 possible ONHS — Mechanics Institute exhibit in the = 8 eae te % : 1867 Paris Exhibition (LAC OFNC Collection, 26 SS ie a Be a2 S April 1866). Their paths had crossed before, in Que- >a Ie a ae os 3 bec City in 1862-1863, McGee had lectured to the Ss1% BS =S The Club continued to struggle financially despite the generally rosy economic picture in the country, as it also had, ironically, during the “Roaring Twenties”. Vol. 118 In the mid-1950s, for example, Treasurer R. J. Moore advised the Council “to go easy, especially keep Excursions & Lectures Committee under control” (LAC OFNC Collection, 29 February 1956). Walton Groves’ assessment was even more brutal, stating that “many of the Club’s activities have been crippled by the delay in publication” and by the need to publish sO many issues in such a short time (LAC OFNC Collection, 30 September 1956). Despite the return to timely publication of the CFN, at the end of the decade Groves still reported that “the financial position of the Club continues to be very grave” at (LAC OFNC Collection, 1 December 1960). Still, the OFNC managed a Special Issue of The CEN, Alice Wilson’s Geology of Ottawa (Wilson 1956). It was likely only possible because Carleton University placed a pre-publication order for 1000 reprints (LAC OFNC Collection, 13 April 1956). Before environmental conservation and the protec- tion of natural biodiversity became major social and cultural phenomena in the Western World in the 1960s, the locally-focused elements within the OFNC had been gradually increasing the organization’s level of involvement in such directions. In the early 1950s, for example, Harrison Lewis met with the Federal District Commission (FDC) to discuss how the OFNC could (and would) provide wildlife protection and enhancement consultation during Gatineau Park dev- elopment. They also explored the idea of an OFNC field station space being provided in an expropriated Gatineau Park building (LAC OFNC Collection, 7 April 1951). An ORNC Gatineau Park Committee was formed under the chairmanship of OFNC 1960-1961 President Winston Mair and made various recom- mendations for wildlife habitat enhancement there. The Federal District Commission also asked the Club to be involved in the establishment of a Brewery Creek bird sanctuary in Hull (Gatineau), presumably in res- ponse to the high profile that site had received from former British High Commissioner Malcolm Mac- Donald’s war time reminiscences in Birds of Brewery Creek (MacDonald 1947) (LAC OFNC Collection, 13 March 1952). Nothing came of that and neither FDC consultation was mentioned again after 1954 (LAC OFNC Collection, 15 December 1954). Conservation priorities (late 1960s-early 1980s) A forerunner of the OFNC Conservation Commit- tee, called the Preservation of Natural History Sites Committee, was established in 1960 with Bill Baldwin as its chair and with locally-active members including biology professor Donald A. Smith, CWS biologist Vic Solman, geologist D. D. Hogarth and botanist W. G. Dore (1912-1996) (LAC OFNC Collection, 15 December 1960). One of its first ventures — opposing the proposed destruction of the vast Mer Bleue peat- land in eastern Ottawa (it was to be used as a regional landfill!) — was highly successful. It led not only to 2004 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 23 FIGURE 15. Members of the Traill Group near Pinks Lake 29. May 1965, one of the last outings before disbanding as a formal group. From left to right are Winifred Anderson, Ruth Resenel, Alice Frith, Sheila Thomson, Bill Thomson, Rowley Frith, Hue McKenzie, Elva MacKenzie, Anne Hanes and unknown. Photograph by Charlotte Dill. Trail & Landscape 16(2): 95 [1982]. protection of that magnificent natural area, but also to a series of ecological and biodiversity investigations of the wetland (Baldwin and Mosquin 1969). Donald Smith played a pivotal role in generating the neces- sary awareness amongst both the naturalist commu- nity and National Capital Commission (NCC) officials of the significance of the bog (Dorais et al. 1974). The protected area — now some 3 500 ha in size — is managed by the NCC for the benefit of its ecological functions and has been declared an internationally significant wetland under the Ramsar Convention on Wetlands (National Capital Commission 1996). In this period local Club members and the general public benefitted from two birding columns in Ottawa newspapers, one in The Ottawa Citizen by Wilfred Bell and the other in The Ottawa Journal by the ap- propriately named John Bird. The latter weekly column, “Bird’s-eye View”, was particularly beautifully written, accurate and insightful. It emphasized the growing level of birding activity in the Ottawa Valley and offered a timely (unofficial) bulletin board of OFNC and conservation events (McNicholl 1994). Certainly the most important single occurrence for the local Club element in this period was the estab- lishment of Trail & Landscape (T&L) (Figure 14). As expressed by the then CFN editor Theodore Mos- quin, it was to be “a newsletter as well as contain[ing] articles of wide appeal in the conservation field” (LAC OFNC Collection, 5 January 1967). The existing OFNC Newsletter wasn’t offering sufficiently comprehensive local coverage, nor could it satisfactorily accommodate the documentation of local conservation-oriented nat- ural environment investigations which were increas- ingly being undertaken by Club members. T&L was an immediate and astonishing hit, not only satisfying a need of active local Club members but generating a huge, unprecedented increase in membership (Figure 6). Anne Hanes (1925-1981) was appointed as the first editor and exercised these duties superbly through 13 volumes (1967 to 1979), always encouraging environ- mental conservation themes wherever possible (Green- wood 1980). Two subsequent editors, Joyce Reddoch (from 1980 to 1989) and Fenja Brodo (from 1991 to 1992 and 1993 to 2001) expertly supervised and directed the develop- ment of long runs of the publication. Reddoch’s term was distinguished by the production of a number of large review articles which she solicited for both their intrinsic natural history value and their importance as reliable references for various conservation applications thereafter. A number of these, such as the reviews of Ottawa District amphibian and reptile species (Cook 24 THE CANADIAN FIELD-NATURALIST 1981, a revision of an earlier series in 1967 solicited by first editor Anne Hanes) and butterfly species (Layberry et al. 1982), continue to be locally referenced, as do all of the natural areas documentation published in T&L. The Club and its members have also been well served by other editors of Trail & Landscape, Elizabeth Morton (1990), Bill Gummer (from 1992 to 1993) and Karen McLachlan/Hamilton (from 2001 to the present). Trail & Landscape quickly became the contempo- rary source for information on local natural environ- ment issues. It has provided an ideal bridge between the recording of natural environment information for its own sake and in directing and assessing conserva- tion priorities (Brunton 1986b). To some, the timeliness and thus effectiveness appears to have suffered, how- ever, after a decision to reduce the publication sched- ule, for manpower and economic reasons, from five issues/year to four, starting with the 1988 volume. The subsequently reduced publication frequency and fewer total pages have coincided with a substantial reduction (by almost half) in the number of articles with direct conservation applications. The huge increase in membership and higher profile of conservation and field-oriented activities in the late 1960s may have set too hectic a pace for some long- time Club stalwarts who had been more concerned with the national publication role. Hoyes Lloyd resigned from the Council at the end of 1967, stating that he felt it inappropriate as an OFNC Honorary Member to also serve as a Club director (LAC OFNC Collec- tion, 6 June 1967). There is no such limitation on Hon- orary Members, however, and a number have contin- ued to serve on Council with distinction for many years. Notable in this regard is 1979 Honorary Member Bill Cody, who has served continuously on Council since December 1947, shortly after becoming The CFN Business Manager earlier that year (LAC OFNC Col- lection, 21 May 1947). And similarly impressive is the contribution of Frank Pope, who has served on Council continuously since 1980 and has occupied the positions of Corresponding Secretary (three years), Treasurer (six years), Vice-President (one year) and President (an unprecedented six years) (Appendix 1). The Canadian Field-Naturalist underwent a signif- icant reconfiguration in January 1970, introducing a new size, format and appearance not just to improve its look but “to [make] its content as relevant as pos- sible to the natural history needs of our time” (Mosquin 1970). And by “relevant”, Editor Ted Mosquin meant strongly conservation oriented. That first issue was graced by a photo with a Timber Wolf — the contem- porary symbol of wilderness protection — crossing a snowy landscape. The issue contained both a variety of articles on Canadian endangered species and a direc- tory of conservation organizations in Canada. Conser- vation applications of carefully-researched ecological and natural diversity investigations have been a hall- mark of The CEFN ever since. Vol. 118 Both as OFNC 1969-1971 President and editor of the CFN from 1967 to 1972, Mosquin was at the fore- front of efforts to enhance the level of OFNC conser- vation action and to engage both the membership and the general public in this. OFNC 1971-1972 President Sheila Thomson and OFNC 1972-1975 President Irwin Brodo were similarly motivated conservation voices in the OFNC, the three of them working togeth- er to persuasively direct Club environmental protection initiatives. Mosquin initiated a series of at times hard- hitting and at times controversial editorials (not the last we were to see in the CFN!), challenging Club members, the larger naturalist community, and public decision-makers to be more effective in protecting natural environments in Canada. In 1972 he resigned the CFN editorship to take on the challenge of found- ing editor of the Canadian Nature Federation’s Nature Canada. The infectious enthusiasm, environmental passion and unquestioned technical expertise of these three activists in concert with other Club leaders of the day such as OFNC 1963-1966 President George McGee (1909-1991) and OFNC 1967-1969 President Hue McKenzie, were an inspiration to a whole generation of new Ottawa-area naturalists (personal observation). For us newcomers, they transformed the understand- ing of the study of the natural environment from a seemingly slightly idiosyncratic, individual endeavour undertaken for personal entertainment to a meaning- ful — even important — and highly integrated calling. What a revelation! The growing commitment to natural environment conservation in the late 1960s and early 1970s saw the re-energizing of OFNC Council and membership support in this area. At the urging of Ted Mosquin and Ed Greenwood and recognizing that the Preservation of Natural History Sites Committee was not broad enough in its mandate, in early 1967 the OFNC estab- lished the Natural Areas Committee to coordinate original field conservation research and to actively search out potential natural reserve sites in the Ottawa Valley (LAC OFNC Collection, 10 April 1967). By 1970, the committee was undertaking a broad program of local field investigations and had provided a basis for the identification of significant natural areas in the Regional Municipality of Ottawa-Carleton’s 1974 Official Plan. So significant was the role and mandate of this group that the Natural Areas Committee was soon being promoted as the most significant of Club committees (LAC OFNC Collection, 8 December 1970). Increasing demand for input into various con- servation issues both locally and beyond, resulted in the establishment of the Research and Briefs Commit- tee (1972-1973), which in 1974 was combined with the Natural Areas Committee to form the Conservation Committee. This committee continues to play an inte- gral role in OFNC and community affairs to the present day. 2004 The OFNC Centennial in 1979 understandably brought forth a great deal of interest in the history of the Club in particular and of the naturalist commu- nity of Ottawa in general. Trail & Landscape provided the major vehicle for sharing this information amongst the local membership. Events culminated with a ban- quet intended to be on the exact 100" anniversary, but as with previous celebrations, it was erroneously held on 19 March (Reddoch 1979a), not the actual centennial date of 25 March. The increased interest in the historical record gen- erated by the Centennial year encouraged a new appre- ciation of the designation of OFNC Honorary Mem- bers. These had been given out only irregularly over the years to honour lengthy, significant service to the OFNC or Canadian natural sciences. Although honor- ary memberships have been awarded more regularly since 1971, the Centennial Year saw a new focus on this acknowledgment of such exceptional contributions (Brunton and Gummer 1987). The conservation initiatives of the 1970s and early 1980s saw a resurgence of more systematic field examinations of uncommon and ecologically significant habitats in the Ottawa Valley such as fens, rivershore communities, and relict woodlands (Dugal 1978; Reddoch 1979b; White 1979). Publication of the first complete, annotated checklist of the vascular plants BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 25 of the Ottawa District also reflected this knowledge (Gillett and White 1978) and contained many of the records newly discovered by Club members. That burst of new knowledge and energy demon- strated that suggestions in the early years of the 20" century that our knowledge of the natural environ- ment of the Ottawa area was all but complete, were badly incorrect. Our understanding of native biodi- versity and of the ecological systems supporting it requires continual inventory and evaluation to enable effective protection and maintenance of this natural heritage. The Club’s conservation activities in this era, led by activist presidents Roger Foxall and Roger Taylor between 1978 and 1982 and by such energetic mem- bers as Loney Dickson, Allan Reddoch, Joyce Red- doch, Albert Dugal, Stephen Darbyshire, and David White, covered an amazing spectrum and exhausting number of subjects. These ranged from a campaign for a conservation area system in the Region of Ottawa- Carleton (now city of Ottawa) to prevention of the importation of Raccoon-dogs into Canada, and from the prevention of 1988 Winter Olympic facility devel- opment within Banff National Parks to consultations with federal government officials on ecological re- quirements in Gatineau Park and National Capital Greenbelt planning. Ficure 16. Excursion of Ottawa Field-Naturalists to the Rideau Trail in October 1979 (photograph by C. Beddoe). Contrast with Figure 9. Long-time member Mary Stuart is second from left. 26 THE CANADIAN FIELD-NATURALIST The Club hosted the annual meeting of the Feder- ation of Ontario Naturalists in 1983, establishing a successful program format that was followed by sub- sequent provincial gatherings for years to come. The OFNC again successfully hosted the provincial orga- nization’s annual conference in 1993 (Pope 1993). A more whimsical contribution is to wildlife con- servation initiated in this period was the ““Seed-a-thon’”’, a fund-raising effort whereby sponsors pledge a par- ticular amount for every species observed on a par- ticular day by a particular birding team, to be applied towards the cost of supporting a network of OFNC public bird feeder stations. Initiated by the Birds Com- mittee on the suggestion of long-time OFNC member Violet Humphries in 1981 (personal communication), this was one of the first of what now are commonplace fund-raising events across Canada (Brunton 1981). There was a human cost to all this activity, how- ever, and many of the lead Club personnel in these matters eventually were called away by the mundane but critical matters of family and careers, moving on to other things and/or were simply exhausted by it all. This was exacerbated in the later years of the 1970s and into the 1980s by new pressures on the time and resources of professional research staff within the federal public service, reducing their availability and energy for involvement in such initiatives as local conservation and research. Only three OFNC presidents since the mid 1970s (D. F. Brunton, R. John and E. Zurbrigg), for example, were professionally involved in natural environment investigation or management (Appendix 1). One of the most satisfying conservation ventures of recent decades has to be the effort begun in the early 1980s to save the massive, Provincially Signifi- cant Alfred Bog east of Ottawa (Cuddy 1983) from destruction by agri-business interests. This battle saw the Club purchase conservation land for the first time (in 1982) so as to have legal standing in the fight to secure proper protective zoning. It was a long and difficult effort but with the effective leadership of OFNC President Frank Pope, considerable financial and material contributions from numerous individual members, and the critical involvement of the Nature Conservancy of Canada, the entire wetland was saved and is now either in protective public ownership or zoned as parkland (Pope 2002). The primary national contribution of the OFNC continued to be led by the publication of the CEFN, al- though as noted, direct involvement in national-scale conservation issues was common enough in this peri- od. Despite a continuing and perhaps even enhanced national/international status and production standard, however, the seemingly unanswerable question of how national or local the journal should be was raised again in a series of editorials in the late 1970s. The appro- priateness and credentials of a citizen-based (‘‘ama- teur”) organization directing the affairs of the journal was severely criticized by Lorraine Smith, Editor 1972- Vol. 118 1981, who suggested that “the burden of being account- able to the scientific natural history community for a national journal ... may now be more than this local club should retain.” [The days of calling the OFNC a “scientific society” were truly gone!] “Therefore... if the journal’s reputation is not maintained and there is no move for a change in publisher, I challenge those who want a top-quality Canadian natural history jour- nal, such as The Canadian Field-Naturalist is now, to initiate a new journal” (Smith 1981). These comments do not explain, however, how the OFNC managed to maintain and enhance such a fine journal for so many years. After much discussion, consultation and review by an Ad Hoc Committee on OFNC Publications com- posed primarily of scientific specialists, the OFNC Council approved (8 December 1982) a new Publica- tions Policy to guide the CEN and other Club publi- cations (Bedford 1983). The Policy reiterated the Club’s continuing, century-long commitment to the scientific integrity and high technical standards of the CFN, as well as the Club’s intention to have the journal remain the official publication of the OFNC. Francis Cook assumed editorship of the CFN from 1962 to 1966, and again in 1981, and has guided the journal to the present day with a steady and depend- able hand, introducing important innovations such as the publication of Canadian status reports for candidate endangered species. There has been a great increase in the amount of material published in each volume during Cook’s tenure (volumes averaged 756 pages over the five years ending in 2002). In the face of the many complications in the publication process, the timely publication of some issues has not occurred. Scientific and technical quality has never been sacri- ficed, however. The 28 volumes edited by Cook repre- sent by far the longest and most prolific service of any CEN editor in the 124-year history of the publication, exceeding by more than a decade the tenure of the next longest-serving editors, Harold Senn and Arthur Gibson (Brunton 1986a). Contemporary Times and the Future (mid- 1980s to date) James Macoun, Percy Taverner and other “nation- als” who worked so hard to enhance the mandate, effectiveness, and reach of The Canadian Field- Naturalist would be relieved to see that the publica- tion has achieved the solid reputation, steady produc- tion, and high scientific standards they hoped to see. An interesting expression of that long-term contribu- tion and durability is demonstrated in the number of original descriptions (diagnoses of new taxa) which have occurred in the pages of the CFN. Over 730 have been published in the first 100 volumes (Brunton 1987). The level of field-oriented investigation of the Otta- wa Valley natural environment that was so productive in the 1970s and early 1980s, however, has been dras- 2004 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB Number of Members | SS ES SSS CY MSY ET | TS prerereeco FiGureE 17. Ottawa Field-Naturalists’ Club membership trend 1975-2003. Numbers of institutional subscribers to The Canadian Field-Naturalist were not calculated within membership totals nor reported separately in Annual Reports after 1971. A 1975 reorganization offered individual members (reported to total 1371 that year) the opportunity to be only non- voting subscribers to the The Canadian Field-Naturalist, dramatically reducing the OFNC membership total reported for 1976 (Erskine 1977) [Individual and institutional subscribers were reported separately in the Annual and Editor’s Reports for The Canadian Field-Naturalist in each volume]. Absolute membership numbers prior to 1976, therefore, are not directly comparable to these of subsequent years (from OFNC Annual Reports, 1975-2002). tically reduced. The appearance of substantially fewer area inventories and record documentation articles in Trail & Landscape reflects that trend. This may be part of an unfortunately strong North American perception in recent decades that continuing, original field in- vestigations are of less importance than in previous times. T&L remains, however, an important and highly relevant source of environmental news and natural environment documentation. Programs intended to actively involve the partici- pation of local members have remained an important part of the OFNC. Dozens of field outings, lectures and special events are undertaken each year involving a diversity of well-informed leaders and speakers. For decades the Club’s Excursions and Lectures Com- mittee has crafted an interesting and instructive program that contributes greatly to the enhancement of members’ skills and to their enjoyment in the field. While field investigations focus on both familiar and exotic local- ities throughout the Ottawa District and the Ottawa Valley, the Club does not confine itself to this area. The spring “migration” of a bus load of OFNC mem- bers to Point Pelee National Park, for example, has become a popular biennial birding event since the 1980s. This is important stuff. The majority of Club mem- bers are locally based and most of us are involved, first and foremost, because we enjoy exploring and learning about our natural world. Providing support for scientifically important undertakings such as the publication of the CFN and the protection and preser- vation of important natural landscapes, is a valuable additional membership benefit. Membership in the OFNC remains an easy and enjoyable way for indi- vidual citizens to support greater understanding of, and protection for, important natural landscapes and features in the Ottawa Valley and beyond. A noticeable change in the role of the local Club in regards to its community has occurred in recent years. The OFNC is much more integrated into environmen- tal decision-making at a municipal and provincial level than it has been at any time in its past. It has become, in an sense, an ecological consultant to that community. While such a role requires seemingly end- less attendance at meetings and reduced time for core field work, it has provided many excellent opportunities for Club representatives to bring accurate ecological information into the discussion of an issue at an early and effective stage of the decision-making process. A 28 THE CANADIAN FIELD-NATURALIST cynic might say that the Club has become somewhat bureaucratized; an optimist would suggest that the Club now more frequently has a seat at decision-making tables across the National Capital Region. It is impor- tant to note, for example, that it was through this per- iod that the final protection of the important Alfred Bog natural area was achieved. Another very positive achievement of the last decade was the establishment of the Louise de Kiriline Law- rence Conservation Action Fund in 1994, founded with a generous bequest from the estate of that outstanding and eloquent field naturalist, Louise de Kiriline Law- rence (1894-1992) (Ainley 1994). It is sustained by further donations and by the profit from the sale of particular OFNC products such as the “Natural Areas” book (Brunton 1988). The fund provides for “stra- tegic and timely expenditures towards the conservation of natural areas” in the Ottawa Valley (Anonymous 1994). A continuing interest for many Club members is the Fletcher Wildlife Garden (FWG), established in 1987 in degraded woodland and regenerating agricul- tural land on National Arboretum property near the Rideau Canal. With the active cooperation and assis- tance of Central Experimental Farm (Agriculture Canada) personnel who are responsible for land man- agement here, the Fletcher Wildlife Garden volun- teers have transformed the site with native plantings, the creation of a pond and through public interpre- tation. The objective is not only to produce on-going, evolving habitat restoration but to demonstrate suita- ble wildlife-friendly landscaping and gardening prac- tices for National Capital residents and visitors alike — to be “a model for urban gardeners” (Conservation Committee 1987). These are all positive features. A disturbing trend in Club affairs, however, has been declining membership over the last two decades (Figure 17). Perhaps not coincidentally, Club membership peaked ca. 1982 to 1985, at the end of the period of greatest conservation- oriented activity. The local membership decline is seemingly in tandem with the decline of original field- based investigations. Nonetheless, the number of OFNC members remains substantial — the Club is still probably the largest regional natural history organi- zation in Canada — but the downward membership trend needs reversing (Figure 17). The reasons for this decline in participation are un- clear. It seems unlikely to be attributable to the ready availability of up-to-date, comprehensive natural en- vironment data on the World Wide Web since the mem- bership decline began well before such technology was widely available. It may, in part, be a function of some wider, societal disinterest in hands-on involve- ment with citizen-based, research-oriented groups. Whatever the reason(s), the decline has been substan- tial and if it continues, will undermine the financial security and effectiveness of the OFNC. Vol. 118 Since the 1990s the digital world has had as dra- matic an impact on the OFNC as it has on so many facets of contemporary Canadian life. We are only just coming to terms with its potential benefits and challenges. The OFNC’s impressive web page (http:// www.ofnc.ca) has allowed for rapid internal commu- nications amongst Club officials, committee members and conservation partners, and for the posting of information and timely news items. Digital commu- nication is only the latest expression of the Club’s commitment to education of our membership and of the general public in regard to important natural envi- ronment features, needs and opportunities. The impor- tance of the Club’s educational programs has been a major theme throughout OFNC history and is ex- pressed in the mission statement of the organization (see frontispiece of this issue). There is every indica- tion that it will remain so into the future. It is not yet clear how the Club will move forward from this foundation of established programs and re- sources in order to stay informed, relevant — and sol- vent — into the future. That’s not a new challenge but is one that must be answered by each generation in response to the circumstances and opportunities of their day. Some things are clear. For one, the Club needs to re-establish a higher level of field-oriented investigation amongst its membership. This will not only maintain and develop the skills and interests of present mem- bers but also inspire and generate new members. Ac- curate and timely field investigation and documen- tation are essential foundations for the maintenance of the high level of technical credibility that the Club has earned over the years. Continuing to do our home- work, getting the facts straight, and presenting such information in a defensible, objective manner are the keys to maintaining and enhancing the OFNC’s impre- ssive record of conservation achievement. As with any organization, regardless of how vener- able, the OFNC must also continue to re-examine what it provides for the individual member. This is perhaps even more critical in an era with an unprecedented number of electronic diversions competing for the attention of both members and partners alike. Will the traditional superb program of excursions, lectures, and publications be enough to inspire and encourage a growing, involved membership? Do the new technol- ogies of our time offer the same manner of new op- portunities that technologies like the railway train, electric lighting, the automobile, and the airplane offered earlier OFNC members? They surely do. What I hope is a clear message from this review of the long, remarkably productive journey of Ottawa naturalists is that the OFNC is not a static, hard-and- fast “thing”. It is a community, an association of like minds that aims to explore and celebrate the magnifi- cent natural world of the Ottawa Valley and of Canada. And as with any community, we in the OFNC have 2004 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB 29 tos FIGURE 18. Record service to The Canadian Field-Naturalist: Left: Francis R. Cook, Editor 1962-1966; Associate Editor (Herpetology) 1972-1981; Editor 1981-present; right: William J. Cody, Business Manager 1948-present. 25 August 2004. Photo courtesy Ron Bedford. and have had the benefit of the skills, insight, and inspirational leadership of numerous individuals and groups over the years. In weak times and strong, the common thread that has bound Ottawa naturalists since 1879 — indeed, since 1863 — is an overwhelming desire to contribute to the protection and enhancement of the natural features and values that make this place so special. At the end of the day it comes down to the fact that James Fletcher, Frank Pope, Percy Taverner, Sheila Thomson, Edward Van Cortlandt, Herb Groh, Gordon Hewitt, and the rest of us are not that different ... we’re all just Ottawa field naturalists. Acknowledgments The daunting challenge of reviewing a manuscript of this nature was undertaken in a timely and helpful manner by a team of naturalists well familiar with the OFNC and OFNC personalities. These reviewers were Ron Bedford, Bill Cody, Francis Cook, Karen McIntosh, Elizabeth Morton, Frank Pope and Joyce Reddoch. They added much to the effort and I gratefully ack- nowledge their input. I remain, however, fully respon- sible for any errors or omissions which may have escaped their sharp eyes. Frank Pope and Francis Cook provided additional factual input throughout the development of the manuscript. This review has also benefitted from the innumerable discussions (and debates!) I have enjoyed with Francis Cook over the last 25 years on numerous subjects related to the his- tory of the OFNC and to the study of Canadian natural sciences. Ron Bedford, Frank Pope, Joyce Reddoch, and Rob Lee (present chair of the Macoun Field Club) were instrumental in obtaining many of the illustra- tions employed in this article. Rob Lee and Joyce Cook scanned several of these. I am happy too, to acknowl- edge two other important players in this effort. First is the late Herb Groh whose clipping file on OFNC personalities and whose interest in the Club’s past inspired my own curiosity. The other gratefully ack- nowledged is the staff at the Library and Archives Canada (formerly the National Archives of Canada) who so efficiently and pleasantly facilitated my research with the OFNC collection and related files held within the walls of that national treasure. Documents Cited (marked * in text) Dorais, M., R. Bouffard, R. Chenier and K. Adamson. 1974. An examination of the past and present status of the Mer Bleue Peat Bog near Ottawa, Ontario. [Unpublished report], Faculty of Science and Engineering, University of Ottawa. 105 pages. 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Macmillan of Canada, Toronto. 283 pages. Dill, C. 1979. The Club Years 1950-1959. Trail & Landscape 13: 112-120. Dill, C. 1982. From Bog Group to Trail[{l] Group 1955 to 1965. Trail & Landscape 16: 94-96. Dore, W. G. 1968. Reprint of “List of plants collected by Mr. Braddish Billings in the vicinity of the city of Ottawa during the summer of 1866”. Plant Research Institute, Department of Agriculture, Ottawa. Dugal, A. 1978. The South Gloucester Conservation Area revisited. Trail & Landscape 12: 47-53. Dymond, J. R. 1939. The fishes of the Ottawa Region. Contributions of the Royal Ontario Museum of Zoology (15). Toronto. 43 pages. Eggleston, W. 1961. The Queen’s Choice, a story of Canada’s Capital. The National Capital Commission, The Queen’s Printer, Ottawa. 325 pages. Erskine, A. J. 1977. Report of Council to The Ottawa Field-Naturalists’ Club. Canadian Field-Naturalist 91: 212-218. Fletcher, J. 1880. Flora Ottawaensis. Transactions of The Ottawa Field-Naturalists’ Club 1: 48-61. 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Trail & Landscape 14: 2. Groh, H. 1955. “Let us now praise famous men” Canadian Field-Naturalist 69: 75-78. Gwyn, S. 1984. The Private Capital. McClelland and Stewart Ltd., Toronto. 514 pages. 2004 Hall, P. 1986. The Central Experimental Farm and The OFNC: a 100 year cross-fertilization story. Trail & Landscape 20: 146-151. Harper, F. 1925. Correspondence [Wood Buffalo]. Canadian Field-Naturalist 39: 45. [Harrington, W. H.] 1887. Editorial Announcement. Ottawa Naturalist 1: 9-10. Harrington, W. H. 1909. Reminiscences of Dr. Fletcher. Ottawa Naturalist 22: 196-205. Humphreys, V. 1979a. Audubon Screen Tours: general his- tory of the tours in Ottawa. Trail & Landscape 13: 99-101. Humphreys, V. 1979b. The OFNC Lodge at Beattie [sic] Point. Trail & Landscape 13: 4-6. Killan, G. 1993. Protected Places, a history of Ontario’s prov- incial park system. Ontario Ministry of Natural Resources, Dundurn Press, Toronto. 630 pages. Kindle, E. M. 1928. 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Mammals of the Ottawa District. Canadian Field-Naturalist 59: 111-132. Reddoch, J. M. 1979a. The birthday party. Trail & Landscape 13: 132-134. Reddoch, J. M. 1979b. Calcareous fens in the Ottawa District. Trail & Landscape 13: 16-27. Reddoch, J. M. 1979c. Favourite OFNC excursion places. Trail & Landscape 13: 71-96. Reddoch, J. M., and A. H. Reddoch. 1997. The orchids in the Ottawa District: floristics, phytogeography, population studies and historical review. Canadian Field-Naturalist 111: 1-185. Regehr, T. D. 1988. Sir Sandford Fleming. Pages 790-791 in The Canadian Encyclopedia (second edition), Volume 2. Hurtig Press, Edmonton. Rothfels, C. 2003. Synopsis of Ontario herbaria. Field Botanists of Ontario Newsletter 16: 7-19. : S[hutt], F. T. 1912. William White. Ottawa Naturalist 26: 13% Small, H. B. [Jr.] 1929. In memoriam, Col. W. P. Anderson. Canadian Field-Naturalist 43: 64. Small, H. B. [Sr.] 1883. Fishes of the Ottawa District. Trans- actions of the Ottawa-Field-Naturalist’s Club 4: 31-49. Smith, L. C. 1981. Editor’s farewell musings. Canadian Field-Naturalist 95: 233-235. Snure, P. 1978. The Club Years, 1940-1949. Trail & Land- scape 12: 34-44. Solman, V. E. F. 1974. Harrison Flint Lewis, 1893-1974. Canadian Field-Naturalist 88: 507-516. Soper, J. H., and E. L. Bousfield. 1982. A tribute to William Kirwan Willcocks Baldwin, M. B. E. (1910-1979). Cana- dian Field-Naturalist 96: 92-97. Taschereau, P. M. 1972. In Memoriam: Herbert Groh. Canadian Field-Naturalist 86: 299-306. Taylor, J. H. 1986. The history of Canadian cities Ottawa: An illustrated history. James Lorimer & Company, Pub- lishers and Canadian Museum of Civilization, National Museums of Canada. Toronto. 232 pages. Taylor, R. 1979. President’s Message. Trail & Landscape 13230-3ilk Travill, A. A. 1988. Sir James Alexander Grant. Page 927 in The Canadian Encyclopedia (second edition), Volume 2. Hurtig Press, Edmonton. Tufts, H. 1898. Notes on the birds of King’s Co., Nova Scotia, [Part I]. Ottawa Naturalist 12: 172-177. Tufts, H. 1899a. Notes on the birds of King’s Co., Nova Scotia — Part II. Ottawa Naturalist 12: 229-233. Tufts, H. 1899b. Notes on the birds of King’s Co., Nova Scotia — Part II. Ottawa Naturalist 12: 259-262. White, D. J. 1979. The flora of Innis Point. Trail & Land- scape 13: 174-177. 32. THE CANADIAN FIELD-NATURALIST White, G. R., and W. L. Scott. 1882. List of birds found in the vicinity of Ottawa City, specimens of which have been shot within the last few years, Appendix to Report of the Ornithological and Oological Branch for the season of 1881. Transactions of The Ottawa Field-Naturalists’ Club 3: 29-34. Whiteaves, J. F. 1876. Obituary notice of Elkanah Billings, F. G. S., paleontologist to the Geological Survey of Can- ada. Canadian Naturalist and Quarterly Journal of Science 8: 251-261. Whyte, R. B. 1880. Annual Report. Transactions of The Ottawa Field-Naturalists’ Club 1: 7-11. Whyte, R. B. 1909. Dr. Fletcher as a botanist. Ottawa Nat- uralist 22: 206-207. Wilson, A. E. 1956. A guide to the geology of the Ottawa District. Canadian Field-Naturalist 70: 1-68. Wily, T. 1869. Swallow Lore. Transactions of the Ottawa Natural History Society (3). Ottawa. Zaslow, M. 1975. Reading the rocks, the story of the Geolog- ical Survey of Canada 1842-1972. Department of Energ Mines and Resources, Ottawa. 599 pages. Received 22 June 2004 Accepted 10 August 2004 END NOTES: ' Number 1 (un-numbered), Braddish Billings’ Ottawa vascular plant list (Billings 1868) was published between March and November 1868; Number 2 (numbered), 500 copies of Sir J. A. Grant’s examination of the surficial geology of the Ottawa Valley (Grant 1868) were published December 1868; Number 3 (numbered), Thomas Wily’s discussions of swallows (Wily 1869) was published De- cember 1869 (LAC OFNC Collection, 10 December 1869; Dore 1968). Much later, Henry Ami reminiscing about events leading up to the formation of the Club described *... how the Ottawa Club originated from the Epping Forest and Field Club, and later Ottawa Naturalists Club ...” (LAC OFNC Collection, 2 February 1924). Ami may, however, have been referring to names applied to informal groupings of field associates, such as the Fletcher, Harrington et al. group which conducted field work together in the Billings Bridge area in the late 1870s (Harrington 1909). There is no further mention of these groups in OFNC records by any of the individuals who, unlike Ami, were actively involved in the formation of the Club. Later, R. B. Whyte erroneously reported this as occurring on 19 March 1879 (Whyte 1880). He may have been referring to an earlier preliminary meeting or to the date of the OLSS Council meeting where Fletcher and Henry Small Jr. obtained an expression of the co-operation and assistance from the parent organization, rather than the formal orga- nizational gathering. Being documented only one year after the fact and by a Club founder and the originator of the idea of the OFNC, this error would quite understandably be accepted as fact and was repeated for over 100 years as the formal starting date of the OFNC (e.g. Groh 1955; Taylor 1979). + Relations between The Ottawa Field-Naturalists’ Club and The Ottawa Literary and Scientific Society appear to have been strained on at least several occasions in the following years. In his 30 November 1886 Inaugural Address, for example, OLSS President W. P. Anderson complained nN Vol. 118 that the OFNC (of which he was a founding member and had been the Treasurer for the previous three years!) was “antagonistic to the Ottawa Literary and Scientific Soci- ety” (LAC OFNC Collection, 3 December 1886). The OFNC and OLSS councils individually discussed the prob- lem, the OLSS Council subsequently stating that they would consider “ ... any proposition which the field Club may wish to make looking for a change in the relations between the Club and the Society but that a joint meeting of the two Councils cannot conveniently be arranged ...” (LAC OFNC Collection, 14 January 1887). Although no such joint meeting or other formal Club action resulted, the OFNC Council continued to meet in the OLSS museum until March 1890. That was despite another minor dust-up in 1889 concerning the OFNC’s offer to cover some of the public lecture programming required of the OLSS in order to maintain its Ontario government grant (LAC OFNC Collection, 6 November 1889). The involvement of women in the OFNC was very much promoted in these early years — in a patronizing if sincere way. The launch of The Ottawa Naturalist was accompa- nied by a special plea for the involvement of women ... “especially will be ladies welcomed to our ranks, and every effort will be put forward to make the excursions and soirees pleasant as well as instructive” (Harrington 1887). At the March 1888 Annual Meeting “... a discussion en- sued as to the eligibility of ladies as officers, and the desireability of having some on the Council. Professor Macoun gave notice that he would at the next General meeting of the Club move that the executive Committee consist of six members, three of whom shall be ladies” (LAC OFNC Collection, 20 March 1888). This was accom- plished by a Constitutional amendment in March 1890. Margaret A. Mills became the first female OFNC Officer when “Miss Mills” was elected Second Vice-President in 1892 (LAC OFNC Collection, 15 March 1892). She was not present at the meeting, however, and was apparently not agreeable to her election, as she resigned effective the next Council meeting 9 days later (LAC OFNC Collection, 24 March 1892), but stayed on as a Club member for several more years. “This [Ottawa] District was then [1880] understood to mean a radius of about twelve miles from the City of Ottawa ... latterly, however, with the general consent of the botanists of the club, this radius has been extended to about 30 miles ...” (Fletcher 1888). The Ottawa District boundary was metricated in 1981 to a 50 km radius circle centred on the Peace Tower of the Parliament Buildings (Anonymous 1981), including landscape within both south- eastern Ontario and southwestern Quebec. The portrait hung for many years in the Geological Survey Museum (later, National Museum of Canada). It now can be seen on display in the Logan Gallery of the Geological Survey of Canada headquarters on Booth Street in Ottawa. The saga of the OFNC library does not end there, however. In 1948 the Ottawa Public Library (OPL) reported to the Club that they had discovered “... the old library of the Club, now deposited in a storeroom adjacent to the main library” and that several hundred dollars worth of Canadian Field-Naturalist (CFN) back numbers were recovered (LAC OFNC Collection, 29 October 1948). The Council agreed to thank the OPL for storing the collection (since 1917!) by donating a set of The CFN back to 1935. Although the OPL apparently offered to continue to store 2004 a oO the 24 shelves worth of material at no cost to the Club, Council determined that all of the natural history and geology books should be sold (LAC OFNC Collection, 22 November 1948). No one was aware, apparently, that the Club had given the library to the OPL back in 1917 and thus was obviously in no legitimate position to sell the material. Nonetheless, the library was sold to “Mr. [Bernard] Amtmann, a dealer in second hand books ...” who assured the Council that OFNC members “would be given first choice before the [collection] catalogues were sent out ... through his extensive mailing list” (LAC OFNC Collection, 19 January 1950). The Club’s initial share of sales before Amtmann moved to Montreal with the collec- tion was $169 (LAC OFNC Collection, 20 November 1950); the following year Amtmann offered the Club $200 for the remaining volumes of the library which he had sold off “on a poundage basis” and for which he claimed to have incurred a considerable loss. The offer was accepted (LAC OFNC Collection, 16 November 1951) ... a sad ending to what must have been a magnificent collection with an intriguing provenance. The portrait by Franklin Brownell was commissioned in late 1911 (LAC OFNC Collection, 18 December 1911) and hung in “a suitable and prominent place” in the Ottawa Public Library in the spring of 1912 (LAC OFNC Collec- tion, 12 March 1912). In the 1920s it was transferred to the National Museum of Canada “for safekeeping” (LAC OFNC Collection, 14 December 1926) where it hung for many years in the herbarium, its actual ownership and the “temporary” nature of its placement at The Museum eventually forgotten. When this history was discovered and brought to the attention of National Museum of Natural Sciences Assistant Director (and former OFNC Vice-Pres- ident) C. G. Gruchy and Agriculture Canada staff in 1985 (personal communication), arrangements were made by The Museum for cleaning and minor restorations to be undertaken. The portrait, still technically owned by the Club and on loan to The Museum, was officially donated to The Farm on 2 June 1986 by OFNC 1986-1988 President W. G. Gummer (1915-1999) as part of the celebrations of 100 years of agricultural research in Canada (Hall 1986). The Museum and the Club had now fully honoured their commitment to the memory of James Fletcher. The portrait presently hangs, appropriately enough, in the William Saunders Building, named after the Central Experimental Farm Director who hired Fletcher as Canada’s first Domin- ion Entomologist and Botanist. The Ottawa Naturalist was published quarterly for a brief period (1889/1890) as a cost-saving measure but reverted to monthly publication (at the urging of Will Harrington) because of reader preference for more frequent produc- BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB w 15 33 tion. This was initiated despite the Club’s request for a provincial publication assistance grant being denied. An avowed Tory, John Macoun proclaimed that “‘politics had intervened with the success of the application and that we would have to learn to vote properly before we could hope to succeed ...” (LAC OFNC Collection, 18 March 1890). The none too subtle shot at the long serving Liberal govy- ernment of Oliver Mowat was quite likely an accurate observation of the politics of that day (Gywn 1984). The Ottawa Naturalist did eventually receive a provincial gov- ernment publication grant (of $300.00), beginning in 1897. But presumably to Macoun’s chagrin, this occurred with the Liberals still in power! Provincial financial assistance was received thereafter until 1924 (LAC OFNC Collection, 3 November 1924), as was generous federal publication support funding for The Canadian Field-Naturalist in the 1980s and 1990s. The Macoun Autobiography was reprinted as an OFNC Special Publication in the Club’s centennial year (Macoun 1979), with an new introduction by Richard Glover and with both editorial notes and a biographical sketch by historian William A. Waiser. P. B. Symes presented his “nearly complete set” of The Canadian Field-Naturalist and its predecessors to the Club in 1923. It formed the basis for “the Club’s set of original issues” (Patch 1923) which is maintained for the purposes of the current CFN editor. The set is now complete (Francis Cook, personal communication). When Pauline Snure was elected in 1948, she was the first woman president of the OFNC, despite the conspicuous affirmative action efforts on behalf of women members by James Fletcher and associates in the 1880s and again in the first decade of the 20" century. It was presumably this 75" anniversary that inspired Herb Groh to develop a biographical clipping file on OFNC Presidents and other officers. The file was presented to the Club in the early 1980s by Pierre Taschereau on behalf of Groh’s widow and is now preserved in the LAC OFNC Collection. The present editor has suggested (personal communication) that this treatment might be overly critical of Senn’s edi- torial tardiness in comparison to the at-times substantial delay of issues of the CFN during his (Cook’s) tenure. While recent delays sometimes approached (though never matched) those of the 1950s, the critical difference between the two was the impact on the Club in their res- pective eras. The substantial publication delay in the early 1950s all but shut down Club productivity. While not to suggest that contemporary publications delays are without impact, they are clearly less threatening than those of 50 years ago. 34 THE CANADIAN FIELD-NATURALIST APPENDIX 1: Ottawa Field-Naturalists’ Club officers (1879-2004) (derived from (LAC OFNC Collection Minute Books (1879-1970) and The Canadian Field-Naturalist listings and contem- porary OFNC Council Minutes) 1879/1880 1880/1881 1881/1882 1882/1883 1883/1884 1884/1885 1885/1886 1886/1887 1887/1888 1888/1889 1889/1890 1890/1891 1891/1892 1892/1893 1893/1894 1894/1895 1895/1896 1896/1897 1897/1898 1898/1899 1899/1900 1900/1901 1901/1902 1902/1903 1903/1904 1904/1905 1905/1906 1906/1907 President W. White J. Fletcher W. R. Riddell* J. Fletcher J. Fletcher H. B. Small Jr. H. B. Small Jr. W. H. Harrington John Macoun R. B. Whyte R. B. Whyte R. W. Ells R. W. Ells R. W. Ells G. W. Dawson G. W. Dawson G. W. Dawson F. Shutt F. Shutt EWE. Prince BE Prince H. M. Ami H. M. Ami R. Bell R. Bell W. T. Macoun W. T. Macoun S. B. Sinclair W. J. Wilson Vice-President(s)! J. Fletcher W. P. Riddell* W. D. LeSueur* R. B. Whyte W. D. LeSueur R. B. Whyte H. B. Small Jr. R. B. Whyte J. F Whiteaves R. B. Whyte F. R. Latchford R. B. Whyte J. Fletcher John Macoun S. Woods R. B. Whyte S. Woods John Macoun S. Woods* C. F. Marsan R. W. Ells H. M. Ami J. Ballantyne R. B. Whyte J. Ballantyne H. M. Ami T. S. MacLaughlin W. H. Harrington* M. A. Mills* F. Shutt R. W. Ells F. Shutt J. Fletcher F. Shutt H. M. Ami John Macoun W. S. O'Dell R. Bell D. A. Campbell W. T. Macoun A. E. Attwood W. T. Macoun A. E. Attwood* A. Halkett W. J. Wilson S. B. Sinclair W. J. Wilson F. Shutt F. Shutt Treasurer R. B. Whyte? W. H. Harrington? W. H. Harrington? W. P. Anderson W. P. Anderson W. P. Anderson T. S. MacLaughlin T. S. MacLaughlin J. Fletcher J. Fletcher J. Fletcher J. Fletcher G. Harmer A. G. Kingston A. G. Kingston .Am "Harrington ig* J. Fletcher J. Fletcher J. Fletcher J. Fletcher A. Gibson A. Gibson A. Gibson A. Gibson A. Gibson Vol. 118 Secretary(ies)” R. B. Whyte? W. H. Harrington? W. H. Harrington? W. H. Harrington W. H. Harrington W. H. Harrington W. P. Anderson W. H. Harrington W. H. Harrington T. S. MacLaughlin T.S. MacLaughlin T. S. MacLaughlin W. H. Harrington H. M. Ami H. M. Ami H. M. Ami A. Halkett A. Halkett A. Halkett W. J. Wilson W. J. Wilson W. J. Wilson W. J. Wilson* W. J. Wilson W. J. Wilson* T. E. Clarke T. E. Clarke T. E. Clarke 2004 1907/1908 1908/1909 1909/1910 1910/1911 1911/1912 1912/1913 1913/1914 1914/1915 1915/1916 1916/1917 1917/1918 1918/1919 19193 1920 1921 1922 1923 1924 1925 1926 1927 1928 1929 1930 1931 1932 1933 1934 1935 1936 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB President W. J. Wilson A. E. Attwood A. E. Attwood A. Halkett* A. G. Kingston* A. McNeill L. H. Newman L. H. Newman A. Gibson A. Gibson H. I. Smith H. I. Smith C. G. Hewitt M. Y. Williams M. Y. Williams R. M. Anderson R. M. Anderson H. Lloyd H. Lloyd H. Lloyd* G. A. Miller* N. Criddle N. Criddle E. M. Kindle E. EF. G. White H. F. Lewis H. F. Lewis C. M. Sternberg C. M. Sternberg M. E. Wilson M. E. Wilson H. Groh Vice-President(s)! A. E. Attwood A. E. Attwood A. Halkett A. Halkett E. FG. Eifrig A. Halkett E. E.G. Eifrig A. G. Kingston L. H. Newman A. Gibson L. H. Newman A. Gibson J. W. Gibson A. Gibson H. I. Smith C. G. Hewitt H. I. Smith C. G. Hewitt H. I. Smith C. G. Hewitt E. D. Eddy M. Y. Williams C. G. Hewitt L. D. Burling M. Y. Williams L. D. Burling P. A. Taverner L. D. Burling R. M. Anderson G. A. Miller H. Lloyd H. Lloyd G. A. Miller G. A. Miller N. Criddle G. A. Miller N. Criddle G. A. Miller* N. Criddle E. M. Kindle* es} = a 5 M Ib, L F F . M. Sternberg* wiePatch= . E. Wilson . M. Sternberg . E. Wilson . M. Sternberg . E. Wilson erie fe) ea19 ke (Viel =" eafevol esli@l Qt = fe) = P. A. Taverner H. Groh P. A. Taverner P. A. Taverner R. DeLury Treasurer Secretary(ies)? A. Gibson T. E. Clarke A. Gibson T. E. Clarke A. Gibson E. Clarke H. Groh J. J. Carter W. T. Macoun D. E. Blackader W. T. Macoun D. E. Blackader W. T. Macoun E. D. Eddy G. LaLacheur G. LaLacheur J. R. Dymond J. R. Dymond* F. W. Waugh E. B. Crampe* C. B. Hutchings C. B. Hutchings C. B. Hutchings B. A. Fauvel B. A. Fauvel B. A. Fauvel B. A. Fauvel B. A. Fauvel W. Lloyd W. Lloyd W. Lloyd W. Lloyd W. Lloyd W. Lloyd W. Lloyd W. Lloyd J. F Watson E. D. Eddy A. Halkett* G. O. McMillan L. D. Burling L. D. Burling C. L. Patch C. L. Patch C. L. Patch C. L. Patch C. L. Patch C. L. Patch J. FE. Wright J. E. Wright J. F Wright J. F. Wright J. F Wright B. A. Fauvel B. A. Fauvel G. S. Postethwaite G. S. Lewis G. S. Lewis G. S. Lewis P. Whitehurst P. Whitehurst 36 1937 1938 1939 1940 1941 1942 1943 1944 1945 1946 1947 1948 1949 1950 1951 1952 1953 1954 1955 1956 1957 1958 1959 1960 1961 1962 1963 1964 1965 1966 1967 THE CANADIAN FIELD-NATURALIST President H. Groh P. A. Taverner A. E. Porsild A. E. Porsild H. G. Crawford H. G. Crawford D. Leechman D. Leechman FE. Banim F. E. Banim W. H. Lanceley W. H. Lanceley P. Snure P. Snure J. W. Groves J. W. Groves R. Frith R. Frith W. K. W. Baldwin W. K. W. Baldwin L. S. Russell L. S. Russell E. L. Bousfield E. L. Bousfield W. W. Mair* D. R. Beckett* D. R. Beckett D.R. Beckett G. McGee G. McGee G. McGee H. Mackenzie Vice-President(s)! P. A. Taverner R. DeLury R. DeLury* H. G. Crawford* A. E. Porsild H. G. Crawford D. Leechman H. G. Crawford D. Leechman D. Leechman D. Leechman F. E. Banim F. E. Banim W. H. Lanceley F. E. Banim W. H. Lanceley W. H. Lanceley A. L. Rand W. H. Lanceley A. L. Rand A. L. Rand P. Snure P. Snure J. W. Groves J. W. Groves R. Frith J. W. Groves R. Frith R. Frith W. K. W. Baldwin K. W. Baldwin Senn K. W. Baldwin E. Bourguignon Russell Bousfield Russell Bousfield B B IL, E Le E ousfield leakney Bousfield Bleakney W. W. Mair D. R. Beckett W. W. Mair D. R. Beckett D. R. Beckett* V. Solman* E. Bousfield W. W. Mair D. A. Smith W. W. Mair G. McGee W. W. Mair G. R. Hanes W. W. Mair F. R. Cook F. R. Cook H. Mackenzie J. C. Woolley T. Mosquin W. H. W. A. 9. a Sy Bas Hf MS ae ssh J E J Treasurer W. Lloyd W. Lloyd W. Lloyd W. Lloyd W. Lloyd C. H. D. Clarke I. Conners I. Conners I. Conners I. Conners C. Frankton C. Frankton C. Frankton C. Frankton R. J. Moore R. J. Moore R. J. Moore R. J. Moore R. J. Moore R. J. Moore R. J. Moore R. J. Moore J. M. Gillett J. M. Gillett J. M. Gillett A. Banning A. Banning A. Banning A. Banning R. D. Wainwright L. G. Howden Vol. 118 Secretary(ies)” P. Whitehurst* C. W. Lounsbury* C. W. Lounsbury C. W. Lounsbury C. W. Lounsbury C. W. Lounsbury J. W. Groves J. W. Groves J. W. Groves J. W. Groves O. H. Hewitt O. H. Hewitt H. J. Scoggan H. J. Scoggan J. Scoggan J. Scoggan J. Scoggan al . Scoggan J. Scoggan . Scoggan J. Scoggan J. Scoggan M. Banfield M. Banfield M. Banfield ee ee eee a M. Banfield A. M. Banfield D. A. Smith A. W. Rathwell A. W. Rathwell A. W. Rathwell A. W. Rathwell 2004 1968 1969 1970 1971 1972 1973 1974 1975 1976 1977 1978 1979 1980 1981 1982 1983 1984 1985 1986 1987 1988 1989 1990 199] 1992 1993 1994 1995 BRUNTON: THE OTTAWA FIELD-NATURALISTS’ CLUB President H. Mackenzie H. Mackenzie T. Mosquin T. Mosquin S. Thomson I. M. Brodo I. M. Brodo E. C. D. Todd E. C. D. Todd R. A. Foxall R. A. Foxall R. Taylor R. Taylor R. Taylor D. F. Brunton D. F. Brunton E. F. Pope E. F. Pope W. Gummer W. Gummer J. Harrison J. Harrison J. Harrison R. John* E. F. Pope ESE Pope E. EF. Pope E. F. Pope Vice-President(s)! T. Mosquin* J. Tener* G. McGee* T. Mosquin W. A. Holland W. A. Holland S. Thomson S. Thomson I. M. Brodo I. M. Brodo E. C. D. Todd E. C. D. Todd E. C. D. Todd R. A. Foxall R. A. Foxall R. Taylor R. Taylor C. Gilliatt H. L. Dickson H. L. Dickson* D. F. Brunton* C. G. Gruchy P. M. Catling C. G. Gruchy* ESE; Pope* P. M. Catling W. Gummer W. P. Arthurs W. Gummer W. P. Arthurs* B. A. Campbell J. Harrison D. F. Brunton J. Harrison D. F. Brunton* K. Strang* J. Harrison R. John K. Strang R. John D. Cuddy E. Fox* S. Blight* vacant M. Murphy M. Murphy D. Moore M. Murphy D. Moore Treasurer 37 Secretary(ies)? L. G. Howden F. M. Brigham F. M. Brigham F. M. Brigham P. Kevin C. Gruchy C. Gruchy P. J. Sims P. J. Sims B. Henson B. Henson B. Henson B. Henson B. Henson P. D. M. Ward P. D. M. Ward P. D. M. Ward P. D. M. Ward P. D. M. Ward F. Valentine F. Valentine* J. Gehr M. Scromeda G. Marston G. Marston G. Marston G. Marston G. Marston A. W. Rathwell A. W. Rathwell A. W. Rathwell A. W. Rathwell A. W. Rathwell A. J. Erskine (RS) A. H. Reddoch (CS) A. J. Erskine (RS) J. D. Lafontaine (CS) A. J. Erskine (RS) C. Gruchy (CS) P. J. Narraway (CS) A. J. Erskine (RS) D. Laubitz (RS) S. Armstrong (CS) D. Laubitz (RS) S. Armstrong (CS) D. Laubitz (RS) V. Hume (CS) D. F. Brunton (RS) E. F. Pope (CS) E. F. Pope (RS) W. Gummer (CS) E. F. Pope (RS) W. Gummer (CS) E. F. Pope (RS) W. Gummer (CS) G. Hamre (RS)* A. Martell (CS)* B. Martin (RS & CS)* A. Martell (CS)* B. A. Campbell (CS)* B. J. Martin (CS) E. Bottomley (RS) B. A. Campbell (CS) M. Coleman (RS) D. F. Brunton (RS)* R. John (RS)* B. A. Campbell (CS) D. Duchesne (RS) M. Aksim (CS) E. Fox (RS) E. Evans (CS) E. Evans (CS) D. Furlong (RS)* C. Firth (RS)* N. Stow (RS)* E. Fox (RS)* E. Evans (CS) C. Clark (RS) S. Gawn (RS) E. Evans (CS) S. Gawn (RS) E. Evans (CS) D. Smythe (RS) E. Evans (CS) 38 THE CANADIAN FIELD-NATURALIST Vol. 118 President Vice-President(s)! Treasurer Secretary(ies)” 1996 D. Moore M. Murphy G. Marston D. Smythe (RS) E. Evans (CS) 1997 D. Moore M. Murphy S. Shaw* D. Smythe (RS) T. Reeve* L. Cairnie (CS) 1998 D. Moore E. Zurbrigg S. Shaw G. McNulty (RS) D. Smythe vacant (CS)* 1999 D. Moore E. Zurbrigg ESE: Pope? G. McNulty (RS) D. Smythe vacant (CS)* 2000 E. Zurbrigg R. John E. F. Pope J. Martens 2001 E. Zurbrigg R. John E. F. Pope K. Allison 2002 E. Zurbrigg R. John E. F. Pope K. Allison 2003 G. McNulty M. R. Murphy E. F. Pope S. L. Bourque G. Marston 2004 M. R. Murphy G. Marston E. F. Pope S. L. Bourque * served only a portion of this term; RS — Recording Secretary CS — Corresponding Secretary ' First and Second Vice-President positions existed in most but not all terms. 2 Secretary and Treasurer positions were combined until 1882; > The OFNC “year” was rearranged in 1919 to coincide with the calendar year rather than the fiscal year, resulting in a 1918-1919 year ending in March 1919, a9 month “year” for the remainder of 1919, and normal calendar years thereafter. * Corresponding Secretary position left vacant for two years then eliminated; Recording Secretary position continues. Continuing Environmental Change — An Example from Nova Scotia EDMUND S. TELFER Scientist Emeritus, Canadian Wildlife Service, Environment Canada Current address: 3582-42 Street, Edmonton, Alberta, T6L 5A1 Canada Telfer, Edmund S. 2004. Continuing environmental change — an example from Nova Scotia. Canadian Field-Naturalist 118(1): 39-44. Information from personal experience, from community elders and published literature served as a basis for evaluating envi- ronmental changes in the District of North Queens and adjacent areas of Southwestern Nova Scotia over the past century. Major events included disappearance of the Caribou (Rangifer tarandus), the arrival of White-tailed Deer (Odocoileus virginianus), the severe reduction of Canada Yew (Taxus canadensis), disappearance of Lynx (Lynx canadensis), a major dieoff of Striped Skunks (Mephitis mephitis), decline of American Beech (Fagus grandifolia), the loss of mature birch (Betula spp.), the severe reduction of Moose (Alces alces), the arrival of the American Dog Tick (Dermacentor variabilis) and Coyotes (Canis latrans), and the restoration of Beaver (Castor canadensis). The proximate cause of many of those changes were plant and animal disease, while the ultimate causes were naturally occurring animal range expansion and human impacts. The warming of the climate over the past 150 years probably played a role. The nature and timing of the events could not have been predicted. Key Words: Nova Scotia, plant and animal diseases, change in biota, climate change, adaptive management. Here, I present and discuss an example of long-term environmental change. It is a narrative account of changes in the occurrence of some mammals and plants in my home area, the District of North Queens, and surrounding portions of the interior of southwestern Nova Scotia. I personally observed these changes or heard about them from my father and other community elders and further document them by reference to the available literature. I discuss the changes in the light of current resource management philosophy. Since my father was born in 1882 and was a teenager in the 1890s, the changes noted cover a century. The District of North Queens lies in the interior of the southwestern peninsula of Nova Scotia and is centred at approximately 65°09'W and 44°18'N. It is an area of scattered farms and homesteads where the population has subsisted for 200 years on a mix of agriculture and forestry. The district contains the western portion of a swarm of glacially-formed drumlin hills that occur in Lunenburg and Queens counties (Roland 1982). Drum- lins provide arable land but comprise a small propor- tion of the land area. Most of the district is rocky, for- ested terrain that, even where cultivatable and produc- tive, was difficult for early settlers to clear. The remain- der of the interior of western Nova Scotia is a complex of water bodies, bogs, fens, barrens and stony soils supporting Acadian forests of mixed conifers and decid- uous tree species (Rowe 1972). When my father was born the area contained considerably more crop agri- culture and improved pasture than today. For the County of Queens as a whole, approximately 30 000 acres (12 140 ha) were under agriculture in 1870 (Canada 1871). This area dropped to 7410 acres (2999 ha) by 1966 (Nova Scotia 1966). However, those were small proportions of the total area, 4.4% in 1870 and 1.1% in 1966. Because the terrain is very complex, and the forest composed of many small stands, with the whole divided into small private land parcels, timber harvest has consisted of cutting small patches or individual trees as a market or need for the wood occurred. However, in recent years there have been larger clearcuts and some conversion of mixed forest to conifer plantations on industrial ownerships. The decline in the area of mixed- wood has been counterbalanced to an unknown extent by reversion of agricultural land to mixedwood forest and by improved fire protection (Telfer 1971). Mammals In the 1880s Caribou Rangifer tarandus, were still common in unsettled parts of Nova Scotia (Rand 1933; Sheldon 1936). An old hunter (the late F. M. Forrest, personal communication) told me of a trip he made when he was 16 years old (that would have been in 1888) to find Caribou. Somewhere at the west side of the present day Kejimkujik National Park he found a herd on a large barren and had a good chance to ob- serve them and to shoot a fat yearling to replenish his provisions. Within 20 years Caribou appear to have been gone from North Queens. Another old woodsman (the late E. B. Smith, Sr., personal communication) told me that when he began to guide sportsmen in the early 1900s there were still broad trails beaten though the woods in all directions in the remote parts of the District that had been made and kept open by Caribou. Sheldon (1936) also commented on the Caribou trails. By my time the Caribou were long gone and their trails had faded away before the encroaching forest. Caribou suffered from logging and burning on their winter ranges and from poorly controlled hunting. However, the probable final cause of Caribou extirpation was 39 40 THE CANADIAN FIELD-NATURALIST meningeal worm, Parelaphostrongylus tenuis, intro- duced by White-tailed Deer, Odocoileus virginianus (Benson and Dodds 1977). The loss of a large mammal like Caribou from the regional fauna quite probably had at least minor im- pacts on other plants and animals living in the area. I have not been able to find any records of any obser- vations of such impacts. However, Caribou move fast and far and feed while moving so they spread their use of vegetation out in time and space (Miller 1982). The relatively small population of Caribou in Nova Scotia would have had slight impact on other species. Moose, Alces alces, were distributed throughout Nova Scotia and were the common game animal in my father’s youth. However, after 1926 they were observed to be dying (Sheldon 1936; Benson 1958). The sick Moose exhibited symptoms similar to those later described by Anderson (1965) as resulting from infestation by the meningeal worm (Parelaphostron- gylus tenuis). This organism was later found in Nova Scotia Moose by Smith et. al (1964). Parelaphostron- gylus tenuis is relatively common in White-tailed Deer. It does little harm to whitetails but may cause heavy mortality among other species of the deer family (Thomas and Dodds 1988). By the 1940s, Moose were dying in large numbers from the condition which had come to be called “Moose Sickness” (Benson and Dodds 1977). The animals showed evidence of paral- ysis, blindness and other neurological symptoms al- though they were often in good physical condition otherwise. Annual reports of the Nova Scotia Depart- ment of Lands and Forests commented on the dead Moose. The 1949 report was typical, “Nearly all Rangers report moose as being in a healthier condi- tion which is perhaps indicated by the decrease in the number of moose which were found dead. This number was 75, a decrease of 74 from last year” (Nova Scotia 1949: 65). Of course only a fraction of the dead Moose would have come to the attention of the authorities. Moose almost vanished from southwestern Nova Scotia by the 1960s but recovered slightly following later declines in deer numbers. Although P. tenuis was found in some Nova Scotia Moose in the 1960s and has been shown to cause a “moose sickness” type of illness, that does not prove that all the animals that perished in the dieoff of Nova Scotia Moose in the 1930s and 1940s died of meningeal worm infection. However, the similarity of the symptoms displayed by the sick Moose suggests that most died of the same cause. Moose have been found to suffer from neurological symptoms where no P. tenuis has been observed in autopsies (Tony Nette, personal communication). Those symptoms have been recently linked to deficiencies of cobalt and vitamin B,, (Frank et al. 2004). However, it seems unlikely that a nutritional deficiency would cause a catastrophic dieoff like that reported in Moose in Nova Scotia in the 1940s, and the coincidence of the dieoff with the Vol. 118 post-introduction boom in White-tailed Deer num- bers suggests that P. tenuis was the principal cause. My father was an adult before deer showed up in North Queens. They had been spreading across Nova Scotia with the help of some introductions, and by the late 1800s their numbers were increasing (Rand 1933). There were many reasons for the increase. Possibly the most decisive was the climatic warming that set in about 1850 (Hawboldt 1952; Lamb 1982). The first deer hunting season in Nova Scotia was in 1916 and by the 1930s deer were common (Sheldon 1936; Benson and Dodds 1977). Over 40 000 were reported killed in Nova Scotia in one year in the early 1950s — more than two for every square mile in the province (Benson and Dodds 1977). Deer remained numerous until the unusu- ally severe winter of 1955-1956 caused a major dieoff of deer in western Nova Scotia. Since then they have fluctuated with winter conditions. Beavers, Castor canadensis, were not mentioned by my father as a species that he encountered in North Queens as a young man in the 1890s and the early 1900s. The animals had been trapped almost to extirpa- tion in Nova Scotia during the preceding century. How- ever, in 1907 Beavers were given complete protection (Wood 1973) and the Department of Lands and For- ests staff began live-trapping and transplanting them from the few remaining populations to unoccupied parts of the province. One cold autumn dusk when I was about five years old my father took me on a short walk to a small lake where Beavers had been released. We saw a new dam blocking the outlet stream and a new house with fresh mud on it. The Beavers had made themselves at home. They soon became abundant in North Queens and remained so until after 1945. Limited trapping was allowed after that time, reducing Beaver numbers and the dense population of animals had reduced their food supply along streams and lakes. However, Beavers have continued to occur in substantial numbers in North Queens (Wood 1973). Hunting with hounds was a favourite activity of my father and his friends. They hunted Red Foxes (Vulpes vulpes) and what they referred to as “wildcats”. Most of the cats were Bobcats (Lynx rufus) but my father also described killing cats that were over six feet long from rear toes to front toes when held up by their hind legs. Those large, rangy animals were probably Lynx. More (1873) stated that there were “lucifees” (Loup cervier, Lynx) and “little lucifees” in Queens County. However, the Lynx had become rare or extir- pated by the 1930s (Rand 1933; Sheldon 1936) and is now absent from the region (McCord and Cardoza 1982). According to local tradition, there were no Striped Skunks, Mephitis mephitis, in North Queens when the area was first settled. The species is not mentioned by More (1873) who, as a land surveyor, was very famil- iar with the mammals of the region. It was said that 2004 sometime in the mid-1800s someone brought a dead, frozen animal to a local blacksmith shop. None of the farmers who happened to be there could identify it. However, a few days later someone passed who had worked in the United States and who identified it as a skunk. By my father’s time skunks were common. However, by the 1930s, skunks were gone, probably due to an epidemic of distemper (Dodds 1969) but the decline in agricultural land use may also have had a negative impact on this animal of farms and forest edge. They were found in Nova Scotia only along the New Brunswick border but by the 1960s they had once again begun to spread into central areas of the province (Dodds 1969). None were present in Kejim- kujik National Park in the early 1970s (Wood 1973). However, in 2001 I observed a skunk near Ten Mile Lake, in Queens County. My father would have had difficulty in believing that the howl of the Coyote (Canis latrans) could ever be heard in North Queens. Yet in the 1970s Coyotes ar- rived in Nova Scotia. In North Queens, the popula- tion of White-tailed Deer in Kejimkujik National Park was apparently reduced by Coyotes which were com- monly seen along Park roads and heard on still nights throughout the area (Patterson 1994). The impact of Coyotes not only on deer but on competing Red Foxes, on Snowshoe Hares (Lepus americanus), microtine rodents, grouse (Bonasa umbellus and Canachites can- adensis) and other ground-nesting birds must have been considerable. Plants Before the arrival of White-tailed Deer, the forest floor in mature spruce-hemlock stands was carpeted with Canada Yew, Taxus canadensis. This evergreen shrub is highly palatable to deer and by the 1940s it had been eliminated from the forest. In the 1960s one experienced biologist (the late Harrison F. Lewis, personal communication) told me that the only sur- viving yew bush that he knew of was growing beside the outhouse at a warden cabin where one branch had grown through the wall of the well-ventilated facility and had thus escaped browsing. One of the most important trees of North Queens is the White Pine (Pinus strobus) (Rowe 1972). White Pines were a major resource in North Queens histori- cally (More 1873). However, in my father’s time re- generating White Pine was increasingly attacked by the White Pine Weevil (Pissodes strobi). That insect kills the leaders on the top of the tree, creating crooked stems or, at worst, “cabbage trees” whose lower branch- es continually grow up to take the place of the weevil- killed central stem. White Pine Weevil is a native North American insect. However, the first official records of its presence in the Maritimes date from about 1920 (Martineau 1984). The insect quickly spread throughout the range of the White Pine including North Queens. The spread of the weevil was facilitated by the exten- TELFER: CONTINUING ENVIRONMENTAL CHANGE — AN EXAMPLE 4| sive areas of regenerating forest created by logging, fire and the invasion of old fields by White Pine (Belyea and Sullivan 1956). By my time the value of the White Pine resource in North Queens was greatly reduced and crooked, multi-stemmed pine occupied much space in the forest. White Pine remains an important tree in the district but the weevil attacks have reduced the value of many stems for lumber. The American Beech, Fagus grandifolia, was com- mon in North Queens when my father was a boy. Pure beech stands occupied the ridge tops and beech trees were also scattered throughout the hardwood and mixedwood stands. Beech nuts were always an impor- tant seasonal food source for wildlife. Black Bears (Ursus americanus), especially, sought them out for a high fat and protein food before hibernation. When whitetails arrived in Nova Scotia they quickly became the most important consumers of the dwindling supply of nuts. Loucks (1962) cited evidence that beech was formerly more common in southwestern Nova Scotia. He believed that wildfire had reduced the occurrence of the species. Early in the 1900s the Woolly Beech Scale (Cryptococcus spp.), an insect that spreads Beech Nectria Fungus (Nectria coecinea var. faginata), ar- rived in Nova Scotia, probably an accidental intro- duction from Europe (Boyce 1961). The fungus gains entry to the tree through holes made by the Woolly Beech Scale. Saunders (1970) stated that Woolly Beech Scale entered Nova Scoria from Europe about 1890 and that the fungus also entered North America through Nova Scotia in about 1929. Nectria disease debilitates beech by invading the bark and cambium, eventually killing the tree. Roland and Smith (1969: page 341) noted that “All the beech in N.S. is severe- ly affected by the Nectria beech canker.” Most beech in North Queens was unthrifty and dying by the mid- twentieth century and in all cases moribund stems were thoroughly infested with Nectria cankers. Beech have declined substantially (Telfer 1971). They have been replaced by species like Sugar Maple (Acer saccha- rum) and Red Maple (Acer rubrum), Hemlock (Tsuga canadensis) and Red Spruce (Picea rubens). White Birch (Betula papyrifera) and Yellow Birch (Betula alleghaniensis) were prominent trees in the North Queens forest when my father was a boy. In- deed the old farmhouse in which he was born and died was sheathed with bark of the white birch between the wallboards and the shingles. In the mid-1930s a die- back was observed in the birch (Hawboldt 1952. Mature stands all died. This phenomenon occurred all over northeastern North America. Temperatures were high in the 1930s following a steady rise from Little Ice Age conditions that existed before the 1850s (Hawboldt 1952). One hypothetical cause of birch mortality was that the warming climate altered the species composition of the mycorrhizal flora in the soil to the detriment of species that could enter into symbiotic relationships with birch roots 42 THE CANADIAN FIELD-NATURALIST (Hawboldt 1952). Birch would thus have suffered from nutrient deficiencies. Although birch regeneration sur- vived, live mature trees were very scarce by my time. One would have had to travel a long way to collect enough bark to sheath a house. In the 1960s I worked on forest inventories in a remote part of the District where the mature forest had formerly been dominated by a mixture of birches, Red Spruce and Hemlock (Fernow 1912), but by then the birches were rotten snags and Red Spruce, White Pine, Balsam Fir (Abies balsamifera) and Red Maple were sprouting in the gaps created by their death. Arachnids An invading organism that caused excitement in North Queens in the mid-twentieth century was the American Dog Tick (Dermacentor variabilis). Opinion is divided on whether the ticks were relict in a small area in Western Nova Scotia or if they were introduced. One woodsman told me of encountering ticks along a small section of the Tusket River, near the western end of Nova Scotia, during a canoe trip early in the 1900s (the late F. M. Forrest, personal communi- cation). The distribution pattern shown by Dodds et al. (1969) suggests dispersion from an epicentre of infection. The Tusket River flows through the area of earliest distribution. By the 1930s loggers working in the western part of North Queens began to complain of tick bites. Ticks advanced on a broad front across the district in the late 1940s and the 1950s, causing consternation among the people who were unused to them. Since the 1960s, American Dog Ticks have con- tinued to spread eastward in Nova Scotia (Andrew Hebda, personal communication). Weather Nova Scotia is on the track of the equatorial Atlantic hurricanes. North Queens has been occasionally lashed by at least the fringes of those storms. In 1939, the area was hit by a hurricane that blew down many trees but did limited damage in closed stands. However, in 1954, a powerful hurricane, code named “Edna”, struck North Queens (Hawboldt and Bulmer 1958). Mature and old forests suffered particularly. Hurricane Edna was fol- lowed by a campaign of salvage logging as landown- ers tried to recover as much of the blowdown as pos- sible. The forest openings were invaded by regeneration of Red Spruce, Red Maple, White Pine and particu- larly by Balsam Fir in place of the dominant Hemlock of the former stands. Prediction and Management Causes of some of the observed changes were nat- ural, like birch dieback and distemper, while others were due at least in part to human intervention. There were some introductions of White-tailed Deer to Nova Scotia that probably accelerated their spread. Logging, small farming and attendant wildfires no doubt in- Vol. 118 creased the amount of suitable habitat for deer and also the area occupied by White Pine regeneration, the habitat of the White Pine Weevil. Hunting probably hastened the extirpation of Caribou and Lynx and the near extirpation of beaver. Coyotes may have been able to invade eastern North America partly because European settlers extirpated Grey Wolves, Canis lupus, from regions to the west of Nova Scotia. Ticks and the insect vector and the fungus involved in the beech Nectria disease may have been accidentally introduced. Climatic warming over the past 150 years interacted in a complex way with other causes of ecological change. The most important factors causing change to the terrestrial biota in North Queens operated haphazardly. Most were outbreaks of plant and animal diseases and parasites. Unpredictable outbreaks of disease are sometimes not considered to be very important by re- newable resource managers simply because they are random and basically unforeseeable. Also, the educa- tion of resource managers usually focuses on other topics. However, Hurricane Edna represented another kind of random event. There was sufficient evidence of the occasional occurrence of such severe hurricanes that shrewd ecosystem managers would probably have allowed for their possible occurrence when preparing management plans. The observed ecological changes in North Queens did not damage basic ecological functions like produc- tion, herbivory and predation. However, the suite of species involved changed. White-tailed Deer to some extent replaced Moose and Caribou as consumers while Coyotes became important predators as Lynx had been previously. After decades of absence, Beavers were brought back in numbers by deliberate manage- ment. Energy flow through the ecosystem was unim- paired but flowed through somewhat different chan- nels. The observed changes exemplify the fluidity and unpredictability of nature (Botkin 1990). What might have happened if, beginning in the 1890s, the forests and wildlife of North Queens had been under a modern regime of ecosystem manage- ment? Foresters might have made management plans aiming at production of a steady supply of birch logs. All would have been well for the first fifty years, then they would have found that the birch growing stock was dying and that spruce and fir were taking its place. As time went on the beech would also have been lost as a commercial species. Forest management would have had to be altered, as it indeed was following Hur- ricane Edna when managers were forced to revise their thinking about harvesting techniques (Johnson 1986) and deal with storm-caused openings and changes in forest composition. Conversely, the patchy nature of the storm-damaged forests provided improved habitat for White-tailed Deer and Moose. Wildlife managers would have found a major ungu- late species, the Caribou, suddenly declining. Even if the meningeal worm problem had been known, would <2 2004 managers have tried solve it by exterminating the White-tailed Deer population? Whitetails were by then spread thinly over the entire province. By the time Moose also declined there were so many deer that erad- ication would have been impossible. Even without par- asites, how would biodiversity managers have viewed the invasion of exotic whitetails especially since they were also the culprits in the extirpation of the Canada Yew? Managers would have been largely helpless to protect the yew except possibly by fencing small res- erved patches. Although “ecosystem” preservation is often a suitable management approach to maintain threatened species, it would have been counterproduc- tive in this case because old hemlock/spruce forest is preferred winter habitat for whitetails as well as the site of best yew growth (Schierbeck 1931). Lynx still remain on Cape Breton Island so protection from hunt- ing might have been adequate to recover the species on the mainland of Nova Scotia as it did for the Beaver. Because skunks appear to have died from dis- temper, reintroduction of healthy animals following the dieoff might also have re-established viable popu- lations and prevented local extirpation. Our hypothetical ecosystem managers would have emerged from the hundred-year period somewhat bat- tered. They would have lost a member of the charis- matic megafauna, the Caribou. They would no doubt have been chagrined by the fact that the district had been overrun by the exotic White-tailed Deer and Coy- otes, not to mention the obnoxious dog ticks. Some major tree species would also have been drastically reduced. However, a major understorey shrub and two mammal species might have been rescued and one mammal species, the Beaver, was indeed returned to the area by management. Would the public consider that money spent trying to maintain the ecosystems as they were in the 1890s to have been well spent as part of a worthwhile effort to “preserve” “nature” in the region? Or would managers simply be seen as having cast themselves in front of the juggernaut of inevitable ecological change? In the 1890s the coming changes and their impact could not have been foreseen. If managers at that time had set goals like doubling the sustainable harvest of White Birch and developing an industry to use it, or if they had set out to double the Caribou population to satisfy outfitters, they would have been in for a rough time. Yet, such factory-type production goals are often set in resource management. This example highlights the significance of scale in biodiversity conservation. Would it have been worth- while to try to keep species that are still common else- where in North America, like Caribou, Lynx, Striped Skunks, and Canada Yew, in the district or even in Nova Scotia? At what price should we try to keep everything that was present historically in a local or regional biota? Or should the waxing and waning of species’ ranges be accepted as inevitable? TELFER: CONTINUING ENVIRONMENTAL CHANGE — AN EXAMPLE 43 Although such a comprehensive management ap- proach would have been unthinkable in the 1890s, a program of adaptive management (Walters 1986) would have been appropriate. With the adaptive model in mind, managers might have been aware of the likeli- hood of natural variability and therefore cautious in goal setting. They could have initiated a program of monitoring that might have picked up changes in spe- cies status in their early stages when some prediction of the magnitude of upcoming impacts could have been made. While little could have been done to change the course of events in the short term, a more comprehen- sive and quantitative understanding of the operation of nature might have led to effective interventions later on. The haphazard nature of the changes in the ecology of North Queens during the past hundred years under- lines the difficulty of predicting what will happen next in nature. One is reminded of the soldier’s axiom re- garding shellfire, “You won’t hear the one that gets you”. The example of North Queens demonstrates the inevitability of environmental change and emphasizes the value of a flexible approach to forest and wildlife management. Literature Cited Anderson, R. C. 1965. Cerebrospinal nematodiasis (Pneu- mostrongylus tenuis) in North American cervids. Trans- actions of the North American Wildlife and Natural Res- ources Conference 30: 156-167. Belyea, R. M., and C. R. Sullivan. 1956. The white pine wee- vil: a review of current knowledge. Forestry Chronicle 32: 58-67. Benson, D. A. 1958. Moose “sickness” in Nova Scotia — I. Canadian Journal of Comparative Medicine 22: 244-248. Benson, D. A., and G. D. Dodds. 1977. The deer of Nova Scotia. Department of Lands and Forests, Province of Nova Scotia. 92 pages. Botkin, D. B. 1990. Discordant harmonies — a new ecology for the twenty-first century. Oxford University Press, New York. 241 pages. Boyce, J. S. 1961. Forest pathology, 3" edition. McGraw- Hill Book Company, New York. 572 pages. Canada. 1871. Census of Canada 1870-71. Ministry of Agri- culture, Ottawa (5 volumes). Dodds, D. G. 1969. The striped skunk, Mephitis mephitis (Schrieber), in Nova Scotia. Canadian Field-Naturalist 83: 229-233. Dodds, D. G., A. M. Martell, and R. E. Yescott. 1969. Ecol- ogy of the American dog tick, Dermacentor variabilis (Sag) in Nova Scotia. Canadian Journal of Zoology 47: 171-181. Fernovw, B. E. 1912. Forest conditions in Nova Scotia. Com- mission of Conservation, Ottawa. 93 pages plus maps. Frank, A., J. McPartlin, and R. Danielsson. 2004. Nova Scotia moose mystery — a moose sickness related to cobalt and vitamin B,, deficiency. The Science of the Total Envi- ronment 318: 89-100. Hawboldt, L. S. 1952. Climate and birch “dieback”. Depart- ment of Lands and Forests, Province of Nova Scotia. Bulletin 6. 37 pages. 44 THE CANADIAN FIELD-NATURALIST Hawboldt, L. S., and R. M. Bulmer. 1958. The forest resources of Nova Scotia. Nova Scotia Department of Lands and Forests. 171 pages. Johnson, R. S. 1986. Forests of Nova Scotia — a history. Nova Scotia Department of Lands and Forests and Four East Publications. Halifax. 407 pages. Lamb, H. H. 1982. Climate, history and the modern world. Methuen, London. 387 pages. Loucks, O. L. 1962. A forest classification for the Maritime Provinces. Forest Research Branch. Reprinted from the Proceedings of the Nova Scotian Institute of Science 25: Part 2, 1959-60. Martineau, R. 1984. Insects harmful to forest trees. Multi- science Publications Limited and the Canadian Forestry Service. 261 pages. McCord, C. M., and J. E. Cardoza. 1982. Bobcat and lynx. Pages 728-766 in Wild mammals of North America. Edited by J. A. Chapman and G. A. Feldhamer. Johns Hopkins University Press.1147 pages. Miller, F. L. 1982. Caribou. Pages 923-959 in J. A. Chapman, and G. A. Feldhamer, Wild mammals of North America. Edited by Johns Hopkins University Press.1147 pages. More, J. F. 1873. The history of Queens County, N.S. Nova Scotia Printing Company, Halifax. [Facsimile edition by Mika Studio, Belleville, Ontario, 1972]. 250 pages. Nova Scotia. 1966. Nova Scotia Forest Inventory — Lunen- burg Subdivision, Counties Lunenburg, Queens. Nova Scotia Department of Lands and Forests, Forestry Divi- sion Inventory Section. 36 pages plus tables. Nova Scotia. 1949. Report of the Department of Lands and Forests, 1949. King’s Printer, Halifax Nova Scotia. 96 pages. Patterson, B. R. 1994. Surplus killing of White-tailed Deer, Odocoileus virginianus, by Coyotes, Canis latrans, in Nova Scotia. Canadian Field-Naturalist 108: 484-487. Vol. 118 Rand, A. L. 1933. Notes on the mammals of the interior of western Nova Scotia. Canadian Field-Naturalist 47: 41- 60. Roland, A. E. 1982. Geological background and physio- graphy of Nova Scotia. Nova Scotia Institute of Science, Halifax. 311 pages. Roland, A. E., and E. C. Smith. 1969. The flora of Nova Scotia. The Nova Scotia Museum. 743 pages. Rowe, J. S. 1972. Forest regions of Canada. Department of the Environment, Canadian Forest Service Publication 1300. 172 pages plus map. Saunders, G. L. 1970. Trees of Nova Scotia. Bulletin 37, Nova Scotia Department of Lands and Forests. 102 pages. Schierbeck, O. 1931. Forestry vs. game cover. Canadian Field-Naturalist 45: 28-30. Sheldon, C. 1936. The mammals of Lake Kedgemakooge and vicinity, Nova Scotia. Journal of Mammalogy 17: 207-215. Smith, H. J., R. McG. Archibald, and A. H. Corner. 1964. Elaphostrongylosis in Maritime moose and deer. Canadian Veterinary Journal 5: 287-296. Telfer, E. S. 1971. Changes in carrying capacity of deer range in western Nova Scotia. Canadian Field-Naturalist 85: 231- 234. Thomas, J. E., and D. G. Dodds. 1988. Brainworm, Pare- laphostrongylus tenuis, in moose, Alces alces, and white- tailed deer, Odocoileus virginianus, of Nova Scotia. Cana- dian Field-Naturalist 102: 639-642. Walters, C. 1986. Adaptive management of renewable res- ources. Macmillan Publishing Company, New York. Wood, T. J. 1973. The recent mammals of Kejimkujik Nation- al Park. Proceedings of the Nova Scotia Institute of Sci- ence 27: 43-58 (1971-1973). Received 15 February 2002 Accepted 24 June 2004 Habitat Segregation Among Songbirds in Old-Growth Boreal Mixedwood Forest ENID E. CUMMING 1542 Empress Ave., Saskatoon, Saskatchewan S7K 3G3 Canada Cumming, Enid E. 2004. Habitat segregation among songbirds in old-growth boreal mixedwood forest. Canadian Field- Naturalist 118(1): 45-55. The foraging behaviour of ten species of insectivorous songbirds — Boreal Chickadee (Poecille hudsonicus), Golden-crowned Kinglet (Regulus satrapa), Ruby-crowned Kinglet (R. calendula), Blue-headed Vireo (Vireo solitarius), Tennessee (Vermivora peregrina), Blackburnian (Dendroica fusca), Magnolia (D. magnolia), Yellow-rumped [Myrtle] (D. coronata), Black-throated Green (D. virens), and Bay-breasted (D. castanea) warblers — was observed in the boreal mixedwood forest of Prince Albert National Park in central Saskatchewan. Birds segregated their habitat use by preferentially foraging in different tree species, and through preferential use of different foraging locations (height and position) within trees. White Spruce (Picea glauca) was used more than expected by Yellow-rumped Warblers, and Golden-crowned and Ruby-crowned kinglets. Tennessee and Magnolia warblers used White Birch (Betula papyrifera), more than expected and Boreal Chickadees and Blue-headed Vireos used Balsam Fir (Abies balsamea) more than expected. Boreal Chickadees, Ruby-crowned Kinglets, and Tennessee, Blackburnian and Yellow-rumped warblers all used the bottom part of trees less than expected, while Blue- headed Vireos foraged near the top of trees less than expected. Large inner branches were avoided by Tennessee, Blackburnian and Yellow-rumped warblers, while Bay-breasted Warblers and Blue-headed Vireos avoided small outer twigs. In conifers, Blackburnian Warblers foraged significantly higher in the trees than all other species except Black-throated Green and Bay- breasted warblers. Blackburnian Warblers also foraged significantly higher than Blue-headed Vireos and Magnolia Warblers in deciduous trees. Key Words: Songbirds, behaviour, foraging, boreal forest, old-growth, Prince Albert National Park, Saskatchewan. Behavioural interactions and niche partitioning in songbirds have received much attention in the literature (MacArthur 1958; Sabo 1980; Maurer and Whitmore 1981). For many boreal songbirds, however, much on their basic biology remains poorly understood (Sodhi and Paszkowski 1995). Many of the detailed behavioral descriptions have come from studies conducted in the New England States (Rabenold 1978; Robinson and Holmes 1982; Hunt and Flaspohler 1998). This is like- ly an incomplete description for some birds, as several of the species discussed have their breeding ranges almost entirely restricted to the Canadian boreal forest (Ficken et al. 1996; Rimmer and McFarland 1998). Studies on the southern fringes of the boreal forest, in Maine or Minnesota, are not adequate on their own to understand boreal ecology as many of the plant and ani- mal species are not the same as further north (Rowe 1972; Acton et al. 1998). Even for wide-ranging species, there is evidence that birds do not behave the same way and do not use the same kind of habitat in all parts of their range (Noon et al. 1980; Collins 1983; Robi- chaud and Villard 1999). Recent studies in the boreal forest have focused on habitat associations, and habitat requirements of song- birds (Schieck et al. 1995; Kirk et al. 1996). This infor- mation is urgently needed in light of increased pressure from large-scale forestry in this region (Cummings et al. 1994). There is mounting evidence to show that mature and old boreal mixedwood forest support an entirely different suite of species not found in younger age classes (Stelfox 1995; Cumming and Diamond 2002). These older age classes of forest are currently under a harvesting intensity that is disproportionate to their abundance in the landscape (Cummings et al. 1994; Weyerhaeuser 1998*). The goal of this project was to investigate details of within-stand habitat segregation among boreal song- birds, focusing on the foliage-gleaning songbird guild of old (>120 years) boreal mixedwood forest. This type and age of forest was chosen as it was biologically the most diverse and structurally the most heterogeneous habitat in the region (Erskine 1977; Johnson et al. 1995: Robichaud and Villard 1999). In addition, mature to old (80+ years), mixedwood forest will become increas- ingly rare at the landscape level if current land use practices (i.e., logging, oil and gas exploration and agri- culture) continue at their present rate (Cummings et al. 1994: Hobson et al. 2002; Schneider et al. 2002*). Study Area This study was carried out in the boreal, mixedwood forest of Saskatchewan (Kabzems et al. 1986; Acton et al. 1998), in Prince Albert National Park (53°35'N, 106°00'W) during 1990 and 1991 (see Bayne and Hobson 1997 for description). Three sites were chosen in forest stands that were >40 ha in area and >120 years old. Sites were as similar in topography and vegetative cover as field conditions allowed. Stands were aged by forest inventory maps, and by taking core samples from six of the largest trees on each site. Each study 45 46 THE CANADIAN FIELD-NATURALIST site was surrounded by continuous forest of similar type, but younger age. None of the study sites had been subjected to forest harvesting, and wildfire was the driving force behind forest succession in this area (Weir et al. 2000). The study sites were structurally very heterogeneous, with many tree-fall gaps and many standing dead trees. These canopy gaps allowed for regeneration, which resulted in a wide range of tree sizes and ages. Domi- nant tree species were White Spruce, Picea glauca, Balsam Fir, Abies balsamea, and Trembling Aspen, Populus tremuloides, with scattered White Birch, Betula papyrifera. Average canopy height was 21 metres with a maximum height of 38 metres (Cumming 1995). White Spruce was the only species that reached heights of over 30 metres, with some specimens also having diameters in excess of one metre. Understory trees (<12 metres), were mainly Balsam Fir, while the main shrubby species were Balsam Fir, White Birch sap- lings, and Alder, A/nus sp. Shrub distribution was patchy, with dense patches occurring in tree-fall gaps, and scattered shrubs in the rest of the study area. The most abundant ground cover species were various mosses, Bunchberry (Cornus canadensis), Wild Sarsa- sparilla (Aralia nudicaulis), Ostrich Fern (Matteuccia struthiopteris) and several horse-tail species (Equi- setum Spp.). Methods Observations on bird behaviour took place from late May to early August in 1990 and 1991, and birds were observed between 04:00 (dawn) and 21:00 (dusk). Upon locating a focal bird, continuous obser- vations were made using binoculars until the bird was lost from sight (Altman 1974; Martin and Batson 1986). To avoid possible biases, observations that last- ed less than five seconds or longer than two minutes were not used in the analysis. Bird species, behaviour, tree species and bird position in the tree were record- ed. Bird height was estimated to within three metres. Linear regression analysis (Zar 1996) was used to ensure that estimates accurately reflected clinometer- measured heights (Cumming 1995). In addition to behavioural observation, unlimited distance point counts (Bibby et al. 1982) were con- ducted twice during the breeding season (June), in order to estimate relative abundances of the various bird species. The average number of individuals per point count for each bird species was obtained by taking the maximum number recorded for that species and dividing by the number of point counts. Data from the three sites were pooled and used as a relative index of abundance for the study as a whole. For more details on bird species abundances see Cumming and Diamond (2002). Following the method of MacArthur (1958), con- ifers were divided into 15 zones; 5 vertical and 3 hori- zontal, to determine in which part of the tree birds were foraging (Figure 1). Vertical zones divided the tree into Vol. 118 fifths, with zone | at the top and zone 5 at the bottom. Horizontal zones corresponded to the outer part of the branch with new growth, fresh needles and small twigs (zone T), the middle layer of older needles and larger branches (zone M), and the inner part of the branch, largely devoid of needles with large diameter branches (zone B). A similar system was used for deciduous trees except they were divided into three vertical zones due to their more globular shape (Figure 2). Although many birds exhibit male-female foraging differences (Holmes 1986), data for both sexes were pooled since the focus of the study was on interspecific differences in habitat use and because some species in the study were not sexually dimorphic. Bird foraging behaviour (tree zone use), did not differ significantly between the two species of conifer or between the two species of deciduous trees. Therefore, data for each bird species was pooled based on either coniferous or deciduous trees (Cumming 1995). Vegetation in the study areas was measured using a modified James and Shugart (1970) method. Ten 0.04 ha circles (22 m diameter) were established in each study area, one centered on each of five point count stations and one at 50 metres in a random direc- tion from that point. Within each circle, species, height in meters and diameter at breast height (dbh) in cm for all trees larger than 7 cm dbh were recorded. Percent basal area for each tree species was calculated using dbh values. Basal area more accurately reflected tree availability to birds than numbers of trees, due to heterogeneity in tree size. To check for possible visibility biases in the various tree species, simultaneous confidence intervals (Chi- squared with Bonferroni confidence intervals) were constructed by pooling the number of seconds of ob- servation for all bird species (Byers et al. 1984). Simultaneous confidence intervals were also used to assess whether birds preferred or avoided certain tree species. Chi-squared test with Yates correction (Zar 1996) were used post priori to determine if birds had a preferred foraging zone within trees. Foraging diver- sity was analyzed using the Shannon Index (Shannon and Weaver 1949). To assess niche overlap between species, the proportion of observations each bird species spent in each of the 15 possible tree zones (9 zones in deciduous trees) was compared between each species pair using Morisita’s Index of overlap (see Diamond 1983). With this index, potential niche over- lap can vary from | (complete overlap) to 0 (no over- lap). Foraging height data were analyzed using a 1- way Anova (Proc GLM; SAS Institute 1988), and the results subjected to a Tukey’s HSD pair-wise com- parison test (with P=0.05). Results Tree Species Preference By basal area, White Spruce, Balsam Fir, Trembling Aspen and White Birch comprised 46%, 30%, 20%, and 4%, respectively, of the available arboreal habitat. 2004 CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 47 BOCH (N=128) H=2.25 @ 10- 14.9% [ 5.9.9% LI 0- 4.9% FiGuRE 1. Songbird use of zones in coniferous trees. Shadings indicate frequency of observations within a zone (see legend). N= number of observations, H= Shannon diversity index. See Table 4 for species abbreviations. 48 THE CANADIAN FIELD-NATURALIST White Spruce was used more than expected by Yellow-rumped (Myrtle) Warblers, (Dendroica coro- nata), Golden-crowned (Regulus satrapa) and Ruby- crowned (R. calendula) kinglets and in proportion to its availability by all other species (Table 1). Balsam Fir was used more than expected by Boreal Chicka- dees (Poecile hudsonicus) and Blue-headed Vireos (Vireo solitarius) but less than expected by Tennessee Warblers (Vermivora peregrina). White Birch was used more than expected by Tennessee and Magnolia (Dendroica magnolia) warblers, while Trembling Aspen was used less than expected by Boreal Chickadees, Golden-crowned and Ruby-crowned kinglets, Blue- headed Vireos, Yellow-rumped and Magnolia war- blers (Table 1). Bay-breasted Warblers (Dendroica castanea) used all tree species in proportion to their availability; however, they did exhibit a tendency to prefer White Spruce and avoid White Birch that was approaching significance. Blackburnian (D. fusca) and Black-throated Green (D. virens) warblers were not tested due to insufficient sample size (Brennan and Morrison 1990). Point counts All ten species were found in all three study sites, except Black-throated Green Warbler, which only oc- curred in one of the three study sites. Average bird abundance per point count from most to least abundant was; Magnolia Warbler (1.2), Yellow-rumped Warbler (0.8), Bay-breasted and Tennessee warblers (0.7), Blackburnian Warbler and Ruby-crowned Kinglet (0.4), Blue-headed Vireo (0.3), Boreal Chickadee and Gol- den-crowned Kinglet (0.2) and Black-throated Green Warbler (0.1). Foraging Zones In coniferous trees the most diversely foraging spe- cies (in order) were Yellow-rumped Warblers, Ruby- crowned Kinglets, Magnolia Warblers, and Boreal Chickadees (Figure 1). These species were observed in most tree zones and had the highest diversity indices (Shannon and Weaver 1949). The least diversely forag- ing species were Black-throated Green Warblers, Black- burnian Warblers and Blue-headed Vireos (Figure 1). Vol. 118 In the vertical zones, Ruby-crowned Kinglets, and Tennessee and Yellow-rumped warblers used the bottom of the tree, less than expected, while Boreal Chickadees and Golden-crowned Kinglets used the top and bottom less than expected (Table 2). Black- burnian and Bay-breasted warblers used the bottom two zones less than expected, while Blue-headed Vireos used the top two zones less than expected (Table 2). For the horizontal zones Bay-breasted Warblers and Blue-headed Vireos both used small twigs on the ends of branches (zone T) less than expected, while Tennessee and Blackburnian warblers used small twigs (zone T) more than expected (Table 2). Yellow- rumped Warblers use large inner branches (zone B) less than expected, while Boreal Chickadees, Golden- crowned and Ruby-crowned kinglets and Magnolia Warblers showed no significant difference in their use of inner and outer branches. Black-throated Green Warblers were not tested due to insufficient sample size (Brennan and Morrison 1990). In deciduous trees, only six of the ten species had a large enough sample size to test foraging zone use (Figure 2). In deciduous trees, Blackburnian Warblers used the top of the trees more than expected. Yellow-rumped, Magnolia and Blackburnian warblers used zone B less than expected, and Tennessee Warblers used zone T more than ex- pected (Table 2). Niche Overlap In conifers, niche overlap varied from 0% between Blue-headed Vireos and Blackburnian Warblers to 95% between Yellow-rumped Warblers and both Magnolia Warblers and Boreal Chickadees The four most diversely foraging species, Boreal Chickadees, Ruby-crowned Kinglets, and Yellow-rumped and Mag- nolia warblers, all overlapped each other by >80%. In deciduous trees, the most foraging overlap was between Yellow-rumped and Blackburnian warblers and between Magnolia and Tennessee warblers; both species pairs overlapped by 88% (Table 3). The least amount of overlap in deciduous trees (40%), was be- tween Black-throated Green and Bay-breasted warblers. TABLE |. Bird use of different tree species compared to the tree’s availability (y? test with Bonferroni confidence intervals, DF=3 for all species). Use Bird Species Boreal Chickadee Golden-crowned Kinglet > Expected Balsam Fir White Spruce Ruby-crowned Kinglet White Spruce Blue-headed Vireo Balsam Fir Tennessee Warbler White Birch Magnolia Warbler White Birch Yellow-rumped Warbler White Spruce Bay-breasted Warbler - Significance: *< 0.05, **<0.01, ***<0.001 < Expected ye Pp Trembling Aspen 43.3 ss Trembling Aspen 10.1 A Trembling Aspen 529 : Trembling Aspen 6.3 ‘i Balsam Fir 50.7 tte Trembling Aspen 10.9 Trembling Aspen DAD Ee = 5.0 n.s. 2004 CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 49 YRWA MAWA (N=54) (N=64) H=1.85 H=1.85 H=1.90 BLBW (N=53) H=1.91 H=1.35 H=1.25 FiGure 2. Songbird use of zones in deciduous trees. Shadings indicate frequency of observations within a zone (see legend). N= number of observations, H= Shannon diversity index. See Table 4 for species abbreviations. Foraging Height average foraging height of Blackburnian Warblers was In coniferous trees, the average foraging height of Blackburnian Warblers was significantly higher than all other species except Black-throated Green and Bay- breasted warblers (Table 4). Black-throated Green and Bay-breasted warblers both foraged significantly higher than Magnolia Warblers and Blue-headed Vir- eos. Foraging heights of all other species were not sig- nificantly different (Table 4). In deciduous trees, the significantly higher than that of Blue-headed Vireos and Magnolia Warblers but not significantly different from other species. Canopy vs. Subcanopy Use Blackburnian, Black-throated Green, and Tennessee warblers were seen in canopy level trees (>12 metres) in two-thirds or more of all observations (Table 5). At the other extreme, two-thirds of the observations on 50 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Bird use of tree zones where zone use was significantly more or less than expected (x? test with Yates’s correction; DF=!1 for all species). Species Horizontal Zones T,M,B Use OG In Coniferous Trees Boreal Chickadee n.s Golden-crowned Kinglet ns. Ruby-crowned Kinglet ns. Blue-headed Vireo at < 20.9 Tennessee Warbler al > 14.9 Blackburnian Warbler ay > 18.5 Magnolia Warbler ns. Yellow-rumped Warbler B < 14.6 Bay-breasted Warbler at < 10.5 In Deciduous Trees Tennessee Warbler T > 30.3 Blackburnian Warbler B < 24.9 Magnolia Warbler B < 155 Yellow-rumped Warbler B < 233) Vertical Zones IP 1 to5 Use Ye RB sD) < 20.4 nce We) < 14.5 cae 5 < 16.5 sats a Ie) < 1h oe Fata 5 < ig? “ook nas 4,5 < 7.4 mee 3 > 13.7 ie cates 3) < 10.1 ar ae 4,5 < 7.5 ne eae ns. se I > 6.6 * eae ns. bt n.s. Significance: *< 0.05, **<0.01, ***<0.001; T = outer part of branch with new growth, fresh needles and small twigs; M = middle layer of older needles and larger branches; B = inner part of branch, largely devoid of needles with large diameter branches; | to 5: vertical zones, 1 = top, 5 = bottom. TABLE 3. Average (mean + SD) foraging height (metres) in coniferous and deciduous trees. Species marked with * for- aged significantly higher than other species (1-way Anova with Tukey’s HSD test, P=0.05). Bird Species Coniferous Deciduous Boreal Chickadee 6.1 + 3.4 ONS: 2 Golden-crowned Kinglet TOE 229 ODES al Ruby-crowned Kinglet 7.3+44.7 9.5+4.7 Blue-headed Vireo 6.1+2.0 6.3+3.8 Tennessee Warbler SiOFE SE 5S) aed] Blackburnian Warbler esi Bel /a3) ay Thee Spoke Magnolia Warbler DR) Baa 6.9 +5.0 Black-throated Green Warbler 12.7 + 7.8* 11.44+3.8 Yellow-rumped Warbler 7.0+4.2 NIMs Site) Bay-breasted Warbler 10.3 + 4.8* loves Shs Coniferous: Blackburnian Warbler foraged significantly high- er than all but Black-throated Green Warbler and Bay-breasted Warbler. Black-throated Green Warbler and Bay-breasted War- bler foraged significantly higher than Blue-headed Vireo and Magnolia Warbler. Deciduous: Blackburnian Warbler foraged significantly high- er than Blue-headed Vireo and Magnolia Warbler. Magnolia Warblers and Boreal Chickadees were in subcanopy trees (Table 5). Blue-headed Vireos and Ruby-crowned Kinglets were approximately equal in their use of canopy and subcanopy trees, while Golden- crowned Kinglets and Yellow-rumped and Bay-breast- ed warblers tended to be observed slightly more often in canopy level trees (Table 5). Yellow-rumped Warblers were observed in canopy level trees significantly more than Magnolia Warblers (Z=5.2, P<0.001) and Boreal Chickadees (Z=3.4, P<0.001). Discussion Tree Species Preference The four main tree species in the study area (White Spruce, Balsam Fir, Trembling Aspen, and White Birch), were not equally abundant, nor equally large (Cumming 1995). If abundance alone had been used as the criteria for tree availability for foraging then Balsam Fir would have appeared to comprise a larger proportion of available habitat than it actually did. Patches of old, White Spruce-dominated forest are relatively rare in Saskatchewan (Kabzems et al. 1986; Acton et al. 1998); therefore, it was expected that birds which prefer White Spruce would selectively occupy this habitat. The data appear to support this, as all species in the study used White Spruce either in proportion to, or greater than, its abundance. Yellow- rumped Warblers and Golden-crowned and Ruby- crowned kinglets all used White Spruce more than expected. Boreal Chickadees and Blue-headed Vireos used White Spruce in proportion to its abundance and used Balsam Fir more than expected. All five species used Trembling Aspen less than expected (Table 1). This is similar to their behaviour elsewhere, as other researchers have found these five species to be conifer specialists (Keast and Saunders 1991; Ingold and Wallace 1994; Ingold and Galati 1997). Trembling Aspen was the least favored tree; it was used less than expected by half the bird species in the study. Previous research has suggested differences in insect abundance may exist between aspen and other tree species (Ives and Wong 1988). Holmes and Robin- son (1981) suggested structural characteristics of aspen trees and their leaves may impede foraging by insec- tivorous birds. However, Greenburg (1979) felt that Yellow-rumped and Bay-breasted warblers were phys- 5] CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 2004 690 O80 8L0 +9°0 cL0 180 Iv'0 19°0 IL°0 IL0 cs'0 ty 0 IL'0 09°0 cL'0 SLO depisag % ISLIOAY LS'0 970 OS'0 0c 0 0 Or'0 Lv'0 19°0 cs'0 IAHd £9°0 cL ‘0 Or'0 ce, $90 09°0 $80 c8'0 TYOD €8°0 870 vr'0 160 18°0 98°0 £60 HOOd S80 eL0 Or'0 OS'0 cL'0 $80 cL'0 bro OS'0 09°0 MNLd ‘JQ[SULY POUMOIN-UIP[OH =[PYOD ‘[sury poumo.s-Aqny =~PyYOr ‘sapeyoyD [eolog =Ho9oW OdIIA poproy-anlg_ =[AH “1o[qieM Ud9ID poyeoryi-yor[g =MNL “lorqieay ueluingyorlg = a714 Ja[qIVAA DOSSOUUaL =VWMALL “alqiey paysvoig-Aeg =VMaA_ “lorqieay vlpousep] =VMVIW “Jorgen, poduini-Moppox =VMUA 8L°0 870 ce 0 OS'0 Md 1d 09°0 880 98°0 OS '0 IL0 S80 VMAL MNLE Md Ta VMAL a VMdd cL0 = VMVI €8'0 cL'0 TR VMUA $90.1] snonplsap Uy IAHd IXOD HOO Oa MNILG Mad VMAL = VM c8'0 = VAVIN cL 0 £60 = VMUA $991] SNOIOJIUOD UT VMdd VAVW VMUA — satoeds pig ‘(XOPUT S,PIUSLIOJ]) $99.9 SNONPIOap puke sNOJa}JIUOS UT satoads UdaMJOq de]IDAO SUISRIOY Pp ATAVL Sy THE CANADIAN FIELD-NATURALIST TABLE 5. Bird use of canopy versus subcanopy trees, from most to least canopy using species. Number of Observations Bird Species Canopy Subcanopy Blackburnian Warbler 62 ii Black-throated Green Warbler iL) 5 Tennessee Warbler 80 41 Yellow-rumped Warbler 94 63 Bay-breasted Warbler 73 55 Golden-crowned Kinglet 16 12 Ruby-crowned Kinglet Dil, 25 Blue-headed Vireo 24 25 Boreal Chickadee 29 50 Magnolia Warbler 60 126 ically adapted to foraging in conifers and both their body size and foraging habits impeded foraging in deciduous trees. Tennessee and Magnolia warblers both used White Birch more than expected. Morse (1989) felt that many songbirds preferred various Betula species for mechanical reasons; they had smaller leaves than other trees (especially Fagus, Acer and Populus) and were easier for gleaning songbirds to reach. Other studies have found various species of birch have higher den- sities of lepidopteran larvae than other trees (Holmes and Robinson 1981; Holmes and Shultz 1986). An insufficient sample size prevented testing of Black- throated Green Warbler’s tree species preference. It is notable, however, that 50% of the foraging observa- tions on this species were in White Birch, even though these trees represented only 4% of the available habitat. Other researchers (Holmes and Robinson 1981; Robi- chaud and Villard 1999) have also found that Black- throated Green Warblers are partial to birch and use it greater than expected by chance alone. Tennessee Warblers used Balsam Fir less than ex- pected, while Blackburnian and Black-throated Green warblers were never observed in this tree. Although Blackburnian and Black-throated Green warbler use of this tree could not be tested, the fact that Balsam Fir comprised 30% of the available trees suggests that both birds were avoiding it. Blackburnian and Black-throated Green warblers were consistently ob- served high in the canopy, so Balsam Fir with an average height of 11 + 5.6 metres may have been too short to attract these species (Cumming 1995). Branch structure may also have played a role in some birds’ avoidance of fir. Differences in branch shape have been attributed to several bird species’ preference of Red Spruce (Picea rubens) over White Spruce (Morse 1989; Parrish 1995). Balsam Fir branches were less ridged than those of White Spruce (personal observa- tion). Birds which preferentially foraged on thin outer twigs appeared to have more difficulty foraging on the tips of Balsam Fir branches than they did in White Vol. 118 Spruce (personal observation). This was particularly noticeable in Tennessee Warblers because this species focused most of its foraging activities on the outer twigs and new growth in both coniferous and decid- uous trees. In contrast, birds which foraged in many tree zones (Yellow-rumped, Magnolia, and Bay-breast- ed warblers) appeared to be less affected by branch structure and they used Balsam Fir in proportion to its availability. Foraging Boreal Chickadees were the only permanent resi- dents in the study and one of the few passerines that are resident in the boreal forest (Ficken et al. 1996). It is considered an advantage for birds to be flexible in their foraging habits when they live in an unpredic- table environment, like the boreal forest (Hunt and Flaspohler 1998). Boreal Chickadees were diverse foragers, using most tree zones and overlapping more than 80% with Ruby-crowned Kinglets and Yellow- rumped, Magnolia, Bay-breasted and Tennessee warb- lers. Other studies found Boreal Chickadees had an average foraging height of five metres, and used all parts of the tree branch (Sabo and Holmes 1983; Ficken et al. 1996). Similarly, I found Boreal Chicka- dees foraged at an average height of six metres and showed no significant difference in their use of inner and outer branches. They did however, use the top and bottom of trees less than expected. Boreal Chick- adees may have avoided the tops of trees because this zone was heavily used by several other species. Boreal Chickadees have been shown elsewhere to shift their foraging activity lower in the trees due to competition from Black-capped Chickadees (Vassallo and Rice 1981). Boreal Chickadees also tended to forage in small subcanopy trees more than any other species except Magnolia Warblers. Ruby and Golden-Crowned kinglets were similar in their behaviour; however, Ruby-Crowned Kinglets were usually higher in the trees and were more diverse foragers. Rabenold (1978) found that where Ruby- crowned Kinglets and Golden-crowned Kinglets oc- curred together, Ruby-crowned Kinglets were more diverse in their foraging activity and occurred higher in the canopy than Golden-crowned Kinglets. Franz- reb (1984) felt this was due to social dominance of Ruby-crowned over Golden-crowned kinglets. No interspecific social interactions were observed in this study; however, Ruby-crowned Kinglets were twice as abundant as Golden-crowned Kinglets and may have influenced the behaviour of the latter through numerical dominance. Large inner and middle branches were favoured by both Blue-headed Vireos and Bay-breasted Warblers; this was an area of the trees that was seldom used by other species. There was only moderate foraging over- lap (47%) between Blue-headed Vireos and Bay- breasted Warblers, as they foraged at significantly dif- ferent heights. Foraging on large inner and middle 2004 branches is similar to the behaviour these two species have displayed elsewhere (Williams 1996; James 1998). Both Blackburnian and Black-throated Green war- blers foraged high in the trees (significantly higher than most other species), and had larger foraging overlap with each other (85%) than they did with any other species. These warblers appeared to be specialized in a tree-top niche, and have been identified in many stud- ies as feeding, singing, and nesting high in conifer- ous trees (Morse 1993, 1994). Indeed, Morse (1994) found that Blackburnian Warblers were unlikely to be found in forest without at least some conifers over 18 metres tall. Unlike MacArthur (1958) and Morse (1989), I found no evidence that Black-throated Green Warblers were socially dominant over any other spe- cies in the study area. This may have been because Black-throated Green Warblers were the least abundant bird in the study and only occurred in one of three study sites. Similar to their behaviour elsewhere, Tennessee Warblers were observed at a variety of heights in all tree species, but foraged mainly on the terminal foli- age (Rimmer and McFarland 1998). Foraging on the tips of the branches and using many different trees may help Tennessee Warblers (a Vermivora warbler) fit into a community (spruce-fir forest) that already has several coexisting species of Dendroica warblers (Morse 1989). Tennessee Warblers also used White Birch more than any other species except Magnolia and Black-throated Green warblers. Greenburg (1979) felt that small warblers were better able to forage on deciduous foliage than larger warblers for the physical reasons. Tennessee, Magnolia and Black-throated Green warblers were smaller than most species, except for the two kinglets, and Boreal Chickadees, which were conifer specialists. Therefore, they may have been using a food resource that the larger Yellow-rumped and Bay-breasted warblers would have trouble exploiting. I found Magnolia Warblers used subcanopy trees more than any other species and had the lowest aver- age foraging height of all species observed. This was similar to Hall (1994), who found Magnolia Warblers used mainly young conifers and were seldom found at any great height. Magnolia Warblers displayed no significant difference in their use of inner, middle and outer parts of branches. This was likely because they most often foraged young trees; there was far less difference in branch structure between inner and outer parts of branches in young trees than there was in large old trees (personal observations). Yellow-rumped Warbler showed the greatest diver- sity of foraging habits. They used almost every tree zone in both coniferous and deciduous trees. Yellow- rumped Warblers have been noted elsewhere as being generalists and this flexibility is thought to help them withstand adverse conditions better than their con- geners (Rabenold 1978; Hunt and Flaspohler 1998). Unlike their behaviour elsewhere (MacArthur 1958; CUMMING: HABITAT SEGREGATION AMONG SONGBIRDS 53 Morse 1989), Yellow-rumped Warblers did not use mainly the bottom and inner branches. Yellow-rumped Warblers occurred most often in the middle and outer branches at mid to upper heights in both conifers and deciduous trees, the same zone frequented by Black- throated Green Warblers in other studies (MacArthur 1958; Morse 1989). Morse (1971) and Howe (1979) felt that the presence or absence of Black-throated Green Warblers caused shifts in Yellow-rumped War- bler foraging activity. I observed no social interactions between Yellow-rumped and Black-throated Green warblers, and as Yellow-rumped Warblers were eight times more abundant, there was likely little interspe- cific competition between these two species. Indeed, Yellow-rumped Warblers experienced the most overlap, and likely the most interspecific competition, from Magnolia Warblers, Boreal Chickadees, and Ruby- crowned Kinglets (Table 4 and Figure 1). Niche Overlap Northern regions, such as the boreal forest, are thought to have such large pulses of seasonally avail- able food that for songbirds food saturation occurs (Rabenold 1983; Wiens 1989). This seasonal pulse of food, combined with a high number of neo-tropical migrants, allows greater niche overlap between song- birds than occurs in other habitats (Rabenold 1978, 1983; Wiens 1989). Rabenold (1978) found that spruce- fir forests in particular had a high number of songbirds which were “...broadly overlapping congeners”. Although the present study and Rabenold’s were not directly comparable, many of the species were the same between the two studies and results from this study support his theory of generalist stacking. Species with the most generalized foraging behaviour had the high- est amount of niche overlap, while those with more specialized foraging behaviour had little niche overlap. Canopy vs. Subcanopy Availabilty of canopy versus subcanopy trees was not measured; therefore, I could not test whether birds were significantly selecting one over the other. At the two extremes, however, it was apparent that Black- burnian Warblers were using very tall trees almost exclusively, while Magnolia Warblers were using most- ly young regenerating trees. I tested Yellow-rumped Warblers’ use of canopy and subcanopy trees against Magnolia Warblers’ and Boreal Chickadees’ because they had a 95% foraging overlap (in conifers) with both species (83% with Magnolia Warbler in deciduous trees). A Z-test (in results) found Yellow-rumped Warblers used tall (canopy) trees sig- nificantly more than the other two species. I felt this was a significant difference between these species, as they used the available habitat in a similar way and they did not forage at significantly different heights. Magnolia and Yellow-rumped warblers were also the two most abundant species in study. Previous research has shown old-growth boreal mixedwood forest such as those studied here to harbor 54 THE CANADIAN FIELD-NATURALIST unique communities not seen in pre-rotation age forest (Erskine 1977; Kirk et al. 1996: Cumming and Dia- mond 2002). This diverse songbird community appears to be caused by an interaction between a structurally diverse habitat and a mixture of generalized and specil- ized foraging behaviours of the bird species involved. Although this study did not examine multiple age class- es of forest, it would appear that the community dyna- mics seen here may be fairly unique due to canopy gap dynamics that tend to be associated with old-growth forest (Hunter 1990). This type and age-class of forest has not received the attention it deserves from either environmental impact studies or long-term manage- ment plans for the boreal forest. The scientific commu- nity needs to pay greater attention to the contribution old-growth boreal mixedwood forest makes to bio- diversity on both a regional and national scale. Acknowledgments This study was carried out in partial fulfillment of the requirements for a M.Sc. degree at the University of Saskatchewan. Parks Canada allowed field work to be carried out in Prince Albert National Park. Funding was provided by the Canadian Wildlife Service, Prince Albert National Park, Nature Saskatchewan, and the Saskatchewan Wildlife Habitat Development Fund. 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Gill. The Academy of Natural Sciences, Philadelphia, Pennsylvania, and The American Ornithologists’ Union, Washington, D. C. Zar, J. H. 1996. Biostatistical analysis. Third edition. Pren- tice-Hall. Englewood Cliffs, New Jersey. Received 10 March 2001 Accepted 20 May 2004 Effects of Mid-winter Snow Depth on Stand Selection by Wolverines, Gulo gulo luscus, in the Boreal Forest JONATHAN D. WRIGHT! and JEssICA ERNST Ernst Environmental Services, Box 753, Rosebud, Alberta TOJ 2T0 Canada Present address: Box 648, East Coulee, Alberta TOJ 1BO Canada Wright, Jonathan D., and Jessica Ernst. 2004. Effects of mid-winter snow depth on stand selection by Wolverines, Gulo gulo luscus, in the boreal forest. Canadian Field-Naturalist 118(1): 56-60. Wolverines (Gulo gulo luscus) in a study area in the boreal upland forests of northwestern Alberta and northeastern British Columbia (approximately 57°N) were noted to be limited to upland landscapes, despite abundant food in adjacent lowland landscapes. Snow-tracking suggested that the species was selecting for the densest climax conifer stands for travel in search of food. It was hypothesized that snow depth was a limiting factor for Wolverines in the boreal forest during midwinter, and that they selected for this stand-type because of the buffering effect of this type of canopy on ground snow-depths. A series of snow-depth measurements were collected. Snow depths collected along Wolverine trails were very significantly lower than random snow depths collected under upland canopy (F = 32.84, df = 1, P << 0.010). There was a significant buffering effect on snow depth indicated for upland canopy (F = 11.1, df = 1, P < 0.010), while adjacent lowland canopy had no significant buffering effect on snow depth (F = 3.45, df = 1, P > 0.05). Wolverines were hypothesized to be limited to upland landscapes in the study area because of the buffering effect on snow-depth of the stand types found there, and not for reasons of food availability. Climax conifer stands were interpreted as being of high importance to Wolverine survival during winter. Conser- vation implications include the detrimental effect on Wolverine populations likely to result from current timber harvesting practices in the boreal forest. Key Words: Wolverine, Gulo gulo luscus, snow depth, boreal forest, stand, landscape, climax, buffering effect, timber harvest- ing, Alberta, British Columbia. The Wolverine, Gulo gulo luscus, remains among the least understood of the world’s forest mesocarnivores. Recent research focusing on the life history of the species in cordilleran landscapes is beginning to fill knowledge gaps. However, landscape features docu- mented as being of high importance to Wolverines in mountainous regions — for example, the alpine zone (Eric Lofroth, personal communication [see Acknow- ledgments section for affiliation]) — are unavailable to forest populations. In the boreal forest, Wolverines are believed to be declining from densities that appear to be lower than in any of the other landscapes in which they occur (Banci 1994). As they are less observable in the forest due to the unbroken forest canopy, Wolverine study in the forest ecoprovinces is an even greater challenge than in alpine habitats. It is therefore not surprising that Banci (1994) has suggested that stand-level and land- scape scale “habitat use by Wolverines in forests has not been adequately investigated”, and that research is needed to study the “habitat needs of Wolverine in forests, because there is no sound basis for developing habitat management prescriptions at the stand level”. In forested areas of the northwest, the Wolverine is the furbearer about which wildlife agencies have the greatest concern (Bill Johnson, personal communica- tion). There has been agreement among researchers that Wolverine “habitat is probably best defined in terms of adequate year-round food supplies in large, sparsely inhabited wilderness areas, rather than in terms of particular types of topography or plant associations” (Kelsall 1981)*. Seasonal shifts by Wolverine in cor- dilleran landscapes from the alpine zone in the summer to subalpine forest in the winter have been attributed to the availability of food (Banci 1994), or are thought to be related to avoidance of high temperatures or humans (Hornocker and Hash 1981). While snow depth has been investigated as a limiting factor for Fishers, Martes pennanti (Krohn et al. 1995; Raine 1983), it has not been implicated as a factor affecting habitat selection in the Wolverine. This paper presents the hypothesis that Wolverines in the boreal forest are limited by mid-winter snow con- ditions, which in turn affects stand, and in this case, landscape selection. Methods Snow tracking is being increasingly recognized as a reputable scientific tool in wildlife studies, management and conservation, with efforts being made to establish standardized terminology, institute university courses on the subject, and to establish networks of trackers throughout the North American continent (Rezendes 1999; Zielinski and Kucera 1995). Snow tracking may be the only practical way of learning details of Wol- verine habits and habitat use, as such details are not adequately provided by radio-telemetry studies (John Krebs, personal communication; Eric Lofroth, per- sonal communication). 56 2004 Wolverine and other furbearer activities were mon- itored during three winter seasons (1997-2000) utiliz- ing snow-tracking methods. Approximately 34 000 km were traveled by truck, snowmobile, cross-country skis and on foot, in search of furbearer tracks, with an emphasis on locating Wolverines. The tracking efforts covered approximately 1100 km/? in the region of the border country of Alberta and British Columbia known as “Chinchaga” (after the Chinchaga River), at approx- imately 57° north latitude. The tracking area offered conditions especially conducive to this type of study, in that the landscape could be conveniently divided into distinct upland and lowland components. These com- ponents offered markedly different and readily divis- ible stand characteristics as a result of the “Great Chin- chaga Fire” which blanketed the area during the 1950s (Don Williams, personal communication), burning all but some upland ridges. Fifty years following the fire, two distinct forest types predominate: early-to-mid seral second-growth of predominantly aspen or pine in the lowlands, with more limited late-seral, climax or “over-mature” stands predominated by White and Black spruce (Picea spp.), and mostly dead and dying Populus spp. in the uplands. The lowland forest may be further characterized as forming a mosaic broken by broad expanses of relatively open Black spruce fen and willow (Salix sp.) muskeg. The upland forest cover is by comparison more continuous. The area is thoroughly criss-crossed by open linear corridors at varying stages of regeneration that were cleared to conduct seismology work, and have been implicated in possible predator/prey imbalances. Seis- mology work and subsequent oil and gas exploration and extraction activities have been traditional in the area since the 1950s (Brody 1981). The area is expe- riencing a surge of such activities at present, and there was virtually no segment of the area covered in track- ing Wolverines that was not impacted by these activ- ities at some point during the three winter seasons. Moose (Alces alces) reach some of their highest densities in North America in the area (Brody 1981), being most heavily distributed in the lowlands, and frequent in the uplands. Groups of Woodland Caribou (Rangifer tarandus caribou) are frequent at low den- sities in the lowlands. These are the only two com- mon ungulate species in the region, and they offer a generous prey-base for the area’s healthy Grey Wolf (Canis lupus) population, whose kills in turn offer a ready food source for the infrequent Wolverines. When Wolverine tracks were located, the individuals were fore-tracked (older trails) or back-tracked (fresh trails) in order to gain insights into important and little-known details of the creature’s use of the boreal forest, including landscape and stand use. Tracks were accessed in the morning by snowmobile or truck, and followed on foot as long as daylight permitted, with the exception of one overnight excursion conducted in order to assess advantages and disadvantages to the WRIGHT AND ERNST: STAND SELECTION BY WOLVERINES 57 tracking process of remaining in situ. Pertinent details of the Wolverine’s behavior were logged in field note- books during the tracking event as well as being photographically documented. Snow depth measurements (in centimetres) were taken over a time frame of approximately | week, char- acterized by below-freezing temperatures, between periods of snowfall. Snow depths were taken at loca- tions determined using random numbers generated by a portable computer (calculator). The number indicated the number of paces to be taken on the ground, either within a grid-square (random snow depths), or along a Wolverine or Lynx (Lynx canadensis) trail, at which point a snow-depth was taken. Bes following snow-depth data were recorded: Ten random depths on open upland sites (to quantify “non-buffered” upland snow depth) with a grid-cell size of 1 km?; e 27 random depths in forested upland sites (“buffered” upland depths) with a grid-cell size of 1 km; e Ten random depths on open lowland sites (to quantify non-buffered lowland snow depth) with a grid-cell size of 1 km’; e 25 random depths on forested lowland sites (buffered lowland depths) with a grid-cell size of 1 km?; e 46 random depths along documented Wolverine trails at random paced intervals; e 29 random depths along Lynx trails (for comparison purposes) at random paced intervals. The following comparisons were made utilizing ANOVA tests of significance: e Snow depths along Wolverine trails versus random for- ested upland snow depths; e Random forested upland snow depths versus random open upland snow depths (= “buffering effect” of upland canopy on snow depth); e Random forested lowland snow depths versus random open lowland snow depths (= buffering effect of lowland canopy on snow depth): e Snow depths along Lynx trails versus random forested lowland snow depths (for comparison purposes). Crusting of snow typically occurs in the area of this study during late winter to early spring, and occasion- ally in late fall, depending on temperatures. Crustless (powder) conditions are the norm in this region for the majority of the snow season. Snow conditions were noted as being of a crustless nature during the collec- tion of this data. Wolverine reactions to linear corridors such as access roads and seismic lines were documented as encountered, with special attention being paid to the effect that snow conditions on the corridors (compact- ed versus undisturbed) had on Wolverine response. Efforts were made to locate Wolf kills in both representative landscapes in order to monitor such important food sources for utilization by Wolverines. Results Wolverine tracks were located on ten occasions over three winters (for a mean of one Wolverine track en- 58 THE CANADIAN FIELD-NATURALIST countered per 3400 km of searching). Thirteen sepa- rate tracking events resulted, totaling 20 linear km of off-corridor (forest) tracking, or 26.6 total off-cor- ridor km using Magoun’s (1985) estimate of adding 33% to arrive at non-linear distance traveled. Wolver- ines were tracked for an additional 12.3 km on linear corridors offering conditions of compacted snow, for a total estimated tracking distance of 38.9 km. Seven Wolf kills (Moose) were located, two in the upland landscape and five in the lowland landscape. These kills were monitored for Wolverine use. An additional road-killed Moose in the lowland landscape was monitored for Wolverine use. Of these, Wolver- ine were documented as utilizing one of the upland Wolf kills. Despite frequent available food in the lowland land- scape, Wolverine tracks were located only in the upland landscape. Wolverines were revealed to have remained in the uplands throughout the tracking events. One Wolverine tracked was noted to have traveled through a burned-over upland area regenerating to Lodgepole Pine (Pinus contorta). The second-growth pine formed a low (approximately three-metre), dense canopy. Heavy use of the burn by Snowshoe Hares (Lepus americanus), provided compacted trails on which the Wolverine traveled through this second-growth. Wolverines often encountered linear corridors during their travels. Wolverine diverged from their line-of- travel under the forest canopy to travel on 100% of the linear corridors encountered that offered conditions of compacted snow (n=17), for distances ranging from 3 — 3000 m. Wolverines did not travel on any of the corridors encountered that had undisturbed snow cover (n = 16), choosing instead the most direct route across in 100% of instances noted. There were two instances where an individual Wol- verine paralleled an undisturbed east-west corridor from just within the south-facing edge of the canopy. The individual crossed the respective corridors in- volved to select the south-facing aspect for travels of 470 m and 30 m, before diverging deeper into forest. Exposure to the sun had acted in concert with the buffering effect of the canopy to reduce snow depth from 32 cm on the corridor, to 4 — 11 cm under the canopy’s edge. Snow depths along routes selected for travel by Wolverines were significantly less than random snow depths under the upland canopy (F = 32.84, df = 1, P << 0.010). TABLE 1: Comparison of random snow depths Wolverine Trails Range 13-47 cm Mean 31 cm Standard Deviation 9.03 Vol. 118 Random upland snow depths under the canopy were significantly less than snow depths in open upland areas (F = 11.1, df = 1, P< 0.010), whereas random lowland snow depths under canopy were not significantly dif- ferent from snow depths in open lowland sites (F = 3.45, df = 1, P > 0.05). Snow-depths along routes selected for travel by Lynx were not significantly different from random snow depths under the lowland canopy, where Lynx data were collected (F = 2.86, df = 1, P > 0.05). When Wolverine tracks were discovered exiting the forest onto access routes, the Wolverines involved had invariably been traveling through dense to extremely dense coniferous cover. While subsequent travel on compacted linear corridors would frequently take a Wolverine through a variety of stand types, the Wol- verines invariably selected similarly dense coniferous cover for re-entry into the forest. Within the forest, Wolverines were frequently, even continuously, ob- served to alter their direction of travel (n = 29) to select for routes that offered an even slightly denser canopy cover (increased buffering effect on snow). Of the ran- dom upland sites, 70% were estimated to represent a similar cover type to that selected by Wolverines for travel, as compared to 20% of random lowland sites. Discussion Popular and even scientific literature of past decades reflected the level of misconception surrounding the Wolverine. The Wolverine was seen as unique among wildlife in being best adapted to survive during the harshest months, to the point of finding winter a sea- son of ease (Rausch and Pearson 1972). The animal’s adaptations to a winter landscape were provided as evidence of this: the dense, luxurious coat (which may give the illusion of good underlying physical condition in winter); the enormous, well-haired “snowshoe” feet cited as enabling the animal to coast effortlessly on top of deep snow; the preference for the harshest wilderness areas. In fact, in an ongoing study in the foothills of northern British Columbia, Wolverines in winter were often found upon examination to be in a state of energetic stress approaching thresholds of star- vation (Don Reid, personal communication); a large percentage of dissected Wolverines have empty gastro- intestinal tracts (Banci 1994). Unlike such sympatric predators as Martens (Martes americana), and col- oured foxes (Vulpes vulpes), Wolverines are too large to survive on small prey (Banci 1994), and must there- fore wander widely, as indicated in all studies, in search Random Lowland Locations Lynx Trails 23 — 52 cm 17-54 cm 40 cm 36 cm Wes) 8.2 2004 of less reliable yet more bountiful sources of food such as large ungulate carrion. While recent research is presenting a different pic- ture of Wolverines and their habits, apparent contra- dictions still abound. One of the more notable of the apparent contradictions in Wolverine behavior is ad- dressed by this paper: Why do Wolverines select for wide-open landscapes in some parts of their range and the densest of cover in other areas such as the boreal forest? During the tracking of Wolverines in the boreal forest at Chinchaga, patterns of behavior became ap- parent. The species limited itself to the upland land- scape despite abundant food in the lowlands, con- tradicting literature which suggested that wilderness conditions and a ready supply of food governed the Wolverine’s presence or absence, rather than specific landscape and stand features (Kelsall 1981*; Banci 1994). The observation that Wolverines in the study area were limiting themselves to a specific landscape (at least during winter) was supported by conversa- tions with the area’s fur trappers and wildlife agents, who noted that: (1) over many decades of trapper experience in the Chin- chaga area, Wolverines were encountered in heavily timbered upland terrain (Les Sharp, personal com- munication); (2) in a winter trapper’s experience over 13 years on a lowland landscape trapline adjacent to uplands where Wolverines were tracked during this study, not a single Wolverine was caught, nor tracks ever detected (Larry Smith, personal communication). (3) in other lowland traplines in the area with long tra- ditions of use none were recalled to have ever yielded Wolverine pelts (Bill Johnson, personal communi- cation). During the tracking efforts, it became quickly appar- ent to me that the Wolverines were continuously sel- ecting, where feasible, for the path of least snow cover. When faced with even a slight thinning of the forest canopy (= deeper snow conditions), the animals were observed to have paused, selected the densest alter- native timber, and altered their route to follow this “path of least resistance” in terms of snow depth. The Wolverines would only cross such openings if: (1) a compacted Showshoe Hare trail crossed the deeper snow, which they would then select; (2) alternative routes were not available without extensive detour, in which case the Wolverine would cross the deeper snow over the shortest straight-line path, usually at a walking gait. Wolverines in the area seemed to favor a “2x” or “3x” lope (Zielinski and Kucera 1995) when less impeded by snow, suggesting that this was their most efficient gait in terms of energy expenditure versus distance traveled. They would immediately switch to a less efficient walk, however, when faced with unavoid- able stretches of deeper snow. In comparison, Lynx were observed not to avoid deeper snow, readily crossing open areas of the deepest snow conditions, as encountered. This is supported WRIGHT AND ERNST: STAND SELECTION BY WOLVERINES 59 not only by the snow depth data, which showed Lynx (in direct contrast with Wolverine) to have selected paths of insignificantly less snow depth, but also by the fact that Lynx were abundantly found in the lowland landscape where the buffering effect of the forest canopy was demonstrated to be insignificant. Wolverines traveling together did so single-file for an estimated 95% of the distance traveled, traveling in tandem only along ridges where snow cover was neg- ligible. It would be interesting to discover if social travel in this species represents a survival strategy, and if the individuals involved alternate as lead-individual. That Wolverines at Chinchaga were not selecting the densest canopy conditions in order to avoid detection by man or predators is obvious for the following reasons: (1) In 100% of cases, they followed open linear corridors offering compacted snow conditions when encountered (including winter roads up to 18 m wide; freshly opened seismic lines; snowmobile trails; all-terrain vehicle tire tracks; wind-swept ice on creek-beds bisect- ing open muskeg; compacted otter runways on open creek beds) for travel of distances up to 3 km; (2) Wolverines are more abundant on the open tundra (characterized by hard-crusted snow conditions) than they are in the forest (Don Reid, personal communi- cation; Banci 1994); (3) Wolverines seem to prefer the open alpine zones during snow-free seasons in cordilleran landscapes (Whitman et al. 1986; Banci 1994; Don Reid, personal com- munication; Eric Lofroth, personal communication). The hypothesis presented by this paper is that Wol- verines in the mid-winter boreal forest are limited by snow conditions. Conifer-dominated climax stand types offer the highest buffering effect on snow conditions underfoot. In this study area, such stand-types are only found extensively in upland landscapes, which accounts for the absence of Wolverines in the lowlands, despite abundant food there. Wolverines travel great distances in search of unpredictable food (Banci 1994) while under conditions of energetic stress, necessi- tating that they pay the strictest attention to energy expenditure while traveling. To travel in conditions of deep powder (unbuffered) snow is to increase risk of starvation, and so such conditions are avoided where alternatives exist. While the relatively enormous feet of the Wolverine, providing low foot loads of 22 g/cm? (Knorre 1959) are of great advantage when crusts begin to form (Eric Lofroth, personal communication), they have little effect in the midwinter powder snows of the northwest characterizing this study, as indicated by shifts to an inefficient walking gait in deep, midwin- ter snow. It is further suggested here that the size of the Wolverine’s feet, rather than being an indication of their level of mastery of winter conditions, is in- stead just the opposite, an adaptation indicative of their extreme level of sensitivity to snow depths as a result of specialized foraging strategies combined with rela- tively large body size leading to conditions of energetic stress. While one could argue that Lynx, with their equally large feet and specialized foraging strategies, 60 THE CANADIAN FIELD-NATURALIST should therefore show a similar sensitivity to snow depths not indicated in this study, it should be remem- bered that the Lynx data herein were collected during years of high hare numbers. Perhaps during the low hare cycle, Lynx must also become more sensitive to snow depths. Conclusions During midwinter, Wolverines in the boreal upland forests of northwestern Alberta and northeastern British Columbia demonstrate preferences at the stand level and landscape scale that are apparently unrelated to food availability, consistently selecting for stand types that offer the greatest available buffering effect on ground snow depth. Wolverines require the type of boreal forest habitat considered optimum for Martens, the species with which they may be considered most highly sympa- tric. However, Wolverines appear to be even more dependent on climax conifer forests during winter than are Martens. Climax conifer cover appears to be of high importance to Wolverines in the boreal forest because of the buffering effect this stand type has on snow depth. Forest practices that remove climax growth may create conditions that are not conducive to Wolverine conservation. As current logging practices in the boreal forest are geared towards precisely this type of stand removal, timber harvesting may be a key factor in appa- rent population declines. Conditions following removal of climax cover may not become favorable again for Wolverine habitation for many decades (Don Williams, personal communication). The situation may be im- proved in the case of immediate regeneration to Pinus spp., however, with a minimum estimated lapse of 20 years (several Wolverine generations) before conditions become again conducive to Wolverine travel, if not other life history components of this species. Efforts should be made to preserve linkages of climax conifer cover between more extensive areas of intact climax forest habitat. Acknowledgments Funding, accommodations, and ongoing support for this project were provided by Pioneer Natural Res- ources Canada Inc., to whom the authors are very deeply grateful. We would also like to thank the fol- lowing (in order they appear in text) for their per- sonal communications: Eric Lofroth, British Columbia Ministry of Environment, Lands and Parks, Victoria, British Columbia; Bill Johnson, Alberta Environment, Peace River, Alberta; John Krebs, Columbia Basin Vol. 118 Fish & Wildlife Compensation Program, Nelson, Bri- tish Columbia; Don Williams, Alberta Environment, Manning, Alberta; Don Reid, British Columbia Minis- try of Environment, Lands and Parks, Smithers, British Columbia; and the late Les Sharp, as well as Larry Smith, fur-trappers in the Chinchaga region of Alberta. Documents Cited (marked with * in text) Kelsall, J. P. 1981. Status report on the Wolverine, Gulo gulo, in Canada in 1981. (Committee on the Status of Endangered Wildlife in Canada (COSEWIC), Ottawa, Ontario. 47 pages. Literature Cited Banci, V. 1994. Wolverine. Pages 99-127 in American marten, Fisher, Lynx and Wolverine in the western United States. Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon and W. J. Zielinski. USDA Forest Service General Technical Report RM-254. Brody, H. 1981. Maps and dreams; Indians and the British Columbia frontier. Douglas & McIntyre. Vancouver, British Columbia, and Toronto, Ontario. Hornocker, M. G., and H. S. Hash. 1981. Ecology of the Wolverine in northwestern Montana. Canadian Journal of Zoology 59: 1286 — 1301. Knorre, E. P. 1959. Ecology of the elk. Jn Transactions of the Pechora-Ilych State Game Preserve by G. A. Novikov. Translated by The Canadian Wildlife Service 1966. 3895. (7): 324 pages. Krohn, W. B., K. D. Elowe, and R. B. Boone. 1995. Relations among Fishers, snow and martens: development and eval- uation of two hypotheses. The Forestry Chronicle 71(1): 97-105. Magoun, A. J. 1985. Population characteristics, ecology, and management of Wolverines in northwestern Alaska. Ph. D. thesis, University of Alaska, Fairbanks, Alaska. 197 pages. Raine, R. M. 1983. Winter habitat use and responses to snow-cover of Fisher (Martes pennanti) and marten (Martes americana) in southeastern Manitoba. Canadian Journal of Zoology 61: 25-34. Rausch, R. L., and A. M. Pearson. 1972. Notes on the Wolverine in Alaska and the Yukon Territory. Journal of Wildlife Management 36: 249-268. Rezendes, P. 1999. Tracking & the art of seeing; how to read animal tracks and sign. Firefly Books, Willowdale, Ontario. 336 pages. Whitman, J. S., W. B Ballard, and C. L. Gardner. 1986. Home range and habitat use by Wolverines in south- central Alaska. Journal of Wildlife Management 50: 460- 462. Zielinski, W. J., and T. E. Kucera. 1995. American Marten, Fisher, Lynx, and Wolverine: survey methods for their detection. USDA General Technical Report PSW-GTR- lsyie Received 19 July 2001 Accepted 26 May 2004 Wolverine, Gulo gulo luscus, Resting Sites and Caching Behavior in the Boreal Forest JONATHAN D. WRIGHT and JESSICA ERNST Ernst Environmental Services, Box 753, Rosebud, Alberta TOJ 2T0 Canada 'Present address: Box 648, East Coulee, Alberta TOJ 1B0 Canada Wright, Jonathan D., and Jessica Ernst. 2004. Wolverine, Gulo gulo luscus, resting sites and caching behavior in the boreal forest. Canadian Field-Naturalist 118(1): 61-64. Wolverine (Gulo gulo luscus) caches and resting sites were examined in a study area in the boreal upland forests of northwestern Alberta and northeastern British Columbia (approximately 57°N). Cache sites were in climax, or ““overmature” stands of Black Spruce (Picea mariana) or mixed-wood of high complexity, dominated by conifers, and in which the Trembling Aspen (Populus tremuloides) and Balsam Poplar (Populus balsamifera) component consisted of mostly dead or dying trees characteristic of such old growth in the boreal uplands. Sites offered relatively good visibility of the surrounding stand. Sites were never located in the dense to extremely dense homogenous spruce stands documented as being favored for travel by Wolverines in the study area. The better used cache complexes were accessed by numerous weil-used trails made by the Wolverines themselves. Caches consisted of the bones, hide and hair of Moose (Alces alces) believed to have been killed by Grey Wolves (Canis lupus). Caches were classified as “simple caches” composed of a single feeding site and/or excavation and “cache complexes” involving one or more feeding “stations”, latrines, resting sites, and climbing trees that may have been used as avenues of escape from competitors/predators. Resting sites were located atop the snow in relatively open locations that offered good visibility of the surroundings. Climax stands were implicated as being of importance to Wolverine caching behavior. Conservation implications include the detrimental effect on Wolverine populations likely to result from current timber harvesting practices | | | in the boreal forest. Key words: Wolverine, Gulo gulo luscus, cache, resting site, climax, boreal forest, stand, Alberta, British Columbia. Knowledge gaps exist involving many aspects of Wolverine life history in forests, including caching behavior and selection of resting sites (Banci 1994). This paper presents observations of these aspects of Wolverine activities as documented over three winters (1997-2000) of snow-tracking efforts in the boreal upland forests of the border between northwestern Alberta and northeastern British Columbia, at approxi- mately 57°N. Methods Wolverine and other furbearer activities were moni- tored during three winter seasons (1997-2000) utilizing snow-tracking methods. The tracking efforts covered approximately 1100 km? in the region of the border country of Alberta and British Columbia known as “Chinchaga” (after the Chinchaga River). Tracking is being increasingly recognized as a useful scientific tool in wildlife studies, management and conservation, with efforts being made to establish standardized termi- nology, to institute university courses on the subject, and to establish networks of trackers throughout the North American continent (Rezendes 1999; Zielinski and Kucera 1995). Snow tracking may be the only practical way of learning details of Wolverine habits and habitat use, as such details are not adequately pro- vided by radio-telemetry studies (Eric Lofroth, personal communication). Approximately 34 000 km were traveled by truck, snowmobile, cross-country skis and on foot, in search of furbearer tracks over the course of the three winters, with an emphasis on locating Wolverines. Markedly different and readily divisible stand characteristics are a feature of the area as a result of the “Great Chinchaga Fire” which blanketed the region during the 1950s (Don Williams, personal communication), burning all but the upland ridges. Fifty years following the fire, two distinct forest types predominate: early-to-mid seral second growth of predominantly aspen or pine in the lowlands, with more limited late-seral, climax or “over- mature” stands predominated by White Spruce and Black Spruce (Picea spp.), and mostly dead and dying Populus spp. in the uplands. The lowland forest may be further characterized as forming a mosaic broken by broad expanses of relatively open Black Spruce fen and open Willow (Salix spp.) muskeg. The upland forest cover is by comparison more continuous. Moose (Alces alces) reach some of their highest densities in North America in the area (Brody 1981), being most heavily distributed in the lowlands, and frequent in the uplands. Groups of Woodland Caribou (Rangifer tarandus caribou) are frequent at low densi- ties in the lowlands. These ungulates offer a generous prey base for the area’s healthy Grey Wolf (Canis lupus) population, whose kills in turn offer a ready food source for the infrequent Wolverines. When Wolverine tracks were located, the individ- uals were fore-tracked (older trails) or back-tracked (fresh trails) in order to gain insights into important and little-known details of the creatures’ use of the 61 62 THE CANADIAN FIELD-NATURALIST boreal forest, including landscape and stand use. Tracks were accessed in the morning by snowmobile or truck, and followed on foot as long as daylight permitted, with the exception of one overnight excursion con- ducted in order to assess advantages and disadvan- tages to the tracking process of remaining in situ. Pertinent details of the Wolverines’ behavior were logged in field notebooks during the tracking event as well as being photographed. Results Wolverine tracks were located on ten occasions over three winters (for a mean of one Wolverine track en- countered per 3400 km of searching). Thirteen sep- arate tracking events resulted, each lasting approxi- mately six hours, and totaling 20 linear km of off- corridor (forest) tracking, or 26.6 total off-corridor km using Magoun’s (1985) estimate of adding 33% to arrive at non-linear distance traveled. Wolverines were tracked for an additional 12.3 km on linear corridors offering conditions of compacted snow, for a total estimated tracking distance of 38.9 km. Five cache sites and three resting sites were encountered and docu- mented using a GPS unit, camera, and sketches in field notebooks. Despite frequent available food in the lowland land- scape, Wolverine tracks were located only in the up- land landscape. Wolverine were revealed to have re- mained in the uplands throughout the tracking events. Wolverine often encountered linear corridors during their travels. Wolverine diverged from their line of travel under the forest canopy to travel on 100% of the linear corridors encountered that offered conditions of compacted snow (n=17), for distances ranging from 3 to 3000 m. In one instance, two Wolverines traveling together followed a compacted (snowmobile) trail on a linear corridor for 1270 m to bring them within 150 m of a cache site, at which point they exited the trail at right angles to follow the shortest linear distance directly to the cache. All five caches observed consisted of bones and/or sections of the hides of Moose (Alces alces), all of which were believed to have stemmed from the Wol- verine’s scavenging of Grey Wolf (Canis lupus) kills. Bones and hide are believed to be a very important component of the Wolverine’s winter economy (Banci 1994; Haynes 1982). Cache-sites appear to be closely frequented until the nutritive value is exhausted, judg- ing from the well-used system of trails and tracks of varying ages in the vicinity of such sites. Cache sites encountered ranged from those consist- ing of a single bone process or an excavation (n = 2), to those better described as “cache complexes” (n = 3). Such complexes were believed to stem from the scay- enging of a single kill, resulting in a series of smaller caches spread over areas which ranged from highly localized to widely radiating. Cache sites were in cli- max, or “overmature” stands of Black Spruce (Picea mariana) or mixed-wood of high complexity, domi- Vol. 118 nated by conifers in which the Trembling Aspen (Populus tremuloides) and Balsam Poplar (Populus balsamifera) component consisted of mostly dead or dying individuals characteristic of such old growth in the boreal uplands. Sites offered relatively good visi- bility of the surrounding stand. Sites were never locat- ed in the dense to extremely dense homogenous spruce stands documented as being favored for travel by Wolverines in the study area, perhaps because of the limited field of vision in such stands. The better-used cache complexes were accessed by numerous well-used trails made by the Wolverines themselves, and had certain features in common, including: e areadily accessible spruce tree (Picea spp.) of rela- tively large dbh (diameter at breast height) selected and well-used for climbing (perhaps as refuge from Wolves, or a safe area to digest between feedings); ° aresting site characterized as a depression in the snow formed by the Wolverine’s body and with a good view of surroundings; e one or more latrines — specific off-trail sites the Wol- verine visited to eliminate wastes; e aradiating series of “feeding areas” characterized by areas approximately 1.5 m* of well-compacted snow, on which bones and bone fragments or sections of Moose-hide were found, but not both, and distinguish- able from a kill site by the lack of Moose stomach- content remains; e excavations [often in the snow, but on more than one occasion (n = 3) in the earth beneath the overhanging boughs of a very large spruce tree], that likely had contained fragments of cached food. A detailed description of the cache and resting sites presented on a site-by-site basis follows, in order as encountered. Cache #1 Cache # I was a widely radiating cache complex. The original Wolf kill site was believed to have oc- curred on a linear corridor regenerating to Lodgepole Pine (Pinus contorta) located at 57° 27° 20” N and 120° 08° 26” W. Old Wolf tracks lead to and from this central location, and Wolverine trails of varying ages, all apparently more recent than the Wolf tracks, were present. Shards of Moose bone were located at a feeding area here. Three additional caches connected by Wolverine trails and radiating from the kill site combined to comprise the cache complex. (1) At 57° 27° 20” N and 120° 08’ 18” W a cache con- sisting of Moose hide remains and an abundance of Moose hair was located. (2) At 57° 27’ 17” N and 120° 08’ 18” W a fresh excava- tion in the earth under the boughs of a very large spruce tree was located. This cache was believed to have been visited by a Wolverine within several hours prior of its discovery. (3) At 57° 27° 22” N and 120° 08’ 06” W a cache consist- ing of bone shards and fragments was located. The shards were larger than those examined at the kill site. Further components of this cache included an excava- tion in the snow down to the sphagnum ground-cover and a latrine. 2004 This cache complex was not characterized by a climbing tree, which was a notable component of some other cache sites. The Wolverine tracked to this cache complex was dragging what was believed to be a trap by one of its left feet. This hindrance apparently ren- dered it incapable of climbing, as nowhere during the tracking event was it observed to have done so, whereas other Wolverines tracked climbed regularly. The trap appeared to affect the individual’s locomo- tion as well; unlike other Wolverines tracked, this in- dividual never varied its gait from a “2x lope” (Zielinski and Kucera 1995). Cache # IT Cache # II was located 150 m from a linear corridor with compacted access (snowmobile trail) that was followed by two Wolverines traveling together, as mentioned, for 1270 m, before exiting the corridor at a right-angle to head directly to the cache. This cache is best described as a localized complex, and appeared to represent the richest of the observed caches as evidenced by heavy use. It was located in a climax stand of somewhat stunted Black Spruce. Four feeding stations were located here, characterized by the existence of bone shards and larger fragments and processes of bone. There were three latrines, two con- taining white scats and one containing brown scats. Resting Site # 1 A resting site, slightly elevated, was located at the north edge of the complex. The resting site consisted of a 43 x 35 cm oval depression atop the snow, at the base of the remains of a short (30 cm), well-wea- thered spruce stump. Snow in the depression was well- compacted. The resting site was relatively open, away from the boles of living trees, and offered a good view of the cache complex and its approaches. It was accessed by four well-used trails, at approximately the four points of the compass. A few guard hairs were present in the depression, along with a few dribbles of urine at the edge of the east access. A climbing tree (Black Spruce) of approximately 14 m was located in a dense clump of smaller spruce at the southeast edge of the cache complex. The tree was larger than the stand’s average and heavily used, as evidenced by claw-marks on the trunk, heavy debris of bark atop the snow at the tree’s base, and the degree of trampling of snow within the spruce clump. Climb- ing on the tree appeared to terminate at a point near its apex where there was a “witches’ broom” formation estimated to be large enough to support a resting Wol- verine; it may have been an additional resting site. No additional trees in the area of the cache complex bore any evidence of having been climbed by a Wolverine. Resting Site # 2 A second resting site was discovered by following the trail of the two Wolverines after leaving Cache # 2. Approximately 200 m from the cache, this resting site again consisted of an oval depression (58 x 43 cm) WRIGHT AND ERNST: WOLVERINE RESTING SITES AND CACHING 63 atop the snow at the base of a relatively large Tama- rack (Larix laricina) tree. Snow in the depression was well-compacted. This resting site was characterized by having more canopy cover than the previous site, although the cover was by no means dense. There was no cache in the immediate vicinity of this resting site, and no apparent climbing tree. Cache # III Cache # III was located at 57° 26’ 84” N and 120° 07° 69” W and is best termed a “simple cache”, rather than a cache-complex. This cache consisted of a single large bone-process and a well-gnawed Moose’s hoof. The cache was located by following the previ- ous two Wolverines, which evidently were aware of, or scented, this cache, as evidenced by their diverting their course of travel by approximately 50 m to reach the cache. There were no other features characteristic of other caches present at Cache # III. Cache # IV Cache # IV located at 57° 27’ 25” N and 120° 06’ 81” W is best termed a cache-complex, although not extensive. Certain features lacking at the last (simple) cache were present at this one. This cache consisted of a collection of bone fragments located on a single feeding platform under a relatively open canopy in a high-complexity climax mixed-wood stand. The cache was accessed by numerous well-used Wolverine trails. No latrines were discovered at this cache. Approximately 10 m from the main cache was an excavation in the earth under the overhanging boughs of a very large spruce tree, accessed by a Wolverine trail, and virtually identical to the excavation discoy- ered at Cache # I. This excavation, too, was deemed to have contained cached food that had been recently recovered by a Wolverine. Cache # IV was further characterized by a well-used climbing tree (spruce) of larger than average dbh, located approximately 4 m from the feeding-station. This cache was located approximately 10 m from a linear corridor with compacted access (snowmobile trail) that was used as a conduit for 700 m in a nor- therly direction by the Wolverine pair upon exiting the cache. This cache had been visited recently by a Marten (Martes americana). Cache #V Cache # V was located in very large, climax mixed- wood dominated by spruce, near the base of a ridge adjacent to Resting Site # III. This was a simple cache, consisting of a single fresh excavation in the earth beneath the overhanging boughs of a large spruce, much as described for caches I and IV. The excava- tion was also believed to be for the purpose of recov- ering cached food. Resting Site # 3 Resting Site # 3 was located along the Wolverine pair’s trail at 57° 27° 72” N and 120° 05’ 11” W. As in the case of the other resting sites observed, this 64 THE CANADIAN FIELD-NATURALIST one consisted of an oval depression atop the snow approximately the same dimensions as the last pre- vious two sites (measurements not taken due to this similarity). The site was mid-slope on a prominent ridge covered by a stand of very large mixed-wood dominated by spruce. The site was relatively open, and offered good view down and along-slope, and only slightly less so upslope. Within 1 m of the resting site was a medium-sized pine tree which had been well- climbed to a height of approximately 4 m. Tracks indicated that this resting site may have been utilized by a third Wolverine which was travel- ing down slope. It appeared as though this third Wolverine left the resting site to travel approximately 15 m along-slope to meet the Wolverine pair as they traveled upslope. The single Wolverine may have util- ized the climbing-tree to observe the approaching pair. A confusion of tracks resulted where the three Wolverines apparently met. It is believed that all three animals subsequently traveled upslope together. Discussion Caches were located in climax stands of less density than those apparently favored for travel by Wolverines in the study area. Visibility of approaching competi- tors/predators (Wolves) may be the deciding factor in both cache-site and above-snow resting-site selection. That the caches were all located in climax timber re- inforces the importance of this stand-type to Wolver- ine. Present forest practices that result in the harvest- Vol. 118 ing of climax stands may negatively impact Wolver- ine populations. Acknowledgments Funding, accommodations and ongoing support for this project were provided by Pioneer Natural Res- ources Canada Inc., to whom the authors are deeply grateful. Literature Cited Banci, V. 1994. Wolverine. Pages 99-127 in American marten, Fisher, Lynx and Wolverine in the western United States. Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, L. J. Lyon and W. J. Zielinski. USDA Forest Service Gen- eral Technical Report RM-254. Brody, H. 1981. Maps and dreams; Indians and the British Columbia frontier. Douglas & McIntyre. Vancouver, Bri- tish Columbia; Toronto, Ontario. Haynes, G. 1982. Utilization and skeletal disturbances of North American prey carcasses. Arctic 35: 266-281. Magoun, A. J. 1985. Population characteristics, ecology, and management of Wolverines in northwestern Alaska. Ph.D. thesis. University of Alaska, Fairbanks, Alaska, 197 pages. Rezendes, P. 1999. Tracking & the art of seeing; how to read animal tracks and sign. Firefly Books, Willowdale, Ontario. 336 pages. Zielinski, W. J. and T. E. Kucera. 1995. American marten, Fisher, Lynx, and Wolverine: survey methods for their detection. USDA General Technical Report PSW-GTR- ilyt/s Received 19 July 2001 Accepted 26 May 2004 Seasonal Home Ranges of Raccoons, Procyon lotor, Using a Common Feeding Site in Rural Eastern Ontario: Rabies Management Implications SARAH C. ToTTon,! RICHARD C. ROSATTE,? ROWLAND R. TINLINE,? and LAURA L. BIGLER‘ '340 Second Avenue West, Owen Sound, Ontario N4K 4L7 Canada ? Ontario Ministry of Natural Resources, P.O. Box 4840, Peterborough, Ontario K9J 8N8 Canada > Geographic Information Systems Laboratory, Queen’s University, Kingston, Ontario K7L 3N6 Canada + Zoonotic Disease Section, Diagnostic Laboratory, College of Veterinary Medicine at Cornell University, P.O. Box 5786, Upper Tower Road, Ithaca, New York 14852-5786 USA Totton, Sarah C., Richard C. Rosatte, Roland R. Tinline, and Laura L. Bigler. 2004. Seasonal home ranges of Raccoons, Procyon lotor, using a common feeding site in rural eastern Ontario: Rabies management implications. Canadian Field-Naturalist 118(1): 65-71. Thirteen adult Raccoons (Procyon lotor) (six females, seven males) that fed at a garbage dump north of Kingston, Ontario were radio-tracked from 21 June to 16 October 1995 to assess their seasonal home ranges and movements. Average Minimum Convex Polygon (MCP) summer and fall home ranges for the collared Raccoons were 78.4 ha (SD=46.2 ha) and 45.6 ha (SD=29.7 ha), respectively. Average grid cell summer and fall home ranges for the collared Raccoons were 143.3 ha (SD=40.0 ha) and 116.9 ha (SD=24.9 ha), respectively. Summer ranges of the Raccoons were significantly larger than fall ranges using both the MCP method (P=0.05) and the grid cell method (P=0.073). Yearling Raccoons travelled an average summer maximum distance from the dump of 2608 m (SD=1964, n=3), more than double the distance of adults (22 yr) at 1239 m (SD=547, n=10). The population density for the study area in late August 1995 was estimated at | Raccoon/12 ha based on an effective area surrounding the dump of 234 ha. Home range and movement data may be useful to design a strategy to control Raccoon rabies in Ontario. Key Words: Raccoon, Procyon lotor, rabies, communal feeding, disease transmission, field study, home range, telemetry, Ontario. Raccoon (Procyon lotor) rabies was first reported in Ontario, Canada, during July 1999 (Wandeler and Salsberg 1999; Rosatte et al. 2001). Point infection control methodologies are currently being used (since 1999) in Ontario to contain the outbreak to a small area (Rosatte et al. 2001). As well, a Raccoon rabies model is being developed in Ontario to assist with the control of the disease by predicting the rate of spread of Raccoon rabies both temporally and spatially. Know- ledge on Raccoon home ranges, movements and popu- lation dynamics in Ontario is needed to develop and validate the rabies model so that it reflects the actual sequence of events that occur during a Raccoon rabies epizootic/enzootic (Broadfoot et al. 2001). The spatial distribution of food is known to influ- ence contact rates in Raccoons (Procyon lotor) (Sei- densticker et al., 1988). Clumped food resources such as garbage dumps may increase potential contact rates of Raccoons as these resources cause members of a population to congregate from a wide area. This in turn may influence rabies transmission in the popula- tion (Seidensticker et al. 1988). Range of movements of Raccoons using common feeding sites may be a useful indicator of potential rabies spread. Such infor- mation can be used to design effective baiting strate- gies to vaccinate these animals against rabies as well as provide input for the development of rabies models. Home ranges of Raccoons tend to shift due to sea- sonal changes in behavior and therefore must be calcu- lated separately for each season (Kauffmann 1982). Summer is the family rearing period when lactating mothers and their offspring travel together and it is also the main dispersal period for yearling males (Mech et al. 1968; Fritzell 1978). Fall is a time when the juve- niles may disperse and Raccoons prepare for the com- ing winter dormancy period (Shirer and Fitch 1970). In this study, home range was defined using criteria of White and Garrott (1990) as the area within which the animal normally moved in a specified time frame, in this case the summer and fall of 1995. In this study, movements of Raccoons using a com- mon feeding site in rural eastern Ontario, Canada, were determined by radio-telemetry to assess the size of their summer and fall home ranges. The same Raccoons on which contact data were obtained in the Totton et al. (2002) study were used. In addition, two Raccoons, caught at a smaller feeding site (compost heap) were tracked periodically to determine their daytime rest- ing sites. The population density of Raccoons in this study was also measured as it influences home range and contact rate. Study Area and Methods Trapping took place at a private garbage dump (44°34’N, 76°20’W) and at a compost bin on the grounds of the Queen’s University Biological Station 40 km north of Kingston, Ontario (44° 35’N, 76° 19° W). The area surrounding the dump consisted of farm land (livestock), forest, marsh, and cottages (most of which were only occupied during the summer). The entire 65 66 THE CANADIAN FIELD-NATURALIST study area was about 460 ha. Fifteen Raccoons (eight females and seven males) were collared between 20 May and 27 June 1995. Raccoons were captured us- ing Tomahawk #106 (Tomahawk Live-trap Company, Tomahawk, Wisconsin, USA), and Havahart #1079 (Havahart Live Trap Company, Niagara Falls, Ontario, Canada) live-traps. All Raccoons were ear-tagged (numbered size | and 2, National Band and Tag Com- pany, Newport, Kentucky), vaccinated against rabies (Imrab® inactivated rabies vaccine, Merieux, Inc., Athens, Georgia, USA) and canine distemper (Fromm D, modified live virus, SOLVAY animal health, Inc., Mendota Heights, Minnesota, USA). They were im- mobilized by intramuscular injection of ketamine hy- drochloride (Rogar/STB Inc., London, Ontario, Can- ada) and xylazine hydrochloride [Rompun] (Bayvet, Rexdale, Ontario, Canada) [30 mg/kg body weight ketamine, 10:1 ratio ketamine:rompun]. We deter- mined their sex and extracted a first premolar tooth for age determination by cementum analysis (Johnston et al. 1987). Each animal was then fitted with an adjus- table radio-collar [151 to 152-Mhz] (Lotek Engi- neering Inc., Newmarket, Ontario, Canada) and rel- eased at its point of capture. The radio-tracking system consisted of a four- element Yagi antenna, 151 MHz (FM) transmitters mounted on whip antenna collars (Lotek Engineering, Newmarket, Ontario), one programmable hand-held receiver (Lotek model SRX-400; Lotek Engineering, Inc., Newmarket, Ontario) that operated in the 151- 152 MHz range, one hand-held compass, and a four- wheel drive pick-up truck. Animals were given at least seven days to acclimatize to their collars before radio-tracking began, in accordance with White and Garrott’s (1990) recommendations. The tracking per- iod lasted from 21 June to 16 October 1995 with attempts being made to locate each Raccoon two to three times per week. Since only one telemetry receiver was available, sequential rather than simultaneous bearings had to be taken. A maximum interval between first and last bearings of 10 min was set to minimize telemetry error caused by animal movement (except for bearings taken during the day when the animals were inactive, at which time the interval may have been longer). In a study by Gert and Fritzell (1996), 23% of the locations came from triangulations with between-bearing intervals in excess of 8 min. For this reason, the 10 min cut-off was deemed reasonable for this study. Continuous radio-tracking (i.e., location of the ani- mals at least every 15 min (Harris et al. 1990)) was not feasible with only one receiver; therefore, for this study, discontinuous tracking was performed. Location estimates were made for each animal three or four times between dusk and dawn at roughly 2-h intervals, and once during the following afternoon to determine day- time resting sites. The tracking schedule was construct- ed by randomly selecting six of the collared animals Vol. 118 trapped at the dump for one given tracking night. The remaining Raccoons were then tracked on the next scheduled night. A different set of Raccoons was ran- domly chosen for the following tracking night and so on. Dates of tracking nights were randomly selected for each week. Three types of location estimate techniques were used: scanning, triangulation, and homing. Scanning involved tuning into the collar frequencies of the dump animals while the researcher sat in the middle of the dump area. Data from telemetry accuracy tests indi- cated a mean transmitter-receiver distance of 240 m+ 30 m (n=20) when the signal was picked up at a gain of 10. A Raccoon was therefore considered to be in a radius of this distance from the center of the dump area if its signal was detected from there at a gain of 10 or less. Most of the locations for the dump animals were obtained by triangulation. This technique involves tak- ing directional bearings from two to three different receiver sites at known locations and using these to estimate the true location of a remote transmitter on the animal’s collar (White and Garrott 1990). Accuracy tests were performed to determine the error associated with locations estimated by triangu- lation in this study. Bearing accuracy has two compo- nents: bias (the average difference between the true bearing and the bearing estimated by the receiving system for a series of receiver-transmitter locations), and precision, which is the standard deviation of these errors (White and Garrott 1990). The bias was 9° and was significantly different from O° (t=3.61, n=63, P<0.001; one sample t-test (Zar 1996)). Therefore 9° was subtracted from all bearings. Precision of the sys- tem was + 20°. Screening criteria derived from telemetry accuracy tests were applied to all bearings to eliminate errors due to signal bounce. All bearings which did not inter- sect with other bearings taken on the same transmit- ter were removed. All bearings taken from receiver sites associated with large errors (high levels of sig- nal bounce) were removed from subsequent analysis. Also, bearings taken when the transmitter-receiver dis- tance was >2000 m were removed. In addition, all loca- tions involving distances between transmitter and rec- eiver of over | km were inspected for plausibility. A computer program called TRIANG was used to estimate animal locations from triangulated bearing pairs and to calculate the distance between receiver and transmitter for each bearing. TRIANG did not compute animal locations when three bearings were taken. In this case, locations were determined by plot- ting the bearing angles in AutoCAD and estimating the centre of the triangle created by the intersection of the bearings. Locations for two of the collared Raccoons were usually obtained by homing (White and Garrott 1990). Locations of animals determined by homing are not affected by error in the telemetry system; however, they 2004 are affected by the researcher’s ability to pinpoint the den location on a map (White and Garrott 1990). All locations determined by homing in our study were plotted by hand on 1:10 000 maps of the area to +50 m to obtain Universal Trans Mercator Co-ordinates (UTMC). Home ranges were estimated using the minimum convex polygon (MCP) method (Mohr 1947), since this is the only home range method that is strictly com- parable between studies (Harris et al. 1990). Ranges for both seasons were combined to compare degree of overlap between seasons. Because it is advantageous to use more than one home range estimate technique (Voigt and Tinline 1980), the grid cell method of home range analysis (Siniff and Tester 1965) was also used. Size of the grid cells was chosen to reflect radio fix accuracy based on the results of accuracy tests. In order to enclose the uncertainty area associated with scanning the dump for transmitter signals (and this was larger than the area associated with triangu- lation) a grid square would have to measure 480 m on each side. Therefore this was the size of grid square (23 ha) used in estimating grid cell home ranges. The grid was oriented by centering a grid square over the dump site. Summer home ranges were calculated from teleme- try data collected in June, July, and August; Septem- ber and October fixes were used to calculate fall home ranges. Two female Raccoons were excluded from the analysis because insufficient locational fixes (<16) were acquired during the study. A Mann-Whitney U test (Zar 1996) and Statistica Version 6.0 software (StatSoft Inc., Tulsa, Oklahoma) were used to test for statistical differences in Raccoon home range size. Raccoons were grouped into regular and occasional dump visitors based on the number of nights each animal was seen at the dump out of a total of 35 observation nights in a concurrent behavioral study (Totton et al. 2002). Raccoons were defined as regu- lars if they were observed at the dump on >51% of all observation nights and occasionals if they were seen at the dump on <51% of the observation nights. Since only one telemetry receiver was available, and hence discontinuous locational fixes had to be ob- tained, detailed analysis of the movement patterns of raccoons in this study was not possible. Trapping to estimate the Raccoon population size took place at the dump from 28 August to 19 Sep- tember 1995. At this time of year, juveniles are larger and easier to trap and handle than they are earlier in the summer (Seidensticker et al. 1988). The number of Raccoons in the study area was estimated using a modified Petersen Index (Begon 1979). Density of the dump population was not based solely on the area of the trapping site (garbage dump = 2.3 ha) because it was evident from telemetry data that some Rac- coons were travelling from a much wider area to feed at the dump. Instead, using the number of Raccoons TOTTON, ROSATTE, TINLINE, AND BIGLER: RACCOONS 67 calculated by the methods above, estimates were made of the crude density of the population as defined by Seidensticker et al. (1988) based on the average maxi- mum width of the dump Raccoons’ summer (MCP) home ranges (1530 m). Since Seidensticker et al. (1988) did not specify how this distance was used to calculate overall area, a square was centred over the dump with each side equal to the distance calculated and this was used for calculation of crude density. Its area was 234 ha. Results For locations determined by triangulation, distance between the observer and estimated transmitter loca- tion ranged from 8.3 m to 1837.9 m and averaged 330.5 m (SD=239.3 m). Uncertainties in Raccoon position for triangulated bearings (tan 20° x trans- receiver distance) ranged from +3.0 m to +668.9 m with a mean of +120.3 m and standard deviation (SD) of 87.1 m (n=1181). Overall locational uncertainty, including that associated with scanning and homing techniques as well as triangulation, was +146.6 m (n=1110). No statistical differences were detected between male and female ranges for values calculated using either the MCP method (P=0.78) or the grid cell meth- od (P=0.26). Therefore, male and female data were pooled to compare summer and fall ranges. As well, only three yearlings were trapped at the dump in this study and home range data were available for two of those. Consequently, statistical comparisons between adult and yearling home range sizes were not per- formed. No differences could be detected between home ranges of Raccoons that regularly versus occa- sionally visited the dump. Average Minimum Convex Polygon (MCP) summer and fall home ranges for the collared raccoons were 78.4 ha (SD=46.2 ha) and 45.6 ha (SD=29.7 ha), respectively (Table 1). Average MCP summer/fall home range overlap was 31.2 ha (SD=17.2). Average grid cell summer and fall home ranges for the collared Raccoons were 143.3 ha (SD=40.0 ha) and 116.9 ha (SD=24.9 ha), respective- ly (Table 1). Average grid cell home range overlap between summer and fall was 95.8 ha (SD=27.5). Summer ranges of the Raccoons were significantly larger than fall ranges using both the MCP method (P=0.05) and the grid cell method (P=0.073). The most widely ranging Raccoon (a male year- ling) in the study was originally trapped at the dump on 27 June. It was later located by telemetry near a farmhouse 4 km northeast of the dump on 16 July. By 18 July, it was visually identified at the dump site again where it remained until 8 August. The only other yearling of the dump Raccoons wandered a maximum of 769 m from the dump during the summer. The aver- age distance of the farthest fix from the dump during the summer for the yearling Raccoons was 2608 m (SD=1964, n=3) and for the adults (2 2 yr), 1239 m 68 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Summer (June to August) and fall (September to October) home ranges of 13 Raccoons which fed at a rural Ontario garbage dump and at a compost heap in 1995!. Method Number Summer of fixes range (summer) (ha) mean (SD)? mean (SD) MCP SZe (ile) 78.4 (46.2) Grid Cell 44 (6.7) 143.3 (40.0) Number Fall Area of overlap of fixes range of ranges (fall) (ha) mean (SD) mean (SD) mean (SD) 34.1 (9.6) 45.6 (29.7) Sil= (GU) 33a (92) 116.9 (24.9) 95.8 (27-5) 'n=13 raccoons {7 males — 6 adults and | yearling; 6 females — 5 adults (all lactating) and 1 yearling} 2 §D=Standard Deviation TABLE 2. Home range, movements and density of Raccoons in different areas of North America Location Habitat Home Range Movements (km?) (km) Ontario rural 0.5-4.0 4-45 North Dakota rural 0.2-49 1-24 Minnesota rural 7-12 >3 Wisconsin Toronto urban 0.4 3Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8W 9W2 Canada ‘Strawberry Isle Research Society, Box 213, Tofino, British Columbia VOR 2Z0 Canada >Present address: World Wide Fund for Nature, GPO Box 528, Sydney, NSW 2001 Australia McAlpine, Donald F., Stan A. Orchard, Kelly A. Sendall, and Rod Palm. 2004. Status of marine turtles in British Columbia waters: a reassessment. Canadian Field-Naturalist 118(1): 72-76. Marine turtles in British Columbia have previously been considered off course stragglers. Here we document 20 new reports for Green Turtles, Chelonia mydas, and Leatherback Turtles, Dermochelys coriacea, for the province. Until recently there had been no concerted effort to acquire data on marine turtle abundance or frequency off British Columbia. Observations presented here allow a reassessment of marine turtle status in British Columbia waters. We suggest Green Turtles and Leatherbacks should be considered rare vagrants and uncommon seasonal residents, respectively, off British Columbia and that they are a natural part of the British Columbia marine environment. Key Words: Green Turtle, Chelonia mydas, Leatherback Turtle, Dermochelys coriacea, British Columbia, status. There are few published reports for marine turtles in British Columbia waters. Marine turtles in the prov- ince have generally been considered “straggler[s]” (Kermode 1932) or “off course” (Gregory and Camp- bell 1984). Nussbaum et al. (1983) ignore marine tur- tles in their treatment of northwest amphibians and reptiles, and marine turtles are not included in a recent field guide to northwest reptiles “because sea turtles rarely visit these shores” (St. John 2002). During the preparation of a review of the status and conservation of marine turtles in Canadian waters (McAlpine et al. in press) recent occurrences of Green Turtles off Brit- ish Columbia were encountered (McAlpine et al. 2002). Here we document additional new reports for Green Turtles, Chelonia mydas, as well as Leatherback Tur- tles, Dermochelys coriacea, for British Columbia. These new records nearly double the number of published occurrences for marine turtles from the province. When considered in the context of a recent compilation for adjacent Alaska (Hodge and Wing 2000), these obser- vations suggest a reassessment of sea turtle status in British Columbia waters is warranted. The precipitous decline in Pacific Leatherback numbers (Spotila et al. 2000), so serious it has led to a call for a moratorium on long-line fishing in the Pacific Ocean (Anonymous 2002), also provides a timely context. Based on terms used to document bird status in Alaska, Wing and Hodge (2002) have proposed a standardized terminology to describe marine turtle occurrences. This terminology, ranging through seven categories from accidental to abundant, is loosely based on number of occurrences only. Although useful, this system ignores the ecological context for occurrence. Here we suggest modifying the system of Wing and Hodge (2002) by adding, where appropriate, one of four descriptors to their categories: resident, seasonal 72 resident, migrant, or vagrant. Resident refers to turtles present year round, seasonal resident refers to turtles present only at certain times of the year, migrant refers to turtles moving through a broad area to another loca- tion, vagrant refers to turtles occurring outside their normal resident or migratory range. We use this system, in conjunction with that of Wing and Hodge (2002), to describe marine turtle status in British Columbia. There are 11 published reports for Green Turtles (Carl 1955; Radovich 1961 cited in Stinson 1984; Hodge and Wing 2000; McAlpine et al. 2002) from British Columbia and 14 reports of Leatherbacks (Ker- mode 1932; Carl, 1944, 1960; MacAskie and Forester 1962). Published and unpublished marine turtle reports for British Columbia prior to 1982 are summarized in Stinson (1984). Table | lists one unpublished British Columbia Green Turtle report, included by Stinson (1984) and six new ones; three unpublished Leatherback reports included in Stinson (1984) and 10 new Leatherback occurrences. There are currently 26 British Columbia reports for the Leatherback accompanied by sufficient locality data to map (McAlpine et al. in press), as well as other reports which are not site-specific. All of the 16 British Colum- bia Green Turtle reports can be map plotted (McAlpine et al. 2003). Several of the recent Green Turtle records are supported by photographic evidence (Figures 1- 3), as are four of the Leatherback sightings (Figure 4; Table 1). The skull in Figure | is readily identified as that of C. mydas using the figures in Wyneken (2001). This stranding is reported in issue Number 2 of KSM Wutsiin, the official newletter of Hartley Bay School. Unfortunately, the skeletal remains of this turtle were not retained. Detailed necropsies carried out on the Green Point and Matlahaw Point C. mydas are now archived at the Royal British Columbia Provincial 2004 MCALPINE, ORCHARD, SENDALL, AND PALM: MARINE TURTLES 73 TABLE |. Recent occurrences of marine turtles from British Columbia waters. Date Location Chelonia mydas 22 November 1996 Hartley Bay early November 1998 mouth of Tlell R early November 1998 5 km S of Tlell R 6 November 2001 4km N of Tlell R 20 December 2001 Greene Point 21 January 2002 Matlahaw Point Dermochelys coriacea mid-July 1970 Sharbau Island summer 1977 Off Ucluelet September 1977 Off Ucluelet 30 August 1981 Skidiyate Inlet Hectate Strait Esperanza Inlet 1 September 1981 16 September 1982 August 1993 Clerke Point 26 May 1996 Mt Douglas Park May 1997 Kyuquot Sound September 1997 SE of Langara Island Spring 1998 Tofino 6 September 2000 __ off shore 4 August 2001 off Langara Island 'PC = personal communication. 3525) S336) SS nS9: SS esi) 49° (3' 49° 23' S130; 49° 55' 49° (0! Sul: 53° 20' 49° 52' 50° 05' 49° 08' 50° 05' 54° 12' 49° 07' 48° 43' 54° 18' 18" 129215) 131° 56' 131° 54' ISH aps) 125° 43' 126° 29' 128° 45' 126° 38' 125° 50’ 131 730% 130° 30' 126° 44! 127° 48' 123° 58' W27e213\ 132° 58' 1252753: 127° 26' 1335210; Latitude(N) Longitude(W) Comments Carcass, Figure | Carcass, PC!: W. Flood to DFM Carcass, PC: W. Flood to DFM Fresh carcass, carapace 78.7 cm, Figure 2 Fresh carcass, male, carapace 68.7 cm, Figure 3 Carcass, male, carapace 69.9 cm Stinson (1984) Stinson (1984) Free swimming, RBCPM photo number 695 Alive in salmon gillnet, RBCPM photo number 749 Stinson (1984) RBCPM photo number 826 Feeding on Cyanea, pc: J. Watson to G. Ellis Free swimming/Times-Colonist, 1 June, page B2 Carcass, PC: G. Jamieson to DFM Free swimming, large numbers of Vellela sp. present, PC: S. Buchanan to S. Stebbins Decomposed carcass, PC: to SAO Free swimming, PC: K. Morgan to DFM Free swimming, PC: E. Simkin to G. Ellis, Figure 4 Ficure 1. Skeletal remains of a dead Green Turtle stranded at Hartley Bay, British Columbia, 22 November 1996. The skull in this photo can be readily identified as that of C. mydas (Hartley Bay School photo). 74 THE CANADIAN FIELD-NATURALIST Museum and the Strawberry Isle Research Society, Tofino. A report prepared by a veterinary pathologist following examination of the Green Point Turtle states this turtle died from a bacterial infection to which it may have been predisposed following exposure to environmental stressors, such as low water tempera- ture. The animal was also suffering from a moderately severe case of pneumonia. The Matlahaw turtle ap- peared healthy and cause of death was not determined. Gregory and Campbell (1984) suggested the single live Green Turtle from British Columbia reported by Carl (1955) was “probably just off course”. On the basis of minimum thermal tolerance in cheloniid turtles and the few additional reports of live C. mydas. McAlpine et al. (2002) concurred, considering the species “accidental” in the province. Kermode (1932) identified the Leath- erback as a “‘straggler” in the North Pacific, Carl (1944) reported the species as “wandering” to British Colum- bia waters, and Cook (1981*) considered Leatherback Turtles observed in Canada were either “migrants or strays”. Stinson (1984), however, suggested occurrences of sea turtles in the northeastern Pacific could not be dis- missed as accidental but are influenced by temperature anomalies in the ocean. While she found sightings of marine turtles occur regularly during years of normal ocean temperatures, greater numbers of turtles were observed than expected when sea temperatures were above the mean. Hodge and Wing (2002) noted that in Alaska marine turtle occurrences are almost equally divided between warm-water and normal-water years. Most noteworthy was the complete lack of turtles in cold-water years. Whether the increased number of recent reports reflects a real increase in the prevalence of marine turtles off British Columbia, or simply increased public awareness and reporting, is unknown. However, based on their standardized occurrence terminology for < ESTOORG Ata oy AR RY a ARAN et os Ga FIGURE 2. This 78.7 cm Green Turtle came ashore fresh dead on 6 November 2001, 4 km N of the Tlell River, Queen Charlotte Islands, British Columbia (V. Flood photo). Vol. 118 FIGURE 3. This 68.7 cm male Green Turtle came ashore fresh dead at Green Point, Vancouver Island, British Col- umbia, on 20 December 2001. (P. Clarkson/Parks Can- ada photo). marine turtles, Wing and Hodge (2002) report that marine turtles occur too often in Alaska waters to be considered accidental. These authors consider marine turtles a natural part of the Alaskan marine environ- ment. Likewise, Stinson (1984) concludes marine turtles are a regular part of the environment in the northeast Pacific. Based on the frequency of reports, Wing and Hodge (2002) considered the Green Turtle and Leatherback Turtle as rare and uncommon, respec- tively, in Alaska. Stinson (1984) identified a marine turtle season of July to September in the northeast Pacific Ocean and stated September was the critical marine turtle month in British Columbia waters. How- ever, nearly all of her British Columbia marine turtle observations are of Leatherbacks. Currently, the sea- sonal pattern of occurrence that appears to be emerg- ing, both in British Columbia and Alaska, suggests that D. coriacea is indeed most frequent during the July to September period, while C. mydas is most likely to be found stranded in the northeast Pacific during October to December. It should be noted that most British Columbia Green Turtle observations have been of stranded turtles, often recently deceased. Hodge and Wing (2000) observe that the low frequency of hard shell turtles from Alaska waters indicates these turtles are straying beyond their toler- able range, while they consider Dermochelys in Alaska as occupying marginal habitat that may not be con- sistently used from year to year. The accumulating number of marine turtle observations from British Columbia suggests that these species are even more frequent in British Columbia than Alaska. Following 2004 MCALPINE, ORCHARD, SENDALL, AND PALM: MARINE TURTLES 75 Ficure 4. Leatherback turtle photographed 4 August 2001 off Langara Island, Queen Charlotte Islands, British Columbia (Erik Simkin photo). Wing and Hodge (2002), and our own suggestions above, we describe C. mydas as a rare vagrant and and D. coriacea as an uncommon seasonal resident in British Columbia waters. However, we would add there has been no effort to methodically collect infor- mation from dead stranded turtles nor, until a recent initiative by the Vancouver Aquarium (C. Sbrocchi, personal communication to DFM), to acquire sightings data on marine turtle abundance or frequency off British Columbia. Anecdotal information on several Leatherback occurrences is included in the KSM Wutsiin newsletter noted above, indicating canvas- sing local communities, and especially native fishers, could be a source of information on marine turtle occurrences in British Columbia. Additionally, Glen Jamieson (Pacific Biological Station, personal commu- nication to DFM) reports that Leatherbacks are “periodically seen off the west coast of [Vancouver Island]” and Frank Bernard (Pacific Biological Station, personal communication to SAO) reported that he had seen as many as six Leatherbacks in a day 80-320 km off the west coast of Vancouver Island. Reports of the Pacific Ridley, Lepidochelys olivacea, and the Logger- head, Caretta caretta, for Alaska (Hodge and Wing 2000) suggest focused investigations may reveal these species also occur off the British Columbia coast. Acknowledgments We thank the following individuals for sharing British Columbia marine turtle observations and infor- mation with us and allowing us to include their reports here: Scott Buchanan, Barry Campbell, Graeme Ellis, Glen Jamieson, Marilyn Joyce, Wayne and Virginia Flood, Ken Morgan, Carla Sbrocchi, Eric Simkin, Jane Watson, Kris Willcock, and the late Frank Bernard. In particular, we would like to thank Eric Hill and Simone Westgarth of the Hartley Bay School for their efforts in obtaining information that allowed identifi- cation of the Hartley Bay turtle. Documents Cited (marked * in text) Cook F. R. 1981. Status report on the Leatherback Turtle, Dermochelys coriacea. Committee on the Status of En- dangered Wildlife in Canada, Ottawa, 17 pages. Literature Cited Anonymous. 2002. Moratorium on Pacific long-lining could save leatherbacks. Marine Turtle Newsletter 97: 26. Carl, G. C. 1944. The reptiles of British Columbia. Hand- book Number 3. British Columbia Provincial Museum, Victoria, British Columbia. Carl, G. C. 1955. The green turtle in British Columbia. Report of the Provincial Museum of Natural History and Anthropology, Victoria, British Columbia 1954: B77-78. 76 THE CANADIAN FIELD-NATURALIST Carl, G. C. 1960. The reptiles of British Columbia. Hand- book Number 3. British Columbia Provincial Museum, Victoria, British Columbia. Gregory, P. T., and R. W. Campbell. 1984. The reptiles of British Columbia. Handbook number 44. British Colum- bia Provincial Museum, Victoria, British Columbia. Hodge, R. P., and B. L. Wing. 2000. Occurrences of marine turtles in Alaska waters 1960-1998. Herpetological Review 31: 148-151. Kermode, F. 1932. A remarkable capture of Leatherback Turtles off Bajo Reef, near Nootka Sound, West Coast of Vancouver Island, British Columbia. Report of the Prov- incial Museum of Natural History for the year 1931. Victoria, B.C. Pages 6-7. MacAskie, I. B., and C. R. Forrester. 1962. Pacific Leather- back off the coast of British Columbia. Copeia 1962: 646. McAlpine, D. F., M. C. James, J. Lien, and S. A. Orchard. In press, Status and conservation of marine turtles in Canadian waters. Jn Ecology, conservation and status of reptiles in Canada. Edited by C. R. Seburn and C. A. Bishop. Herpetological Conservation 3. McAlpine, D. F., S. A. Orchard, and K. A. Sendall. 2002. Recent occurrences of the green turtle from British Colum- bia waters. Northwest Science 76: 185-188. Vol. 118 Nussbaum, R. A., E. D. Brodie Jr., and R. M. Storm. 1983. Amphibians and reptiles of the Pacific Northwest. University of Idaho Press, Moscow, Idaho. Radovich, J. 1961. Relationship of some marine organisms of the northeast Pacific to water temperatures, particularly during 1957 through 1959. California Fish and Game, Fisheries Bulletin 112: 1-62. Spotila, J. R., R. D. Reina, A. C. Steyermark, P. T. Plotkin, and F. V. Paladino. 2000. Pacific Leatherback Turtles face extinction. Nature 405: 529-530. Stinson, M. L. 1984. Biology of sea turtles in San Diego Bay, California and in the northeastern Pacific Ocean. MSc. thesis, San Diego State University, California. St. John, A. 2002. Reptiles of the Northwest: British Colum- bia to California. Lone Pine Publishing, Edmonton, Alberta. Wing, B. L., and R. P. Hodge. 2002. Occurrence terminology for marine turtles. Marine Turtle Newsletter 95: 15-16. Wyneken, J. 2001. The anatomy of sea turtles. U.S. Depart- ment of Commerce, NOAA Technical Memorandum NMEFS-SEFSC-470. 172 pages. Received 27 February 2003 Accepted 20 November 2003 | | Introduced Marine Species in the Haida Gwaii (Queen Charlotte Islands) Region, British Columbia N. A. SLOAN and P. M. BARTIER Parks Canada, Gwaii Haanas National Park Reserve and Haida Heritage Site, P.O. Box 37, Queen Charlotte, British Columbia, VOT 1S0 Canada Sloan, N. A., and P. M. Bartier. 2004. Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) region, British Columbia. Canadian Field-Naturalist 118(1): 77-84. This historical review of a marine area’s introduced species was facilitated by geo-referenced marine species inventories of the Haida Gwaii (Queen Charlotte Islands) region. One plant, 14 invertebrate, and two fish introduced species have been recorded since the early 20" century from the marine waters around Haida Gwaii. Records of species occurrences are listed and mapped, and modes of introduction are discussed. It will be important to continue documenting areas’ introduced species locations to track the progress of invasions that could affect local marine ecosystem well-being. Key Words: introduced species, marine, Haida Gwati, Queen Charlotte Islands, British Columbia. *“... the control of alien marine species is in its infancy.” (Bax et al. 2001) Introduced (non-indigenous) marine species are of global concern (Bax et al. 2001). The dynamism and connectivity of marine ecosystems, mariculture and shipping facilitate species’ introductions. The introduc- tion of some species may be followed by rapid local dispersal of propagules and appreciable ecological con- sequences (Grosholz 2002). Introduced species know]- edge is more developed for land and freshwater than for marine ecosystems (Ruiz et al. 2000; Simberloff 2000; Bax et al. 2001). Carl and Guiguet (1958) were the first to broadly document species introductions in Pacific Canada. Cur- rent awareness of the problem is growing in the region, and Levings et al. (2002) have reviewed introduced marine species found in the Strait of Georgia, south- ern British Columbia. Using the 30 introduced mollusks along the Pacific coast of North America as an example, Carlton (1992) reported the main modes of introduction as: (1) asso- ciated with introduced Japanese and Atlantic oysters imported for mariculture (approximately 27 species), (2) via vessel hull fouling and organisms living in wood, and (3) from overseas ships’ ballast water releases. More recently, Chapman et al. (2003) reported that along the U.S. Pacific coast, 24 of the 37 marine and estuarine bivalve species commercially available as sea- food are alien species. We report on the documented presence of introduced marine plants, invertebrates and fishes in the Haida Gwatii (Queen Charlotte Islands) region of northern British Columbia. No introduced marine birds or mam- mals have been recorded. The plant and invertebrate records, gleaned from the literature and museum col- lections, are archived in the geographic information system (GIS) databases of Gwaii Haanas National Park Reserve and Haida Heritage Site (Sloan and Bartier 2000; Sloan et al. 2001). We excluded introduced species whose total geographic range encompasses the Haida Gwaii region, although, of course, their presence could be inferred. An example is the amphipod Coro- phium acherusicum from Asia and now known from the northern mainland British Columbia coast (Bousfield and Hoover 1997). The American Shad (Alosa sapidis- sima) have long been known from the whole northeast Pacific (Welander 1940), but only recently from Haida Gwaii waters (Workman et al. 1996). Atlantic salmon (Salmo salar) observations were from the Atlantic sal- mon Watch Program web site: http://www.pac.dfo-mpo. gc.ca/sci/aqua/AS WP_e.htm. The fish records are not yet in our database. Results and Discussion For Haida Gwaii, accidental (or active) introductions of one plant, 14 invertebrate, and two fish species asso- ciated with fishery or mariculture development are summarized in Table 1 and illustrated in Figures | and 2. Likely some of these species have spread northward to Haida Gwaii from the more developed southern mainland British Columbia and U.S. coasts where they first became established. There have been introductions of species to British Columbia targeted for mariculture, such as Pacific Oyster (Crassostrea gigas) from Japan, plus their attached associates (Quayle 1988) or parasites (Bower et al. 1994). For example, the seaweed Sargas- sum muticum is now ubiquitous coast-wide, including Haida Gwaii (Figure 1). It was accidentally introduced into southern British Columbia in the early 20" century attached to Pacific oysters. Another example of collateral introduction in British Columbia is the parasitic copepod Mytilicola orientalis, likely introduced via Pacific Oyster stock and now widely infesting Native Littleneck Clams (Protothaca staminea) and Butter Clams (Saxidomus giganteus) in southern British Columbia (Bower et al. 1994). TL Vol. 118 THE CANADIAN FIELD-NATURALIST 78 6L6] YOITED €L6] UeMO) L661 0418 866] duINOg pur aidsay]iH 0007 ‘Te 12 UeOD LO6] OFlRH 0007 ‘Te 19 UROD 6L6] UOITE) P96I aAeng 6L61 UOIeED 100@ ‘Te 39 ueOlS C86l SugsIy C66] UOIIeD pue udyoD 9€6] Jaseay 6L61 VOID QOOZ JonARg pur uvO[S 000c [491d 6L6] WOE S90INOS uMouyuUN UOTNGLASIp JUdLIND {punos nseL— £96] AIAINS £66] & Ul Poyeoo] 919M OUOU — INOGIeH UspeN pue jJoO[U] Jossey| yur yoRo QOO'ST peonpo.nut (OAC) epeuRD suvddQ pure SalJOYysty — 796] punog |Jouusy pur Jo[Uy aesoprys ‘JOSSLIA] JJO “JO[UT JOSseA UT SoqTS ainjpNd ie. 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SUD pur Yst{ fo juouniedag eiuiojyeED 1007 ‘Te 39 UROTS 0007 ZIny pue soulH C66 UOYIeD pure uayog L007 ‘Te 19 UeoTS 0007 Ziny pur soury C66] UoITeD pur usyoD 1007 ‘[e 19 UPOTS C66] UOITeD pure uayod) C86] UIMIG pue ysneurary 7661 YSNeurary L007 ‘Te 39 UROTS 6L6] WORE) peoidsapim Ajayi] ‘ayquyrear JOA JOU JIB SPIOIAI HRMD epIey q—>—a_x«=sSemuqa=nasaouOnmnma*Tleon0oeESEEES=$ SS (panuluod) "| ATAVL 80 THE CANADIAN FIELD-NATURALIST Langara Island Parry Passage Cryplosula pallasiana Lininodrilus monothecus Cryptosula pallasiana Graham Rennell Sound Island Rose Masset Spit Delkatla Vol. 118 ° Pelonaia corrugata Trechus oblusus Limnodrilus nonothecus Kumdis Bay Yakoun Bay a Skidegate Inlet Moresby Island Cartwright Sound Tasu Sound Bowerbankia gracilis Sabia conica Ciona intestinalis Legend a Sargassum mulicum Mya arenaria Ampithoe vallida ® Other invertebrate sightings as noted FiGure |. Map of Haida Gwaii showing locations mentioned in the text and collection sites of introduced marine plant and invertebrate species, based on data from Sloan and Bartier (2000) and Sloan et al. (2001). Mariculture-associated Introductions In 1977, concern over introduced aquatic species issues stimulated establishment of the federal-prov- incial Fish Transplant Committee (recently renamed Introductions and Transfers Committee — ITC) under Bowerbankia gracilis _ Schizoporella unicornis ® Tubularia crocea mandates from the federal Fisheries Act and the Bri- tish Columbia Fisheries Act and British Columbia Wild- life Act (BC 1990). The ITC evaluates potential risks to the environment associated with introductions or transfers of either finfish or invertebrates (“shellfish’’) Hecate Strait Hotspring Island Dolomite "Burnaby" Narrows Harriet Harbour Houston Stewart Channel 2004 SLOAN AND BARTIER: INTRODUCED MARINE SPECIES 8] 4 row )\_¢ Alaskaj British Columbia Dixon Entrance K Hecate x Strait ‘ —__ Queen None Recorded Charlotte Sound 1 - 100 | 101 - 1,000 a > 10,000 ¥ *\ x \ FiGurE 2. Map of the northern British Columbia and southeast Alaska regions showing the density of Atlantic salmon obser- vations. All data are from Fisheries and Oceans Canada’s Atlantic Salmon Watch Program: http://www.pac.dfo-mpo.gc. ca/sci/aqua/ASWP_e.htm. [accessed May, 2004]. British Columbia data are from 1987 to 2002 and partitioned according to Pacific Fishery Management Areas, and the Alaska data are from 1990 to 2002 and partitioned according to the Alaska Department of Fish and Game Area Polygons. into British Columbia marine or freshwaters. Besides _ genetic material into native species. Further, there are the species themselves and their attached associates, concerns over potential ecological displacement of there is the risk of introducing diseases, parasites or native species. The ITC issues licences to introduce 82 THE CANADIAN FIELD-NATURALIST aquatic species into British Columbia or to transfer species between domestic water bodies. Shellfish mariculture remains a potential avenue of species introduction to Haida Gwaii. A Masset-based group has investigated the potential for local shellfish mariculture for coastal community economic develop- ment. Leased culture operations in Skidegate Inlet and Rennell Sound remain active for Pacific Oyster culture. These operations are unlikely a threat of introducing oysters as local waters are too cold for oyster breeding although they are suitable for growth of certified disease-free juveniles (spat) from culture in southern British Columbia. However, local waters may not be too cold for oysters’ parasites or other associated species. From 1997 to 2001, pilot raft culture sites were tested to grow certified disease-free Weathervane Scallop (Patinopecten sp.) hybrid (native x Japanese) spat (B. Mark, Masset, personal communication). Only the Rennell Sound site remains active for scallop cul- ture (R. Lozon, Queen Charlotte City, personal com- munication). The potential for species introduction is likely low, because no successful settlement of hybrid Weathervane Scallops has occurred in British Colum- bia in the last decade (Island Scallops Ltd., Qualicum Beach, personal communication). Ships’ Ballast Water Gauthier and Steel (1998) reported that Canada was receiving approximately 52 million tonnes of ballast water from foreign shipping annually with little pro- tective policy or regulation. Indeed, the major vector of introductions to Pacific North America has been from shipping (Ruiz et al. 2000). Concerning Pacific Canada, Levings (1999) mentioned the Canadian Bal- last Water Management Guidelines issued by Transport Canada in 2000 with an Annex (II) for the Pacific coast aimed at preventing introduction of non-indige- nous aquatic organisms (http://www.tc.gc.ca/Marine Safety/). These Guidelines will become Regulations under the Canada Shipping Act. The Vancouver Port Authority has had a mandatory ballast water program since 1997. It is based on the assumption that mid-ocean ballast water exchange, with water containing pelagic species not likely adapted to coastal conditions, decreases likelihood of introducing viable species into port waters (Levings et al. 2004). Port authorities are now finding, however, that such ex- change criteria are only partially effective (C. Levings, Fisheries and Oceans Canada (DFO), personal com- munication). Other British Columbia ports invoking ballast water management are Nanaimo and New West- minster. The north coast of British Columbia, with an active deep-water international port in Prince Rupert, however, has no ballast water program. Further, Lev- ings et al. (2004) express concern about “intracoas- tal” transport. For example, from Haida Gwaii there is on-going barge and self-dumping log barge traffic with Vol. 118 southern British Columbia and there has been relative- ly recent international vessel traffic for mine ore con- centrates from Tasu Sound (mine closed 1973) and Harriet Harbour (mine closed 1968). The ecological effects of introduced species such as invertebrates on the Pacific coast are poorly studied (Carlton 1992). It is sobering to reflect that, once in- troduced, marine species may be difficult to control and their ecosystem consequences may be damaging (Simberloff 2000; Grosholz 2002). An example is the European Green Crab, Carcinus maenas, introduced to the San Francisco Bay area in 1989. The Green Crab has since been recorded from Esperanza Inlet on the northwest coast of Vancouver Island (Jamieson et al. 2002). As active predators, Green Crabs could affect British Columbia intertidal fauna, as they have in California (Grosholz 2000). Times and attitudes have changed about introduc- tions of marine species. In the 1980s, DFO discussed introduction of the large, predatory Atlantic lobster (Homarus americanus) around Haida Gwati. Barber (1983) concluded that up to 7700 km? of Hecate Strait area south of Skidegate Inlet, as well as Masset Inlet, were suitable for lobster introduction. Now, such ini- tiatives are contrary to agency mandates for sustain- able, ecosystem-based management. Among introduced marine species, Atlantic Salmon (Salmo salar) currently has the highest ecosystem and socio-political profile in British Columbia (Gross 1998; Volpe et al. 2001). Important issues include escapement and persistence in river systems facilitating competition for spawning habitat with native salmo- nids. Further, the threat of continuing introductions helps animate the vigorous debate over expansion of salmon farming in British Columbia. Although Atlantic Salmon have been reported from Haida Gwaii coastal waters, there are no salmon farms in Haida Gwaii, and there are no records at this time of Atlantic salmon from within Haida Gwaii rivers and streams (V. Fradette, DFO, personal communication). This overview of a marine region’s introduced spe- cies was facilitated through marine species inventories from the historical literature of the Haida Gwaii region. We do not claim that it is complete, but it is a start. It will be important to continue documenting introduced species reports to track the progress of introductions that could affect local marine ecosystem well-being. Acknowledgments We thank C. D. Levings of Fisheries and Oceans Canada (DFO) for reviewing an early draft and G. E. Gillespie (DFO) for information on fishes. Literature Cited Austin, W. C. 1985. An annotated checklist of marine in- vertebrates in the cold temperate northeast Pacific. Khoya- tan Marine Laboratory, Cowichan Bay, British Columbia. 3 volumes. 682 pages. 2004 Barber, F. G. 1983. Lobster transplant to Masset Inlet. Can- adian Technical Report of Fisheries and Aquatic Sciences 1181. 6 pages. Bax, N., J. T. Carlton, A. Mathews-Amos, R. L. Haedrich, F. G. Howarth, J. E. Purcell, A. Rieser, and A. Gray. 2001. The control of biological invasions in the world’s oceans. Conservation Biology 15: 1234-1246. BC (British Columbia Ministry of Agriculture, Fisheries and Food). 1990. Federal and Provincial Transplant Reg- ulations. Aquaculture Factsheet Bulletin (Aquaculture and Commercial Fisheries Branch) 8-1. 3 pages. Bousfield, E. L., and P. M. Hoover. 1997. The amphipod superfamily Corophioidea on the Pacific coast of North America. Part V. Family Corophidae. Corophiinea, new subfamily. Systematics and distributional ecology. Amphi- pacifica 2(3): 67-139. Bower, S. M., S. E. McGladdery, and I. M. Price. 1994. Synopsis of infectious diseases and parasites of commer- cially exploited shellfish. Annual Review of Fish Diseases 4: 1-199. California Department of Fish and Game. 2002. A survey of non-indigenous aquatic species in the coastal and estuarine waters of California. Office of Spill Prevention and Response. [http://www.dfg.ca.gov/ospr/index.html]] Carl, G. C., and C. J. Guiguet. 1958. Alien animals in British Columbia. British Columbia Provincial Museum Handbook 14. 94 pages. Carlton, J. T. 1979. History, biogeography, and ecology of the introduced marine and estuarine invertebrates of the Pacific coast of North America. Unpublished Ph.D. thesis, University of California, Davis, California. 904 pages. Carlton, J. T. 1992. Introduced marine and estuarine mol- lusks of North America: an end-of-century perspective. Journal of Shellfish Research 11: 489-505. Chapman, J. W., T. W. Miller, and E. V. Coan. 2003. Live seafood species as recipes for invasion. Conservation Biology 17: 1386-1395. Coan, E. V., P. V. Scott, and F. R. Bernard. 2000. Bivalve seashells of western North America: marine bivalve mol- lusks from Arctic Alaska to Baja California. Santa Barbara Museum of Natural History Monographs 2. 764 pages. Cohen, A. N., and J. T. Carlton 1995. Nonindigenous aquatic species in a United States estuary: a case study of the bio- logical invasions of the San Francisco Bay and Delta. United States Fish and Wildlife Service, Washington, D.C. and National Sea Grant College Program, Connecticut Sea Grant, Groton. National Technical Information Service Report PB96-166-525. Springfield, Virginia. 246 pages. Cowan, I. M. 1973. Sabia conica (Schumacher) on the Pac- ific coast of North America. Veliger 16: 290. Druehl, L. D. 2000. Pacific seaweeds a guide to common seaweeds of the west coast. Harbour Publishing, Madeira Park, British Columbia. Erséus, C. 1982. Taxonomic revision of the marine genus Limnodriloides (Oligochaeta: Tubificidae). 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One species with two biologies: Atlantic salmon (Salmo salar) in the wild and in aquaculture. Can- adian Journal of Fisheries and Aquatic Sciences 55 (Sup- plement 1): 131-144. Harbo, R. M. 1997. Shells and shellfish of the Pacific North- west: a field guide. Harbour Publishing, Madeira Park, British Columbia. Hines, A. H., and G. M. Ruiz. 2000. Biological invasions of cold-water coastal ecosystems: ballast-mediated intro- ductions in Port Valdez/Prince William Sound, Alaska. Regional Citizens’ Advisory Council of Prince William Sound, Valdez, Alaska. 340 pages. Huntsman, A. G. 1912. Holosomatous ascidians from the coast of western Canada. Contributions to Canadian Biol- ogy 1906-1910:103-185. Jamieson, G. S., M. G. G. Foreman, J. Y. Cherniawsky, and C. D. Levings. 2002. European green crab (Carcinus maenas) dispersal: the Pacific experience. Pages 561-575 in Crabs in cold water regions: biology, economics and management. Alaska Sea Grant College Program AK- SG-02-01. : Kavanaugh, D. H. 1992. Carabid beetles (Insecta: Coleop- tera: Carabidae) of the Queen Charlotte Islands, British Col- umbia. Memoirs of the California Academy of Sciences 16: 113 pages. Kavanaugh, D. H., and T. L. Erwin. 1985. Trechus obtusus Erichson (Coleoptera: Carabidae), a European ground beetle, on the Pacific coast of North America: its distribu- tion, introduction, and spread. Pan-Pacific Entomologist 61: 170-179. Levings, C. D. 1999. Review of current practises to reduce the risk of introducing non-indigenous species into Pacific region via ballast water. Canadian Stock Assessment Secretariat Research Document 99/211. 13 pages. Levings, C. D., J. R. Cordell, S. Ong, and G. E. Piercey. 2004. The origin and identity of invertebrate organisms being transported to Canada’s Pacific coast by ballast water. Canadian Journal of Fisheries and Aquatic Sciences 61: 1-11. Levings, C. D., D. Kieser, G. S. Jamieson, and S. Dudas. 2002. Marine and estuarine alien species in the Strait of Georgia, British Columbia. Pages 111-131 in Alien in- vaders in Canada’s waters, wetlands, and forests. Edited by R. Claudia, P. Nantel and E. Muckle-Jeffs. Natural Resources Canada, Canadian Forest Service, Ottawa. McKinnell, S., A. J. Thompson, E. A. Black, B. L. Wing, C. M. Guthrie I, J. F. Koerner, and J. H. Helle. 1997. Atlantic salmon in the North Pacific. Aquaculture Res- earch 28: 145-157. Mecklenburg, C. W., T. A. Mecklenburg, and L. K. Thors- teinson. 2002. Fishes of Alaska. American Fisheries Society, Bethesda, Maryland. 1037 pages. Quayle, D. B. 1964. Distribution of introduced marine mol- lusca in British Columbia waters. Journal of the Fisheries Research Board of Canada 21: 1155-1181. 84 THE CANADIAN FIELD-NATURALIST Quayle, D. B. 1988. Pacific oyster culture in British Colum- bia. Canadian Bulletin of Fisheries and Aquatic Sciences 218. 241 pages. Ruiz, G. M., P. W. Fofonoff, J. T. Carlton, M. J. Won- ham, and A. H. Hines. 2000. Invasion of coastal marine communities in North America: apparent patterns, pro- cesses, and biases. Annual Review of Ecology and System- atics 31: 481-531. Simberloff, D. 2000. No reserve is an island: marine reserves and nonindigenous species. Bulletin of Marine Science 66: 567-580. Sloan, N. A., and P. M. Bartier. 2000. Living marine legacy of Gwaii Haanas. I. Marine plant baseline to 1999 and plant-related management issues. Parks Canada Technical Reports in Ecosystem Science 27. 104 pages. Sloan, N. A., P. M. Bartier, and W. C. Austin. 2001. Living marine legacy of Gwaii Haanas. II. Marine invertebrate baseline to 2000 and invertebrate-related management 1s- Vol. 118 sues. Parks Canada Technical Reports in Ecosystem Sci- ence 35. 331 pages. Volpe, J. P., B. R. Anholt, and B. W. Glickman. 2001. Competition among juvenile Atlantic salmon (Salmo salar) and steelhead (Oncorhynchus mykiss): relevance to inva- sion potential in British Columbia. Canadian Journal of Fisheries and Aquatic Sciences 58: 197-207. Welander, A. D. 1940. Notes on the dissemination of shad, Alosa sapidissima (Wilson), along the Pacific coast of North America. Copeia 1940: 221-223. Workman, G. D., J. Fargo, B. Beall, K. L. Yamanaka, and V. Haist. 1996. R/V W.E. Ricker assemblage survey of Hecate Strait, May 23-June 9, 1995. Canadian Data Report of Fisheries and Aquatic Sciences 974. 94 pages. Received 27 December 2002 Accepted 11 June 2004 Ruby-throated Hummingbird, Archilochus colubris, Entanglements in Burdock, Arctium spp., at Delta Marsh, Manitoba HEATHER L. HINAM!, SPENCER G. SEALY, AND Topp J. UNDERWOOD Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 'Present Address: Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada Corresponding author sgsealy @cc.umanitoba.ca Hinam, Heather L., Spencer G. Sealy, and Todd J. Underwood. 2004. Ruby-throated Hummingbird, Archilochus colubris, entanglements in burdock, Arctium spp., at Delta Marsh, Manitoba. Canadian Field-Naturalist 118(1): 85-89. Exotic burdock (Arctium spp.) pose a risk of mortality for small native birds, such as the Ruby-throated Hummingbird (Archi- lochus colubris), which may become entangled in its burrs. At Delta Marsh, Manitoba, we found 11 hummingbirds and five individuals of four species of songbirds entangled on burdock in the dune-ridge forest over a 20-year period. Entangled birds were mostly migrants. Most hummingbirds caught were juvenile males, whereas the few songbirds were mostly adult males. We suspect that hummingbird entanglements resulted from an attraction to the purple flowers of burdock, but aggressive interactions with conspecifics and other factors may have been involved. Birds may be at a higher risk of entanglement at important migratory stopover sites, such as Delta Marsh, where both burdock and large numbers of birds are concentrated in a small area. Key Words: exotic plants, burdock, Arctium spp., fatal entanglement, Ruby-throated Hummingbird, Archilochus colubris, pas- serines, mortality, Delta Marsh, Manitoba. Exotic species often exert negative effects on the survival of native flora and fauna (Atkinson 1989). This is exemplified by the relationship between burdock (Arctium spp.) and small birds and bats in North Amer- ica (McNicholl 1988, 1994). Four species of burdock were introduced from Eurasia into North America in the early 1600s (Gross et al. 1980; Harms 2001). All four species of burdock develop seed heads covered with burrs that stick to passing animals, thus dis- persing the seeds (McNicholl 1988). These burrs are hazards for small birds and bats because individuals may become entangled in the hooked bracts and die (McNicholl 1988, 1994). The effect of such entangle- ments on bird populations may be magnified when the plants are concentrated in small areas where birds nest or forage during migration. For many songbirds, the dune-ridge forest that sepa- rates the south shore of Lake Manitoba from Delta Marsh (50°11 N, 98°19’ W), Manitoba, is an important breeding area (Goossen and Sealy 1982; MacKenzie et al. 1982) and migratory stopover site (den Haan 1996). Three species of burdock (A. minus, A. lappa, A. tomentosum) have been documented in the area, but Common Burdock (A. minus) is the most abundant, occurring along the edge of the marsh and in the ridge forest, as well as in ditches and at other disturbed sites (Shay 1999). The composition of the understory of the ridge forest has changed over the last 30 years and the amount of burdock apparently has increased (Kenkel and Graham 1994; S.G. Sealy, personal obser- vations). With this apparent increase, Delta Marsh has the potential to become a trap line for small birds such as the Ruby-throated Hummingbird (Archilochus colu- bris). Indeed, this species has been one of the most frequently discovered entangled in burdock (McNicholl 1988, 1994; Raloff 1998; Nealen and Nealen 2000). Here we document additional cases of mortality of Ruby-throated Hummingbirds and songbirds due to entrapment on the seed heads of burdock at Delta Marsh, Manitoba. Entanglements Sealy and co-workers have studied songbird popu- lations in the ridge forest at Delta Marsh since 1973, but it was not until 1983 that the first bird was found entangled on burdock. The decomposed remains of a Ruby-throated Hummingbird were discovered that spring, but death had occurred the previous year. Since then, 10 more hummingbirds have been found, all but one in August 1985 (Table 1). One of these was alive and released, leaving behind several feathers stuck to the burrs (Table 1). From 1977 to 1979, 1981 and 1983, hummingbirds were banded in conjunction with gen- eral songbird banding in the ridge forest from mid-May through the end of August. The timing of the entan- glements in 1985 coincided with the fall migration of hummingbirds through the ridge forest in August (Figurel). However, except for the live hummingbird, the dates of entanglement only approximate the time of death because the rates of decomposition and mummi- fication are not known. Thus, some hummingbirds could have been killed earlier in August. Despite sporadic searching since 1985, only one other entangled hummingbird was found, in 2002 (Table 1, Figure 2), but since 1994, five individuals of four other species have been found entangled fatally (Table 2). Another record consisted of only a few feathers from an unknown passerine species, possibly 85 86 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Records of Ruby-throated Hummingbirds entangled in burdock at Delta Marsh, Manitoba. Date Age/Sex! Condition Specimen No.” 19 May 1983 Unk/unk Decomposed: overwintered, wings, ventral feathers attached 4986 15 August 1985 Unk/unk Released, feathers stuck on burdock 29817 18 August 1985 HY/female Recently caught (intact) 4982 18 August 1985 HY/male Recently caught (intact) 4983 18 August 1985 Unk/unk Partially decomposed: attached by ventral and wing feathers 4985 19 August 1985 HY/female Recently caught 4981 19 August 1985 AHY/male Partly decomposed 4984 19 August 1985 AHY/male, — Facing each other with burrs in between HY/male (fresh and mostly intact) 4987 A,B 21 August 1985 HY/male Stuck across burr (fresh, mostly intact) 4980 14 September 2002 HY/female Decomposed, sternum visible 499] "Unk = unknown, HY = hatch year, AHY = after hatch year. Bird specimens deposited in the vertebrate collections of the Manitoba Museum, Winnipeg, Manitoba, Canada. >Feathers deposited in the University of Manitoba Zoology Museum, Winnipeg, Manitoba, Canada. TABLE 2. Records of songbird species entangled in burdock at Delta Marsh, Manitoba. Species Date Age/Sex! Condition Specimen No.” Unknown passerine 15 August 1985 Unk/unk Feathers stuck on burr 29823 Golden-crowned Kinglet 30 October 1994 Unk/male Wings and breast attached (fairly intact) 4976 Ruby-crowned Kinglet 2 November 1996 AHY/male Left wing caught (fresh, intact) 4977 Common Yellowthroat May 1999 AHY/female | Decomposed: overwintered, caught by one leg 4978 13 May 2000 AHY/male Decomposed: overwintered, caught by breast, both wings 4979 Yellow-rumped Warbler? —_10 October 2002 HY/unk Fresh, caught by right foot, feathers of right flank, left wing 4988 'Unk = unknown, HY = hatch year, AHY = after hatch year. *Bird specimens deposited in the vertebrate collections of the Manitoba Museum, Winnipeg, Manitoba, Canada. Feathers deposited in the University of Manitoba Zoology Museum, Winnipeg, Manitoba, Canada. ‘Banded 14 September 2002. a sparrow (C. Dove, personal communication). This bird may have become entangled and freed itself; alter- natively it may have been preyed upon or scavenged. Two Common Yellowthroats (Geothlypis trichas), a species that nests at Delta Marsh, were found in spring after apparently becoming entangled the previous year. One Golden-crowned Kinglet (Regulus satrapa) and one Ruby-crowned Kinglet (R. calendula), both boreal forest breeders and late migrants, were found freshly dead in the fall. In October 2002, a freshly dead Yellow-rumped Warbler (Dendroica coronata) was dis- covered less than one month after it had been banded in the ridge forest. All four of these species have been found trapped on burdock elsewhere (McNicholl 1988, 1994). The sex and age of birds were identified where pos- sible by characteristics of the plumage, feathers and bills (Pyle 1997). Of the hummingbirds entangled, five were males and three were females, whereas three songbirds were males and one was a female (Tables 1, 2). Six hummingbirds were juveniles and two were adults (Table 1). By contrast, three songbirds were adults and only one was a juvenile (Table 2). Discussion Most birds entangled in burdock at Delta Marsh likely were migrants. Although Ruby-throated Hum- mingbirds nest at Delta Marsh (Underwood and den Haan 2000) in small numbers, all entanglements were discovered during their fall migration period through Delta Marsh (Figure 1). Among the other species killed on burdock, only the Common Yellowthroat breeds commonly at Delta Marsh (Underwood and den Haan 2000). The preponderance of migrants in our sample of burdock mortalities is interesting. Several small songbirds, such as the Least Flycather (Empidonax minimus) and Yellow Warbler (Dendroica petechia), nest at high densities in the ridge forest (Goossen and Sealy 1982; Briskie and Sealy 1989; S. G. Sealy, un- published data), but neither species has been recorded entangled in burdock there, although the Least Fly- catcher is a documented victim elsewhere (Underwood and Underwood 2001). Individuals of both species have been observed perched on stems of burdock (Sealy and Underwood, personal observations) and Sealy ob- served two male Yellow Warblers, one chasing the other 2004 14 12 2) x2) = 2 = 10 = & & 8 =) a Gen CO: .6 ten ® £ 5s 4 a 2 0 1 4 if; 10 HINAM, SEALY, AND UNDERWOOD: HUMMINGBIRD ENTANGLEMENTS IN BURDOCK 87 13 16 19 22 August FicureE 1. Number of Ruby-throated Hummingbirds found caught on burdock at Delta Marsh, August 1985 (1 = 9), in relation to the number of hummingbirds banded per day throughout 1977 (n = 7), 1978 (n = 37) and 1979 (n= 4), 1981 (n = 23) and 1982 (n = 42). that hit seed heads during the chase. Although they became stuck, they extricated themselves within sec- onds of impact. This observation notwithstanding, accidental strikes on burdock seed heads likely account for few mortalities. This may explain the infrequency of entanglements among the resident songbirds, which have not been observed foraging on burdock flowers or arthropods attracted to them. It is commonly assumed that birds become entangled while foraging (McNicholl 1988). Although humming- birds feed mainly while hovering (Robinson et al. 1996), their small size may be the reason they frequent- ly become entangled. As noted by Nealen and Nealen (2000), Ruby-throated Hummingbirds prefer reddish flowers (Robinson et al. 1996; but see Miller and Miller 1971), although they do not specialize on a particular species (Bertin 1982). Because burdock flowers are with- in the red spectrum (Gross et al. 1980), hummingbirds may be attracted to them in the ridge forest where there are few of the hummingbird’s putatively preferred spe- cies (Robinson et al. 1996; Shay 1999). Birds may also become entangled while foraging for insects that inhabit the flowers or the seed heads. Insects may comprise up to 60% of the diet of Ruby- throated Hummingbirds and individuals have been known to glean larval lepidoptera and other insects from the surface of plants (Robinson et al. 1996). Over 20 species of insects from three Orders, Lepidoptera, Coleoptera and Hymenoptera, have been found on burdock seed heads (Mulligan and Kevan 1973; Gross et al. 1980). The kinglets and warblers may have been attempting to take insects from the burrs when they became entangled (see Needham 1909). Entanglements may also result from social inter- actions between individuals. The deaths of two male hummingbirds on the same burr cluster (Table 1, MM 4987A and B) may have resulted when one individ- ual attacked the other at the flower cluster and they both became entangled as the interaction ensued. Sealy watched a male Ruby-throated Hummingbird attack another male as it hovered by a seed head. The hov- ering bird became entangled by one leg, but extri- cated itself within seconds. The attacker had already flown away. Wind may also cause entanglements, buf- feting birds against seed heads as they move through the area (McNicholl 1988). Thus, identifying the factor or combination of factors that promote individual bird entanglements is difficult. 88 THE CANADIAN FIELD-NATURALIST FIGURE 2. Ruby-throated Hummingbird entangled in burdock found 14 September 2002, Delta Marsh (photograph by T. J. Underwood). The large number of hummingbird entanglements in burdock over a short period in August 1985 seems unusual. Burdock apparently has increased in abun- dance on the ridge forest (Kenkel and Graham 1994) since our studies began at Delta in 1973. This suggests that the frequency of entanglement should have in- creased or at least remained constant. However, search effort has varied widely since 1985 and migration rates and weather patterns affecting bird movements through the ridge forest vary from year to year. Hence, it is difficult to assess from occasional records the overall effect of burdock at Delta Marsh on migrating birds. Nevertheless, the growing number of reports of burdock-related deaths in birds (e.g., McNicholl 1994; Raloff 1998; Underwood and Underwood 2001) sug- gests that this type of mortality may be more important than originally believed, particularly at places like Delta Marsh, and King’s Park in Winnipeg (Underwood and Underwood 2001), where burdock and large numbers of migrating birds are concentrated. Further study of the interaction between birds and burdock and the pos- sibly fatal consequences for birds should result in a better understanding of the effects of this exotic plant species on bird populations. Acknowledgments We thank Janis Klapecki for cataloguing bird speci- mens in The Manitoba Museum and Carla Dove, Vol. 118 Smithsonian Institution, for attempting to identify feathers. Some specimens were collected by G. C. Biermann, J. V. Briskie, H. E. den Hann, K. A. Hobson, E. Kiviat, A. Mcllraith, and R. M. Underwood. H. E. den Haan provided details on the banding of the Yel- low-rumped Warbler from the files of the Delta Marsh Bird Observatory and assisted with age/sex identifica- tion of specimens. We thank the reviews, A. J. Erskine and M. K. McNicholl, for their comments on the man- uscript. Accommodation and facilities were provided by the Delta Marsh Field Station (University of Mani- toba). Officers of the Portage Country Club permitted us to conduct some of the field work on their property. Funding was provided by the Natural Sciences and Engineering Research Council of Canada. Literature Cited Atkinson, I. 1989. Introduced animals and extinctions. Pages 54-75 in Conservation for the twenty-first century. Edited by D. Western and M. C. Pearl. Oxford University Press, Oxford. Bertin, R. I. 1982. The Ruby-throated Hummingbird and its major food plants: Ranges, flowering phenology, and migration. Canadian Journal of Zoology 60: 210-219. Briskie, J. V., and S. G. Sealy. 1989. Determination of clutch size in the Least Flycatcher. Auk 106: 269-278. den Haan, H. 1996. Delta Marsh Bird Observatory interim report: 1996. University of Manitoba Field Station (Delta Marsh) Annual Report 31: 81-84. Goossen, J. P., and S. G. Sealy. 1982. Production of young in a dense nesting population of Yellow Warblers, Dendroica petechia, in Manitoba. Canadian Field-Naturalist 96: 189-199. Gross, R. S., P. A. Werner, and W. R. Hawthorn. 1980. The biology of Canadian weeds 38. Arctium minus (Hill) Bernh and A. lappa L. Canadian Journal of Plant Science 60: 621- 634. Harms, V. L. 2001. Burdocks in Saskatchewan. Blue Jay 59: 92-98. Kenkel, N. C., and K. Graham. 1994. Population biology and the control of Common Burdock (Arctium minus (Hill) Bernh.) at the University Field Station, Delta Marsh. University of Manitoba Field Station (Delta Marsh) An- nual Report 29: 112-116. MacKenzie, D. I., S. G. Sealy, and G. D. Sutherland. 1982. Nest-site characteristics of the avian community in the dune-ridge forest, Delta Marsh, Manitoba: A mullti- variate analysis. Canadian Journal of Zoology 60: 2212- D223% MeNicholl. M. K. 1988. Bats and birds stuck on burdock. Prairie Naturalist 20: 157-160. MeNicholl, M. K. 1994. Additional records of birds caught on burdock. Ontario Birds 12: 117-119. Miller, R. S., and R. E. Miller. 1971. Feeding activity and color preference of Ruby-throated Hummingbirds. Condor 73: 309-313. Mulligan, G. A., and P. G. Kevan. 1973. Color, brightness and other floral characteristics attracting insects to the blos- soms of some Canadian weeds. Canadian Journal of Botany 51: 1939-1952. Nealen, H. J., and P. M. Nealen. 2000. Ruby-throated Hum- mingbird death by Common Burdock (Arctium minus). Wilson Bulletin 112: 421-422. 2004 Needham, J. G. 1909. Kinglets captured by burdocks. Bird- Lore 11: 261-262. Pyle, P. 1997. Identification guide to North American birds. Part I. Slate Creek Press, Bolinas, California. Raloff, J. 1998. Botanical *Velco’ entraps hummingbirds. Science News 154:244. Robinson, T. R., R. R. Sargent, and M. B. Sargent. 1996. Ruby-throated Hummingbird, Archilochus colubris. In The Birds of North America, no. 204. Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadel- phia, and The American Ornithologists’ Union, Washing- ton, D.C. Shay, J. M. 1999. Annotated vascular plant species list for the Delta Marsh, Manitoba and surrounding area. Univer- HINAM, SEALY, AND UNDERWOOD: HUMMINGBIRD ENTANGLEMENTS IN BURDOCK 89 sity of Manitoba Field Station (Delta Marsh), Occasional Publication 2. Underwood, T. J., and H. E. den Haan. 2000. Checklist of the birds of Delta Marsh, third edition. [Delta Marsh Bird Observatory, Portage la Prairie, Manitoba]. Underwood, T. J., and R. M. Underwood. 2001. Observa- tions on burdock-killed birds in King’s Park, Winnipeg, MB. Blue Jay 59: 64-69. Received 7 March 2003 Accepted 23 April 2004 Multiple Mating Results in Multiple Paternity in Richardson’s Ground Squirrels, Spermophilus richardsonii James F. Hare!, GLENDA Topp!, and WENDY A. UNTEREINER? ' Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada * Botany Department, Brandon University, Brandon, Manitoba R7A 6A9 Canada Hare, James F., Glenda Todd, and Wendy A. Untereiner. 2004. Multiple mating results in multiple paternity in Richardson’s Ground Squirrels, Spermophilus richardsonii. Canadian Field-Naturalist 118(1): 90-94. Microsatellite DNA primers developed from Columbian Ground Squirrels (Spermophilus columbianus) were used to establish paternity in a Manitoba population of Richardson’s Ground Squirrels (Spermophilus richardsonii). Primers resolving variation at six microsatellite loci allowed ascription of paternity to 32 of 85 offspring born among litters of 15 breeding females sampled. While the failure to unambiguously document paternity for all juveniles precludes the use of these data to address questions of sperm competition and male mating success, the results do provide direct evidence that multiple mating by female Richardson’s Ground Squirrels results in multiple paternity within litters. Key Words: Richardson’s Ground Squirrel, Spermophilus richardsonii, mating, microsatellite DNA, multiple paternity, Manitoba. Differences in gamete size, and hence the differen- tial investment made in gametes by males and female ultimately define the sexes (Parker et al. 1972) and have promoted the evolution of disparate reproductive strate- gies. Males commonly enhance their fitness by obtain- ing copulations with more than one female (Bateman 1948), maximizing the propagation of like copies of their genes by siring as many offspring as possible. Conversely, the enhancement of female fitness is typi- cally achieved through choosiness, whereby females mate only with males providing the greatest access to resources (Verner and Willson 1966; Thornhill 1976; Stanford 1995) and/or who are the most fit and con- tribute good genes to the female’s progeny (Zahavi 1975; Weatherhead and Robertson 1979; Hamilton and Zuk 1982). At a proximate level, anisogamy also promotes sex-differential mating behaviour. The rela- tively few, large, nutrient-rich eggs produced by females necessitate only a single male ejaculate containing vast numbers of small, nutrient-poor spermatozoans to achieve fertilization, and thus female reproductive suc- cess may not increase through multiple mating. Despite these fundamental predictions following from anisogamy, multiple mating by females is com- mon in animals and can prove beneficial to females for various reasons. Obtaining multiple male mates may ensure fertilization (Hoogland 1998), enhance access to resources (Gray 1997), promote parental care on the part of males (Davies et al. 1996), confuse males as to paternity and thus reduce the probability of infanti- cide (Hrdy 1977), increase female fitness by ensuring genetic compatibility between the female’s eggs and at least some of the sperm (Zeh and Zeh 1996) or in- cite sperm competition (Gomendio and Roldan 1993), whereby the fitness of the female’s offspring is en- hanced by virtue of the good genes contributed by the victorious spermatozoans (Evans and Magurran 2000). Where multiple mating occurs, it is critical to un- derstand its implications to both males and females. Advances in molecular biology have revolutionized the study of reproductive behaviour, and revealed that observations of male/female association and mating behaviour may not be representative of parentage in free-living animal populations (Quinn et al. 1987). Methods for determining parentage (see Fleischer 1996 for a review) have also become more powerful and ac- cessible, thereby providing researchers with an oppor- tunity to obtain comparative data which address both the proximate and ultimate questions pertaining to re- productive behaviour. Such a comparative picture has begun to emerge for the ground-dwelling squirrels (Spermophilus spp.), where multiple mating commonly results in multiple paternity within litters, and the first male to mate with a given female often sires a disproportionate number of offspring within a litter. Hanken and Sherman (1981) reported that 78% of 27 Belding’s Ground Squirrel (S. beldingi) litters for which paternity could be deter- mined using polymorphic blood proteins were multiply sired by between two and three males. Similarly, 50% of eight Thirteen-lined Ground Squirrel (S. tridecem- lineatus) litters were sired multiply, with the first male to mate siring 75% of offspring in those litters (Foltz and Schwagmeyer 1989). Boellstorff et al. (1994) de- tected multiple paternity in 88.9% of nine litters of California Ground Squirrels (S. beecheyi) and Murie (1995) reported multiple paternity in 15.8% of 165 Columbian Ground Squirrel (S. columbianus) litters. Murie (1995), however, predicted that his data likely underestimated the incidence of multiple paternity given that only five polymorphic loci were used, and only one of those had more than two alleles. Indeed, using a more comprehensive and variable series of microsatellite primers (Stevens et al. 1997), multiple 90 2004 paternity was detected in 64% of litters born to fe- male Columbian Ground Squirrels that had mated mul- tiply (Stevens, Strobeck and Murie unpublished data). Of those litters, 77% were sired by two males, 21% by three males, and 2% by four males. Further, there was a distinct first-male advantage, with 65% of the offspring sampled from among 77 litters sired by the first male to mate, and a progressively declining pro- portion of paternity success for subsequently mating males (Stevens, Strobeck, and Murie unpublished data). Despite the widespread occurrence of multiple pater- nity and a first-male mating advantage, deviations from that pattern are apparent among ground squirrels. Using both protein isozymes and DNA fingerprinting (see Burke 1989), Lacey et al. (1997) revealed that multiple paternity among Arctic Ground Squirrels (S. parryi) was relatively rare; it occurred in only | (9%) of 11 litters examined and the first male to mate sired the majority (90%) of the offspring in that litter. A departure from the first-male mating advantage has been detected in Idaho Ground Squirrels (S. brun- neus), Where Sherman (1989) reported a last-male mating advantage (a minimum of 66 — 100% of pups sired by the last/longest attending male) in the five of seven litters (71.4%) where multiple paternity was evi- dent. Additional data for other Spermophilus species will prove useful in determining whether these species are unique in their apparent departures from the gen- eral pattern, and thus how these expressions of multiple mating and sperm precedence impact the behaviour and fitness of males and females. Richardson’s Ground Squirrels (S$. richardsonii) are locally abundant throughout grazed areas of the Great Plains (Michener and Koeppl 1985). Females mate multiply (Michener and McLean 1996), and al- though this may result in multiple paternity within lit- ters (van Staaden et al. 1994, Michener personal com- munication), direct evidence of multiple paternity is lacking. Such data would contribute to our understand- ing of reproductive behaviour, and open new avenues for research into the contributions of paternal kinship to social interactions (e.g., Holmes and Sherman 1982) and patterns of dispersion among relatives in nature. We applied primers developed by Stevens et al. (1997) to prospective parents and offspring from litters of 15 breeding females in a Manitoba population of Rich- ardson’s Ground Squirrels to test whether multiple mating results in multiple paternity within litters. Methods Study site, research subjects, and sample collection Research was conducted between 8 April and 26 July 2002 on a 1.5-ha section of a larger 5-ha cattle pasture near Westbourne, Manitoba (50°10.190'N, 98°38.103'W). Richardson’s Ground Squirrels on that site were live-trapped in National or Tomahawk traps baited with peanut butter and were marked for per- manent identification with numbered metal ear tags HARE, TODD, AND UNTEREINER: MULTIPLE MATING Q| (Monel #1 fish fingerling tags). Each squirrel was also given a distinctive mark on its dorsal pelage with black hair dye (Clairol Hydrience 52 Black Pear]). Trapping was conducted during the mating season (12 through 25 April 2002) to determine mating dates of females and to monitor the reproductive condition of all squirrels. After mating had ended, trapping con- tinued to assess gestational progress in females, and a 2-mm tissue punch was taken from one pinna of each adult (10 males, 18 females). Samples were placed in 1.5-ml microcentrifuge tubes in 95% ethanol, and stored at 4 — 7°C prior to DNA extraction and subse- quent molecular analysis. Fifteen females for which mating and parturition dates were known, and for whom ear punches had been obtained, and nest-burrow en- trances located were chosen to provide litters for pater- nity analysis. All juveniles in those litters were trapped within three days of their first emergence from their natal burrow, marked with ear tags and dye marks, and tissue sampled as described for adults. Animal research was conducted in accordance with the guidelines for the use of animals in research set forth by the Canadian Council on Animal Care. Molecular analysis of paternity Individual ear punches were placed in 1.5-mL micro- centrifuge tubes, centrifuged in a speed-vac for 5 min to remove the excess ethanol and digested at 55°C for 4 — 6 hrs in 70 uL of a sterile cell lysis solution (10 mM Tris-HCL, 100 mM EDTA, 2% SDS, pH 8.0) containing 3 — 4 uL Proteinase K (10 mg/mL). Following the addition of 4 uL of RNase A, the tubes were mixed by inversion and incubated at 37°C for 30 min. Proteins were precipitated by adding 23.5 uL of 7.5 M ammonium acetate to each tube of cell lysate and placing the samples on ice for 30 min. Tubes were centrifuged at 14000 rpm for 3 min and the superna- tant was transferred to a new 1.5-mL tube containing 70 wL isopropanol. Samples were mixed by inversion, stored at -20°C for several hours and centrifuged at 14000 rpm for 2 min. Following the removal of the supernatant, the pelleted DNA was washed in 75 UL of ice-cold 75% ethanol and centrifuged at 14000 rpm for 1 min. The ethanol was removed with a pipette and the tubes containing the DNA were placed in speed-vac for 5 — 10 min. The dried pellets were re- hydrated in 40 uL of sterile TLA buffer (10 mM Tris- HCL, 0.1 mM EDTA, pH 8.0) and stored at -20°C. Amplifications were performed on a GeneAmp® PCR System 9700 (Applied Biosystems, Foster City, CA) in a 25-uL volume containing ~ 200 ng DNA, 0.8 mM dNTP, PCR buffer (0.1 M Tris-HCI pH 8.3, 0.5 M KCl, 15 mM MgCl, 0.005 g gelatin/10 mL buffer), 0.5 units of Taq polymerase, and 0.2 UM con- centrations of each of seven primers (GS3, GS12, GS14, GS20, GS22, GS25, GS26) described by Stevens et al. (1997). The cycling conditions were 94°C for 2 min followed by two cycles of 94°C for 30 s, 58°C for 20 s, 72°C for 5 s, and 33 cycles of 94°C for 15 s, 54°C for 92 THE CANADIAN FIELD-NATURALIST 20 s and 72°C for 5 s followed by a final extension at 72°C for 30 s. PCR products were diluted (1:1) with sterile water and | uL of each diluted product was resolved by polyacrylamide gel electrophoresis on an ABI 373 Automated Sequencer (Applied Biosys- tems). Microsatellites were visualized and sized using GeneScan® 3.1 and Genotyper® 2.1 software (Applied Biosystems). Results Of the seven primers employed, one (GS22) result- ed in multiple banding patterns and was excluded from subsequent analysis. The remaining six primers resolved meaningful variation among individuals and were used in establishing familial relationships among the individuals sampled. Based upon the observed fre- quencies of the three to six alleles evident at those six loci (Table 1), the probability that two randomly drawn unrelated individuals will be identical at all loci is | in 9577 (from equation in Patekau and Strobeck 1994), Further, the probability that an unrelated male will be excluded as sire given knowledge of the mother’s geno- type is 0.91 (from equation in Chakravarti and Li 1983). All calculations assume that the loci examined are not linked, and provide highly conservative estimates of exclusion given that they are based on pooled data from dams, sires, and their offspring. Paternity was definitively ascribed to at least one juvenile within litters of all 15 dams included in the study. In total, however, unique identification of the male sire was achieved for only 32 of the 85 juveniles that emerged in those litters. Based upon those iden- tifications, multiple sires were detected in 12 of the 15 litters examined (80%), with a minimum of two sires in 11 and a minimum of three sires in one of those 12 litters (2.1 + 0.1 sires/litter; mean + SE). Discussion The microsatellite primers developed by Stevens et al. (1997) proved effective in resolving multiple pater- nity within litters of Richardson’s Ground Squirrels. Indeed, with multiple sires in 80% of the 15 litters Vol. 118 examined, multiple paternity is at least as common in Richardson’s Ground Squirrels as it is in other Spermo- philus species with the exception of California Ground Squirrels. Definitive genetic evidence of multiple paternity in the present study suggests that the use of the primers described by Stevens et al. (1997) in future studies of the reproductive and social behaviour of Richardson’s Ground Squirrels would prove quite fruitful. In concert with data on mating behaviour of the sort presented in Michener and McLean (1996), such studies could address the outcome of sperm competition and its rela- tion to mating order (Sherman 1989), the role of aggres- sion and male territoriality in affecting male mating success (Schulte-Hostedde and Millar 2002), and the factors underlying mate choice such as the effects of symmetry (Penton-Voak et al. 2001), body size (Gwynne 1981), paternal effort (Huber et al. 2002), and genetic relatedness (Bateson 1983; Chapman and Crespi 1998). The use of these microsatellite loci to assess relatedness among individuals (Queller et al. 1993) would also allow insight into the role paternal kinship plays in influencing social behaviour (Widdig et al. 2002). Additional markers, however, would be necessary to comprehensively document the paternity of each and every individual. Microsatellite primers that were not employed in the present study, such as GS17 and GS34 (Stevens et al. 1997) or the 13 Idaho Ground Squirrel microsatellite primers described by May et al. (1997), would likely prove useful in this regard. The alleles identified in our Manitoba population (Table 1) differ from those described for the Richard- son’s Ground Squirrels examined by Stevens et al. (1997) in evaluating the efficacy of their primers on other Sciurids (see Table 2 in Stevens et al. 1997). Given that geographic variation, the primers employed here should also prove useful for studies of dispersal (Chapuisat et al. 1997), interpopulation differences related to conservation efforts (Patekau and Strobeck 1994), and perhaps even in refining the taxonomy of the ground-dwelling squirrels (Gill and Yensen 1992). TABLE 1: Characteristics of microsatellite loci (from Stevens et al. 1997) applied in paternity analysis of Richardson’s Ground Squirrels. Results are based upon allele frequencies from 10 adult male, 15 adult female, and 85 juvenile squirrels. Expected heterozygosity and probability of identity were calculated according to formulae presented in Patekau and Strobeck (1994), and probability of exclusion was calculated following the methods of Chakravarti and Li (1983). Locus Alleles GS3 2215229523 18233 GS12 147, 149, 152, 154 GS14 242, 244, 246 GS20 220; 222.237 GS25 138, 142, 144, 146, 148, 154 GS26 109, 111, 113, 115 Expected Probability Probability heterozygosity of identity of exclusion 0.630 0.178 0.347 0.672 0.128 0.416 0.549 0.298 0.253 0.105 0.803 0.053 0.769 0.057 0.557 0.480 0.336 0.232 2004. HARE, TODD, AND UNTEREINER: MULTIPLE MATING 93 Acknowledgments Fleischer, R. C. 1996. Application of molecular methods to We are grateful to Curtis Strobeck and Lindsey the assessment of genetic mating systems in vertebrates. Carmichael of the University of Alberta for making Pages 133-161 in Molecular Zoology: Advances, Strategies, Fa and Protocols. Edited by J.D. Ferraris and S. R. Palumbi. the primers available to us and providing guidance on the use of the Genotyper® software, respectively. Gail Michener facilitated our work by allowing Mag- dalena Kujath to obtain samples from her study pop- ulation that proved useful in piloting the application of this technique. Our understanding of microsatellite methodology was enhanced through conversations with Nate Lovejoy, Ross McGowan, and Jan Murie, while Andrea Angus provided sound advice and assis- tance in the lab. We thank Jennifer Cassin, Kristin Melnyk, Melanie Sharpe, and David Wilson for capable field assistance and the Johnson family of Westbourne, Manitoba, for allowing us access to the squirrels on their land. Funding for this work was provided in part by a stipend from Human Resources and Development Canada in the form of a Summer Career Placement grant for Kristin Melnyk, and by the Natural Sciences and Engineering Research Council of Canada in the form of a University Summer Research Award to Glenda Todd, and operating grants to James Hare and Wendy Untereiner. Research infrastructure was also provided by a Canada Foundation for Innovation Res- earch Development Fund award to Wendy Untereiner. This is paper number 301 of the Delta Marsh Field Station. Literature Cited Bateman, A. J. 1948. Intra-sexual selection in Drosophila. Heredity 2: 349 — 368. Bateson, P. 1983. Optimal outbreeding. Pages 257 — 277 in Mate choice. Edited by P. Bateson. Cambridge University Press, Cambridge. Boellstorff, D. E., D. H. Owings, M. C. T. Penedo, and M. J. Hersek. 1994. Reproductive behaviour and multiple paternity of California ground squirrels. Animal Behaviour 47: 1057-1064. Burke, T. 1989. DNA fingerprinting and other methods for the study of mating success. Trends in Ecology and Evo- lution 4: 139-144. Chakravarti, A., and C. C. Li. 1983. The effect of linkage on paternity calculations. Pages 411-422 in Inclusion prob- abilities in parentage testing. Edited by R. H. Walker. American Association of Blood Banks, Arlington, Virginia. Chapman, T. W., and B. 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Received 29 May 2003 Accepted 15 June 2004 Demographic Patterns and Limitation of Grey Wolves, Canis lupus, in and Near Pukaskwa National Park, Ontario S. ANNE FORSHNER!?, PAUL C. PAQUET?, FRANK G. M. Burrows*, GRAHAM K. NEALE, KEITH D. WADE®. and WILLIAM M. SAMUEL 'Department of Biological Sciences, University of Alberta, Edmonton, Alberta T6G 2E9 Canada *Mailing address: P.O. Box 10, Jasper National Park, Jasper, Alberta TOE 1E0 Canada Faculty of Environmental Design, University of Calgary, Calgary, Alberta T2N 1N4 Canada 4Bruce Peninsula National Park, Tobermory, Ontario NOH 2RO Canada ‘Garcia and Associates, Bozeman, Montana 59718 USA °Pukaskwa National Park, Heron Bay, Ontario POT 1RO Canada Forshner, S. Anne, Paul C. Paquet, Frank G. M. Burrows, Graham K. Neale, Keith D. Wade, and William M. Samuel. 2003. Demographic patterns and limitation of Grey Wolves, Canis lupus, in and near Pukaskwa National Park, Ontario. Canadian Field Naturalist 118(1): 95-104. In response to concern regarding the growth and long-term viability of the wolf population in and near Pukaskwa National Park, a study of demographic patterns and limitation of radio-collared wolves (Canis lupus) was completed between 1994 and 1998. The mean annual finite rate of increase (0.96) suggested that population growth of wolves was limited and declining slightly. Small pack sizes, high cumulative mortality, and low reproductive success also suggested a declining population. Two limiting factors, ungulate biomass and human-caused mortality, were examined to determine the importance of each in limiting the population growth of wolves. Ungulate biomass was involved because occurrence of natural-caused mortality was high (9 of 17 wolves) compared with other studies. In addition, consumption rates were low and similar to other studies where starvation and other signs of malnutrition were noted. Further, Moose densities in the study area were low to moderate and below thresholds indicating nutritional stress for wolves. Occurrence of human-caused mortality was high (8 of 17 wolves) suggesting that it was also an important limiting factor, particularly given the low availability of ungulate biomass and reproduction noted in this study. Based on present demographic patterns, ungulate biomass, and human-caused mortality, the wolf population likely will remain at present low densities or continue to decline. Key Words: Canis lupus, wolves, limitation, demographic patterns, Pukaskwa National Park, Ontario. In 1996, researchers in the Greater Pukaskwa Eco- system (GPE) (Figure 1) in Ontario postulated that two limiting factors were negatively affecting the growth and long-term viability of the Grey Wolf (Canis lupus) population (Burrows et al. 1996"). These limiting fac- tors were low availability of ungulate prey and high mortality from human causes. Managers in Parks Can- ada Agency were concerned because wolves are a native species in the GPE. Thus, Parks is legislated to protect and further, to ensure the long-term viability of that population (Parks Canada 2000). This prompted a study of wolves in the western half of the GPE, in- _ cluding Pukaskwa National Park (PNP). In this paper, we quantify population limitation of _ wolves and examine the importance of ungulate bio- mass and human-caused mortality in limiting the popu- lation growth of wolves, 1994-1998. For these pur- _ poses, we review and discuss demographic data on wolf _ densities, population growth, reproduction, and mortal- ity. Further, we report and discuss data on the avail- ability of prey and rates of kill and consumption. ' * References marked with asterix (*) are listed in a separate Documents Cited section following Acknowledgements, all others are in Literature Cited. Study Area The study area comprised 4500 km? in the western half of the GPE on the north shore of Lake Superior in Ontario (48°N and 85°W) (Figure 1). The area includes PNP (1878 km?) but also adjacent land with intensive forestry, gold mines, towns and associated infrastruc- ture. Two distinct physiographic regions, coastal and inte- rior, occur within the study area. The coastal region is characterized by rugged topography with elevations ranging from 189 to 650 m. Many lakes and rivers occur in the area, creating a patchy landscape. The interior region is a flat plateau characterized by a heavily erod- ed landscape of mountains previously scoured by con- tinental glaciers (Poitevin et al. 1989"). Mean annual precipitation is 74 cm along the coast and 64 cm inland. Winter and summer temperatures range from —13C° — 14.6C° for the coastal area and —17C° — 15.9C° inland (Poitevin et al. 1989"). Ice cov- er on Lake Superior ranges annually from 5-100% (Skibicki 1994’). Vegetation on the coast along Lake Superior and in- land is mixed with associations of Balsam Fir (Abies balsamea), Jack Pine (Pinus banksiana), White Birch (Betula papyrifera), White Spruce (Picea glauca), Black Spruce (Picea mariana), Eastern White Cedar 95 96 THE CANADIAN FIELD-NATURALIST Lake Superior Legend | __ Greater Pukaskwa Ecosystem M8 Pukaskwa National Park wolf study area Highway 17 N Peete earier en eaten : 20 km M |Prepared by: Parks Canada a | Pukaskwa National Park 2001 —_L. Parent Vol. 118 FicureE |. Location of the wolf (Canis lupus) study area within the Greater Pukaskwa Ecosystem, Ontario, Canada (center 48°N, 85°W). (Thuja occidentalis), and Trembling Aspen (Populus tremuloides), with occasional Red Maple (Acer ru- brum) and other hardwoods more locally abundant in the southeastern corner of the study area. Predatory mammals included Grey Wolf, Black Bear (Ursus americanus), Red Fox (Vulpes vulpes), Lynx (Lynx canadensis), River Otter (Lontra canadensis), Fisher (Martes pennanti), American Marten (Martes americana), Mink (Mustela vison), and Weasels (Mus- tela spp.). Coyotes (Canis latrans) were rare except around towns. Moose (Alces alces) were the primary and most abundant ungulate prey species for wolves. Woodland Caribou (Rangifer tarandus tarandus) were few and concentrated in small bands along the coast of Lake Superior (Bergerud 1985). Numbers ranged from 6-14 in PNP, 1993-1997 (Wade 1993*, 1995*, 1997", 1999*). White-tailed Deer (Odocoileus virginianus) were rare in the GPE. Methods Capture and handling We attempted to locate wolves in as many packs as possible. Wolves were captured with modified leg- hold traps in summer (7 = 21) and by using a net-gun from a helicopter in early winter (n = 5). All wolves | 2004 Legend A Heron Bay @ Mobert First Nation © Marathon tx White River see22 Swallow River Pack —-—- Bremner River Pack —--— White River Pack == Cascade Lake Pack —-— Neys Pack |_| Rein Lake Pack — — Black River Pack (Cee Be} 0 _20 km | Prepared by: Parks Canada te Pukaskwa National Park 2001 _L. Parent iiaebn * ° ~ 2 annneeeed FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 97 i Highway 17 Ficure 2. Annual (1 April — 31 March) home ranges of wolves in the study area, 1994-1998. Home ranges are 95% MCP. Years for home ranges of packs were Swallow and Bremner River Packs, 1997-1998; White River Pack, 1996-1998; Cascade Lake Pack, 1995-1996; Neys Pack, 1995-1997; Rein Lake Pack, 1994-1998; Black River Pack, 1994-1996, 1997-1998. were immobilized with Telazol® (tiletamine hydro- chloride (HCL) and zolazepam HCL, A.H. Robins Co., Richmond, Virginia). Rectal temperature, pulse, and respiration of wolves were closely monitored throughout the procedure. Immobilized wolves were examined for injuries, equipped with conventional VHF transmitters (Lotek®, Newmarket, ON), weighed, sexed, and aged. A committee for care of wild animals approved all capture and handling operations (Wildlife Animal Care Committee, Ontario Ministry of Natur- al Resources, 1994-1997). Biotelemetry The target frequency for locating each radio-collared wolf was four times/month in summer (April-Octo- ber) and six to eight times/month in winter (Novem- ber-March). Wolves were located by plane using a portable receiver (Lotek® SRX-400), right-left switch boxes, and paired three-element Yagi antennae mount- ed on the wing struts of a fixed-wing aircraft (Cessna 185). Wolf location was recorded with a Global Posi- tioning System (Garmin® 75 Aviation). For all loca- tions, transmitter frequency, observer, date, time of location, number of wolves, color of wolves, and pre- sence of pups were recorded. Mean error of telemetry (difference between observed and true location) was calculated by using data we collected when regularly locating stationary transmitters placed throughout the study area. Location data were downloaded into a Geo- graphical Information System (GIS [Tydac SPANS*]) for display and analysis of wolf movements. Home Ranges Ranges V® software (Kenward and Hodder 1996°) was used to calculate annual (1 April — 31 March) sizes of home ranges. To represent these areas we used relocations of packs and 95% minimum convex polygons (MCP) (Mohr 1947). All obvious extrater- ritorial forays and dispersals were excluded from the analyses (Ballard et al. 1997). We assumed home ranges were defined when the observation-area curve formed an asymptote (Kenward and Hodder 1996") and locations were obtained throughout the year. For each pack we used one radio-collared wolf/year to represent the annual home range of the pack. This is reasonable as locations from one wolf indicate position of the entire pack when a high degree of association exists among pack members (Kolenosky 98 THE CANADIAN FIELD-NATURALIST and Johnston 1967; Fuller and Keith 1980; Fritts and Mech 1981; Ciucci et al. 1997). This condition was confirmed in this study by aerial observations of packs during telemetry flights. Accuracy of locations for the entire study was 150 m, which was the mean error of telemetry obtained by all participants. Accordingly, we changed the fix reso- lution from the Ranges V® default of 1 m to 150 m. This resolution was used to set the width of the bound- ary strip that was included in polygon edges and areas (Kenward and Hodder 1996"). We left the scal- ing parameter at the software default of 1 m, which means that each coordinate unit was | m from the next. Density, pack sizes, and population growth Density of wolves/1000 km? was calculated by determining intra-pack densities (number of wolves in pack/home range size) and averaging these den- sities/year (Potvin 1987; Bjorge and Gunson 1989; Okarma et al. 1998). The number of wolves in a pack was based on the maximum number of wolves ob- served in mid-winter (15 January-15 February). We defined a pack as a group of two or more wolves that traveled together for more than one month (Messier 1984). In two cases we had insufficient data to deter- mine the sizes of home ranges, thus we followed Mes- sier (1985) and used data from previous or subsequent years. Population growth or the mean annual finite rate of increase was calculated based on the ratio of succes- sive yearly estimates of density (Fuller 1989). Reproduction We did not observe wolves at dens during this study. Dense vegetation and the secretive nature of wolves precluded accurate visuals of wolf groups until Octo- ber or November, at which time pups were difficult to distinguish physically from adults. Hence, successful year-specific reproduction was ascertained when: (1) pups were captured in spring; or (2) a pack increased in size from March to the following December, pro- viding that sites of focal activities (e.g., pup-resting areas) were observed in the intervening time (Messier 1985). Unsuccessful reproduction (i.e., no or failed reproduction) was ascertained when: (1) a pack did not demonstrate focal activity sites in the summer; or (2) a pair remained together from March to the follow- ing December (Messier 1985). Results are reported for each pack by year. Moose density To examine availability of ungulate biomass to wolves, we used Moose density (moose/1000km?) based on aerial surveys using stratified random sampl- ing (Gasaway et al. 1986*). More specifically, from 1993 to 1999 a single Moose density was calculated for PNP and the three Wildlife Management Units (21A, 21B, 33) surrounding PNP where wolf packs were distributed. There was little or no change in Moose Vol. 118 density among yearly estimates (Burrows 2001), thus we averaged results from 1993-1999 for each area. Rates of kill and consumption of prey by wolves The rates of killing and consumption of large prey by wolves in four packs were studied by daily aerial and ground observation, January-March 1998. The Bremner River Pack was located 57 times between 18 January and 27 March 1998 (69 days) and the Rein Lake Pack was located 57 times between 8 Janu- ary and 26 March 1998 (79 days). Other packs locat- ed were the White River Pack, 22 times between 11 February and 20 March 1998 (38 days) and the Swal- low River Pack, 22 times between 9 February and 22 March 1998 (42 days). To calculate the kill rate, we recorded the number of animals killed by wolves/ tracking period and the number of wolves present at the kill (Messier 1985). Prey killed were located from the air and from ground-based tracking. At kill sites, we confirmed prey species, time, and cause of death. For only the largest pack of wolves (Bremner River), in addition to aerial locations, we simultane- ously snow-tracked wolf movements and collected scats to determine if all kill sites were found with the aerial telemetry. Technicians at Big Sky Laboratory (PO Box 0776, Florence, Montana 59833-0776) iden- tified prey remains by macroscopic examination and comparison with known material and hair-scale im- pressions (Adorjan and Kolenosky 1969°). In this analysis we considered only tracking sessions where pack locations were not separated by >54 hr. There were a few exceptions, however, where loca- tions were separated by 72 hr. These periods were retained in the analysis because wolves made a kill or visited one of several garbage dumps the day they were relocated making it unlikely that we missed a kill. Nonetheless, kill rates in this study should be considered minimums as wolves were not relocated every day and some small prey such as deer (fawns and adults), Caribou calves, Beaver, and other smaller prey items may have been missed. It is unlikely, how- ever, that we missed many of these kills because White-tailed Deer and Caribou were rare in the study area. We report kill rates as ungulates killed/wolf/day. Rates of consumption were calculated based on kill rates and average weights of wolves and prey. We calculated the whole weight of wolves based on the average from radio-collared adults and other wolves found dead in the study area. The average edible weights of Moose and beaver prey were assumed to be 330, 261, 114, and 13 kg for adult male Moose, adult female Moose, young-of-the-year Moose and Beaver, respectively (Peterson 1977; Thurber and Peterson 1993). We assumed the average weight of a White-tailed Deer was 105 kg for an adult male (Kolen- osky 1972; Forbes and Theberge 1996). Eighty % of the adult deer carcass was considered edible (Pimlott 1967; Forbes and Theberge 1996). All consumption rates are reported as kg prey/kg wolf/day. 2004. FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 99 TABLE 1. Sizes of annual home ranges and home range areas/wolf of seven packs of Grey Wolves (Canis lupus) in the study area, 1994-1998. Early Late winter winter number number 100% 95% Number of wolves of wolves MCP MCP of radio Area/wolf Year Pack in pack* in pack? (km?)° (km?)° fixes (km?)¢ 1994-1995 Black River® 6 4 283 156 53 26 Rein Lake 3 3 310 249 38 83 1995-1996 Black River® 4 4 548 388 5 97 Rein Lake 3 3 600 533 58 178 Neys* 3} 4 269 244 96 8] Cascade Lake! 1 204 170 26 170 1996-1997 Rein Lake® 2 | 561 Syi/ 39 279 Neys* 6 4 113 101 96 17 White River 2 2 407 345 Sy) 173 1997-1998 Black River® 4 3 468 450 74 113 Rein Lake® 2 1 692 600 87 300 White River 2, 2 589 498 65 249 Bremner River 6 3 760 644 93 107 Swallow River 5 3 567 500 62 100 Mean 8}55) DEG 455 388 63 139 SE 0.5 0.3 52 48 6.3 25 “Maximum pack size, 15 January-15 February. > Maximum pack size, March. © Sizes of home ranges were described using the minimum convex polygon method (MCP) (Mohr 1947). ‘Based on 95% MCP. © Packs that used town dumps. ‘Home range size and home range area/wolf were not included in mean because the areas were not fully defined; i-e., the observation-area curve was asymptotic but locations were not obtained throughout the year. Mortality and survival of radio-collared wolves We completed survival analysis for radio-collared wolves from 20 August 1994 to 31 December 1998. Wolves were re-located from time of capture until mortality or the radio-signal disappeared. For known deaths we estimated the date of mortality to the near- est day using evidence from the field. When evidence was unavailable, day of mortality was deemed the midpoint of the interval between the last day the wolf was known alive and the day it was discovered dead. The cause of mortality was often identified on site and when possible, confirmed by necropsy. We calculated the cumulative mortality of radio- collared wolves (n = 25) using the Kaplan-Meier prod- uct limit estimator and Minitab (Version 12) software. One of 26 captured wolves was shot by a trapper while in the research trap and is not included in the analysis. Cause of mortality was described using %. We assumed the proximate cause of death was the ultimate cause of death. We were unable to assess the relative importance of other factors that may have been involved. Results Twenty-six adult wolves were captured and then radio-collared (n = 25) or tagged (n = 1) from 1994- 1997. These animals represented seven packs and one lone wolf. Two of seven packs occurred almost exclusively in the park and all wolf packs were radio- collared in the study area. There were no other wolf packs in the study area during this study. We followed two packs in 1994-1995, four in 1995-1996, four in 1996-1997, and six in 1997-1998. The average mass of adult female wolves (n = 11) was 26.9 + 1.4 kg and that of adult males (n = 14) was 36.5 + 2.8 kg. Home ranges Sizes of annual home ranges (Figure 2) of seven packs across 13 pack-years were adequately described in this study (Table 1). Estimates accurately repre- sented areas used by wolves because sizes of annual home ranges were not correlated with number of relocations (r, = 0.52, 0.05 > P > 0.02). Home range sizes of packs and home range areas/wolf were vari- able. The average annual home range size was 388 + SE 48 km2 (95% MCP, n = 13, range = 101-644 km’) 100 (Table 1). The average home range area/wolf was 139 + SE 25 km? (95% MCP, n = 13, range = 17-300 km?) (Table 1). Density, pack sizes, and population growth Wolf density did not change over time; recorded densities were 7.9, 9.6, and 7.2 wolves/1000 km? in 1995-1996 (n = 4 packs), 1996-1997 (n = 4), and 1997- 1998 (n = 6), respectively. Density declines, however, if the Neys pack (Figure 2) is excluded from the calculations. This pack exclusively used dumps for food (Krizan 1997) and the home range was much smaller compared with all other packs in the study (Table 1). Accordingly, wolf densities were 7.1, 5.9, and 5.9 wolves/1000 km? in 1995-1996 (n = 3 packs), 1996-1997 (n = 3), and 1997-1998 (n = 5). Average mid-winter (15 January — 15 February) pack size was 3.5 + SE 0.5 wolves (n = 14 pack-years) (Table 1). This average declined in late winter (March) to 2.7 + SE 0.3 (n = 14 pack-years). The number of wolves in all except two packs remained stable or declined, 1994-1998. Numbers fluctuated annually in the Neys and Swallow River Packs (Table 2). Accord- ingly, the mean annual finite rate of increase from 1995-1998 was 0.96. Reproduction From spring 1994 to spring 1998, wolves reproduced successfully in eight of 22 pack-years (36%) (Table 3). This was a maximum estimate of successful repro- duction. In two of eight pack-years, we assumed wolves had reproduced because large numbers of wolves were noted in the packs in the following early fall and winter. Moose density Average densities of Moose varied among manage- ment units. Management Unit 33 had the highest Moose density (0.285 + 0.03-0.07 moose/km? 90% CI) followed by Unit 21A (0.225 + 0.02-0.03), 21B (0.220 + 0.02-0.03), and PNP (0.153 + 0.03-0.08). Rates of kill and consumption The Swallow River (3 wolves) and Bremner River Packs (5) killed and consumed more ungulates than the White River (2) and Rein Lake Packs (2). The kill rates for each pack respectively were 6.8, 3.4, 0.0, and 0.0 ungulates/wolf/day. Consumption rates were 0.21, 0.11, 0.0, and 0.0 kg prey/kg wolf/day. The White River and Rein Lake Packs did not kill any ungu- lates; however, both packs scavenged from various sources. The White River Pack scavenged from Moose that were killed by vehicles or trains, from other wolf kills, and from snare sets. The Rein Lake Pack scav- enged from refuse in the town dump for White River (Figure 2). Mortality and survival of radio-collared wolves As of 31 December 1998, 17 of 26 wolves radio- collared or tagged from 1994 to 1998 were dead, only four were confirmed alive, and five were missing. Eight wolves died from human causes: trains killed three, THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Highest numbers of wolves in packs in the study area, 1994-1998. The number of sightings is in parentheses. Pack size* Pack 1994 1995 1996 1997 1998 Rein Lake 3(4) = 3(3) 1(2) 1> 0 White River D (2) ee (A) 2(2) 2(9) 2(2) Bremner River 3(3)P Cascade River 1(4) 0 Black River 4Q) VAC) w4e@) 3(4) Swallow River 5 (D)o 3(3) 4(1) Neys 3(4) — 4(2) 2(2)) 4) * Maximum numbers of wolves seen in March. > Pack sizes were confirmed by track-counts made from the ground. TABLE 3. Reproductive success of wolf packs, 1994-1998. Year Pack 1994. 1995 1996 1997 =1998 Black River +4 4 - + + White River “pb = = = = Rein Lake +b = = = = Neys + - ? Bremner River ~ + Swallow River = = “+ = reproduced successfully; — = did not reproduce successfully. > This pack likely reproduced successfully because many wolves (10) were seen in late fall (November — December). three were snared, and two shot. Nine wolves died from natural causes: two starved, two were killed by other wolves, four died from disease (three from mange and one from blastomycosis), and one died from un- known natural causes. We assumed this last wolf was not killed by humans because we were in a remote area, there were no signs of humans in the area, and we found no bullets, snares, or other human devices. Survival of radio-collared wolves decreased between one and three years post-collaring. Wolves had a 32% (SE 0.10) chance of dying in the first year, a 30% (SE 0.15) chance of dying in the second year, and a 57% (SE 0.26) chance of dying in the third year. Median survival time was 689 days or 1.9 years post-collaring. Discussion Population limitation of wolves The growth rate of the wolf population in the study area was limited from 1995-1998. The mean annual finite rate of increase, 0.96, indicated a 4% rate of decline. This rate of increase is not unique, however, and similar rates recorded from other populations have varied from 0.93-2.40 (Theberge and Strickland 1978; Fritts and Mech 1981; Ballard et al. 1987; Hayes et al. 1991"; Messier 1991; Pletscher et al. 1997). \ | i { 2004 In addition to the estimated rate of growth, there are a number of other factors that suggest the wolf popula- tion was declining slightly. First, pack sizes were small and generally declining. Mean pack size (3.5 wolves + SE 0.5) was much smaller than the average of 10 wolves for packs that hunt moose in North America (Mech 1970). Furthermore, five of seven packs in this study remained stable or declined in size from 1994 to 1998. If this population were increasing in size, the number of wolves within packs would likely increase. This happened in the Yukon where rapid increases in pack sizes of colonizing wolves were the primary means by which an intensively reduced wolf population reached their pre-reduction densities (Hayes and Harestad 2000). High cumulative mortality of wolves is the second factor that suggested a declining population. We com- pared the cumulative rate of mortality from the first year (32%) of our study with annual rates of mortal- ity from other studies. There is little agreement among researchers on the annual rate of mortality that causes a population decline in wolves. However, Fuller (1989) reviewed several wolf studies across North America and concluded that populations would stabilize with an overall annual mortality rate of 35%. Given this, it appears the mortality rate in this study was sufficient to account for the slightly declining rate of growth in this study. Coupled with high mortality of adult wolves, low reproductive success of wolves in this study suggested a population decline. Wolves reproduced successfully in only 36% of possible occasions compared with 45-93% noted in other areas (Messier 1985; Potvin 1987; Peterson et al. 1998; Hayes and Harestad 2000). We were unable to determine if wolves produced pups that died soon after birth or whether whelping occurred at all. Lack of denning, however, suggests no pups were produced. Limiting factors We examined the importance of two factors that lim- ited the growth of the wolf population in this study: un- gulate biomass and human-caused mortality. Ungulate biomass is commonly reported to limit growth of other wolf populations (Mech 1977a; Fuller and Keith 1980; Packard and Mech 1980; Keith 1983; Messier 1985; Peterson and Page 1988) and data from this study suggest it was important. The strongest data rep- resented the occurrence of natural-caused mortality. In this study, more than half (9 of 17) of radio- collared wolves died from natural causes, which is high compared with other North American studies (Peter- son 1977; Carbyn 1982; Peterson et al. 1984; Ballard et al. 1987; Hayes et al. 1991”). Starvation and intra- specific aggression were responsible for four of nine (24%) wolves dead in this study and have been report- ed common in other populations where ungulate bio- mass is low. For instance, in southwestern Quebec, FORSHNER ET AL.: GREY WOLVES IN PUKASKWA NATIONAL PARK 10] Messier (1985) noted wolves with fewer prey available incurred more deaths from natural causes, namely starvation and intraspecific aggression. In that area, similar to our study area, moose density was 0.23 moose/km? and there were no other ungulate species present. Similarly, Mech (1977a) noted occurrence of starvation and intraspecific aggression increased as prey availability declined in Minnesota. Disease was the other natural cause of death ob- served in this study. Four of nine wolves (24%) died from either sarcoptic mange or blastomycosis. Blasto- mycosis is enzootic in Minnesota (Schlosser 1980) and Wisconsin (Sarosi et al. 1979; McDonough and Kuzma 1980) but until now (Krizan 2000; Paquet et al. 2001), had not been reported from other wolf populations across North America. This level of disease-related mortality has not been reported in any other popula- tions of wolves. In other populations, disease accounts for 2-21% of wolf mortality (Carbyn 1982; Peterson et al. 1984; Fuller 1989; Ballard et al. 1997) and is often not even reported (Messier 1985; Ballard et al. 1987; Potvin 1987; Hayes et al. 1991°; Meier et al. 1995; Pletscher et al. 1997). The only other study where disease clearly affected a wolf population was in Alaska where rabies accounted for 21% of wolf mor- tality and was a significant factor in the decline of the population (Ballard et al. 1997). Disease cannot be linked with certainty to low ungulate biomass but wolves that lack food should be more vulnerable to disease than those with more food available. Furthermore, food shortage leading to nutritional stress could combine with disease factors to increase the significance of otherwise innocuous or sub-lethal conditions (Brand et al. 1995). We also examined rates of consumption of ungu- late prey to determine the importance of ungulate bio- mass as a limiting factor. Consumption rates for three of four packs in this study were low (Bremner River, Rein Lake, and White River). These packs consumed <0.13 kg/kg wolf/day, which Mech (1977b) deter- mined is the minimum rate of consumption required for all wolves to survive and rear pups successfully. Two packs killed no ungulates and relied on scav- enging to survive (White River and Rein Lake Packs). These data suggest that at least three of four packs could have been limited by food. Indeed, in the White River Pack, the dominant female failed to reproduce the following spring and was extremely emaciated (mass = 23.5 kg) when killed by other wolves later in the summer. Similarly, the Rein Lake Pack was reduced to one wolf by winter 1997. She did not reproduce the following summer and to survive, she scavenged mainly from the town dump for White River (Figure 2). She was dead as a result of mange by December 1998 (mass = 28.5 kg). The Bremner River Pack may have been limited by food but data were not strong. One wolf dispersed in summer 1998 and died from unknown natural causes. Other wolves could have dis- 102 persed and died later because pack numbers dropped from nine to three over the winter 1997-1998 (un- published data). Some members of this pack, however, did survive and reproduce two years in a row. Most of the rates of consumption in this study are similar to those from other areas where starvation and other signs of malnutrition of wolves were noted. For instance, Messier (1987) noted more deaths of wolves from malnutrition in areas of low density of moose (0.23 moose/km7) where wolves had 0.05 kg/kg wolf/ day (based on kill rate of 1.7 kg/wolf/day and wolf mass of 32.3 kg). Peterson and Page (1988) noted starvation and other indicators of severe nutritional stress in an area of high Moose density (1.9 moose/ km?) when food availability fell below 0.12 kg/kg wolf/day (based on kill rate of 4.0 kg/wolf/day and wolf mass of 32.3 kg). As a final method to assess the importance of un- gulate biomass, we examined density of Moose, the main prey for wolves in this study. Moose density was low to moderate (0.153-0.285 moose/km?) and similar to Moose densities in other areas where wolves were nutritionally stressed. Messier (1987) found that in areas where Moose density dropped below 0.4 moose/km*, wolves were nutritionally stressed. He also reported that below 0.2 moose/km? wolf packs could not subsist and (or) reproduce successfully (Messier 1985). The second limiting factor we examined was human- caused mortality. Besides ungulate biomass, it is the other most commonly reported factor that limits the growth of wolf populations (Gasaway et al. 1983; Keith 1983; Peterson et al. 1984; Fuller 1989; Noss et al. 1996; Paquet et al. 1996"). In other areas where human-caused mortality was considered the primary limiting factor, it accounted for 69-80% of all mortal- ity (Peterson et al. 1984; Ballard et al. 1987, 1997). In our study area, human causes accounted for only 47% of mortality of adult radio-collared wolves. None- theless, human-caused mortality is likely still impor- tant, particularly given the low ungulate biomass and reproduction noted in this study. Gasaway et al. (1983) found that in areas with low ungulate biomass, har- vest levels as low as 20% can limit wolf populations. Fuller (1989) found that wolf populations with low productivity can withstand less overall mortality be- cause there are fewer pups, which often make up dis- proportionate amounts of harvests. In conclusion, the population growth of wolves in this study area was limited and declined slightly based on (i) mean annual finite rate of increase; (ii) small and generally declining pack sizes; (iii) high cumulative mortality; and (iv) low reproductive success. Based on these demographic patterns, low availabil- ity of ungulate biomass and existing levels of human- caused mortality, this population likely will remain at present low densities or continue to decline. This situa- tion is challenging to managers for Parks Canada Agen- cy because the study area, which includes a National THE CANADIAN FIELD-NATURALIST Vol. 118 Park, may not have a highly productive source popu- lation for wolves. Further, protection for wolves out- side the park is limited because few restrictions exist regarding the nature, timing, and extent of wolf har- vesting. Acknowledgments Field research was funded by Parks Canada. We thank all PNP wardens and staff for field and techni- cal support, particularly K. Wade, S. Sutton, and C. Strong. Additionally L. Parent provided much GIS expertise. Thanks also to J. Whittington and R. Whit- tington for field support and pilots W. Roberts and M. Robb for many safe hours of flying time. Thanks to Helicopter Wildlife Management Team for skillful capturing and collaring of animals. All animal treat- ment procedures were approved by the Wildlife Animal Care Committee, Ontario Ministry of Natural Resources (Permit Numbers 1994-13, 1995-13, 1996- 13, 1997-13). We thank J. Whittington, B. Dobson, and M. Boyce who incisively reviewed later drafts of this manuscript. S.A. 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Journal of Mammalogy 74: 879-889. Received 13 March 2003 Accepted | March 2004 Estimation of Seed Bank and Seed Viability of the Gulf of Saint Lawrence Aster, Symphyotrichum laurentianum, (Fernald) Nesom Joni F. Kemp and CureisTIAN R. LACROIX! Department of Biology, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island C1A 4P3 Canada 'Author to whom correspondence should be addressed. Kemp, Joni F., and Christian R. Lacroix. 2004. Estimation of seed viability of the Gulf of Saint Lawrence Aster, Symphyotrichum laurentianum (Fernald) Nesom. Canadian Field-Naturalist 118(1): 105-110. The Gulf of St. Lawrence Aster, Symphyotrichum laurentianum, is a member of the family Asteraceae and is listed as “threatened” by COSEWIC (Committee on the Status of Endangered Wildlife in Canada). This rare and vulnerable halophyte grows in only a few locations in New Brunswick, Prince Edward Island, and the Magdalen Islands, Quebec. As an annual, S. laurentianum relies exclusively on its seeds to survive to the next generation. The goal of this study was to estimate the quantity of viable S. laurentianum seeds in the persistent and transient seed banks at selected sites in Prince Edward Island. Overall, the number of seeds in the transient and persistent seed banks is low. The greatest concentration of seeds was found near the surface of the soil. In addition, only a small proportion of those seeds tested positive for viability based on Tetrazolium staining. Of the seeds in the persistent and transient seed banks combined, 53% were viable whereas only 2% of the seeds in the persistent seed bank were viable. Population surveys were also completed at the five known sites (both extinct and extant) in Prince Edward Island National Park. All sites showed signs of decline based on population estimates dating back to 1993. The Covehead Pond site showed the greatest decline: from 250-300 individuals in 1993 to only 10 individuals in 2002. The population at Dune Slack also showed a dramatic decrease from approximately 65 000 in 1999, to 2 200 individuals in 2002. Monitoring of this plant and the development of a management plan for the species are critical to its survival. Key Words: Symphyotrichum laurentianum, Aster laurentianus, Gulf of St. Lawrence Aster, seed viability, Tetrazolium, transient and persistent seed banks, rare plant, COSEWIC, Prince Edward Island National Park. The Gulf of St. Lawrence Aster, Symphyotrichum laurentianum (Fernald) Nesom, is a member of the family Asteraceae and is listed as “threatened” accord- ing to COSEWIC (Committee on the Status of Endan- gered Wildlife in Canada 2004*). It is an annual plant that relies exclusively on its seeds to survive until the next germination season, which makes this plant very vulnerable. S. /aurentianum is a rare halophyte only found in a few areas in Prince Edward Island (PEI), New Brunswick (NB), and the Magdalen Islands in Quebec. It grows in salt marshes that are only flooded during spring and high tides. The St. Lawrence Aster has smooth, hairless, and fleshy linear-lanceolate to spatulate, generally sessile leaves (Figure 1). It can grow up to 30 cm tall and can bear | to over 700 flower heads (Houle and Haber 1990). Each flower head is subtended by leafy bracts and is composed exclusively of white to pink disk florets surrounded by a white pappus (Stewart 2000* and Figure 1). Seeds are enclosed in achene fruit walls when they are dispersed. In 1990 fewer than 1000 plants were known to exist in PEI, NB and Quebec (Magdalen Islands) combined. However, further research and surveys have produced several new sites. While several sites in PEI and Que- bec contain more than 1000 individuals each, the St. Law- rence Aster is still vulnerable to extinction (Gilbert et al. 1999*; Stewart and Lacroix 2001). Recent research conducted by Stewart (2000*) showed that population sizes could suddenly decline dramatically. The extent of the persistent seed bank of a plant must be known in order to fully understand the germina- tion ecology of the species (Baskin and Baskin 1998). In addition, the viability of a plant’s seeds must also be maintained in order for the plant to be successful (Fenner 2000). A seed bank is a reserve of viable, un- germinated seeds in the soil. Seeds are considered vi- able if they have the ability/potential to germinate under favourable conditions (Baskin and Baskin 1998). There are two categories of seed banks: transient and persis- tent. A transient seed bank refers to a short-term storage of seeds, that is seeds that are viable for one year or one germination season. The seeds in the transient seed bank therefore germinate in the first year after dispersal. Any seeds that are dormant and survive to subsequent ger- mination seasons are considered to be part of the per- sistent seed bank. The persistent seed bank contains seeds that remain viable in the soil for at least two ger- mination seasons, making this a long-term reserve of seeds that are at least one year (or one germination sea- son) old. A knowledge of both the extent of the seed banks and the germination potential of seeds of S. lau- rentianum constitutes crucial baseline data to develop a management plan for the species. The specific goals of this study were: (1) to estimate the quantity of seeds in the seed bank (both transient and persistent) for S. lJawrentianum at each of the select- ed sites in PEI in order to evaluate the ability of the plant to maintain current populations, and (2) to test the 105 106 THE CANADIAN FIELD-NATURALIST Vol. 118 FiGurE |. Representative photographs of Symphyotrichum laurentianum, pre-flowering (A) and post-flowering (B). viability of seeds in the persistent seed bank and the transient seed bank. Materials and Methods Population Surveys Five sites within the PEI National Park (Figure 2) . Were visited. Total population counts were made at the Covehead Pond and East Marsh A sites because of the small number of plants at these locations. The Western Wetland site was also surveyed to confirm that no asters were growing there. Population estimates were recorded for Dune Slack and East Marsh B sites be- cause of the large population sizes. These estimates were accomplished by using a grid system. The gener- al area where the plants were growing was subdivided into 0.5 m? sections. A random number of sections was selected and all plants within these grids were counted. These counts along with the total area of the site were used to extrapolate the approximate number of plants found at each site. Collection of Soil Samples Soil cores were taken from five sites within PEI National Park (Figure 2), the only location in the pro- vince where the plant is found: Covehead Pond, Dune Slack, East Marsh A, East Marsh B and Western Wet- land. Three transects were run at each site except Cove- head Pond. Twenty coring points were flagged along three transects in Dune Slack. Due to sampling limita- tions imposed by Parks Canada, only ten coring points were flagged along three transects at East Marsh A and a further ten along the three transects at East Marsh B. Similarly, ten coring points were also flagged along the three transects for Western Wetland. Eleven coring points were flagged at Covehead Pond in a slightly different manner because of the physical nature of the site. Four transects were run from the water’s edge of the pond to the high tide mark along the traditionally heavily populated side of the pond. Eight coring points were flagged evenly among these transects and the re- maining three points were spaced evenly along the other side of the pond where the plants were also reported to grow. Two sets of soil cores were taken from each site: one during late June/early July, and another during late August/early September. These dates were chosen to correspond to the type of seed bank that was sampled. The late June/early July samples were taken before the emergence of the 2002 season plants, to ensure that the transient as well as persistent seed banks were includ- ed. The late August/early September samples were taken after all the seeds that were going to germinate that season had germinated. Consequently, only the persistent seed bank was included in this specific sam- ple date. All the core samples were taken using an Oakfield “LS” 36” soil sampler with an effective coring length of 24 cm and an inner diameter of | inch. In order to study the vertical distribution of seeds in the soil, each soil core was divided equally into a top, middle, and bottom section. Each section of the core was labelled, placed into a separate bag, and put in a freezer set at -4°C until processing which took place two weeks to three months later. All seeds found in the soil samples were tested for viability. Viability Testing Seeds were tested for viability using 2,3,5-triphenyl tetrazolium chloride (TTC) (Grabe 1970). The TTC test is based on a colour reaction: any seeds that are 2004 KEMP AND LACROIX: SEED BANK AND SEED VIABILITY 107 FIGURE 2. Map of all Prince Edward Island sites used in this study. Study sites are depicted as squares on both maps. All five sites shown in (A) are located in the national park: 1. Covehead Pond, 2. Western Wetland, 3. Dune Slack, 4. East Marsh B, and 5. East Marsh A. (B) Location of sites on Prince Edward Island with black square highlighting the specific area shown in (A). viable will stain pink/red. Complete achenes (contain- ing seeds) were placed on a piece of filter paper in a petrie dish, and left overnight to soak in distilled water. The next day, when the seeds were fully imbibed with water, they were dissected out of the achene fruit wall and seed coat. After removal of the seed coat, the embryos were placed in a 1% TTC solution for two hours at 35°C, as prescribed for dicotyledonous seeds. The embryos were then evaluated for viability according to the scheme developed in Grabe (1970). As a general rule, the radicle, shoot tip and cotyledons must stain pink/red in order for the seed to be con- sidered viable. Statistical Analysis The viability of seeds in the soil was compared graphically for vertical distribution and sampling dates. Statistical analysis was not possible on these data be- cause there was no replication. Due to the vulnerable status of the plant, sampling was limited by restrictions on collecting imposed by Parks Canada. The vertical distribution of seeds and fruit walls within the soil was compared using Kruskal-Wallis tests (non parametric equivalent of ANOVA). All data from the five sam- pling sites on PEI were pooled for analysis. 108 Results Population Sizes Population sizes for all known PEI sites are shown in Table 1. All populations experienced a decrease in numbers, the most noticeable of these being the pop- ulation at the Dune Slack site with more than 95% decline. The smallest population at the Covehead Pond site also experienced a decrease from 243 individuals in 1999 to only 10 individuals in 2002. Seed Banks A total of 122 cores were taken from PEI sites. Of these, 32 contained at least one seed. There were sig- nificantly more seeds in the top portion of those cores than in any other portion of the soil samples (Figure 3; Kruskal-Wallis test; P < 0.05, n=45 for each por- tion). The non-random distribution of the seeds, as evi- denced by the highly variable number of seeds found in the core samples, is also highlighted in Figure 3. This may be related to where the plants clustered the year before, a factor we were not able to determine prior to establishing coring sites. Seed Viability There was a noticeable decline in the percentage of viable seeds among the top, middle, and bottom sec- tions of the soil cores taken on PEI. We found that 28.4% of the seeds in the top section of the cores were viable compared to 20% of the seeds in the middle sec- tion of the cores. None of the seeds in the bottom section of the cores were viable. There was a marked decline in the percentage of via- ble seeds between the late June/early July sample and the late August/early September sample (Figure 4). The percentage of viable seeds in the persistent seed bank (2%) is practically non-existent when compared to the viability of the seeds in the persistent and tran- sient seed banks combined (53%). This means that the majority of viable seeds germinate within one year of being produced. THE CANADIAN FIELD-NATURALIST Vol. 118 Discussion Population Sizes Symphyotrichum laurentianum populations on PEI continue to show dramatic fluctuations and it can be very difficult for an annual plant to rebound from these declines in population size. Although the Dune Slack site was considered one of the two largest populations on PET]; its rate of decline, if maintained, will bring the population down to a size comparable to the smallest populations. A comparatively small population at Covehead Pond has been relatively stable at 150 to 300 individuals for the past ten years but the last popu- lation count found only ten individuals which, cou- pled with the low viability of seeds in the seed banks, indicates that this site is on the verge of extinction if preventive measures are not taken. The Island Nature Trust, a local conservation orga- nization, conducted a survey of suitable areas on PEI in 2002 to locate new populations of S. /aurentianum. Several prime candidate habitats were located but un- fortunately no asters were found at those sites (Mac- Quarrie, personal communication). Seed Banks There appears to be a lack of mixing of the soil in the salt marshes which causes a clustering of seeds in the top sections of the soil. This trend is obvious in the vertical distribution of seeds shown in Figure 3. Seeds that remain trapped in the upper layer of the soil are more likely to get exposed to the elements thereby potentially affecting the viability of the embryo. One of the most meaningful results from this study is the fact that the viability of seeds in the persistent seed bank is practically non-existent. This can be dev- astating for an annual plant such as S. laurentianum. Without the long-term storage of viable seeds, this plant is vulnerable to short term changes in its habitat, both natural and anthropogenic. TABLE |. Comparison of population sizes for Symphyotrichum laurentianum on Prince Edward Island. Population GPS Location Covehead Pond 46° 25' 49.39" N 63° 09' 07.98" W 46° 24' 47.88" N 63° O1' 16.89" W Western Wetland Dune Slack 46° 24' 51.49" N 62° 59' 41.98" W East Marsh A 46° 24' 55.97" N 62° 58' 43.89" W East Marsh B 46° 24' 55.27" N 62° 59' 06.11" W * estimate 1992 1999 2002 168 243 10 425 0 0) 15 000 — 20 000* 65 250* 2 200* 133 50 000 — 60 000* 25 000 — 60 000* (Sites A and (Sites A and 44 100* B combined) B combined) | ) 2004 40 ; 2 Qa. 5 om 30 th, o jo gat ° io ‘0 o ep) — * 2 mn o a E =— yea Top Middle Bottom Portion of the Core (N= 32) FiGuRE 3. Box plots showing median (centre line) and 25" and 75" percentile for numbers of seeds found in cores from all Prince Edward Island soil samples that con- tained seeds. Outliers (*) are included to show the non-random distribution of the seeds. During the summer of 2002, the Dune Slack site was heavily flooded during a storm. This caused the site, and consequently the maturing aster plants, to be smoth- ered in eel grass. Since the Dune Slack population had already experienced a dramatic decrease in size between 1999 and 2002, the necessary number of mature viable seeds needed to survive this natural short-term event may not be available. Covehead Pond is an example of a site that is very likely to become extinct due to anthropogenic causes. This site is isolated from Covehead Bay by a road. The culvert that is presently in place did not allow proper drainage from the pond during the summer of 2002 and caused this site to be flooded. The ten plants that were 100 a n xo} o o no o a 60 & > es fo} oO RB 40 4 = oO 2 a) ae 20 | ) i) 1 2 3 Persistent and Transient Persistent N=89 N=93 Seed Bank Type FiGurE 4. The percentage of viable seeds in the seed banks on Prince Edward Island. Persistent and transient seed banks = all late June/early July samples on Prince Edward Island. Persistent seed bank = all late August/ early September samples on Prince Edward Island. KEMP AND LACROIX: SEED BANK AND SEED VIABILITY 109 seen at this site in 2002 were flooded by water and also smothered by eel grass. This occurred at a crucial time during seed maturation. Only one plant was found at the Covehead site in 2003. Status of the Species In 2003, there were 441 Canadian species at risk, and 140 of these were plants (Committee on the Status of Endangered Wildlife in Canada 2004%*). Until recently, there has been very little in terms of Federal legislation in place to protect these species. The Species at Risk Act (SARA) which came into effect in 2002 will hopefully facilitate this process especially with regard to developing a plan for the recovery of every extir- pated, endangered, or threatened species. The results of this study show a disturbing trend: the estimated number of seeds in the seed banks is low, the viability of the seeds in those seed banks is also low (especially in the persistent seed bank), and the popu- lations are declining. The Gulf of St. Lawrence Aster is currently under review and it is important that all available information on the biology of the species be considered (especially information pertaining to seed banks). The status of this plant has recently been up- dated to “threatened” which is defined as “a species that is likely to become endangered if nothing is done to reverse the factors leading to its extirpation or ex- tinction” (Committee on the Status of Endangered Wildlife in Canada 2004*). This new designation con- firms the urgent need to design a management plan for S. laurentianum to ensure that it does not become ex- tinct. Although harsh weather patterns leading to strong winds and flooding cannot be prevented (i.e., extreme flooding of the Dune Slack site during summer of 2002), alternate measures such as transplanting ex situ plants into affected sites should be considered. Pre- liminary results show that seedlings of S. laurentianum can be grown from seed and transplanted from cul- ture medium to soil with a relatively high success rate (Stewart 2000*). However, a population decline and loss of suitable habitat due to anthropogenic activities can be prevented and further steps. such as ensuring proper drainage at a site such as Covehead Pond, should be taken. Acknowledgments This research was partially funded by an NSERC (National Sciences and Engineering Research Council) operating grant to C. Lacroix. We would like to thank Parks Canada for allowing us access to the sites to collect samples. Thank you to Kate MacQuarrie for her comments on this manuscript. Documents Cited (marked * in text) Committee on the Status of Endangered Wildlife in Can- ada. 2004. COSEWIC assessment and update status report on the Gulf of St. Lawrence Aster, Symphyotrichum lauren- tianus in Canada. Ottawa. 39 pages. Retrieved May 2004, froma http://www.cosewic.gc.ca 110 Gilbert, H., J. Labrecque, and J. Gagnon. 1999. La situa- tion de l’aster du Saint-Laurent (Aster laurentianus, syn. : Symphyotrichum laurentianum) au Canada. Ministére de l'Environnement Direction de la conservation et du patri- moine écologique, Québec. Stewart, S. E. 2000. Micropropagation of Aster lauren- tianus. Unpublished honours thesis, University of Prince Edward Island, Charlottetown, Canada. Literature Cited Baskin, C. C., and J. M. Baskin. 1998. Seeds: Ecology, bio- geography, and evolution of dormancy and germination. Academic Press, San Diego. Fenner, M. 2000. Seeds: The ecology of regeneration in plant communities. CABI Publishing, New York. THE CANADIAN FIELD-NATURALIST Vol. 118 Grabe, D. E., Editor. 1970. Tetrazolium testing handbook for agricultural seeds: contribution Number 29 to the hand- book on seed testing. Tetrazolium Testing Committee of the Association of Official Seed Analysts. Houle, F., and E. Haber. 1990. Status of the Gulf of St. Law- rence Aster, Aster laurentianus (Asteraceae), in Canada. Canadian Field-Naturalist 104: 455-459. Stewart, S. E., and C. R. Lacroix. 2001. Germination poten- tial, updated population surveys and floral, seed and seed- ling morphology of Symphyotrichum laurentianum, the Gulf of St. Lawrence Aster, in the Prince Edward Island National Park. Canadian Field-Naturalist 115: 287-295. Received 2 June 2003 Accepted 3 June 2004 Consumption of Shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus JONATHAN W. Moore! and G. J. KENAGY? ' Department of Biology, University of Washington, Box 351800, Seattle, Washington 98195 USA; e-mail: jwmoore @u. washington.edu * Burke Museum and Department of Biology, University of Washington, Box 353010, Seattle, Washington 98195 USA e-mail: kenagy @u.washington.edu Moore, Jonathan W., and G. J. Kenagy. 2004. Consumption of shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus. Canadian Field-Naturalist 118(1): 111-114. In an investigation of the dietary habits of Arctic Grayling (Thymallus arcticus) we found that two individuals out of 93 sampled in southwestern Alaska (approximately 59°N, 159°W) contained a total of five shrews (Sorex spp.). These shrews contained enriched levels of nitrogen stable isotopes, suggesting utilization of nutrients derived from salmon. We hypothesize that normally terrestrial shrews accidentally enter streams while foraging along the productive riparian zones of creeks with high densities of salmon. Shrews are apparently susceptible to opportunistic predation by resident stream fishes, including Arctic Grayling, when they enter the streams. Key Words: Arctic Grayling, Thymallus arcticus, diet, salmon-derived nutrients, shrews, Sorex spp., stable isotopes. The Arctic Grayling (Thymallus arcticus) is a com- mon nearctic freshwater fish in lakes and streams at high latitudes (Nelson 1994). Grayling typically feed on a variety of aquatic and terrestrial invertebrates in the water column or air-water interface (Armstrong 1986). In addition, Arctic Grayling occasionally supple- ment their diets with salmon eggs and small fish (Armstrong 1986). Despite existing knowledge of Arctic Grayling biology and its regional importance, the ecol- ogy and behavior of this species remain not well un- derstood, including the extent to which Arctic Grayling exploit terrestrial sources of prey (Armstrong 1986). As part of an extensive study of the aquatic and asso- ciated terrestrial ecosystem and its relationship to sal- mon in southwestern Alaska (Schindler et al. 2003), we had the opportunity to investigate dietary habits of Arctic Grayling. We discovered evidence of apparently rare consumption by Arctic Grayling of small terres- trial mammals of the Order Insectivora, shrews of the genus Sorex. During summer 2001 we sampled resident fishes in five streams that flow into Lake Nerka in the Wood _ River-Tichick State Park, Dillingham County, south- western Alaska (approximately 59°36’N, 159°05’W), by conventional and fly angling. The study streams sup- | port large runs of Sockeye Salmon (Oncorhynchus _ nerka). Captured fish were anesthetized, marked with subcutaneous tags, weighed, measured, and released. _ We obtained diet samples by back-flushing the stomach with a gastric lavage. Diet samples were preserved in | ethanol and transported back to the University of Washington for analysis. _ We sampled the stomach contents of 93 Arctic Gray- | ling from five tributaries of Lake Nerka. Two of the 93 Arctic 2001 Grayling contained visible evidence of the body parts of shrews (Figure 1, Table 1). One Arctic Grayling (length 394 mm, mass 591 g), captured on 13 August in the Little Togiak River contained the remains of three shrews. A second Arctic Grayling (length 391 mm, mass 522 g), captured on 25 August in Elva Creek contained the remains of two shrews. Nothing else about the two observed Arctic Grayling was unusual other than their stomach contents. Although the fish were relatively large, they were not extreme, being longer than about 70% of the other Arctic Gray- ling. In addition to the shrews, the diets of these two fish contained aquatic and terrestrial insects typical of the diets of the other Arctic Grayling sampled at this same time and location. The five alcohol-preserved shrews (Sorex spp.) from the diet samples of two Arctic Grayling were deposit- ed in the Burke Museum mammal collection. Three shrews (UWBM 74151, 74152, 74154) were sufficient- ly intact that we could examine individual teeth in the skull, which is critical for species identification. We used the keys of Nagorsen (1996), based on body and skull measurements and dental characteristics, particu- larly the relative length of the third unicuspid. In two cases, digestion had progressed too far to allow species determination by these characters. We subsequently referred tissue samples of all five specimens to Eric Waltari, Idaho State University, for molecular (cyto- chrome-b) identification, according to protocols used in his systematic investigation of Sorex, as developed by Demboski and Cook (2001). Molecular identification confirmed the three initial morphological identifica- tions and provided definitive identification for the two previously unidentifiable individuals (UWBM 74153, 74155) The two shrews from Elva Creek were identified as Sorex monticolus and Sorex cinereus (Figure 1, upper). The S. cinereus specimen had a wet weight of 3.3 g IU 112 and was fairly intact except for digestion of the fur away from the skin. Due to more extensive digestion, the S. monticolus specimen could not be measured and weighed accurately, but its intact skull and dentition allowed a positive identification (Table 1). The three shrews from the Little Togiak River (Fig- ure 1, lower) included a highly intact S. cinereus with considerable fur still attached to its skin (Figure 1, lower, center). The other two shrews were extensively digested and could only be identified by molecular analysis, as S. cinereus (left) and S. monticolus (right). In summary, the two Arctic Graylings from two dif- ferent streams both had fed on shrews of two species (Table 1). The presence of the shrews Sorex monticolus and Sorex cinereus in this region of southwest Alaska ly- ing to the north of Bristol Bay and east of the Kilbuck Mountains is consistent with the known geographic ranges of shrews (Hall 1981) and a more recent dis- tributional analysis (S. O. MacDonald and J. A. Cook, personal communication). Shrew abundance and diver- sity are high in this area, and three other species are also present: S. hoyi, S. tundrensis, and S. yukonicus. Although some species of the genus Sorex live and feed primarily in water (Beneski and Stinson 1987), none of the five Sorex species in the study area is pri- marily aquatic. However, all five occur to some extent in riparian habitat, where their success is enhanced by the aquatic component of the food chain. As a result of this habitat association, the shrews are potentially accessible to predatory fishes in these waters. The ten- dency for shrews of the genus Sorex to enter water, whether for foraging or dispersal, has been reviewed by Hanski (1986). In addition to intentional entry into the water, it is also possible that these shrews occa- sionally enter the water accidentally, resulting from their normal activities near the water’s edge. Shrews may be attracted to the riparian zones of these creeks because of the potential foraging oppor- tunities offered by abundant salmon carcasses. Ceder- holm et al. (1999) list shrews, including Sorex cinereus, as known scavengers of salmon carcasses. Evidence from stable isotopes ('°N) from our study (Table 1) and Ben-David et al. (1998) show that shrews (Sorex spp.) living near streams containing anadromous sal- THE CANADIAN FIELD-NATURALIST Vol. 118 mon runs bear the enriched isotopic signature of feed- ing directly or indirectly on salmon, for example, on insects that feed on salmon carcasses. We analyzed the stable isotope signature of muscle tissue from each shrew. Because anadromous salmon have a relatively high isotopic signature (6 °N = 11-13) compared to most other potential food sources, isotopes have been used to trace salmon nutrients through both aquatic and terrestrial food webs (Kline et al. 1990; Ben- David et al. 1998). The abundance of '°N in shrew muscle tissue was measured by combusting several mg of ground muscle tissue from each shrew in a mass spectrometer by the Stable Isotope Lab at the Univer- sity of California, Davis. Stable isotopes were expressed as a delta (6) value—the deviation from an isotope standard (atmospheric nitrogen). In other words, SION) = (GSNae Nie ee) ON reenact) XoLOOO! We used a two-sample t-test to compare 6°N of shrews in diets of Arctic Grayling to published values of isotopes from six shrews that were collected more than 500 m from streams with salmon in southeast- ern Alaska by Ben-David et al. (1998). The five shrews we collected from Arctic Grayling diets were relatively enriched, averaging 5.88 + 0.47 6 PN (mean + standard error; Table 1), higher than shrews collected from sites over 500 m from Alaskan salmon-bearing streams, that averaged 4.5 + 0.3 6 !SN (ty9 = 2.24, P < 0.10) (Ben-David et al. 1998). This relative enrichment suggests that the shrews we found in the Arctic Grayling diets may have been feeding directly on salmon carcasses or indirectly, for example, by feeding on insects that had been feeding on salmon carcasses. The high productivity of these riparian zones, especially along streams with high densities of salmon, probably contributes posi- tively to the success and survival of shrews, the tiniest of mammals, in this extreme northern environment. The skewed distribution of the number of shrews contained per Arctic Grayling is evidence that con- sumption of shrews is not a random event. We assessed whether Arctic Grayling consumption of shrews was random by comparing the observed frequency distri- bution of the number of shrews in Arctic Grayling stomachs to the expected distribution, assuming pre- dation to be random. We calculated expected frequen- sample TABLE |. Characteristics of shrews removed from the stomachs of two Arctic Grayling. Sorex spp 6 ©N! Location S. cinereus? 4.73 Elva Creek S. monticolus* SP) Elva Creek S. monticolus* 6.33 Little Togiak River S. cinereus® 7.42 Little Togiak River S. cinereus* 5.70 Little Togiak River Body dimensions? Specimen number 96-38-11 mm UWBM 74151 indeterminable UWBM 74152 indeterminable UWBM 74153 88-38-12 mm UWBM 74154 indeterminable UWBM 74155 ' 8I5N is the stable isotope of nitrogen and can be used to trace salmon-derived nutrients. ? Dimensions are total length, tail length, and hind foot length, in mm. > Identified both by dentition and cytochrome-b analyses. “Identified by only cytochrome-b analysis because of missing/digested dentition. 2004 MoOoRE AND KENAGY: CONSUMPTION OF SHREWS BY GRAYLING FiGurRE 1. Shrews removed from the stomachs of two Arctic Grayling, in various stages of digestion. Upper: Shrews from a 522 g Arctic Grayling sampled at Elva Creek (59°34’N, 159°05’W) — Sorex monticolus (UWBM 74152) left and Sorex cinereus (UWBM 74151) right. Lower: Shrews from a 591 g Arctic Grayling at Little Togiak River (59°36’N, 159°04’W) — Sorex cinereus (UWBM 74155) left, S. cinereus (UWBM 74154) center, and Sorex monticolus (UWBM 74153) right. cy distributions using the Poisson function in Matlab 5.313, assuming an average number of shrews per Arctic Grayling of 0.054 (5 shrews in 93 Arctic Gray- ling). We encountered fish with either two or three shrews in their stomachs, but no fish with only one shrew. Based on a random expectation for Arctic Gray- ling consumption, there was a 0.12 probability of sam- pling an Arctic Grayling with two shrews and only a 0.003 probability for three shrews. Thus we conclude that Arctic Grayling consumption is not random, and we speculate that this is due to individual differences in either the effectiveness or motivation of individual Arctic Grayling as shrew predators or scavengers. It may also be due to differences in the probability of shrews entering the feeding sites of specific Arctic Grayling. Although Arctic Grayling are typically considered to be specialists on aquatic and terrestrial insects (Arm- strong 1986), our observation indicates that some Arctic Grayling are opportunistic feeders, capable of consuming small mammals. Shrews, typically 4-8 g (Nagorsen 1996), represent a meal for Arctic Graylings that provides a large amount of energy compared to that of typical individual invertebrate food items. Con- sumption by Arctic Grayling of small mammals, in- cluding shrews, has been reported by several previous authors. Alt (1978*) reported an unspecified small number of shrews in diets of Arctic Grayling in the Fox River of western Alaska. De Bruyn and McCart (1974*) found that seven out of 136 Arctic Grayling contained a single shrew in the Firth River, Yukon, Canada. A European study of the grayling Thymallus thymallus indicated a 10% incidence of small mam- mals in the diet, particularly the shrew Sorex araneus, and the capture of these fish at night by bait angling suggested their ability as nocturnal hunters of semi- aquatic prey such as shrews (Teplov 1943). Miller (1946) reported that one out of 102 Arctic Grayling at Great Bear Lake, Canada, contained juvenile lem- mings (rodent) of an unspecified species in the diet. Reed (1964*) also found juvenile lemmings in two Arctic Grayling out of 1300 individuals sampled in the Tanana River drainage, Alaska. Terrestrial food sources often subsidize the diets of freshwater fish, and these food sources can include small mammals (for example, Brown Trout [Salmo trutta], Cochran and Cochran 1999; and Largemouth Bass [Micropterus salmoides|, Hodgson and Kinsella 1995). Although Arctic Grayling are traditionally con- sidered insectivores, our report demonstrates that they also opportunistically consume shrews that venture into freshwaters. Acknowledgments We thank the Alaska Salmon Project of University of Washington for logistic support and the National Science Foundation and ARCS foundation for finan- cial support. D. E. Schindler and M. D. Scheuerell provided insightful discussions and helped collect fish diets. M. D. Scheuerell and J. I. Jones prepared the shrew muscle samples fer isotopic analysis, which were run by D. Harris. We thank E. Waltari for molec- ular identification of the shrews and R. Rausch for advice on the observation and literature. 114 Documents Cited (marked * in text) Alt, K. T. 1978. Inventory and cataloging of sport fish and sport fish waters of western Alaska. Alaska Department of Fish and Game. Federal Aid in Fish Restoration, Annual Performance Report, 1977-1978. Project F-9-10, 19 (G-I- P): 36-60. de Bruyn, M., and P. J. McCart. 1974. Life history of the grayling (Thymallus arcticus) in Beaufort Sea Drainages in the Yukon Territory. /n Fisheries research associated with proposed gas pipeline routes in Alaska, Yukon and Northwest Territories. Edited by P. J. McCart. Canadian Arctic Study, Ltd., Calgary, Biological Report Series 15. 39 pages. Reed, R. J. 1964. Life history and migration patterns of Arc- tic grayling, Thymallus arcticus (Pallus), in the Tanana River drainage of Alaska. Alaska Department of Fish and Game Research Report Number 2: 8-30. Literature Cited Armstrong, R. H. 1986. A review of Arctic grayling studies in Alaska, 1952-1982. Biological Papers of the Univer- sity of Alaska, Number 23: 1-110. Ben-David, M., T. A. Hanley, and D. M. Schell. 1998. Fertilization of terrestrial vegetation by spawning Pacific salmon: the role of flooding and predator activity. Oikos 83: 47-55. Beneski, J. T., and D. W. Stinson. 1987. Sorex palustris. Mammalian Species 296: 1-6. Cederholm, C. J., M. D. Kunze, T. Murota, and A. Sibatani. 1999. Pacific salmon carcasses: essential contributions of nutrients and energy for aquatic and terrestrial ecosystems. Fisheries 24: 6-15. Cochran, P. A., and J. A. Cochran. 1999. Predation on a meadow jumping mouse, Zapus hudsonius, and a house mouse, Mus musculus, by brown trout, Salmo trutta. Canadian Field-Naturalist 113: 684-685. THE CANADIAN FIELD-NATURALIST Vol. 118 Demboski, J. R., and J. A. Cook. 2001. Phylogeography of the dusky shrew, Sorex monticolus (Insectivora, Soricidae): insight into deep and shallow history in northwestern North America. Molecular Ecology 10: 1227-1240. Hall, E. R. 1981. The mammals of North America. John Wiley, New York. 1175 pages. Hanski, I. 1986. Population dynamics of shrews on small islands accord with the equilibrium model. Biological Journal of the Linnean Society 28: 23-36. Hodgson, J. R., and M. J. Kinsella. 1995. Small mammals in the diet of largemouth bass, revisted. Journal of Fresh- water Ecology 10: 433-435. Kline, T. C., Jr., J. J. Goering, O. A. Mathisen, P. H. Poe, and P. L. Parker. 1990. Recycling of Elements Trans- ported Upstream by Runs of Pacific Salmon: I. 6 '°N and 6 C Evidence in Sashin Creek, Southeastern Alaska. Canadian Journal of Fisheries and Aquatic Sciences 47: 136-144. Miller, R. B. 1946. Notes on the Arctic grayling, Thymallus signifier Richardson, from Great Bear Lake. Copeia 1946: 227-236. Nagorsen, D. W. 1996. Opossums, shrews, and moles of British Columbia. University of British Columbia Press, Vancouver. 169 pages. Nelson, J. S. 1994. Fishes of the World. Third edition. John Wiley and Sons, New York. 600 pages. Schindler, D. E., M. D. Scheuerell, J. W. Moore, S. M. Gende, T. B. Francis, and W. J. Palen. 2003. Pacific salmon and the ecology of coastal ecosystems. Frontiers in Ecology and the Environment |: 31-37. Teplov, V. P. 1943. The significance of the common shrew (Sorex araneus L.) and some other vertebrates in the nutri- tion of Thymallus thymallus L. [in Russian]. Zoological Journal 22: 366-368. Received 9 June 2003 Accepted | June 2004 Unusual Behavior by Bison, Bison bison, Toward Elk, Cervus elaphus, and Wolves, Canis lupus L. Davip Mecu!, Rick T. McINTYRE2? and DouGLAS W. SMITH? 'U.S. Geological Survey, Northern Prairie Wildlife Research Center, 8711 - 37" Street, SE, Jamestown, North Dakota 58401 - 7317 USA; Mailing address: The Raptor Center, 1920 Fitch Avenue, University of Minnesota, St. Paul, Minnesota 55108 USA * National Park Service, Yellowstone Center for Resources, P. O. Box 168, Yellowstone National Park, Wyoming 82190 USA 3 Mailing address: General Delivery, Silver Gate, Montana 59081 USA Mech, L. David, Rick T. McIntyre, and Douglas W. Smith. 2004. Unusual behavior by Bison, Bison bison, toward Elk, Cervus elaphus, and Wolves, Canis lupus. Canadian Field-Naturalist 118(1): 115-118. Incidents are described of Bison (Bison bison) in Yellowstone National Park mauling and possibly killing a young Elk (Cervus elaphus) calf, chasing wolves (Canis lupus) off Elk they had just killed or were killing, and keeping the wolves away for extended periods. During one of the latter cases, the Bison knocked a wolf-wounded Elk down. Bison were also seen approach- ing wolves that were resting and sleeping, rousting them, following them to new resting places and repeating this behavior. These behaviors might represent some type of generalized hyper-defensiveness that functions as an anti-predator strategy. Key Words: Bison, Bison bison, Wolt, Canis lupus, Elk, Cervus elaphus, Yellowstone National Park. Bison (Bison bison) are known to be aggressive to directed the capturing and radio-tagging of the Wolves, other ungulates including Elk and even to kill Elk allowing them to be located and observed. (Cervus elaphus) calves (Chapman 1937: 148; Rush 1942: 225; McHugh 1958: 143; Mahan 1977). Little Observations is known about the causes and circumstances of this 1. 30 May 1995 — Bison Attack Elk Calf aggressiveness except for the scattered reports docu- A recently born Elk calf that separated from its menting it. Bison also defend themselves aggressively mother became mixed up in the middle of a Bison herd when attacked by Wolves (Canis lupus) (Carbyn and southwest of the Lamar Picnic Area. During the early Trottier 1988; Carbyn et al. 1993; MacNulty 2002), stages of this observation, filmmaker Ray Paunovich although Bison have been reported to be indifferent to saw a Bison knock the calf down and then repeatedly close, but non-attacking, Wolves (Catlin 1876-1: 254 butt it to the ground. cited by McHugh 1938; Goodwin 1939: 369 cited by RM watched a group of Bison encircle the calf, then McHugh 1938; and Soper 1941: 403; Carbyn et al. lick and sniff it. The calf got up and tried to walk off, 1993). but several Bison chased it. One Bison butted the calf in We also have observed the same types of behavior of _ the side and knocked it down. Twelve additional Bison Bison toward wolves both when being attacked and came over to sniff it. The calf got up and ran off, but not being attacked in Yellowstone National Park. How- was chased and knocked down again by the Bison. ever, we have observed other types of Bison aggres- As long as the calf remained down or stood still, the _ siveness toward Elk and toward Wolves that has not — Bison just sniffed and licked it, but whenever it ran off, _ been reported, and we have seen another possible case they chased it and knocked it down. This sequence _ of Bison killing an Elk calf. Because our observations __ took place at least four times. A cow Bison later butted | considerably extend what is known about Bison aggres- _ the Elk calf several times as it lay on the ground. Then siveness and because observation of any ungulates chas- _ the herd walked away. The calf was motionless as the | ing predators are rare (Berger 1979), we report them Bison moved off and was possibly dead from all the below and propose an explanation for the possible adap- _ butting. RM then left, but Paunovich thought he saw tiveness of the aggressiveness. some movement from the calf after RM left. Whether __ Both Elk and Bison have inhabited the park for many the calf survived is unknown. _ decades, but Wolves were reintroduced to the ecosys- 2. 6 June 2001 — Bison Displace Wolves from Kill Bea an P73) and 1996 (Bangs et al. 1998). Wolves in "at 0529 hrs, RM spotted a fresh adult Elk kill with | Yellowstone prey primarily oo Elk (Mech et al. 2001) four yearling Druid Peak pack Wolves next to it south- _ but do kill Bison as well (Smith et al. 2000). RM made east of the Lamar Valley Picnic Area. The carcass was { a peo auons in ns northeastem En of Yellowstone intact and the Wolves must have just killed it. Three ational Park through binoculars and spotting scopes Wolves were about to start feeding. _ with 60x power and recorded them by tape recorder: A Bison bull walked up to the carcass and sniffed its _ LDM independently observed and recorded with pad head. The Wolves went to the rear of the carcass and and pencil the observation on 20 March 2002. DWS HS 116 started tugging on that section as the bull continued to sniff the head area. This seemed to be the Wolves’ first feeding on the kill. Three more Bison bulls approached the carcass. One Wolf at the carcass walked off a few steps with a tucked tail, then came right back and snarled at the nearest bull. The bulls walked toward the three Wolves, and the Wolves backed off. Two of the bulls sniffed the Elk’s head. A Wolf sneaked back in and fed on the rear end. Soon six bulls were at the site. They had a stand-off with the three Wolves at 0537 hrs. When one Wolf tried to approach the carcass, a bull ran at it, and the Wolf backed off. The other bulls walked toward the rest of the Wolves, and they scattered. The six bulls kept the three Wolves away from the carcass through 0554 hrs. At that time, the bulls left the site, and the Wolves moved to the carcass and fed. The three Wolves were still feeding on the kill when RM left at 0615 hrs. In Wood Buffalo National Park, Alberta, mixed herds of Bison were observed forcing wolves away from wolf-killed Bison calves about 1 and about 4 months old (L. N. Carbyn, personal communication). 3. 3 March 2002 — Bison Attack Wounded Elk Calf This incident was observed and video-taped by cine- matographer Shane Moore in the Blacktail area of Yellowstone National Park. The tape was viewed by RM who transcribed the details summarized below. The incident began when a herd of Bison ran 0.15 km downhill to a 9-month-old Elk calf with blood on its neck. Several Coyotes (Canis latrans) were near- by. The Bison herd gathered near the Elk, and over a 4-hr period, the Bison sniffed, licked, chased, and but- ted the Elk as the animal tried to seek refuge from the Coyotes by running into the Bison herd. The Bison but- ted the calf throughout the episode on 21 occasions during 40 minutes of videotaping, usually in the rear or side, and once knocked it into the air. At times the Elk was bedded or lying on its side when the Bison butted it. Partway through the video, it was apparent that the Elk’s abdomen was wounded, and an organ was hang- ing out. Moore believed that wound resulted from when he saw a Bison charge the Elk at full speed and hit it very hard in the side. Although the rest of the Bison herd then blocked the view, the calf stayed down for 20 minutes and when it arose, its abdomen had been ripped open. Toward the end of the four-hour observation, the Bison drifted away from the wounded and bedded calf. As the calf continued to weaken, two Coyotes attacked it. Three additional Coyotes joined the attack, and the five Coyotes soon killed the calf. 4. 20 March 2002 — Bison Displace Wolves from Kill Eleven members of the Druid Peak pack (1 1-months old and 23-months old; no adults) had been chasing Elk in the Hellroaring Creek area of Yellowknife Na- tional Park. At about 0931 hrs, the Wolves targeted a single cow Elk and pulled her down. A herd of about THE CANADIAN FIELD-NATURALIST Vol. 118 39 Bison rushed to the downed Elk, chased the Wolves off, and surrounded the Elk tightly (0933 hrs). Sud- denly the Elk leaped up. Two Wolves approached the Elk and nipped at it. The Elk ran through the Bison herd, and the two Wolves pursued it. After about 30 m the Elk fell. The Wolves were on it again for about 30 sec when the Bison chased them off again (0936 hrs). Later the Elk arose and almost immediately collapsed. The Elk arose a third time, ran a short distance and col- lapsed. She later tried to stand again but could only get up on her hind legs. A Bison butted the Elk’s rear end and knocked her down, and the Bison herd surrounded the Elk again and kept the Wolves away. Each time a Wolf approached, a Bison would chase it 15-30 m. After a few minutes, the Wolves headed away up a hill and bedded (0946 hrs). The Elk, while surrounded by Bison, kept raising her head and trying to get up. Usually the Bison stood closely around her but sometimes jumped back a few meters. The Bison remained tightly around the Elk from about 0939 hrs on. By about 1135 hrs, the Elk’s head was no longer up and her body lay flat; she seemed dead. Ravens landed on or next to her, although the Bison still surrounded her closely. Bison licked or sniffed (we could not determine which) the carcass intermittently for several minutes. The Wolves had returned at 0955 hrs and hung around 15-60 m from the Bison herd, and at various times some tried to reach the downed Elk. Each time, one or more Bison would skirmish with them and try to drive them off. About 1253 hrs, the Bison started moving away from the Elk, and by 1305 hrs Wolves moved in. However, the Bison quickly returned and ran the Wolves off. This happened several times when Bison were both east and west of the Elk. Eventually, however, the Bison all grouped east of the Elk, and the 11 Wolves began to feed. The Bison then charged the Wolves and ran them off. Then the Wolves returned and chased the Bison away. In the next few minutes there were two addition- al standoffs at the carcass, and both times the Bison chased away the Wolves. Such skirmishes continued as Wolves and Bison surged toward the Elk and each other, but gradually the Bison began to head a few meters farther east, and the Wolves became bolder. By 1313 hrs, the Wolves controlled the carcass and remained there and fed, while the Bison drifted off eastward. By 1350 hrs most of Wolves were done feeding and slept on a hill above the carcass, although individual Wolves fed later as well. 5. 22 March 2002 — Bison Roust Sleeping Wolves In the same general area as observation 4 was made, eight of the 11 Druid Peak Pack Wolves seen on 20 March were sleeping at 0909 hrs when six Bison lead- ing a larger herd (probably the same 39 seen there two days earlier) approached them. The Wolves arose, moved 100 m, and lay down again. Two minutes later, the Bison approached them to within 3 m, and the 2004 Wolves again moved off 100 m and lay down. At 0918 hrs, a Bison approached one of the Wolves, which then confronted it; the other Wolves joined in harassing the Bison for 30 seconds. The Wolves then left at 0920 hrs and moved 90° and a few hundred meters and lay down on a rocky ridge. Eventually the Bison followed and rousted the Wolves from there. Several more times the Bison followed the Wolves and rousted them out of their beds. 6. 16 April 2002 — Bison Bulls Displace Wolves from Kall At 0826 hrs, RM saw 11 Druid pack Wolves (the breeding male, two yearlings, and eight 1 1-month-old pups) chase Elk about 1.6 km west of the Lamar Ran- ger Station. They pulled down a cow Elk at 0858 hrs. A few Bison bulls started moving toward them from the north. Two Bison ran into the site and chased the Wolves off. A yearling and two pups came right back to the carcass. One of the bulls returned to the site and chased off one of the Wolves. At 0900 hrs several Wolves were feeding as four Bison bulls stood near the carcass. Two bulls charged the Wolves and drove them off, then stood close to the carcass. At that point, five Wolves watched the carcass from 35 m away. The two bulls charged the breeding male and another Wolf, which fled eastward. The other Wolves ran to the kill and fed. The breeding male cir- cled around and joined them. A bull then walked to the carcass, and the feeding Wolves backed off a meter or so. The other nearby Bison approached the site. That bull charged the Wolves, and they ran off but came right back. Both bulls then stood next to the Elk carcass and sniffed it. The bigger bull charged the Wolves, but as before the Wolves soon stopped, and the breeding male and three others returned to the site. The two additional Bison bulls came toward the car- cass from the west. One of the bulls shook his head, and some Wolves backed off a step or two. Other Wolves stood their ground and faced the bull which was about 6 m away. The bull shook his head again, but the Wolves continued to feed and ignored him. The breeding male Wolf stepped away and ate some snow. When he returned to the carcass, a bull chased him off. The Wolf came right back. The same bull, which was the largest and most aggressive of the four bulls there, then drove all the Wolves off. The breed- ing male Wolf returned, and the big bull charged him. A second bull was standing a meter or so from the carcass. The big bull chased the breeding male Wolf southward, then returned to the site. The two bulls kept the Wolves from the kill until the bulls walked a short distance away at 09:12; the Wolves immedi- ately ran in. All 11 fed or were near the site. As the two bulls walked off to the north a pup followed them. Around 0920 hrs, three bulls approached the car- cass from the northeast as the Wolves continued to MEcH, MCINTYRE, AND SMITH: UNUSUAL BEHAVIOR BY BISON Ki, feed. One pup headed westward, but the others stood their ground. The bulls turned back before they reached the site. A fourth bull chased a Wolf to the north of the carcass. One bull then ran in toward the carcass, but the Wolves countercharged, drove him away, and resumed feeding. A fifth bull was north of the carcass but did not yet approach it. A sixth bull approached the kill, and a pup circled around to his rear and followed him. Another pup joined the first pup behind the bull. Two more of the original bulls approached the carcass from the north. The lead bull walked by the north side of the carcass with a raised tail. As another bull approached the site, the seven feeding Wolves backed off. At 0927 hrs three bulls remained at the carcass keeping the Wolves away. The breeding male Wolf stood nearby. He and a pup left, and the breeding male bedded north of the site at 0929 hrs. Eight other Wolves stood near the carcass. At 0930 hrs two bulls started sparring with each other and began to drift away from the carcass. The breeding male Wolf and a yearling walked to the kill and fed. In the next minute other Wolves joined them. All six bulls remained away from the site, and the Wolves fed unmolested. Discussion The aggressiveness shown by Bison toward Elk and toward non-attacking Wolves is puzzling but might be some kind of hyperdefensive or hyperaggressive behav- ior. Predator harassment by several species of ungu- lates has been documented, and various explanations have been offered (summarized by Berger 1979), each involving ultimate reduction of predation risk. These explanations do not account for Bison attacking Elk, however. The most common element in our observa- tions and those reported earlier of Bison attacking Elk was the presence of a weak or downed Elk and the butting, sniffing, and mobbing of it. In the Elk inci- dents involving Wolves, perhaps the presence of the predators was only incidental and made the Bisons’ aggressiveness appear directed at keeping Wolves away from the carcasses or injured animals. In other words, our observations might involve two phases of hyperaggressive behavior: (1) a general aggressiveness toward various intruders, perhaps as generalized anti-predator behavior that causes the Bison to approach intruders, and (2) continued aggressive- ness toward intruders that do not flee, e.g., wounded animals or animals seeking refuge in the Bisons’ herd because these animals persist in remaining nearby. This would also include Wolves that are trying to feed on carcasses near the Bison herd. Bison are so large and powerful that when a herd acts aggressively, it can charge and attack any animal with impunity, includ- ing Wolves and Grizzly Bears (Ursus arctos). Thus it may be generally advantageous for them to advertise this ability by harassing intruders persistently without making fine distinctions as to type. 118 Acknowledgments This study was supported by the Biological Res- ources Division, U.S. Geological Survey, and the Yellowstone Center for Resources. Literature Cited Bangs, E. E., S. H. Fritts, J. A. Fontaine, D. W. Smith, K. M. Murphy, C. M. Mack, and C. C. Niemeyer. 1998. Status of gray wolf restoration in Montana, Idaho, and Wyoming. Wildlife Society Bulletin 26: 785-798. Berger, J. 1979. “Predator harassment” as a defensive strat- egy in ungulates. American Midland Naturalist 102: 197- 199. Carbyn, L. N., and T. Trottier. 1988. Descriptions of wolf attacks on bison calves in Wood Buffalo National Park. Arctic 41: 297-302. Carbyn, L. N., S. M. Oosenbrug, and D. W. Anions. 1993. Wolves, bison and the dynamics related to the Peace-Atha- basca Delta in Canada’s Wood Buffalo National Park. Circumpolar Research Series (4). Canadian Circumpolar Institute, University of Alberta. Catlin, G. 1876. Illustrations of the manners, customs, and condition of the North American Indians. Two volumes. Chatto & Windus, London, U.K. 530 pages. Chapman, W., and L. Chapman. 1937. Wilderness wan- derers. Charles Scribner’s Sons, New York. 318 pages. THE CANADIAN FIELD-NATURALIST Vol. 118 Goodwin, G. C. 1939. The bison. Chapter XX in North American Big Game. Edited by Alfred Ely, H. E. Anthony, and R. R. M. Carpenter. Charles Scribner’s Sons, New York. 533 pages. MacNulty, D. R. 2002. The predatory sequence and the influ- ence of injury risk on hunting behavior in the wolf. Master’s thesis, University of Minnesota, St. Paul, Minnesota. Mahan, B. R. 1977. Harassment of an elk calf by bison. Canadian Field-Naturalist 91: 418-419. McHugh, T. 1958. Social behavior of the American buffalo (Bison bison bison). Zoologica 43: 1-41. Mech, L. D., D. W. Smith, K. M. Murphy, and D. R. MacNulty. 2001. Winter severity and wolf predation on a formerly wolf-free elk herd. Journal of Wildlife Manage- ment 65: 998-1003. Rush, W. M. 1942. Wild animals of the Rockies. Harper and Brothers, New York. 296 pages. Smith, D. W., L. D. Mech, M. Meagher, W. E. Clark, R. Jaffe, M. K. Phillips, and J. A. Mack. 2000. Wolf-bison interactions in Yellowstone National Park. Journal of Mam- malogy 81: 1128-1135. Soper, J. D. 1941. History, range and home life of the north- ern bison. Ecological Monographs 11: 347-412. Received 11 August 2003 Accepted 4 June 2004 Notes Morphology of Female Woodland Caribou, Rangifer tarandus caribou, in Saskatchewan W. JAMES RETTIE Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada Present address: Northeast Science & Information Section, Ontario Ministry of Natural Resources, Ontario Government Complex, Highway 101 East, P.O. Bag 3020, South Porcupine, Ontario PON 1HO Canada Rettie, W. James. 2004. Morphology of female Woodland Caribou, Rangifer tarandus caribou, in Saskatchewan. Canadian Field- Naturalist 118(1): 119-121. I obtained morphological measurements from captured female Woodland Caribou in central Saskatchewan. I found that only girth was a good predictor of body mass in adult animals. I also determined that Woodland Caribou from Saskatchewan, though similar in size and mass to alpine Woodland Caribou in Yukon, are larger than migratory Woodland Caribou and smaller than forest dwelling Woodland Caribou from western Alberta and Yukon. Key Words: Woodland Caribou, Rangifer tarandus caribou, morphology, Saskatchewan. Body sizes and body masses of subspecies and pop- ulations of Rangifer tarandus (Eurasian Reindeer and North American Caribou) vary considerably (e.g., Dauphiné 1976; Thomas 1982; Reimers 1983; Chan- McLeod et al. 1995) in response to a wide range of cli- matic and other environmental conditions (Skogland 1983; Reimers 1983). In North America, there is also a high degree of morphological variation within the Woodland Caribou subspecies (R. t. caribou). The best described populations of Woodland Caribou are those in Yukon (Gauthier and Farnell 1986; Kuzyk et al. 1999) and in Québec and Labrador (Parker 1981; Huot 1989). However, the subspecies has a more or less continuous distribution across northern Canada and the variations in climate, topography, and plant and animal communities across the range of the subspecies might be expected to favour different body sizes and masses in different areas. In particular, Skogland (1983) argued that morphology was density-dependent and related to the factors limiting population growth. Both Skogland (1983) and Chan-McLeod et al. (1999) reported that female R. tarandus that had reproduced successfully were significantly lighter the following winter than those that had not. Here, I present the first description of Woodland Caribou from central Saskat- chewan and a discussion of their morphology relative to limiting factors. Study Area and Methods As part of research into population dynamics (Rettie and Messier 1998) and behaviour (Rettie and Messier 2000, 2001), I captured 43 female Woodland Caribou in central Saskatchewan (approximately 54° N to 55°30’ N and 104° W to 109° W). All animals were captured in winter (dates between 13 December and 14 March) between March 1992 and January 1995. At the time of capture I used a flexible steel tape to measure the following parameters from most animals: total body length, body length to base of tail, girth (all to the nearest cm), mandible, and metatarsal lengths (to the nearest 5 mm). Methods followed those of Dauphiné (1976), modified for application to live animals. I mea- sured body mass to the nearest kilogram for ten animals using an electronic load scale (Senstek, Saskatoon, Saskatchewan) suspended from a tripod or beneath a helicopter. I also extracted a tooth from most animals for aging. Further details on capture and aging appear in Rettie and Messier (1998). Animal capture and handling procedures followed animal care protocol number 920092 of the University of Saskatchewan. Using natural log transformed values from the eight adult animals for which I had complete sets of data as well as body mass, I applied a stepwise multiple linear regression to assess the relationship of body measure- ments to body mass (p = 0.05 to enter, p = 0.10 to exit) and then applied the resulting equation to the morphometric data for all animals. I then tested for differences in calculated body mass for adult females that were accompanied by a calf at time of capture and those that were without a calf. All statistical analyses were conducted using SPSS for Windows Version 10.0.7 (SPSS Inc. 2000). Results and Discussion I had morphometric data from 34 adult animals (>31 months old at time of capture) and from three yearlings (19-22 months old at time of capture). The summary statistics for all measurements appear in 119 120 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |: Morphological measurements and actual and calculated body mass of adult female Woodland Caribou in central Saskatchewan. Parameter n Actual body mass 9 Calculated body mass” 32 Body length to base of tail 33 Total body length 33 Metatarsal length 30 Shoulder to hoof tip 15 Girth 32 Mandible length 32 “ Body mass kg calculated using the equation arising from the regression analysis: e mean + | sd range 131 + l6kg 96 -156kg LS eye like 96 —-160kg LOLS =p isiem 165 -—217cm 200)=9 = lsicm 177. —230cm 43.0 + 14cm 41.0 — 46.0 cm Il). ee W/m 102. —126cm AS) eee ikem 115) —154cm S14 ei ple8 cm 28.0 — 35.0cm 1.76 In Girth — 3.78 TABLE 2: Morphological measurements and actual body mass of yearling female Woodland Caribou in central Saskatchewan. Parameter n Actual body mass ] Body length to base of tail 3} Total body length 3 Metatarsal length 3 Girth 3 Mandible length 3 Tables 1 (adult animals) and 2 (yearling animals). When compared to Yukon Woodland Caribou, the animals in Saskatchewan are almost as tall at the shoul- der (115 cm vs. 116 cm), are the same girth (129 cm), but are not as long (191 cm vs. 206 cm) as the alpine animals measured by Kuzyk et al. (1999). Saskatchewan caribou are smaller than forest dwelling Woodland Caribou from Yukon (Kuzyk et al. 1999) and from western Alberta (Gauthier and Farnell 1986) in all comparable parameters. From the regression analysis, I concluded that only girth was significantly related to body mass (p =0.04, r? = 0.78). I used the resulting equation, where girth is measured in cm: calculated body mass (kg) = e(!-76/Girth — 3.78 to calculate body mass for all adult animals for which I had girth measurements. The calculated body mass values appear in Table 1. Figure 1 shows the relationship between girth and body mass and its 95% prediction interval. Untransformed data are presented in the figure while analyses were based on natural log transformed data. The wide prediction interval in Figure | suggests that further sampling is required to reduce uncertainty in the girth-body mass relationship. The relationship is particularly weak for smaller girths where the relationship is influenced by a single obser- vation. Though the summarised measurements present a reference point for future studies on morphological variation in Woodland Caribou, the prediction of body mass from body measurement and the equation pre- mean + | sd range 98 kg GQ es. 3) con 170 —1175 cm 185i lem 182 —1188 cm 41.7 + 0.6cm 41.0 -— 142.0cm See eee Src 110 —1116cm 28.2 + 2.0cm 26.0 — 130.0cm sented should be made cautiously (Cattet et al. 1997). Finally, I did not detect a difference in calculated body mass between adult animals with and without a calf-at- heel at time of capture (¢-test, t= 0.30, df = 30, p = 0.76). Skogland (1983) argued that body sizes in R. tar- andus were density-dependent, the larger body size 200 body mass = @(1-76!n Girth - 3.78) *=0.78 180 r S52 P=0.004 - 160 aa Body mass (kg) XD oO 110 120 130 140 150 Girth (cm) FiGurE 1. Relationship between girth (cm) and body mass (kg) for adult female Woodland Caribou in Saskatchewan. Solid line represents the equation derived through multiple linear regression; dashed lines represent the 95% prediction interval of the equation. Filled squares represent data used to derive the equation. 2004 in North American Caribou being attributed to their low densities, a consequence of limitation by predation rather than by food resources. Amongst Woodland Caribou populations, the George River Caribou Herd in Quebec and Labrador is food limited and contains the smallest individuals (Parker 1981; Huot 1989). The body sizes of Woodland Caribou in Saskatchewan are larger than George River Herd animals and support the argument that Saskatchewan populations are not food limited (Rettie and Messier 1998). As with the larger animals in Caribou populations in western Al- berta and Yukon (Edmonds 1988; Kuzyk et al. 1999), predation is the likely proximate limiting factor for Saskatchewan Woodland Caribou populations (Rettie and Messier 1998). Skogland (1983) also suggested that predator in- duced constraints on lifetime reproductive success led to larger body size in North American Caribou by de- laying reproductive maturity in favour of increased somatic growth. However, yearling female Woodland Caribou in Saskatchewan were not only pregnant (Rettie and Messier 1998) but larger than both adult Reindeer (Skogland 1983) and adult migratory Wood- land Caribou (Huot 1989). Relative to observations in Norway, there does not appear to be a trade off being made between reproduction and somatic growth. In- stead, Saskatchewan Woodland Caribou appear able to mature as yearlings and still achieve large adult body sizes. My failure to detect differences in body mass between reproductive classes may represent a lack of difference, result from a poor predictive ability of my equation for body mass, or be a consequence of inter- annual variation as reported by Chan-McLeod et al. (1999). Literature Cited Cattet, M. R. L., S. N. Atkinson, S. C. Polischuk, and M. A. Ramsay. 1997. Predicting body mass in polar bears: is morphometry useful? Journal of Wildlife Management 61: 1083-1090. Chan-McLeod, A. C., R. G. White, and D. E. Russell. 1995. Body mass and composition indices for female barren- ground caribou. Journal of Wildlife Management 59: 278-291. NOTES Chan-McLeod, A. C., R. G. White, and D. E. Russell. 1999. Comparative body composition strategies of breeding and nonbreeding female caribou. Canadian Journal of Zoology 77: 1901-1907. ; Dauphiné, T. C. 1976. Biology of the Kaminuriak population of barren-ground caribou. Part 4. Growth, reproduction and energy reserves. Canadian Wildlife Service Report Series Number 38. Edmonds, E. J. 1988. Population status, distribution, and movements of woodland caribou in west central Alberta. Canadian Journal of Zoology 66: 817-826. Gauthier, D. A., and R. F. Farnell. 1986. Comparison of caribou physical characteristics from Yukon and neigh- boring caribou herds. Rangifer Special Issue 1: 131-135. Huot, J. 1989. Body composition of the George River caribou (Rangifer tarandus caribou) in fall and late winter. Cana- dian Journal of Zoology 67: 103-107. Kuzyk, G. W., M. M. Dehn, and R. S. Farnell. 1999. Body- size comparisons of alpine- and forest-wintering wood- land caribou herds in the Yukon. Canadian Journal of Zoology 77: 1017-1024. Parker, G. R. 1981. Physical and reproductive characteristics of an expanding woodland caribou population (Rangifer tarandus caribou) in northern Labrador. Canadian Jour- nal of Zoology 59: 1929-1940. Reimers, E. 1983. Growth rates and body size differences in Rangifer, a study of causes and effects. Rangifer 3: 3-15. Rettie, W. J., and F. Messier. 1998. Dynamics of woodland caribou populations at the southern limit of their range in Saskatchewan. Canadian Journal of Zoology 76: 251-259. Rettie, W. J., and F. Messier. 2000. Hierarchical habitat selection by woodland caribou: its relationship to limit- ing factors. Ecography 23: 466-478. Rettie, W. J., and F. Messier. 2001. Range use and movement rates of woodland caribou in Saskatchewan. Canadian Journal of Zoology 79: 1933-1940. Skogland, T. 1983. The effects of density dependent res- ource limitation on size of wild reindeer. Oecologia 60: 156-168. SPSS. 2000. SPSS for windows Version 10.0.7 SPSS Inc., Chicago. Thomas, D. C. 1982. The relationship between fertility and fat reserves of Peary caribou. Canadian Journal of Zoology 60: 597-602. Received 3 December 2002 Accepted 21 May 2004 122 THE CANADIAN FIELD-NATURALIST Vol. 118 Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in Southeastern Idaho LisA M. Coss! and VINCENT A. CoBB? Department of Biological Sciences, Idaho State University, Pocatello, Idaho 83209 USA Current Addresses: 'Department of Biology, Cumberland University, Lebanon, Tennessee 37087 USA *Department of Biology, Middle Tennessee State University, Murfreesboro, Tennessee 37132 USA Cobb, Lisa M., and Vincent A. Cobb. 2004. Occurrence of parasitoid wasps, Baeus sp. and Gelis sp., in the egg sacs of the wolf spiders, Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in southeastern Idaho. Canadian Field- Naturalist 118(1): 122-123. Egg sacs of the wolf spiders Pardosa moesta and Pardosa sternalis were sampled for two years during June to September in southeastern Idaho. Parasitoid wasps, Baeus sp. (Sceleonidae) and Gelis sp. (Ichneumonidae), were observed in the egg sacs of both Pardosa species. Of 322 egg sacs examined, 14.6% were parasitized. Parasitism of egg sacs occurred throughout most of the Pardosa egg sac-carrying season. Key Words: Wolf Spider, Pardosa moesta, P. sternalis, egg sac, predation, parasitic wasps, Baeus sp., Gelis sp., parasitoid, reproduction. Parasitic wasps are well known predators of spiders and may serve as either ectoparasites (e.g., externally feeding on the body) or endoparasites (e.g., feeding on eggs and developing young within egg sacs) (Foelix 1982). Larval endoparasites that consume spiders eggs are classified as parasitoids (Roberts and Janovy 2000) with the most speciose family of parasitoid wasps be- ing the Ichneumonidae (Godfray 1994). To parasitize a spider egg sac, wasps insert their ovipositor into the wall of the egg sac and lay eggs which develop into larvae and predate the spider eggs (Austin 1985). Female wolf spiders (Lycosidae) have the conspic- uous behavior of carrying their egg sacs attached to their spinnerets. Although it would appear that such behavior could provide substantial maternal care, para- sitism of egg sacs still occurs. The wasps Baeus (Scel- ionidae) and Gelis (Ichneumonidae) have been found in previous studies to be common parasitoids of Lyco- sidae egg sacs (Kaston 1948; Edgar 1971a; Austin 1985). Wolf spiders in the genus Pardosa are common ground-dwelling spiders throughout much of the north- ern hemisphere (Edgar 1971b; Dondale and Redner 1990; Buddle et al. 2000). Their reproductive period may last several months (Edgar 1971b; Cobb 1992; Buddle 2000) and some species may produce multi- ple clutches during a single season (Edgar 1971b; Wolff 1981). The main focus of this study was an investigation of the reproductive ecology of P. moesta (Banks) and P. sternalis (Thorell) in eastern Idaho. Cobb (1992) sug- gested that in southeastern Idaho both species have a tendency to produce multiple clutches within a season. During the study, P. moesta and P. sternalis females and their egg sacs were collected between one and five times per month from June through September of 1990 and 1991. All collections occurred in a two hectare meadow near the base of Scout Mountain (Caribou Na- tional Forest, Bannock County, Idaho) at an elevation of approximately 2000 m. During the late spring and early summer the meadow was marshy, fed by a moun- tain stream. However, in late summer and early fall the meadow was dry and had been grazed by cattle. For each spider and corresponding egg sac we collected a variety of morphometric data, which included female mass, clutch mass, and clutch size. While examining the egg sacs we discovered several of them were para- sitized. Identifiable parasitoids of P. moesta and P. stenalis egg sacs were wasps of the families Scelionidae and Ichneumonidae. P. M. Marsh, of the USDA Agricultural Research Station, Beltsville, Maryland, identified the scelionid specimens as Baeus sp., and the ichneumonids at Gelis sp. All but two of the identifiable parasitoids for both Pardosa species were Baeus sp. Two egg sacs of P. sternalis were found with a single Gelis sp. each. The Gelis were collected on 23 June and 22 July 1991. Pupal parasitoids inside the egg sacs could not be identified to species. Because not all individ- ual parasitoids could be identified, other parasitoid genera may have been present in the egg sacs. Of 134 egg sacs for P. moesta and 188 egg sacs for P. sternalis, 47 were infected with parasitoids (14.6% parasitism overall). Within these infected egg sacs 281 parasitoids were counted. Parasitism rate was 12.7% for P. moesta and 16% for P. sternalis (Table 1). Egg sacs were found parasitized from June to Sep- tember for P. moesta and P. sternalis. During mid and late May 1990, a sample of egg sacs was collected from P. sternalis but none contained parasitoids. All of the P. sternalis egg sacs in the May collection contained eggs only and no spider instar stages, indicating egg | | 2003 TABLE |. Number of parasitized egg sacs for Pardosa moesta (n = 134) and Pardosa sternalis (n = 188) from southeastern Idaho. Data are represented as number of egg sacs with para- sitoids (number of egg sacs sampled). P. moesta P. sternalis Month 1990 199] 1990 199] June 0 (23) = 8 (37) Le CS) July 1 (40) 1 (9) 7 (45) Be (60) August 12 (25) = 6 (15) IP (G21) September 0 (27) 3(10) Dea alisia) Die (i) sac formation had begun only recently. The number of parasitoids per egg sac varied considerably. Mean (+ SE) number of parasitoids per egg sac for P. moes- ta was x = 5.1 + 1.4 (range = 2 - 15) in 1990 and x = 6.5 + 2.3 (range = 1 - 12) in 1991. For P. sternalis, the number of parasitoids per egg sac was x = 5.7 + 1.6 (range = | - 37) in 1990 and x = 8.4 + 4.6 (range = 1 - 35) in 1991. This study indicates that parasitism occurred through- out the egg carrying season and was highest in Aug- ust 1990 and September 1991. Interestingly, these high parasitism rates correspond to times when Pardosa in this region are producing smaller egg clutches (Cobb 1992). Several species of Pardosa have their greatest reproductive output early in the reproductive season and produce smaller egg sacs and fewer eggs later in the season (Eason 1969). Potentially, these higher para- sitism rates in the late summer could have resulted in selection for greater spring reproduction in Pardosa. Although parasitism has been documented for several species of Pardosa (Eason et al. 1967), recorded levels of parasitism are uncommon and variable. Eason (1969) indicated egg parasitism rates of less than one percent in P. lapidicina in Arkansas, while Edgar (1971a) ob- served levels that ranged from 3% to 35% in Scotland. Our data indicate variable and sometimes high levels of parasitism as well. Such high levels of parasitism have been shown to reduce the number of spiders in the field (van Baarlen et al. 1994) as well as the re- cruitment of young (Edgar 1971a). Although we have documented seasonal variation in parasitism rate, we do not know if these Pardosa populations are being negatively impacted by parasitoids. Acknowledgments We thank P. M. Marsh of the U.S. Department of Agriculture for identifying parasitoid wasps and J. Redner of the Canadian Department of Agriculture for the initial identification of spider specimens. Funding NOTES for this project was provided partly by a grant to L. Cobb from the Idaho State University Graduate Res- earch Committee. Literature Cited Austin, A. D. 1985. The function of spider egg sacs in rela- tion to parasitoids and predators with special reference to the Australian fauna. Journal of Natural History 19: 359- 370. Buddle, C. M. 2000. Life history of Pardosa moesta and Par- dosa mackenziana (Araneae: Lycosidae) in central Alberta, Canada. Journal of Arachnology 28: 319-328. Buddle, C. M., J. R. Spence, and D. W. Langor. 2000. Succession of boreal forest spider assemblages following wildfire and harvesting. Ecography 23: 424-436. Cobb, L. M. 1992. Reproductive tactics of two sympatric species of Pardosa (Araneae, Lycosidae). M.S. thesis. Idaho State University, Pocatello, Idaho. 59 pages. Dondale, C. D., and J. H. Redner. 1990. The insects and arachnids of Canada. Part 17. The wolf spiders, nursery- web spiders, and lynx spiders of Canada and Alaska (Araneae: Lycosidae, Pisauridae, and Oxyopidae). Agri- culture Canada Publication 1856. Eason, R. R. 1969. Life history and behavior of Pardosa lapidicina Emerton (Araneae: Lycosidae). Journal of the Kansas Entomological Society 42: 339-360. Eason, R. R., W. B. Peck, and W. H. Whitcomb. 1967. Notes on spider parasites, including a reference list. Journal of the Kansas Entomological Society 40: 422-434. Edgar, W. D. 1971a. Aspects of the ecology and energetics of the egg sac parasites of the wolf spider Pardosa lugu- bris (Walckenaer). Oecologia 7: 155-163. Edgar, W. D. 1971b. The life-cycle, abundance and seasonal movement of the wolf spider, Lycosa (Pardosa) lugubris, in central Scotland. Journal of Animal Ecology 40: 303-322. Foelix, R. F. 1982. Biology of spiders. Harvard University Press, Cambridge, Massachusetts. Godfray, H. C. J. 1994. Parasitoids: Behavioral and evolu- tionary ecology. Princeton University Press, Princeton, New Jersey. Kaston, B. J. 1948. Spiders of Connecticut. State Geology and Natural History Survey of Connecticut Bulletin 70. 1020 pages. Roberts, L. S., and J. Janovy Jr. 2000. Gerald D. Schmidt and Larry S. Robert’s Foundations of Parasitology. 6" edi- tion. McGraw-Hill, Boston, Massachusetts. Van Baarlen, P. 1994. Eggsac parasitism of money spiders (Araneae: Linyphiidae) in cereals, with a simple method for estimating percentage parasitism of Erigone spp. eggsacs by Hymenoptera. Journal of Applied Entomology 118: 217-223. Wolff, R. J. 1981. Wolf spiders of the genus Pardosa (Ara- neae: Lycosidae) in Michigan. Great Lakes Entomologist 14: 63-68. Received 4 March 2002 Accepted 14 June 2004 124 THE CANADIAN FIELD-NATURALIST Vol. 118 The Northern True Katydid, Pterophylla camellifolia (Orthoptera: Pseudophyllidae), at Ottawa, Ontario STEPHEN J. DARBYSHIRE Agriculture and Agri-Food Canada, Central Experimental Farm, Saunders Building #49, Ottawa, Ontario K1A 0C6 Canada Darbyshire, Stephen J. 2004. The True Katydid (Prerophylla camellifolia) at Ottawa, Ontario. Canadian Field-Naturalist 118(1): 124-126. Five males of the Northern True Katydid (Pterophylla camellifolia) are reported from Ottawa, Ontario (approximately 45°25°N, 75°42’W), in 2001 and 2002 at three separate locations. Based on its distribution and habits extralimital occurrences in Ottawa are likely due to chance introduction. Key Words: Northern True Katydid, Pterophylla camellifolia, Ontario, distribution, insect dispersal. In mid-August 2001 two male Northern True Katy- dids, Pterophylla camellifolia (Fabricius), were heard calling at a site in an urban residential area of eastern Ottawa, Ontario (location 1, Figure 1). Calling contin- ued until about 9 October when daily minimum tem- peratures fell below freezing. Calling perches in trees were located between 5 and 10 metres above the ground. Several species of trees were utilized, including Sugar Maple (Acer sacharrum Marsh.), Norway Spruce (Abies balsamea (L.) Mill.) and Red Oak (Quercus rubra L.). Over the calling period they moved short distances (less than 50 metres horizontally) from one tree to another, but were never heard calling together in the same tree (it was assumed that the two males heard repeatedly were the same two individuals). Prerophylla camellifolia is not usually associated with coniferous trees and the Norway Spruce was occupied for only a single night. a S ee j EX ON rs: | 3 x sci "3s Ke heal 5 . aN ; Ss ae / N wes ee — ng Lei Exe oy LS ig Y BS, ny 7 3 mY : CAH) Be fh =e 7s _ die opie = a—_—_= 1 km BAN Ere Ye Bx == rh (= Al roy a8 4 oe \ « & at ay ‘ \ ‘ Ww fe SEV\NAVe Oa Sees CZ SEA 2 times the average mass of a usual clutch (mean = 580.3 g, Arnold 1988). Therefore, it is possible that each female produced the entire clutch for these large broods, but it seems improbable. The possibility of intra-specific pre-HBA (hatch brood amalgamation) is supported by Stotts and Davis (1960) who reported two instances of American Black Duck females laying eggs in the nest of another fe- male. They (Stotts and Davis 1960: 145) also com- mented that “Eleven others [clutches] may have been the result of similar parasitism (a total prevalence of 1.8 percent). An example of inter-specific pre-HBA also was recorded by Stotts and Davis (1960) who documented that an American Black Duck female began laying in a Mallard (Anas platyrhynchos) nest that contained five eggs. The Mallard laid four more eggs before deserting, but the American Black Duck laid 11 eggs to equal a 20-egg mixed clutch. Also, they reported that two American Black Duck females nested within 46 cm of each other on an offshore duck blind in Chesapeake Bay, Maryland; one female gradually incorporated the other’s clutch into her own and incubated all eggs (Stotts and Davis 1960: 142). Large clutches associated with ducks nesting on islands, however, may be related to high nest density (i.e., 25.2 — 35.7 nests/ha on Bodkin Island, Chesapeake Bay, Krementz et al. 1991); but is uncom- mon elsewhere (e.g., 0.06 — 0.12 nest/ha in Maine bogs (Coulter and Miller 1968: 35). Because nest sites are not limiting in Maine or across the breeding range, nesting females are widely dispersed, except on islands (Coulter and Miller 1968), nests are well hidden (Bent 1923), and females are secretive when returning to nests in twilight hours (J. R. Longcore, unpublished data), it seems improbable that our large broods resulted from intra-specific pre-HBA (Beau- champ 1997). The third explanation for the large broods is intra-specific post-HBA. For this scenario, two addi- tional clutches of 10-12 eggs must have hatched at the same time and ducklings in those broods were then acquired by the females that we observed. We know that two other American Black Duck broods (i.e., of five and 10 ducklings) used Snake Flowage at the same time as the large broods. To account for this scenario of post-HBA, six American Black Duck broods of similar age must have been associated with these two wetlands. Concentration of broods on high quality 2004 wetlands is common (Longcore et al. 1998); indeed these two wetlands also supported one Green-winged Teal (Anas crecca), three Wood Duck (Aix sponsa), and four Hooded Merganser (Lophodytes cucullatus) broods. For the 20-duckling brood, JRL recorded on 24 June that some ducklings appeared to be in different age Classes (i.e, I[b and IIc), which suggests post-HBA; however, the duckling brood of 18-22 appeared as all the same age to DGM. The two brood-rearing females were large, extremely attentive, and especially adept in eliciting rapid responses from ducklings by uttering a few low calls. Although females are capable of laying large clutches and the remote possibility of nest para- sitism (pre-HBA) exists, we conclude that the most plausible explanation for these two large broods was intra-specific post-HBA. Acknowledgments We thank P. E. Malicky for assistance during brood surveys and J. M. Eadie, T. W. Arnold, and A. J. Erskine for review and helpful suggestions on drafts of the manuscript. Documents Cited (* marked in text) Gollop, J. B., and W. H. Marshall. 1954. A guide for aging duck broods in the field. Mississippi Flyway Council Technical Report, Minneapolis, Minnesota. 14 pages. Literature Cited Arnold, T. W. 1988. Life histories of North American game birds: a reanalysis. Canadian Journal of Zoology 66: 1906- 1912. Beauchamp, G. 1997. Determinants of intraspecific brood amalgamation in waterfowl. Auk 114: 11-21. Bent, A. C. 1923. Life histories of North American Wild Fowl. Order Anseres (Part). Smithsonian Institution, U.S. National Museum, Government Printing Office, Washing- ton, D.C. Coulter, M. W., and W. R. Miller. 1968. Nesting biology of Black Ducks and Mallards in northern New England. Ver- mont Fish and Game Department Bulletin 68-2. 74 pages. Cowardin, L. M., V. Carter, F. C. Golet, and E. T. LaRoe. 1979. Classification of wetlands and deepwater habitats of the United States. U.S. Fish and Wildlife Service, FWS/ OBS-79/31. 103 pages. NOTES 13] Eadie, J. McA., F. P. Kehoe, and T. D. Nudds. 1988. Pre- hatch and post-hatch brood amalgamation in North Ameri- can Anatidae: a review of hypotheses. Canadian Journal of Zoology 6: 1709-1721. Krementz, D. G., V. D. Stotts, D. B. Stotts, J. E. Hines, and S. L. Funderburk. 1991. Historical changes in laying date, clutch size, and nest success of American black ducks. Journal of Wildlife Management 55: 462-466. Longcore, J. R., and J. R. Ringelman. 1980. Factors affect- ing waterfowl breeding density and productivity estimates in the northeast. Transactions of the Northeast Section, The Wildlife Society 37: 169-181. Longcore, J. R., D. A. Clugston, and D. G. McAuley. 1998. Brood size of sympatric American black ducks and mal- lards in Maine. Journal of Wildlife Management 62: 142- 151. McAuley, D. G., and J. R. Longcore. 1988. Survival of juve- nile Ring-necked Ducks on wetlands of different pH. Jour- nal of Wildlife Management 52: 169-176. Norton, S. A. 1980. Geological factors controlling the sen- sitivity of aquatic ecosystems to acid precipitation. Pages 521-531 in Atmospheric sulfur deposition, environmental impact and health effects: Proceedings of the Second Life Sciences Symposium, Potential Environmental and Health Consequences of Atmospheric Sulfur Deposition, Gatlinburg, TN, October 14-18, 1979. Edited by D. S. Shriner, C. R. Richmond, and S. E. Lindberg. Ann Arbor Science Publishing, Ann Arbor, Michigan. Reed, A. 1970. The breeding ecology of the Black Duck in the St. Lawrence estuary. Ph.D. dissertation, Laval Uni- versity, Quebec, Canada. 175 pages. Skutch, A. F. 1967. Adaptive limitation of the reproductive rate of birds. Ibis 109: 579-599. Stotts, V. D., and D. E. Davis. 1960. The Black Duck in the Chesapeake Bay of Maryland: breeding behavior and biology. Chesapeake Science 1: 127-154. Tufts, R. W. 1986. Birds of Nova Scotia. Third Edition. Nimbus Publishing and Nova Scotia Museum, Halifax, Nova Scotia. Zammuto, R. M. 1986. Life histories of birds: clutch size, longevity, and body mass among North American game birds. Canadian Journal of Zoology 64: 2739-2749. Received 28 February 2002 Accepted 15 June 2004 132 THE CANADIAN FIELD-NATURALIST Vol. 118 Observations of Interactions between Puma, Puma concolor, and Introduced European Red Deer, Cervus elaphus, in Patagonia WERNER T. FLUECK CONICET (Consejo Nacional de Investigaciones Cientificas y Tecnoldégicas), Postal address: DeerLab, C.C. 176, 8400 Bariloche, Argentina; e-mail: deerlab@infovia.com.ar Flueck, Werner T. 2004. Observations of interactions between Puma, Puma concolor, and introduced European Red Deer, Cervus elaphus, in Patagonia. Canadian Field-Naturalist 118(1): 132-134. Direct observations of interactions between native Puma (Puma concolor) and introduced European Red Deer (Cervus elaphus) in Patagonia are discussed with respect to the absence of evolutionary sympatry. Although the founding stock of European Red Deer had been lacking natural predation pressure for considerable time due to the previous extinction of large predators, these observations suggested that inherent antipredator behavior of European Red Deer toward this novel predator, once detected, was effective and may partially explain the success of European Red Deer as an invasive species. Puma behavior supported the view that they are a generalist predator which opportunistically utilizes new prey species like European Red Deer. Key Words: European Red Deer, Cervus elaphus, Puma, Puma concolor, introduced species, predation, behavior, interaction, Argentina. The role of large predators in regulating northern cervids continues to be debated. This function of large predators has to be viewed from a historical-evolu- tionary angle; how predators affect prey populations in modern human-modified settings can be expected to differ greatly (Berger 1998; Flueck 2000). The evolu- tion of large predators appears to be a systemic func- tional response as shown by re-evolving ecomorphs among very distant taxonomic groups. Furthermore, ecologically complete northern systems had several predator species with different hunting strategies and several prey species which thus prevented coevolution (Flueck 2000). Parts of Patagonia represent a natural experiment in south-temperate Argentina where Euro- pean Red Deer (Cervus elaphus elaphus) were intro- duced during the 1920s (Flueck and Smith-Flueck 1993) to areas with native Puma (Puma concolor). As the Puma has only occurred in the Americas, these two species have no evolutionary history of sympatry. Fur- thermore, the native large predators had been extinct for a considerable time in areas of Europe where the European Red Deer originated, and immediate memory of such predators was thus absent. Hornocker’s (1970) work showed that direct obser- vations of attempts by Puma to capture prey was near- ly impossible and very few reports have been made previously, mostly stories by hunters (see Robinette et al. 1959). A few additional accounts are based on interpretation of tracks in the snow (Robinette et al. 1959). During many years of studying Puma, Seiden- sticker et al. (1973: 31) never observed Mule Deer (Odocoileus hemionus) or American Elk (Cervus ela- phus nelsoni) mobbing a Puma, and I found no refer- ence to a cervid fighting off a Puma successfully. Here I report on direct observations of interactions between Puma and European Red Deer. Study Area The study area is in the western foothill portion of the province of Neuquén, Argentina (40°58'S, 71°12'W). The topography is primarily mountainous with most features formed by glacial processes. The dominant climate is temperate with main precipitation occur- ring between April and September. There is an abrupt precipitational gradient from west to east due to the rain shadow effect of the Andes which results in a strongly defined vegetation structure and floristic com- position. The study site is between 900 and 1200 m elevation and represents the ecotone between forest and steppe. Patches of forests are characterized by “Nire” trees (Nothofagus antarctica) and “Ciprés” (Austrocedrus chilensis) at lower elevations and are re- placed by “Lenga” trees (Nothofagus pumilio) at high- er elevations. Forest patches at lower elevations alter- nate with wet grasslands (“mallines”) with abundant growth of herbaceous plants, whereas at high elevation they are replaced by grass-dominated steppe of “Coirén Amargo” (Stipa speciosa var. major) and “Coirén Dul- ce” (Festuca pallescens), with variable occurrence of brush species like “Neneo” (Mulinum spinosum), “Calafate” (Berberis spp.) and “Espino Negro” (Colle- tia spinosissima). Riparian areas also contain galleries of small trees like “Radal” (Lomatia hirsuta), “Maitén” trees (Maytenus boaria) and “Laura” (Schinus pata- gonicus). Methods and Results Observations of interactions between European Red Deer and Puma were recorded while stalking alone during field work to collect deer for reproductive stud- ies. Here I report on two such incidents. On 10 November 1999 at 20:15 h I was returning to camp at a pace faster than stalking as it was getting 2004 late and the deer had not yet descended into the flat open feeding areas below surrounding hills which they use for cover during midday hours. The area I tra- versed was semi-open with patches of grassland inter- spersed with some patches of trees like Laura, Radal, and Cypress, typical of the acetone here. There were also patches of brush, predominantly Espino Negro and Calafate. As I was about to pass a stand of Radal, which was approximately 10 m in diameter, I heard the alarm bark of a European Red Deer cow about 20 m away coming from within the stand. I stopped imme- diately, believing that she must have seen me through the Radal trees, particularly as the same had happened only a short while earlier with a cow which was bed- ded in a similar tree patch. I thus waited in antici- pation that this animal might step out of the trees in curiosity, in which case I would have collected her as well. Looking with binoculars, I could see her move behind the trees and affirmed that she had not yet left. Suddenly, she barked again and came charging to- ward me through the patch of trees. Once she was in sight, I saw that she was pursuing an adult Puma, and it was clear that the barking was aimed at the predator and not at me. The Puma stopped shortly after get- ting out into the open, but as the cow came charging again, the Puma took off and the cow chased the cat around a circular patch of Espino Negro of about 7 m in diameter and | m in height. The fast chase took the animals around this bush patch twice, until the Puma apparently stopped out of sight behind the bush. The cow then stopped in front of it and commenced bark- ing again. Finally the cow trotted back toward the patch of trees on the same path where she had come from, only to turn around suddenly and charge toward the patch of bush, stamping in front of it and barking con- tinuously. Four times she repeated this behavioral pat- tern of trotting away and turning back, but when she finally reduced her aggressiveness and appeared about to retreat, I got ready to collect her. As she turned to enter the trees and was obviously walking away from the scene, I shot, which caused her to run through the tree patch on the same path on which she had origi- nally come out. After picking up my gear, I came around a bush and went towards the trees. Immediately, I saw the Puma jump out of a tree behind that patch of bush, and follow the deer on the same path through the trees. I then entered the tree patch, having to hunch down to pass through the trail. As I reached the other side, still in a hunched position, I saw the head of a Puma appear at about 15 m from behind a patch of bunch grass; it must have heard me coming through the trees. Looking through the binoculars, I determined it was a cub of the previous year. Still looking through my binoculars, I saw the head of the mother Puma pop up, and without hesitating she approached me to with- in 10 m. Apparently this was the first she knew of my presence and she began to snarl, which could be NOTES 133 clearly heard. After 30 seconds of staring straight at me, she sat down, but kept looking directly at me and snarl- ing periodically. She then appeared to relax in the sense that she would start to look from side to side, blink- ing slowly, and she was no longer in an alert position. As the sun had set and I needed to collect infor- mation on the cow before dark, I decided after about 5 minutes to move on ahead. I stood up which im- mediately put the Puma into alert mode. She stood up, started to snarl continuously and stared at me. As I made a first few steps towards her, it became clear that she would rather move back than hold her posi- tion, so I continued to move forward. She retreated as much as I would move forward. Approximately 5 m out into the open, I could again see the cub which immediately retreated into the brush upon seeing me. At 15 m from the trees I found the dead cow, while the female Puma stood 10 m away still snarling. As I started to make brisker movements, like taking off the backpack, she retreated farther away and out of sight behind the brush. The thorax of the European Red Deer had already been torn open by the Puma. The European Red Deer cow was of average body size for this region (Flueck, unpublished data), except girth (116 cm) which was 10% below average. She had absolutely no fat reserves (sternum, rump, omen- tum, kidney). This was probably due to extreme drought conditions during spring through fall of 1999. Body conditions in fall of 1999 were so low that only 56% of adult females conceived (Flueck 2001) as compared to 100% in previous years. This 14-year-old female had given birth to a calf 1-2 days earlier as evidenced by the still enlarged uterus and the large but hard udder. Furthermore, in the trees I found remains of a calf which had been partially scavenged at least a day earlier; cause of death could not be determined. The second observation occurred on 17 November 1996, while I was stalking in an area of hilly grassland interspersed with brush. At 18:15 h, I heard a Euro- pean Red Deer cow bark on the other side of a canyon and shortly after, I observed her running nervously back and forth in a semi circle at a patch of Espino Negro brush, about 5x10 m in area and | m tall. Then a calf cried and approximately 5 minutes later cried again. It was obvious that the female had a prob- lem with the calf. Shortly after, | saw an animal body appear from behind the brush patch and enter it. A few minutes later, the female stopped barking and trotted away down the hill. I then went to the spot and as I slowly approached the brush about where the ani- mal had entered earlier, an adult Puma fled crossing the brush patch. Right at the edge was a dead calf which was barely eaten. The Puma had dragged it from some distance when crossing the brush patch. The calf, a male, had body measurements indicating it weighed 11 to 12 kg (Flueck, unpublished data), and several features indicating it was only a few days old (first incisors were out only 4 mm; umbilical scab). 134 Discussion Puma in Patagonia are large predators and compar- able to or surpass the largest races known from the nearctic (Iriarte et al. 1990). Franklin et al. (1999) reported average weights for female and male Pata- gonian Puma as 47.5 and 75.8 kg, respectively. Female Puma are known to increase their kill rate several fold when accompanied by cubs and can be expected to increase their aggressiveness. It is thus of interest that a European Red Deer female at the energetically most stressful period right after birthing coupled with nutritional stress from the drought was able to chase a female Puma which had young, up a tree. It indicates that the defense behavior of European Red Deer towards a discovered novel predator like Puma was effective in preventing the death of the mother. In the first case, the cause of the earlier death of the calf is unknown, and as it is common for a cow to remain in the vicinity of a recently born calf after it dies, it ex- plains the aggressive behavior toward the Puma which likely could have prevented predation on the offspring. In the second case, the mother could not prevent pre- dation of her calf by Puma; however, the learning event might enhance the probability that a future offspring might survive. Innate antipredator behavior of intro- duced European Red Deer therefore, appears success- ful in assuring adequate numbers of offspring to survive against the native Puma hunting strategy. The loss of exposure to large predators for the founding stock in their original habitat did not prevent the estab- lishment of the founding populations in Patagonia, indicating that rapid learning to defend against Puma must have occurred. Other cervids had become naive to their large predators which became locally extinct, but adapted quickly once these predators reappeared. This was clearly shown for Roe Deer (Capreolus capreolus), when Lynx (Lynx lynx) were reintroduced (Breitenmoser and Haller 1993), and Moose (Alces alces) with offspring after reestablishment of Wolves (Cants lupus) and Bears (Ursus arctos) (Berger et al. 2001). On a larger scale, it is also clear that Puma in Pata- gonia, as the major predator of introduced European Red Deer, are not able to prevent them from increas- ing numerically (Flueck et al. 2003). Furthermore, as Puma and deer occur across extensive landscapes with no human-related mortality factors for Puma, I pos- tulate that intrinsic mechanisms are responsible for preventing Puma populations from reaching densities where they would exert a control over the deer popu- lations (Flueck 2000). Puma prey frequently on Euro- pean Red Deer indicating their generalistic and oppor- tunistic feeding behavior. Although about 80% of the natural mortality of adult female deer was due to Puma, annual survival rate of these deer remained high at 91% (Flueck et al. 2004). THE CANADIAN FIELD-NATURALIST Vol. 118 These observations support existing studies that the Puma behaves as a generalist predator which can oppor- tunistically adapt to new prey species. Furthermore, they demonstrate that European Red Deer have an ample repertoire of innate antipredator behavior which may partially account for their success as an invasive species. Acknowledgments I am grateful for the cooperation received from var- ious landowners and the Administracion de Parques Nacionales de Argentina. Thanks also to J. Smith-Flueck and P. S. Gibson for useful comments on the manu- script. Literature Cited Berger J. 1998. Future prey: some consequences of the loss and restoration of large carnivores. Pages 80-100 in Behavioral Ecology and Conservation Biology. Edited by T. Caro. Oxford University Press, Oxford, England. Berger, J., J. E. Swenson, and I. L. Persson. 2001. Recol- onizing carnivores and naive prey conservation lessons from Pleistocene extinctions. Science 291: 1036-1039. Breitenmoser, U., and H. Haller. 1993. Patterns of predation by reintroduced European lynx in the Swiss Alps. Journal of Wildlife Management 57: 135-144. Flueck, W. T., and J. M. Smith-Flueck. 1993. Uber das in Argentinien angesiedelte Rotwild (Cervus elaphus, L., 1758): Verbreitung und Tendenzen. Zeitschrift fiir Jag- dwissenschaft 39: 153-160. Flueck, W. T. 2000. Population regulation in large northern herbivores: evolution, thermodynamics, and large predators. European Journal of Wildlife Research 46: 139-166. Flueck, W. T. 2001. Pregnancy rates of introduced red deer in Patagonia, Argentina after a period of drought. Ecologia Austral (Argentina) 11: 17-24. Flueck W. T., J. M. Smith-Flueck, and C. M. Naumann. 2003. The current distribution of red deer (Cervus elaphus) in southern Latin America. European Journal of Wildlife Research 49(2): 112-119. Flueck, W. T., J. M. Smith-Flueck, and N. A. Bonino. In press. A preliminary analysis of cause-specific and capture-relat- ed mortality, and survival of adult red deer in northwestern Patagonia. Ecologia Austral. Franklin, W. L., W. A. Johnson, R. J. Sarno, and J. A. Iriarte. 1999. Ecology of the Patagonia puma Felis con- color patagonica in southern Chile. Biological Conser- vation 90: 33-40. Hornocker, M. G. 1970. An analysis of mountain lion preda- tion upon mule deer and elk in the Idaho primitive area. Wildlife Monographs 21: 1-39. Iriarte J. A., W. L. Franklin, W. E. Johnson, and K. H. Redford. 1990. Biogeographic variation of food habits and body size of the American puma. Oecologia 85: 185-190. Robinette, W. L., J. S. Gashwiler, and O. W. Morris. 1959. Food habits of the cougar in Utah and Nevada. Journal of Wildlife Management 23: 261-273. Seidensticker, J. C., M. G. Hornocker, W. V. Wiles, and J. P. Messick. 1973. Mountain lion social organization in the Idaho primitive area. Wildlife Monographs 35: 1-60. Received | September 2000 Accepted 4 June 2004 2004 NOTES 135 Premieres mentions de la Couleuvre mince, Thamnophis sauritus septentrionalis, au Québec JEAN-FRANCOIS DESROCHES! et RICHARD LAPARE* 'Collége de Sherbrooke, Département des Techniques d’écologie appliquée, 475 du Parc, Sherbrooke (Québec), J1K 4K1 Canada 2Ministére des Transports du Québec, Direction de Laval-Mille-fles, 1725 boulevard Le Corbusier, Laval (Québec), H7S 2K7 Canada Desroches, Jean-Francois, et Richard Laparé. 2004. Premiéres mentions de la Couleuvre mince, Thamnophis sauritus septentrionalis, au Québec. Canadian Field-Naturalist 118(1): 135-137. Deux Couleuvres minces, Thamnophis sauritus septentrionalis, ont été trouvées les 12 et 13 aoat 2003, dans le comté de Pontiac en Outaouais, au Québec. II s’agit des premieres mentions de cette espéce dans la province. Des détails concernant les spécimens trouvés et les habitats sont présentés, de méme qu’une breve discussion sur la présence de cette espece au Québec. Trois autres Couleuvres minces ont été trouvées en 2004, dans le méme secteur, le méme qu’une autre a I’ fle-du-Grand-Calumet, a environ 30 km au nord-ouest. Two Ribbonsnakes, Thamnophis sauritus septentrionalis, were found on 12 and 13 of August 2003, in Pontiac County, Outaouais region, in Québec. These are the first records of the species for the province. Details on the specimens and habitats in which they were found are presented, as well as a brief discussion on the presence of Ribbonsnakes in Québec. Three additional Ribbonsnakes were found in 2004, in the same area, and another one on Ile-du-Grand-Calumet, about 30 km northwest. Mots-Clés : Couleuvre mince, Thamnophis sauritus septentrionalis, aire de distribution, premieré mention, habitat, Québec. Key Words: Northern Ribbon Snake, Thamnophis sauritus septentrioncilus, distribution, first record, habitat, Québec. La Couleuvre mince (Thamnophis sauritus) est pré- sente dans une grande partie de l’est de 1’ Amérique du Nord, de la Floride jusqu’en Ontario (Conant et Collins 1998). Au Canada, elle est représentée par la sous-espéce T. s. septentrionalis, laquelle se rencontre dans le sud de l’Ontario et sous forme de population disjointe en Nouvelle-Ecosse (Cook 1984). Malgré sa répartition limitrophe au Québec, cette couleuvre n’avait jamais été rapportée dans cette province. 7! s. septentrionalis mesure au plus 96,5 cm a l’age adulte et ressemble a premiére vue a la Couleuvre rayée de Pest (Thamnophis sirtalis sirtalis), présentant comme celle-ci un corps foncé marqué de trois rayures longi- tudinales généralement jaunes. On distingue la Cou- leuvre mince de la Couleuvre rayée par la position de ses rayures latérales, situées sur les 3*°™° et 4°™° ran- gées d’écailles du corps, son corps plus élancé et sa queue comparativement plus longue, et l’écaille pale bien définie devant chaque ceil (Cook 1984). La colora- tion des flancs, sous les rayures latérales, est souvent marron. La Couleuvre mince est considérée menacée en Nouvelle-Ecosse et en situation préoccupante en Ontario (COSEPAC 2002*), tandis que la Couleuvre rayée est tres commune dans ces provinces tout comme au Québec. Les 12 et 13 aoat 2003, dans le cadre d’un travail d’inventaire faunique effectué par le Ministére des Transports du Québec, dans un secteur circonscrit du comté de Pontiac, en Outaouais, nous avons décou- vert deux spécimens de la Couleuvre mince. Dans les deux cas les couleuvres se trouvaient sur une route de sable et de gravier et ont été apercues a partir d’un véhicule en mouvement. Le temps était ensoleillé et chaud, avoisinant les 30° Celsius. Le premier spécimen de la Couleuvre mince a été observé le 12 aoat a 18h10, sur le chemin Bristol Mines, a 1,2 km a l’ouest de la jonction avec le che- min Pontiac Station (45°29'29"N; 76°20'12"O), a environ 2 km au nord de la riviére des Outaouais. Il s’agissait d’un adulte mort écrasé, mesurant 57,6 cm du museau au cloaque, pour une longueur totale (in- cluant la queue) de 80,9 cm. Malgré que la couleuvre semblait avoir été tuée plusieurs heures auparavant, il a été facile de l’identifier par la position des rayures latérales et la couleur marron des flancs. Le second individu a été trouvé le 13 aodt a 16h53, sur un chemin reliant les immenses tas de débris miniers a la riviére des Outaouais (45°28'38"N; 76°20'15"O), a environ | km au nord de cette riviére. C’était un sub- adulte mesurant 29,7 cm du museau au cloaque, pour une longueur totale (incluant la queue) de 45,5 cm. Ce spécimen était vivant et 4 découvert sur le che- min. Des photographies des deux Couleuvres minces ont été prises par les auteurs. Les spécimens ont été déposés au Musée canadien de la nature (CMNAR 35485, 35486). 136 L’habitat ot furent trouvées les Couleuvres minces est majoritairement forestier avec présence de milieux humides. Dans le premier cas, il s’agit dune forét mélangée a dominance de feuillus, ou poussent Acer rubrum, Populus grandidentata, Populus tremuloides, Fraxinus sp. et Thuja occidentalis. En bordure nord du chemin se trouve un étang 4 Quenouilles (Typha sp.) dense d’environ 250 m de longueur par 20 m de largeur. Dans le second cas, il s’agit d’une forét hu- mide a dominance de feuillus. On y trouve Betula papyrifera, Prunus sp., Acer rubrum, Populus balsa- mifera, Larix laricina, Fraxinus sp. et Alnus incana ssp. rugosa. A 40 m au nord, de chaque cété du ch- emin, un marécage est adjacent a la route. On y retrou- ve des arbres morts en milieu inondé. Des recherches effectuées dans les habitats avoisinants n’ ont pas per- mis d’observer d’autres Couleuvres minces. Quatre Couleuvres rayées de l’est, soit deux vivantes et deux mortes, ont été observées sur la route durant I’ inven- taire. En 2004, trois (3) autres Couleuvres minces adultes ont pu étre observées dans le méme secteur, qui couvre 18 km?, et une (1) autre sur I’ Ile-du-Grand-Calumet, a quelque 30 km au nord-ouest. La premiere a été trouvée le 9 juin en bordure d’un étang a quenouilles pres d’un chemin de terre (45°29'10"N; 76°20'56"0). La seconde a été observée le 14 juin dans une flaque d’eau en asséchement, bordée de saules (Salix sp.), a 10 m de la forét (45°29'40"N; 76°22'00"0). La troisiéme a été vue le 12 juillet sur une hutte de Castors (Castor canadensis) (45°31'51"N; 76°18'43"0). Le spécimen de I’[le-du-Grand-Calumet a été trouvé le 11 aofit sure une hutte de Castors (45°41'12"N; 76°38'31"0). Des photographies des Couleuvres minces ont été prises par l’un des auteurs (JED). La présence de la Couleuvre mince au Québec n’est pas surprenante puisqu’on la retrouve en Ontario a proximité, de l’autre c6té de la riviére des Outaouais (Tobias et Evans 1979; Cook 1981; Oldham et Weller 2000*). Il est plut6t curieux qu’elle n’ait pas été rapportée plus tot dans la province. II s’agit bien sir d’une région peu habitée et sous-inventoriée par les biologistes, mais plusieurs inventaires spécifiques y ont été réalisés notamment pour les tortues. Il apparait pos- sible que sa ressemblance avec la Couleuvre rayée de Vest, tres commune dans la région, ait contribué a la garder dans |’anonymat durant toute cette période. La Couleuvre mince devient donc la 38'*™° espéce de l’herpétofaune au Québec et la 8°™° en ce qui con- cerne les couleuvres. La liste des couleuvres du Qué- bec comportait sept (7) espéces depuis 1927, année ot la Couleuvre brune (Storeria dekayi) fut découverte a Montréal (voir les données détaillées dans : Bider et Matte 1991). Les additions récentes a l’herpétofaune québécoise concernent les autres groupes (anoures, urodéles et tortues). Ce sont la Salamandre sombre des montagnes (Desmognathus ochrophaeus), décou- verte au Québec en 1988 (Alvo et Bonin 2003), la THE CANADIAN FIELD-NATURALIST Vol. 118 Tortue musquée (Sternotherus odoratus) en 1989 (Chabot et St-Hilaire 1991) et la Rainette faux-grillon boréale (Pseudacris maculata) en 1991 (Bider et Matte 1994). L’addition de la Couleuvre mince a la liste des reptiles du Québec revét une importance particuliére puisqu’il s’agit sans aucun doute d’une espéece trés rare dans la province et qu’elle est déja considérée en situa- tion précaire en Ontario et menacée en Nouvelle-Ecosse. Des inventaires supplémentaires seront nécessaires afin de préciser sa répartition et son statut au Québec. Fi- nalement, cette couleuvre vient probablement clore la liste de ’herpétofaune du Québec, si l’on se fie a la répartition actuellement connue des espéces. Remerciements Nous tenons a remercier le Ministére des Transports du Québec, et plus particulierement M. Ahmed Kho- dari, chef de service du Service des inventaires et du Plan, Direction régionale de |’Outaouais, pour avoir permis la réalisation de l’inventaire faunique au cours duquel les mentions de Couleuvres minces ont pu étre faites. Nous remercions également MM. Daniel St- Hilaire et Michel Lalancette, de la Société de la faune et des parcs du Québec, pour le prét des cartes topo- graphiques. Les spécimens de 2004 ont été trouvés par Mathieu Ouellette, Josiane Bergeron, Louis-Philippe Gagnon et Benoit Rivard. Documents cités (identifiés par * dans le texte) COSEPAC. 2002. Comité sur la situation des espéces en péril au Canada. http://www.cosepac.gc.ca/index.htm. Oldham, M. J., et W. F. Weller. 2000. Ontario Herpe- tofaunal Atlas. Natural Heritage Information Centre, Ontario Ministry of Natural Resources. http://www. mnr.gov.on.ca/MNR/nhic/herps/ohs.html (a jour le 09- 11-2002). Littérature citée Alvo, R., et J. Bonin. 2003. Rapport sur la situation de la Salamandre sombre des montagnes (Desmognathus och- rophaeus) au Québec. Société de la faune et des parcs du Québec, Direction du développement de la faune. Québec. 32 pages. Bider, J. R., et S. Matte. (Compilé par). 1991. Atlas des amphibiens et reptiles du Québec 1988-1989-1990, ver- sion 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 pages. Bider, J.-R., et S. Matte. 1994. Atlas des amphibiens et rep- tiles du Québec. Société d’ histoire naturelle de la vallée du Saint-Laurent et ministére de l’Environnement et de la Faune du Québec, Direction de la faune et des habitats. Québec. 106 pages. Chabot, J., et D. St-Hilaire. 1991. Premiére mention de la Tortue musquée, Sternotherus odoratus, au Québec. Cana- dian Field-Naturalist 105 : 411-412. Conant, R., et J. T. Collins. 1998. A Field Guide to Rep- tiles and Amphibians of Eastern and Central North Ameri- ca. Third edition, expanded. The Peterson Field Guide Series. Houghton Mifflin Company, Boston. 616 pages. 2004 Cook, F. R. 1981. Amphibians and reptiles of the Ottawa District. Trail & Landscape 15 : 75-109. Cook, F. R. 1984. Introduction aux Amphibiens et Reptiles du Canada. Musée national des sciences naturelles et Musées nationaux du Canada. Ottawa, Canada. 211 pages. Desroches, J.-F., et David Rodrigue. 2004. Amphibiens et reptiles du Québec et des Maritimes. Editions Michel Quintin, Waterloo, Québec. 288 pages. NOTES 137 Revisée 16 Aott 2004. Tobias, T. N., et R. E. Evans. 1979. Range extension of the Ribbon Snake in Ontario. Trail & Landscape 13 : 180-181. Re¢u 20 aout 2003 Acceptée 27 fevrier 2004 Book Reviews ZOOLOGY Amphibian Decline: An Integrated Analysis of Multiple Stressor Effects Edited by Greg Linder, Sherry K. Krest, Donald W. Sparling. 2003. SETAC North America, 1010 North 12th Avenue, Pensacola, Florida 32501-3367 USA. xxi + 345 pages. U.S.$98 Cloth. This volume contains presentations at the Workshop on the Global Decline of Amphibian Populations sponsored jointly by the Society of Environmental Toxicology and Chemistry (SETAC) and The Johnson Foundation 1-23 August 2001 at Racine, Wisconsin. Fifteen papers are included as 10 chapters, one with A-B and another with A-E designations for no appar- ent reason. Canadian data and perspective on the problem of declines are not neglected. Of the 35 contributing authors listed, 31 are from the United States but four are Canadian: Christine Bishop (Canadian Wildlife Service, Delta, British Columbia), David Cunnington (“freelance”’, Victoria, British Columbia), Martin Ouellet (Redpath Museum, McGill University, Mon- treal, Quebec), Bruce D. Pauli (Canadian Wildlife Ser- vice, Hull, Quebec). Ouellet is one of nine coauthors of Chapter 6 “Biotic Factors in Amphibian Population Declines”, and Bishop, Cunnington, and Pauli are half of the six coauthors of Chapter 7 “Physical Habitat and Its Alteration: A Common Ground for Exposure to Environmental Stressors”. None of the individual chapters is a particularly easy read and their results or conclusions often are neither clear-cut nor universal in their application. However, collectively, they effectively and repeatedly bring out the complications in separating climatic, chemical pol- Birds of Africa: From Seabirds to Seed-eaters By Chris and Tilde Stuart. 1999. MIT Press, Five Cambridge Center, Cambridge, Massachusetts, USA. 176 pages. U.S. $29.95 Cloth. This is an attractive book that appears to be in- tended as a typical “coffee table” volume. It is large in size (22 x 28 cm). profusely illustrated and very well laid out, with between 2 and 9 illustrations on each two-page spread. The plates themselves range from full page to quite tiny (under 4 cm’). The text is presented in 12 chapters plus an intro- duction, species list, and index. Each chapter is devoted to a group of bird families with some feature in com- mon. Thus we have “Birds of the Oceans’, “Birds of Inland Waters and the Coastline”, “LBJS”, and so on. Chapters then deal with each family in turn (grouped lutant, and biotic pressures, many of which doubtlessly combine to produce documented or supposed losses or declines. The texts are heavily laden with flow and modelling in 18 figure diagrams, 18 tables, 5 matrices and even two “test boxes” to visually present concepts and consequences. Each chapter has its own biblio- graphy. The concluding summary chapter effectively stresses the continuing, unresolved nature of the problems, but, in an inevitable rather self-congratulatory manner, arrives at the conclusion that this workshop was vital to promoting a much-needed exchange of expertise on various aspects of real and potential amphibian dec- lines and that it has created a network for further discussion and exchange. Although one could argue that this was happening anyway, the workshop did usefully focus discussions and the book spreads the state of knowledge. It is useful both for what research has been done and for what future approaches might be productive. One can hardly disagree with the funda- mental conclusion that further sharing of information as it develops will be essential to adequately defining future measures. If, eventually, there ever will be effec- tive conservation of the world amphibian fauna it will depend on a better understanding of the causes and reality of the declines. FRANCIS R. COOK Canadian Museum of Nature, Ottawa, Ontario KOG 1RO Canada in the sequence of their common names), giving the number of species that occur in Africa together with a brief account of the family characteristics, and some- thing about a sampling of the species. Occasionally there are boxes which treat a particular topic in some detail; for example, “The Dilemma of Island Birds”. The main body of the text enlivens the chronicle of numbers and species descriptions by concentrating on interesting anecdotes and “gee whiz” facts about the species under discussion. The authors are African and obviously have had considerable experience with the continent’s bird life. On the whole their statements seem accurate, although I was unable to check some of them, but some are incorrect. For example, Great Skua does not breed in the Southern Hemisphere, Kelp 138 2004 Gull is not an African endemic, and the Australian Ibis is not conspecific with the Sacred Ibis. For me this book has some rather irritating problems that in sum seriously detract from its interest. Overall there is a lack of cohesion, mainly resulting from the organization of the text. The arbitrary chapter group- ings inevitably run into families that do not quite fit the category, or fit into more than one. I would be in- clined to place gulls and terns in the coastal waterbird chapter, lots of the “Seed-eaters” are classic LBJ’s, and “Terrestrial Birds” results in a strange collection of species with Pittas following Ostrich. This is ac- centuated by the layout within the chapters, with (for example) “Avocets and Stilts” at the beginning of Chapter 3, “Oystercatchers” and “Plovers and lap- wings” in the middle separated by Pelicans, and the “Waders or shorebirds” at the end. True, the system- atic list at the end is in order, but this does little to balance the sense of randomness that one gets from the text itself. Probably the authors would point out approaches of this kind are not unusual for books of the genre: they are not, after all, basic reference texts. Unfortu- nately, attractive though the layout is, there are prob- lems with the plates as well. In many cases the tinier The Firefly Encyclopedia of Birds Edited by Christopher Perrins. 2003. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 Canada. 656 pages. U.S.$59.95. Naturalists across the continent strive to ever in- crease their knowledge of local wildlife, and all love to hear about wildlife from other localities. Birdwat- chers, for example, can become very knowledgeable about their local species, and the zealous ones will even study species from other areas within the same country, continent or even from around the world. But the vast array of bird species is overwhelming, and it takes a serious encyclopedia to pull them all together. Luckily for us, The Firefly Encyclopedia of Birds edited by Christopher Perrins is just what we needed — a good collection of text and photos of the birds of the world all in one volume. Nicely present- ed with tons of sharp photos illustrating everything from specific behaviours to simply jaw-dropping “‘cool- ness” (including several underwater pictures such as common murres on page 280-281, common kingfisher on page 368, and American dipper on page 521), the book is backed up by solid text summarizing well the basic tenets of ornithology. Moreover, each chapter BooK REVIEWS eo) illustrations are difficult to see properly, many of the poses are poor, and the selection of subjects pictured leaves something to be desired. Opening the book at random I find pp. 130-1 show two different pictures of Arrow-marked Babblers, one of the Southern Pied Babbler, and one of a Wallcreeper. The latter only occurs as a vagrant in North Africa, and the plate shows little of this species’ remarkable wing pattern, so it simply looks like a gray bird. Africa does have a creeper species, and the text on these pages also in- cludes the broadbills (Eurylaimidae). These species are not illustrated at all. In summary, this book brings together a large num- ber of photographs of African birds, some of them very good, together with a compilation of interesting facts and anecdotes on African bird families. My quarrels with content are really irrelevant if this is all you are looking for, or if you find this book attractive enough in itself to buy. Those who desire a thorough overview of the African avifauna should look else- where. CLIVE E. GOODWIN 1 Queen Street, Suite 405, Cobourg, Ontario K9A 1M8 Canada focuses on a bird family (the large number of species precludes a species-by-species analysis), and is accom- panied by a “factfile”. This sidebar summary provides a general description of size and habits, as well as a broad distribution map and my favorite, a visual com- parison of bird size to a human. The book also details several interesting specific behaviors, such as pigeon homing (pages 294-295), or the nest parasitism of Cuckoos (pages 316-317), but nothing on the degredation of the tundra by Snow Geese, a topic of high importance and actuality in North America. It also discusses extinct birds such as the Elephant Bird (page 21), the Dodo (page 291), and a short paragraph on the Passenger Pigeon (page 293). The book provides a fantastic introduction to birds from all around the world, is extremely well presented, very reasonably priced ($75.00 CAN, hardbound), and constitutes a worthy, if not essential, addition to the library of all bird enthusiasts. SERGE LARIVIERE 226 Rang Bois-Joly, St-Apollinaire, Québec GOS 2E0, Canada 140 Birds of Belize By H. Lee Jones. 2004. University of Texas Press, P.O. Box 7819, Austin, Texas 78713-7819 USA. x + 444 pages. U.S.$60.00 Cloth, U.S.$34.95 paperback. This first edition covers the 574 species of birds recorded in the old British colony of Honduras, and now called Belize. Virtually all the birds are shown in colour plates, with accompanying text and 234 range maps. The book has a brief introduction to the biogeography and climate of Belize and a how-to- use chapter. This latter is very important because it explains how the author has chosen to organize the book and select the “common” names, as well as the names in Maya, Spanish and the aboriginal language. The range maps, at the back of the book, mostly cover the resident species and are large enough to be a useful guide. The text is precise and clear and contains much useful information. For example, the call of the North- ern Potoo is described in terms that should enable anyone to recognize it. This is the key characteristic in separating the Common from the Northern Potoo. Another good example is the description of the Double- toothed kites, explaining the “Distinctive puffy white undertail coverts that spread to the sides of the rump .... This is a very good field character and this description captures it well. The artwork is by Dana Gardiner, the same artist that illustrated the Birds of Costa Rica by Stiles and Skutch. I found his style a little stiff, giving the birds a flat appearance and missing something of their jizz. These new renderings, while similar to those in Birds of Costa Rica, are an improvement. The Wedge-billed Woodcreeper illustration more clearly depicts its re- markable tail. The Emerald Toucanet is the correct subspecies, Aulacorhynchus prasinus prasinus, show- ing a white throat. However, I did have difficulties Herpetology: Third Edition By F. Harvey Pough, Robin M. Andrews, John E. Cadle, Martha L. Crump, Alan H. Savitsky and Kentwood D. Wells. 2004. Pearson Prentice Hall, Upper Saddle River, New Jersey, USA. ix + 726 pages. U.S.$93. In the preface to this edition of Herpetology, the authors state, “understanding amphibians and reptiles as organisms requires a perspective that integrates their morphology, physiology, behavior, and ecology and places that information in a phylogenetic context.” The authors have done a commendable job at fulfill- ing this daunting task. The text book is divided into four parts and each part is subdivided into chapters. Part one (What are amphibians and reptiles?) explores the field of her- petology, the place of amphibians and reptiles in ver- tebrate evolution, systematics and diversity of amphib- ians and reptiles, and biogeography. Part two (How do THE CANADIAN FIELD-NATURALIST Vol. 118 with some plates, but I think this relates more to the printing than the artwork. For example, the depiction of Cedar Waxwing is far too intense and the elaenias are much too dark. The depiction of the Violet Sab- rewing is deep blue. I have never seen one in the field that did not look violet (as described accurately in the text and depicted accurately in Birds of Costa Rica) under any light conditions. In particular I checked the depictions of Scaly-breasted Humming- bird (poorly done in Birds of Costa Rica) and it is better, but I still think a novice would have trouble in the field using the plate for identification purposes. The nomenclature is fairly standard. The author retains Rock Dove for Columba livia (instead of the new “Rock Pigeon” — thereby reducing confusion with the orange-brown Rock Pigeon of South Africa, Columba guinea). | had to use the scientific name of Phaethornis longirostris to be sure the author’s Long- billed Hermit was the same as Stiles and Skutch’s (Western) Long-tailed Hermit. I am still confused as to the status and distribution of Passerini’s and Cherrie’s Tanager (a recent split of Scarlet-rumped Tanager). Only Paserrini’s Tanager is listed for Belize, which I believe is correct. This guide is just under 6” x 9” x 1.25” (15 x 22 x 3 cm) and is too large for the average pocket. Al- though I have some reservations, | still think this book will make a useful field guide, especially if you use the text in conjunction with the plates. This book rep- resents a good stride forward in literature for this section of the Americas. If it is to be used in the field I would be tempted to take along a Mexican guide as a second reference. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario, K1J 6K5 Canada they work?) has chapters on temperature, energetics, reproduction, locomotion and feeding. Part three (What do they do?) covers movements, communication, mat- ing systems, diets and species assemblages. The fourth part (What are their prospects for survival?) discuss- es conservation of amphibians and reptiles in a single chapter. How does this edition differ from previous editions? The second edition was published in 2001 and hence this edition includes many references to recent pub- lications. There is also a greater emphasis on phylo- genetic analyses, particularly in the early chapters on systematics. This edition also features colour photo- graphs and colour distribution maps for the families of amphibians and reptiles. The colour maps do not really add much, but the colour photographs are effec- tive. A chapter on biogeography is one of the biggest 2004 changes in content in this edition. The authors have also wisely split the chapter on reproduction into two chapters, one each for amphibians and reptiles. The section on conservation has also been expanded, re- flecting the growing concern over many species in both groups. It is hard to find much wrong with this exhaustive examination of the current state of herpetology. The one conspicuous absence is a glossary, something that would surely be important in any comprehensive text- book such as this. Overall, the combined talents of Lizards: Windows to the Evolution of Diversity By Eric R. Pianka and Laurie J. Vitt. 2003. University of Cali- fornia Press, 2120 Berkeley Way, Berkeley, California, USA. xii + 333 pages. U.S.$45 Cloth. Individually, each author of this volume has more than 30 years of lizard study in various areas of the world. These include the southwestern (both) and southeastern (Vitt) United States, as well as Africa and Australia (Pianka) and Central and South America (Vitt). Each has included a personalized capsule bio- graphy of his background and interests, each empha- sizing what first attracted him to lizards. This book is number 5 in the University of Califor- nia Press series Organisms and Environments. The introduction explains that it grew from a resolve by authors 10 years ago. At that time, while working on a third lizard ecology symposium volume, they envi- sioned production of a comprehensive semi-popular book on the group. In a review of the earlier work, the 1994 Lizard Ecology: Historical and Experimental Perspectives, Robert W. Murphy (1995. The Canadian Field-Naturalist 109(1): 135-136), while generally praising its standards of “highest academic acumen”, thought it “not necessarily well suited for bedside reading by the merely curious. And it will not make a good coffee table book as there are no high-gloss photographs”. The latter criticism has been particularly well res- ponded to in the new volume. It is not only coffee- table size in design (28.6 x 22.5 cm) but has the req- uisite high-gloss photographs scattered throughout. These, often emphasizing activity or displays, splen- didly illustrate the diversity of behaviour as well as form and pattern for a well-chosen variety of lizards. There are also a scattering of graphs, tables, diagrams, and maps to illustrate particular points. Included are evolutionary relationships between groups of lizards, biotic and abiotic factors affecting a lizard’s well- being, avenue of heat gain and loss for a diurnal bask- ing lizard, comparison of mean percent use of the seven most important prey categories by neotropical and desert lizards, prey size vs lizard size, independent evolution of body type in Jamaica and Puerto Rico, and population size for species of land iguanas on islands of the Caribbean, to choose a few at random. Book REVIEWS 14] six experts with diverse research interests is hard to beat. After two previous editions, the authors have resolved most of the inevitable errors and contradic- tions resulting from integrating the writing of so many people. It is hard to imagine a better single volume overview of these fascinating creatures. DAVID SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1S0 Canada The introduction raises the question: “what good are lizards?” This is first summarily dismissed with a curt retort, emphasizing the authors’ abhorrence of anthropocentrism, “what good are people?”. Subse- quently, the instructive value of lizards in relation to the ecosystem is attributed to their multitude of forms and the variety of habitats they have successfully occupied. They can thus effectively serve as “model” organisms for broad understanding of ecology and the diversity of animal life, a theme also emphasized in the earlier symposium volumes. Following the introduction are three major parts with seven, six, and two sections: (1) Lizard Life- styles: Evolutionary history and phylogeny, Getting around in a complex world, Lizards as predators, Es- caping predators, Social behavior, Reproduction and life history, and Reflections of a real world; (2) Liz- ard Diversity: Iguanians, From Geckos to Blind Liz- ards, From Racerunners to Night Lizards, Skinks, From Girdled Lizards to Knob-scaled Lizards, Monsters and Dragons of the Lizard World; (3) Synthesis: Historical perspective, Lizards and humans. In the initial section it is pointed out that as a group “lizards” is para- phyletic due to excluding the snakes. The latter are omitted as they traditionally have been treated as a separate equal group, though now recognized in con- temporary clarification as just an offshot within vara- noid lizards (as shown in Figure 1.4, page 16). The major partitions are followed by a four-page Taxonomic Summary (with number of species in each family or subfamily (if it has been subdivided) listing all included genera). The species totals are given (by major subdivisions) as [guania 1340+; Scleroglossa: Gekkota 973+, Incertae sedis 151+; Autarchoglossa (excluding snakes but inclusive of Scincomorpha: Lacertoidea, Scincoidea: Anguimorpha [including Varoidea]) 1745+; or collectively over 4200. The book concludes with a three-page Glossary (abiotic to zygo- dactyly), an 18-page references section, and a nine- page index. Throughout, the book is a pleasant mix of formal and informal styles. Scientific discussions have shuffled among them separate blocks of personal observations, often field experiences of one or the other author. As stated (page 7) “Throughout this book, we continu- 142 ally return to questions we asked as children in an attempt to encourage readers to open their minds and to ask questions”. This responds splendidly to the ear- lier reviewer’s wish for a text “well suited for bed- side reading by the merely curious”. Unfortunately, the large coffee table format and the glossy reflective pages make it awkward to handle in bed and read by bedside light. But as an up-to date comprehensive survey of world lizards it has no equal. It provides Raptors of Western North America Brian K. Wheeler. Princeton University Press, 41 William Street, Princeton, New Jersey 08540-5237 USA. 544 pages, 625 colour photographs, 56 maps, U.S.$49.50 Cloth This book is both a tour de force and a labour of love, and the immediate new standard for diurnal raptors in western North America. Raptors are known for their variable plumages, and standard field guides cannot cover the almost infinite permutations and com- binations; hence, erroneous identifications are legion among regular birders, and even among raptor enthu- siasts. No full-time museum, government or university employee has dared to undertake a book of this scope; it is all the more astounding that this hiatus was filled by a truck driver. More incredibly, Wheeler has devel- oped the skills to take the great majority of the pho- tographs himself. His experience and enthusiasm are evident throughout. He has reviewed the literature carefully and has been given access to unpublished information from graduate students and others; for example, numbers of Northern Goshawk and Bald Eagle territories are provided for each state. There are nearly two pages about the ten-year cycle of the Snow- shoe Hare and Northern Goshawk, absent from most other raptor books; goshawks peaked at Duluth, Min- nesota, during the irruption years of 1972, 1982, 1992 and 2001, mostly adults fleeing the mixed forest fol- lowing each Snowshoe Hare crash. Excellent historical accounts include notably that for the California Condor. There are four excellent glos- saries: general, anatomy and feather, plumage and molt, and displays. The detailed descriptions of raptor plu- mages are unprecedented and unequalled. Fortunately, important points are highlighted in bold, making the tedious mass of description more user-friendly. Readers will be amazed by the remarkable variabil- ity of plumage in species such as the Ferruginous Hawk (25 photos), Rough-legged Hawk (39 photos), and Swainson’s Hawk (40 photos), and especially the Red-tailed Hawk; variants within the latter species are depicted in 82 photographs, 22 pages of verbal des- cription and six maps. THE CANADIAN FIELD-NATURALIST Vol. 118 great perspective of their global importance, a fact strictly Canadian naturalists, who have only six spe- cies belonging three families of non-snake lizards to observe in their entire country, have little first-hand chance to appreciate. FRANCIS R. COOK Canadian Museum of Nature, Ottawa, Ontario K1P 6P4 Canada John Economidy has contributed clear, accurate range maps for 33 species, six for the Red-tailed Hawk alone. The maps are incredibly precise, but I noticed four minor errors for Saskatchewan species: the Big Muddy area of Saskatchewan is omitted from the Golden Eagle; the two southern year-round localities for Bald Eagle should be winter only; recent small southerly extensions of Osprey and Northern Goshawk ranges are not shown; the two races of the Merlin are not known to overlap between Saskatoon and Prince Albert. The maps also give no indication of range changes in the past. Since this veritable encyclopaedia of western raptors (an eastern counterpart became available simultane- ously) is too heavy for most backpacks and too large for most glove compartments, most of us will use it as a reference in our libraries. One hesitates to quibble about the first edition of such a superb book. Yet the title is misleading, since owls are not included; inclu- sion of the term “diurnal” would have made the title more accurate. The ground squirrel has not lost half its range in western Canada, as it has in Idaho (page 521). Use of dieldrin, not DDT, was coincident with the early 1960s crash of the Merlin on the Canadian prairies (page 447). Use of “very uncommon” some- times contradicts his definitions in the introduction. I detected few errors, but “verses” in place of “versus” jarred me (page 199). Wheeler has been involved in two previous hawk identification books, each excel- lent, each with William S. Clark as co-author. The first was Hawks in the Peterson Field Guide series and the second, A Photographic Guide to North American Raptors. For most general birders, either would suffice, although many plumages are omitted from both of the smaller books. If you are a raptor aficionado, don’t let the cost of the new book deter you; the 603 colour photographs easily justify the price and the 33 colour maps are more detailed than anything previously available. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8, Canada I 2004 Raptors of Eastern North America By B. Wheeler. 2002. Princeton University Press, 41 Williams Street, Princeton, New Jersey, USA. xv + 439 pages, U.S.$45.00 Cloth Let me begin by saying this is a wonderful book. It gives a very detailed account of all the 26 full species of hawks, eagles and falcons occurring in eastern North America. It is profusely illustrated with extremely good, frame-filling, crisply focused photographs. These range from nine photographs for the Short-tailed Hawk to an incredible 82 of the Red-tailed Hawk. The text, des- pite its somewhat clipped English, is very informative. All of the known and recognizable sub-species are included, both in the text and, where valid, in the photographs. The level of detail given in the plumage descriptions is far greater than in any other book I have read. The range maps are sized as appropriate to the species they cover. For widespread birds the map depicts all of eastern North America. However, larger scale maps are used for such localized birds as Flor- ida’s Snail Kite. This is easily the best photographic-style guide I have seen and is a real testament to the author’s dedi- cation. Thus it is the most thorough guide to this group of North American birds you can purchase. The full- color, superb photographs alone are worth the price of the book. The author has included all the plumages, races, and colour variations possible. They cover a bewildering range of plumages for species that can be annoyingly difficult to identify in the field. This makes it an almost essential book for serious bird- watchers and ornithologists and a great resource for beginners trying to grapple with the plumage variabil- ity. I do not consider this book to be a portable field guide, but a reference work. It is a little too large and Turtles and Tortoises By Vincenzo Ferri. 2002. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 Canada. 255 pages. $24.95 This compact little book is a translation of the 1999 Italian publication, Tutto Tartarughe e Testuggini. It can best be thought of as a photographic guide to 152 species of turtles — roughly half of the species alive today. The book opens with an introduction covering the biology, evolution, classification and conservation of turtles. The bulk of the book is occupied by species accounts. The species accounts are grouped geograph- ically into oceans (sea turtles) and six terrestrial areas: Palearctic, Afrotropical, Oriental, Nearctic, Neotropical and Australian regions. There are 152 numbered spe- cies accounts and additional unnumbered accounts sprinkled throughout the book. It is unclear why some species accounts are not numbered, although the un- numbered accounts do not have a colour photo (they BOOK REVIEWS 143 heavy (due to the thick, high-quality paper). More important, this would be too precious a book to risk damaging in the field. I searched through for errors and did not notice any. In fact, I was impressed by the thoroughness of the coverage. For example, the author has correctly includ- ed the spotty northern locations for Osprey in Lab- rador. However, I do have one major problem with this vol- ume. It is a companion volume to Raptors of Western North America, a slightly larger book covering 33 species. Thus, there are 23 species common to each book. The species accounts and photograph are almost identical, except that the range maps are different. This means there is a good deal of unnecessary repetition. I can only assume that some marketing guru felt that two versions would sell better than one. Combining into one book would increase the size from 544 to about 600 pages to allow for the text for two addition- al species and the eastern range maps for all species (plus a little added to the index.). This would presum- ably increase the price to around U.S.$60. Each book is a worthwhile purchase on its own. If you buy the western book you will miss the accounts for Snail Kite only. If you buy the eastern guide you will lose the accounts for nine western raptors. If you buy both you will get more than 75% repetition, in essence wasting U.S.$30, to get the coverage of one extra species plus the relevant range maps. If you can afford only one book, buy the western guide. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario, K1J 6K5 Canada do have a colour illustration) or map. The species ac- counts are brief (some less than 100 words), with most ranging from half a page to a full page in length. Each account has standardized subheadings: Family, Distri- bution and habitat, and Characteristics. Some of the accounts also have a “Situation” subheading describing conservation issues. Perhaps because it is a translation, this book is plagued with errors. Translation-type errors include some unusual common names. For example, the Bog Turtle (Clemmys muhlenbergii; now Glyptemys muh- lenbergii) is called Muhlenberg’s Turtle. That may well be the English equivalent of the Italian common name, but that term is not used in North America. Similarly, the Painted Turtle (Chrysemys picta) is referred to as the Painted Tortoise. In addition, the genus is mis- spelled “Chrysemis.” Errors in content also abound. The author asserts that Eunotosaurus is the earliest known ancestor of turtles, yet this theory is no longer 144 widely accepted. It is also stated that some turtles can hibernate for 2-3 months, a gross understatement of the 6+ months that some turtles spend in hibernation. The species accounts are also fraught with errors or misleading information. For some North American species, the author lists individual U.S. states where the species is present, yet the list is rarely complete. The Blanding’s Turtle (Emydoidea blandingii) account omits any mention in the text or the map of the Nova Scotia population. The author accepts the Mississippi Map Turtle (Graptemys pseudogeographica kohni) as a species (G. kohni), even though the species account THE CANADIAN FIELD-NATURALIST Vol. 118 suggests it is a subspecies. There are also some curi- ous omissions. There are six species accounts for mem- bers of the genus Graptemys yet the most wide-ranging species, the Northern Map Turtle (G. geographica) is not included. Overall, one has the sense that the text was assembled quickly, or by using outdated reference materials, and that it was not reviewed by a scientific expert. Enjoy the photos but don’t believe everything you read. DAvID SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1SO Canada What Good are Bugs? Insects in the Web of Life By Gilbert Waldbauer. 2003. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts, USA. 384 pages. U.S'$29295 Waldbauer has written several other popular books about insects, and these have been met with high acclaim; I have not read those tomes, and so see his most recent work with unbiased eyes. What good are bugs? focuses on the interactions insects have with other animals and plants, both living and dead. Several chapters in each category illustrate the myriad ways in which insects, wittingly or unwit- tingly, are key to earth’s ecosystems as we know them today. Topics range from seed dispersal to recycling dead animals and the control of animal and plant populations. Truly, the scope of this book is close to all-encompassing, and is written in a style that will not be condescending to the informed naturalist or biologist, but will be readily consumable by the bud- ding naturalist, too. There are a few factual problems, as well as some opinions that are more typical of the entomophobic component of the public, that were surprising and dis- appointing to see in a book which promotes insects. Several times Waldbauer has insects “attacking” plants, when he really means “feeding on” plants. No one would ever say that deer or rabbit attack plants, it’s no different with insects; they are just feeding on the plants, a point that naturalists, biologists, and surely this author should understand. “Attacking plants” is an expression used by certain people or industries to incite action, to justify eradication, to gain sympathy from the uninformed, and should itself be eradicated from our vocabulary. The book ends with a chapter-by-chapter listing of selected readings. I like this method of listing refer- ences since the reader can readily choose among works only in the topic of interest. Overall, this is a book well worth having. RANDY FE. LAUFF Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada Belugas in the North Atlantic and the Russian Arctic Edited by M.P. Heide-Jorgensen, @.Wiig, and D. G. Pike. 2002. NAMMCO Science Publication 4, The North At- lantic Marine Mammal Commission, Polar Continental Centre, N-9296, Tromso, Norway NOK 150. 270 pages. The greater part of this symposium is devoted to those beluga whales that migrate through Baffin Bay and Davis Strait between the eastern part of the Canadian Arctic and West Greenland, and are difficult to follow across the deep water (and impossible in the dark season). Moreover, it is a difficult publica- tion to review because it represents “work in progress’’, the results of which are not yet fully understood by the authors themselves. Three main techniques of study were (1) mitochondrial molecular genetics from tissue sampling of restrained or dead animals; (2) aerial pho- tographic surveys; (3) tracking individuals by means of satellite radio tags attached to implants in the dorsal ridge of temporarily trapped belugas. The results are too tentative to summarize easily. Many groups of bel- ugas observed in summer in arctic estuaries are essen- tially matrilines, that is, adult females with young ani- mals of both sexes, while most adult males may move independently in different ways. There is also the prac- tical concern that catches of belugas at southwest Green- land, where there is most open water and the largest catching boats, appear to exceed recruitment. The exact area from which these animals come is unknown. After this it is a pleasure to move on to simpler studies! An overall survey of Russian and Siberian arctic waters from the mainland was carried out in the course of ice studies for belugas, narwhals and Greenland or Bowhead whales, plus the few Grey whales that enter the Arctic Ocean. There is now little hunting for them in this half of the Arctic. Lastly, a summary is given of up-to-date knowledge of the numerical status of belugas that inhabit, year- round, the less than 200 linear km of the St. Lawrence 2004 estuary in Quebec. This is the most biologically pro- ductive part because of year-round vertical water mix- ing. Aerial photographic surveys of numbers are there- fore simple to carry out, but it is necessary to measure the percent of animals diving too deeply to be regis- tered visually or by camera form the air. M. C. S. Kingsley and team did this using a helicopter hover- ing at an altitude that did not disturb the animals. The correction factor turned out to be more than two, so that estimates of numbers needed to be more than doubled, giving a total of more than 1200. Moreover, a series of estimates made since even casual hunting ended, ca. 1979, up to the last recorded survey in 1998, showed that the population has been growing slowly. I add to this review some more recent information that several belugas, including one radio-tracked animal, moved in autumn 2002 from SE Hudson Bay to off The Firefly Encyclopedia of Insects and Spiders Edited by Christopher O’Toole. 2002. Firefly Books Ltd., Willowdale, Ontario I will first state my biases and say that I’m not a big fan of encyclopaedias. Having got that out of the way, I must say this book was engaging, excellently laid out, and an absolute joy to read. The photographs ranged from very good to excellent to amazing. The full-frame images of even small insects startled me with their clarity. Diagrams were liberally used, effective, and also of exceedingly high quality. Each chapter was written by one or more experts, and I enjoyed reading them all. The majority of insect orders are covered, as are most of the arachnids (not just the spiders as the title alludes). Although this book could not serve as a university text in entomology, if I were to offer a bug course at the high school level, I would use this book. I have seen many entomology texts, and none are as visu- ally appealing as this book. This book would not sub- stitute for a field guide because of its size, but the qual- ity of the text, illustrations and photos far exceeds any of the field guides I’ve used. Interspersed with the taxonomic accounts which dominate the book were photo stories, fact files and special features. These articles ranged from less than a page to two pages and provided a different tangent on insect and arachnid life. As with the rest of this book, these articles were well done. BooK REVIEWS 145 Nain, Labrador (D. W. Doidge, personal communi- cation), showing that therefore some animals “bleed” from the Arctic into the Labrador Current. These must be the source of infrequent but not rare animals which reach the east coast of Newfoundland (Curren and Lien. 1998. Canadian Field-Naturalist 112(1); 28- 31). These were noted by Kingsley in this book, but have not been tracked further. Animals coming from the St. Lawrence estuary are also known to “bleed” into waters around the Maritime Provinces as far as the Bay of Fundy and even to New Jersey (various references). It therefore seems very unlikely that the St. Lawrence population is isolated genetically from the main Arctic population of belugas. D. E. SERGEANT Box 745, Hudson Heights, Quebec JOP 1JO Canada The weaknesses were very few. My underlying feeling was that I couldn’t be certain as to the in- tended audience. The wording is usually so straight- forward that children from eight or nine years old should be able to read it; however, technical ter- minology creeps in now and again, and would likely give those same early readers quite a pause. For the most part, the technical lingo is kept to a minimum, and is often isolated from the main text in‘ sidebars. This book is billed as an encyclopaedia, and the publishers I think were striving to market the book to a wide audience. My only quibble about the actual content was that the minor taxa were mentioned only in a list along- side the more dominant taxa, and not given their own words or photographs to describe them. I realize that no book that features the ricinuleids more than the beetles will ever sell, especially to the general public, but a photograph and a quarter page or so of text for each of these minor taxa would have been appropriate. Overall though, a great reference book, with won- derful visual appeal. RANDY F. LAUFF Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5 Canada 146 BOTANY Biotic Forest Communities of Ontario By Norman Duncan Martin and Norma M. Martin. 2003. Commonwealth Research. Belleville, Ontario, Canada. 195 pages. $10.00 paper. Martin and Martin set out to identify the basic for- est communities of Ontario, describe their biotic com- position, and consider the successional relationships among them. While they draw on an impressive amount of personal research and a broad literature survey, the book lacks focus and therefore may have a limited value to a general audience. A fundamental component of any scientific study is that the work must be repeatable. When presenting study results it is therefore essential that the methods are Clearly laid out. This is not the case here. The read- er is informed that the authors tallied vegetation in transects in various types of forest. A single map of transect locations is provided, with numerous exam- ples of data sheets. No details are provided to explain the selection of transect locations nor which aspects of the vegetation were sampled or how. There is also no explanation of the analysis itself, other than to acknowledge the influence of the “schools of inter- pretation” of Clements, Curtis, Hills, Whittaker and others. These were indeed important workers in this field, but listing their names doesn’t allow the reader to critically examine the results of the current study. The bulk of the book is devoted to descriptions and tables illustrating the ten forest community types identified by the authors. There are interesting obser- vations here, but the sheer quantity of poorly organized data makes it difficult to appreciate. Most of the data is presented as tables showing various measures of THE CANADIAN FIELD-NATURALIST Vol. 118 abundance in selected samples. The only graph pre- pared by the authors is labelled as showing a “dis- cernable pattern with characteristic variability”. The discernable patterns are lines overlaid on the plotted points without any statistical support. This would not be acceptable in an undergraduate ecology class, let alone a scholarly thesis. It is unfortunate that the authors make no reference to Lee et al. (1998), which has become the standard forest classification system for Ontario. It may be in- teresting to contrast the ecological land classification (ELC) of Lee et al. with Martin and Martin’s forest communities. If their data could be used to refine or correct the ELC system it would be far more useful than it is as a stand-alone study. The most interesting part of this book is the rela- tionship between forest types and their fauna. The authors have collated a great deal of their own data as well as data from published and unpublished sources. A more thoughtful analysis of this aspect of their study might produce a more enduring contribution to the study of forest ecology in Ontario. Literature Cited Lee, H., W. Bakowsky, J. Riley, J. Bowles, M. Puddister, P. Uhlig, and S. McMurray. 1998. Ecological land classification for Southern Ontario: first approximation and its application. Ontario Ministry of Natural Resources, Southcentral Science Section, Science Development and Transfer Branch. SCSS Field Guide FG-02. TYLER SMITH 5900 rue Monkland, Apartment 10, Montreal, Quebec, H4A 1G1 Canada Les champignons des arbres de l’est de l’ Amerique du Nord By Bruno Boulet. 2003. Les Publications du Québec, Sainte- Foy, Quebec, Canada. 727 pages. $49.95 The author and a number of colleagues have invest- ed a considerable effort to produce this detailed treatise on the principal wood-inhabiting fungi (mostly poly- pores) of eastern North America. The 19 x 24 cm, hardcover book is heavy but the binding is robust. The book contains nearly 500 color pictures, typically 6 or 7 on a page, of very good quality. There are three principal sections. First there is a brief commentary on the vegetation of Quebec, espe- cially in relation to the geographic and host distribution of the wood-inhabiting fungi. Then 118 pages are devoted to a discussion of the state of our knowledge of the polypores, including their medicinal properties, edibility, and traditional and commercial uses. The role of these fungi as forest pathogens and their influence on the management of forests are reviewed and various tree defects caused by these fungi are depicted in color. The second section of the book introduces the reader to the taxonomy, nomenclature and classification of the polypores, as well as their distinctive macroscopic and microscopic features. The latter are in some in- stances essential in naming specimens. This section concludes with an explanation of the most common means of identifying and naming a specimen, i.e., the botanical dichotomous key, and a Key including over 400 taxa. About 120 of the 400 are mushrooms that are briefly described in the key and 76 of them are shown in a color photo. Many are truly wood-inhab- iting fungi but a few are mycorrhizal (Lactarius and Russula) and others are incidental inhabitants of well- rotted woody debris. The third and the major part of the book presents essential information that allows recognition of 169 species, observations on their biology and ecology, color pictures, and distribution maps. Readers should be cautious in their interpretation of the distribution maps because when I compared 8 maps with distri- butions in two references from the bibliography, 1.e., 2004 numbers 142 and 145, each map was deficient. Two specific examples are (1) reference 142, titled Alba- trellus in Michigan, includes A. caeruleoporus, A. confluens, A. cristatus and A. peckianus but Michigan is not shaded on the maps for those species, and (2) shading on the map for Sistotrema confluens covers southern Quebec and Nova Scotia but reference 145 notes its presence in those provinces as well as in Michigan, North Carolina, New Hampshire, New York, Vermont and Wisconsin. Two new species, Auricularia americana Parm. et I. Parm. ex Audet, Boulet et Sirard and Polyporus long- iporus Audet, Boulet et Sirard, are proposed. Several species are reported for the first time in North America, for example, Antrodiella pallasii, Postia alni, P. folli- culocystidiata, P. ptychogaster, Phellinus cinereus and Polyporus tubaeformis. Although two names, Postia minisculoides and P. subpendula, are proposed as new combinations, if their basionyms, i.e., the initial name Book REVIEWS 147 given to the fungus and its place of publication, are not in the book the new combinations are not validly published. There is a picture (plate 15 D) labeled Punc- tularia strigosozonata, that shows a typical fruit body of the orange crust fungus, Phlebia radiata. The book concludes with a glossary containing ap- proximately 250 terms with their English equivalent and a definition that often includes a reference to a page where the term is used or illustrated, an index of French and English common names, an index of sci- entific names, and a bibliography of 450 entries. The book is recommended to mycologists, forest pathologists, forest ecologists, and naturalists. It is a significant contribution to our knowledge of the wood- inhabiting fungi of eastern North America. J. GINNS 1970 Sutherland Road, Penticton, British Columbia V2A 8T8 Canada Manual of Vascular Plants of Northeastern United States and Adjacent Canada: Second Edition By Henry A. Gleason and Arthur Cronquist. 2004. The New York Botanical Garden Press, 200th Street and Kazimiroff Boulevard, Bronx, New York 10458-5126 USA. 993 pages. U.S.$69.00. Cloth. The second edition of this most useful flora was first printed in 1991. Subsequent printings have taken place in 1993, 1996, 1998, 2000 and 2002. The seventh printing which has a slightly larger page (15 cm x 23 cm, rather than 14 cm x 21.5 cm) has a slightly larger type- face. The text pages have exactly the same page numbers as the earlier printings. Some corrections were made in the 1993 volume. In 1999 nineteen individuals con- tributed numerous corrections but these corrections could not be included in the 2000 and 2002 printings without having an electronic version. One was finally made for the 2004 printing. This new volume has an interesting but almost hid- den drawing of a Tulip-tree, Liriodendron tulipifera, on the front cover. The introductory pages start with a Table of Contents which includes a list of the families in taxonomic order with their page numbers. This is followed by a Foreword by Patricia K. Holmgren and Noel H. Holmgren and short bibliographies of Henry Allan Gleason and Arthur Cronquist by Noel H. Holmgren together with photographs of them. The Glossary which preceded the synoptic keys in the earlier printings now follows the main text. This is followed by the Index to Common Names and the Index to Scientific Names which in the earlier volumes were combined. This new volume is a most welcome step ahead with the numerous changes and corrections, and The New York Botanical Garden Press is to be congratulated even though it is still called the Second Edition. WILLIAM J. Copy Biodiversity, National Program on Environmental Health, Agriculture and Agri-Food Canada, Research Branch, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada The Wild Orchids of North America, North of Mexico By P. Martin. 2003. University Press of Florida, 15 Northwest 15th Street, Gainesville, Florida. USA. $27.95 paper, U.S.$45.95 cloth. This book is a special version of a check list. The orchids included are all the recorded species found above the U.S.—Mexican border north to the Arctic and Greenland. This covers 223 species plus 24 sub- species and varieties. Additionally, this takes in 103 growth and color forms, 24 hybrids, and introduced species. The species are arranged alphabetically by sci- entific name, so the first entry is the charming little Spotted Orchid Amerorchis rotundifolia. The author gives the genus, synonyms, misapplied names, typical common name, references and range. The author also adds any appropriate comments. For each species there is a5 x 7 cm photograph of the flower and a line draw- ing, generally of the whole plant. In some cases the line drawing is of the flower only which is a duplication that does not add information. In addition to the over 60 species that can be found in Canada, and the familiar genera (Platanthera, Cyprepedium etc.), there are several genera that | normally associate with the tropics (Vanilla, Laelia, Epidendrum). While many of these are escapees from cultivation, there are a number that are native species. Not surprisingly, most of these tropical epiphytes are to be found in Florida. Indeed, I was surprised to see 148 how many species of Vanilla (of vanilla ice-cream fame from Vanilla planifolia) that are native to Florida. The author uses the term “waif” to denote random individual occurrence. An example of a waif is Laelia rubescens, first seen in Florida in 1999. This abundant and attractive Central American species is a popular garden plant and a likely escapee. The accompanying photograph shows a white blossom. All the wild L. rubescens | have seen were pale lavender. The species coverage and their current status are both accurate and up-to-date. This book contains the split between the Spotted Orchid, Cyprepedium gutta- tum (Alaska and NWT), and the Yellow Spotted Orchid, C. yatabeanum (Kodiak Island). It includes the New- foundland orchis (Platanthera albida) of Newfoundland and Greenland. The book does not contain any habitat information, nor anything of the plants’ biology. Gen- erally, nothing is given on abundance and the distribu- THE CANADIAN FIELD-NATURALIST Vol. 118 tions are very generic (e.g., Manitoba east to New- foundland south to Texas and Georgia). It would be exciting to see an expanded version of this book con- taining descriptive text on habitat and biology, accom- panied by useable range maps. This would create an encyclopedia of North American orchids. In the mean- time, this book is a very handy reference and will make a good field guide. The book’s size precludes it fitting a pocket, but it will slip easily into your back pack. As well as the usual glossary and bibliography, the author includes a well-organized key. This, combined with the clear format, makes it a no-nonsense, practical guide for botanists, naturalists and orchid enthusiasts. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1iJ 6K5 Canada Trees of the Carolinian Forest: A Guide to Species, Their Ecology and Uses By Gerry Waldron. Boston Mills Press, 132 Main Street, Erin, Ontario NOB 1TO Canada. 274 pages. $24.95. The Carolinian zone of southern Ontario is home to more than 1600 plant species. It is also the most dense- ly populated area of the country. As a consequence, Canada’s most biologically diverse forests are also among the habitats most threatened by development. Anyone with an interest in the appreciation and con- servation of this natural heritage will do well to read Gerry Waldron’s Trees of the Carolinian forest. Mr. Waldron has succeeded in producing a beautiful book that is at once a pleasure to read and quite informa- tive. The subtitle, “A guide to species, their ecology and uses” is somewhat misleading — this is much more than a field guide. Waldron draws on a variety of his- torical and scientific sources to set the scene: what is the Carolinian zone? how is it related to other eco- logical regions? and how have successive human cul- tures altered this region? With the ecological context established, Waldron treats each of 73 tree species not as individuals but as members of a community. The book starts with an ecological history of the Carolinian zone, from glaciation through settlement by indigenous and European humans up to the present. While far from an presenting an exhaustive review of post-glacial colonisation, Waldron includes enough detail to distinguish his treatment from the usual cli- chéd summary that appears in books of similar scope. We learn of the massive hemlock dieback 5000 years back, and that beech was averaging 20 kilometres per century as it crept north. This is a fascinating subject, and it’s a shame that Waldron includes no references to his sources here. While in-text citations would be overly pedantic, including a few key references such as Pielou (1991) would be worthwhile. Waldron uses quotations from the journals of early settlers and survey crews to illustrate both the appear- ance of the “virgin” forest and the attitudes of Euro- peans to their new homeland. Once again, he piqued my curiosity, but in this case there are perhaps no read- ily accessible published sources he could refer the reader to for further information. A short discussion of the definition of the term “Carolinian” follows. Waldron covers the topic in five pages — a clearly presented summary of an important concept. This is typical of the book as a whole. The author quite capably distills complex ideas into simple language, without sacrificing accuracy in the process. With the context established, Waldron devotes some forty pages to a discussion of biodiversity, ecological communities, and our role in their protection, and especially, their restoration. He obviously brings a lot of experience to bear on the subject. Most books on restoration focus on technical details — how, where, and when to plant a tree, etc. Refreshingly, Waldron starts by examining why (and why not) to plant trees as part of a restoration. In a region where restoration of endangered prairie habitat often begins with the removal of trees planted as part of misguided natu- ralization programs, this is an important discussion. That said, he acknowledges the difficulty in pursuing a “do-nothing” approach, and offers suggestions for accelerating natural successional processes. These are presented as ideas to consider, not as ready-made pre- scriptions for restoration success. And so it is that on page 112 of this tree guide that the actual species treatments begin. Each species is allotted two pages. This includes the usual description of their habitat and appearance, with insights into their use in restoration and urban plantings. Unfortunately, Waldron has chosen to arrange the species alphabet- ically by common name. This may simplify things for the beginner, but it has the unfortunate consequence of separating walnut (Juglans nigra) from butternut (Juglans cinerea) and poplar from aspen (both Popu- 2004 lus). Taxonomic arrangements are a very useful tool for learning about the relationship between species, but only if we use them! The book is rounded out with 32 colour plates. These include a variety of habitat, habit, and close-up photographs of Carolinian trees. Some of them are quite instructive, such as the close-ups of the bark cross-sections of red and white elm, or the collection of acorns from different oak species. Others are simply beautiful images of the authors favourite organisms. One small improvement would be the inclusion of a scale for the close up shots. Taken as a whole, this book is an incredible resource for naturalists in southern Ontario. It will serve equally ENVIRONMENT Book REVIEWS 149 well as an introduction to the ecology of the Carolin- ian forest for the general naturalist and as an idea book for restorationists and land managers. I look forward to the publication of a companion volume on the shrubs of the Carolinian forest, hinted at in the introduction. Literature Cited Pielou, E. C. 1991. After the ice age: the return of life to glaciated North America. University of Chicago, Chicago. TYLER SMITH 5900 rue Monkland, Apartment 10, Montreal, Quebec, H4A 1G1 Canada Good News for a Change: How Everyday People are Helping the Planet David Suzuki and Holly Dressel. 2003. Greystone Books, #201 — 2323 Quebec Street, Vancouver, British Columbia V5T 4S7 Canada. 399 pages. $24.95, U.S.$16.95 This is a paperback re-issue of Suzuki and Dressel’s 2002 book originally titled Good News for a change: hope for a troubled planet. Either way, the book is cleverly titled to attract readers who are tired of doom and gloom environmental books. The concept is praise- worthy: bring together in one place success stories of the many and varied ways people around the world are changing the way they do things, for the benefit of their communities, their environment and their offspring. At the same time, the authors do not shy away from the bad news. Yes there are wonderful changes taking place but they are up against very seri- ous problems and a frightening inertia embedded in our social and economic systems. The authors have set a tremendous challenge for themselves in tackling the full sweep of environmental problems we face and in trying to represent emerging solutions from around the planet. They organize this unwieldy subject thematically with chapters on busi- ness practices, democracy, biodiversity, water, food, forests, fisheries, and air pollution. The final chapter “Breaking out of the Box” addresses the effect of our global culture on humans as natural beings and the importance of reconnecting what we do with our deep- est values. Most chapters focus on one or two case studies examined in some depth with additional exam- ples of related projects in other parts of the world. Examples are drawn from village co-operatives, family- run businesses, multi-national corporations, farms, in- digenous societies and non-profit groups from such diverse places as India, Africa, Germany, Brazil, USA, Indonesia and many others. Also included are a list of organizations to contact, detailed endnotes, a brief bibliography and a reasonably detailed index. The book is a little lax in explaining the political back- ground to some of its examples, especially Canadian ones, and a map plotting the locations for major exam- ples would have been beneficial. Throughout Good News there is an admirable effort to synthesize, and several key themes are followed. Aligning our activities with natural environmental systems results in double dividends — the costs are lower, the benefits are greater and extend beyond eco- nomics. A resource is used sustainably when it is man- aged by a stable community that is economically de- pendent on it and exerts local control and local ownership. Sustainable practices must be tailored to the specific situation; the people who follow them are humble and are constantly learning from their mis- takes. Many small projects are more effective and more efficient than a single large one. We are all indigenous to this planet. If I have any quibble with this book it is that there isn’t enough of it. I want to know more about how Judy Wicks manages to make a living (and a rather substantial one) running a social activist restaurant. I want to understand how a small check dam on an in- termittent stream in India can reverse desertification caused by forest removal. I want to hear from the efforts that haven’t worked, because I don’t believe as easy at the book makes it seem. There is no question we need more books like this, perhaps ones targeted at specific issues. Henry Mintzberg (1994) argues for the importance of identifying “emergent strategies” (new strategies that emerge spontaneously at any level of an organization) and helping them spread. That in essence is what Suzuki and Dressel have done. Literature Cited: Mintzberg, H. 1994. The Rise and Fall of Strategic Planning. Free Press, Simon and Schuster, New York, USA. 454 pages. CAROLYN SEBURN Seburn Ecological Services, 920 Mussell Road, RR 1, Oxford Mills, Ontario KOG 1S0 Canada 150 Whose Bird? By Bo Boelens and M. Watkins. 2004. Yale University Press, P.O. Box 209040, New Haven, Connecticutt 06520-9040 USA. x + 400 pages U.S.$35 paper. Have you ever been curious as to who was the Bonaparte of Bonaparte’s Gull fame? Was it really the feared Corsican tyrant? Now you can get a book that will answer that question and many more. The authors have researched about 1400 people who have, at one time or another, given their names to bird spe- cies. There is a cameo biography for each individual. As the book gives the accepted English names, there is a very slight bias to the coverage. European and American naturalists dominate the list, but to be fair Professor Ijima (jima’s Warbler, Phylloscopus tjimae) gets his credit. Montezuma does not fare as well. He gets a single sentence biography stating he was Em- peror of the Aztecs. Actually, Montezuma I, Emperor of Mexico, was a remarkable man. He was a victorious general before he succeeded his uncle as emperor in 1436. He is only recognized for his Quail, with no mention of his strikingly handsome Oropendola. In contrast, Californian Alberto Treganza (Treganza’s — now Great Blue — Heron) gets almost a whole page. Many of the names have fallen by the wayside. The Bancroft’s Night Heron is now called by the more descriptive name of Yellow-crowned Night Heron. The loss of some names is. a benefit. | would have trouble spelling and pronouncing Sjéstedt (Sjostedt’s Owlet, now Barred Owlet). No disrespect to Mr. Helmut Sick (Sick’s Manakin), but I think the Golden-crowned Manikin would prefer its new name. However, I am sad to see Lear’s Macaw is now an Indigo Macaw. Edward Lear, mostly known for his nonsense poems, was a remarkable Victorian bird artist who deserves a memorial. William MacGillivray’s name lives on as THE CANADIAN FIELD-NATURALIST Vol. 118 MacGillivray’s Warbler (Oporornis tolmia) but his son’s bird, MacGillivray’s Petrel, is now the Fiji Petrel, but scientifically is Pseudobulwaria macgillivrayi (The generic name honours the Reverend James Bulwer.). Some of these names are still current. Thekla Lark, named after the dying daughter of a German natural- ist is still a Thekla Lark. Of the 17 species named for the pioneering Dutch ornithologist Coenraad Temmink, six still remain. I am delighted that the Adélie Penguin’s name remains intact as it really suits this charming bird. Be thankful that Admiral Jules-Sebastien-César Dumont D’Urville named it for his wife and not himself. There are some odd stories in the book. I was par- ticularly titivated by the tale of Colonel George Montagu (Montagu’s Harrier) who was cashiered from the British army for “provocative marital skirmishing” and thereby took up the study of science. I will never look at the harrier the same way again! There are some names that are conspicuously absent. Roger Tory Peterson, Peter Scott (Scott’s Oriole is named for Winfield Scott) and James Fisher were probably born too late in history. Linneus or Linné, who invented the nomenclature system, was alive in the right era, but remains un-honoured. As well as the cameos, the authors have collected together those people who shared a characteristic in common, such as diplomats, Germans or those who died of gunshot. Many of the entries have a black- and-white portrait of the person featured. For those of you who love trivia or are keen naturalists preferably both, this is a fun book. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Forest Dynamics and Disturbance Regimes: Studies from Temperate Evergreen-Deciduous Forests By Lee E. Frelich. 2002. Cambridge University Press, 40 West 20th Street, New York, New York 10011-4221 USA. 266 pages. GBP50 (U.S.$80). Disturbance is ubiquitous in forest ecosystems. Forested landscapes are best viewed as an integration of climatic, biotic, edaphic and geomorphic processes that determine the character of disturbance events occurring Over a wide range of temporal and spatial scales. Disturbed by the extremes of either catastroph- ic, stand-replacing events that may include fire, in- sect outbreak, and extensive windthrow, or periodic, small-scale gap processes mediated by fungal patho- gens, forests are in constant flux when viewed from a landscape perspective. Such a wide range in the per- iodicity, intensity and scale of disturbance events, and the diversity of bio-edaphic interactions create a complex, fluid, heterogeneous landscape. Lee Frelich, founder and director of the University of Minnesota Center for Hardwood Ecology, intro- duces the reader to the significant disturbances that have shaped, and continue to shape, the hemlock- hardwood forests of the northern regions of the Lake States (Minnesota, Wisconsin, and Michigan). For the past two decades Dr. Frelich has dedicated himself to understanding the stand- and forest-level dynamics of these deciduous-to-boreal transition forests. He forms part of a long tradition of university and govern- ment (United States Forest Service) forest ecology research, much of which is scattered in scientific jour- nals and government reports. Forest Dynamics and Disturbance Regimes provides for the first time, in an engaging, well-illustrated, and synthetic format, the fruit of this rich research legacy. 2004 “Under what conditions do forests change or stay the same?” Thus might one summarize the intent of this book. Introductory chapters set the scene by des- cribing the Great Lakes temperate forests and their disturbance regimes dominated by fire, wind, insect outbreaks and mammalian herbivory. Of significance to the practicing forest ecologist will be the chapter on sampling and interpretative techniques used to detect and interpret forest disturbance regimes. Emphasis is placed on the use of tree radial increment patterns as a valuable source of insight into stand disturbance history. Frelich emphasizes the critical role played by dis- turbance in both stand development and forest suc- cession. He properly distinguishes between stand development and succession, both of which are often confused in the literature. Disturbance will always ini- tiate a new cycle of stand development in the regen- erating, post-disturbance forest. However, disturbance may or may not initiate a species change or a new successional sequence. Consideration is also given to the differing effects of disturbance at both the stand- and landscape-level. This distinction is important, especially given the wide temporal and spatial scales at which disturbances may occur. Furthermore, instability at the stand level may be interpreted as stability at the landscape level. Inter- pretation often depends on the scale of investigation. A particular strength of this work is Frelich’s ability to engage the complex interaction of different distur- bances. Frelich not only introduces the wide diversity of temporal and spatial patterns of forest change, but even more importantly, highlights often counter-intuitive insights into forest change and continuity. I found the following particularly noteworthy: (1) the nonlinear response of forest species composition to disturbance severity, (2) the cause and development of patchy hard- wood-softwood mosaics, (3) clarifying taxonomy of the concept of old-growth, (4) how different forest types BooK REVIEWS 15] can exist on relatively homogeneous sites, and (5) the multiple successional pathways open to any particu- lar forest type. Frelich’s final chapter summarizes the notion of forest stability. It provides conceptual models of forest response to disturbance, 3-D models of suc- cession in different forest types, and a final classifica- tion of four different types of forest landscape. This work is particularly important as humans con- tinue to “disturb” forests, especially by commercial forestry. Before any claims can be made about the desirability of the changes created by human interven- tions, it is essential to properly comprehend the range of natural forest disturbance regimes and the associ- ated changes in forest structure and tree species com- position. The book addressees the scientific community and would properly of greatest interest to forest ecologists and all students of forest change. The judicious mix of empirical case studies, hypothetical examples and conceptual models helps the reader to think “beyond the box.” The many line drawings, flow charts and black-and-white photographs help to clarify the dif- ferent concepts. As one is reminded in the subtitle, this book focuses exclusively on the temperate evergreen-deciduous for- ests (of the Lake States). While it is certain that many of the concepts developed from research in this forest type are applicable to other forest types, it is wise to resist any quick and easy transfer of ideas. Forests grow in conditions that span a wide ecological spectrum, a situation that often resists our human tendency to categorize and classify. Be that as it may, this book provides rich and substantive insight into this well- studied — and much-loved — forest region at the decid- uous-boreal interface. JOHN McCartny, S.J. St. Mark’s College, University of British Columbia, 5935 Iona Drive, Vancouver, British Columbia V6T 1J7 Canada Snowshoes & Spotted Dick; Letters from a Wilderness Dweller By Chris Czajkowski. 2003. Harbour Publishing Co. Ltd., PO Box 219, Madeira Park, British Columbia VON 2HO Canada. 298 pages. Paper U.S.$24.95 This book is a one-sided account of the wilderness experiences of a woman of extraordinary independ- ence, as she writes letters to a friend named Nick. Chris Czajkowski is an author and wilderness guide who has chosen a life of relative and geographic iso- lation on remote property where she lives three quarters of the year in the coast mountains of British Colum- bia, 480 kilometres north of Vancouver. There is no telephone or mail delivery and the radio only works on clear days. The computer used while writing the book draws the electricity required to function from solar panels. The visitors to the eco-tourism business “Nuk Tessli Alpine Experience” are brought in by float plane, along with any supplies and news from the outside world. This, the fourth book by the same author, is her account of the incredible effort to build a third cabin on the property. Its format of letters scribed to a friend in Germany is generously interspersed with some textual description, a few black-and-white pictures and hand- drawn sketches. The reader is drawn in by expressive language evok- ing in the imagination pictures of the breathtaking vistas, and natural surroundings so that you feel almost as though you have been there, perhaps as a visitor to “Nuk Tessli”. The added mystery of “what is Spotted Dick?” is a cute grab but is just a little overdone. The easy flow of the language makes the 298 pages an easy read, despite the occasional construction jargon. sy The actual process of building a log cabin in the woods, particularly in such a remote location, is fascinating. The cabin is built in several steps, over numerous months, using hand tools, chainsaws, and, occasion- ally, some friends. There are a few drawbacks from the enjoyment of this book though, especially without having read the first three. The first is that in this book there are a number of interesting stories that are alluded to but not told in detail because they’ve been recorded in the previous books. The second is that the letters are all to Nick and because it does not include the letters written from him, it feels a bit like being in the same room with someone on the phone — you hear half the conversation and although you get the gist of the THE CANADIAN FIELD-NATURALIST Vol. 118 whole, you feel the missing of the other half. The third and most perplexing is the lack of description of the characters or their relationship with each other. The reason this is perplexing is because the author herself describes this criticism received from a book review- er — and then simply passes it off with “The trouble is, I live with the people I write about. Experience has shown that no matter how innocuous a portrait I paint, the subject will find something about which he or she is unhappy.” While I am sure this could be true, the book would earn a wider audience if it were not written for those people alone. DAWN BURNETT Jacques Whitford Limited, 1 Union Street, Elmira, Ontario N3B 3J9 Natural Grace: The Charm, Wonder, & Lessons of Pacific Northwest Animals & Plants By William Dietrich. 2003. University of Washington Press, 1326 Fifth Avenue, Suite 555, Seattle, Washington 98101- 2604 USA. 236 pages. U.S.$19.95 Natural Grace is a contemporary example of the medieval book of beasts called a bestiary. Bestiaries were very popular during the Medieval Period, and focused on the life of nature as a model or paradigm for human behavior. These books were not scientific in the contemporary sense of the term, but instead combined religious and moral teaching with a close observation of nature, zoological commentaries, and fabulous and fictitious creatures. Rather than being studied in and of itself, nature was considered symbol- ic of both the virtue and moral life of human beings. While Deitrich’s work certainly provides the reader with a more sophisticated scientific understanding of nature, the subtitle of the book betrays its “bestiary” lineage as the reader is invited to appreciate the charm, wonder, and lessons of a variety of creatures and natural phenomena. This collection of essays is adapted from William Dietrich’s popular articles in the Seattle Times’ North- west Magazine. The author divides his work among four themes: (1) the common and ubiquitous creatures that we often take for granted, (2) the itty-bitty world that ranges from soil to spiders, (3) the rhythmic, clockwork world of tectonics and tides, and finally, (4) the iconic symbols of the Pacific Northwest includ- ing, among others, the bald eagle and the killer whale or orca. Dietrich is gifted with the contemplative eye. Follow his gaze and you will enter a world of delight and amazement. We live side by side with other creatures and are immersed within the workings of nature. But often, we are oblivious to the ways of our environs, so intent and fascinated are we with our own creations and fabrications. Dietrich pries apart our fabricated world and invites us to direct our gaze out beyond the confines of culture. He calls us to a long loving gaze on the other of nature. This contemplative gaze is a virtue that strengthens with time and practice. Dietrich is a patient and humor- ous guide, gently leading the reader to grow in knowl- edge of the natural world; knowledge, not simply of the biological or physical facts, but of the beauty, sig- nificance, and yes, even mystery inherent in the sub- ject of our gaze. Of special interest is the manner by which the author weaves together culture and nature. Whether it be the forces of nature that define the boundaries of human culture and provide the myriad free ecological services that allow us our cities and farms, or the manner in which particular species have become embedded in our folk and cultural lore, the reader is left with the knowledge that we are depend- ent on the “others” of nature far beyond our physical survival. Human ignorance instills fear, and fear breeds vio- lence and destruction. Natural Grace sets a contrary path. Knowledge of the other, of nature, can lead to intimacy, and intimacy can be the foundation of love. Love, in turn, leads to mutuality, care, and concern for the other. Dietrich’s melodious writing can indeed lead one to greater love of nature and hopefully, action on behalf of nature. This book is a lovely blend of nature and culture, of scientific vulgarization and social lore. You will learn much about the natural history of the Pacific Northwest. As well, I think, you will learn about how we (and maybe you yourself) relate to the natural world, and how that relationship has changed with time, and how it may or must change in the future. Given Dietrich’s place of habitation (Seattle), this book focuses on the State of Washington, with excur- sions south to Oregon and north to British Columbia. So, if you call the Pacific Northwest your home, you will learn much about your “neighbours.” However, regardless of one’s geography, Dietrich’s work is more a work of perspective, rather than of content. His actors live and work in the Pacific Northwest, but the perspective shared is universal. No matter where you 2004 live, the long, loving, contemplative gaze on the real is always a possibility. The 12" century Cistercian monk, Saint Bernard of Clairvaux, once said; “I have discovered that you will find far more in the forests than in books; trees and stones will teach you that which you cannot learn from any master.” Turn a discerning eye to the crea- Book REVIEWS 153 tures and processes that surround you and you will enter a world of marvel and beauty. Let Natural Grace be your companion and guide. JOHN McCarthy, S.J. St. Mark’s College, University of British Columbia, 5935 Iona Drive, Vancouver, British Columbia V6T 1J7 Canada Tales from the Underground: A Natural History of Subterranean Life By David W. Wolfe. 2001. Perseus Publishing, 5500 Central Avenue, Boulder, Colorado 80301 USA. 221 pages. $27.50 Tales from the Underground is a MUST read for any naturalist interested in what’s happening under our feet. The book is engaging, well-written, and filled with information that is truly awesome. In the introduction, author David Wolfe, Associate Professor of Plant Ecology at Cornell University, in- vites his “subterranean-impaired” readers to use their imaginations and join him on a dive trip into a mys- terious world scientists are only just beginning to un- derstand. Wolfe launches the journey with a description of the various soil “profiles”, also known as layers or horizons, and the organisms that inhabit them. Some of the life forms I had never heard of; others I had never thought about in the way Wolfe describes them. Take plants, for example. Wolfe starts by praising their display of foliage aboveground. He then points out that plants are unique because they simultane- ously inhabit both the surface and sub-surface realms. He calls them the “great mediators between the two realms”, and explains their functions both above and below the ground, with a focus on their buried other half: roots. In Part I of the book, “Ancient Life’, Wolfe covers a lot of ground. He discusses theories of the origin of life, the basic elements necessary for the evolution of life, Earth’s advantage as a life-generating planet, and speculation pointing to the subterranean realm as the place where life began. He also introduces “extremo- philes”, creatures that live in severe conditions thou- sands of feet underground, where pressure and tem- peratures are extraordinarily high, and where there is neither light nor oxygen — the postulated setting for the beginning of life. Wolfe also describes Carl Woese’s discovery of the amazing genetic diversity of these extremophiles, and discusses how that discovery changed the universal tree of life. Originally composed of bacteria, plants, fungi, animals and protozoa, the tree of life now has only three branches — bacteria, archaea (the extremo- philes) and eukarya (with plants and animals as two small twigs). It’s a fascinating and humbling picture. In Part I, “Life Support for Planet Earth”, Wolfe dedicates an entire chapter to Charles Darwin and the lasting influence of the renowned scientist’s meticu- lous and painstaking work. Wolfe also portrays certain inconspicuous subterranean creatures and the impor- tant roles they play in cycling essential elements and facilitating the flow of energy — processes that sustain life on this planet. He also discusses the dual roles played by soil with respect to plant and animal dis- eases; some soil microbes cause diseases, while others provide powerful antidotes. The third part of the book, “The Human Factor”, describes the enormous footprint of Homo sapiens on the planet. Wolfe tells the sobering, appalling story of human activities and their devastating impacts on wildlife such as prairie dogs, black-footed ferrets, and burrowing owls, and on soil integrity in general. Des- criptions of soil erosion and the 1930s dust bowl crisis are particularly gripping. Yet despite the doom and gloom, Wolfe ends on an optimistic note. The very last sentence of the book reads, “It is my hope that as more of us become aware of the life beneath our feet, and its relevance to our well-being, we will be inclined to work together to maintain the biological integrity of the underground, and preserve some of what we find there for future generations” — words we have heard before in one form or another, but good to see repeated in this context. Tales from the Underground is David Wolfe’s first book, and I hope it isn’t his last. He has a pleasant and easy-to-read writing style, and manages to convey sci- entific concepts so that even a non-scientific type like me can understand. Tales from the Underground has found a permanent place on my bookshelves. R. SANDER-REGIER RR5 Shawville, Quebec JOX 2YO Canada 154 NEw TITLES +Available for review * Assigned for review Zoology * Amphibian Decline: An integrated analysis of multi stressor effects. By Greg Linder. 2004. SETAC North Amer- ica, 1010 North 12th Avenue, Pensacola, Florida. U.S.$98 Cloth. Annotated Checklist of Birds of Chile. By M. Marin. 2004. Lynx Edicions, Barcelona, Spain. 144 pages. U.S.$15.75 [in English and Spanish]. Biology of Sharks and their Relatives. By J. Carrier, J. Mu- sick and M. Heithus. 2004. CRC Press, 2000 NW Corporate Boulevard, Boca Raton, Florida. U.S.$99.95. * Birds of Belize. By H. L. Jones. 2004. University of Texas Press, Austin, Texas. 317 pages, U.S.$60.00 Cloth, $34.95 Paper. Birds of South Asia. By P. Rasmussen and J. Anderton. 2004. Lynx Edicions, Barcelona, Spain. U.S.$95 Cloth. Birds of Spain. 2004. Lynx Edicions, Barcelona, Spain. 296 pages. U.S.$49 [in English and Spanish]. Birds of the Raincoast. By H. Thommasen and K. Hutchings. 2004. Harbour Publishing, P.O. Box 219, Madeira Park, Brit- ish Columbia. 260 pages. U.S.$24.95. The Exotic Amphibians and Reptiles of Florida. By W. Meshaka, B. Buttergield and J. Hauge. 2004. Krieger Pub- lishing P.O. Box 9542 Melbourne, Florida 32902-9452. 166 pages. U.S.$34.50. * Hawks and Owls of the Great Lakes Region & Eastern North America. By C. Earley. 2004. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto, Ontario, M2H 3K1. 128 pages. $16.95 Paper, $24.95 Cloth. + Orca — Visions of the Killer Whale. By Knudtson. 2004. Greystone Books, Vancouver, British Columbia. xvii +110 pages, $19.95. Pacific Reef and Shore. By R. Harbo. 2004. Harbour Pub- lishing, P.O. Box 219, Madeira Park, British Columbia. 80 pages. U.S.$9.95. Reproductive Biology and Early Life History of Fishes in the Ohio River Drainage. By T. Simon and R. Wallus. 2004. CRC Press, 2000 NW Corporate Boulevard, Boca Raton, Florida U.S.$119.95. * Self-Portrait with Turtles: A Memoir. Edited by David M. Carroll. 2004. Houghton Mufflin Co., 222 Berkeley Street, Boston, Massachusetts 02116. 192 pages. 40 black-and-white line drawings and halftones. $23.00 Hardcover. Botany * Arboretum America: a Philosophy of the Forest. By Diana Beresford-Kroeger. 2003. University of Michigan Press, 839 Greene Street, P.O. Box 1104, Ann Arbour, Michigan. 196 pages. U.S.$52.50. Botanical Latin: Fourth Edition. By William T. Stearn. 1992 [Re-issued as a paperback 2004] Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon, USA. 560 pages. Paperback, U.S.$29.95. * Manual of Vascular Plants of Northeastern United States and Adjacent Canada. By H. Gleason. 2004. New York THE CANADIAN FIELD-NATURALIST Vol. 118 Botanical Gardens, 200" Street and Kazimiroff Boulevard, Bronx, New York. 993 pages. U.S.$69.00. The Orchid in Lore and Legend. By Luigi Berliocchi. 2000 [Re-issued as a paperback 2004] Edited by Mark Griffiths. 2000 Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon, USA. 200 pages. Paperback, U.S.$19.95. The Secrets of Wildflowers. By J. Saunders. 2004. The Lyons Press, 246 Goose Lane, P.O. Box 480, Guilford, Connecticut. 304 pages. U.S.$24.95. Wildflowers of the Seacoast in the Pacific Northwest. By J. Trelawny. 2004. Harbour Publishing, P. O. Box 219, Madeira Park, British Columbia. 80 pages. U.S.$12.95. Environment Arctic National Wildlife Refuge. By S. Banerjee. 2004. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia. 176 pages. U.S.$36.95. + The Bird Almanac. By D. Bird. 2004. Key Porter Books, 70 The Esplanade, Toronto, Ontario, Canada. xvii + 460 pages, not illustrated. $24.95. * Dancing at the Dead Sea — Tracking the World’s Envi- ronmental Hotspots. By Alanna Mitchell. 2004. Key Porter Books, 70 The Esplanade, Toronto, Ontario MSE 1R2 Canada. 259 pages. $26.95. Estuarine Research, Monitoring and Resource Protection. By M. Kennish. 2004. CRC Press, 2000 NW Corporate Boulevard, Boca Raton, Florida. U.S.$119.95. Important Transboundary Belarussian-Lithuanian and Lithuanian-Russian Wetlands. By L. Raudonikis, A. Skura- tovich, L. Baléiauskas, E. Drobelis, D. Grishanov. 2003. Botanikos institutas, Vilnius, Lithuania. * The History of Ornithology in Virginia. By D. Johnston. 2004. University of Virginia Press, P.O. Box 400318, Charlottesville, Virginia. x + 219 pages. U.S.$35. The Lewis and Clark Columbia River Water Trail: A Guide for Paddlers, Hikers, and Other Explorers. By Keith G. Hay. 2004. Timber Press, 133 SW 2nd Avenue, Suite 450, Portland, Oregon, USA. 264 pages. Paperback, U.S.$19.95. * Politics of the Wild - Canada & Endangered Species. Edited by K. Beazley and R. Boardman. 2004. Oxford University Press, 70 Wynford Drive, Don Mills, Ontario, Can- ada. x + 254 pages, not illustrated. $29.95. * A Primer of Ecological Genetics. By J. Conner and D. Hartt. 2004. Sinauer Associates, Box 407, Sunderland, Massa- chusetts, USA. 207 pages. U.S.$34.95. + Resource and Environmental Management in Canada. Edited by B. Mitchell. 2004. Oxford University Press, 70 Wynford Drive, Don Mills, Ontario. x + 600 pages, $29.95. * Whose Bird? By B. Boleus and M. Watkins. 2004. Yale University Press. 384 pages. U.S.$35.00 paper. Children’s books Saving Birds: Heroes around the world. By P. Salmansohn and S. Kress. 2004. Tilbury House, Gardiner, Maine. 40 pages. U.S.$16.95. King Of Fish: The Thousand Year Run of Salmon. By E. Bauer. 2003. Voyageur, Stillwater, Minnesota. 160 pages. U.S.$29.95. News and Comment Point Pelee Natural History News 3(4) The Winter 2003 issue, volume 3, number 4, pages 53-68, contains: Point Pelee: Its Evolution and Structure (Alan S. Trenhaile) — Noteworthy Bird Records: September to Novem- ber 2003 (Alan Wormington) — More Cave Swallows: The Fall of 2003 (Alan Wormington) — Point Pelee Christmas Bird Count: December 15, 2003 (Sarah E. Rupert) — Fox Squirrels at Point Pelee: 1977, 1990 and 1997 (Alan Wormington) — In the Field (Ross’s Gull: Second Record for Point Pelee; “Harlan’s” Red-tailed Hawk: New to Point Pelee). The Point Pelee Natural History News was edited by Alan Wormington (e-mail: wormington@juno.com). Editorial Assis- COSEWIC Assessment Results November 2003 This update on Canadian Species at Risk issued by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) is 44 pages and contains: About COSEWIC, (mandate, membership, definitions) — Summary Tables [COSEWIC species designated in five “risk” categories and in Not at Risk, and Data Deficient categories (Tables 1-3)] — Results of November 2003 COSEWIC meeting (Tables 4-5) — COSEWIC Assessment Results — Explanation of sym- bols — Species examined in five “risk” categories (Table 6), Not at Risk (Table 7), and Data Deficient (Table 8) — Record tants were Gordon D. Harvey and Michelle T. Nicholson; web site www.wincom.net/~fopp/Natural_History_News.htm. This is the last issue as the Board of Directors of the publisher, The Friends of Point Pelee, has decided to terminate the pub- lication due mainly to the lack of material being submitted. All past issues are available in the Nature Nook Book Store in Point Pelee National Park’s Visitor Centre or by mail from the Friends of Point Pelee, 1118 Point Pelee Drive, Leaming- ton, Ontario N8H 3V4 Canada; phone 519-326-6173; e- mail: fopp@ wincom.net. of Status Re-examinations — Record of Name Changes. Listed are 12 extinct, 21 extirpated, 160 endangered, 108 threat- ened, and 140 of special concern. Of the 441 forms in these categories, 65 are mammals, 58 birds, 31 reptiles, 19 amphib- ians, 78 fishes, 16 arthropods, 19 molluscs, 140 vascular plants, 9 mosses, and 6 lichens. In addition, 32 forms have been considered and found not at risk, and 29 to be data deficient. This publication is available from COSEWIC Secretariat, 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 (61) Number 60, February 2004. Contents: A Workshop on the Amphibians of Madagascar at Giand (Franco Andreone) — Amphibian Colonization of Mitigation Wetlands in Nebraska (Mark M. Peyton) — Annual Meeting Report from Canada (David Galbraith) — A Study of the Plethodontid Salamander Populations at Raccoon Creek State Park, Pennsylvania (Jen- nifer D. Haney and Mary S. Kostalos) — Reports on DAPTF Seed Grants — Froglog Shorts — New Book on Amphibian Declines (Amphibian Declines: An Integrated Analysis of Mul- tiple Stressor Effects. 2003. SETAC Press, Pensacola, Florida: http://www.setac.org/pubs. html). 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, and Arizona State University. The newsletter is edited by John W. Wilkinson, Department of Biological Sci- ences, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom; e-mail: daptf@open.ac.uk. Fund- ing for Froglog is underwritten by the Detroit Zoological Institute, P. O. Box 39, Royal Oak, Michigan 48068-0039, USA. Publication of issue 61 was also supported by Peace Frogs www.peacefrogs.com and by RANA and the US National Science Foundation grant DEB-0130273. The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and Reptile Conservation Network — Reseau Canadien de Conservation des Amphibiens et des Reptiles 8(1) January 2004 Contents: Editor’s Note: thanks to sponsors TD Bank Financial Group, TD Friends of the Environment, The Pelee Island Winery, Parks Canada [Kerrie Serben] — A note about our acronym [David Galbraith] — CARCNET/RECCAR 2003: The Pelee Isand Experience [David Galbraith] — Summary of Species at Risk Proceedings from the 8th Annual Meeting of CARCNET/RECCAR and the 3rd Annual Pelee Island Winery Endangered Species Festival (ESF) [Kim Barrett and Ben Porchuk] — The Lake Erie Watersnake: A 23-year Perspective on Microevolution, Population Status, and Recovery Planning [Richard B. King] — Loving Nature to Death [Ron Brooks] — Another Leap into Learning [Sara L. Ashpole] — Books to Get: Turtles [Anita Baskin-Salzberg and Allen Salzberg]; A guide to creating vernal ponds — Endangered Northern Leopard Frog Reintroduced into East Kootenay After 20 Year Disappearance — Frogs and Toads of Canada CD — Current CARCNET/RECCAR Board of Directors — New Species of Salamander Found in Revel- stoke [Larry. Halverson] — Paper on Traffic Mortality and Turtles [reviewed by David Seburn] — Coming soon... CARCNET 2004 September 24-27 Edmonton — Member- ship in CARCNET/ RECCAR. LDS) 156 For information on membership in the Canadian Amphibian and Reptile Conservation Society/Reseau Canadien de Con- servation des Amphibiens et des Reptiles ($10 students, $16 non-students] contact Bruce Pauli, Canadian Wildlife Service, THE CANADIAN FIELD-NATURALIST Vol. 118 National Wildlife Research Centre, Carleton University, Raven Road, Ottawa, Ontario K1A 0H3. Web site: http://www.carc net.ca/ Annotated List of the Arctic Marine Fishes of Canada In 2004 Fisheries and Oceans Canada, Central and Arctic Region, Winnipeg, Manitoba, printed a limited number of copies of the Annotated List of the Arctic Marine Fishes of Canada by B. W. Coad and J. D. Reist as Canadian Manu- script Report of Fisheries and Aquatic Sciences 2674, iv + 112 pages. A primary list covers the 189 species represent- ing 115 genera in 48 families of marine and anadromous fishes in Canadian Arctic and marine waters. Given under scien- tific name, authority, and description date, are the Common Name, Provinces/Territories of occurrence, Ecozone, Distribu- tion (ocean and extralimital), Numbers, Size (maximum), and Biology (depth, food, commercial importance). An extralimitial list gives 83 species occurring in adjacent areas that might eventually be recorded in Canadian Arctic waters annotated as eastern (eastern Davis Strait, southwest Greenland) and western (western Beaufort Sea of Alaska from Point Barrow Marine Turtle Newsletter (103) January 2004. 32 pages: ARTICLES: Behavior of Green Sea Turtles in the Presence and Absence of Recreational Snork- ellers — Turtle Strandings in the Southern Eritrean Red Sea — First Confirmed Occurrence of Loggerhead Turtles in Peru — A Harness for Attachment of Satellite Transmitters on Flatback Turtles — Notes: Nesting of the Hawksbill Turtle in Shidvar Island, Hormozgan Provance, Iran — MEETING REPORTS — OBITUARIES (Henry H. Hildebrand 1922-2003; C. Robert Shoop 1935-2003) — ANNOUNCEMENTS — NEWS & LEGAL BRIEFS — RECENT PUBLICATIONS. eastwards). A brackish water list gives 36 species whose life is usually spent entirely in fresh waters but which may enter estuaries and seashore pools in the Canadian Arctic. There are 3 tables (species by province and territory, by ecozones, and by distribution (Alaska, Atlantic, Eurasia)); 2 maps (Can- adian Arctic waters and ecozones with this area); and two appendicies: one giving an alphabetical listing of scientific names of families with common names in English, French, Inuktitut and Inuvialuktun and one giving an alphabetical listing of species by scientific name with notation of new to Arctic, new to Canada, and page number in text. Copies of this report are available from Fisheries and Oceans Canada Central and Arctic Region, Winnipeg, Mani- toba R3T 2N6 Canada: e-mail reistj@dfo-mpo.ge.ca [Cata- logue number Fs 97-4/26/2674E, ISSN 0706-6473]. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Exeter, Exeter EX4 4PS United Kingdom; e-mail MTN @seaturtle. org; Fax +44 1392 263700. Subscriptions and donations towards the production of both the MTN and its Spanish edition NTM [Noticiero de Tortugas Marinas] should be made online at or c/o SEATURTLE.ORG, 11400 Classical Lane, Silver_Spring, Maryland 20901 USA. Ontario Natural Heritage Information Centre Science and Information Newsletter 9(1) Winter 2004 Contents of this 20 page issue: FEATURE ARTICLES: Decade of Achievement (NHIC 10" anniversary celebrated 9 Decem- ber 2003) — A Decade of Ontario Botanical Dsicoveries; 2003 PROGRAM HiIGLicuts: NHIC Staff Participate in James Bay Coastal Vegetation Study — Notes on the Vegetation and Flora of Flooding River Shores — A Profile of Natural Areas Recently Added to the Natural Areas Database; 2003 ProJEcT HIGHLIGHTS: NHIC Conducts Turtle Surveys — NHIC Coordinates Odonata and Reptile Surveys — Update on the Great Lakes Conservation Blueprint Projects — Identifying Key Natural Areas and Linkages in Southern Ontario — NHIC’s Southern Region Element Occurrence Prioritization Project; NEws AND NorTEs: Species at Risk: A Diversity of Designations — What’s in a Name? Changes to the Provincial Amphibian and Reptile Lists — NHIC Attends Pelee Island Events — NatureServe Canada and Parks Agency sign MOU — NatureServe Participates with CWS on Spe- cies at Risk Information Needs Analysis — Additions to the COSEWIC list — Minister visits NHIC; Book REvVIEws: NHIC STAFF INFORMATION. Mailing address for Natural Heritage Information Centre, Ontario Ministry of Natural Resources, 300 Water Street, 2nd Floor, North Tower, P.O. Box 7000, Peterborough, On- tario K9J 8M5, Canada; www.mnr.gov.on.ca/MNR/nhic.cfm. Web page: http://www.mnr.gov.on.ca/MNR/nhic.html 2004 NEWS AND COMMENT 157 Erratum Canadian Field-Naturalist 117(1) Ballard, Warren B., Matthew A. Cronin, Martin D The abbreviation for concentrations in micrograms per 5 9 . 5 a . x z ; Robards, and William A. Stubblefield. 2003. Heavy gram of dry weight should be corrected in two places. , ‘ ; . x 0 as J. c rd ae 5 te pS) =f; k Pa ; Ore metal concentrations in Arctic Foxes, Alopex lagopus, in On page 120 “mg/g” left column line 18, and right the Prudhoe Oil Field, Alaska. Canadian Field-Naturalist column line 21, should be “g/g”. 117(2): 119-121. Erratum Canadian Field-Naturalist 117(2) Lindquist, E. S., C. F. Aquadro, D. McClearn, and K J. 117(2): 184-189. McGowan. 2003. Field identification of the mice On page 4, Figure 1, 1A was repeated for 1B. The correct Peromyscus leucopus noveboracensis and P. maniculatus 1B is shown below with IA. gracilis in central New York. Canadian Field-Naturalist Distance from P. leucopus Distance from P. leucopus O 4 2 38 4 5 6 Distance from P. maniculatus Distance from P. maniculatus FiGurRE 1. Classification of P. |. noveboracensis and P. m. gracilis using the discriminant-function coefficients given in Table 2. Figure la shows the classification of adults only. Figure 1b shows the classification of adults and juveniles. Open and closed symbols denote P. m. gracilis and P. |. noveboracensis, respectively. In Figure 1b, P. m. gracilis is indicated by open circles and squares (adult and juvenile, respectively), and P. |. noveboracensis by closed triangles and stars (adult and juvenile, respectively). 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 read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, 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. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit, to the Editor, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before itis submitted. All authors should have read and approved it. Institutional or contract approval for the publication of the data must have been obtained by the authors. Also authors are ex- pected to have complied with all pertinent legislation regard- ing the study, disturbance, or collection of animals, plants or minerals. The place where voucher specimens have been de- posited, and their catalogue numbers, should be given. Lati- tude and longitude should be included for all individual local- ities where collections or observations have been made. Print the manuscript on standard-size paper, doublespace throughout, leave generous margins to allow for copy mark- ing, and number each page. For Articles and Notes provide a bibliographic strip, an abstract and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names should be omitted except in taxonomic manuscripts or other papers involving nomenclatural problems. “Standard” com- mon names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in the text or in a sepa- rate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “British” or “American” spellings are acceptable in English but should be consistent within one manuscript. 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Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for pub- lication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by paying $80 for each page, plus $15 for each illustration (any size up to a full page), and up to $80 per page for tables (depending on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Business Manager. Reprint order forms are included when galley proofs are sent to authors. If grant or institutional funds are not available, club members and subscribers may apply for a waiver of charges for the first five pages. Limited joumal funds are available to help offset publi- cation charges to authors with minimal financial resources. Requests for financial assistance should be made to the Busi- ness Manager when the manuscript is accepted. Reprints An order form for the purchase of repents will accompany the galley proofs sent to the authors. FRANCIS R. Cook, Editor 13078 Land O’Nod Road RR 3 North Augusta, Ontario KOG IRO Canada 158 TABLE OF CONTENTS (concluded) Volume 118 Number 1 Record high wolf, Canis lupus, pack density L. DAVID MECH AND SHAWN TRACY Extraordinary size and survival of American Black Duck, Anas rubripes, broods JERRY R. LONGCORE AND DANIEL G. MCAULEY Observations of interactions between Puma, Puma concolor, and introduced European Red Deer, Cervus elaphus, in Patagonia WERNER T. FLUECK Premiéres mentions de la Couleuvre mince, Thamnophis sauritus septentrionalis, au Québec JEAN-FRANCOIS DESROCHES ET RICHARD LAPARE Book Reviews ZooLoGy: Amphibian Decline: An Integrated Analysis of Multiple Stressor Effects — Birds of Africa: From of Seabirds to Seedeaters — The Firefly Encyclopedia of Birds — Birds of Belize — Herpetology: Third Edition — Lizards: Windows to the Evolution of Diversity — Raptors of Western North America — Raptors of Eastern North America — Turtles and Tortoises — What Good are Bugs? Insects in the Web of Life — Belugas in the North Atlantic and Russian Arctic — The Firefly Encyclopedia of Insects and Spiders Botany: Biotic Forest Communities of Ontario — Les champignons des arbres de |’est de l’ Amerique du Nord — Manual of Vascular Plants of Northeastern United States and Adjacent Canada: Second Edition — The Wild Orchids of North America, North of Mexico — Trees of the Carolinian Forest: A Guide to Species, Their Ecology and Uses ENVIRONMENT: Good News for a Change: How Everyday People are Helping the Planet — Whose Bird? — Forest Dynamics and Disturbance: Studies from Temperate Evergreen-Deciduous Forests — Snowshoes & Spotted Dick: Letters from a Wilderness Dweller— Natural Grace: The Charm, Wonder, & Lessons of Pacific Northwest Animals & Plants — Tales from the Underground: A Natural History of Subterranean Life NEw TITLES News and Comment Point Pelee Natural History News 3(4) — COSEWIC Assessment Results November 2003 — Froglog (61) — The Boreal Dip Net 8(1) — Annotated List of the Arctic Marine Fishes of Canada — Marine Turtle News- letter (103) — Ontario Natural Heritage Information Centre Science and Information Newsletter 9(1) Winter 2004 — Erratum: The Canadian Field-Naturalist 117(1) — Erratum:The Canadian Field- Naturalist 117(2) Advise to Contributors Mailing date of the previous issue 117(4): 16 September 2004 2004 PA 129 138 146 149 153 THE CANADIAN FIELD-NATURALIST Volume 118 Number 1 200 ——_—— Ore NET EE UUE 125 Years’ Anniversary: The Ottawa Field-Naturalists’ Club: origins and history DANIEL F. BRUNTON l Articles Continuing environmental change — an example from Nova Scotia EDMUND S. TELFER 39) Habitat segregation among songbirds using the southern boreal mixedwood forest ENID E. CUMMING 45 Effects of mid-winter snow depth on stand selection by Wolverines, Gulo gulo luscus, in the boreal forest JONATHAN D. WRIGHT AND JESSICA ERNST 56 Wolverine, Gulo gulo luscus, resting sites and caching behavior in the boreal forest JONATHAN D. WRIGHT AND JESSICA ERNST 61 Seasonal home ranges of Raccoons, Procyon lotor, using a common feeding site in rural eastern Ontario: rabies management implications SARAH C. TOTTON, RICHARD C. ROSATTE, ROWLAND R. TINLINE, AND LAuRA L. BIGLER 65 Status of marine turtles in British Columbia waters: a reassessment DONALD F. MCALPINE, STAN A. ORCHARD, KELLY A. SENDALL, AND RON PALM 2 Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) region, British Columbia N. A. SLOAN AND P. M. BARTIER Vil Ruby-throated Hummingbird, Archilochus colubris, entanglements in burdock (Arctium spp.) at Delta Marsh, Manitoba HEATHER L. HINAM, SPENCER G. SEALY, AND Topp J. UNDERWOOD 85 Multiple mating results in multiple paternity in Richardson’s Ground Squirrels, Spermophilus richardsonii JAMES F. HARE, GLENDA Topp, AND WENDY A. UNTEREINER 90 Demographic patterns and limitations of wolves, Canis lupus, in and near Pukaskwa National Park, Ontario S. ANNE FORSHNER, PAUL C. PAQUET, FRANK G. M. BURROWS, GRAHAM K. NEALE, AND KEITH D. WADE 95 Estimation of seed bank and seed viability of the Gulf of Saint Lawrence Aster, Symphyotrichum laurentianum (Fernald) Nesom — Joni FE. KEMP AND CHRISTIAN R. LACROIX 105 Consumption of shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus JONATHAN W. Moore AND G. J. KENAGY Hi Unusual behavior by Bison, Bison bison, toward Elk, Cervus elaphus, and Wolves, Canis lupus L. Davip MEcH, RICK T. MCINTYRE, AND DOUGLAS W. SMITH 115 r Notes Morphology of female Woodland Caribou, Rangifer tarandus caribou, in Saskatchewan W. JAMES RETTIE 119 Occurrence of parasitoid wasps in the egg sacs of Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in southern Idaho LisA M. COBB AND VINCENT A. CoBB 12D The Northern True Katydid, Pterophylla camellifolia (Insecta: Orthroptera: Pseudophyllidae), at Ottawa, Ontario STEPHEN J. DARBYSHIRE 124 (continued on inside back cover) ISSN 0008-3550 The CANADIAN — FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 118, Number 2 April-June 2004 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; te encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining 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. 1 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 2004 Council President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor Francis R. Cook Cendrine Huemer To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, 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. 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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 members (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: April—June 2004 (April 2005). Cover: Brown Bear, Ursus arcts, watches Gray Wolf, Canis lupus, in the Katmai National Park and Preserve in southwest Alaska. See paper by Smith, Partridge, and Shoen pages 247-250 (Kent Fredrikson photograph). The Canadian Field-Naturalist Uxiijeees2y Volume 118, Number 2 April-June 2004 Morphology and Population Characteristics of Vancouver Island Cougars, Puma concolor vancouverensis STEVEN F. WILSON!, APRYL HAHN, AARON GLADDERS, KAREN M. L. Gou, and DAvip M. SHACKLETON Wildlife Research Group, Agroecology, Faculty of Agricultural Sciences, University of British Columbia, 270-2357 Main Mall, Vancouver, British Columbia V6T 1Z4 Canada 'Present address: EcoLogic Research, 406 Hemlock Avenue, Gabriola Island, British Columbia VOR 1X1 Canada Wilson, Steven F., Apryl Hahn, Aaron Gladders, Karen M. L. Goh, and David M. Shackleton. 2004. Morphology and population characteristics of Vancouver Island Cougars, Puma concolor vancouverensis. Canadian Field-Naturalist 118(2): 159-163. Cougars are a management concern on Vancouver Island because they are a top predator and because there have been frequent attacks on humans on the island. However, little is known about Cougar ecology in the Pacific Northwest of North America. We studied Cougar morphology and population characteristics as part of a larger study in two areas on Vancouver Island. We derived a multivariate measure of body size to describe changes with age and sex. Body size was similar in the two study areas. Survival rates for adult females were higher than those reported elsewhere; however, hunters avoided shooting females in general, and radio-collared Cougars in particular. Litter size at first detection was lower than reported in many other studies and may be related to food availability. Key Words: Cougars, Puma concolor vancouverensis, morphology, survival, mortality, natality, Vancouver Island, British Columbia. The Mountain Lion, or Cougar (Puma concolor van- couverensis), 1s one of the two main large predatory carnivores in the forests of Vancouver Island, British Columbia, Canada. It also has a long history of inter- actions with humans as a nuisance (livestock predation) and as a game species (Beier 1991). The incidence of Cougar attacks on humans in North America is also highest on the island. However, little is known about the ecology of Cougar in this part of its range, mainly because this subspecies inhabits dense forests typical of coastal British Columbia. We examined Cougar mor- phology and population characteristics in two study areas on Vancouver Island to investigate the effects of age, sex, and location on body size, and to determine survival rates and reproductive characteristics. Methods Research was conducted on a 700-km? area near Northwest Bay (NWB), British Columbia, and on a 1000-km? area centred on the Adam and Eve River valleys (AE) approximately 300 km northwest of NWB (Figure 1). Principal habitats at NWB consisted of different seral stages of Douglas Fir (Pseudotsuga men- ziesii), Western Hemlock (Tsuga heterophylla), and Western Redcedar (Thuja plicata) forests. The AE study area was more rugged and dominated by exten- sive stands of unlogged forest, particularly at higher elevations. Habitats at lower elevations were dominated by Western Hemlock and Western Redcedar while spe- cies at higher elevations included Mountain Hemlock (Tsuga mertensiana) and Yellow Cedar (Chamaecy- paris nootkatensis). Understory vegetation on both study areas was typically dense and consisted of Salal (Gaultheria shallon), ferns (several genera) and Vac- cinium species communities. Both study areas had cool summers and mild winters, with temperatures of 0-10°C for 4-6 months/year and annual precipitation of 1700-5000 mm/year (Meidinger and Pojar 1991). AE was cooler and wetter, with a greater proportion of precipitation falling as snow. Cougars were captured between 4 March 1991 and 1 September 1996 at NWB and from 18 April 1997 to 12 September 1998 at AE. Local houndsmen were contracted to track and tree Cougars. Treed Cougars that could safely be immobilized were darted with a mixture of ketamine hydrochloride (3 mg/kg estimat- ed total weight) and medetomidine hydrochloride (0.1 mg/kg). Cougars were sexed, measured (see below), and aged according to tooth replacement and wear, and by gum recession (Ashman et al. 1983; Laundré et al. 2000). Adult Cougars were fitted with VHF radio col- lars (Telonics, Inc., Mesa, AZ). Immobilizations were reversed with atipamezole (0.15 mg/kg). Morphologi- cal data from hunter-killed Cougars in and near AE were also included in our analyses. 159 160 126° W 124° W 50° N Bay (NWB) FiGureE |. Location of cougar study areas on Vancouver Island, British Columbia, Canada. Morphological measurements taken included: total weight (using a 100 kg spring balance to the nearest 1 kg), neck circumference at the base of the skull, chest circumference immediately behind the front legs, body length from nose to base of tail, tail length from base to tip of the last vertebrae (all to the nearest cm with a cloth tape measure), canine length from gumline to tip, and front and rear pad widths (to the nearest mm with a cloth tape measure; AE only). We used principal component analysis (PCA) to derive a single variable to represent body size, based on the correlation matrix of morphological variables (Statistica 1995). We used this body size measure in- stead of total weight to describe the size of Cougars because it was less condition-dependent that using total weight alone. Where Cougars were captured >1 time, we included data from only the most recent capture. We used an analysis of covariance to test for differences in body size (log-transformed) between sex and study area, using age as a covariate. We also calculated means (+2 SE) for each morphological variable (by sex), but excluded Cougars <2 years old to minimize skews in distributions caused by the smaller body sizes of juvenile Cougars. We calculated survival estimates with the staggered entry design of Pollock et al. (1989), which is based on the Kaplan-Meier product limit estimator (Kaplan and Meier 1958). This method estimates annual sur- vival rate as the product of weekly survival rates (1 — d/r), where d is the number of animals that die and r THE CANADIAN FIELD-NATURALIST Vol. 118 is the number of animals “at risk” in a weekly period. Cougars carrying functioning radio collars comprised the “‘at risk” sample for each week. The design was “staggered entry” because not all Cougars carried col- lars at the same time. Cougars were added to the week- ly sample as they were collared, and were removed as they died, had their collars removed, or when their col- lars stopped functioning. Analysis started with the collaring of the first Cougars in the two study areas. Annual survival estimates were based on 52 consecu- tive, seven-day periods. There were sufficient data to analyse adult female Cougar survival; however, sample sizes were small, and the length of the study was relatively short in relation to the life expectancy of Cougars. Therefore, our estimates should be interpreted with caution. We report the reproductive characteristics of col- lared females anecdotally because data were insuffi- cient to calculate population growth rates; specifical- ly, we had few data on birth intervals and survival to maturity. We report litter sizes (42 SE) when they were first seen, so litter sizes at birth may have been larger, although in at least three litters, kittens were seen when <10 days old (eyes not open). Results Thirty Cougars were measured at NWB (17 females and 13 males) and 26 at AE (15 females and 11 males). We used total weight, neck and chest circumfer- ence, and total length (body length + tail length) in the final PCA to derive a body size variable. All four variables loaded strongly and positively on the first axis, which explained 83% of the variation in the dataset. As a result, we used the first PCA axis as the index of body size (Table 1). We did not include canine length and body and tail lengths as separate variables in the analysis because doing so did not significantly increase the variation described in the first axis. Body size differed between males and females (P < 0.000), but not between study areas (P = 0.736; Figure 2). Differences between males and females were also evident in univariate means of the morphologi- cal measurements (Table 2). Mean survival rates were similar for collared Cou- gars on both study areas (Table 3). Known causes of female mortality at NWB included intraspecific kill- ings (2) and animal control (3). The cause of one fe- male mortality at NWB and both female mortalities at AE were unknown. Two mortalities of collared male TABLE |. Correlation between morphological variables and the first principal component axis (also known as “factor load- ings”). The axis (based on the correlation matrix) was used as an index to describe body size of Vancouver Island Cougars in subsequent analyses. Factor loadings of morphological variables Neck Chest 0.889 0.927 Total Weight 0.994 Body length Eigenvalue 3.314 % variation explained 0.878 82.8 2004 WILSON, HAHN, GLADDERS, GOH, and SHACKLETON: VANCOUVER ISLAND COUGARS 16] Males: y = 1.45Ln(x) - 0.54 R? = 0.70 Females: e 6 y = 0.52Ln(x) - 1.09 R? = 0.34 UF T 0 2 4 T li 6 8 10 Age (years) FiGure 2. Relationship between body size (derived from the principal component analysis on morphological measurements, see text) and age for male and female Cougars in the two Vancouver Island study areas. Logarithmic trend lines are presented for males (squares @) and females (diamonds @) , pooled by study area. The body size measurement was used because it was less condition-dependent than using total weight alone. Cougars were investigated at AE; one was shot by hunt- ers and the other was suspected to have been shot illegally. At NWB, mean litter size at first detection was 1.9 + 0.1 (n= 16). One female was known to have a litter of three kittens, and there were 12 other litters of two kittens and 3 of one kitten when first observed. Fe- males were known or suspected to give birth during all months from March-October. Consecutive litters were recorded for two females; each had litters 21 and 23 months apart. Litter size at AE was 1.8 + 0.7 (n = 5). There was one litter of three, and two litters each of one and two kittens. Females gave birth in January, August, and September. No collared females at AE had more than one litter during the study period. Discussion Results presented in this paper are based on small absolute sample sizes; however, Cougars are rare and secretive, and inferences about their morphology and behaviour must often be based on few animals. The TABLE 2. Means and standard errors of morphological measurements taken on captured Vancouver Island Cougars. Males Measurement n Mean Total Weight (kg) 13 Ss Neck (cm) 12 39.5 Chest (cm) 12 VS) Body length (cm) 13 128.2 Tail Length (cm) 13 73.0 Front pad Width (mm) 9 57.9 Hind pad Width (mm) 9 50.4 Canine Length (mm) 10 28.6 Females 2 SE n Mean 2 SE Si 22 39.1 Pel 3.0 23 394 13} 3.5 23 65.4 1.6 6.6 23 120.9 Bill 4.5 23 ORI 3} 1.9 12 Syl I) 33 12 37.9 9.1 es) 23 26.8 2.0 162 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 3. Annual and mean survival rates of female Vancouver Island Cougars collared in two study areas: Northwest Bay (NWB) and Adam and Eve rivers (AE). Annual estimates were based on a staggered-entry design, starting with the first Cougars collared at NWB and AE in March 1991 and April 1997, respectively. Study Area Year n NWB 1991-1992 1992-1993 1993-1994 1994-1995 1995-1996 AE 1997-1998 1998-1999 1999-2000 ISSAP Sy Sap ays densities of Cougars in our study areas also suggest that significant proportions of the populations were captured (2.6-7.3 Cougars/100 km? at NWB and 1.4- 2.0 Cougars/100 km? at AE; S. Wilson, unpublished data). Our sample sizes were similar to, or larger than, those in many comparable studies (e.g., n = 22, Hemker et al. 1984; n = 68, Ross and Jalkotzy 1992; n= 76, Lindzey et al. 1994; n = 34, Spreadbury et al. 19962 n=13;, Prankhin;et ‘al. 1999: n= 21, Pierce;ct al. 2000). Few researchers have published Cougar morpholog- ical characteristics (c.f Kohlmann and Green 1999; Grigione et al. 2002); however, mean measurements of Vancouver Island Cougars in this study were smaller than those reported by Cowan and Guiguet (1965) for 14 adult male (mean total length 241 cm, tail length 89 cm, mean total weight 73 kg) and 7 adult female (mean total length 206 cm, tail length 79 cm, mean total weight 46 kg). Vancouver Island Cougars are slightly smaller than those found elsewhere in British Columbia (Cowan and Guiguet 1965). Survival estimates are critical for population man- agement, but few studies report them. Using the same method and similar sample sizes, Lindzey et al. (1988) reported mean survival rates (S = 0.731) for female Cougars in a largely unhunted population in southern Utah that were lower than those we calculated for Vancouver Island Cougars (mean for both study areas S = 0.885). Hunting is an important component of mortality in Cougar populations where hunting is al- lowed (Hemker et al. 1984; Logan et al. 1986; Ross and Jalkotzy 1992). The Vancouver Island population is no exception; however, we were in frequent contacts with hunters in our study areas and know they often treed radio-collared cougars but chose not to shoot them, even though we did not request this. This might have inflated the survival rates calculated for radio- collared Cougars. Researchers and managers should be aware of this bias when calculating survival esti- mates from similar radio telemetry studies where hunters “lend a hand” by not shooting study animals. Survival -2 SE +2 SE Mean 1 1 0.75 0.47 1 0.82 0.56 1 0.88 0.64 1 0.89 0.80 0.53 1 1 0.83 0.50 ff 0.88 Animal control by British Columbia Wildlife Branch Control Officers was a significant source of human- caused mortality at NWB, where the human popula- tion density was higher than at AE. Intraspecific killing has been reported as an important source of mortality elsewhere (Lindzey et al. 1988; Spreadbury et al. 1996), but in our study it was detected only at NWB where Cougar population density was significantly higher than at AE. Litters in both of our study areas were smaller than those reported in southeastern British Columbia (x = 3.1; Spreadbury et al. 1996), southwestern Al- berta (x = 2.2; Ross and Jalkotzy 1992), Wyoming (x = 2.7; Logan et al. 1986), and southern Utah (x = 2.4; Lindzey et al. 1994). Other studies have recorded births in most months of the year (Ross and Jalkotzy 1992; Lindzey et al. 1994), with a peak in late summer and early fall (Lindzey et al. 1994). Our observations at AE were similar. The pattern was dif- ferent at NWB, with no births recorded in winter and no obvious peaks in births during the spring-fall period. Columbia Black-tailed Deer (Odocoileus hemionus columbianus) are the Vancouver Island Cougars’ pri- mary prey. Indices of deer abundance declined 55% from 1991-1996 at NWB and 38% from 1995-1999 at AE (British Columbia Ministry of Environment, Lands and Parks, unpublished data); therefore, small litter sizes among Vancouver Island Cougars may be a result of low food availability. Also, Black-tailed Deer are smaller than mainland Mule Deer (Odocoileus hemi- onus; Shackleton 1999). This highlights the importance of maintaining prey populations in management of Cougars on Vancouver Island. Acknowledgments This research was supported by Forest Renewal British Columbia, British Columbia Ministry of Envi- ronment, Lands, and Parks (MELP), Habitat Conserva- tion Trust Fund, Science Council of British Columbia, MacMillan Bloedel Limited, TimberWest Limited, Western Forest Products Limited, BC Hydro, Mountain 2004 Equipment Co-op, and the University of British Col- umbia. We thank M. DeLaronde, S. Foster, D. Koshow- ski, J. Leo, R. Ramcharita, H. Robinson, C. Wainwright, K. Wickert, and local houndsmen for assistance with fieldwork. For logistic support we also thank G. Brun- ham, K. Brunt, T. Hamilton, D. Janz, and H. Schwantje of MELP, as well as many staff of MacMillan Bloedel Limited in Nanaimo and Campbell River, British Columbia, and of TimberWest Limited in Nanaimo, B.C. S. Kohlmann, F. Scott and D. Nagorsen made helpful comments on the manuscript. This paper is dedicated to the memory of the late K. Atkinson, who conceived and supported the project from the outset. Literature Cited Ashman, D., M. Christensen, M. Hess, G. Tsukamoto, and M. Wickersham. 1983. The mountain lion in Nevada. Nevada Department of Fish and Game, Federal Aid in Wildlife Restoration Final Report W-48-15. Beier, P. 1991. Cougar attacks on humans in the United States and Canada. Wildlife Society Bulletin 19: 403-412. Cowan, I. McT., and C. J. Guiguet. 1965. The mammals of British Columbia. British Columbia Provincial Museum Handbook Number 11, Victoria, British Columbia. 414 pages. Franklin, W. L., W. E. Johnson, R. J. Sarno, and J. A. Ariarte. 1999. Ecology of the Patagonia puma Felis con- color patagonica in southern Chile. Biological Conservation 90: 33-40. Grigione, M. M., P. Beier, R. A. Hopkins, D. Neal, W. D. Padley, C. M. Schonewald, and M. L. Johnson. 2002. Ecological and allometric determinants of home-range size for mountain lions (Puma concolor). Animal Conservation 5: 317-324. Hemker, T. P., F. G. Lindzey, and B. B. Ackerman. 1984. Population characteristics and movement patterns of cou- gars in southern Utah. Journal of Wildlife Management 48: 1275-1284. Kaplan, E. L., and P. Meier. 1958. Nonparametric estimation from incomplete observations. Journal of the American Statistics Association 53: 457-481. WILSON, HAHN, GLADDERS, GOH, and SHACKLETON: VANCOUVER ISLAND COUGARS 163 Kohlmann, S. G., and R. L. Green. 1999. Body size dynam- ics of cougars (Felis concolor) in Oregon. Great Basin Naturalist 59: 193-194. Laundré, J. W., L. Hernandez, D. Streubel, A. Altendorf, and C. L. Gonzalez. 2000. Aging mountain lions using gum-line recession. Wildlife Society Bulletin 28: 963-966. Lindzey, F. G., Ackerman, B. B., Barnhurst, D., and Hemker, T. P. 1988. Survival rates of mountain lions in southern Utah. Journal of Wildlife Management 52: 664- 667. Lindzey, F. G., W. D. Van Sickle, B. B. Ackerman, D. Barn- hurst, T. P. Hemker, and S. P. Laing. 1994. Cougar popu- lation dynamics in southern Utah. Journal of Wildlife Management 58: 619-624. Logan, K. A., L. L. Irwin, and R. Skinner. 1986. Character- istics of a hunted mountain lion population in Wyoming. Journal of Wildlife Management 50: 648-654. Meidinger, D. V., and J. Pojar. 1991. Ecosystems of British Columbia. Special Report Series 6, British Coiumbia Ministry of Forests, Victoria, British Columbia. 330 pages. Pierce, B. M., V. C. Bleich, and R. T. Bowyer. 2000. Social organization of mountain lions: does a land-tenure system regulate population size? Ecology 81: 1533-1543. Pollock, K. H., S. R. Winterstein, C. M. Bunck, and P. D. Curtis. 1989. Survival analysis in telemetry studies: the staggered entry design. Journal of Wildlife Management S31 Ross, P. I., and M. G. Jalkotzy. 1992. Characteristics of a hunted population of cougars in southwestern Alberta. Journal of Wildlife Management 56: 417-426. Shackleton, D. M. 1999. The Hoofed Mammals of British Columbia. Volume 3: The Mammals of British Columbia. Royal British Columbia Museum, Victoria, British Colum- bia. 268 pages. Spreadbury, B. R., K. Musil, J. Musil, C. Kaisner, and J. Kovak. 1996. Cougar population characteristics in south- eastern British Columbia. Journal of Wildlife Manage- ment 60: 962-969. Statistica. 1995. Statistica users manual volume III. Statsoft Incorporated, Tulsa, Oklahoma. 782 pages. Received 7 May 2001 Accepted 5 November 2004 Conservation Evaluation of Lemmon’s Holly Fern, Polystichum lemmonii, a Threatened Fern in Canada” GEORGE W. DoucLas!: 2 ‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information Branch, P.O. Box 9993 Stn Prov Govt, Victoria, British Columbia V8W 9R7 Canada > Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada Douglas, George W. 2004. Conservation evaluation of Lemmon’s Holly Fern, Polystichum lemmonii; a threatened fern in Canada. Canadian Field-Naturalist 118(2): 164-168. In Canada, Lemmon’s Holly Fern, Polystichum lemmonii, is restricted to the Baldy Mountain area on the eastern side of the Okanagan River valley in south-central British Columbia. This population represents the northern limits of the species which ranges south through northern Idaho, Washington and Oregon to northern California. In British Columbia, P. lemmonii is associated with ultramafic rocky ridges within a montane forest at an elevation of 1900 m. The population in the Baldy Mountain area is relatively small, unprotected and potentially imperilled by mining exploration, forest road construction or wildfires. Key Words: Lemmon’s Holly Fern, Polystichum lemmonii, threatened, distribution, population size, British Columbia. Lemmon’s Holly Fern, Polystichum lemmonii Underw. [taxonomy and nomenclature follow Douglas et al. (1998a, b; 2000)], is a member of a cosmopolitan genus of over 175 species (Smith and Lemieux 1993; Wagner 1993). It is one of eight Polystichum species occurring in British Columbia (Ceska 2000) and nine occurring in Canada (Cody and Britton 1989; Wagner 1993). Generally, American authors treated P. lem- monii as a synonym of P. mohrioides (Bory) C. Presl. until Wagner (1979) demonstrated that the North Am- erican plant was different from the South American plant. Polystichum lemmonii was first recorded in Canada by Cody and Britton (1984). Polystichum lemmonii 1s an evergreen, perennial, tufted fern arising from a short, stout rhizome (Figure 1; Ceska 2000). The decumbent to ascending fronds are 10-40 cm long, 3-7 cm wide and 2-pinnate. The 20-35 pinnae on each side of the rachis are ovate with rounded pinnules. The ultimate segments are entire or weakly toothed. The round sori are attached near the midvein with entire or minutely toothed indusia. In British Columbia, P. lemmonii may be confused with either Kruckeberg’s Holly Fern (P. kruckebergii) or Mountain Holly Fern (P. scopulinum). It may be * The field work for the Polystichum lemmonii project was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002). This information formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Douglas 2003*) and the subsequent assess- ment of threatened (COSEWIC 2003*). The present paper also includes more recent information that will be used in a National Recovery Strategy for P. lemmonii (Doug- las 2005). FiGurE 1. Mlustration of Polystichum lemmonii (line drawing from Ceska 2000). 164 2004 ———~ KAMLOOPS SPENCES BRIDGE ARMSTRONG Sw PRINCETON PENTICTON KEREMEOS | ) BRITISH ‘& . COLUMBIA FIGURE 2. The small square indicates the location of Polystichum lemmonii in British Columbia. DOUGLAS: LEMMON’S HOLLY FERN IN CANADA 165 distinguished from the latter two species by the lack of spines on the teeth of the ultimate segments of the pinnae (Hitchcock et al. 1969; Wagner 1993; Ceska 2000). North American and Provincial Ranges Polystichum lemmonii ranges from south-central British Columbia, sporadically south through Wash- ington and Oregon to northern California (Smith and Lemieux 1993; Wagner 1993). In Canada, it is known only from the Baldy Mountain area in the Okanagan River valley in south-central British Columbia (Figure 2; Ceska 2000; Douglas et al. 2002). Habitat In western North America, Polystichum lemmonii occurs on sites where ferromagnesian or ultramafic rocks outcrop (Kruckeberg 1969; Wagner 1993). There are a number of ultramafic rock outcrops in western British Columbia but only the two small, adjacent ridges, at an elevation of 1900 m, in the Baldy Moun- tain area support the latter species (Figure 3). These dunite rock outcrop habitats are also characterized by shallow soils thus creating xeric microclimates that exclude many nearby species adapted to more mesic microclimates or non-ultramafic soils (Figure 4). These ridges, therefore, have a typically depauperate ultra- mafic flora and lack a tree cover in contrast to the FicureE 3. Aerial view of the ultramafic east ridge. Most Polystichum lemmonii plants occur on the north side of these east-west oriented ridges. The west ridge is slightly longer. 166 surrounding montane forests. The most prominent species on the ridge include Common Juniper (Juni- perus communis), Indian’s Dream (Aspidotis densa), Alpine Sandwort (Minuartia obtusiloba) and Yarrow (Achillea millefolium). Biology There is a limited amount of information on the biology and ecology of Polystichum lemmonii. The ultramafic habitat, however, has been well-studied by Kruckeberg (1969). Plants of ultramafic substrates are adapted to tolerate low levels of calcium, nitro- gen, phosphorus and molybdenum and high levels of magnesium, chromium and nickel (Kruckeberg 1969). Wagner (1979) has provided some genetic informa- tion. P. lemmonii is a tetraploid (2n = 82) and thought to be one of the parents of both P. kruckebergii and P. scopulinum. Sporophytes of P. lemmonii, as with most evergreen fern species, often retain significant numbers of mature spores over the winter that are then released the fol- lowing spring (Farrar 1976). PR. lemmonii also grows vegetatively by subterranean rhizome elongation often resulting in large clumps of clones. Because of the dry site conditions, which are not ideal for spore germination or gamete fertilization, most reproduction is probably by rhizome elongation (Walker 1979). Prothalli were not observed at the site. Long-distance dispersal of spores of P. lemmonii is evident by the distance to the nearest locations of the species in the adjacent state of Washington in the United States. P. lemmonii occurs in the Twin Sisters Range of Washington and in the Wenatchee Mountains (Kruckeberg 1969), a distance of about 205 km to the southwest and 225 km to the south of Baldy Moun- tain, respectively. The Tulameen River ultramafic site, where both P. kruckebergii and P. scopulinum occur (Kruckeberg 1969; Douglas and Labrecque 2003%*), does not contain P. lemmonii even though the Tula- meen River site is halfway between Baldy Mountain and the Twin Sisters Range. Reproduction of P. lem- moni at the Baldy Mountain area is evident since about 30 percent of the 853 plants counted were relatively young (plant tufts less than four cm wide). Population Attributes The population of P. lemmonii occurs on two adja- cent, rocky ridges. The ridges, which are about 280 and 200 m long by 50 m wide, are separated by a distance of 160 m. A 2001 count of all plants in the popula- tion by the author revealed a total of 853 plants over 0.72 ha. A collection by D. M. Britton in 1987 (at Department of Agriculture, Ottawa) mentions a popu- lation size of “perhaps a thousand plants”. This would indicate that the population has remained relatively stable for at least 15 years. THE CANADIAN FIELD-NATURALIST Vol. 118 Provincial, National and Global Ranks The British Columbia Conservation Data Centre has ranked this species as S1 and placed it on the British Columbia Ministry of Sustainable Resource Management Red-list (Douglas et al. 2002). This is the most critical category for imperilled rare native vascular plants in British Columbia. A rank of S1 is considered “critically imperilled because of extreme rarity (5 or fewer occurrences or very few remaining individuals) or because of some factors making it especially vulnerable to extirpation or extinction” (Douglas et al. 2002). Since the species is restricted to British Columbia, the National rank is N1. Globally, Polystichum lemmonii is ranked G4 and is frequent to common in its range and apparently secure. Threats and Protection The most direct threat to Polystichum lemmonii is mining exploration. At the present time the entire area is occupied by active mining claims and explora- tion could occur with short notice. Additional threats include the potential use of the rock outcrop for forest road construction and the possibility of intense wild- fires. Extremely high forest fuel loads in the the area may lead to wildfires similar to that experienced in the region in 2003. Introduced species are of no con- cern at this site due to the ultramafic properties of the soils. The population in the Baldy Mountain area is on public land but is not part of a protected area. It is conceivable that this area could qualify as a Wildlife Habitat Area but this status has yet to be proposed. Polystichum lemmonii could be a candidate species for protection under the provincial Wildlife Amend- ment Act as it is currently Red-listed by the British Columbia Conservation Data Centre. Evaluation The British Columbia Conservation Data Centre considers Polystichum lemmonii to be threatened/en- dangered in British Columbia (Douglas et al. 2002) and the Committee on the Status of Endangered Wild- life in Canada has assessed the species as threatened (COSEWIC 2003). Just over 850 plants are known from a single site in Canada at the Baldy Mountain area of south-central British Columbia. The prognosis for this species is not good since ultramafic rock out- crops often attract mineral exploration and the entire area is covered by active mineral claims. These rock outcrops could also be of potential use as a quarry for road building materials. The extremely high forest fuel loads in the adjacent area could also support wildfires. Establishment of a Wildlife Habitat Area and removal of the active mineral claims would remove the major threats at the site. 2004 DOUGLAS: LEMMON’S HOLLY FERN IN CANADA 167 FicureE 4. Polystichum lemmonii plants are conspicuous among the low vegetation on the ultramafic soils. Acknowledgments I would like to thank Kathy Paige for her partici- pation in the fieldwork and Louise Bacon and Jenifer Penny for office assistance. Graham Nixon, British Columbia Geological Survey, kindly provided infor- mation on ultramafic rock types. Documents Cited (marked * in text) COSEWIC. 2003. COSEWIC assessment and status report on Lemmon’s holly fern Polystichum lemmonii in Canada. Committee on the Status of Endangered Wildlife in Can- ada. www.cosewic.gc.ca, Ottawa, Ontario. 13 pages Douglas, G. W. 2003. COSEWIC Status report on Lemmon’s holly fern Polystichum lemmonii in Canada, in COSEWIC assessment and status report on Lemmon’s holly fern Polystichum lemmonii in Canada. Committee on the Status of Endangered Wildlife in Canada. www.cosewic.gc.ca, Ottawa, Ontario. 13 pages. Douglas, G. W., and J. Labrecque. 2003. COSEWIC Status report on Mountain Holly Fern or Polystic des Rochers, Polystichum scopulinum in Canada. Unpublished report. Ottawa, Ontario. 20 pages. Douglas, G. W. 2005. National Recovery Strategy for Lem- mon’s Holly Fern (Polystichum lemmonii). In preparation. Ottawa, Ontario. Literature Cited Ceska, A. 2000. Pteridophytes. Pages 260-343 in Illustrated flora of British Columbia. Volume 5. Dicotyledons (Salica- ceae through Zygophyllaceae) and Pteridophytes. Edited by G. W. Douglas, D. Meidinger and J. Pojar. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests. Victoria, British Columbia. 427 pages. Cody, W. J., and D. M. Britton. 1984. Polystichum lemmonii, a rock shield-fern new to British Columbia and Canada. Canadian Field-Naturalist 98: 375. Cody, W. J., and D. M. Britton. 1989. Ferns and fern allies of Canada. Agriculture Canada Research Branch, Publica- tion 1829/E. Ottawa, Ontario. 430 pages. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare native vascular plants of British Columbia. Second edition. Province of British Columbia. Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus- trated flora of British Columbia. Volume 5. Dicotyledons (Salicaceae through Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests. Victoria, British Columbia. 389 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. Illustrated flora of British Columbia. Volume 1. Gymno- sperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests. Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 2. Dicotyledons (Balsaminaceae through Cuscutaceae). Bri- 168 tish Columbia Ministry of Environment, Lands and Parks and Ministry of Forests. Victoria, British Columbia. 401 pages. Farrar, D. L. 1976. Spore retention and release from over- wintering fronds. American Fern Journal 66: 49-52. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1969. Vascular plants of the Pacific Northwest. Part 1. Vascular Cryptogams, Gymnosperms and Mono- cotyledons. University Washington Press, Seattle, Wash- ington. 914 pages. Kruckeberg, A. R. 1969. Plant life on serpentine and other ferromagnesian rocks in northwestern North America. Syesis 2: 14-114. Smith, A. R., and T. Lemieux. 1993. Dryopteridaceae. Pages 91-94 in The Jepson manual: Higher plants of California. THE CANADIAN FIELD-NATURALIST Vol. 118 Edited by J. C. Hickman. University of California Press. Berkeley, California. 1400 pages. Wagner, D. H. 1979. Systematics of Polystichum in western North America north of Mexico. Pteridologia 1: 1-64. Wagner, D. H. 1993. Polystichum. Pages 290-299 in Flora of North America North of Mexico — Volume 3. Magno- liophyta: Magnoliidae and Hamamelidae. Edited by Flora of North America Editorial Committee. Oxford University Press Inc. New York, New York. 475 pages. Walker, T. G. 1979. The cytogenetics of ferns. In The experi- mental Biology of ferns. Edited by A. F. Dyer. Academic Press, London, England. Received 15 October 2002 Accepted 1 September 2004 Conservation Evaluation of the Pacific Population of Tall Woolly-heads, Psilocarphus elatior, an Endangered Herb in Canada* GEORGE W. DouGLas! and JEANNE M. ILLINGWORTH? ‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information Branch, P.O. Box 9993 Stn Prov Govt, Victoria, British Columbia V8W 9R7 Canada 23537 Savannah Ave., Victoria, British Columbia V8X 1S6 Canada Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada Douglas, George W., and Jeanne M. Illingworth. 2004. Conservation evaluation of the Pacific population of Tall Woolly- heads, Psilocarphus elatior, an endangered herb in Canada. Canadian Field-Naturalist 118(2): 169-173. In Canada, Psilocarphus elatior occurs in British Columbia, Alberta and Saskatchewan. This paper examines the status of the Pacific populations located on southeastern Vancouver Island in southwestern British Columbia. The Pacific population consists of 12 recorded sites of which only five have been confirmed since 1993. In British Columbia, P. elatior is associated with dried beds of vernal pools and other open, moist depressions at lower elevations. In British Columbia, P. elatior populations occur in large numbers at only two of the seven locations. Key Words: Tall Woolly-heads, Psilocarphus elatior, endangered, distribution, population size, British Columbia. Tall Woolly-heads, Psilocarphus elatior (A. Gray) A. Gray” [Taxonomy and nomenclature follows Doug- las (1998) and Douglas et al. (1998; 1999a, c; 2001a b), is a member of a genus of five species which occur in the Americas (Cronquist 1950). It is one of three species occurring in British Columbia and Canada (Douglas 1998). Psilocarphus elatior is a small, erect, annual plant up to 15 cm tall (Figure 1; Douglas 1995, 1998). The opposite leaves are silky-hairy, linear-oblong, entire on the margins and 1.2 to 3.5 mm long. The flowers form solitary spherical heads in the leaf axils or at the tips of the stems or leaf branches. The flowers lack involucres but have involucral-like leaves at their bases. Each of the outer (female) threadlike ray flowers has a well developed, 2.4 to 3.8 mm long receptacle bract. These bracts have a translucent appendage below the summit. In British Columbia, P. elatior may be distinguished from Slender Woolly-heads (P. tenellus var. tenellus) by its erect habit, larger heads and receptacular bracts. In the field, young specimens of P. elatior may also be confused with young specimens of Lowland Cudweed (Gnaphalium palustre). Close examination will reveal * Field work for the Psilocarphus elatior project was fund- ed by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002). These data formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Douglas, et al. 1999b) and the subsequent assessment of endangered (COSEWIC 2004*) for the Pacific population. The present paper also includes more recent information from research funded by the British Columbia Conserva- tion Data Centre. that most of the leaves of the latter are alternate and the flower heads have densely woolly involucres, broader ray flowers and a non-spherical shape. North American and Provincial Ranges Psilocarphus elatior ranges from southwestern Brit- ish Columbia, southeastern Alberta and southwestern Saskatchewan in Canada, south through Idaho, Wash- ington and Oregon to northern California (Cronquist 1955; Scoggan 1979; Morefield 1993; Douglas 1998). In British Columbia, it is restricted to south-eastern Vancouver Island in south-western British Columbia (Figure 2; Douglas 1998; Douglas et al. 2002). Habitat Psilocarphus elatior inhabits dried beds of vernal pools, ephemeral lake edges (Figure 3) and other open moist, often disturbed sites at lower elevations. The sites are often level and generally slightly depressed. Psilocarphus elatior is not found within particular communities and often occupies sites where other spe- cies are sparse. There does not appear to be a consis- tent association with other plants. The Somenos Lake locality near Duncan supports the largest and most vigourous population of P. elatior (Table 1). The plants grow on the moist shoreline of the lake and appear when the lake level recedes in late spring or early summer. Major associates at this site include Slender-beaked Sedge (Carex athrostachya), One-sided Sedge (C. unilateralis), Brass Buttons (Cotula coronopifolia) and Skunkweed (Navarretia squarrosa). At Uplands Park, in the Municipality of Uplands near Victoria, five subpopulations occur in a large ephemeral meadow. Associated species include Bent- grass (Agrostis sp.), California Oatgrass (Danthonia 169 170 THE CANADIAN FIELD-NATURALIST Figure 1. Illustration of Psilocarphus elatior (line drawing by Elizabeth J. Stephen in Douglas [1995, 1998]). californica), Toad Rush (Juncus bufonis), Perennial Ryegrass (Lolium perenne) and Small Hop-clover (Trifolium dubium). Introduced species are more abun- dant here than at the Somenos Lake site and appear to be increasing yearly. The Christmas Hill site, although smaller than the previous two sites, is in relatively good condition. Native species in this habitat include Carolina Meadow-foxtail (Alopecurus carolinianus), Green- sheathed Sedge (Carex feta), Heterocodon (Hetero- codon rariflorum), and Scouler’s Popcornflower (Plagiobothrys scouleri). The two remaining populations, at Scafe Hill and Cattle Point, occur in small depressions. These popu- lations are probably the least stable of the extant pop- ulations. The Cattle Point populations are absent dur- ing some years but the seed bank appears to remain. Psilocarphus elatior is considered to be a vernal pool specialist (Keeley and Zedler 1998). They define vernal pools as “precipitation-filled seasonal wetlands inundated during periods when temperature is suffi- cient for plant growth, followed by a brief water- logged-terrestrial stage and culminating in extreme desiccating soil conditions of extended duration”. The Vol. 118 f ss L PORT VANCOUVER “iw ‘fp ISLAND O UCLUELET SIDNEY FIGURE 2. The location and status of Psilocarphus elatior sites in British Columbia (0 — extirpated sites, e — re- cently confirmed sites, Hi — present status unknown). species is able to outcompete grassland species due to its tolerance of inundation and aquatic/wetland species due to its tolerance of soil desiccation and heat during summer drought. Biology Other than general habitat information, there is little in the literature regarding the biology and ecology of Psilocarphus elatior. It is likely, however, that this plant shares many of the same traits that are typical of the genus. 2004 DOUGLAS AND ILLINGWORTH: PACIFIC POPULATION OF TALL WOOLLY-HEADS 17] Ficure 3. A dense population of Psilocarphus elatior on the edge of Somenos Lake. The prominent sedge is Carex athrostachya. This site is usually submerged until late spring. Lack of structures attractive to insects and animals, and an interpretation of the floral structure indicate these inconspicuous, woolly annuals may self-polli- nate. Cronquist (1950) suggested that in the genus Psilocarphus, the position of the receptacular bracts, together with the position of the corolla and stigmas, effectively guide these latter structures towards the central flowers. Since it is the central flowers alone that produce pollen, and since the wool and leaves sur- rounding the head appeared to prevent pollen loss by wind, self-pollination is indicated. It is also possible that pollen may not be essential for seed-production and that asexual reproduction may be occurring, al- though chromosome counts would be necessary to verify this. Seed dispersal also appears limited. Since the ach- enes are much smaller than the enclosing bracts, the only obvious means of dispersal is by water or wind. Table 1. Locations and Population Sizes for Psilocarphus elatior in British Columbia. Collection Site Last Observation Cloverdale (Victoria) 1887 Cedar Hill (Victoria) 1887 Ucluelet 1909 Roberts Bay (Sidney) 1913 Swartz Bay (Sidney) 1931 Francis-King Regional Park (Victoria) 1962 University of Victoria (Victoria) 1966 Cattle Point, Uplands Park (Victoria) 1993 Scafe Hill (Victoria) 1996 Uplands Park (Victoria) 1998 Christmas Hill (Victoria) 2001 Somenos Lake (Duncan) 2002 Collector/ Population Observer (no./area) Macoun Extirpated Macoun Extirpated Macoun Extirpated Macoun Extirpated Groh Extirpated Melburn Unknown Turner Extirpated Ryan 200/10 m? Roemer 20/? m? Douglas 40 000+/1 200+ m? Douglas & Penny 450/72 m? Douglas & Douglas 100 000+/45 m? 172 The plant’s habitat in vernal pools may also permit the bracts and achenes to be transferred by the muddy feet of waterfowl and other animals (Cronquist 1950). Further studies are required to determine many as- pects of the population dynamics of P. elatior includ- ing the average life-cycle of the species, the frequency and requirements for seed germination and survival, and its competitive ability with other species. Population Attributes Psilocarphus elatior has been recorded from 12 sites in southwestern British Columbia (Table 1). Five of these have been confirmed since 1993. The status of the remaining populations is unknown and many are believed extirpated. Population areas range from 10 m? to over 1200 m? while numbers of plants range from 20 to over 100 000. In the majority of cases, popula- tion trend analysis is not available because of limited demographic data. Provincial, National and Global Ranks Provincially, P. elatior is ranked S2 by the British Columbia Conservation Data Centre (Douglas et al. 2002) which indicates this species to be “imperiled because of rarity (typically 6-20 extant occurrences or few remaining individuals) or because of some fac- tor(s) making it vulnerable to extirpation or extinction.” Nationally the species is ranked N3, while globally it has a rank of G5. The latter rank indicates it is “frequent to common to very common; demonstrably secure and essentially ineradicable under present con- ditions”. Threats and Protection Habitat destruction is the greatest threat to the exist- ing populations of Psilocarphus elatior. Some sites occur in areas subjected to heavy pedestrian tram- pling or are at risk from private development. Threats to the continued survival of this species are compound- ed by the lack of biological and ecological information which create difficulties in site management. Four of the P. elatior populations are partially pro- tected by their location in municipal or regional parks or special protected areas (Somenos Lake). The latter probably has the greatest degree of protection since it is administered by British Columbia Parks and falls under the Park Act. Most of the municipal or regional parks receive little active management at the present time, at least with respect to their rare plants. Park enhancement projects, road and trail developments and heavy recreational use by humans often result in the destruction of the native vegetation and rare plant spe- cies. Psilocarphus elatior could be a candidate species for protection under the provincial Wildlife Amendment Act as it is currently Red-listed by the British Colum- bia Conservation Data Centre (Douglas et al. 2002). Most of the populations contain small numbers of plants. Once a population becomes small, it becomes THE CANADIAN FIELD-NATURALIST Vol. 118 more susceptible to demographic and environmental variation and loss of genetic variability. In some cases, small populations are at risk of inbreeding depres- sion, genetic drift and loss of fitness (Primack 1998). Evaluation The British Columbia Conservation Data Centre considers Psilocarphus elatior to be threatened/endan- gered in British Columbia (Douglas et al. 2002a) and the Committee on the Status of Endangered Wildlife in Canada has assessed the species as endangered (COSEWIC 2003). Most of the populations are small and not viable. With limited knowledge of the plants biological and ecological requirements, this species is vulnerable to extirpation in British Columbia. Without research on growth requirements and further demo- graphic information, the stability of the present popu- lations will remain unknown. Studies are also neces- sary to determine if the present habitats are necessary for the successful growth of P. elatior or if these plants are simply outcompeted elsewhere. The limited num- ber of individuals also reduces the potential for gen- etic variation which may be necessary to respond to environmental changes in the future. Acknowledgments We thank Sylvia Douglas, Jenifer Penny, and Michael Ryan for their field assistance. Documents Cited (marked * in text) COSEWIC. 2004. Canadian Species at Risk. Committee on the Status of Endangered Wildlife in Canada. www.cose wic.gc.ca, Ottawa, Ontario. 13 pages. Douglas, G. W., J. Gould, and J. M. Illingworth. 1999. Status report on the Tall woolly-heads, Psilocarphus elatior (A. Gray) A. Gray. Unpublished report. Committee on the Status of Endangered Wildlife in Canada, Ottawa. 19 pages. Literature Cited Cronquist, A. 1950. A review of the genus Psilocarphus. Research Studies of the State College of Washington 18: 71-89. Cronquist, A. 1955. Vascular plants of the Pacific Northwest. Part 5: Compositae. University of Washington Press, Seat- tle, Washington. 343 pages. Douglas, G. W. 1995. The sunflower family (Asteraceae) of British Columbia. Volume 2 — Astereae, Anthemideae, Eupatoreae and Inuleae. Royal British Columbia Museum, Victoria, British Columbia. 393 pages. Douglas, G. W. 1998. Asteraceae. Pages 96-392 in Illustrat- ed flora of British Columbia. Volume 1. Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). Edited by G. W. Douglas, G. B. Straley, D. Meidinger, and J. Pojar. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002. Rare native vascular plants of British Columbia. Second edition. Province of British Columbia, Victoria, British Columbia. 358 pages. 2004 Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- trated flora of British Columbia. Volume 3. Dicotyledons (Diapensiaceae through Onagraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- trated flora of British Columbia. Volume 4. Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Col- umbia Ministry of Forests, Victoria, British Columbia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar 2001a. Illus- trated flora of British Columbia. Volume 6. Monocotyle- dons (Acoraceae through Najadaceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Colum- bia. 361 pages. Douglas, G. W., D. Meidinger, and J. Pojar 2001b. Ilus- trated flora of British Columbia. Volume 7. Monocotyle- dons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998. Illustrated flora of British Columbia. Volume 1. DOUGLAS AND ILLINGWORTH: PACIFIC POPULATION OF TALL WOOLLY-HEADS 173 Gymnosperms and Dicotyledons (Aceraceae through Asteraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Keeley, J. E., and P. H. Zedler. 1998. Characterization and global distribution of vernal pools. /n Ecology, conserva- tion, and management of vernal pool ecosystems: Proceed- ings from a 1996 Conference. Edited by C. W. Witham, E. T. Bauder, D. Belk, W. R. Ferren Jr., and R. Ornduff. California Native Plant Society. Sacramento, California. Morefield, J. 1993. Psilocarphus. Page 329 in The Jepson manual: higher plants of California. Edited by J. C. Hick- man. University of California Press, Berkeley, California. 1400 pages. Primack, R. B. 1998. Essentials of Conservation Biology. 24 Edition. Sinauer Associates Inc. Sunderland, Massa- chusetts. Scoggan, H. J. 1979. The flora of Canada. Part 4 — Dico- tyledoneae (Loasaceae to Compositae. National Museum of Natural Sciences, Ottawa, Ontario. Publication in Botany Number 7. Pages 1117-1711. Received 15 October 2002 Accepted | September 2004 Conservation Evaluation of Howell’s Triteleia, Triteleia howellii, an Endangered Lily in Canada* GEORGE W. DouGLas!:? and JENIFER L. PENNY! ‘Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information Branch, P.O. Box 9993 Stn Prov Govt, Victoria British Columbia V8W 9R7 Canada ? Current address: Douglas Ecological Consultants Ltd., 6230 North Road, Duncan, British Columbia V9L 6K9 Canada Douglas, George W., and Jenifer L. Penny. 2004. Conservation evaluation of Howell’s Triteleia, Triteleia howellii, an endangered lily in Canada. Canadian Field-Naturalist 118(2): 174-178. In Canada, Triteleia howellii is restricted to Quercus garryana stands and grass-dominated meadows on southeastern Van- couver Island in southwestern British Columbia. Nine sites have been confirmed in recent years while three other sites are considered extirpated. These Canadian sites represent the northern range limits of 7: howellii. Threats to existing populations vary in intensity. Although most populations are protected to a certain extent from direct habitat destruction, introduced species pose a serious potential threat to the continued existence of most populations. Managing sites for 7: howellii is difficult because little information is available regarding the general biology of this species. Key Words: Howell’s Triteleia, Triteleia howellii, endangered, distribution, population size, British Columbia. Howell’s Triteleia, Triteleia howellii (S. Wats.) Greene [taxonomy and nomenclature follow Douglas et al. (1998a, b; 1999a, b; 2000, 2001a, b)], some- times treated as T. grandiflora Lindl. var. howellii (S. Wats.) Hoover, is a member of a genus of 14 species in North America (Keator 1993). Three species occur in British Columbia and Canada (Scoggan 1979; Pojar 2001). Triteleia howellii is a perennial herb from a deep, straw-coloured, fibrous-scaly, nearly globe-shaped, bulb-like corm (Figure 1; Pojar 2001). The erect, flow- ering stem is 20-50 cm tall with one or two smooth, slender, linear basal leaves. The leaves are 20-40 cm long, 3-8 mm wide, sheathed at the base and have en- tire margins. The flowers consist of six whitish to blue, vase-shaped to narrowly bell-shaped, fused segments forming a 1.5-2 cm long tube. The corolla lobes, which are about as long as the tube, are in two, spreading, petal-like whorls, about as long as the tube. The outer three are broadly lanceolate, the inner three are oblong- egg-shaped and all are slightly ruffled. The fruit con- sists of a stalked, egg-shaped capsule containing black rounded seeds. Triteleia howellii is similar in appearance to its close relative 7: grandiflora. It is distinguished from the latter * The field work for the Triteleia howellii project was funded by the British Columbia Conservation Data Centre. The results appear in the British Columbia Conservation Data Centre database and a rare plant manual (Douglas et al. 2002). This information formed the basis for a Committee on the Status of Endangered Wildlife in Canada status report (Douglas 2003*) and the subsequent assessment of Endangered (COSEWIC 2003*). The present paper also includes more recent information, funded by the Nature Conservancy of Canada and the Habitat Conservation Trust Fund, that will be used in a National Recovery Strategy for T: howellii (Douglas and Smith 2005). by its flat filaments which are attached at the same level on the perianth tube (Pojar 2001). The filaments of 7: grandiflora, in contrast, are not flat and are attached at two levels on the perianth tube. North American and Provincial Ranges Triteleia howellii ranges from southwestern British Columbia, south through Washington and Oregon to northern California (Barkworth 1977a; Keator 1993). In Canada, 7: howellii is known only from southeast- ern Vancouver Island (Figure 2; Pojar 2001; Douglas et al. 2002). Habitat In British Columbia T- howellii occurs on rock out- crops, in Garry Oak (Quercus garryana) woodlands, Garry Oak/Arbutus (Arbutus menziesii) stands and occasionally in highly disturbed sites dominated by weeds in private yards and on roadsides. In the highly disturbed sites, dominants include Orchard Grass (Dac- tylis glomerata), Cheat Grass (Bromus tectorum), Com- mon Vetch (Vicia sativa), Rip-gut Brome (Bromus rigidus), Soft Brome (B. hordeaceus), Perennial Rye- grass (Lolium perenne), and Pacific Sanicle (Sanicula crassicaulis var. crassicaulis), all introduced except for Sanicula. At the highest quality site, in the Quercus garryana woodland at the Cowichan Garry Oak Preserve, the habitat is classified as a Quercus garryana/Dactylis glomerata plant community (Douglas et al. 2002*) and is characterized by deep, dark soils up to a metre in depth. It is likely that prior to understory dominance by D. glomerata in this (Figure 3), and other Quercus stands of the region, this plant community would have fallen within the Q. garryana/California Brome (Bro- mus carinatus) community type (Roemer 1972). An 174 2004 FiGure 1. Illustration of Triteleia howellii (Line drawing in Pojar 2001). extremely rich low shrub and herb stratum is present during the spring. The most prominent species in the Cowichan Garry Oak Preserve Garry Oak stand are Sanicula crassicaulis var. crassicaulis and Dactylis glomerata (Douglas et al. 2002*). Other species with DOUGLAS AND PENNY: HOWELL’S TRITELEIA IN CANADA Wes x Ay 9 Fie: \ comoxS SER 3 ie Nil ¢ AV f =, South g T Wellington NANAIMO*\4> \ VANCOUVER ISLAND FiGureE 2. Distribution of Triteleia howellii in British Col- umbia (o — extirpated sites, e — recently confirmed sites). moderate to high constancies associated with T. how- ellii include Common Camas (Camassia quamash), Bromus spp., Broad-leaved Shooting-star (Dodecatheon hendersonii_ ssp. hendersonii), Cleavers (Galium aparine), and Common Snowberry (Symphoricarpos albus). A marked change in composition takes place by mid-summer. Many of the conspicuous native plants (e.g., Great Camas (Camassia leichtlinii), C. quamash, Dodecatheon hendersonii, and Yellow Montane Violet (Viola praemorsa ssp. praemorsa)) have completed their yearly life cycle and have essentially disappeared. Perennial grasses that were not recognizable or had not initiated growth in the spring and numerous intro- 176 THE CANADIAN FIELD-NATURALIST TABLE |. Locations and population sizes for Triteleia howellii in Canada Collection Last Site Observation Oak Bay (Victoria) 1912 Saanich Arm (Victoria) 1919 Witty’s Lagoon Regional Park (Metchosin) 1999 Gordon Head (Saanich) 1999 Cowichan Garry Oak Preserve (Duncan) 1999 Cowichan River Estuary (Duncan) 2001 Thetis Lake Regional Park (View Royal) 2002 Uplands Park (Victoria) 2003 Mt. Tzuhalem, base of (along Khenipsen Road) 2003 Albert Head Lagoon Regional Park (Metchosin) 2003 Horth Hill Regional Park (North Saanich) 2003 Beacon Hill Park (Victoria) 2004 Somenos Lake (Duncan) 2004 Williams Head Road (Metchosin) 2004 duced annuals, well adapted to the drier soils, domi- nate the understory. At this time, Dactylis glomerata and Vicia species are the most prominent species with greatly increased mean covers. Other prominent species in mid-summer include the native grasses, Bromus carinatus and Alaska Oniongrass (Melica subulata), and the introduced grasses, Barren Brome (Bromus sterilis) and Kentucky Bluegrass (Poa pra- tensis). Triteleia howellii also occurs in a Quercus garryana — Arbutus menziesii stand. The shrub layer is more prominent at this site and is dominated by Tall Oregon- grape (Mahonia aquifolium) and Oceanspray (Holo- discus discolor). Major associates include Hairy Honeysuckle (Lonicera hispidula), Bromus rigidus, Galium aparine, Small-flowered Nemophila (Nemo- phila parviflora) and Hedgehog Dogtail (Cynosurus echinatus). Biology There is little information known about the biology or ecology of Triteleia howellii throughout its range. Reproduction is by division of the corm, by produc- tion of numerous cormlets, and by seed (Barkworth 1977b). Population Attributes Triteleia howellii has been collected at 14 sites in Canada, all of which are located on southeastern Vancouver Island (Table 1). Of the 14 sites, 11 have been confirmed since 1999 while the status of the re- maining three sites is unknown and are likely extir- pated. Population areas range from small (one m7) to over three or four hectares, while plant numbers range from a single plant to over 430 plants (Table 1). Little information is available on population trends. The sites that have recently been examined show the populations Vol. 118 Collector/ Population Observer (no./area) Beaven Extirpated Newcombe Extirpated Douglas & Penny 43/200 m? Fontaine 51/5 m? Douglas 430/3-4 ha Douglas 62/3 m? Ceska 1/1 m?* Penny & Fairbarns Not seen since 1917, 2003 search unsuccessful, probably extirpated Janszen 6/.05 m?* Roemer 8/2 m? Janszen 3/1 m?* Fairbarns ca 200/12 m? Douglas & Richards 90/140 m? Milne 14/?m? are apparently stable although numbers of flowering plants may vary. Provincial, National and Global Ranks The British Columbia Conservation Data Centre has ranked this species S2 and placed it on the British Columbia Ministry of Sustainable Resource Manage- ment red-list (Douglas et al. 2002). This is the most critical category for imperilled rare native vascular plants in British Columbia. A rank of S2 is consid- ered “critically imperilled because of rarity (typically 6-20 extant occurrences or few remaining individuals) or because of some factor(s) making it vulnerable to extirpation or extinction.” Since the species is restrict- ed to British Columbia, the National rank is N2. Glo- bally, Triteleia howellii is ranked G3G4, indicating that, although the presently known sites are less than 100, it is more likely that this species is frequent to common in its range and apparently secure. Threats and Protection The most direct and immediate threat to Triteleia howellii is habitat destruction. This is of particular con- cern in the grass-dominated meadows often associ- ated with the Quercus garryana communities that are restricted to the southeastern side of Vancouver Island and some of the Gulf Islands. This type of vegetation was much more common before colonization by European settlers. This destruction has continued to the present resulting in the elimination of almost all sites occurring outside parks or ecological reserves. Historically, Q. garryana communities and grass- dominated meadows have always been heavily influ- enced by human activity, especially fires. Roemer (1972) believed that without human interference some of these stands would have eventually been replaced by Douglas-fir forests. 2004 DOUGLAS AND PENNY: HOWELL’S TRITELEIA IN CANADA 177 FiGure 3. Habitat of Triteleia howellii in a Quercus garryana stand in the Cowichan Garry Oak Preserve near Quamichan Lake. Dactylis glomerata is the dominant grass in this late summer photo. The suppression of fire within the past century may also have contributed to the decrease of Triteleia how- ellii populations. Most of the sites in which T: howellii has been collected were likely maintained in the past as a result of periodic fires, both natural and unnatural. In the past, aboriginal peoples probably set fire to these stands to maintain them as an important habitat for wildlife (Roemer 1972). Since that time, these sites have experienced little disturbance, resulting in the in- vasion and expansion of many other species, especial- ly introducted grasses. The introduction of European species has resulted in substantial, and probably irreversible, changes not only to the grass-dominated meadows associated with Quercus garryana, but also to the rocky xeric sites north and west of Victoria where Triteleia howellii has been collected in the past. One of the most devas- tating species is Scotch Broom (Cytisus scoparius), which has become a dominant shrub on xeric, exposed sites throughout much of southeastern Vancouver Island and the Gulf Islands. Much of the vegetation is now dominated by introduced grasses. These species include Early Hairgrass (Aira praecox), Sweet Vernal- grass (Anthoxanthum odoratum), Cynosurus echinatus and Dactylis glomerata. Some of the populations contain very few plants. Once a population becomes small, it becomes more vulnerable to demographic and environmental vari- ation and loss of genetic variability. In some cases, small populations are at risk of inbreeding depression, genetic drift and loss of fitness (Primack 1998). The population with the best protection is located at the Cowichan Garry Oak Preserve where the general public is excluded. This five-hectare stand near Quami- chan Lake on Vancouver Island represents the best example of a Quercus garryana woodland in the prov- ince and probably one of the best in the Pacific North- west. The stand contains a relatively low number of exotic species and has active management plans in place. This site also contains the largest known popu- lation of T; howellii in the province (Douglas et al. 2002*). A number of Triteleia howellii populations are also in small regional parks in the Greater Victoria area. These include populations at Beacon Hill, Witty’s Lagoon, Thetis Lake, and Horth Hill. Most of these parks receive little active management at the present time, at least with respect to their rare plants. Park enhancement projects, road and trail developments and heavy recreational use by humans often result in the destruction of the native vegetation and rare plant species. Triteleia howellii could be a candidate species for protection under the provincial Wildlife Amendment 178 Act as it is currently red-listed by the British Colum- bia Conservation Data Centre. One of the populations (Somenos Lake) of T: howellii is protected by the Park Act since it is currently administered by Parks BC. Evaluation The British Columbia Conservation Data Centre considers Triteleia howellii to be threatened/endangered in British Columbia (Douglas et al. 2002a) and the Committee on the Status of Endangered Wildlife in Canada has assessed the species as endangered (COSEWIC 2003). About 1000 flowering plants have been recorded in recent years from 11 locations. Only four of these populations can be considered viable since the remaining populations cover areas of less than 5 m?. The prognosis for this species may not be good since, since only one population, at the Cowichan Garry Oak Preserve, is included in an active management plan. Acknowledgments We thank Sharon Hartwell, Marie Fontaine, and Lora May Richards for their assistance with field work. We also thank Adolf and Oluna Ceska, Matt Fairbarns, Moralea Milne and Hans Roemer for information on populations at several sites. Documents Cited COSEWIC. 2003. COSEWIC assessment and status report on Howell’s Triteleia, Triteleia howellii in Canada. Com- mittee on the Status of Endangered Wildlife in Canada. www.cosewic.gc.ca, Ottawa, Ontario. 16 pages Douglas, G. W. 2003. COSEWIC Status report on Howell’s Triteleia, Triteleia howellii in Canada, in COSEWIC as- sessment and status report on Howell’s Triteleia, Triteleia howellii in Canada. Committee on the Status of Endan- gered Wildlife in Canada. www.cosewic.gc.ca, Ottawa, Ontario. 16 pages. Douglas, G. W., J. L. Penny, and R. E. Maxwell. 2002b. Composition, phenology, stand structure and soils of a Quercus garryana (Garry Oak) woodland community at Quamichan Lake, Vancouver Island, British Columbia. British Columbia Conservation Data Centre, Ministry of Sustainable Resource Management, Victoria, British Columbia. 52 pages. Douglas, G. W., and S. J. Smith. 2005. National multi-species recovery strategy for species at risk in Garry Oak wood- lands. In preparation. Ottawa, Ontario. Literature Cited Barkworth, M. E. 1977a. The taxonomic status of Brodi- aea howellii Watson (Liliaceae). Northwest Science 51: 139-148. Barkworth, M. E. 1977b. Intraspecific variation in Brodi- aea douglasii Watson (Liliaceae). Northwest Science 51: 79-90. Douglas, G. W., D. Meidinger, and J. L. Penny. 2002a. Rare native vascular plants of British Columbia. Second edition. Province of British Columbia. Victoria, British Columbia. 359 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999a. Illus- trated flora of British Columbia. Volume 3. Dicotyledons THE CANADIAN FIELD-NATURALIST Vol. 118 (Diapensiaceae through Onagraceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Colum- bia. 423 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 1999b. Illus- trated flora of British Columbia. Volume 4. Dicotyledons (Orobanchaceae through Rubiaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Colum- bia. 427 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2000. Illus- trated flora of British Columbia. Volume 5. Dicotyledons (Salicaceae to Zygophyllaceae) and Pteridophytes. British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 389 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2001a. Illus- trated flora of British Columbia. Volume 6. Monoco- tyledons (Acoraceae to Najadaceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests. Victoria, British Columbia. 361 pages. Douglas, G. W., D. Meidinger, and J. Pojar. 2001b. Illus- trated flora of British Columbia. Volume 7. Monocotyle- dons (Orchidaceae to Zosteraceae). British Columbia Ministry of Sustainable Resource Management and British Columbia Ministry of Forests, Victoria, British Columbia. 379 pages. Douglas, G. W., G. B. Straley, and D. Meidinger. 1998a. Illustrated flora of British Columbia. Volume 1. Gymno- sperms and Dicotyledons. (Aceraceae through Astera- ceae). British Columbia Ministry of Environment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 436 pages. Douglas, G. W., G. B. Straley, D. Meidinger, and J. Pojar. 1998b. Illustrated flora of British Columbia. Volume 2. Dicotyledons. (Balsaminaceae through Cuscutaceae). British Columbia Ministry of Envi- ronment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 401 pages. Keator, G. 1993. Triteleia. Pages 1206-1208. In The Jepson manual: Higher plants of California. Edited by J. C. Hickman. University of California Press. Berkeley, Cali- fornia. 1400 pages. Pojar, J. 2001. Liliaceae. Pages 270-318. Jn Illustrated flora of British Columbia. Volume 6. Monocotyledons (Acora- ceae to Najadaceae). Edited by G. W. Douglas, D. Meid- inger, and J. Pojar. British Columbia Ministry of Environ- ment, Lands and Parks and British Columbia Ministry of Forests, Victoria, British Columbia. 361 pages. Primack, R. B. 1998. Essentials of Conservation Biology. 2™4 Edition. Sinauer Associates Inc. Sunderland, Massa- chusetts. Roemer, H. L. 1972. Forest vegetation and environments of the Saanich Peninsula, Vancouver Island. Ph.D. thesis. Uni- versity of Victoria, Victoria, British Columbia. 405 pages. Scoggan, H. J. 1979. The flora of Canada. Part 4. Dicotyle- doneae (Loasaceae to Compositae). National Museum of Natural Sciences, Ottawa, Ontario. National Museum of Natural Sciences Publications in Botany (7): 1117-1710. Received 15 October 2002 Accepted | September 2004 Conservation Evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada TYLER W. SmITH!, GEORGE W. DOUGLAS? AND ALLAN G. HARRIS? 'Royal Botanical Gardens, P. O. Box 399, Hamilton, Ontario L8N 3H8 Canada Present address: Plant Science, McGill University, Raymond Building, 21 111 Lakeshore Road, Ste. Anne de Bellevue, Quebec H9X 3V9 Canada Conservation Data Centre, British Columbia Ministry of Sustainable Resource Management, Terrestrial Information Branch, P. O. Box 9993 Station Provincial Government, Victoria, British Columbia, V8W 9R7 Canada Present address: Douglas Ecological Consultants Ltd., 62030 North Road, Duncan, British Columbia V9L 6K5 Canada 3Northern Bioscience, 136 S. Hill Street, Thunder Bay, Ontario P7B 3V1 Canada Smith, Tyler W., George W. Douglas, and Allan G. Harris. 2004 Conservation Evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada. Canadian Field-Naturalist 118(2): 179-184. In Canada, Lipocarpha micrantha has been documented at eight locations in Quebec, Ontario, and British Columbia. Four of these populations have apparently been extirpated. The remaining populations, ranging from 120 to approximately 40000 plants, are all northern disjuncts from the main range of this species. Threats to these populations include water level regulation and shoreline development. Considering the threats to the habitat of Lipocarpha micrantha, and the small size of most of the remaining populations, it has been designated an Endangered species in Canada. Key Words: Small-flowered Lipocarpha, Lipocarpha micrantha, British Columbia, Ontario, Quebec, endangered, distribution, population size. Small-flowered Lipocarpha, Lipocarpha micrantha (Vahl.) G. Tucker, is the only member of a mainly trop- ical genus to occur in Canada. While it appears under this name in the Flora of North America (Tucker 2002), some recent treatments place it in the genus Hemicar- pha Nees (Gleason and Cronquist 1991). Lipocarpha, including Hemicarpha, is generally accepted as a dis- tinct genus. However, taxonomic clarification of closely related genera in the Cyperaceae may require nomen- clatural revisions, pending the outcome of ongoing research (Muasya et al. 2002). Lipocarpha micrantha is a caespitose annual sedge, 2-20 cm tall with narrow (0.5 mm) leaves up to 10 cm long. The 1 to 3 subsessile flower spikes are borne at the top of the stem. The spikes are ovoid, 2—6 mm long, with numerous perfect flowers concealed behind spi- rally imbricate scales. The inflorescence is subtended by 2 to 3 leaf-like bracts, the lowest of which looks like a continuation of the stem (Figure 1: Hitchcock et al. 1969; Gleason and Cronquist 1991; Douglas and Ceska 2001; Tucker 2002). The densely tufted plants resemble the seedlings of many other sedge species that occur in the same habitat. In Canada Lipocarpha micrantha is perhaps most simi- lar in aspect to Awned Cyperus (Cyperus squarrosus L.), another small annual sedge. Lipocarpha micrantha is distinguished from this and all other Cyperus spe- cies by its spirally arranged flowers. Biology Lipocarpha micrantha is only visible during a short period each year. It germinates in late summer, when dropping water levels expose the open sandy habitat it requires. Flowering and fruiting occur in August and September. High water levels may prevent germination from occurring, with the population remaining dormant until appropriate conditions occur. We do not know how long dormant seeds may remain viable, but field observations suggest they can persist at least two years. Migrating waterfowl may be a vector for long-distance dispersal. Distribution Lipocarpha micrantha ranges from Brazil to Canada. North of Mexico, the main range extends from Texas, north to northwestern Ontario, and east to the east coast of the United States. Disjunct populations have been recorded east and west of the main range, in Alabama, Arizona, British Columbia, California, Florida, Ken- tucky, Maine, New Mexico, New York, North Carolina, and Quebec (Tucker 2002). In Canada, Lipocarpha micrantha occurs at Osoyoos Lake and Okanagan Lake in British Columbia (Figure 2), and at Rainy Lake and Lake of the Woods in north- western Ontario (Figure 3). Populations along the De- troit River in southern Ontario and along the north shore of Lake Champlain in Quebec have been extirpated. Habitat Lipocarpha micrantha grows on sandy beaches and interdunal swales that are subject to seasonal flooding, but are protected from high waves or strong currents. It is usually found in areas of very sparse vegetation, and apparently is intolerant of competition from other plant species. These habitat conditions are maintained by fluctuating water levels. While Lipocarpha micrantha 179 180 THE CANADIAN FIELD-NATURALIST Vol. 118 Ficure 1. Illustration of Lipocarpha micrantha (Line drawing by permission from Hitchcock et al. 1969). requires seasonal low water levels to germinate and flower, periodic high water is required to prevent more vigorous species from dominating its shoreline habitat (see Keddy and Reznicek 1986 for a discussion of the relationship between water level fluctuations and wetland vegetation). Common associates of Lipocarpha micrantha in- clude a variety of Cyperus, Bidens, and Salix species. Cyperus squarrosus has been noted as an associate at every station in Canada. Population Trends and Limiting Factors Of eight documented populations (Ceska and Ceska 1980; Oldham and Crins 1988; Sabourin et al. 1992*; Oldham 1996; Harris et al. 2000*; Oldham 2000; Smith et al. 2002*) of Lipocarpha micrantha in Canada, only four persist (Table 1). The Lake Champlain population, in Missisquoi Bay, Quebec, was discovered in 1953 (Louis-Alphonse 3458, specimen at MT; herbarium acronyms follow Holmgren et al. 2003*), and was documented through- out the 1950s. However, a search in 1989 failed to find any plants (Sabourin et al. 1992*). Most recently, TWS searched the site in 2002 but did not find any plants despite the presence of appropriate habitat and moder- ate water levels. This site is adjacent to a campground and recreational activity may have contributed to the demise of this population. Water quality may also have been a factor, as a river flowing into Lake Champlain at this location smells strongly of raw sewage. 2004 —~ —— ———> \ SMITH, DOUGLAS, AND HARRIS: SMALL-FLOWERED LIPOCARPHA 181 KAMLOOPS SPENCES BRIDGE I i) aw m4 7 _) \ENDERBY _/ ARMSTRONG / . Okanagan ; Z Te camoce KEREMEOS, BRITISH COLUMBIA NE Lake . j Ree NCETON ~y PENTICTON J On Ficure 2. Distribution (stars) of Lipocarpha micrantha in British Columbia. The oldest records of Lipocarpha micrantha in Can- ada are collections from the Detroit River shoreline, south of Windsor, Ontario, in 1892 (Macoun 28668, specimen at CAN) and 1901 (Macoun 7594, speci- mens at TRTE, GH). The populations represented by these collections have presumably been destroyed by shoreline development. The only recent records from this region are from the vicinity of Holiday Beach Con- servation Area, near Amherstburg. This population was discovered in 1984, when 15 plants were observed (Oldham and Crins 1988). The population was still present in 1987 (Sabourin et al 1992*), but no plants were found during two searches by TWS in 2001. The habitat at Holiday Beach has been seriously degraded: dense mats of algae covered the beach, except in areas where it had been cleared away by heavy machinery. 182 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Locations and Population Sizes for Lipocarpha micrantha in Canada. Site Last Observer (Last Population Observation Searcher) (number/area) Osoyoos Lake (British Columbia) 1980 Ceska Extirpated Osoyoos Lake, 2001 Douglas 30 000-50 000/2.3ha Osoyoos Indian Reserve (British Columbia) Okanagan Lake, 2002 Klinkenberg 120/2 m?* Sun Oka Beach Provincial Park (British Columbia) Sable Island (Ontario) 2001 Harris 1 800/0.4ha Poundnet Bay (Ontario) 2000 (see text) Harris 75/0.00lha Holiday Beach (Ontario) 1987 (2001 ) Oldham (Smith) Extirpated Detroit River (Ontario) 1901 Macoun Extirpated Missisquoi Bay (Quebec) 1957 (2002 ) Louis-Alphonse Extirpated (Smith) The Lake of the Woods population, at Sable Island, Ontario, was discovered in 1995, and several thousand plants were noted at that time (Oldham, personal com- munication 2001). Surveys in 2001 documented approx- imately 1800 plants. The long sandspit island provides large areas of suitable habitat in Sable Island Prov- incial Nature Reserve. As a consequence of the ephemeral nature of its habitat and its ability to remain dormant in unfavour- able years, Lipocarpha micrantha may persist un- detected at locations searched thoroughly by botanists. An unsuccessful search during highwater, when much potential habitat is temporarily unavailable, is there- fore not justification for declaring a population extir- pated. This was the case at Poundnet Bay on Rainy Lake, Ontario, in 2001, where water levels were well above average. No Lipocarpha micrantha plants were found there during 2001 A population of 75 plants was discovered at this location in 2000. When water levels return to average or below this area will provide habitat for Lipocarpha micrantha. It is known from several beaches on the Minnesota side of Rainy Lake. Lipocarpha micrantha was first documented on Osoyoos Lake, British Columbia, by Ceska and Ceska (1980). Of the two populations noted, one has been eradicated by shoreline development. The British Col- umbia Conservation Data Centre has been monitor- ing the second population since 1991. In 2001 a total of between 30 000 and 50 000 plants were invento- ried. This is the highest recorded population at that site, and it is the largest population in Canada. The high number is attributed to excellent growing conditions, and also to the higher search intensity in 2001. If con- ditions remain stable at this site the population should continue to thrive. However, local development plans at the site, including a casino and marina, would likely destroy about 50 to 60% of the remaining habitat. A third, small (20 plants over 2 m*) population was dis- covered on the shoreline of Okanogan Lake in 2002. Special Significance of the Species Lipocarpha micrantha has very specific habitat requirements: open, sandy shorelines, protected from strong waves, with limited competition from other plants. It is extremely sensitive to alteration of both water levels and shoreline structure. As such it may be an important indicator of wetland quality. Protection Lipocarpha micrantha has a NatureServe (2002*) global rank of G5 or “secure”. In the United States, Lipocarpha micrantha is listed as Endangered in Connecticut, Maryland, New Jersey, New York, and Pennsylvania, and it is listed as Threatened in Maine and Ohio (United States Department of Agriculture, Natural Resources Conservation Service 2002*). In Canada, Lipocarpha micrantha is ranked as N1 or “critically imperilled”, and is listed as Threatened in Ontario (NHIC, 2003*) and Endangered/Threatened in British Columbia (Douglas et al. 2002). Both extant Ontario populations of Lipocarpha micrantha are in a Provincial Nature Reserve and a Provincial Conservation Reserve. As such they are pro- tected from shoreline development, but are subject to water level regulation. The extant populations in British Columbia occur on an Indian Reserve (Osoyoos Lake Indian Reserve #1) and in a Provincial Park (Sun Oka Beach Provincial Park). There is currently no specific provincial rare species legislation in place for the pro- tection of endangered/threatened vascular plants in British Columbia. At the federal level the Species at Risk Act protects COSEWIC-listed plants on federal lands (which includes Indian Reserves). It also empow- ers the federal government to protect habitat outside federal lands if the province fails to protect listed spe- cies or their habitat; these mechanisms are discretionary. Evaluation of Status The status of Lipocarpha micrantha in Canada is precarious. The largest, apparently stable, population at Osoyoos Lake in British Columbia is threatened by planned development. Should this development proceed, approximately half of the remaining Lipocarpha mic- rantha plants in Canada will be physically destroyed, and the remaining habitat may no longer support the species. The two remaining Ontario populations are 2004 SMITH, DOUGLAS, AND HARRIS: SMALL-FLOWERED LIPOCARPHA 183 FiGure 3. Distribution of Lipocarpha micrantha in Ontario and Quebec. Sable Island is shown as a circle, Ponies: Bay as a triangle. Squares denote extirpated populations. relatively small, both in number and area, and there- fore are vulnerable to environmental disturbance. If development at any of the remaining populations pro- ceeds without consideration for this species it will very likely be extirpated from Canada. In particular, altera- tion of natural water level cycles could result in the destruction of the habitat required by Lipocarpha mic- rantha. The authors’ recommendation that Lipocarpha micrantha be uplisted to Endangered was accepted by COSEWIC in November 2002. Acknowledgments The authors gratefully acknowledge the following people for assistance with this report. Kara Brodribb, Wasyl Bakowsky, Dan Dufour, Rob Foster, Ron Hall, Erica Oberndorfer, Mike Oldham, Kathy Paige, Amy Tesselin, and Christine Thuring assisted with field sur- veys. Wasyl Bakowsky, Brian and Rose Klinkenberg, Jim Meeker, Mike Oldham, Tony Reznicek and David Szymanski provided location and habitat data. Erica Oberndorfer, Jenifer Penny and Shyanne Smith assist- ed with the production of maps and compilation of status data. Graham Rose (Big Creek Hunt Club) gra- ciously granted access to their properties for survey work. This paper is based on a status report submitted to the Committee On the Status of Endangered Wildlife in Canada (COSEWIC), September 2001. The distri- bution map for Ontario and Quebec, Figure 3, was provided by COSEWIC. Funding was provided by COSEWIC, the George Cedric Metcalf Foundation, Royal Botanical Gardens, and the British Columbia Conservation Data Centre. Documents Cited Harris, A. G., M. J. Oldham, R. F. Foster, and W. D. Bakowsky. 2000. Preliminary life science inventory of Rainy Lake, Ontario. Rainy Lake Conservancy, Rainy Lake, Ontario. Holmgren, P. K, N. H. Holmgren, and L. C. Barnett. 2003. Index Herbariorum [web application]. Updated October 1, 2003. New York Botanical Garden, Bronx, New York. Available at: http://www.nybg.org/bsci/ih/ih.html. Natural Heritage Information Centre. 2003. Natural Heri- tage Information Centre Database (http://www.mnr.gov. on.ca/MNR/nhic/data.cfm). Natural Heritage Information Centre, Ontario Ministry of Natural Resources. NatureServe 2002. NatureServe Explorer: An online ency- clopedia of life [web application]. 2002. Version 1.6. Arling- ton, Virginia, USA: NatureServe. Available at http://www. natureserve.org/explorer. Sabourin, A., M. J. Oldham, and D. Paquette. 1992. Status report on the Small-flowered Lipocarpha, Lipocarpha micrantha (syn. Hemicarpha micrantha) in Canada. COSE- WIC, Ottawa, Ontario. 184 Smith, T. W., G. W. Douglas, and A. G. Harris. 2002. Update COSEWIC Status Report on Small-Flowered Lipo- carpha Lipocarpha micrantha (syn. Hemicarpha micran- tha). COSEWIC, Ottawa, Ontario. United States Department of Agriculture, Natural Re- sources Conservation Service. 2002. The PLANTS Data- base, Version 3.5 (http://plants.usda.gov). National Plant Data Centre, Baton Rouge, Louisiana. Literature Cited Ceska, A., and O. Ceska. 1980. Additions to the flora of Brit- ish Columbia. Canadian Field-Naturalist 94: 69-74. Douglas, G. W., and A. Ceska. 2001. Cyperaceae. Pages 12-199 in Illustrated flora of British Columbia. Volume 6. Monocotyledons (Acoraceae to Najadaceae). Edited by G. W. Douglas, D. Meidinger and J. Pojar. 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Phylogenetic relationshipsin Cyperus L. s.l. (Cyperaceae) inferred from plastic DNA sequence data. Botanical Jour- nal of the Linnean Society 138: 145-153. Oldham, M. J. 1996. Interesting plant records from north- west Ontario. Ontario Natural Heritage Information Centre Newsletter 3. Oldham, M. J. 2000. Botany Highlights. Ontario Natural Heritage Information Centre Newsletter 6: 10-11. Oldham, M. J., and W. J. Crins. 1988. New and significant records of Ontario sedges (Cyperaceae). Canadian Field- Naturalist 102: 500-507. Tucker, G. C. 2002. Lipocarpha. Pages 195-197 in Flora of North America North of Mexico. Volume 23. Cyperaceae. Edited by Flora of North America Editorial Committee. Oxford University Press Incorporated, New York, New York. 608 pages. Received 13 November 2003 Accepted 5 April 2004 Immobilization of Clover-trapped White-tailed Deer, Odocoileus virginianus, with Medetomidine and Ketamine, and Antagonism with Atipamezole JosHUA J. MILLSPAUGH!, BRIAN E. WASHBURN!, TAMARA M. MEYER?, JEFF BERINGER?, and LONNIE P. HANSEN? ‘Department of Fisheries and Wildlife Sciences, University of Missouri, 302 Anheuser-Busch Natural Resources Building, Columbia, Missouri 65211 USA *Missouri Department of Conservation, Conservation Research Center, 1110 South College Avenue, Columbia, Missouri 65201 USA Millspaugh, Joshua J., Brian E. Washburn, Tamara M. Meyer, Jeff Beringer, and Lonnie P. Hansen. 2004. Immobilization of Clover-trapped White-tailed Deer, Odocoileus virginianus, with medetomidine and ketamine, and antagonism with atipamezole. Canadian Field-Naturalist 118(2): 185-190. We evaluated the effectiveness of immobilizing Clover-trapped White-tailed Deer (Odocoileus virginanus) with medeto- midine hydrochloride (HCl) and ketamine HCI during winter and summer by monitoring immobilization intervals and vital signs. In winter, we captured deer in Clover traps in | 4-ha research enclosure for relocation to another on-site enclosure (n = 5). In summer, we captured free-ranging deer in Clover traps to attach radio-collars (n = 4). We administered an estimated 0.055 mg/kg medetomidine HCI and 2.5 mg/kg ketamine HCI to adult (> 1.5 years of age) deer and 0.06 mg/kg medetomidine HC] and 2.5 mg/kg ketamine HCl to subadult (< 1.5 years of age) deer. We used an intramuscular injection of atipamezole HC] as the antagonist at a rate of 0.275 mg/kg for adults and 0.3 mg/kg for subadults > 30 minutes post-induction. Mean induction time in winter was 11.2 minutes (SE = 2.5, range = 5.4 — 24.2) and 6.5 minutes (SE = 0.8, range = 6.2 — 7.5) in summer. After atipame- zole HCl injection, the mean time to walking was 17.1 minutes (SE = 3.5, range = 7.5 — 41.5 minutes) in winter and 11.3 minutes (SE = 3.8, range = 4.7 — 13.5) in summer. Rectal temperature was relatively constant throughout immobilization; however rectal temperatures of 5 deer (7 = 3 in winter; n = 2 in summer) exceeded 40°C, a sign of hyperthermia. Respiration rate and pulse rate peaked at about 20 minutes post-medetomidine HCI and ketamine HCl injection, then generally declined thereafter. No mortalities were observed in our study. Medetomidine HCI and ketamine HCI doses for Clover-trapped White-tailed Deer pro- vided satisfactory induction times, sufficient level of anesthesia for short-distance relocation or radio-collar Eucte alee and were effectively reversed with an IM injection of atipamezole HCl. Key words: White-tailed Deer, Odocoileus virginianus, atipamezole, capture, chemical restraint, Clover trap, deer, ketamine, immobilization, medetomidine, Missouri. White-tailed Deer (Odocoileus virginianus) have been chemically immobilized with Telazol® (1:1 tile- tamine hydrochloride (HCl) and zolazepam HCl) and xylazine HCI (Schultz et al. 1992; Kilpatrick and Spohr 1999), ketamine HCl and xylazine HCI (Mech et al. 1985; Farley et al. 1986; Kreeger et al. 1986; Ballard et al. 1998; Kilpatrick and Spohr 1999), etorphine and xylazine HC] (Presnell et al. 1973; Presidente et al. 1973; Nielsen 1982), xylazine HCI alone (Gibson et al. 1982), phencyclidine HCl (Dean et al. 1973), and succinylcholine chloride (Wesson et al. 1974; Jacobsen et al. 1976). Kreeger (1996) recommended 4.4 mg/kg Telazol® and 2.2 mg/kg xylazine HCl to immobilize White-tailed Deer; 0.125 mg/kg yohimbine HCl was the recommended antagonist. Recently, Kilpatrick and Spohr (1999) used a 4.8:3.9 mg/kg dose of Telazol®: xylazine HCI to dart free-ranging White-tailed Deer. Alternative drugs recommended by Kreeger (1996) include ketamine HCl and xylazine HCl, etorphine, xylazine alone (for calm deer only), and a combina- tion of medetomidine HCI and ketamine HCl. Medetomidine HCl depresses the central nervous system and acts similarly to xylazine HC] (Jalanka and Roeken 1990), but with greater affinity to alpha,- adrenoreceptors (Klein and Klide 1989; Jalanka and Roeken 1990; Kreeger 1996). Ketamine HCl, an anes- thetic, is often combined with a tranquilizer or seda- tive to improve induction and recovery (Haigh 1982; Kreeger 1996). Medetomidine HCl and ketamine HC] alone or in combination with other drugs have been suc- cessfully used to immobilize a diversity of large ungu- lates including Reindeer (Rangifer tarandus tarandus) (Ryeng et al. 2001, 2002), Mule Deer (Odocoileus hemionus) (Caulkett et al. 2000), Mule Deer/White- tailed Deer hybrids (Caulkett et al. 2000), Sika Deer (Cervus nippon) (Tsuruga et al. 1999), gemsbok (Oryx gazella) (Grobler et al. 2001), Roan Antelope (Hippo- tragus equinus) (Citino et al. 2001), Blue Duiker (Cephalophus monticola) (Bailey et al. 1995), Moose (Alces alces) (Arnemo 1995), Tigers (Panthera tigris) (Miller et al. 2003), Impala (Aepyceros melampus) (Bush et al. 2004), European Mink (Mustela lutreola) Four- nier-Chambrillon et al. 2003) and Red Deer (Cervus elaphus) (Arnemo et al. 1994). Notable among the advantages listed in these and other studies is the ability to reverse medetomidine HC] 185 186 with an intramuscular (IM) injection of atipamezole HCl (Tsuruga et al. 1999; Haulena et al. 2000). Atipa- mezole HCI is an extremely efficient alpha,-adren- ergic antagonist compared with yohimbine HCl and tolazoline HCl (Kreeger 1996) and effectively reverses medetomidine HCl in many wildlife species (Tsuruga et al. 1999; Haulena et al. 2000; Grobler et al. 2001). If medetomidine HCl and ketamine HC] were as effi- cient and safe as other immobilizing agents (e.g., Tela- zol® and xylazine HCl) and if an IM injection of ati- pamezole HCI was a safe and effective antagonist, this combination could prove efficacious in field studies. Our objective was to determine the effectiveness and safety of immobilizing Clover-trapped White-tailed Deer with medetomidine HCI and ketamine HCI, and the feasibility of reversing this combination with an IM injection of atipamezole HCl, during winter and sum- mer by monitoring immobilization intervals and vital signs. To our knowledge, an evaluation using mede- tomidine HCI and ketamine HCl to immobilize free- ranging White-tailed Deer and antagonism by ati- pamezole HCI has not been previously published. Methods Our review of the medetomidine HCI and ketamine HCl combination took place in winter and summer 2001. In summer and winter, we trapped White-tailed Deer in Clover traps (McCullough 1975). During win- ter we trapped deer at the Charles W. Green Conser- vation Area, located near Ashland, Missouri. Traps were baited with corn, set each evening, and checked at sunrise. All winter-trapped and immobilized deer (n = 5) were captured within a 4-ha research enclo- sure for relocation to another on-site 4-ha enclosure. These deer were not habituated to humans. In summer, we trapped White-tailed Deer at the Thomas S. Baskett Wildlife Research and Education Area, located near Ashland, Missouri. Traps, baited with salt, alfalfa, and corn, were set each evening and checked at sunrise. We captured and immobilized summer-trapped deer (n = 4) for purposes of radio- collar attachment. These deer were free-ranging ani- mals and not habituated to humans. In winter and summer, we immobilized adult deer (> 1.5 years of age) (n = 2 in winter; n = 3 in sum- mer) using an estimated 0.055 mg/kg medetomidine HCl and 2.5 mg/kg ketamine HCI; 0.06 mg/kg mede- tomidine HCI and 2.5 mg/kg ketamine HCI was ad- ministered to subadult deer (< 1.5 years of age) (n = 3 in winter; n = 1 in summer). Adult deer, estimated to be 60 kg, were given 3.3 mg of | mg/ml Domitor® (medetomidine HCl; Orion Corporation, Orion— Farmos, Espoo, Finland) and 150 mg of 100 mg/ml Ketaset® (ketamine HCl; Fort Dodge Laboratories, Inc., Fort Dodge, Iowa, USA). Subadult deer, estimat- ed to be 30 kg, were given 1.8 mg of 1 mg/ml mede- tomidine HCl and 75 mg of 100 mg/ml ketamine HCl. We injected drugs IM into the biceps femoris with a THE CANADIAN FIELD-NATURALIST Vol. 118 hand syringe. Sex and age (adult or subadult) were recorded and during winter each deer was marked with a plastic cattle ear tag in one ear for later identification. In summer, deer were fitted with radio-transmitters. Following sedation in winter and summer, we ap- plied an ophthalmic ointment and blindfolded the deer. In winter, we relocated deer to a different on-site 4-ha enclosure. Two or three field assistants placed the sed- ated deer into the rear of a vehicle (range from 100 to 300 m away), which was driven to the release enclo- sure (< | km driving distance). Each deer was carried into the enclosure (< 30 m away), and placed in a ster- nal recumbent position. We reversed the medetomidine HCI and ketamine HCI combination with an IM injection of Antisedan® (atipamezole HCI; Orion Corporation, Orion—Farmos, Espoo, Finland) into the biceps femoris with a hand syringe at a rate of 0.275 mg/kg (16.5 mg of 5 mg/ml atipamezole HCl) for adults and 0.3 mg/kg (9 mg of 5 mg/ml atipamezole HCl) for subadults. We visually monitored all deer until they departed the area. During winter and summer, we attempted to monitor immobilization intervals and vital signs at 5-minute intervals. For all deer, we recorded time of medetomi- dine HCI and ketamine HCl injection, induction (time from injection to time animal was handled), atipame- zole HCl administration, “head up” (time when the animal first lifted its head), “standing” (time when the animal first stood up), and “walking” (time when the animal successfully departed the area). We also recorded respiration rate (breaths/minute), rectal tem- perature (°C), and pulse rate (beats/minute) at 5-min- ute intervals beginning at the time of medetomidine HCI and ketamine HCl injection for respiration rate and beginning 10 minutes post-medetomidine HCl and ketamine HCl injection for temperature and heart rate. Results Nine deer (1 = 5 in winter, including 2 female sub- adults, 1 male subadult, | female adult, and 1 male adult; n = 4 in summer, including 3 adult females and 1 female subadult) were immobilized using the drug combination described above and either relocated or equipped with a radio-collar. No mortality has been observed 10 months post-winter immobilization and 5 months post-summer immobilization. Mean induction time was 11.2 minutes (SE = 2.5, range = 5.4 — 24.3) in winter and 6.5 minutes (SE = 0.8, range = 6.2 — 7.5) in summer. Time to atipamezole HCl injection averaged 54.4 minutes (SE = 3.7, range = 37.7 — 79.2) in winter and 33.4 (SE = 1.3, range = 30.4 — 33.8) in summer. After atipamezole HCl injection, the mean time to head up was 11.4 minutes (SE = 2.9, range = 4 — 27.5) in winter and 9.3 minutes (SE = 0.7, range = 8.4 — 9.9) in summer. Mean time to standing was 15.9 minutes (SE = 3.6, range = 4.4 — 41) in win- ter and 10.5 minutes (SE = 3.2, range = 4.7 — 12.3) in summer. The mean time to walking was 17.1 minutes 2004 MILLSPAUGH, WASHBURN, MEYER, BERINGER, AND HANSEN: IMMOBILIZATION OF WHITE-TAILED DEER 187 (SE = 3.5, range = 7.5 — 41.5 minutes) in winter and 11.3 minutes (SE = 3.8, range = 4.7 — 13.5) in summer. With the exception of rectal temperatures, other vital signs were considered normal (Table 1). Rectal temperatures were stable from 10 — 30 minutes post- medetomidine HCl and ketamine HCl injection in summer and winter (Table |). Temperatures of 1 sub- adult female captured in winter were 37.4°C and 36.2°C at 70 and 110 minutes post-induction, respectively; thus, she was within 1.2°C of becoming hypothermic (defined as < 35°C; Kreeger 1996; DelGuidice et al. 2001). No attempt was made to increase body temper- ature of this individual prior to antagonism with atipa- mezole HCl. After atipamezole HCI administration, it took that individual 41.5 minutes to depart, the max- imum time observed in our study. Rectal temperatures of 5 deer (n = 3 in winter, m = 2 in summer) exceeded 40°C, a sign of hyperthermia. No attempt was made to decrease body temperature of these animals prior to antagonism. Respiration rates showed little variability in summer and winter and were generally in the upper 20’s to low 30’s (breaths/minute) (Table 1) peaking at about 20 minutes post-medetomidine HCl and ketamine HCl injection. Pulse rates peaked about 20 minutes post-medetomidine HCl and ketamine HCl injection at 95 beats/minute (SE = 3.6) in winter and 103 beats/minute (SE = 1.4) in summer and declined thereafter to 74 beats/minute (SE = 5.6) in winter and 88 beats/minute (SE = 3.1) in summer at 30 minutes post-medetomidine HCI and ketamine HCl injection (Table 1). No other adverse side effects were noted. During summer captures, response to IM injection of atipamezole HCl was predictable, as previously described by Jalanka and Roeken (1990). Within 3 — 6 minutes of atipamezole HCl injection, “ear-twitch- ing” occurred, followed by leg extensions after an addi- tional 3 — 6 minutes, and “head up” 2 minutes there- after. Standing followed within another 2 minutes and the animal departed almost immediately with good muscle coordination. Discussion Medetomidine HCl] and ketamine HCl doses for Clover-trapped deer provided satisfactory induction times, produced a sufficient level of anesthesia for short-distance relocation or radio-collar attachment, and were effectively reversed with an IM injection of atipamezole HCI. Small dosage volume, ease of prepa- ration and predictable responses to sedation and to the antagonist make this combination a useful alterna- tive to drug combinations that may require prolonged recovery. Medetomidine HCl and ketamine HCl provided induction times similar to those reported in other studies and with other drugs. For 13 captive White- tailed Deer in Minnesota, it took 2 to 35 minutes (medi- an = 8, SE = 1.2) from the time of xylazine HCI and ketamine HCl administration before deer lost the ability to stand (Mech et al. 1985). Our mean induction time in winter (x = 11.2 minutes, SE = 2.5, range = 5.4 — 24.3, n = 5) was about double the median time of 6.2 minutes (range = 0.5 — 17.3) reported by Jalanka and Roeken (1990) for 28 captive White-tailed Deer housed at the Helsinki Zoo, but similar to our summer mean time of 6.5 minutes (SE = 0.8, range = 6.2 — 7.5). In the Helsinki Zoo study, deer were given an average of 61 ug/kg (SD = 14, median = 58, range = 37 — 98) medetomidine HCI and a mean ketamine HCI dose of 1.6 mg/kg (SE = 0.3, median = 1.5, range = 1 — 2.3). The time to walking after the IM injection of ati- pamezole HCI was similar to that for deer in other studies reversed with yohimbine HCl, but was less TABLE |. Mean + SE (N) vital signs of White-tailed Deer immobilized with medetomidine hydrochloride (HCl) and ketamine HCl, and antagonized with atipamezole HCl during winter and summer 2001 in mid-Missouri for purposes of relocation (winter) and radio-collar attachment (summer). Adults were immobilized with 0.055 mg/kg medetomidine HCI and 2.5 mg/kg ketamine HCl and subadults were immobilized with 0.06 mg/kg medetomidine HCl and 2.5 mg/kg ketamine HCl. Adults were antagonized with 0.275 mg/kg of atipamezole HCI and subadults were antagonized with 0.03 mg/kg of atipamezole HCI. We have not reported data where information was collected on < 2 deer. Time 0 is the time of medetomidine HCI and ketamine HCl injection. Winter Summer Time Respiration Rate Pulse Rate Temperature Respiration Rate Pulse Rate Temperature (minutes) (breaths/minute) (beats/minute) (c) (breaths/minute) (beats/minute) (°C) 0 PH pd =e PEMA G)) — - 23 + 1.4 (4) - ~ 5 DifeD) 7D) -3)()) - - 23 + 2.2 (4) - - 10 28.8 + 2.9 (5) 87 +3.6(4) 41.1405 (2) 31+2.4 (4) 98+2.0(4) 39.9+ 1.2 (4) 15 29.6 + 3.2 (5) 88 +43(4) 40.9+0.8 (4) 31+2.2(4) 101+1.9(4) 39.8414 (4) 20 SI =n 244) 94.7+3.6(3) 40.62 1.2 (3) 30+2.6(4) 103+1.9(4) 39.8+1.4(4) 25 Posy \oe2 Syil (G) 86 +3.5(4) 404+1.2 (4) 30+3.1(4) 101424(4) 39.6+1.3 (4) 30 29.3 + 3.6 (4) Ase) 16\(2) wae 3 9-3 -t ile 9 (2) 32 + 2.9 (4) 88+3.1(4) 39.3+1.2 (4) 35 24 +2.4(2) Diet) ED (2) - 24 + 0.0 (2) ~ = 40 - — ~ 24 + 0.0 (2) - ~ 45 Syd" Tes 3}5) (3) - - — — = 50 PR: a2 3)3} (8) - 40.1 + 1.2 (2) - - - 188 variable. An intravenous injection of yohimbine HCl following xylazine HCl and ketamine HCl immo- bilization resulted in an adult male walking in 1.5 min- utes (SE = 0.5, n = 2 immobilizations) to 26.5 minutes (SE = 11.5, n = 4 immobilizations) for an adult fe- male (Mech et al. 1985). For 22 White-tailed Deer (18 free-ranging captured using drop nets and 4 cap- tive) immobilized with xylazine HCl and ketamine HCl, an IM injection of yohimbine HCI produced a mean recovery time of 11.6 minutes (SE = 2.3) (Wallingford et al. 1996). Hsu and Shulaw (1984) used an IV injection of yohimbine HCI and reported a mean recovery time of 4.4 minutes (SD = 5.4 minutes) for xylazine HCl-immobilized deer. With the exception of rectal temperature, the vital signs observed in this study were within the normal range of reported values for White-tailed Deer. Mautz and Fair (1980) reported pulse rates of a 46-kg adult White-tailed Deer female in July that was lying, stand- ing/walking, and running at 65, 74, and 106 beats per minute, respectively. Pulse rates observed in this study were similar to predicted walking pulse rates of White- tailed Deer (Moen 1978: 722), yet higher than those rates reported by Jalanka and Roeken (1990: 267). Rectal temperatures of five deer were above 40°C, a sign of hyperthermia (Kreeger 1996). Average rectal temperatures in our study in winter (39.3°C at 30 minutes, SE = 1.9, n = 2) and summer (39.3°C at 30 minutes, SE = 1.2, n = 4) are similar to the maximum rectal temperatures reported by DelGuidice et al. (2001:1151) for White-tailed Deer captured by Clover trap and immobilized with xylazine HCI and ketamine HCI during the winter in Minnesota. Also in Minne- sota, Rogers et al. (1987) reported rectal temperatures of two female free-ranging White-tailed Deer fawns in all seasons during various activities ranged between 38.2° and 40.1°C. Rectal temperatures of 3 adult male White-tailed Deer in Mississippi, averaged 39.3°C during late August and September and 38.6°C in early December (Demarais et al. 1986). As suggested by DelGiudice et al. (2001), corrective actions (i.e., packing snow around the animal) should be taken when immobilized animals become hyperthermic. Consequently, rectal temperatures should be monitored and protocols should be established to determine when corrective actions should begin (DelGiudice et al. 2001). Both medetomidine HCl and ketamine HCl have wide safety margins, produce calm inductions in several artiodactyls, are safe for humans (Jalanka and Roeken 1990), and have not caused any apparent detrimental effects in pregnant females (Jalanka 1993). Few adverse side effects were noted by Jalanka and Roeken (1990) after 1240 immobilizations with medetomidine HCl, ketamine HCl, and antagonism with atipamezole HCl. Worth noting, some ruminants became resedated be- tween 30 and 240 minutes post-IV reversal with ati- pamezole HCI (Jalanka and Roeken 1990) and unre- THE CANADIAN FIELD-NATURALIST Vol. 118 markable ruminal tympany was common in ruminants prior to atipamezole HCI administration (Jalanka and Roeken 1990; Jalanka 1993). A disadvantage of the medetomidine HCl, ketamine HCl, and atipamezole HC] combination is cost. In U.S. currency, cost to immobilize each adult deer was $34.72 ($32.76 for medetomidine HCI and $1.96 for ketamine HCl) and $18.85 for each subadult deer ($17.87 for medetomidine HC] and $0.98 for ketamine HCl). Atipamezole HCI cost $33.22 per adult deer and $18.14 for each subadult deer. In contrast, Kilpatrick and Spohr (1999) reported a cost of $8.44/deer for Telazol® and xylazine HCl immoblized deer and $6.34/ deer for ketamine HCI and xylazine HCI without lyo- philizing costs. With lyophilizing, costs were $10.05/ deer for Telazol® and xylazine HCl and $16.49 for ketamine HCl and xylazine HCI (Kilpatrick and Spohr 1999). Consequently, use of medetomidine HCl, keta- mine HCl, and atipamezole HCI may be cost-prohib- itive in studies requiring immobilization of many ani- mals. However, given the desirable properties discussed above, including rapid reversal that requires less man- power time, these desirable qualities may outweigh drug costs. Also, the ability to administer an antagonist IM may be advantageous in field studies (Wallingford et al. 1996). We recommend researchers investigate the utility of reversing the less costly and popular Telazol®/xylazine HCl and ketamine HCl/xylazine HCl combinations with an IM injection of atipame- zole HCl. The combination of medetomidine HCl, ketamine HCl, and atipamezole HCl reported herein provided an effective level of anesthesia for Clover-trapped White-tailed Deer. We recommend the use of 0.055 mg/kg medetomidine and 2.5 mg/kg ketamine HCl to immobilize adult Clover-trapped White-tailed Deer and 0.06 mg/kg medetomidine HCl and 2.5 mg/kg ketamine HC] for subadults. Furthermore, we recom- mend 0.275 mg/kg and 0.3 mg/kg of atipamezole, injected IM, to reverse this combination in adults and subadults, respectively. Acknowledgments We thank R. Houf, T. Mong, and numerous volun- teers for their assistance in the field. Missouri Depart- ment of Conservation, Conservation Research Center, provided Clover traps. A University of Missouri Res- earch Board grant, the University of Missouri, and the Missouri Department of Conservation supported this project. Literature Cited Arnemo, J. M. 1995. 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International Wildlife Veterinary Services, Inc., Laramie, Wyoming. 342 pages. Kreeger, T. J., G. D. DelGiudice, U. S. Seal, and P. D. Karns. 1986. Immobilization of white-tailed deer with xylazine hydrochloride and ketamine hydrochloride and antagonism by tolazoline hydrochloride. Journal of Wild- life Diseases 22: 407-412. Mautz, W. W., and J. Fair. 1980. Energy expenditure and heart rate for activities of white-tailed deer. Journal of Wildlife Management 44: 333-342. McCullough, D. R. 1975. Modification of the Clover Deer Trap. California Fish and Game 61: 242-244. Mech, L. D., G. D. DelGiudice, P. D. Karns, and U. S. Seal. 1985. Yohimbine hydrochloride as an antagonist to xyla- zine hydrochloride-ketamine hydrochloride immobiliza- tion of white-tailed deer. Journal of Wildlife Diseases 21: 405-410. Miller, M., M. Weber, D. Neiffer, B. Mangold, D. Fonte- not, and M. Stetter. 2003. Anesthetic induction of captive tigers (Panthera tigris) using a medetomidine-ketamine combination. Journal of Zoo and Wildlife Medicine 34: 307-308. Moen, A. N. 1978. Seasonal changes in heart rates, activity, metabolism, and forage intake of white-tailed deer. Journal of Wildlife Management 42: 715-738. Nielsen, L. 1982. Electronic ground tracking of white- tailed deer chemically immobilized with a combination of etorphine and xylazine hydrochloride. Pages 165-174 in Chemical immobilization of North American wildlife. Edited by L. Nielsen, J. C. Haigh, and M. E. Fowler. Wisconsin Humane Society, Inc., Milwaukee, Wisconsin. 447 pages. Presidente, P. J. A., J. H. Lumsden, D. R. Presnell, W. A. Rapley, and B. M. McGraw. 1973. Combination of etor- phine and xylazine in captive white-tailed deer: I. Effects of hematologic, serum biochemical and blood gas values. Journal of Wildlife Diseases 9: 342-348. Presnell, K. R., P. J. A. Presidente, and W. A. Rapley. 1973. Combination of etorphine and xylazine in captive white- tailed deer: I. Sedative and immobilization properties. Journal of Wildlife Diseases 9: 336-341. 190 Rogers, L. L., A. N. Moen, and M. L. Shedd. 1987. Rectal temperatures of 2 free-ranging white-tailed deer fawns. Journal of Wildlife Management 51: 59-62. Ryeng, K. A., J. M. Arnemo, and S. Larsen. 2001. Determination of optimal immobilizing doses of a mede- tomidine hydrochloride and ketamine hydrochloride com- bination in captive reindeer. American Journal of Veter- inary Research 62: 119-126. Ryeng, K. A., S. Larsen, and J. M. Arnemo. 2002. Mede- tomidine-ketamine in reindeer (Rangifer tarandus taran- dus): Effective immobilization by hand- and dart-admin- istered injection. Journal of Zoo and Wildlife Medicine 33: 397-400. Schultz, S. R., M. K. Johnson, and W. A. Forbes. 1992. Immobilization of captive white-tailed deer with mixtures of Telazol® and Rompun. Proceedings of the Annual Con- ference of Southeastern Association of Fish and Wildlife Agencies 46: 29-36. THE CANADIAN FIELD-NATURALIST Vol. 118 Tsuruga, H., S. Masatsugu, T. Hiroshi, J. Kiyoe, and K. Koichi. 1999. Immobilization of sika deer with medeto- midine and ketamine, and antagonism by atipamezole. Journal of Wildlife Diseases 35: 774-778. Wallingford, B. D., R. A. Lancia, and E. C. Soutiere. 1996. Antagonism of xylazine in white-tailed deer with intramus- cular injection of yohimbine. Journal of Wildlife Diseases 32: 399-402. Wesson, J. A., P. F. Scanlon, and R. E. Mirarchi. 1974. Immobilization of white-tailed deer with succinylcholine chloride: success rate, reactions of deer and some physio- logical effects. Proceedings of the Annual Conference of the Southeastern Association of Game and Fish Commis- sion 28: 500-506. Received 2 March 2002 Accepted 3 October 2004 reer Predation on Two Mule Deer, Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the Southern Canadian Rocky Mountains Dore Poszic!, CLAYTON D. Apps?, and ALAN Disb? ' Alhardstr. 15, 28757 Bremen, Germany ? Aspen Wildlife Research Inc., 2708 Cochrane Road N.W., Calgary, Alberta T2M 4H9 Canada (corresponding author) 3 Parks Canada, P. O. Box 220, Radium Hot Springs, British Columbia VOA 1MO0 Canada Poszig, Dérte, Clayton D. Apps, and Alan Dibb. 2004. Predation on Two Mule Deer, Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the southern Canadian Rocky Mountains. Canadian Field-Naturalist 118(2): 191-194. A male Canada Lynx (Lynx canadensis) killed two Mule Deer (Odocoileus hemionus) in the southern Canadian Rocky Mountains in January 1999 and made use of the kills for 28 days. Canada Lynx predation on ungulates has been reported but is rare, and accounts have been brief. We detail the lynx behaviour associated with the kills and their consumption. An infra- red monitor and attached camera were used to register daily activity at the kill site. We speculate on the factors that may have influenced this opportunistic predation event. Key Words: Canada Lynx, Lynx canadensis, infrared monitor, Mule Deer, Odocoileus hemionus, predation, British Columbia. The Canada Lynx (Lynx canadensis), hereafter referred to as lynx, preys mainly on Snowshoe Hares (Lepus americanus) and other small mammals, but rare accounts of ungulate predation have been noted (e.g., Saunders 1963; van Zyll de Jong 1966; Parker et al. 1983; Stephenson et al. 1991). However, these accounts do not describe lynx behaviour associated with the kills or their consumption. We give a detailed description of two Mule Deer (Odocoileus hemionus) kills made by a lynx and its subsequent use of them for 28 days. The observations were made in the course of an in-depth field study of lynx ecology in the south- ern Canadian Rocky Mountains of southeastern British Columbia and southwestern Alberta. While snow-tracking on 18 January 1999, we found two Mule Deer that had been killed by a radio-col- lared adult male Canada Lynx in Kootenay National Park, British Columbia (51° N, 116° W). One was a doe aged by cementum annuli to be 3.5 years, and we estimated the other deer to be a fawn (teeth not com- pletely erupted). The kills were separated by a distance of 10 m and were located at 1580 m elevation on a 15° slope of northwest aspect in a closed-canopy forest of Lodgepole Pine (Pinus contorta), Engelmann Spruce (Picea engelmannii), and Subalpine Fir (Abies lasio- carpa). The understory was sparse, although the imme- diate area around the kill site had some woody debris to a depth of about | m and some conifer thickets. The snow depth at the site was 90 cm and lynx penetration was 9-14 cm. The fawn apparently was killed on 14-15 January, judging from daily radio-telemetry fixes of the lynx and accumulated snow cover. The shoulders, rib cage, part of the neck, upper front legs and organs had been consumed. The front and rear had been severed and lay twisted and partially cached in a hollow in the snow. The doe was killed on 18 January between about 1300 and 1500 h. On initial inspection, no snow had accu- mulated on the carcass, and the blood had not yet co- agulated or frozen. The deer was found lying on its side with legs extended, and no part had yet been consumed. Teeth of both deer were in good condition, and no deformities of their legs or hooves were apparent. The doe appeared to be in good physical condition. Both deer had major wounds on the dorsal side of their necks, and the spacing of obvious tooth-punctures matched that of a lynx (Figure 1). The cause of death of the fawn could not be determined because it had been con- sumed to a great extent and was not intact. On the doe, several puncture wounds were located in two small areas on the dorsal to slightly lateral area of the neck, 10-15 cm behind the ears. Claw marks on the deer’s back and shoulders suggested that the lynx had leapt onto the deer and had been “riding” while biting it. Evidence of predation of the fawn was obscured by recent snow, but old blood stains 2 m from the carcass suggested that it had been killed and not scavenged. When removing the mandible of the fawn, we found pieces of undigested Subalpine Fir in its mouth, sug- gesting that the animal had been browsing shortly before it died. Tracks in the snow provided a clear record of how the lynx encountered and stalked the doe. The lynx approached this deer from a slightly higher elevation than where both kills were found, partially circling the site for about 100 m. It briefly stopped at the edge of a small opening about 50 m from the kill site, and then again 25 m uphill on a knoll. Moving downhill in a normal gait, the lynx used several fallen logs for cover, but was clearly stalking for the last 10 m. It then made three bounds before its attack, as the doe stood next to a thicket of mixed-age trees. The deer appeared to succumb <4 m from the attack site. The lynx then dragged the carcass about 7 m to its final location. LOM 192 THE CANADIAN FIELD-NATURALIST Vol. 118 FiGure 1. Neck wound (arrows indicate puncture locations) resulting in the death of an adult female Mule Deer by a Canada Lynx in the southern Canadian Rocky Mountains, January 1999. Six site visits conducted over the three weeks fol- lowing the initial find indicated that the lynx first fed on the doe’s neck, continued to feed on the shoulders and rib cage, and finally ate the rear and internal organs. Deer hair had been removed and was positioned around and on top of the kill. There were several well-used lynx beds that appeared to be strategically located upslope, <120 m from the kill site, with clear down- hill views. The hindquarters of the fawn were moved to 4 different locations between our site visits, and each time were cached under snow and branches. It is possible that this was prompted by our inspections. On 29 January we installed an infrared monitor (Trailmaster™ 1500, Goodson and Associates Inc., Kansas, with attached Olympus camera) at the doe’s kill site to document the 24-hour activity of the lynx at the site. The monitor was programmed to register all beam interferences, while the remote camera was set to take a photo upon beam interference, with <1 photo/15 min and between 0900 h and 1630 h to avoid flash disturbance. Because the lynx no longer appeared to be feeding on the fawn, we expected that all subse- quent feeding events would be registered. Twenty photos of the lynx were obtained and showed that only this animal fed on the carcass (e.g., Figure 2). In addition, no other predator was detected at the site during inspections that occurred about every four days. lynx activity at the kill peaked around mid- night and in the late morning (Figure 3). The lynx abandoned the kill site after 28 days, at which time the head and lower side of the doe’s car- cass were frozen into the ground. Scratch marks sug- gested that the lynx had unsuccessfully tried to remove some snow to access parts of the deer. Upon incidental recapture 16 days later, the lynx weighed 15.9 kg, 18.6% more than at his previous and original capture in November 1996, while skeletal measurements had not changed (C. D. Apps, unpublished data). Our account is consistent with the observations by Stephenson et al. (1991) of lynx ambushing ungulate prey at close range. They reported a lynx remaining on a Caribou (Rangifer tarandus) kill for 42 days. In our study, the Lynx apparently abandoned the site be- cause it could not make use of the carcass remains, and it might have stayed longer had conditions permitted. Previous reports of lynx predation on ungulates have been associated with the low phase of the approx- imate 10-year population cycle of Snowshoe Hares, the lynx’s primary prey. The reliance of lynx on alter- nate prey is greatest at this time (Mowat et al. 2000). Although the deer kills we report occurred during an assumed Snowshoe Hare population peak, hare densi- ties in the study area were comparable to boreal regions during the cyclic low (Apps 2000; C.D. Apps, unpub- lished data). 2004 Poszic, APPS, AND DIBB: PREDATION ON MULE DEER By A LYNX 193 = itl FIGURE 2. Remote camera photo of a Canada Lynx at the carcass of a Mule Deer it killed in the southern Canadian Rocky Mountains, January 1999. 80 10 a Frequency of Monitor interference ~9 70 . =—@—Frequency of Days with > 0 Monitor Interferences 8 60 Uf 50 6 3 g S 40+ 5 s S a we 4 30 3 20 2 10 00:00- 02:00- 04:00- 06:00- 08:00- 10:00- 12:00- 14:00- 16:00- 18:00- 20:00- 22:00- 01:59 03:59 05:59 07:59 09:59 11:59 13:59 15:59 17:59 19:59 21:59 23:59 Time of day (h) Ficure 3. Lynx activity while feeding on a Mule Deer kill over 11 days in the southern Canadian Rocky Mountains, January 1999. Total frequency (1 min. intervals; n = 344) and number of days that the Canada Lynx triggered an infrared monitor (monitor interference) per 2-h period are shown. 194 We further note that the relatively high elevation and deep snow of the kill site made it an atypical loca- tion for Mule Deer during mid-winter. We speculate that the deer may have been avoiding other predators, most notably Gray Wolves (Canis lupus) that travel primarily through the valley bottom. Thus, we expect that this predation was opportunistic, perhaps influ- enced by relatively low densities of Snowshoe Hares, the age class and sex of the lynx (adult male), and the vulnerability of the deer. Acknowledgments The Southern Canadian Rockies Lynx Project was jointly funded by the Habitat Conservation Trust Fund through the British Columbia Ministry of Water, Land and Air Protection (formerly Environment, Lands and Parks), Parks Canada, and the Columbia Basin Fish and Wildlife Compensation Program. Additional financial contributions were provided by the Friends of Kootenay National Park, Husky Oil, World Wild- life Fund Canada, Fairmont Hotels and Resorts (for- merly Canadian Pacific Hotels & Resorts), and the Bow Valley Naturalists. Contract supervision, input, and direction were provided by A. Dibb of Parks Can- ada, J. Krebs of the Columbia Basin Fish and Wildlife Compensation Program, and A. Fontana, R. Neil, G. Tipper, R. Forbes, and I. Teske of BC Environment. THE CANADIAN FIELD-NATURALIST Vol. 118 Literature Cited Apps, C. D. 2000. Space use, diet, demographics, and topo- graphic associations of Lynx in the southern Canadian Rocky Mountains. Pages 351-371 in Ecology and conser- vation of Lynx in the United States. Edited by L. F. Rug- giero, K. B. Aubry, S. W. Buskirk, G. M. Koehler, C. J. Krebs, K. S. McKelvey, and J. R. Squires. University Press of Colorado, Boulder, Colorado. Mowat, G., K. G. Poole, and M. O’Donoghue. 2000. Ecology of Lynx in northern Canada and Alaska. Pages 365-306 in Ecology and conservation of Lynx in the United States. Edited by L. F. Ruggiero, K. B. Aubry, S. W. Buskirk, G. M. Koehler, C. J. Krebs, K. S. McKelvey, and J. R. Squires. University Press of Colorado, Boulder, Colorado. Parker, G. R., J. W. Maxwell, L. D. Morton, and G. E. J. Smith. 1983. The ecology of the Lynx (Lynx canadensis) on Cape Breton Island. Canadian Journal of Zoology 61: 770-786. Saunders, J. K. Jr. 1963. Food habits of the Lynx in New- foundland. Journal of Wildlife Management 27: 384-390. Stephenson, R. O., D. V. Grangaard, and J. Burch. 1991. Lynx, Felis lynx, predation on Red Foxes, Vulpes vulpes, Caribou, Rangifer tarandus, and Dall Sheep, Ovis dalli, in Alaska. Canadian Field-Naturalist 105: 255-262. van Zyll de Jong, C. G. 1966. Food habits of the Lynx in Alberta and the Mackenzie District, N.W.T. Canadian Field-Naturalist 80: 18-23. Received 8 September 2002 Accepted 25 October 2004 Effect of Fire Intensity and Depth of Burn on Lowbush Blueberry, Vaccinium angustifolium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, Production in Eastern Ontario Luc C. DUCHESNE! and SUZANNE WETZEL2? 'Rorest BioProducts Inc., 945 Queen Street East, Sault Ste Marie, Ontario P6A 2E5 Canada >Natural Resources Canada, Canadian Forestry Service, 1219 Queen Street East, Sault Ste Marie, Ontario P6A 3A5 Canada 3Corresponding author at swetzel@NRCan.gc.ca Duchesne, Luc C., and Suzanne Wetzel. 2004. Effect of fire intensity and depth of burn on Lowbush Blueberry, Vaccinium angusti- folium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, production in eastern Ontario. Canadian Field-Naturalist 118(2): 195-200. The effects of prescribed fire intensity and depth of burn were investigated on Lowbush Blueberry (Vaccinium angustifolium) and Velvet Leaf Blueberry (Vaccinium myrtilloides) stem density, blueberry production and the number of blueberries/stem in a clear-cut Jack Pine, Pinus banksiana, ecosystem of eastern Ontario. Blueberry production and stem density were significantly (P < 0.001) increased by low intensity prescribed fires of 597 and 1268 kW/m. In contrast, prescribed fires of medium and high intensities did not affect blueberry production and stem density. The number of blueberries/stem was not affected (P = 0.056) by prescribed burning, two years after treatment. Pearson’s multiple correlation analysis showed that blueberry production (R: -0.683, P < 0.01), stem density (R: 0.733, P < 0.01) and the number of blueberries/stem (R: 0.803, P < 0.01) correlated with depth of burn. As well, blueberry production (R: 0.507, P < 0.05) and stem density (R: -0.504, P < 0.05) correlated with fire intensity. Depth of burn was a better predictor of berry production and stem density than fire intensity. These results suggest that only low intensity fires with little penetrating effect in the ground should be used to manage blueberry crops. Key Words: Lowbush blueberry, Vaccinium augustifolium, Vaccinium myrtilloides, wildfire, nontimber forest products (NTFP). Nontimber forest products (NTFP) are all botanical commodities harvested from the forest excluding in- dustrial timber use (Duchesne et al. 2000). NTFP are critical to the economy of forest and rural communi- ties by providing food as well as supplemental income, which often improves the standard of living of rural and First Nation communities (Brubaker 1997; Du- chesne et al. 2001; Mohammed 1999). Despite the socio-economic importance of this industry there has been little scientific research conducted for its support, presumably because of its novelty. However, the long- term success of the NTFP industry depends on acquir- ing precise knowledge regarding the availability, and sustainability of wild harvests as well as strategies to domesticate and manage many of the most sought after NTFP species. Species such as Lowbush Blueberry (Vaccinium angustifolium) and Velvet Leaf Blueberry (Vaccinium myrtilloides) need to be further domesti- cated and/or managed in natural ecosystems to meet the growing demand from the food and nutraceutical industries (Duchesne et al. 2001). This paper describes research conducted to improve blueberry management. Canada is the world’s largest producer of wild blue- berries (Lowbush and Velvet Leaf blueberries) with an annual output of $42 425 million in farm gate value resulting in production of 43 511 tonnes of frozen blueberries (Agriculture and Agri-Food Canada 2002). The demand for Lowbush and Velvet Leaf blueberries is such that management of natural or established sites is conducted to promote optimal yields ranging from 3360 to 8967 kg/ha (Mohammed 1999). Given that blueberries are worth approximately $1.00/kg to farm- ers, the annual output of managed fields ranges from $3360 to $8967/ha, which is far superior to the aver- age timber growth of Canada’s forests of 1.59 m°/ha/ year (Lowe et al. 1996) with an approximate value of $100/ha/year. Pruning of blueberry fields, either by mowing or burning, is used routinely to maximize blueberry pro- duction (Badcock 1958; Eaton 1958; Eaton and White 1960) as it stimulates the development of new shoots from shallow rhizomes (Van Hoefs and Shay 1981). In turn, new shoots are preferred over old shoots as the ratio of flower buds to leaf buds is greater on new shoots than on older shoots (Hall et al. 1972). Fire pruning of blueberry fields is generally accomplished by using straw as a fire carrier or tractor mounted oil or propane gas burners that emit a constant flame (Blatt et al. 1989). Burning also has the added advantage of reducing problems with insects and disease (Blatt et al. 1989; Smith and Hilton 1971). Although fire is frequently used for the management of Lowbush and Velvet Leaf blueberries (Blatt et al. 1989), which form one commercial crop, there is a lack of information about the relationship between fire behaviour and its impact on berry production. Indeed, current prescrip- tions aim at using fires that kill all stems (Blatt et al. 1989). However, fire behaviour varies a great deal even during prescribed burns, which in turn affects the eco- logical outcome of post-fire ecosystems (McRae et al. 2001). 195 196 Frontal fire intensity (or Byram’s fireline intensity) is a measurement of the quantity of energy liberated as a fireline moves through an ecosystem, and is the most encompassing physical measurement of fire behavior related to ecological impacts (reviewed by Alexander 1982 and McRae et al. 2001). Fire intensity is some- times used together with depth of burn (Miller 1977), which is a measurement of heat penetration into the soil. However, both frontal fire intensity and depth of burn are relatively new concepts in fire ecology and have never been applied to blueberry management. Hence, the objective of this study was to investigate the effect of fires of different intensities and depth of burn on Lowbush and Velvet Leaf blueberry pro- duction and vegetative growth. Materials and Methods Study site The study area is located at Frontier Lake (46°00’N, 77°33’ W) in a Jack Pine, Pinus banksiana, stand in eastern Ontario within the middle Ottawa section (L.4c) of the Great Lakes — St. Lawrence Forest region (Rowe 1972). As reported by Herr et al. (1994), the site is near the Petawawa Research Forest, and is rela- tively flat, with a difference in elevation of approx- imately 4 m over 1.0 km. The surface deposit is a fine-grained and deep sand (10-30 m deep) (Gadd 1962) and the soil is a humo-ferric podzol. The study site was selected because of its uniformity in tree com- position and topography. It was harvested in 1942 and 1943 leaving residual standing timber with a stump diameter of 17.5 cm or less. Dendrochronological analysis of dominant trees and snags with multiple fire scars suggests that the study site sustained sev- eral fires, with the most recent in 1943, presumably from broadcast slash burning following harvesting (E. Stechishen, personal communication). THE CANADIAN FIELD-NATURALIST Vol. 118 Presently, the stand comprises a mix of Jack Pine (Pinus banksiana), Red Pine (Pinus resinosa) and White Pine (Pinus strobus) with the Red and White pine forming an emergent layer (Herr et al. 1994). Although the Jack and Red pine are of similar ages, the Jack Pine has higher relative density and is the dominant tree species on the site. Other plant species found at the site include serviceberry (Amelanchier sp.), Sweet Fern (Comptonia peregrina), Wintergreen (Gaultheria procumbens), Sheep Laurel (Kalmia an- gustifolia), Ground Pine (Lycopodium complanatum), Cow Parsnip (Maianthemum canadense), Hairy Solo- mon’s Seal (Polygonatum pubescens), Bracken Fern (Pteridium aquilinum ) and Sand Cherry (Prunus pumila). Treatments In the summer of 1990, an area of 150 m x 1000 m was clear-cut with the slash left in place. The site was divided into 40 plots of 35 m x 70 m with 8 m fire- guards established around each plot. In 1991, ten plots were burned under different indices of the Canadian Forest Fire Weather Index (FWI) system (Table 1) resulting in frontal fire intensities that varied from 597 to 21 305 kW/m and depth of burn that varied from 0.37 to 2.78 cm (McAlpine 1995). This range of fire intensity and depth of burn far exceeds the range of fire behaviour currently used in blueberry management (unpublished observations). Pre- and post-burn fuel loads were measured for both slash and duff fuels (McRae et al. 1979). Fire rate of spread was measured with a pin grid network on each plot; fire arrival times at each pin were recorded to provide distance and time information. Fuel con- sumption was determined as the difference between these two values. Fire intensity was calculated using Byram’s (Byram 1959: cited in McAlpine 1995) TABLE |. Fire weather', fire behaviour parameters! and blueberry production from thirteen 35 m x 75 m plots in a clear cut Jack Pine after prescribed burning of different fire intensities at Frontier Lake Experimental site in Eastern Ontario. Plot Date Depth! of Intensity! Blueberries” Stem density* Blueberries/stem* Number of burn! FWI! — Burn (cm) (kW/m) (g/m?) (stems/m?) (N/plant) 2 08/08/91 3.1 0.37 597 492.86 a (167.4) 73.2 a (8.5) 7.53 a (3.4) 7 08/08/91 3.6 0.38 1 268 242.18 a (79.44) 44.2 b (9.8) 5.52 a (1.6) 44 06/24/91 15.8 1.79 2 305 10.81 b (7.6) 5.9 c (0.7) 2.26 a (1.8) 40 07/10/91 4.3 2.78 4 844 114 b, (©@:9) 9.4 c¢ (2.9) 0.90 a (0.69) 32 07/12/91 9.6 1.85 7 600 8.78 b (8.7) 6.7 c (5.6) 0.53 a (0.46) 26 06/14/91 11.0 3.50 10 941 4.96b (4.8) 9.9 ¢ (7.1) 3.40 a (2.9) 46 06/24/91 17.4 1.95 12 065 ZO Mb Gels) 4.0 c (1.9) 3.10 a (2.7) 36 07/12/91 3.9 Dail 13 202 27.20b (8.9) 12.1 c (4.6) 2.59 a (1.13) 43 07/10/91 8.8 2.64 20 334 0.53b (0.24) 5.7 c (4.1) 0.50 a (0.42) 24 06/14/91 21.0 1.79 21 305 5.33b (0.48) 13.8 c¢ (2.4) 0.04 a (0.06) Controls? — = — 9.01b (2.6) 5.9 c (1.48) 0.78 a (0.25) ' from McAlpine (1995). * within column means followed by a different letter are significantly different at P < 0.05 determined by ANOVA. Observed significance level was adjusted with the Bonferroni procedure. > controls consisted of three unburned plots adjacent to the burned over plots. 2004 intensity equation I=HwR, where I is the intensity of the fire (kW/m), H is the fuel low heat of combustion (assumed to be 18 000 kJ/kg), w is the weight of fuel consumed in the active front (kg/m?) (all fuel con- sumed was assumed to have been burned by the active fire front), and R is the rate of spread (m/sec). To determine depth of burn, ten 50 cm steel pins mounted with horizontal markers were used in each plot to mark the top of the soil litter layer prior to each burn. Immediately after the burn, the distance between the horizontal marker and the top of the litter layer was measured with a ruler and called depth of burn. Two years following prescribed burning, there were 45 plant species in our research plots. The most impor- tant species in terms the biomass and frequency (in decreasing order of importance) were P. aquilinum, V. angustifolium, V. myrtilloides, Amelanchier sp., P. pumila, Comptonia peregrina, Kalmia angustifolia, Yellow Panic-grass (Panicum xanthophysum), and Houton’s Sedge (Carex houghtonii) (Tellier et al. 1995, 1996). Blueberry collection To compare the effect of fire intensity and depth of burn, blueberries were collected two years after pre- scribed fire from the ten burned over plots and from three control plots. Controls consisted of three clear- cut 35 m X 70 m plots adjacent to the burned-over plots. Blueberries were not collected in subsequent years because of logistical reasons limiting access to the research plots. For this study, three 3 m x 3 m quadrats were ran- domly established in each of the 35 m x 70 m plots. The perimeter of each of the 3 m x 3 m quadrats was delineated with a string tied 40 cm above ground and all berries within the 3 m x 3 m quadrats were picked, advancing from the edge of the plot inwards. Ripe, over-ripe, green, and deformed blueberries were in- cluded in the harvest. All plots were harvested on 13 July and 14 July 1993. The harvested berries were kept in plastic coolers in the shade until the end of the col- lection day and their fresh weight determined in the laboratory the same evening. The bags were then placed in drying ovens at 50°C for 5 days before weighing. Because the moisture content of blueberries (83.1 + 0.8 %) was comparable among all plots (P = 0.22) and consistent with published data (Usui et al. 1994), the results were expressed in terms of fresh weight only. As well, no attempt was made to distinguish between V. myrtilloides and V. angustifolium as the frequency of these species (1:15- 1:30, myrtilloides: angustifoli- um) was not significantly different among treatments (S051): To compare the effect of fire intensity and depth of burn on the vegetative abundance of V. myrtilloides and V. angustifolium, the number of stems of both species was determined two years after prescribed fire from the ten burned over plots and three control plots. Once the berries had been removed from the 3 m x 3 m DUCHESNE and WETZEL: EFFECT OF FIRE INTENSITY AND DEPTH OF BURN ON BLUEBERRY 197 quadrats, the number of stems was recorded for the entire quadrat and expressed as stems/m/?. As well, the number of blueberries/stem was com- pared among the prescribed burns and the control. For this, 100 stems were selected randomly outside the 3 m x 3 m quadrats and the number of blueberries determined for each stem. This protocol was repeated in each of the ten prescribed burn plots as well as each of the three control plots. Statistical analyses Analysis of variance (ANOVA) was conducted to compare blueberry production, stem density and blue- berry abundance/stem among the prescribed burns and the controls (Systat 1997). For statistical analyses the three control plots were pooled as one treatment whereas each of the ten burned over plots was treated as a separate treatment (McAlpine 1995). Means were compared using the Bonferroni procedure (Systat 1997). Pearson’s multiple correlation analysis was con- ducted to determine the relationship between frontal fire intensity, depth of burn, berry production and the number of blueberries/stem (Systat 1997). Results Blueberry production (g/m?) differed significantly (P < 0.001) among treatments. The greatest blueberry production was observed in prescribed fires of 597 and 1268 kW/m whereas there were no significant differences among blueberry production of the other treatments, including controls (Table 1). Likewise stem density/ha differed among treatments (P < 0.001) and was greatest in the fires of 597 and 1268 kW/m. There was no difference in the number of blueberries/plant among the other treatments including the controls (P = 0.056). Pearson’s multiple correlation analysis showed that fire intensity correlated (P < 0.05) with blueberry pro- duction, and stem density (Table 2). Depth of burn correlated (P < 0.01) with blueberry production, stem density and blueberries/plant. Depth of burn showed better correlations with blueberry production, number of blueberries/plant and stem density than fire inten- sity (Table 2). Discussion Several studies have demonstrated that the response of ecosystems varies greatly with frontal fire intensity and depth of burn. Fire intensity has been shown to correlate with P. banksiana regeneration (Weber et al. 1987) and post-fire abundance of P. aquilinum (Tellier et al. 1995). Low fire intensities stimulated Corylus cornuta sprouting and height growth of Rubus ideaus (Johnston and Woodard 1985). Biomass nutrient reten- tion (Duchesne and Tellier 1997) and soil seed bank and competing vegetation dynamics along with P. resinosa and P. strobus seedling performance are also affected by prescribed burning intensity (Tellier et al. 1995, 1996; Whittle et al. 1997). Soil microbial activity and 198 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Pearson’s multiple correlation analysis among fire intensity, depth of burn, blueberry biomass, number of blueberries/ stem, and stem density. Intensity Blueberries/ha Intensity 1.000 Blueberries/ha -0.507* 1.000 Depth of burn 0.517% -0.683** Blueberries/stem -0.461 0.803** Stem density -0.504* 0.766** * significant at P < 0.05 ** significant at P < 0.01 diversity were also affected by fire intensity (Duchesne and Wetzel 2000; Staddon et al. 1998a, 1998b). Fire has been used for hundreds of years in blue- berry management by Indigenous people (Chapeskie 2001) and by farmers (Badcock 1958; Blatt et al. 1989), mostly using spring fires. This investigation presents the first comprehensive data on the effect of fire intensity and depth of burn on natural blueberry production, which are indicated as critical factors to consider in blueberry management. Indeed, the current findings show that fire intensity must be kept low in order to increase blueberry productivity using pre- scribed fire. As well, care must also be given to reduce depth of burn, which has a greater impact on blueberry production than fire intensity (Table 2). Whereas fire intensity is a measure of the caloric energy liberated by the fire front (Alexander 1982), depth of burn integrates the interaction between soil moisture conditions and fire intensity (Miller 1977) and is more closely linked to the response of blueberries to fire than fire intensity. Since blueberry shoots arise from shallow rhizomes and portions of the above ground stem not killed by the fire (Flinn and Wein 1977; Martin 1955: Minore 1975), deep heat penetration in the humus layers is likely to reduce stem production. Similar conclusions have been reached in Ponderosa Pine (Pinus ponder- osa) forests where depth of burn influenced greatly understory species regeneration (Armour et al. 1984) which is consistent with the concept that depth of buried propagules, together with heat penetration, is critical in post-fire survival (reviewed by Whittle et al. 1997). In turn, the differences observed in this inves- tigation in post-fire blueberry stem densities, and the negative correlation between stem density and depth of burn may be ascribed to the deleterious effect of fire on the underground rhizomes as was demonstrated on Blue Huckleberry (Vaccinium globulare) (Miller L977): In practice, blueberry production can be maximized by burning in the spring of every second year, presum- ably to maintain a high ratio of flower buds to leaf buds (Hall et al. 1972). In this investigation, the number of blueberries/plant, which was hypothesized to be an Depth of burn Blueberries/stem Stem density 1.000 =O ei 1.000 -0.724** 0.471 1.000 estimate of the flower bud to leaf bud ratio, was not significantly different among treatments but inversely correlated with fire intensity. These results cannot be explained by conventional thinking regarding flower bud formation (Hall et al. 1972), as the number of blue- berries/plant correlated with depth of burn. However, Smith and Hilton (1971) speculated that Lowbush Blueberry performance after fire pruning might result from the stimulative effects of the nutrients in ash deposited on the surface soil during burning. Further investigation is needed to assert the relevance of this hypothesis in blueberry flower biology. Although this investigation shows that low intensity fires have a positive influence on blueberry production, natural blueberry ecosystems support a variety of wild- fire characteristics (Moola et al. 1998; Usui et al. 1995). Hence the continuous effect of repeated fires should be investigated on the long term productivity and biologi- cal conservation of such ecosystems. In particular, it will be important to assess the impact of burning every second year, which is the recommended practice, on soil nutrients (Smith and Hilton 1971). Future work should also be conducted to investigate co-manage- ment of blueberries with timber values (Duchesne et al. 2001). Documents Cited (marked * in text) Agriculture and Agri-Food Canada. 2002. Canadian fruit situation and trends. http://www.agr.gc.ca/misb/hort. Literature Cited Alexander, M. E. 1982. Calculating and interpreting forest fire intensities. Canadian Journal of Botany 60: 349-357. Armour, C. D., Sc. C. Bunting, and L. F. Neuenschwander. 1984. Fire intensity effects on the understory in ponderosa pine forests. Journal of Range Management 37: 44-49. Badcock, A. C. 1958. Prescribed burning and the growing of blueberries. Pages 53-56 in Symposium on Prescribed Burning in Forestry, Agriculture and Wildlife Management. Published by the Newfoundland Research Committee, St. John’s. Sponsored by The Newfoundland Department of Mines and Resources and Memorial University. Blatt, C. R., I. V. Hall, K. I. N. Jensen, W. T. A. Neilson, P. D. Hildebrand, N. L. Nickerson, R. K. Prange, P. D. Lidster, L. Crozier, and J. D. Silbey. 1989. Lowbush blue- 2004 berry production. Agriculture Canada Publication 1477/E. 56 pages. Brubaker, D. 1997. Value-added forestry and aboriginal com- munities: the perfect fit. National Aboriginal Forestry Association, Ottawa, Ontario. 114 pages. Chapeskie, A. J. 2001. Northern homelands, northern fron- tier: linking culture and economic security in contemporary livelihoods in boreal and cold temperate forest communities in northern Canada. Pages 31-44 in Forest communities in the third millennium: linking research, business, and policy toward a sustainable non-timber forest product sector. Edited by I. Davidson-Hunt, L. C. Duchesne and J. C. Zasada. USDA Forest Service, North Central Re- search Station. General Technical Report NC-217. Duchesne, L. C., J. C. Zasada, and I. Davidson-Hunt. 2000. Nontimber forest products in Canada: Scope and research needs. Forestry Chronical 76: 743-746. Duchesne, L. C., I. Davidson-Hunt, and J. Zasada. 2001. The future of the non timber forest products industry. Pages 149-152 in Forest Communities in the Third Mil- lennium: First international conference on non-timber forest products in cold temperate and boreal forests. Kenora Ont. Oct 1-4, 1999. Edited by I. Davidson-Hunt, L. C. Duchesne, and J. C. Zasada . USDA Forest Service, North Central Research Station, General Technical Report NC-217. Duchesne, L. C., and R. Tellier. 1997. Effect of prescribed burning intensity and disk trenching scarification on the nutrient status of aboveground non-crop vegetation in a Jack Pine (Pinus banksiana Lamb.) clear-cut. Forestry Chronical 73: 711-714. Duchesne, L. C., and S. Wetzel. 2000. Effect of clear-cutting, prescribed burning and scarification on litter decomposi- tion in an Eastern Ontario jack pine (Pinus banksiana) eco- system. International Journal of Wildland Fire 9: 195-201. Eaton, E. L. 1958. Blueberry burning and botanical relation- ship. Pages 57-58 in Symposium on Prescribed Burning in Forestry, Agriculature and Wildlife Management. Pub- lished by the Newfoundland Research Committee, St. John’s. Sponsored by the Department of Mines and Resources and Memorial University. Eaton, L., and R. G. White. 1960. The relation between burning dates and the development of sprouts and flower buds in the lowbush blueberry. American Society of Hor- ticultural Science 76: 338-342. Flinn, M. A., and R. W. Wein. 1977. Depth of underground plant organs and theoretical survival during fire. Canadian Journal of Botany 55: 2550-2554. Gadd, N. R. 1962. Surficial geology — Chalk river, descriptive notes. Marginal notes to map 1131A, Canadian Geological Survey Committee, Ottawa, Ontario. Hall, I. V., L. E. Aalders, L. P. Jackson, G. W. Wood, and C. L. Lockhart. 1972. Lowbush blueberry production in Canada. Canadian Department of Agriculture Publication 1477. Herr, D. G., L. C. Duchesne, R. Tellier, R. S. McAlpine, and R. L. Peterson. 1994. Effect of prescribed burning on the ectomycorrhizal infectivity of a forest soil. Interna- tional Journal of Wildland Fire 4: 95-102. Johnston, M., and P. Woodard. 1985. The effect of fire sever- ity level on postfire recovery of hazel and raspberry in east- Central Alberta. Canadian Journal of Botany 63: 672-677. Lowe, J. J., K. Power, and S. L. Gray. 1996. Canada’s forest inventory 1991: the 1994 version. An addendum to Canada’s forest inventory, Natural Resources Canada, DUCHESNE and WETZEL: EFFECT OF FIRE INTENSITY AND DEPTH OF BURN ON BLUEBERRY 199 Canadian Forest Service, Pacific Forestry Centre, Victor- ia, British Columbia. Information Report. BC-X-362E. Martin J. L. 1955. Observations on the origin and early dev- elopment of a plant community following a forest fire. Forestry Chronicle 31: 154-160. McAlpine, R. S . 1995. Testing the effect of fuel consump- tion on fire spread rate. International Journal of Wildland Fire 5: 143-152. McRae, D. J., M. E. Alexander, and B. J. Stocks. 1979. Measurement and description of fuels and fire behavior on prescribed burns: A handbook. Department Environment, Canadian Forest Service, Sault Ste Marie, Ontario. Infor- mation Report O-X-287. McRae, D. J., L. C. Duchesne, B. Freedman, T. J. Lynd- ham, and S. Woodley. 2001. Comparisons between wild- fire and forest harvesting and their implications in forest management. Environmental Review, 9: 223-260. Miller, M. 1977. Response of blue huckleberry to pre- scribed fires in a western Montana Larch-Fir forest. USDA Forest Service Research Paper, INT-188, Intermontane Forest and Range Experimental Station, Ogden, Utah. 33 pages. Minore D. 1975. Observations on the rhizomes and roots of Vaccinium membranaceum. Pacific Northwest Forest and Range Experimental Station. Research Note PNW-261, USDA. 5 pages. Mohammed, G. H. 1999. Nontimber forest products in Ontario: an overview. Ontario Forest Research Institute, Ontario Ministry of Natural Resources. Forest Research Information Paper Number 145. Moola, F. M., A. U. Mallik, and R. A. Lautenschlager. 1998. Effects of conifer release treatments on the growth and fruit production of Vaccinium spp. in northwestern Ontario. Canadian Journal of Forest Research 28: 841-856. Rowe, J. S. 1972. Forest regions of Canada. Canadian For- estry Service Publication Number 1300. Smith, D. W., and R. J. Hilton. 1971. The comparative effects of pruning by burning or clipping lowbush blueberries in northeastern Ontario. Journal of Applied Ecology 8: 781- 789 Staddon, W. J., L. C. Duchesne, and J. T. Trevors. 1998a. Impact of clear-cutting and prescribed burning on microbial diversity and community structure in a Jack Pine (Pinus banksiana Lamb.) clear-cut. World Journal of Microbiol- ogy and Biotechnology 14: 119-123. Staddon, W. J., L. C. Duchesne, and J. T. Trevors. 1998b. Acid phosphatase, alkaline phosphatase and arylsulphatase activities in a jack pine ecosystem after clear-cutting, pre- scribed burning, and scarification. Biology and Fertility of Soils 27: 1-4. Systat. 1997. Systat 7.0. Evanston, Ilinois. Tellier, R., L. C. Duchesne, J. C. Ruel, and R. S. McAlpine. 1995. Effets de l’intensité du brilage dirigé et de la scarifi- cation sur la diversité des especes végétales dans un peu- plement de pin gris (Pinus banksiana Lamb.). Ecoscience 2: 160-167. Tellier, R., L. C. Duchesne, J. C. Ruel, and R. S. McAlpine. 1996. Effets du brilage dirigé et du scarifiage sur |’ établis- sment des semis et sur leur interaction avec la végétation concurrente. Forestry Chronicle 71: 621-626. Usui, M., Y. Kakuda, and P. Kevan. 1995. Composition of energy values of wild fruits from the boreal forest of northern Ontario. Canadian Journal of Plant Science 74: 581-587. 200 Van Hoefs, M. E. G., and J. M. Shay. 1981. The effects of shade on shoot growth of Vaccinium angustifolium Ait after fire pruning in southwestern Manitoba. Canadian Journal of Botany 59: 166-174. Weber, M. G., M. Hummel, and C. E. Van Wagner. 1987. Selected parameters of fire behavior and Pinus banksiana Lamb. regeneration in eastern Ontario. Forestry Chronicle 63: 340-346. THE CANADIAN FIELD-NATURALIST Vol. 118 Whittle, C. A., L. C. Duchesne, and T. Needham. 1997. The impact of broadcast burning and fire severity on species composition and abundance of surface vegetation in a jack pine (Pinus banksiana) clear-cut. Forest Ecology and Man- agement 94: 141-148. Received 12 June 2002 Accepted 24 September 2004 Spring Dispersal Patterns of Red-winged Blackbirds, A gelaius phoeniceus, Staging in Eastern South Dakota H. J. Homan!, G. M. Linz!, R. M. ENGEMAN?, and L. B. PENRyY! 'U.S. Department of Agriculture, National Wildlife Research Center, 2110 Miriam Circle, Bismarck, North Dakota 58501- 2502 USA 2 U.S. Department of Agriculture, National Wildlife Research Center, 4101 LaPorte Avenue, Fort Collins, Colorado 80521 - 2154 USA Homan, H. J., G. M. Linz, R. M. Engeman, and L. B. Penry. 2004. Spring dispersal patterns of Red-winged Blackbirds, Agelaius phoeniceus, staging in eastern South Dakota. Canadian Field-Naturalist 118(2): 201-209. Red-winged Blackbirds (Agelaius phoeniceus) are very abundant summer residents throughout the Prairie Pothole Region of central North America. In late summer they assemble in post-breeding flocks that cause significant amounts of agricul- tural damage, particularly in sunflower fields near natal sites. In April 2001, we aerially color-marked ~370 000 Red-winged Blackbirds near Badger, South Dakota (44°48'N, 97°21'W), to determine if migrants staging here were summer residents in sunflower production areas ~350 km to the northwest. We measured patterns of migratory dispersal by collecting birds in 54 randomly selected blocks in the northcentral U.S. and the Prairie Provinces of Canada. The marked specimens (n = 33) were categorized into three polygons based on analyses of banding and re-sighting data and proximity to concentrated sunflower production. We estimated that 82% of the migrants that had staged in eastern South Dakota resided. within or on the periphery of the sunflower growing area. These birds probably stay near their breeding territories until at least late August and cause early damage to sunflower, which comprises the majority of damage. Resident birds in Alberta and most of Saskatchewan (18%) might arrive too late in the damage season to impact the sunflower crop significantly. Key Words: Agelaius phoeniceus, Red-winged Blackbird, breeding range, color-marking, dispersal patterns, northern Great Plains, spring migration, sunflower damage. Several million hectares of wetlands have been drained in the Prairie Pothole Region (Dahl 1990*), but high densities of semipermanent wetlands remain in North Dakota (0.88/km?, Reynolds et al. 1997*). These are preferred nesting habitat of Red-winged Black- birds (Agelaius phoeniceus) and make North Dakota the center of abundance for this species. During the mid-1990s, the breeding population in North Dakota started gaining rapidly in size. Red-winged Blackbird indices from the North American Breeding Bird Sur- vey (BBS) are at historical highs for the state (Figure 1). From 1994-2002 North Dakota’s average BBS index of all routes in the Prairie Pothole Region ranged from 199 to 250 Red-winged Blackbirds per route. North Dakota led all other U.S. states and Canadian provinces during this period in annual indices of Red- winged Blackbirds (Sauer et al. 2004*). Conflicts bet- ween wildlife and agriculture often occur because of local overabundance (Garrott et al. 1993; Conover 2002), and the growth in numbers of Red-winged Blackbirds has exacerbated a long-term struggle bet- ween sunflower producers and blackbirds (Linz and Homan 1998). Oilseed sunflower is a preferred food of Red-winged Blackbirds (Linz et al. 1984; Homan et al. 1994). It is also an important economic crop in North Dakota. In 2001, 75% of total oilseed sunflower production in North America came from North Dakota (National Agricultural Statistics Service 2004*). With increased depredation pressure on the crop caused by population expansion of Red-winged Blackbirds, many growers probably dropped sunflowers from their field rotations (Lamey and Luecke 1994). Since 1995, the number of sunflower fields has steeply declined in cen- tral and southeastern North Dakota, areas renowned for their high levels of sunflower production (National Agricultural Statistics Service 2004*). Statewide field damage surveys have that blackbird damage to sunflower ranges from $4 to 11 million (U.S. dollars) in the Prairie Pothole Region (Hothem et al. 1988). Based on current population estimates and bioenergetic models, Red-winged Blackbirds cause $3 million of the $5.5 million lost yearly to blackbirds in the core area of sunflower production (Peer et al. 2003). Most of the damage occurs between mid-August and early September, when the calorie content of im- mature sunflower achenes is low and the birds must eat more to reach satiation (Cummings et al. 1989; Con- ner and Hall 1997). During this early damage period, when >75% of total damage occurs, Red-winged Blackbirds account for 80% of the blackbird species observed in sunflower fields (Cummings et al. 1989). This period is before en masse blackbird migration, and the majority of these birds are of local origin. Most will remain within 200 km of their natal sites until molt completion (or near completion) in late August and early September (Dolbeer 1978; Besser et al. 1983; Linz et al. 1983). The damage can be quite serious in central and southeast North Dakota and northeast South Dakota, areas of concentrated production of sunflow- ers (National Agricultural Statistics Service 2004*) and 201 202 plentiful wetlands to attract nesting Red-winged Black- birds (~22 breeding pairs/km7, Linz et al. 2002). By late summer, the Red-winged Blackbird population responsible for most of the sunflower damage in the Prairie Pothole Region is about 39 (+8.8) million (Peer et al. 2003). Reducing excessive field damage (e.g., >10% per field) has proven an enormous challenge. The U.S. Department of Agriculture (USDA) uses an integrated management approach of lethal and nonlethal methods. Most of the available methods are used infrequently because of unmanageable logistics, little or no efficacy, or low cost:benefit ratios (Linz and Hanzel 1997). Mechanical frightening agents (e.g., scarecrows, pyro- technics, and propane cannons) help alleviate light predation pressure, but only work effectively if the dur- ation of the damage period is less than the habituation period (Cummings et al. 1986). Unfortunately, black- birds habituate quickly to frightening agents, particu- larly if the crop is a preferred food in an area of limited alternate foods (Ward 1979). The taste repellent, methyl anthranilate, is also used by producers, but the con- centration in the commercial product (BirdShield™) is below the known repellency threshold, despite the formulation’s high cost (Werner et al. 2005). Experi- ments with avicide were tried in sunflower fields with ongoing damage. Avicide-treated rice (DRC-1339) was placed in small plots opened in the fields. This approach failed because (1) blackbirds foraged almost exclusively on the standing heads of sunflower and would not go to the ground to forage in the plots, (2) precipitation necessitated numerous rebaitings, and (3) nontarget avian risks increased substantially over the baiting period because of migration (Schaaf 2003; Linz et al. 2000). Two nonlethal methods showing some promise are reduction of dense stands of cattail (Typha spp) in wetlands near sunflower (Linz et al. 2004a) and strategic placement of lure crops to attract foraging blackbirds away from sunflower fields (Linz et al. 2004). Cattail management has been a USDA program for over 10 years. The program is small com- pared to existing coverage of cattail (Ralston et al. 2004); and it also faces year-to-year logistical chal- lenges because of instability in roost locations across years combined with cattail’s ability to rapidly invade, colonize, or re-colonize wetlands (Homan et al. 2003). Research on efficacy of lure crops is just beginning and time will be needed to assess this method. Sometimes population management may be the most realistic and effective method for reducing persistent damage caused by overabundant wildlife populations (Garrott et al. 1993). The definition of overabundance is subjective and based upon one’s interests, likes, and dislikes; and therefore is controversial. It has been defined generally as that point of abundance where all positive values of the species have been overwhelmed by negative values created by the sheer numbers of indi- viduals (Conover 2002). The definition implies both a THE CANADIAN FIELD-NATURALIST Vol. 118 spatial and temporal component, as populations are seldom ubiquitous or always aggregated. The defini- tion is fitting, at least from an agricultural perspective, in the sunflower growing areas of North Dakota and northeastern South Dakota. The USDA has drafted an Environmental Impact Statement (EIS) that addresses the conflict between sunflower producers and black- birds in North Dakota, South Dakota, and Minnesota (Federal Register 2001*). Several management options were included in the draft EIS. The research described herein addresses one of the options being considered: population management of spring migrating Red- winged Blackbirds in northeastern South Dakota. This part of South Dakota is one of the most northward sites where the population is still grouped together enough to make population management a feasible alternative. Because blackbird damage to sunflower is a local- ized phenomenon occurring within the broader region of the northern Great Plains (Peer et al. 2003), it is nec- essary to identify specific origins of local populations before implementing any strategy of population man- agement. Although leg-banding has provided valu- able insights into migratory timing and dispersal of Red-winged Blackbirds (Dolbeer 1978), this method draws its samples randomly from regional populations. Moreover, it is labor intensive in both its capturing and ringing methods; these drawbacks combined with natural annual mortality of Red-winged Blackbirds cause data to be parsed over several years because of extremely low rates of band returns (<0.5%, Besser et al. 1983; Gammell et al. 1986; Stehn 1989). An alterna- tive method of tracking movements, aerial mass color- marking, provides a means for rapid accumulation of data on movements of local populations within a single season framework. Recovery rates from mass color- marking projects depend on the number marked com- pared to the size of the study population (Johns et al. 1989): however, thousands of birds can be marked in a single application often yielding a rate of return 10 to 20 times that of leg-banding. Aerial mass color- marking has been used to track spring movements of Red-winged Blackbirds from northeastern Missouri and eastern South Dakota (Knittle et al. 1987; Knittle et al. 1996), short late-summer movements of Red- winged Blackbirds in central North Dakota (Linz et al. 1991), and movements of Red-winged Blackbirds between winter roosts in the southcentral U.S. (Harsch 1995*). The marking formulation is environmentally safe at the concentrations used (Bills and Knittle 1986; Knittle and Johns 1986). In April 2001, we used aerial mass-marking to iden- tify migrating Red-winged Blackbirds staging in large cattail-dominated wetlands in northeastern South Da- kota. Millions of spring migrating blackbirds annu- ally stage at this traditional stopover site between late March and April (Sawin 1999). Our objective was to assess migratory dispersal to breeding territories north of the marking site. Previous research has shown that 2004 HoMaAn, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 203 250 200 North Dakota L 5 “, a 150 ree % = Fi g NL * Saskatchewan 7) oe ‘ — « Le = 5 a _Manitoba 8 100 rid PO eee ee a ims ” i a Fd a ” © Alberta = 50 0 < 2 2 2 ND 2 £ Year Ficure 1. Three-year rolling averages of indices from the North American Breeding Bird Survey from 1967 to 2003. The in- dices are from routes in the Drift Prairie, Glaciated Missouri Plateau, and Aspen Parklands, which are the three major physiographic regions in the Prairie Pothole Region used most frequently by Red-winged Blackbirds. color-marked male Red-winged Blackbirds staging in eastern South Dakota during spring were in part destined for breeding grounds in or near concentrated sunflower production; however, these data were col- lected over 15 years ago (Knittle et al. 1987; Knittle et al. 1996). Population management programs, when used at sites far from the locality where actual damage occurs, must be based on a thorough and contempo- rary knowledge of movements of individual popula- tions (Dolbeer 1978). We believed it necessary to repli- cate the marking studies because changes may have occurred in migratory patterns. Additionally, no data exist on dispersal movements of females, and these data would increase our understanding of the popula- tion’s migration patterns. Methods Marking During March 2001, we monitored cattail-dominat- ed wetlands in six counties in eastern South Dakota for arrival of migratory flocks of blackbirds. Three wet- lands held migratory roosts large enough (>10 000 birds) to warrant aerial mass-marking. All three roosts were within a 32-km radius of Badger, South Dakota (44°48'N, 97°21'W). Roost size was estimated by counting blackbirds as they entered the roost on the evening of the spray (Meanley 1965; Arbib 1972). Spe- cies compositions and sex compositions were estimat- ed by randomly selecting and identifying groups of individual birds as they entered (Dolbeer et al. 1978). A fixed-wing agricultural spray plane applied the marker near dark at an altitude of 20—30 m (Knittle et al. 1987). The following morning we collected birds as they departed the sprayed roost and later examined them for fluorescent marks using a long-wave (360 nm) ultraviolet light. We calculated the number of marked male and female Red-winged Blackbirds in the roost by multiplying the proportion of marked birds in the sample by the estimated number of males and females using the roost on the night of the spray. The marker formulation was a 416-L solution con- sisting of 208 L Carboset® (an acrylic adhesive; Nov- eon™ Incorporated, Cleveland, Ohio), 102 L food- grade propylene glycol, 7.7 kg fluorescent organic pig- ment (DayGlo® Color Corporation, Cleveland, Ohio), 1 L each of a defoamer and surfactant, and 106 L water (Knittle et al. 1996). We used CP® nozzles set at 5 (400 micron droplets). Spray volume was 28L/ha delivered at 1.8 kg/cm’. In spray form the solution dries in 2-3 minutes adhering to the feather surface. 204 Knittle and Johns (1986) found that 5% of marked birds lost their marks within six weeks; however, particles lodged in the barbules can remain until molt (Knittle et al. 1996; Otis et al. 1986). Sampling and Collections We grouped the study area into 140 blocks, each consisting of four 1°-latitude x 1°-longitude units. Two units were randomly selected from each block. To en- sure adequate sample sizes per effort, only those units within regions that averaged 230 Red-winged Black- birds per BBS route were selected (Sauer et al. 2004); 37 units in Canada and 19 units in the U.S. were chos- en in this manner. No units south of the 1° x 1° unit containing the marking sites were sampled. The sam- ple units covered an extensive area (>1.0 x 10° km?) bounded by 44—57° N latitude and 95—119° W longi- tude (Figure 2). Southwestern North Dakota, central Minnesota, central and western South Dakota, and southeastern Montana were excluded because BBS counts were <30 (Sauer et al. 2004). We assigned the 1° x 1° units to one of three disper- sal polygons: core, peripheral, and outside-peripheral (Figure 2). The shape and area of the polygons were based on an analysis of banding and re-sighting data, physiography, and proximity to the area of concen- trated sunflower production (Stehn 1989). The core polygon encompassed the breeding range of birds hav- ing a high probability of causing early damage to sun- flowers. The peripheral polygon contained the sur- rounding breeding range of birds that were capable (with only short pre-migratory movements) of entering the core area during the early damage period (mid- August — early September) (Dolbeer 1978; Gammell et al. 1986). The outside-peripheral polygon contained the birds farthest removed from the core area and en- compassed the rest of the study area not covered by the core and peripheral polygons. The 1° x 1° units were placed in core and peripheral polygon categories if any of the unit’s area intercepted the polygon’s bound- ary. We used a GIS to place the polygons on a geo- referenced grid of the 1° x 1° units. We used shotguns (12-gauge with either #6 or #7/ steel shot) to collect male and female Red-winged Blackbirds on breeding territories during June in the U.S. and June and early July in Canada. The wings were removed from the specimens and placed in an envelope with the GPS coordinates of the collection site. The wings were later viewed for fluorescent pig- ments using a 360-nm ultraviolet lamp under 1.75 mag- nification. The markings on tagged birds are usually subtle and invisible to the naked eye under natural light. To eliminate false positives, only splash marks and individual particles firmly attached to the feathers were considered conclusive evidence of a valid mark (Knittle et al. 1987). Blood samples were removed on- site from randomly selected specimens and used for microsatellite genetic analysis in a separate study (Williams et al. 2004). The wings were frozen and shall THE CANADIAN FIELD-NATURALIST Vol. 118 be kept for future research on primary feather chem- istry and its relationship to geographic origins (Ed- wards and Smith 1984). The data were analyzed using descriptive statistics, likelihood ratio chi-square analyses (G-tests), and non- parametric tests. We used G-tests to assess differences between (1) the proportion of marked birds (sexes com- bined) in the sample with the expected proportion of marked birds in the study area; (2) the proportions of marked males and females in the sample with the ex- pected proportions of marked males and females in the study area; and (3) the proportions of marked birds within core, peripheral, and outside-peripheral poly- gons. The first two tests used extrinsic expected fre- quencies based on estimates of marking efficiency at the staging site and population size in the study area during the breeding season. The third comparison used intrinsic frequencies generated exclusively from the collections (Sokal and Rohlf 1981). Population size in the study area was estimated using a combination of stable age structure analysis from banding data and linear regression analysis of BBS indices and inde- pendently determined density estimates (Stehn 1989). The resultant regression equation was Y = 0.3X, where X was the average BBS count within each ecological stratum in the study area and Y was the population den- sity per km? (see Aldrich 1963; Robbins et al. 1986). We used the Wilcoxon Rank Sum test to compare ranked differences between males and females in dis- tances traveled from the marking site to the collecting sites. Distance measurements were made using a GIS. Results We applied fluorescent marker seven times, tagging an estimated 235 500 male and 131 100 female Red- winged Blackbirds over the period 1-23 April. This period covered the Red-winged Blackbird migration through eastern South Dakota in 2001. We collected 2398 males and 2060 females on breeding territories, of which 33 (0.74%) were marked (Table 1). The overall recovery rate of 0.74% was similar to the expected recovery rate of 0.96% (G = 1.1, P = 0.30), based on total number of birds marked in April and the estimated population size of breeding Red-winged Blackbirds in the sampled collection area (38.1 x 10°, from Stehn 1989) which included the population in the outside-peripheral polygon. The ratio of marked males : females in the sample (f = 23 : 10) was near the expected ratio of 22 : 11 (G = 0.07, P = 0.80). Proportions of marked birds collected in the three polygons differed (G = 6.44, P = 0.04), with the per- centages of marked birds in the core polygon (1.03%) and peripheral polygon (0.93%) nearly three times greater than the percentage collected in the outside- peripheral polygon (0.35%) (Table 1). The outside- peripheral polygon contributed 69% of the total G- value. Of the 33 marked recoveries, 15 (46%) were collected in the core polygon and 12 (36%) were collected in the peripheral polygon. 2004 Homan, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 205 Outside-peripheral polygon Peripheral polygon Core polygon Axel study block © Female N A Male A 3x Marking site 300 0 300 Kilometers a okt FiGurRE 2. Locations of 33 marked Red-winged Blackbirds (in relation to core, peripheral- and outside-peripheral polygons) after dispersing from a spring staging site in eastern South Dakota in April 2001. TABLE 1. Color-marked male (©’) and female (Q) Red-winged Blackbirds collected during June and July 2001 in core, peripheral, and outside-peripheral polygons after migratory dispersal from a staging site in eastern South Dakota during April 2001. Number Number Number Percent units? Collected Marked Marked Polygon ‘os O Total ‘oi fe) Total ol Q All Core 14 793 668 1 461 9 6 15 eit} 0.90 1.03 Peripheral 15 681 605 1 286 10 2 12 1.47 0.33 0.93 Outside 27 924 787 Lala 4 D 6 0.43 0.25 0.35 Total 56 2 398 2 060 4 458 2B 10 33 0.96 0.48 0.74 4No birds were collected in two units 206 No marked birds were collected north of 53° N. Recovery distances were similar between sexes (T = 0.18, P = 0.86), with the median location 519 km (range: 20-1353) from the marking sites (48°23'N, 100°27'W). The median azimuth from the marking sites was 325°. The median distance and direction of marked females placed them on the northwestern side of the core polygon (48°23'N, 100°27'W; whereas, males were 70 km due west (48°30'N, 101°23'W), in the peripheral polygon. For all recoveries combined, the median location was 50 km northeast of Minot, North Dakota (48°14'N, 101°18'W). Discussion The seven sprays were evenly distributed through- out the migratory period and probably yielded an accurate depiction of the migratory distribution pat- terns of Red-winged Blackbirds. Similar to the results of previous banding and color-marking experiments in eastern South Dakota, our data showed that most birds followed the Prairie Pothole Region as they traveled northwest to their breeding territories (Besser et al. 1983; Knittle et al. 1987; Knittle et al. 1996). Although Red-winged Blackbirds are not confined to breeding in this region, they are found here in their highest densities. A statewide population census of breeding birds in North Dakota showed that the Prairie Pothole Region (comprised of several physiographic regions, but mainly the Drift Prairie and Glaciated Missouri Plateau) contained 70% of the state’s Red- winged Blackbird population (Stewart and Kantrud 1972). Unlike parameters such as median distance and direction, both of which can be influenced by differ- ences in breeding densities over the study area, the pro- portions of marked birds within the three polygons were distribution measures mathematically independent of density. It was thus possible to consider our results as reflections of behavioral phenomena, such as fidelity to migratory pathways and philopatry. The marked population was apparently not distributing itself ran- domly in the study area. The low proportion of marked birds in the outside-peripheral polygon suggested that eastern South Dakota may be used mainly by migrants already near their final destination, and as a result, traveling relatively short distances after departure from the staging site. We speculate that other staging sites might exist (perhaps much farther to the west), and these should have birds destined for breeding territories in Montana, southern Alberta, and Saskatchewan. Marking of large roosts farther west (for example, a large blackbird roost exists near Great Bend, Kansas [38° 22'N, 98° 49'W]) may demonstrate that western migratory pathways exist distinct from the one used by blackbirds in eastern South Dakota. Certainly, the very low proportion of marked birds in the outside-periph- eral polygon, compared to the other two dispersal polygons, suggests a different population of birds. THE CANADIAN FIELD-NATURALIST Vol. 118 There were two other marking studies done in the general vicinity of our marking site. These were done at Lake Thompson, South Dakota (44°23'N, 97°33'W), approximately 30 km to the southwest. Both were similar in design, scope, and sampling effort to our study, except that only males were collected. One study was conducted in 1983 (Knittle et al. 1987), the other in 1985 (Knittle et al. 1996). Even though geopolitical boundaries were used to analyze returns instead of dis- persal polygons, comparisons of results among and between studies can still be made. Results were similar between our study and the 1983 study in two impor- tant facets: (1) the recovery rate of marked birds in Canada was substantially lower than the U.S. recov- ery rate (4.4 vs. 7.0%, Knittle et al. 1987), and (2) proportions of marked birds collected in western North Dakota, central North Dakota, and northeastern South Dakota were greater than other regions. By contrast, results diverged in the 1985 study, both from our results and from those from the 1983 study. The expected rate of recovery in the 1985 study was much less than the observed recovery rate (5.2 vs. 14.6%, Knittle et al. 1996). To obtain a 14.6% recovery rate, the male population would have had to have been 6.7 x 10°, a number 65% lower than what was estimated in the study area during that time (19.0 x 10°, Stehn 1989). Saskatchewan’s Red-winged Blackbird population had indeed been in steep decline for nearly a decade, as evidenced by BBS indices (Figure 1); however, trends for Alberta, Manitoba, and North Dakota were either stable or increasing. It is unlikely for such a conspicuous species that a 65% decline would be missed in the BBS surveys. Of course, the number of birds marked in 1985 could have been underestimated by this magnitude, but an even more anomalous result in the 1985 study argues against this; the percentage of marked birds in Canada was greater than the per- centage marked in the Prairie Pothole Region below 49° N (17 vs. 16%). Usually, recovery rates are inverse- ly related to the distance traveled from a marking site because unmarked populations arriving by other routes from other areas dilute the proportion of marked birds. Although it is conceivable that the Red-winged Black- bird population marked at Lake Thompson in 1985 migrated beyond traditional nesting areas and immi- grated into Canada, perhaps seeking better wetland conditions, band recoveries collected over several dec- ades have shown that Red-winged Blackbirds usually return to within 100 km of their natal sites (Nero 1956; Dolbeer 1978). If Red-winged Blackbirds were not philopatric, but nomadic or irruptive, we contend that large fluctuations in BBS counts would occur among physiographic regions comprising the Prairie Pothole Region. No significant correlations existed in yearly changes in BBS indices between North Dakota and the three Prairie Provinces within our study area, and perusal of average BBS indices over a 36-year period does not seem to indicate irregular migratory aegnnce behavior (Figure 1). The extraordinarily high overall recovery rate of marked birds in the 1985 study may have resulted from improper identification of marks (i.e., false positives) caused by environmental contam- ination with fluorescence pigments used for identifying insecticide formulations and fertilizers (Knittle et al. 1987). We believe that our data from 2001 and the data from the 1983 study have provided the most accurate analyses of dispersal patterns of migrating Red-winged Blackbirds in eastern South Dakota. Summary and Conclusions In landscapes with numerous wetlands, blackbird damage to sunflowers averages 25% per field. One- quarter of the fields receive >10% damage (Linz et al. 2000). At the 10% damage level, agricultural producers start dropping sunflower from their crop rotations and replacing it with other oil crops, such as soybean and canola (North Dakota Agricultural Statistics 1990*; Lamey and Luecke 1994). There is no evidence that the rapid population growth of Red-winged Blackbirds has directly affected other bird species; however, the potential for indirect effects exists. Sunflower fields were used by 49 nonblackbird species, representing all the major feeding guilds, during the late-summer and early fall migrations (Schaaf 2003). Sunflower fields have several characteristics that could make them excel- lent migratory stopover sites for rest, cover, and pro- curement of energy reserves (Hutto 1998; Petit 2000). First, they provide large blocks of sturdy herbaceous cover with a dense canopy and rank understory in a landscape that by mid-August is mostly barren north of 46°N because of the harvest of small grains (Schaaf 2003). Second, sunflower fields provide a diverse sel- ection of foods, including seeds of weedy plants in the furrows and numerous invertebrates in and around the fields (Schulz 1978; Charlet et al. 1997; Schaaf 2003). This attracts avian granivores and insectivores (and of course, the carnivores). Soybean and canola fields, which often replace sunflower, have neither the beneficial vegetative structure nor the energy and are a less productive habitat for birds (Linz et al. 2004d). Moreover, these crops are “clean cropped,” unlike sun- flower, which is often not plowed until May of the fol- lowing year. Sunflower stubble was used by 33 species of migrating birds during early spring in North Dakota (Galle et al. 2004). It was used more frequently during spring migration than fields of soybean, small grain, corn, and sorghum. Thus, loss of sunflower fields in the Prairie Pothole Region could have unforeseen ecological impacts on migrating nonblackbird species. We estimate that 82% of the Red-winged Blackbirds that stage in eastern South Dakota during spring migra- tion reside either within or on the periphery of the sun- flower growing area. If population management of the spring breeding population in the northern Great Plains is deemed the best of the EIS alternatives, we suggest that the most efficacious strategy would be to HoMAN, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 207 manage the population in eastern South Dakota. Lastly, it may be possible to avoid impacting the Canadian migrants, which comprised 18% of the birds staging in eastern South Dakota, by not applying management over the entire migration. Canadian Red-winged Black- birds should be among the last of the regional popula- tions of Red-winged Blackbirds to arrive at the stag- ing site (Weatherhead and Bider 1979). The arrival of Yellow-headed Blackbirds (Xanthocephalus xantho- cephalus), a nontarget blackbird species, usually signi- fies that the Red-winged Blackbird migration is nearing its end and may be a good indicator to cease manage- ment operations. We recommend future research on (1) genetic markers to identify local populations of Red-winged Blackbirds residing in Canada and the U.S.; (2) color-marking spring migrants in flyways farther west; (3) the relationship between timing of arrival at stag- ing sites and distance traveled to breeding territories; and (4) color-marking pre-migratory roosts [or other means of identification, such as genetics or mineral analyses of primary feathers] northwest of the core sunflower growing area prior to fall migration. Acknowledgments Personnel from North Dakota/South Dakota Wildlife Services and North Dakota State University helped collect the birds. We thank the following for their sup- port and cooperation on this project: the Departments of Agriculture of North Dakota and South Dakota; the U.S. Fish and Wildlife Service; Minnesota Department of Natural Resources; North Dakota Game and Fish Department; South Dakota Department of Game, Fish and Parks; South Dakota Department of Environment and Natural Resources; Montana Department of Fish, Wildlife and Parks; Canada Customs and Revenue Agency; Canada Department of Foreign Affairs and International Trade; the Environmental Protection Branch of Environment Canada; the Wildlife Branch of Manitoba Conservation; the Environment Branch of the Alberta Natural Resource Service; the Fish and Wildlife Branch of Saskatchewan Environment and Resource Management; the Manitoba Wildlife Animal Care Committee and Animal Care Committees within other provincial, federal, and state governments; the National Sunflower Association; and private landown- ers in the U.S. and Canada. Our research was conduct- ed under the auspices of Quality Assurance Study Pro- tocol QA-776, as amended; as such, safety guidelines, animal care, and scientific validity were reviewed, approved, and permitted by the National Wildlife Research Center, the U.S. Fish and Wildlife Service, the Environmental Protection Branch of Environment Canada, and the natural resource agencies in the states and provinces within the study area. Use of commercial products by the U.S. Government does not constitute an endorsement of these products. 208 Documents Cited (marked * in text) Dahl, Thomas E. 1990. Wetlands losses in the United States 1780’s to 1980’s. U.S. Department of the Interior, Fish and Wildlife Service, Washington, D.C., United States Geo- logical Survey, Northern Prairie Wildlife Research Center, Jamestown, North Dakota. http://www.npwrc.usgs.gov/ resource/othrdata/wetloss/wetloss.htm (Version 16JUL97). Federal Register. 2001. USDA, Animal Plant Health Inspection Service. 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Linz. 1986. A method for estimating turnover in spring blackbird roosts. Journal of Wildlife Management 50: 567-571. Homan, LINZ, ENGEMAN, and PENRY: DISPERSAL PATTERNS OF BLACKBIRDS 209 Peer, B. D., H. J. Homan, G. M. Linz, and W. J. Bleier. 2003. Impact of blackbird damage to sunflower: bioener- getic and economic models. Ecological Applications 13: 248-256. Petit, D. R. 2000. Habitat use by landbirds along nearctic- neotropical migration routes: implications for conservation of stopover habitats. Studies in Avian Biology 20: 15-33. Ralston, S. T., G. M. Linz, and W. J. Bleier. 2004. Quanti- fication of cattail (Typha spp.) in the Prairie Pothole Region of North Dakota in relation to blackbird damage to sunflower. Pages 417-422 in Proceedings of the 16'" International Sunflower Conference. Edited by G. Seiler. International Sunflower Association, Paris, France. Robbins, C. S., D. Bystrak, and P. H. Geissler. 1986. The Breeding Bird Survey: its first fifteen years, 1965-1979. 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Received 30 July 2002 Accepted 29 October 2004 Changes in Loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) Populations in the Lower Matanuska-Susitna Valley, Alaska TAMARA K. Mitis!* and BRAD A. ANDRES2* ' Department of Biological Sciences, University of Alaska Anchorage, 3211 Providence Drive, Anchorage, Alaska 99508 USA 2 U. S. Fish and Wildlife Service, Nongame Migratory Bird Management, 1011 East Tudor Road, Anchorage, Alaska 99503 USA > current address: U. S. Fish and Wildlife Service, Nongame Migratory Bird Management, 1011 East Tudor Road, Anchorage, Alaska 99503 USA * current address: Division of Migratory Bird Management, U. S. Fish and Wildlife Service, P.O. Box 25486, DFC-Parfet, Denver, Colorado 80225-0486 USA; corresponding author Mills, Tamara K., and Brad A. Andres. 2004. Changes in loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) populations in the lower Matanuska-Susitna Valley, Alaska. Canadian Field-Naturalist 118(2): 210-214. More than two-thirds of the human population of Alaska resides in the south-central portion of the state, where its continued growth is likely to affect some wildlife populations negatively. To assess changes in waterbird populations in this region, we compared counts of Common Loons (Gavia immer), Pacific Loons (G. pacifica), and Red-necked Grebes (Podiceps grisegena) made on Matanuska-Susitina Valley lakes. In general, the number of lakes occupied by loon or grebe pairs decreased between 1987 and 1999. Decreases in the number of lakes occupied by Common Loons were less drastic in the northwest region of the study area than in the southeast region; human development is greater in the southeastern portion of our study area. Contrary to lake occupancy, the percentage of lakes that fledged Common Loon chicks remained stable between years. Because the human population is expected to continue to grow, proactive management of lake use and lakeshore development, coupled with monitoring of loon and grebe occupancy and productivity, is needed to ensure the persistence of these waterbird populations in the lower Matanuska-Susitna Valley. Key Words: Common Loon, Gavia immer, Pacific Loon, Gavia pacifica, Red-necked Grebe, Podiceps grisegena, breeding occupancy, productivity, south-central Alaska. The stability of many bird populations mainly de- pends on the stability of the environments they inhabit (Newton 1998). Nesting loons (Gavia spp.) and grebes (Aechmophorus spp., Podiceps spp.) are often nega- tively affected by habitat degradation and increased levels of human-related disturbances (Riske 1976; McIntyre 1978; Evers 2003). Common Loons (Gavia immer), Pacific Loons (G. pacifica), and Red-necked Grebes (Podiceps grisegena) breeding on lakes in the Matanuska-Susitna Valley (Mat-Su Valley), Alaska, are encountering many changes to their local environment. The area’s human population has tripled in the last two decades, and the current annual growth rate is 3.3% (McKibben and Nelson 1999*). Within the Mat-Su Valley, population growth has been greatest in the southeast, the area closest to Alaska’s largest city (Anchorage; McKibben and Nelson 1998*, 1999*),. The growing human population in the Mat-Su Valley has begun to encroach upon the freshwater habitat favored by breeding loons and grebes. Development of lake shorelines for houses and increased recreational use of lakes are suspected of having detrimental effects on loon and grebe populations in the Mat-Su Valley (Tankersley 1987*, Fair 1998*). Concern over the susceptibility of local loon popu- lations to anthropogenic pressures prompted the forma- tion of a volunteer program, the Alaska Loon Watch, in 1985. Under the direction of the Alaska Department of Fish and Game (ADF&G), volunteers collected data on loon and grebe presence and breeding activity on lakes throughout the Mat-Su Valley (see Fair 1998*). Using data collected through the Alaska Loon Watch program and our own observations, we assessed changes in lake occupancy, distribution, and productivity of loons and grebes breeding in the Mat-Su Valley between 1987 and 1999. Based on the significant growth of the human population in the region, we predicted that lake occupancy and productivity by loons and grebes would have decreased over time. Furthermore, we predicted that the distribution of loons and grebes would shift spatially from southeast to northwest in response to corresponding changes in intensity of shoreline devel- opment and lake recreation. Study Area The Mat-Su Valley of south-central Alaska lies be- tween the Matanuska and Susitna rivers and borders upper Cook Inlet. Over 1000 lakes, ponds, and wet- lands of glacial origin occupy kettles in moraines and ice-stagnation complexes in this 500 km? region (Colazzi et al. 1986°). Lakes are classified as oligo- trophic or mesotrophic with pH levels conducive to support fish species preyed on by loons and grebes (Ruggles 1991). The elevation of the valley floor ranges from tide level to 400 m, but local relief of the area is 210 2004 Uplands are dominated by White Spruce (Picea glauca), Paper Birch (Betula papyrifera), and Quaking Aspen (Populus tremuloides), Black Cottonwood (Populus trichocarpa) and willows (Salix spp.) are common along waterways and on alluvial plains. Maritime in- fluences moderate the climate in the Mat-Su Valley with average mid-summer temperatures of 14°C, annu- al snowfall of 150 cm, and annual precipitation of about 48 cm. We concentrated our survey effort with- in the lower Mat-Su Valley, an area bounded by the MILLS AND ANDRES: CHANGES IN LOON AND GREBE POPULATIONS IN ALASKA 211 communities of Palmer to the southeast, Willow to the northwest, Sutton to the east, and the Susitna River to the west (Figure 1). Methods We surveyed lakes in late May and late August of 1999 to determine occupancy by loons and grebes and productivity of loons. Because of their smaller size and tendency to linger in aquatic vegetation, produc- tivity of Red-necked Grebes was not recorded. Of \ Palmer 3) ¥ x ips L ys > So & (a) = c Sans Eklutna oars 1 5 ys iG co N L b agg “ Sn kee Sy 0 he 2, ie \ BN - « s * 4 €s V » ? 4 p as ae ya a .. OK 3 A Woe CRS e s Oy eh ¥ a s S on : F Ae) i Sy Z wall, ot ree ° ° 7€ é =, ~- a © 10 20 Kilometers Fee | : ne ea FiGurE 1. Location of the lower Matanuska-Susitna Valley, Alaska, where lake occupancy and productivity of loons and grebes were assessed, in the delineated area, in 1987 and 1999. D2 219 lakes surveyed in 1987 by biologist Nancy Tankersley (ADF&G) and volunteers of the Alaska Loon Watch program, we determined that 139 lakes had adequate coverage in both years to allow a mean- ingful comparison. In 1999, observers scanned each lake with binoculars from several vantage points along the lakeshore to ensure complete lake coverage. Lakes were scored for the presence or absence of single loons or loon pairs during late May and for the presence or absence of loon chicks during late August. Field meth- ods were similar in 1987. A lake was considered oc- cupied if a loon pair was found on a lake surveyed on or before 5 July, or if chicks were discovered on the lake. Only lakes surveyed after 25 June were used to determine productivity. To detect changes in the spatial distribution of loons over the 12-year period. We divided the study area was divided into two regions of differing human pop- ulation densities. The Little Susitna River runs east to west and divides the study area into southeastern (Was- illa, high human population) and northwestern portions (Willow, low human population). Lake physiographic features are comparable between regions, and lakes vary similarly in size, food availability, and shoreline topography (Ruggles 1991). We used McNemar’s symmetry test for matched pairs to compare temporal and geographic differences in lake occupancy and productivity (Agresti 1996 pages 226-229). Alternative hypotheses were constructed to indicate a decline in parameter estimates between 1987 and 1999; no statistical analysis was conducted when n <3 lakes. We used P-values to examine strength of differences between years and regions. Standardized normal standard errors of differences were also cal- culated (Agresti 1996 page 228). Results The number of lakes occupied by any loon or grebe species decreased significantly between 1987 and 1999 across the entire study area (Table 1). Decreases in lake occupancy between years were consistent among Common Loons, Pacific Loons, and Red-necked Grebes (Table 1). Differences in occupancy of lakes by loon or grebe pairs, however, were not uniform among regions of the Mat-Su Valley. Decreases in the num- ber of lakes occupied by any loon or grebe pair and by Common Loons were less drastic in the northwest region of the study area (Willow) than in the southeast region (Wasilla; Table 1), where development is greater. Decreases in lake occupancy by Pacific Loons tended to be greater in the northwest section (P = 0.1094), but conclusiveness of results was hampered by small sample sizes (Table 1). Point estimates of decreases in occupancy by Red-necked Grebe pairs were similar between regions, but variability was much higher in the northwest region (Table 1). Contrary to lake occupancy, the percentage of lakes that fledged Common Loon chicks remained stable between 1987 and 1999. Information on productivity THE CANADIAN FIELD-NATURALIST Vol. 118 of Red-necked Grebes was not collected in 1987, and sample sizes were too small to permit analysis of changes in productivity for Pacific Loons. Productivity of Common Loons was similar on lakes in both regions of the study area. Discussion Temporal and spatial changes in the population dis- tribution of loons and grebes in the Mat-Su Valley from 1987 to 1999 are likely attributable to the concurrent rise and spatial settlement of the human population. Significant declines in lake occupancy by Common and Pacific loons and Red-necked Grebes indicated that fewer birds are establishing breeding territories at area lakes. Most of the lakes that are no longer used by breeding loons and grebes (65%) were located in the southeastern portion of the study area — an area that has also experienced the greatest human population growth. Common Loons have been extirpated from lakes within the nearby Municipality of Anchorage, and Pacific Loon populations are decreasing there (Fair 1998*). Such changes in lake occupancy may be indicative of declining loon and grebe populations or may reflect large-scale emigration due to the loss of suitable nesting habitat. Common Loons inhabiting boreal regions of Alaska, however, appear to be stable (Groves et al. 1996). Population declines of Common Loons and Red- necked Grebes throughout North America have oc- curred most frequently at the southern boundaries of their breeding ranges where habitat quality has become marginal owing to development (De Smet 1987; Mc- Intyre 1988). In the Mat-Su Valley, a spatial shift in lake occupancy from southeast to northwest may be linked to habitat quality decreases in the same spatial direction. Habitat in the southeastern portion of the study area is likely poorer in quality compared to the northwest, where fewer people live and more lakes remain undeveloped. The stability of lake occupancy to the northwest implies that habitat quality is optimal, a fact supported by the rapid reoccupation of vacant territories (sensu Newton 1998). The negative trends in lake occupancy may also reflect changes in the demographics of the Mat-Su Valley loon and grebe populations. Loons and grebes are long-lived species known to return to breed on the same lake territories year after year (McIntyre 1988; Stout and Neuchterlein 1999). The combination of a long lifespan and strong site-fidelity may have caused individuals to occupy lakes that had deteriorated in quality over several years. When existing occupants died or left, the vacancies were not filled, and lake occupancy declined over time. Thus, lakes may have lost loons and grebes not suddenly through abandon- ment by resident breeders, but by failure of new birds to replace them after the death of residents. Although habitat in the southeastern region may be less suitable during selection of a territory, loons may remain productive if they choose to nest there. Greater 2004 MILLS AND ANDRES: CHANGES IN LOON AND GREBE POPULATIONS IN ALASKA ZAS TABLE |. Occupancy and productivity of breeding pairs of loons and grebes on lakes in high (Wasilla) and low (Willow) human population regions of the lower Matanuska-Susitna Valley, Alaska, in 1987 and 1999. Species — stage Percentages of lakes occupied Difference (SE) area both years 1987 only 1999 only 1999 — 1987 P-value’ Any loon or grebe - occupancy entire area (n = 86) 67 Di, 6 -21 (6) 0.0005 Wasilla (n = 52) 71 29 0 -29 (6) <0.0001 Willow (n = 34) 62 23 15 -8 (10) 0.2905 Common Loon - occupancy entire area (n = 63) 59 33 8 =25) | + (7) 0.0012 Wasilla (n = 36) 58 42 0 -42 (8) <0.0001 Willow (n = 27) 59 22 19 -4 (12) 0.5000 Pacific Loon - occupancy entire area (n = 17) 41 47 12 -35 (17) 0.0547 Wasilla (n = 10) 40 50 10 -40 (21) 0.1094 Willow (n = 7) 43 43 14 -29 (26) 0.3125 Red-necked Grebe - occupancy entire area (n = 33) 76 24 0 -24 (7) 0.0039 Wasilla (n = 25) 76 24 0 -24 (9) 0.0156 Willow (n = 8) 75 25 0 -25. (15) 0.2500 Common Loon - productivity entire area (n = 34) 24 38 38 OP Fas) 0.5775 Wasilla (n = 21) 29 33 38 5 (18) 0.5000 Willow (n = 13) 15 46 38 -8 (25) 0.5000 Pacific Loon - productivity entire area (n = 10) 80 0 20 ~ - Wasilla (n = 7) 86 0 14 - - Willow (n = 3) 67 0 33 - - ' one-sided test, 1987 > 1999. * small sample sizes precluded reasonable statistical testing. breeding success for Common Loons in the south- eastern region may reflect differences in the age struc- ture of the population. Across numerous species, non- breeders and non-territory holders consist mainly of sub-adult birds (Smith 1976; Birkhead et al. 1986). Because replacements for vacant territories in the north- west are likely drawn from the non-territorial cohorts (Ruggles 1991), these loons would be expected to have lower productivity than experienced territorial loons on southeast region lakes. Tolerance of human activity by experienced territory holders may also explain the differences in Common Loon breeding success between the regions. Common Loons are known to be relatively flexible in behavior and may acclimate themselves to low-level human dis- turbance (Titus and VanDruff 1981; Heimberger et al. 1983; McIntyre 1988). Years of breeding experience and gradual habituation to the subtle progression of human disturbance would likely result in higher breed- ing success for loons that established territories years ago. Loons may also benefit by nesting in areas where humans are moderately active. Recreational activities and residents who watch over loons may deter preda- tors from taking young or eggs. Reports from Alaska Loon Watch volunteers often include stories of resi- dents protecting adults and chicks by discouraging Bald Eagles and other predators (Tankersley 1987*, Mills, personal observation). This interference, com- bined with the defensive behavior of an adult loon, may be intense enough to dissuade additional preda- tion attempts. Changes in the distribution of loons and grebes warrant concern about the future of these populations within the region. The human population is expected to continue to increase in the Mat-Su Valley (Alaska Department of Natural Resources 1998), which could result in further changes to the lake habitat used by loons and grebes. Although degradation of lake habi- tats are likely widespread across south-central Alaska, only a small portion of North America’s populations of Common Loon, Pacific Loons, or Red-necked Grebes occur in south-central Alaska. People residing in south- central Alaska, however, value living with wildlife and believe it makes their community “interesting and special” (Alaska Department of Fish and Game 2000). Therefore, proactive management of lake use and lake- shore development, coupled with continued monitoring of loon and grebe populations, is needed to improve the stability of the freshwater habitats and ensure the continued persistence of loons and grebes in the Mat- Su Valley of Alaska. 214 Acknowledgments We sincerely thank Nancy Tankersley and all the Alaska Loon Watch volunteers for their devoted inter- est and effort. This project was funded by the U. S. Fish and Wildlife Service, Region 7 Nongame Migra- tory Bird Program, the North American Loon Fund, and the Alaska State Parks. Special thanks to Rick Mills for assisting in field work. Helpful manuscript comments were provided by A. J. Erskine. Documents Cited (marked * in text) Colazzi, E. J.. M. A. Maurer, and S. J. Carrick. 1986. Reconnaissance of surface water resources in the Houston area, Alaska. Alaska Division of Geological and Geo- physical Surveys, Public-Data File 86-62. 19 pages. Evers, D. C. 2003. Status assessment and conservation plan for the Common Loon (Gavia immer) in North America. Unpublished report, U. S. Fish and Wildlife Service, Hadley, Massachusetts. 80 pages. Fair, J. 1998. The status of loons in Anchorage and the lower Matanuska-Susitna Valley of Alaska: a summary report of Alaska Loon Watch 1985-1997. Unpublished report, Anchorage Audubon Society, Anchorage, Alaska. 36 pages. McKibben, B., and R. Nelson. 1998. Matanuska-Susitna Borough planning department — 1998 facts. Palmer, Alaska. 15 pages. McKibben, B., and R. Nelson. 1999. Matanuska-Susitna Borough planning department — 1999 facts. Palmer, Alaska. 35 pages. Tankersley, N. 1987. Alaska Loon Watch 1987. Unpublished report, Alaska Department of Fish and Game, Anchorage, Alaska. 27 pages. Literature Cited Agresti, A. 1996. An introduction to categorical data analysis. John Wiley and Sons, Inc., New York, New York. 290 pages. Alaska Department of Fish and Game. 2000. Living with wildlife in Anchorage: a cooperative planning effort. Anchorage, Alaska. 138 pages. Alaska Department of Natural Resources. 1998. Alaska’s outdoor legacy. Statewide comprehensive outdoor recre- ation plan 1992-1996. Anchorage, Alaska. 80 pages. THE CANADIAN FIELD-NATURALIST Vol. 118 Birkhead, T. R., S. F. Eden, K. Clarkson, S. F. Goodburn, and J. Pelatt. 1986. Social organization of a population of Magpies, Pica pica. Ardea 74: 59-68. Dale, R. F. 1956. The climate of the Matanuska Valley. U. S. Weather Bureau, Technical Paper Number 27. Washington, D. C. 26 pages. De Smet, K. D. 1987. Organochlorines, predators and repro- ductive success of the Red-necked Grebe in southern Manitoba. Condor 89: 460-467. Groves, D. J., B. Conant, R. J. King, J. I. Hodges, and J. G. King. 1996. Status and trends of loon populations sum- mering in Alaska, 1971-1993. Condor 98: 189-195. Heimberger, M., D. Euler, and J. Barr. 1983. The impact of cottage development on Common Loon reproductive success in central Ontario. Wilson Bulletin 95: 431-439. McIntyre, J. W. 1978. The Common Loon: Part I. Popu- lation in Itasca State Park, Minnesota, 1957-1976. Loon 50: 38-44. McIntyre, J. W. 1988. The Common Loon: Spirit of northern lakes. University of Minnesota Press, Minneapolis. 228 pages. Newton, I. 1998. Population limitation in birds. Academic Press Inc., San Diego, California. 597 pages. Riske, M. E. 1976. Environmental and human impacts on grebes breeding in central Alberta. Ph. D. thesis, University of Calgary. 482 pages. Ruggles, A. K. 1991. Habitat selection by common loons (Gavia immer) in the Matanuska—Susitna Valley, Alaska. M. S. thesis, University of Alaska, Fairbanks. 98 pages. Smith, S. M. 1976. Ecological aspects of dominance hier- archies in Black-capped Chickadees. Auk 93: 95-107. Stout, B. E., and G. L. Neuchterlein. 1999. Red-necked Grebe (Podiceps grisegena). In The Birds of North Amer- ica, 465. Edited by A. Poole and F. Gill, The Academy of Sciences, Philadelphia, Pennsylvania, and The American Ornithologists’ Union, Washington, D. C. 31 pages. Titus, J. R., and L. W. VanDruff. 1981. Response of the Common Loon to recreational pressure in the Boundary Waters Canoe Area, northeastern Minnesota. Wildlife Monographs 79:1-60. Received 30 December 2002 Accepted 2 November 2004 The Barred Owl, Strix varia in Alberta: Distribution and Status LISA TAKATS PRIESTLEY Beaverhill Bird Observatory, Box 1418, Edmonton, Alberta T6B 2X3; e-mail: lisa@beaverhillbirds.com Priestley, Lisa Takats. 2004 The Barred Owl, Strix varia, in Alberta: Distribution and Status. Canadian Field-Naturalist 118(2): 215-224. Barred Owl distribution and status in Alberta were investigated using over 300 individual records (1912 through 1999) that were collected from literature, museum/zoo specimens, nest cards, bird surveys, volunteer raptor banders, and naturalists. Barred Owls were distributed throughout much of the boreal forest, aspen parkland, foothill, and mountain ecoregions of Alberta. Fifty-four breeding records (46 nests) were found. Barred Owls were associated with older forests and had a clumped distri- bution, predominantly along water where larger Balsam Poplar trees provide nesting sites. According to Alberta’s “Status of Alberta Wildlife 2000” criteria, the Barred Owl should be assessed as Sensitive. There is no evidence that Barred Owls have expanded their range in Alberta in the last 100 years; rather, they have maintained their distribution. Key Words: Barred Owl, Strix varia, distribution, status, Alberta, volunteer surveys, nests. The Barred Owl, Strix varia, is widely distributed throughout North America, ranging from the east coast to the western Canadian provinces (American Ornithologists’ Union 1998; Johnsgard 1988; Mazur and James 2000). It is found from the southern tip of Florida to southeastern Alaska. In Canada, the Barred Owl is a permanent resident in woodlands of British Columbia, through central and western Alberta, central Saskatchewan, south-central and southeastern Mani- toba, central and southern Ontario, southern Quebec, New Brunswick, Prince Edward Island, and the Mari- time provinces (Godfrey 1986). There is little pub- lished historical information available on the Barred Owl in western Canada, except for British Columbia (Boxall and Stepney 1982). However, recent studies have found this owl to be more common in western Canada than originally believed (Campbell et al. 1990; Mazur and James 2000; Mazur et al. 1997; Takats 1998). The Barred Owl is still considered rare in northern regions of Alberta (McGillivray 1996). It was found in the boreal forest region north of Edmonton, in the foothills/montane forests of western Alberta, and in Jasper National Park. The detailed status and distri- bution of the Barred Owl, however, is poorly -docu- mented in the province (Boxall and Stepney 1982; Kirk and Hyslop 1998). There has been considerable debate over the theory that the Barred Owl has only recently extended its range into the western states, Canada, and Alaska, although their relative rarity in the west has been assumed from the paucity of histor- ical records (Shea 1973; Taylor and Forsman 1976; Leder and Walters 1980; Boxall and Stepney 1982; Sharp 1989; Houston and McGowan 1999). It has also been suggested that their numbers are increasing in the boreal forest due to their increasing tolerance of pre- dominantly coniferous forests (Boxall 1986). In many parts of its range, the Barred Owl is depend- ent on large areas of remote forest with mature and old growth trees for nesting, roosting, and foraging (Paris 1947; Elody 1983; Devereux and Mosher 1984; McGarigal and Fraser 1984; Elody and Sloan 1985; Allen 1987; Bosakowski et al. 1987; Johnson 1987; Dunbar et al. 1991; James 1993; James et al. 1995; Mazur et al. 1997; Mazur et al.1998). In the United States it has been reported to nest in interior portions of expansive, mature woodland (Allen 1987). The typical Barred Owl nest is in a cavity in a large living or dead tree or in the top of a broken snag. There are records of Barred Owls using more conspicuous plat- form nests (i.e., stick nests) built by squirrels or other large birds (Mazur et al. 1997), as well as ground nests (Robertson 1959; Postupalsky et al. 1997), but these are potentially less productive than cavity nests (Pos- tupalsky et al. 1997) and are generally quite rare. Alberta Environmental Protection (1996) placed the Barred Owl on the Yellow B list in the Status of Alberta Wildlife report. The Yellow B list includes species that are: (1) naturally rare but not in decline, (2) naturally rare and have clumped breeding distribu- tions, or (3) associated with habitats or habitat elements that are, or may be, in decline. The status of Canada’s remaining old growth forests is of growing concern because of the high rate of harvest (Ellis 1993). In the year 2000, the Barred Ow] was placed on the Sensitive species list (Alberta Sustainable Resource Develop- ment 2000). The first step in managing wildlife populations is having knowledge of distribution and abundance (Mosher and Fuller 1996). The first objective of this paper was to compile all historic Barred Owl records (published and unpublished) for Alberta, in order to determine the past and present distribution in the prov- ince. The second, based on recent studies, was to evalu- ate this species’ status and general habitat use (as it relates to status) in Alberta. The third was to establish whether evidence exists to support the suggestion of range expansion into the west. Pas) 216 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 1: Barred Owls banded in Alberta: date, location, age, sex, and name of bander (1966-1999). Date Location Age-Sex* May 1966 Edmonton L-U April 1987 Water Valley SY-U November 1988 SSA Ste U-U April 1988 Deerland AHY-F June 1989 Deerland L-U May 1990 Deerland L-U May 1991 Niton L-U May 1991 Deerland L-U May 1992 Deerland L-U May 1994 Millarville AHY-F June 1994 Tawatina L-U June 1994 Uncas L-U June 1994 Calling Lake AHY-F June 1995 Solomon Creek AHY-F August 1996 Gregg Lake AHY-U May 1995 Water Valley AHY-F May 1995 Water Valley AHY-M May 1995 Uncas AHY-F May 1995 Uncas L-U May 1995 Tawatina L-U November 1995 Redwater AHY-M November 1995 Se lae2° U-U April 1995 Calling Lake AHY-M June 1995 Calling Lake AHY-F June 1996 Tawatina AHY-F June 1996 Tawatina L-U May 1996 Calling Lake AHY-M May 1996 Calling Lake AHY-M May 1996 Calling Lake AHY-F June 1996 Calling Lake AHY-M July 1996 Calling Lake AHY-M August 1996 Calling Lake AHY-M May 1997 Tawatina L-U May 1998 Calling River L-U May 1998 Calling Lake L-U June 1998 Uncas L-U June 1999 Uncas L-U June 1999 Vinca L-U * Age — L=nestling/fledgling, AH Y=after hatch year, SY=second year, Sex — U=unknown, M=male, Methods Information on Barred Owls in Alberta was col- lected from a variety of sources, which included: pub- lished literature, museum collections (National Museum of Canada, National Museum of Natural Sciences (now Canadian Museum of Nature), American Museum of Natural History, Cornell University Museum of Verte- brates, Royal Ontario Museum, University of Calgary Museum of Zoology, University of Alberta Museum of Zoology, Provincial Museum of Alberta, Alberta Breeding Bird Atlas (Federation of Alberta Natural- ists), Avian Raptor Nest Cards (Alberta Sustainable Resource Development/Beaverhill Bird Observatory 1999*), banding records (Canadian Wildlife Service Bird Banding Office), and personal communications with Provincial and National Parks personnel. Records Number Bander 3 E. Jones D. Collister Pletz/B. Gehlert Cromie Cromie Collister Cromie Cromie Stepnisky/G. Court Takats Takats Collister Collister Cromie Cromie Cromie Roper Halmazna Court Sissons/G. Court Cromie Cromie Olsen/R. Sissons Olsen/R. Sissons Olsen/R. Sissons Olsen/R. Sissons Olsen/R. Sissons Olsen/R. Sissons Cromie Cromie Cromie Cromie Cromie Pletz BWW PD FB DR OD EOD DBD i OD OD OD OD i i Fae Ragen re Ce ee ee eG eee OSs GS eee ee eae F=female from the following volunteer programs were also col- lected: Alberta Bird Records 1983-1988 (Alberta Or- nithological Records Committee), Breeding Bird Sur- veys, Christmas Bird Counts and May Species Counts, volunteer owl survey programs (Edmonton Ow! Prowl, Alberta Owl Prowl, Alberta Nocturnal Owl Survey), dead raptors turned in to Alberta Sustainable Resource Development, and unpublished data including field notes. All georeferenced Barred Owl sightings and nest locations were entered into the Biodiversity Species Observation Database (BSOD) (Alberta Sustainable Resource Development/Alberta Conservation Associ- ation 1998*). Maps comparing distributions from 1912 to 1975 and 1976 to 1999 were created in ArcView (ESRI GIS Mapping and Software). 2004 PRIESTLEY: BARRED OWL IN ALBERTA Ficure 1. Photo of a Barred Owl pausing before entering a typical nesting cavity (photo by Gordon Court). Results Of 320 reports of Barred Owls, 297 of these were entered into the BSOD database (23 records lacked information on date, location, and/or observer). The first record of a Barred Owl in Alberta was from a specimen collected in 1912, from the Calgary area, though this has been considered a mislabeled speci- men (Houston and McGowan 1999). The first live Barred Owl recorded in Alberta was one heard calling along the Athabasca River near Fort McMurray in 1934 (Preble 1941). Dead Owls Museum specimens, private collections, and dead owls turned in to Fish and Wildlife constituted 53 Barred Owl locations distributed throughout the prov- ince. The most southeasterly report was a dead owl turned in to Fish and Wildlife in February 1984 from Coronation (Boxall 1986). Morphological information collected on 26 owls shows that the majority were adults (24 of 26), 12 females, 10 males, 4 unknown sex. Boxall and Stepney (1982) reported an unusually high number of dead Barred Owls turned in to Fish and Wildlife during a short period of time (15 indi- viduals from 1982 to 1985). Banding, Literature and Personal Communications Barred Owls have been banded on 38 occasions in Alberta from 1966 to 1999 (CWS Banding Office) by thirteen banders (Table 1). There were 15 clutches of owls banded at nest sites, as well as nine adult females, eight adult males, and three adults of unknown sex. Only one Barred Owl was banded before 1987 (2.6 percent). : There were 42 reports of Barred Owls collected from publications (Preble 1941; Jones 1956; Jones 1966; Salt and Wilk 1958; Salt and Salt 1976; Francis and Lumbis 1959; Jones 1987; Sadler and Myers 1976; Boxall and Stepney 1982; Rintout and Myers 1983; Pintel et al. 1991; Takats 1995) and 63 records collect- ed from personal communications. Only 35 of these records were pre-1975 (33 percent). Jones (1987) sum- marized observations of Barred Owls as being “fairly evenly distributed in Alberta, particularly in Alberta’s northern forests and heavily forested foothill regions of Alberta”. Holroyd and VanTighem (1983) report the Barred Owl as a rare, year-round resident in Banff and Jasper National Parks. They state: “Tt occurs regularly near Jasper townsite especially along the lower Miette River, where it has nested, and near Cot- tonwood Slough (Roy Richards, pers. comm.) and Pyramid Lake (R. Salt, pers. comm.). It has been recorded twice near North Saskatchewan River Crossing (M. Dyer, pers. comm.), twice near Vermilion Lakes Banff, once at Lake Annette (K. VanTighem, pers. comm.) and once near Ma- ligne Canyon (J. Salt, pers. comm.).” Volunteer Surveys Breeding Bird Atlas volunteers reported 65 Barred Owl locations during a five-year survey from 1987 through 1991 (data were collected between February and July of each year). Atlas data showed that Barred Owls were concentrated in the Boreal Forest region, 218 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Barred Owl locations recorded on volunteer owl survey programs (EOP — Edmonton Owl Prowl, AOP — Alberta Owl Prowl, ANOS — Alberta Nocturnal Owl Survey). Date Location Observation Observer 24 February 1988 Big Lake 1 individual EOP-—_ Beck and Beck (1988) 5 March 1988 Cooking Lake 1 individual EOP— _ Beck and Beck (1988) 6 March 1988 Wabamun Creek 1 individual EOP— _ Beck and Beck (1988) 24 February 1989 Big Lake 1 individual AOP-—_ E. Bamford 05 March 1989 Cooking Lake 1 individual AOP—_ R. and M. Berg 06 March 1989 Wabamun Lake 1 individual AOP -—_ S. Jungkind, Belmonte, Gomez, Diener 21 May 1989 Water Valley 1 individual AOP— _ D. Hutchinson, D. Collister 10 April 1998 Lac La Biche pair duetting © ANOS — J. Gammon and P. Okrainec 15 May 1998 Shaw Lake 1 individual ANOS — J. Gammon 18 April 1998 Hillcrest Road 2 individuals ANOS — M. Heckbert and J. Doll 26 April 1998 Sibbald 1 individual ANOS — D. Woodsworth, Kanagawa, Mitchell 27 March 1999 Nojack South 1 individual ANOS — R. Gutsell, R. Wiacek, B. McCulloch and primarily coniferous foothills and montane forests west of Calgary and in Jasper National Park (Semen- chuk 1992). Few records indicated breeding evidence and only one possible breeding record was discovered, north of Lesser Slave Lake at La Crete along the Peace River. During Alberta Christmas Bird Counts from 1965 to 1998, volunteers in 44 count circles recorded 53 Barred Owls. Only 17 (32 percent) of these owls were recorded before 1990. Only four and two owls were recorded during Breeding Bird Surveys and May Species Counts, respectively. There have been three volunteer surveys conducted specifically for owls (Table 2): Edmonton Owl Prowl (1988), Alberta Owl Prowl (1989), and the volunteer Alberta Nocturnal Owl Survey (1998 and ongoing). Seven Barred Owls were found during the first two owl surveys (Beck and Beck 1988, personal commu- nication). Volunteer Alberta Nocturnal Owl surveyors found 13 individual Barred Owls (11 records) on elev- en 10 km transects along roads (16 routes were sur- veyed throughout the province). The Alberta Bird Record contributed 35 more locations of Barred Owls in 1988 (Alberta Ornithological Records Committee). Two intensive studies have been conducted on the Barred Owl in Alberta (Takats 1998; Olsen 1999). Takats (1998) conducted a study on distribution and abundance of the Barred Owl in the Foothills Model Forest (located in the foothills and mountains around Hinton and Jasper). Forty-two different territorial Barred Owls (10 females, 17 males, and 15 unknown sex) were recorded in 1995 and 1996. Of these, seven were paired and the other 28 were single but may have had mates that did not respond to broadcasts (Takats 1998). Surveys were continued in 1997 and 1998 and four additional Barred Owls were located. The density of Barred Owls was determined to be 0.05 and 0.04 owls/km?, in 1995 and 1996, respectively. The habitat used by Barred Owls at 45 calling loca- tions in the Foothills Model Forest was predominant- ly older mixedwood forest containing Trembling Aspen (Populus temuloides), White Spruce (Picea glauca) and Balsam Poplar (Populus balsamifera). Roosting and foraging occurred in a variety of stand types, but were also predominantly older mixedwood Populus sp. and spruce (Takats 1998). A study to evaluate the effects of forest fragmenta- tion on bird communities was initiated in north-central Alberta (Schmiegelow and Hannon 1993). Preliminary work located six Barred Owl territories, and evidence of three breeding pairs was discovered in the Calling Lake area (G. Court, personal communication). Two additional territories and four additional breeding sites were found in a study from 1996 through 1998 (Olsen 1999). During the breeding season, the density of Barred Owls was 0.04 pairs/km?. Owls were found to use old growth mixed forest stands (Olsen 1996). Breeding Records There were 54 breeding records (only three found before 1975). The first evidence of breeding was re- corded in 1949 (Grant 1966), but the first nest was not discovered until 1966, in Edmonton (Jones 1966) (Table 3). Boxall and Stepney (1982) reported eight breeding records, and Semenchuk (1992) reported six confirmed breeding records during the Provincial Breeding Bird Atlas Project (1987-1991). The north- ern-most breeding record was a nest found at 58° 19' latitude and 116° 17' longitude near La Crete (Takats 1995). Some local residents have reported Barred Owls to be present for over 20 years. Breeding records for Barred Owls were not com- mon (n = 54) and only 46 nests have been found (1966 through 1999): 38 (82.6 percent) were in natural cavi- ties (or bowls on top of dead trees that had broken off), two (4.3 percent) were in stick nests, one (2.2 percent) was in a man-made stick nest, and five (10.9 percent) were in nest boxes. In most cases owls nested in older mixedwood stands and used large diameter Populus sp. trees (> 40 cm) (primarily Balsam Poplar) for 2004 TABLE 3: Confirmed breeding records of Barred Owls for Alberta. Man-Made Platform Date Location Nest Type 1949 Lesser Slave Lake Unknown 1966 Edmonton Cavity Nest 1968* Edmonton Cavity Nest 1976* Blue Lake Cavity Nest LOTA* Jasper Cavity Nest 1977* Blue Lake Cavity Nest 1978 Chilver Lake Unknown IO Miette River Cavity Nest 1980* Miette River Cavity Nest 1984** Spruce Grove Cavity Nest 1988 Deerland Cavity Nest 1989 Deerland Cavity Nest 1990 Deerland Cavity Nest 199] Deerland Cavity Nest 199] Niton Cavity Nest 1992 Deerland Cavity Nest 1993 La Crete Cavity Nest 1994 La Crete Cavity Nest 1994 Millarville Unknown 1994 Uncas 1994 Tawatinaw Cavity Nest 1994 Elk Island Stick Nest 1994 Calling Lake Unknown 1994 Miette River Cavity Nest 1995 Lynx Creek Cavity Nest 1995 Miette River Cavity Nest 1995 La Crete Cavity Nest 1995 Tawatinaw Cavity Nest 1995 Cross Lake Cavity Nest 1995 Uncas Nest Box 1995 Calling Lake Cavity Nest 1995 Calling Lake 2 Cavity Nest 1996 Bragg Creek Cavity Nest 1996 Solomon Creek Cavity Nest 1996 Blackcat Ranch Cavity Nest 1996 Miette River Cavity Nest 1996 Tawatinaw Nest Box 1996 Calling Lake Unknown 1997 Cross Lake Cavity Nest 1997 Miette River Cavity Nest 1997 Tawatinaw Cavity Nest 1997 Lac La Biche Stick Nest 1997 Calling Lake Cavity Nest 1998 Uncas Cavity Nest 1998 Solomon Creek Cavity Nest 1998 Calling River Cavity Nest 1998 Calling Lake Cavity Nest 1999 Uncas Nest Box 1999 Tawatinaw Nest Box 999 Fe Vinca Cavity Nest 1999 Cross Lake Nest Box PRIESTLEY: BARRED OWL IN ALBERTA 219 Observer/Reference Grant (1966) Jones (1966) E. Jones (personal communication) Boxall and Stepney (1982) Boxall and Stepney (1982) Boxall and Stepney (1982) Wiseley (personal communication) Richards (personal communication) Richards (personal communication) R. Copeland (personal communication) R. Cromie (Nest Card) R. Cromie (Nest Card) R. Cromie (Nest Card) R. Cromie (personal communication) H. Pletz/B. Gehlert R. Cromie (Nest Card) Takats (1995) Takats (1995) D. Collister (personal communication) R. Cromie (Nest Card) R. Cromie (Nest Card) H. Pletz (personal communication) G. Court (personal communication) Takats (1998) Takats (1998) Takats (1998) A. Miller (personal communication) R. Cromie (Nest Card) R. Cromie (Nest Card) R. Cromie (Nest Card) G. Court (personal communication) G. Court (personal communication) H. Pletz (personal communication) Takats (1998) Takats (1998) Takats (1998) R. Cromie (Nest Card) Olsen (1999) A. Karvonen (personal communication) L. Takats (personal communication) R. Cromie (Nest Card) Olsen (1999) Olsen (1999) R. Cromie (personal communication) L. Takats (personal communication) T. Roper (Nest Card) Olsen/Cromie (Nest Card) R. Cromie (Nest Card) R. Cromie (Nest Card) H. Pletz (Nest Card) R. Cromie (Nest Card) * Boxall and Stepney (1982) ** Nest cut down, two eggs cracked, one survived, owl held at Valley Zoo, Edmonton *** One young Barred Owl was fostered in to this nest. nesting (Takats 1998; Avian Raptor Nest Cards; Olsen 1999). Nest boxes built specifically for Barred Owls were readily taken over in areas where natural cavities were not abundant (G. Court, personal communica- tion; R. Cromie, personal communication). Distribution Most occurrences of Barred Owls were in the boreal forest, foothill, and mountain ecoregions from 1912 through 1974 (Figure 2) and from 1975 to the present (Figure 3). Few owls were recorded in the parkland, 220 THE CANADIAN FIELD-NATURALIST Vol. 118 Barred Owl Occurrences 1912-1974 Medicine Hat FIGURE 2. Barred Owl distribution from 1912 through 1974 (map courtesy of Alberta Conservation Association/Alberta Sus- tainable Resource Development). NR i) 2004. PRIESTLEY: BARRED OWL IN ALBERTA Barred Owl Occurrences 1975-1999 FiGuRE 3. Barred Owl distribution from 1975 through 1999 (map courtesy of Alberta Conservation Association/ Alberta Sustainable Resource Development). 222 and only one individual was recorded in the northern limit of the grassland ecoregion. Detection of Barred Owls appears to have increased after 1975, but there have been only two records outside the range of the early distribution map. Discussion The Barred Owl is distributed over much of Alberta’s forested area. Evidence suggests that Barred Owl dis- tribution in the province has changed little over the last 100 years. As more detailed studies are conducted, as there is an increase in the number of interested naturalists exploring isolated woodlands, and as there is increased access to remote areas, more reports of Barred Owls will accumulate, Oeming (1957) and Jones (1966) concur. Mazur et al. (2000) suggested that Barred Owls are shy of humans, which may account for their secretive nature. In British Columbia the first records of Barred Owls were from Liard Crossing in 1943 and Nechako Low- lands in 1946 (Campbell et al. 1990). Scotter et al. (1985) reported a Barred Owl record from 1977 in the South Nahanni River, Northwest Territories, which shows that Barred Owls have existed northwest of Alberta for over 20 years. As well, the idea that the Barred Owl is adapting to the coniferous boreal forest does not hold true. Barred Owls have shown a clumped distribution in areas that contain large deciduous trees (predominantly riparian areas). They use Balsam Pop- lars for nesting and rely on a more open subcanopy for flight, although the roost cover provided by White Spruce trees is important, particularly in the winter (Takats 1998). Some of the first records of Barred Owls occurred along riparian areas. The Sensitive designation assigned by Alberta Sus- tainable Resource Development (2000) is well founded. Based on recent studies and personal communications with raptor banders and naturalists, the Barred Owl should not be considered rare in Alberta; however, this species does have a clumped breeding distribu- tion. As well, these owls rely on cavities in old, large diameter Populus sp. trees for nesting, and select old and/or mature mixedwood forests (a habitat that is in decline) to fulfill their life requisites (Takats 1998; Olsen et al. 1996*; Olsen 1999; Mazur and James 2000). As older forests are usually targeted first for har- vest, the amount of old growth forest remaining de- creases and stands become increasingly fragmented. Loss of nesting, roosting, and foraging habitat occurs when a forest is clearcut; moreover, Great Horned Owls, Bubo virginianus, are known to favour frag- mented landscapes. There is direct conflict between these two owl species, with the Barred Owl losing out to the larger Great Horned Owl (Bent 1961; Bosa- kowski 1994; Laidig and Dobkin 1995; Takats 1998 and field notes; Olsen 1999; G. Court, personal com- munication). Barred Owl numbers and range are lim- ited by the amount of adequate nesting habitat avail- able where they can reproduce successfully and fledge THE CANADIAN FIELD-NATURALIST Vol. 118 their young without interference from competitors and predators. Acknowledgments Funding for parts of this work was provided by Foothills Model Forest, Alberta Sport, Recreation, Parks and Wildlife Foundation, Alberta Sustainable Resource Development (Fish and Wildlife), Environ- ment Canada (Canadian Wildlife Service), Beaverhill Bird Observatory, and Weldwood of Canada, Hinton. The Bird Banding Office (Canadian Wildlife Service) provided Barred Owl banding information for Alberta. The Federation of Alberta Naturalists provided loca- tions from the Breeding Bird Atlas. The Alberta Con- servation Association/Alberta Sustainable Resource Development provided use of the Biodiversity Species Observation Database (BSOD) to create maps of the distributions. I thank Tony Erskine, Jim Duncan, and Bob Nero for reviewing the submitted manuscript. Gordon Court, Jim Beck, and Geoff Holroyd initially reviewed the manuscript, provided owl location information, and continue to encourage me with my work in raptor ecology. Lance Engley (Alberta Conservation Asso- ciation) provided the maps. Chuck Priestley, Dawn Birn, Kris Kendell, and Jeff Adamyk also provided some editorial comments. Jim Duncan and Kurt Mazur provided the idea to write the paper in the first place. I acknowledge Jim and Barb Beck (the ‘Owl Nuts’) for providing the Edmonton and Alberta Owl Prowl data as well as many other locations. Edgar Jones, Doug Collister, Ray Cromie, Wayne Smith, Hardy Pletz, Bob Gehlert, Ben Olsen, Trevor Roper, and Rick Lauzon (AXYS) provided many owl locations. Thanks also to Wayne Roberts (University of Alberta Museum of Zoology), Michel Gosselin (National Museum of Canada [now Canadian Museum of Na- ture), Mark Steinhilber (Provincial Museum of Alber- ta), and Rusty Copeland (Valley Zoo, Edmonton) for providing information on museum and zoo specimens. Thanks to Howard Troughton for providing the Christ- mas Bird Count information in a ready-to-use file. Special thanks to Stephen Glendinning and Jeff Adamyk, who spent many hours conducting broad- cast surveys for owls in the foothills. Finally, thank you to all the volunteer raptor banders, owl surveyors, and raptor enthusiasts who have contributed most of the information we know about Barred Owls in Alberta. Documents Cited [marked * in text] Alberta Environmental Protection/Alberta Conservation Association. 1998. Biodiversity species observation data- base. 7 floor O. S. Longman Building, 6909-116 Street, Edmonton, Alberta. Alberta Sustainable Resource Development/Beaverhill Bird Observatory. 1999. Avian raptor nest record cards. Edmonton, Alberta. Olsen, B., L. Takats, B. Beck, and J. Beck. 1996. Barred Owl (Strix varia) breeding habitat: draft habitat suitability index model. /n Habitat Suitability Index Models for 35 Wildlife 2004 Species in the Boreal Foothills of West-Central Alberta. Interim Report, Foothills Model Forest, Hinton. 11 pages. 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THE CANADIAN FIELD-NATURALIST Vol. 118 Salt, W. R., and J. R. Salt. 1976. The birds of Alberta, 3" edition. Hurtig Publishers, Edmonton. Salt, W. R., and A. L. Wilk. 1958. The birds of Alberta. Queen’s Printer, Edmonton. Schmiegelow, F. K. A., and S. J. Hannon. 1993. Adaptive management, adaptive science and the effects of forest fragmentation on boreal birds in northern Alberta. Pages 584-598 in Transactions of the 58" North American Wildlife and Natural Resources Conference. Scotter, G. W., L. N. Carbyn, W. P. Neily, and J. D. Henry. 1985. Birds of Nahanni National Park, Northwest Terri- tories. Saskatchewan Natural History Society, Regina, Saskatchewan. Semenchuk, G. P. Editor. 1992. The atlas of breeding birds of Alberta. Federation of Alberta Naturalists, Edmonton, Alberta. 371 pages. Sharp, D. U. 1989. Range extension of the Barred Owl in western Washington and the first breeding record on the Olympic Peninsula. Raptor Research 23: 179-180. Shea, D. S. 1973. Barred Owl records in western Montana. The Condor 76: 222. Takats, D. L. 1995. Rare bird sightings in the boreal forest of northern Alberta. Alberta Naturalist 25: 79-81. Takats, D. L. 1998. Barred Owl habitat use and distribution in the Foothills Model Forest. MSc. thesis, Department of Renewable Resources, University of Alberta, Edmonton, Alberta. 139 pages. Taylor, A. L., and E. D. Forsman. 1976. Recent range exten- sions of the Barred Owl in western North America, in- cluding the first records for Oregon. Condor 78: 560-561. Received 30 July 2002 Accepted 22 June 2004 Use of Eelgrass, Zostera marina, Wrack by Three Species of Ladybird Beetles (Coleoptera: Coccinellidae) in Prince Edward Island DaviD J. GARBARY, SARAH FRASER, CARRIE FERGUSON, and RANDOLPH F. LAUFF Department of Biology, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2WS5 Canada Garbary, David J., Sarah Fraser, Carrie Ferguson, Randolph F. Lauff. 2004. Use of Eelgrass, Zostera marina, wrack by three species of ladybird beetles (Coleoptera: Coccinellidae) in Prince Edward Island. Canadian Field-Naturalist 118(2): 225-228. Large numbers of the introduced ladybird beetle, Coccinella septempunctata L., were present at Wood Islands and Green Point, Prince Edward Island, in wrack consisting primarily of Zostera marina L. (Eelgrass). The wrack occurred in a 0.5 to 1.0 m band parallel to the shore, and was from five to 25 cm thick. The other ladybirds, Propylea quatuordecimpunctata (L.), an introduced coccinellid also found in high numbers, and an individual of the native Hippodamia tredecimpunctata (Say) were found only at Wood Islands. At both sites the ladybird beetles occurred in the mid-intertidal zone along at least 100 m of shoreline, and were absent to rare on the terrestrial vegetation above the high tide mark. At four of the other eight sites surveyed, occasional individuals were present in the wrack, but they were no more abundant than could be observed on landward vegetation. Mean densities of C. septempunctata at the two primary sites were 52 m? (Green Point) and 410 m7? (Wood Islands). Key Words: Coccinella septempunctata, Propylea quatuordecimpunctata, Hippodamia tredecimpunctata, ladybird beetles, Coccinellidae, Zostera marina, Eelgrass, intertidal zone, Prince Edward Island. With over 150 species and subspecies, the Coc- cinellidae of Canada and Alaska are a conspicuous and ecologically important element of the terrestrial biota (McNamara 1991). In addition to the native fauna, there is great interest in the distribution of invasive ladybirds and the subsequent loss of native biodiversity (Gordon and Vandenberg 1991). This is also true in eastern Canada where Hoebeke and Wheel- er (1996) and McCorquodale (1998) and Majka and McCorquodale (in press) reported the spread of intro- duced species in the Maritimes. Despite being a terrestrial family, there are several reports of Coccinellidae from beaches and salt marshes in both freshwater and marine environments (Davis and Gray 1966; Schaefer et al. 1987; Turnock 1996; Pupedis 1997; review by Nalepa et al. 1998). Here we report on the mass occurrence of living ladybird beetles in the marine intertidal of Prince Edward Island and their association with Eelgrass (Zostera marina) wrack. Materials and Methods Ten intertidal locations on Prince Edward Island (Table 1) were visited on 30-31 August 2002 for the examination of Eelgrass wrack as part of a study on Eelgrass decline. An abundant population of ladybird beetles was at site one (Wood Islands), and several hours were spent photographing and quantifying the assem- blage. At subsequent sites three individuals inspected the wrack for a minimum of 15 min along at least 100 m of shoreline. If only a few beetles were observed, their presence was noted. If many were present, densities were calculated using a 20 x 20 cm quadrat. The quad- rat was placed haphazardly on the eelgrass wrack (n = 25) at irregular intervals on the shore. Characteristics of the primary study sites were: The Wood Islands site was a sandy beach with sandstone outcrops. The primary species of seaweeds associated with the wrack were recorded. Green Point was a salt marsh adjacent to an abandoned wharf, and common plants in the marsh were recorded. At Green Point the wrack was almost exclusively Eelgrass. Characteris- tics of other sites are mentioned, where necessary, in the body of the text. The weather on both days was mostly bright and sunny; however, a mild rain fell at Marchwater (1800 h, 30 August) and Belmont Prov- incial Park was extremely windy (0800 h, 31 August). Coordinates of primary sites were determined using a global positioning system (Garmin GPS 12, Olanthe, Kansas; Table 1). Beetles were identified using the keys in Gordon (1985) and by comparison with specimens in the in- sect collection at St. Francis Xavier University. Voucher specimens have been deposited in the Herbarium of St. Francis Xavier University (STFX) and in the insect collection. Results Three species of ladybird beetles, Coccinella sep- tempunctata, Hippodamia tredecimpunctata, and Prop- ylea quatuordecimpunctata, were present in intertidal Eelgrass wrack (Zostera marina) in Prince Edward Island during late August. All species were present at Wood Island; however, only C. septempunctata was collected from the other sites. PIT) 226 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 1. Relative abundance of P. quatuordecimpunctata, H. tredecimpunctata and C. septempunctata in sites across Prince Edward Island, and their corresponding GPS coordinates. “Low” corresponds with an observation of < 5 individuals. Site Coordinates Wood Islands 45°56'N 62°45'W St. Peters Bay 46°26'N 62°28'W March Water 46°29'N 63°44'W Belmont Provincial Park 46°31'N 63°49'W Green Point 46°35'N 63°52'W Cascumpec Bay 46°45'N 64°04'W Linkletter Provincial Park 46°24'N 63°51'W Victoria Provincial Park 46°12'N 63°30'W West River Bridge 46°11'N 63°14'W Pinnette Provincial Park 46°04'N 62°54'W At Wood Islands, C. septempunctata occurred pri- marily in a loose to dense band of Eelgrass wrack, 0.2-1.0 m wide, that formed in the mid-intertidal zone on a sandy beach (Figure 1). No coccinellid beetles were found in the dry wrack in the splash zone nor in the mixed herbaceous vegetation above the high tide mark. A few beetles were also found in small clumps of wrack lower in the intertidal zone that had recently been inundated by the incoming tide and very gentle wave action. The wrack consisted mostly of leaves of Eelgrass and was mixed with several com- mon seaweeds including the brown algae, Fucus ser- ratus, Chorda filum, Chordaria flagelliformis, and the red algae, Chondrus crispus (Irish moss), Palmaria palmata (Dulse) and Furcellaria lumbricalis. The Eelgrass was relatively fresh and had numerous green leaves. The beetles were present on the exposed wrack to about two cm into the wrack mass in loosely packed leaves (Figure 2). Beetle density was 410 + 340 m? (mean + SD). Beetles were absent deep within the 20-30 cm thick wrack bundles. The beetles walked along the surface of the leaves and were single or in clumps of two-five individuals. Occasional beetles were present on the bare sand adjacent to the wrack or on clumps of seaweed (mostly Chorda filum and Fucus serratus) that separated from the main body of wrack. No flying beetles were observed; however, walking beetles occasionally spread their elytra. At Green Point, the Eelgrass wrack accumulated in a 0.5 —2 m band in the mid-intertidal zone of a salt marsh. This was a typical marsh dominated by Spar- tina alterniflora in the lower marsh and a mixture of Abundance Very high Absent Absent Absent High Absent Low Low Low Low S. patens, Scirpus americanus, S. maritimus, Glaux maritimus, Triglochin maritima and Limonium nashii in the mid-intertidal zone. At various sites along the shore there was extensive accumulation of old Eelgrass in the upper intertidal zone. The Eelgrass wrack tend- ed to form a blanket that matted down the surround- ing vegetation or occasionally was suspended up to 30 cm above the ground. The beetles (all C. septempunctata) were found on the Eelgrass in the mid-intertidal zone, with rare indi- viduals on the surrounding grasses and sedges. The insects were common (52 + 8.5 m7) over more than 100 m of shoreline, and were typically single. Beetles were absent on the upper intertidal wrack, and none were observed in the lower intertidal zone. The insects walked on the Eelgrass and did not fly. In addition to the ladybird beetles, one Leptinotarsa decemlineata (Colorado Potato Beetle) was observed on the wrack. At Linkletter and Pinnette Provincial Parks and West River Bridge, 3-5 beetles (all C. septempunctata) were found on Eelgrass in the upper intertidal zone. These sites were primarily sand beaches with either scattered clumps or continuous carpets of Eelgrass up to 2 m wide in the high intertidal zone. At St. Peters Bay, Belmont and Victoria Provincial Parks and Cas- umpec Bay, beetles were absent, despite the presence of extensive Eelgrass wrack. Upon our return to Wood Islands, 28 h after the initial observations, we found that the previous day’s wrack had largely disappeared. A few hundred dead ladybird beetles were present in the remaining wrack and on sandstone outcrops along with a few live individuals. 2004 GARBARY, FRASER, FERGUSON, and LAUFF: USE OF EELGRASS BY LADYBIRD BEETLES Ficure |. Ladybird beetle site at Wood Islands, Prince Edward Island, showing band of intertidal wrack on beach with investi- gator (C.F.) examining Eelgrass. Discussion Although the records for both introduced species (C. septempunctata, P. quatuordecimpunctata) do not represent range extensions, it is disturbing that they did make up essentially all of the coccinellid fauna at the sampled sites. P. quatuordecimpunctata has been recorded in PEI since at least 1994 (Hoebeke and Wheeler 1996). The first records for C. septempunc- tata on PEI (Charlottetown) are from 1982; it is now abundant in the province (Majka and McCorquodale; in press). The two sites where C. septempunctata was abundant shared the feature of having abundant Eel- grass wrack in the mid-intertidal zone. The beetles were generally not found in the older, dry wrack at the upper part of the shore nor in the vegetation above the high tide mark (Wood Islands and Green Point), and the surrounding salt marsh vegetation (Green Point). Examination of the Eelgrass substratum showed no conspicuous populations of invertebrates that might be a suitable food source. In addition, the beetles were sluggish, suggesting exhaustion due to struggling or partial suffocation. The location of the beetles and their behaviour suggest that the they were only recently washed up to the eelgrass wrack, a substrate which allowed them firm footing. Sites with few or no beetles had less fresh Eelgrass wrack. The intertidal substra- tum at these sites tended to be rockier, with better developed populations of seaweed. Although terrestrial, both C. septempunctata and Propylea quatuordecimpunctata have previously been found in coastal habitats including salt marshes in Connecticut, North Carolina and Delaware (Schaefer et al. 1987; Pupedis 1997; Nalepa et al. 1998). Davis and Gray (1966) also reported another ladybird beetle, Naemia serriata, from a salt marsh in North Caro- lina. The mass occurrences of C. septempunctata that we found are different from that in Delaware in which numerous dead and some living individuals were washed ashore following deposition in the ocean and association with seaweed wrack (Schaefer et al. a 2. Coccinella septempunctata on mid-intertidal Eelgrass wrack. Note: The white spots on the leaves are a calcified red alga. FIGURE 228 1987). The North Carolina mass occurrence is similar to that reported by Schaefer et al., and involved mass mortality and shore deposition (Nalepa et al. 1998). Turnock (1996) suggested that accumulations of lady beetles on the shores of Lake Manitoba resulted from wave deposition and subsequent migration up the shore to structures protruding from the sand. Given the localization of C. septempunctata in this study, particularly at Green Point, the Eelgrass seems to have been a fortunate substratum on which the beetles could take hold, and rest prior to dispersal. Although there were conspicuous corpses of lady- bugs in the wrack at Wood Islands on the day after the survey, the vast majority of the population had dis- appeared from the shore. There was no evidence that the beetles had moved onto the grassy field adjacent to the beach. Invasions of Coccinellidae into North America show that these species can cover a large dis- tance (Schaefer et al. 1987). McCorquodale (1998) gives range expansion rates of 31-440 km y"! across North America for four introduced coccinellids in Nova Scotia, although he concedes that not all of this is likely to be long distance flights. Schaefer et al. (1987) comment on the occurrence of C. septempunc- tata on Sable Island, 300 km from mainland Nova Scotia. Thus the 20 km flight across the Northumber- land Strait to or from Nova Scotia is reasonable, espe- cially if a suitable wind is available. Our records of C. septempunctata on the north and south shores of PEI are consistent with eelgrass pro- viding a staging substratum during migration. Migra- tions of Hippodamia convergens cover hundreds of kilometres in single flights in California from San Francisco and the Imperial Valley to the Sierra Nevadas (Hodek 1973). However, the aggregations we observed in Prince Edward Island are more likely reflective of accidental downing of migrating populations as a result of weather conditions, followed by subsequent wash- up. The Eelgrass provided a better refugium than other substrata in the mid-intertidal zone. We suggest that the beetles are only saved by the Eelgrass wrack, and that there is no particular attraction to it as previously described for similar vegetation with C. septempunc- tata (Schaller and Nentwig 2000; Frantsevich and Zolotov 2001). Acknowledgments We thank Christine Nalepa and Chris Majka who provided helpful discussion and literature; the com- ments of two anonymous reviewers were also an asset to this paper. Barry Taylor kindly assisted with plant identification and David McCorquodale verified iden- tifications of beetles in our insect collection. This THE CANADIAN FIELD-NATURALIST Vol. 118 work was supported by a grant from the Natural Sci- ences and Engineering Research Council (NSERC) to D. J. G., funding from the Nova Scotia Museum to R. FE. L., and by NSERC undergraduate awards to S. F and C. F. Literature Cited Davis, L. V., and I. E. Gray. 1966. Zonal and seasonal dis- tribution of insects in North Carolina salt marshes. Eco- logical Monographs 36: 275-295. Frantsevich, L. I., and V. V. Zolotov. 2001. Response to contrast of the seven spot ladybird Coccinella septem- punctata (Coleoptera, Coccinellidae). Journal of Evolu- tionary Biochemistry and Physiology 37: 398-404. Gordon, R. D. 1985. The Coccinellidae (Coleoptera) of America north of Mexico. Journal of the New York Ento- mological Society 93: 1-912. Gordon, R. D., and N. Vandenberg. 1991. Field guide to recently introduced species of Coccinellidae (Coleoptera) in North America, with a revised key to North American genera of Coccinellini. Proceedings of the Entomological Society of Washington 93: 845-864. Hodek, I. 1973. Biology of Coccinellidae. Academia, Prague. 260 pages. Hoebeke, E. R., and A. G. Wheeler. 1996. Adventive lady beetles (Coleoptera: Coccinellidae) in the Canadian Mari- time Provinces, with new eastern U.S. records of Harmonia quadripunctata. Entomological News 107: 281-290. McCorquodale, D. B. 1998. Adventive lady beetles (Coleop- tera: Coccinellidae) in eastern Nova Scotia. Entomologi- cal News 109: 15-20. McNamara, J. 1991. Family Coccinellidae, ladybird beetles. Pages 229-237 in Checklist of beetles of Canada and Alaska. Edited by Y. Bousquet. Publication 1884/E. Agriculture Canada, Ottawa, Ontario. Nalepa, C. A., K. R. Ahlstrom, B. A. Nault, and J. L. Williams. 1998. Mass appearance of lady beetles (Coleop- tera: Coccinellidae) on North Carolina beaches. Entomo- logical News 109: 277-281. Pupedis, R. J. 1997. Aquatic insects of the West River and salt marshes of Connecticut. Bulletin Series, Yale School of Forestry and Environmental Studies 100: 162-177. Schaefer, P. W., R. J. Dysart, and H. B. Specht. 1987. North American distribution of Coccinella septempunctata (Coleoptera: Coccinellidae) and its mass appearance in coastal Delaware. Environmental Entomology 16: 368-373. Schaller, M., and W. Nentwig. 2000. Olfactory orientation of the seven-spot ladybird beetle, Coccinella septempunctata (Coleoptera: Coccinellidae): attraction of adults to plants and conspecific females. European Journal of Entomology OES = 59: Turnock, B. 1996. Lady beetles on the Lake Manitoba beach. University Field Station (Delta Marsh) Annual Report 31: 40-42. Received 17 September 2002 Accepted 15 November 2004 Predicting the Effects of Cerulean Warbler, Dendroica cerulea Management on Eastern Ontario Bird Species JASON JoNES!, WILLIAM J. MCLEISH, and RALEIGH J. ROBERTSON Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6 Canada | Present address, Department of Biology, Vassar College, Poughkeepsie, New York 12604 USA; e-mail: jajones @ vassar.edu Jones, Jason, William J. McLeish, and Raleigh J. Robertson. 2004. Predicting the effects of Cerulean Warbler, Dendroica cerulea, management on eastern Ontario bird species. Canadian Field-Naturalist 1 18(2): 229-234. Single-species habitat management strategies are often undertaken without explicit consideration of their effects on the larger community. Here we explore the potential effects of managing eastern Ontario deciduous forests for the Cerulean Warbler (Dendroica cerulea) by examining its potential as a biodiversity indicator species and as an umbrella species. Our results indicate that the Cerulean Warbler would not be an effective biodiversity indicator, as its distribution across the studied landscape did not coincide with areas of high avian species rich- ness. However, the Cerulean Warbler may be effective as an umbrella species for the maintenance of populations of other canopy-nesting species that require mature deciduous forest habitats. It is hoped that the conclusions reached in Ontario, while perhaps not directly transferable to all parts of the breeding range, encourage other Cerulean Warbler researchers to ask similar questions in their study areas. Key Words: Cerulean Warbler, Dendroica cerulea, conservation utility, biodiversity indicator species, umbrella species, eastern Ontario. Few North American songbirds are receiving the scientific and conservation attention that is currently focused on the Cerulean Warbler (Dendroica cerulea; Robbins et al. 1992; Rosenberg et al. 2002*; Hamel et al. 2004). The Cerulean Warbler has suffered signifi- cant long-term breeding population declines (annual declines of 3.04% over 1966-2000; Link and Sauer 2002) which are largely attributed to habitat destruction on both the breeding and wintering grounds (Robbins et al. 1992). Concern over the long-term health of this species has led to its designation as threatened, rare, or of special concern in the United States and as a Species of Special Concern in Canada (Robbins et al. 1992; Hamel 2000; COSEWIC 2003*). In two separate meeting in 2001 and 2002, a group of academic and governmental scientists, land mana- gers, industry biologists and non-governmental organi- zations formed the Cerulean Warbler Technical Group (CWTG): an effort to develop a proactive, broad-based, multiple stakeholder approach to Cerulean Warbler conservation in both North and Latin America (Hamel et al. 2004). One of the outcomes of these meetings was a breeding grounds research plan that is designed to identify key population limitation factors and ex- plore forest management options for the maintenance and creation of Cerulean Warbler habitat on a range- wide basis. Ironically, the very thing that limits our ability to currently diagnose specific reasons for breeding ground population declines — a general lack of natural and life- history information — may also limit the efficacy of the CWTG research plan. Here, we take advantage of data collected during one of the longest running Ceru- lean Warbler research programs in North America (dating to 1994; Oliarnyk and Robertson 1996; Jones et al. 2000, 2001, 2004; Jones and Robertson 2001: Barg et al. 2005) to make predictions regarding the potential effects of managing deciduous forests for Cerulean Warblers on sympatric bird species in eastern Ontario. Specifically, we ask two questions. One, will the promotion of Cerulean Warblers and Cerulean Warbler habitat promote the preservation of avian diversity? In other words, is the Cerulean Warbler a biodiversity indicator (sensu Landres et al. 1988)? Two, can the Cerulean Warbler act as an umbrella species for other bird species with similar life — and natural histories? The protection of the habitat of the umbrella species ideally results in the protection of the habitat of those species whose requirements are subsumed by those of the umbrella (Launer and Mur- phy 1994; Berger 1997; Simberloff 1998). Study Area and Methods This investigation was conducted at the Queen’s University Biological Station (QUBS), Ontario (44°34'N, 76°20'W), within the Great Lakes-St. Law- rence mixed forest region. Our study area was restricted to approximately 2 600 ha of research tracts managed by QUBS. The landscape in the area is dominated by mature, secondary-growth, lowland mixed deciduous forest, interspersed with rocky outcrops, marshes, lakes and abandoned agricultural fields. 229 230 Bird surveys. In 1997 and 1998, we surveyed birds on QUBS property using variable-circular-plot point counts (Reynolds et al. 1980; Jones et al. 2000). We surveyed 80 stations in 1997 (17 May — 20 June) and 67 stations in 1998 (21 May — 21 June); the 1998 sta- tions were a subset of those sampled in 1997. The point-count stations were located to maximize spatial coverage of the study area. Each point count was 10 min long. Each station was separated by at least 200 m to minimize the potential for double-counting individuals. Point counts were conducted between 0.5 hr before sunrise and 3 hr after sunrise EST in order to sample during peak song activity, and were only conducted under calm weather conditions. For our analyses we included birds detected within 100 m of the plot center. Probability of detection was similar for all species analysed, as the detection thresholds for all but one species (Black-and-white Warbler, Mniotilta varia, 92 m) were beyond 100 m (Jones unpublished data). Data from the first two visits per station each year were used in the analyses to facilitate comparison between years and because two visits are sufficient to confirm the presence or absence of Cerulean Warblers (Jones et al. 2000). Vegetation Surveys. We collected vegetation data at 59 point-count stations in 1997 at five circular sub- plots, each with a radius of 5 m. The first subplot was centered on the point count station center and the other four were located 50 m away in each of the cardinal directions. Within each of the subplots we counted the number of saplings [stems < 3.0 cm diameter at breast height (dbh)], measured the dbh of all stems 2 3.0 cm and grouped them into two size classes (3.0 — 15.0 cm dbh, 15.0 — 30.0 cm dbh). Using an imaginary | m radius cylinder projected upward from the forest floor, we estimated cover within 3 m height intervals from the ground to the top of the canopy. Total cover and per- cent cover of each woody plant species was estimated by eye in each height interval on a scale of 0 to 10 (0 = 0% cover, 10 = 100% cover). Two observers made all cover estimates; the two observers spent several days prior to data collection assuring that their cover estimates were within 10% of one another. For analysis we reduced these cover estimates to two variables: maximum cover below 6 m (understory cover) and maximum cover above 12 m (canopy cover). Each vegetation variable was averaged across subplots to describe the habitat of the point-count station. We measured only those vegetation variables thought to be important to Cerulean Warblers; the importance of these variables has been supported by subsequent re- search (Jones and Robertson 2001; Jones et al. 2001). We also restricted the number of vegetation variables for analytical reasons (see below). Data Analysis. In our analyses, we included only those species known to breed in our study area. We also excluded species that are not adequately sampled by diurnal point counts, such as colonial nesters (e.g., THE CANADIAN FIELD-NATURALIST Vol. 118 Barn Swallow, Hirundo rustica), nocturnal species (e.g., Whip-poor-will, Caprimulgus vociferus), waterfowl (e.g., Wood Duck, Aix sponsa), and waders (e.g., Great Blue Heron, Ardea herodias). (1) Biodiversity indicator evaluation We tested whether the presence of Cerulean War- blers was a predictor of overall bird species richness. For the purposes of these analyses, we defined spe- cies richness as the number of species detected in the first two visits to a point-count station, excluding the Cerulean Warbler if present. We used randomization tests to compare the mean species richness at stations where Cerulean Warblers were present to the expect- ed species richness at a randomly generated sample of points (Chase et al. 2000). In these tests, the mean species richness was calculated for a random sample of point-count stations, with the number of random stations equaling the number of stations where Ceru- lean Warblers were detected in a given year. We iter- ated this procedure 1000 times to generate an expected distribution of mean species-richness values. We then compared the observed species richness associated with Cerulean Warblers and determined its statistical significance. These randomization tests were performed using S-PLUS 4.0 (Mathsoft 1997*). Values reported in the results are means + SE. (2) Umbrella species evaluation The initial step in evaluating the potential of the Cerulean Warbler as an umbrella species was to estab- lish an ecological context. We categorized the bird species detected during our surveys into species groups based on habitat preferences, diet and foraging sub- strate, and nesting substrate. These classifications were based on observations reported in the literature (Ehr- lich et al. 1988; Robbins et al. 1989a; Freemark and Collins 1992; Canterbury et al. 2000); we did not in- clude our survey data in these classifications. In addi- tion, we created a conservation concern grouping that included species which were experiencing statistically significant population declines as indexed by North American Breeding Bird Survey data for 1966-2000 (Sauer et al. 2001*). For the purposes of these analyses we focused on the groups to which Cerulean Warblers belonged: mature forest habitat (n = 13 species), insect- foliage foragers (n = 15), canopy_nesters (n = 12), and species of concern (n = 14). We used two methods to test if the distribution of Cerulean Warblers was representative of the distribu- tions of other species in the same functional group. First, we used the checkerboard score (C-score) devel- oped by Stone and Roberts (1990) to test for non- randomness in presence-absence matrices. One of the reasons we selected this metric of co-occurrence is that is not particularly prone to Type 1 error (Gotelli and Entsminger 2000*) and, unlike other co-occurrence metrics, it allows for overlap in species distributions (Gotelli and McCabe 2002; Feeley 2003). We calcu- lated C-scores (hereafter, observed C-score) for each 2004 JONES, MCLEISH, and ROBERTSON: CERULEAN WARBLER MANAGEMENT TABLE |. Interpretation of the principal components axes from analysis of 5 vegetation variables for 59 point-count stations. Bold-face eigenvalues significant following bootstrap analyses. Axis cvi Ce Wes SPD‘ SSD* LSD! Interpretation of positive axis values PCl 29.2 0.6249 -(.6281 -0.2509 0.0187 0.3195 mature forest with dense canopy REZ 54.1 0.2676 0.2143 0.7024 0.5205 0.3438 mid-succession forest PC3 74.2 0.0036 0.3208 0.1185 -0.7046 0.6218 mid- to late-succession forest PC4 88.5 -0.4737 0.0870 -0.4715 0.4648 0.5742 mature forest with patchy canopy PES 100.0 0.5599 0.6702 -0.4553 0.1280 = -0.1171 early succession forest ‘Cumulative variance explained; "Canopy cover (% cover > 12.0 m); ‘Understory cover (% cover < 6.0 m); ‘Sapling density (stems/m7); ‘Stem density (stems/m7) 3.0-15.0 cm dbh; ‘Stem density 15.0-30.0 cm dbh. functional group to examine if species within each functional group were distributed randomly across the landscape with respect to one another. For each survey year, we used re-sampling techniques to calculate 10 000 C-scores based on the original presence-absence matrix. The observed C-score was then compared with the generated distribution of expected C-scores. A C- score significantly greater than expected indicates that the assemblage is competitively structured; that is, individual species have distinct, and often exclusive, distributions (Stone and Roberts 1990). Conversely, a C-score significantly smaller that expected indicates that there is a degree of cohesion in the distribution patterns of the species included in the matrix. All C- score calculations were carried out using EcoSim 5.0 (Gotelli and Entsminger 2000). For all iterations, the number of species detected at each point-count station was kept consistent with the original matrix and each station was equally likely to be inhabited by a given species. Survey results from 1997 and 1998 were ana- lyzed separately. In our second test, we used logistic regression ana- lyses to predict the probability of occurrence along a habitat gradient for all species within each species group to which Cerulean Warblers belonged. The Red-eyed Vireo (Vireo olivaceus) was not included in these analyses due to its near-ubiquitous distribution. For these analyses we included only the 1997 surveys of the 59 point-count stations for which we collected vegetation data. We generated the habitat gradient by entering all five vegetation variables into a principal components analysis, giving us an approximate 10:1 ratio of sites to variables. We tested the significance of the eigenvalues for each variable within each com- ponent using a bootstrap approach, following the recommendations of Peres-Neto et al. (2003). We drew 1000 bootstrap samples by resampling entire rows with replacement, thereby ensuring that the bootstrap matrices had the same dimensions as the original veg- etation matrix; we conducted a PCA on each of these 1000 matrices. P-values were estimated by the number of bootstrap loadings equal to or less than zero for original loadings that were positive (greater than or equal to zero for negative loadings), divided by 1000. For the purposes of this paper, we considered P = 0.10 as our significance cut-off. The first PC axis (PC1) explained 29% of the variance of the vegetation data (Table 1); positive values along PC1 represented sites of mature deciduous forest and negative values repre- sented early successional forest. We included PC1 as the independent variable in our logistic regression ana- lyses. All these analyses were performed with JMP IN 4.0.2 (SAS Institute Inc. 2000*) and MS-EXCEL using the PopTools add-in module (Hood 2002). We calculated 95% confidence intervals around the logis- tic regression coefficient to facilitate comparison of occurrence probabilities across species. Results (1) Biodiversity indicator — The presence of Ceru- lean Warblers was not a significant predictor of avian species richness in either 1997 or 1998. In other words, observed species richness at Cerulean Warbler point- count stations was not significantly different from spe- cies richness expected by chance, based on bootstrap analyses (1997, observed richness 12.69 + 0.75 spe- cies, expected 13.95 + 0.03, P = 0.12; 1998, observed 9.75 + 0.63, expected 9.82 + 0.03, P = 0.83). (2) Umbrella species — All three of the species groups to which Cerulean Warblers belong exhibited cohesive distributions in both 1997 and 1998, as in- dexed by their C-scores (Table 2). C-scores that are significantly lower then expected are indicative of co- occurrence in the distribution patterns of the species in- cluded in the analyses. Species of conservation concern also exhibited cohesive distributions in both 1997 and 1998 (Table 2). The results of the logistic regression analyses using PC1 indicate that species within each of the species groups to which Cerulean Warblers belonged were distributed differentially along the generated habitat gradient, some more so than others (Table 3). Within the mature forest group, only one of the 12 group spe- cies (Black-and-white Warbler) did not overlap with the Cerulean Warbler confidence intervals. Similarly, only one of the 12 canopy nesting species (Blue Jay, Cyanocitta cristata) did not overlap with the Cerulean Warbler confidence intervals. Conversely, the insect- 232 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Tests for cohesive distributions of functional groups, as indexed by the C-score of Stone and Roberts (1990). Esti- mated values are mean + SE for a generated random distribution based on the actual presence-absence matrix for each group for each year. C-scores that are significantly lower then expected are indicative of co-occurrence in the distribution patterns of the species included in the analyses. 1997 Functional Group Observed Expected Mature forest 80.76 91.94 + 0.03 Insect-foliage 86.02 92.29 + 0.02 Canopy nesters 67.03 71.92 + 0.02 Species of concern 136.11 147.68 + 0.12 1998 P Observed Expected ie 0.0003 41.42 48.13 + 0.02 0.0017 0.0050 52.96 65.15 + 0.02 <0.0001 0.0078 36.63 40.90 + 0.02 0.0068 0.0049 73.32 85.04 + 0.03 0.0001 TABLE 3. Logistic regression coefficients and upper and lower 95 % confidence intervals (C. I.) predicting occurrence across a habitat gradient for members of the mature forest (MF), insect-foliage (IF), canopy nesting (CN) and conservation concern (CC) species groups detected in 1997. The confidence intervals of the species in bold face do not overlap with the confidence interval of the Cerulean Warbler. Species Functional Regression Lower Upper group coefficient Spy Vo Osill, 951701 11: Cerulean Warbler (Dendroica cerulea) all 0.519 0.121 0.917 American Redstart (Setophaga ruticilla) all 0.144 -0.299 0.587 Ovenbird (Seiurus aurocapilla) MF, CC 0.475 0.024 0.926 Black-and-white Warbler (Mniotilta varia) MEF -0.290 -0.645 0.065 Scarlet Tanager (Piranga olivacea) all 0.106 -0.239 0.451 Least Flycatcher (Empidonax minimus) MF, CN, CC 0.116 -0.290 0.522 Yellow-throated Vireo (Vireo flavifrons) MF, IF, CN 0.066 -0.381 0.513 Wood Thrush (Hylocichla mustelina) MF, CC -0.287 -0.791 0.217 Black-throated Green Warbler (Dendroica virens) MF, CN, CC 0.134 -0.272 0.540 Northern Waterthrush (Seiurus noveboracensis) MF -0.195 -0.777 0.387 Warbling Vireo (Vireo gilvus) IF -0.995 -1.936 -0.054 Ruffed Grouse (Bonasa umbellus) MF -0.140 -0.620 0.340 Veery (Catharus fuscescens) MF, CC -0.622 -1.786 0.542 Yellow-rumped Warbler (Dendroica coronata) MF, IF, CN 0.009 -0.483 0.501 Common Yellowthroat (Geothlypis trichas) IEXEE -0.192 -0.555 0.171 Golden-winged Warbler (Vermivora chrysoptera) IEVEGE -0.534 -0.955 -0.113 Indigo Bunting (Passerina cyanea) TENCE -0.535 -0.974 -0.096 Chestnut-sided Warbler (Dendroica pensylvanica) IEF; Ee -0.705 -1.438 0.028 Yellow Warbler (Dendroica petechia) IF -0.232 -0.585 0.121 Black-capped Chickadee (Poecile atricapillus) IF 0.058 -0.279 0.395 Rose-breasted Grosbeak (Pheucticus ludovicianus) Wee -0.725 -1.270 -0.180 Baltimore Oriole (Icterus galbula) LE ENEC 0.281 -0.217 0.779 Blue-grey Gnatcatcher (Polioptila caerulea) IF, CN -0.279 -0.781 0.223 Cedar Waxwing (Bombycilla cedrorum) CN -0.066 -0.440 0.308 Blue Jay (Cyanocitta cristata) CN -0.873 -1.528 -0.218 Eastern Wood-Pewee (Contopus virens) CN, CC -0.202 -0.670 0.266 American Crow (Corvus brachyrhynchos) CN -0.304 -0.835 0.227 foliage and conservation concern groups exhibited less concordance with the Cerulean Warbler distribution relative to the habitat gradient — 43% (6 of 14) and 29% (4 of 14), respectively. Discussion Effective biodiversity indicators tend to be habitat specialists with wide geographic ranges; they also tend to have well-known natural histories (Caro and O’ Do- herty 1999). Despite fitting this profile (Hamel 2000; Oliarnyk 1996; Jones 2000; Barg 2002), our results indicate that the Cerulean Warbler would not be par- ticularly effective as an avian biodiversity indicator in eastern Ontario, as its distribution across the studied landscape did not coincide with areas of high avian species richness. Our co-occurrence and logistic regression results do suggest that the Cerulean Warbler is suited to a role as an umbrella species. Perhaps not surprisingly, the dis- tribution of the Cerulean Warbler was well matched to the distributions of other canopy nesters in mature deciduous forest (e.g., Scarlet Tanager, Piranga oli- vacea). Given the hierarchical nature of habitat selec- tion, broad habitat requirements are likely more impor- 2004 tant in determining species distributions than are spe- cies’ food and nesting requirements (Hutto 1985; Block and Brennan 1993); indeed, we found that the species grouped by diet were more variable in their distri- butions along our succession gradient than were the species grouped by habitat type or nest location. Small body size — and, as a consequence, small home range size (Barg et al. in press) — could limit the Cerulean Warbler’s umbrella suitability, as effective umbrella species tend to have large home ranges (Caro and O’Doherty 1999); however, the Cerulean Warbler’s apparent tendency to live in conspecific aggregations (Hamel 2000) potentially offsets any limitation im- posed by small body size and individual home range size. We do not mean to suggest that the Cerulean Warbler would be a better umbrella than other, more widespread, species such as the Scarlet Tanager. Rath- er, our conclusions suggest that, if habitat manage- ment is directed at Cerulean Warblers, other species will likely benefit. Our co-occurrence and logistic regression results highlight a potential management conflict between mature forest and shrubland species. Although our C- scores indicate significant patterns of co-occurrence within the conservation concern group, there was no overlap between the logistic regression confidence in- tervals of the Cerulean Warbler and shrubland species that are considered to be at risk, such as the Golden- winged Warbler (Vermivora chrysoptera). Given their disparate habitat requirements (i.e., forest edge vs. forest interior), management for Golden-winged War- blers will necessarily conflict with management aimed at maximizing Cerulean Warbler population health. One additional conservation role the Cerulean War- bler may fill, and may be already filling, is that of a flagship species: a species that attracts attention sup- port by virtue of its ‘charismatic’ nature (Simberloff 1998). The Cerulean Warbler is a Neotropical migrant songbird, a group of birds that has been in the conser- vation spotlight over the last 30 years (e.g., Robbins et al. 1989b). The Cerulean Warbler is a beautiful bird with an elusive nature that has a high profile due to extensive public education and activism (Rosenberg et al. 2002) and is highly valued by birders and orni- thologists alike. More recently, the Cerulean Warbler has achieved notoriety as the focus of a legal battle surrounding its candidacy for listing on the U.S. En- dangered Species Act (Ruley 2000). This attention has led to the development of the CWTG and has spurred a great deal of basic research and forest man- agement interest (Hamel et al. 2004). The apparent dependence of this species on large tracts of forest creates a possibility for the Cerulean Warbler to serve as a valuable symbol of the overall health of deciduous forests in eastern North America. Acknowledgments We thank J. Pither and R. Montgomerie for sta- tistical advice. J. Barg, A. Erskine and an anonymous JONES, MCLEISH, and ROBERTSON: CERULEAN WARBLER MANAGEMENT reviewer provided helpful comments on earlier drafts. F. Phelan, M. Phelan, F. Connor and the staff at the Queen’s University Biological Station provided logis- tical support. Funding was provided by B. and K. Jones, Wildlife Habitat Canada, the Eastern Ontario Model Forest Program and an NSERC operating grant to RJR. We also had the support of the World Wildlife Fund of Canada in the form of a MacNaughton Conservation Scholarship to JJ and grants from the Endangered Species Recovery Fund to RJR. This project is part of Natural Legacy 2000, a nationwide initiative in Canada to conserve wildlife and habitats in private and public. We gratefully acknowledge the support of the Government of Canada’s Millennium Partnership Fund. JJ was also supported by an NSERC scholarship, Queen’s Graduate Awards and the Queen’s Bracken Fellowship. Documents Cited [marked * in text] COSEWIC. 2003. COSEWIC Assessment and update status report on the Cerulean Warbler Dendroica cerulea in Canada. Report prepared by J. J. Barg, J. Jones, and R. J. Robertson. Committee on the Status of Endangered Wild- life in Canada. Ottawa, ON. Gotelli, N. J., and G. L. Entsminger. 2000. EcoSim: null models software for ecology. Version 5.0. Acquired Intel- ligence Inc. & Kesey-Bear. Hood, G. 2002. PopTools, Version 2.3. http://www.cse. csiro.au/CDG/poptools/. Mathsoft, Inc. 1997. S-Plus 4.0. Data Analysis Products Division, Mathsoft, Inc. Seattle, Washington. — Rosenberg, K. V., S. E. Barker, and R. W. 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Pages 549-562 in Ecology and conservation of Neotropical migrant landbirds. Edited by J. M. Hagan II and D. W. Johnston. Smithsonian Institution Press, Washington, D.C. Simberloff, D. 1998. Flagships, umbrellas, and keystones: is single-species management passé in the landscape era? Biological Conservation 83: 247-257. Stone, L., and A. Roberts. 1990. The checkerboard score and species distributions. Oecologia 85: 74-79. Received 22 October 2002 Accepted 21 July 2004 Parasite Prevalence in Dark-eyed Juncos, Junco hyemalis, Breeding at Different Elevations HEATHER BEARS Department of Zoology, University of British Columbia, 6270 University Boulevard, Vancouver, British Columbia V6T 174 Canada; e-mail: bears @zoology.ubc.ca Present address: Centre for Applied Conservation Research, 2424 Main Mall, Forest Sciences Building, University of British Columbia, Vancouver, British Columbia V6T 1Z4 Canada Bears, Heather. 2004. Parasite prevalence in Dark-eyed Juncos, Junco hyemalis, breeding at different elevations. Canadian Field-Naturalist 118(2): 235-238. During the summer of 2001, Dark-eyed Juncos (Junco hyemalis) were captured within the lowest (1000 m above sea level) and highest (2000 m asl) elevation extremes of their breeding range in Jasper National Park, Alberta. Blood samples were taken to identify parasite genera, and to test for differences in parasite prevalence among elevations. The most common parasites at either elevation were Haemoproteus spp., Leucocytozoon spp., and Trypanosoma spp. A significantly higher proportion of low- compared to high-elevation birds was infected by at least one of these, supporting the prediction that high-elevation habitats may be refuges from parasites. Key Words: Dark-eyed Juncos, Junco hyemalis, blood parasites, mountains, elevation, Alberta. Birds breeding at high elevations often experience a delayed date of reproductive onset, exposure to ex- treme weather, and scarce or sparsely distributed re- sources (Hamman et al. 1989; Landmann and Winding 1993; Kollinsky and Landmann 1996; Widmer 1999; Bears et al. 2003). Dark-eyed Juncos (Junco hyemalis) show a significant reduction in seasonal reproductive output with increasing breeding elevation from the montane valley (1000 m asl) to the subalpine-alpine treeline (2000 m asl) in Jasper National Park, Alberta (Bears 2002, Bears et al. 2003). Yet, high elevation habitats are not occupied by less competitive age or size classes of juncos, or by later arriving individuals (Bears 2002), all of which are traits that render juncos less successful competitors for territory (Cristol et al. 1990; Grasso et al. 1996). Further, inter-annual return of birds to their site of capture is high, and roughly equivalent among elevations (Bears 2002). Hence, it does not appear that juncos are “forced” to breed at high elevations due to intraspecific competitive exclu- sion from lower-elevation habitat. This led me to ex- plore the idea that high-elevation juncos are compen- sated for decreased seasonal reproductive output over their lifetimes by some unrecognized benefits of breed- ing at high elevations. In other mountain ranges, studies have shown that the blood parasites of birds, and the insect vectors that carry them, tend to be low at upper breeding elevations (Stabler et al. 1974; Braun et al. 1993). Thus, as part of a series of investigations into the benefits of high-elevation breeding, I tested whether the juncos have fewer parasites within higher- versus lower-elevation segments of their breeding range in Jasper. Methods Juncos were captured and monitored at eight 50 — 70 ha study sites in Jasper National Park (52°53'N, 118°3'W), Alberta, from 1 May to 20 August in 2000, and from 15 April to 20 August in 2001. Four sites were located near the lowest elevation within the Park (1000 — 1020 m asl), separated by 5-10 km. Four sites were at the highest elevation at which juncos breed within the park (1950-2100 m asl), and were separat- ed by > 18 km. All high-elevation sites had south or southest aspects. Morphological measurements taken from juncos in these sites (H. Bears, unpublished data) most closely match the Oregon subspecies, Junco hyemalis oregonus (Miller 1941), but some appear to be intergrades between Junco hyemalis oregonus and Junco hyemalis hyemalis, which produces Junco hye- malis cismontanus (Miller 1941). Subspecies desig- nations of Dark-eyed Juncos are a capricious topic based on phenetic rather than genetic data, and they are a taxonomic nightmare in the Canadian Rockies where multiple subspecies meet and hybridize. There- fore, we refer to the birds analyzed here simply as Junco hyemalis. Birds were captured using Japanese mist nets with painted model male juncos as decoys. Juncos were lured towards the net by playing the taped song of a conspecific. Birds were given a numbered Canadian Wildlife Service leg band and colour bands that conveyed sex, age, and site information when caught. Blood samples were taken from each bird caught between | May and 20 August in 2001 only by punc- turing the alar vein with a needle and collecting ca. 40 ul of blood in a heparinized microhematocrit tube. 235 Blood was blown out of microhematocrit tubes into centrifuge tubes and kept on wet ice. Because blood parasites may follow a diurnal periodicity (Gore et al. 1982), the birds selected for parasite identification were caught at approximately the same time each day (7:00-12:00 hrs). Within 10 hours of collection, 5 ul of blood were used to produce three microscope slides per bird, using the methods of Harrison and Harrison (1986) and Bennett (1970). The slides were fixed in 100% methanol immediately. In the laboratory, blood smears were stained with Geimsa stain for 30 min- utes, and rinsed with distilled water followed by acetone under a fumehood (Deviche et al. 2001). Smears were examined under 400 x 10 magnification, and 50 fields of view per slide were classified as negative or posi- tive for various parasites. Parasites were identified to the genus level by examination under high power (1000 x magnification) with oil immersion, using various keys (Pierce 1981; Bennett and Pierce 1988; Burrey-Caines and Bennett 1992; Bennett et al. 1994), ~and by comparing with photos of blood parasites taken from juncos captured in Alaska (supplied by Pierre Deviche, Arizona State University, personal commu- nication). Parasite species were difficult to ascertain with absolute certainty, but the genus level could be resolved without ambiguity. Therefore, we used the genus level when comparing parasite prevalence among elevations. At each site, one bird was caught between the 1* and 5" of each month (May-August), and another between the 11 and 15" of each month, in order to represent sites and time periods within the breeding season equiv- alently. No females were captured using playback and mist-netting methods, and so analyses here deal solely with males. The statistical significance level for all tests conducted was set at a = 0.05. All tests were performed using SPSS 10. All techniques used were approved by the animal care committee of the University of British Columbia (A0-0046), Parks Can- ada (2000-008), and Environment Canada (Banding: 10429 AJ; Collection: BC SCI 2000/067). THE CANADIAN FIELD-NATURALIST Vol. 118 Results Thirty-two high- and 32 low-elevation adult male birds caught across all eight sites were used in these analyses. In addition, 12 fledglings were caught at high elevations (in August) and 12 were caught at low ele- vations (in August). Three parasite genera were found in the blood of juncos: Haemoproteus (Family Plas- modiidae), Leucocytozoon (Family Plasmodiidae) and- Trypanosoma (Family Trypanosomatidae). All data are summarized in Table 1. The most prevalent blood para- site at both elevations was Haemoproteus, followed by Leucocytozoon, and Trypanosoma. A significantly higher proportion of low-elevation birds (66%, 21/32) compared to high-elevation birds (28%, 9/32) were infected by at least one of these parasites (P = 0.002, Fisher’s Exact Test). A higher percentage of low-ele- vation birds (34%, 11/32) compared to high-elevation birds (19%, 6/32) were infected with Haemoproteus spp., but the difference was not significant (P = 0.08, Fisher’s Exact Test). Similarly, more low-elevation birds (25%, 8/32) versus high-elevation birds (9%, 3/32) were infected with Leucocytozoon spp., with a near significant difference (P = 0.07, Fisher’s Exact Test). Of these individuals, 42% (8/19) of low-eleva- tion and 44% (4/9) of high-elevation birds had both Haemoproteus spp. and Leucocytozoon spp. present in their blood. Two cases of infection with Trypano- soma spp. were found in low-elevation birds, and none at high elevations. Thirty-four percent (11/32) of low- elevation and 72% (23/32) of high-elevation birds were not infected with any parasites. There was no relationship between date of capture and infection at low elevations (P = 0.23, r = 0.33; Pearson’s r). How- ever, at high elevations, a low proportion of individ- uals (12.5%, 2/16) caught between 1 May and 15 June were infected with Leucocytozoon spp. and Haemo- proteus spp., but between 15 June and 30 August, a significantly higher proportion were infected (7/16, 44%; P = 0.05, Fisher’s Exact Test). In samples from low elevations not used in this analysis (taken from birds nearby, but not within our study areas) two cases TABLE |. Summary of data on presence and absence of blood parasites in Dark-eyed Juncos from low (~1000 m asl) and high (~2000 m asl) elevations. Adults (ASY) Infection/Parasite Infected Trypanosoma present Leucocytozoon present Haemoproteus present Both Haemoproteus and Leucocytozoon present no parasites Fledglings Parasite Leucocytozoon present Low (N = 32) High (N = 32) 21 9 2 0 8 3 11 6 8 of 19 individuals 4 of 9 individuals 11 23 Low (N = 12) High (N = 12) 2 1 2004 of Plasmodium spp. (family Plasmodiae) were noted. More fledglings were infected with Leucocytozoon spp. at low elevations as compared to at high elevations (17%, 2/12, vs 8.3%, 1/12), but the difference was not significant (P = 0.39, Fisher’s Exact Test). No other blood parasites were observed in fledglings. Discussion High-elevation adult males and fledglings had lower incidences of blood parasites as compared to low-ele- vation adult males and fledglings. If relief from para- sites enables adults to survive longer and breed for more years, then this could aid in equalizing the life- time reproductive success of high- and low-elevation birds. Blood parasites can decrease survivorship by directly increasing susceptibility to predation (Vaughn and Coble 1975). In addition, there may be a trade- off between reproductive effort and the efficiency of the immune response. For instance, parasite loads may increase during the reproductive period, as they did here at high elevations, when breeding adults spend considerable time provisioning their young and in nest and territorial defence, or when breeding effort is increased experimentally (Ots and Horak 1996; Norris et al. 1994; Weatherhead and Bennett 1991, 1992; Rintmaki et al. 1999). A simpler reason for the differential susceptibility observed here might also be due to a difference in the timing of emergence of the insect vectors carrying the parasites. The two most common parasite genera in this study, Haemoproteus and Leucocytozoon, are protozoon para- sites of birds. Haemoproteus spp. are primarily trans- mitted by insects in the dipteran family Ceratopogoni- dae (no-see-ums, sandflies) or Hippoboscidae (louse flies), whereas Leucocytozoon spp. are primarily trans- mitted by simuliids (e.g., black flies) (Greiner and Ritchie et al. 1994; Rosskopf and Woerpel 1996; Rint- maki et al. 1999). Trypanosoma (likely avium), which was rare in our study, is also transmitted by members of the family Simuliidae. The principal effects of Leucocytozoon infections are intravascular haemolytic anemia, weight loss, and sometimes death, whereas Haemoproteus and Trypanosoma in birds are generally less pathogenic (Greiner and Ritchie 1994). Finally, two low-elevation samples (not analyzed as part of the sub-samples selected) contained Plasmodium spp., which is transmitted by mosquitos (family Culicidae). Almost all of these insect vectors were noted to have emerged later at high elevations, perhaps preventing exposure of birds to the parasites for much of the sea- son. High-elevation birds were also at lower densities (Bears 2002), and therefore transmission between birds may have been lower. Results suggest that high-elevation habitats may be of conservation importance in limiting the spread of blood parasites in birds. The Canadian Rockies may play a particularly important role in limiting the rate at which avian diseases travel east and west across BEARS: PARASITE PREVALENCE IN DARK-EYED JUNCOS the Rocky Mountains. Further work, including com- parisons of parasite levels in females, and in other species that breed over wide elevation ranges, is re- quired in order to assess the generality of this parasite refugium hypothesis in the Rocky Mountains of Can- ada. Mechanisms by which high-elevation birds are protected from parasites should also be explored. Acknowledgments Project funding was provided by NSERC, and personal funding by NSERC PGS-A, Environment Canada, Agriculture and Agri-foods Canada, Health Canada, Fisheries and Oceans Canada, and Natural Resources Canada. J. N. M. Smith provided helpful advice in the preparation of this manuscript. S. Stearns encouraged me with the pursuit of parasite quantifi- cation across elevations. I benefited in the field from the help of G. Brown, M. Bandura, and A. Bendzsak. M. Adamson provided microscopy facilities. Thanks to A. J. Erskine and two anonymous referees for reviewing the manuscript. Literature Cited Bears, H. 2002. How and why a small songbird (Junco hye- malis oregonus) breeds over a steep elevation gradient: Shifting life-histories, adaptations and costs and benefits with elevation. M.Sc. thesis, University of British Colum- bia, British Columbia. 80 pages. Bears, H., J. N. M. Smith, and J. C. Wingfield. 2003. Adrenocortical sensitivity to stress in Dark-eyed Juncos (Junco hyemalis oregonus) breeding in low and high eleva- tional habitat. Ecoscience 10: 127-133. Bennett, G. F. 1970. Simple techniques for making avian blood smears. Canadian Journal of Zoology 48: 585-586. Bennett, G. F., and M. A. Pierce. 1988. Morphological forms of the avian Haemoproteidae and an annotated checklist of the genus Haemoproteus. Journal of Natural History 22: 1683-1689. Bennett, G. F., M. A. Pierce, and R. A. Earle. 1994. An annotated checklist of the valid avian species of Haemo- proteus, Leucocytozoon (Apicomplexa: Haemosporina) and Hepatozoon (Apicomplexa: Haemogregarinidae). Systematic Parasitology 29: 61-73. Braun, C. E., K. Martin, and L. A. Robb. 1993. White- tailed ptarmigan (Lagopus leucurus). Pages 1-24 in The birds of North America (68). Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Washington, D.C. 24 pages. Burrey-Caines, J. R., and G. F. Bennett. 1992. The Haemo- proteidae (Apicomplexa: Haemosporina) of the avian fam- ilies Fringillidae and Emberizidae s.]. Canadian Journal of Zoology 70: 1149-1160. Cristol, D. A., V. Nolan, Jr., and E. D. Ketterson. 1990. Effect of prior residence on dominance status of dark- eyed juncos, Junco hyemalis. Animal Behaviour 40: 580- 586. Deviche, P., E. C. Greiner, and X. Manteca. 2001. Seasonal and age-related changes in blood parasite prevalence in Dark-eyed Juncos (Junco hyemalis, Passeriformes). Journal of Experimental Zoology 289: 456-466. Gore, T. C., G. P. Noblet, and R. Noblet. 1982. Effects of pinealectomy and ocular enuculeation on diurnal periodi- 238 city of the Leucocytozoon smithi (Hemosporina) gameto- cytes in the peripheral blood of domestic turkeys. Journal of Parasitology 29: 415-420. Grasso, M. J., U. M. Savalli, and R. Mumme. 1996. Status signalling in Dark-eyed Juncos: perceived status of other birds affects dominance interactions. Condor 98: 636-639. Greiner, E. C., and B. W. Ritchie. 1994. Parasites. Pages 437-478 in Avian medicine: Principles and application. Edited by B. W. Ritchie, G. J. Harrison, L. R. Harrison. Wingers Publishing, Lakeworth, Florida. 350 pages. Hamman, H. J., K. H. Schmidt, and S. Simonis. 1989. Altitude and size of eggs and clutch in the great tit Parus major. Journal fiir Ornithologie 130: 69-74. Harrison, G. J., and L. R. Harrison. 1986. Clinical avian medicine and surgery including aviculture. W. B. Saun- ders Co., Philadelphia, Pennsylvania. 270 pages. Kollinsky, C., and A. Landmann. 1996. Altitudinal distri- bution of male Black Redstarts: Are there age-dependent patterns? Bird Study 43: 103-107. Landmann, A., and N. Winding. 1993. Niche segregation in high-altitude Himalayan chats (Aves, Turdidae): does mor- phology match ecology? Oecologia 95: 506-519. Miller, A. H. 1941. Speciation of the avian genus Junco. University of California Publications in Zoology 44: 173- 434. Norris, K., M. Anwar, and A. F. Read. 1994. Reproductive effort influences the prevalence of haemoprotozoon para- sites in great tits. Journal of Animal Ecology 63: 601-610. Ots, I., and P. Horak. 1996. Great Tits Parus major trade health for reproduction. Proceedings of the Royal Society of London B: Biological Sciences 263: 1443-1447. THE CANADIAN FIELD-NATURALIST Vol. 118 Pierce, M. A. 1981. Distribution and host-parasite checklist of haematozoa of birds in Western Europe. Journal of Natural History 15: 419-458. Rintmaki, P. T., E. Huhta, J. Jokimaki, and D. Squires- Parsons. 1999. Leucocytozoonosis and trypanosomiasis in redstarts in Finland. Journal of Wildlife Disease 35: 603-607. Rosskopf, W. J., Jr., and R. W. Woerpel. 1996. Pet avian conditions and syndromes. Pages 260-282 in Diseases of cage and aviary birds. Edited by W. J. Rosskopf and R. A. Woerpel. Williams & Wilkins; Baltimore, Maryland. 534 pages. Stabler, R. M., N. J. Kitzmiller, and C. E. Braun. 1974. Hematozoa from Colorado birds, IV. Galliformes. Journal of Parasitology 60: 536-537. Vaughn, G. E., and P. W. Coble. 1975. Sublethal effects of three ectoparasites on fish. Journal of Fish Biology 7: 283- 294. Weatherhead, P. J., and G. F. Bennett. 1991. Ecology of Red-winged Blackbird parasitism by haematozoa. Canadian Journal of Zoology 69: 2352-2359. Weatherhead, P. J., and G. F. Bennett. 1992. Resource levels, reproduction and resistance to haematozoon in- fections. Proceedings of the Royal Society of London B: Biological Sciences 265: 1197-1201. Widmer, M. 1999. Altitudinal variation of migratory traits of the garden warbler Sylvia borin. Ph.D. thesis, University of Zurich, Switzerland. Received 24 October 2002 Accepted 4 November 2004 Movements of Subadult Male Grizzly Bears, Ursus arctos, in the Central Canadian Arctic RosBert J. GAu!, Puitip D. MCLOUGHLIN, RAY Case!, H. DEAN CLUFF?, ROBERT MULDERS!, and FRANCOIS MESSIER? 'Wildlife and Fisheries Division, Department of Resources, Wildlife and Economic Development, Government of the Northwest Territories, #600 5102-50th Avenue, Yellowknife, Northwest Territories X1A 3S8 Canada; e-mail: rob_gau@ gov.nt.ca *Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5E2 Canada 3Department of Resources, Wildlife, and Economic Development, Government of the Northwest Territories, North Slave Region, P.O. Box 2668, Yellowknife, Northwest Territories X1A 2P9 Canada Gau, Robert J., Philip D. McLoughlin, Ray Case, H. Dean Cluff, Robert Mulders, and Francois Messier. 2004. Movements of subadult male Grizzly Bears, Ursus arctos, in the central Canadian Arctic. Canadian Field-Naturalist 118(2): 239-242. Between May 1995 and June 1999, we equipped eight subadult male (3-5 yrs old) Grizzly Bears (Ursus arctos) with satellite radio-collars within a study area of 235 000 km?, centred 400 km northeast of Yellowknife, Northwest Territories, Canada. Subadult male annual home ranges were extraordinarily large (average = 11 407 km’, SE = 3849) due, in part, to their movement’s occasional linear directionality. We believe their long-range linear movements may reflect some individuals tracking the migration of Caribou (Rangifer tarandus). Seasonal daily movement patterns were similar to adult males that were previously reported. The areas used by these bears are the largest ranges reported for any Grizzly Bears and the scale of their movements may put individual bears in contact with humans even when developments are hundreds of kilometres from the central home range of an animal. Key Words: Grizzly Bear, Ursus arctos, home range, movements, subadult, central Arctic, Northwest Territories. The population of Barren-ground Grizzly Bears (Ursus arctos) in the Northwest Territories (NWT) exists at a low density in the tundra and is considered “sensitive” after a recent species-at-risk assessment (Government of the NWT, 2000). As such, there is concern for any increase in human presence that might impact on their numbers. Economic activity in the central Arctic of the NWT and Nunavut increased dramatically in the early 1990s with the discovery of diamonds. At present there are three mines (two diamond, one gold) and numerous base-metal, gold, and other diamond developments in the region. To address the effects of these develop- ments, in 1995 the Government of the NWT and the University of Saskatchewan initiated the first multi- faceted research program into the ecology of Barren- ground Grizzly Bears inhabiting the central Arctic. One aspect of our research program was to detail the spatial requirements of the Barren-ground Grizzly Bears in this region (McLoughlin 2000). However, incidental to our primary objectives, we obtained data on large-scale movements of some subadult male bears (3-5 yrs old) that were also captured and monitored. Although many aspects of bear ecology have been well documented, descriptions of dispersal and movement patterns for subadult Grizzly Bears remain rare and difficult to obtain (McLellan and Hovey 2001). While progress with DNA analysis may shed insights on sub- adult dispersal in the future (see Waits et al. 1999; Woods et al. 1999; Woods and Strobeck 2000), current methods to examine dispersal in bears generally re- quire marking young animals to identify when captured or killed at a later date, or radio tracking 2- and 3-year olds captured prior to or immediately following sepa- ration from their mother. Our findings parallel and complement McLoughlin et al. (1999) and McLoughlin et al. (2003). Methods The study area was centred in Canada’s central Arctic (66°10'N, 111°25'W), encompassing approxi- mately 235 000 km? of mainland Nunavut and the NWT (Figure 1). The study area was delineated by the community of Kugluktuk, the Kent Peninsula, Aylmer Lake, Mackay Lake, and Great Bear Lake. We previ- ously noted the biophysical characteristics of the region in Gau et al. (2002). Between May 1995 and June 1999, helicopters were used to search for and capture bears. Bears weighing >110 kg (males) and >90 kg (females) were fitted with tracking devices. There were the minimum weights we considered bears robust enough to wear a collar safely. We considered most two- and three-year-old bears too small and growing too rapidly to be fitted with satel- lite collars. Some telemetry methods (e.g., inserts or breakaways to collar belting, ear-tag transmitters, ex- pandable radio-collars) show promise; however, im- provements are still needed (Costello et al. 2001). Satellite (Service Argos Inc., Landover, Maryland, USA) and conventional VHF radio-telemetry (Telonics Ltd., Mesa, Arizona, USA) were used to obtain spatial information on Barren-ground Grizzly Bears. Most 239 240 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 1. Subadult male Grizzly Bears, and their periods of long-range (>200 km) linear directional movement, captured and collared between 1995 and 1999 in the central Canadian Arctic. Number 95% fixed Mean Daily Movement (km/day) Linear Directional Movements of kernal Late Distance Number Bear Age Year locations range (km?) Spring Summer summer Autumn covered (km) of days G595 4 1995 103 15 899 eS) 10.4 6.2 9) 539 Sif) G600 3 1995 129 22 007 16.6 8.8 10.5 es 7719 59 G612 4 1995 91 4448 24.8 9.7 4.0 Nile ge 471 23 446 33 G618 5 1995 66 4540 6.1 6.7 9.5 5.6 G656 5) 1996 45 32 188 n/a! n/a! n/a! n/a! 273 26 305 40 G657 5) 1996 42 6162 n/a! n/a! n/a! n/a! G689 4 1998 58 3662 n/a! n/a! n/a! n/a! G700 4 1998 64 2349 7.9 59 4.2 ay 201 32 ' <8 locations/season in every season of the year were recorded thus omitted from analysis. collars were designed to transmit approximately 2-5 latitude-longitude locations every two days (8-hour duty cycle) from 1 May to | November. Our calculations and techniques for study of ani- mals, ranges from satellite telemetry locations and rates of movement (km/day) were previously described in McLoughlin et al. (1999) and McLoughlin et al. (2003). Annual home ranges were determined using the 95% isopleth for bears only with 238 locations, so as not to overestimate range size with smaller sample sizes (Seaman et al. 1999). Also, only those animals that transmitted 28 locations/season in every season of the year were included for analysis. We defined seasons according to changes in the diet of Barren-ground Grizzly Bears during the active period (adapted from Gau et al. 2002), including spring (den emergence- 20 June), summer (21 June-31 July), late-summer (1 August-9 September), and autumn (10 September- den entrance). Results Of the male bears that were not in a family group with their mother, 6 of the 45 males handled were too small for collaring. However, we did collar and mon- itor 8 subadult males between 3 and 5 years of age out of a sample of 39 collared males (Table 1). Male subadult movements were extraordinarily large and annual home range averaged 11 407 km? (SE = 3849, range 2349 — 32 188). A unique feature of the subadult movements we observed consisted of periods of long-range (>200 km) linear directional forays. Five of the eight subadults we followed exhibited this linear movement pattern. For example, 4-year-old bear G612 in 1995 moved 471 km from 31 May to 22 June along a northerly trek from the treeline to the Arctic coast. While the other movements may not be as dramatic as G612 in terms of time a distance was covered, long-range linear directional movement appeared common for subadult males in this Barren-ground Grizzly population. Bears G595, G600, G612, and G656 had linear directional movements in the spring; additionally, bears G612, G656, and G700 had autumn linear directional move- ments. The subadult males followed a seasonal daily move- ment pattern similar to adult males (see McLoughlin et al. 1999). Only five bears met our criteria to inves- tigate seasonal movements. Means were 12.6 km/d (SE='3.6), 8:3 km/d' (SE = 0:9), 6:9’ kin/d"\(SE'=' 1:3); and 7.6 km/d (SE = 1.2) for spring, summer, late summer, and autumn, respectively. There was a clear general trend from a high rate of movement (spring) to lower rates for the rest of their active period, although an ANOVA determined no significant differences between seasonal means (F3 ,,= 1.6, P = 0.2). Discussion The annual ranges of adult and subadult Barren- ground Grizzly Bears in the central Canadian Arctic are the largest ranges yet reported for grizzlies in North America (see Table 1, McLoughlin et al. 1999). How- ever, there are very few published data about subadult Grizzly Bear movement patterns either to refute or sup- port our claim (McLellan and Hovey 2001). Jonkel (1987), along with Mace and Waller (1997), noted that dispersing immature brown bears tended to have smaller home ranges than adult male bears. However, LeFranc et al. (1987), along with Nagy et al. (1983), summarized examples where subadults, believed dis- persing from maternal home ranges, had home ranges as large or larger than those of adult males. Regardless, the magnitude of the movement patterns we observed eclipses other results that have been previously pub- lished. We have gleaned valuable insights into the move- ment patterns of Canadian Barren-ground Grizzly Bears. Mean annual ranges were 7245 km? for adult males and 2100 km? for females, with no difference in 2004 GAU, MCLOUGHLIN, CASE, CLUFF, MULDERS, and MESSIER: MOVEMENTS OF BEARS 241 Kent Peninsula ‘CORONATION GULF Kugluktuk hie de oa i \ MacKay axe a Aylmer =z) 250 km co Nunavut Northwest 7 Erritories Lake FiGureE |. Location of the study area in Canada’s central Arctic. The treeline indicates the approximate northern limit of coniferous forest in the region. the ranges for females of differing family status (Mc- Loughlin et al. 2003). The large disparity between adult males and females (5145 km7?), and especially between subadult males and females (9307 km2), is in- teresting since females introduce younger male bears only to a seemingly small portion of the land they will eventually use in the central Arctic. Although we can expect a high degree of movement and population over- lap among males in the central Arctic (McLoughlin et al. 2002), it is possible that some of the subadult male ranges we recorded were somewhat inflated. We sus- pect that some bears in this study tracked the spring migration of Caribou (Rangifer tarandus), a behaviour previously suspected in northern Alaska for Barren- ground Grizzlies (Reynolds and Garner 1987). At the daily and seasonal movement rates we ob- served for subadult males, because of their similarity to adult male daily movement rates, it would be possible for subadult males to have home range sizes typical of male adults in the region. However, the unique fea- ture of the subadult movements we observed appeared to be their occasional linear directionality. Other than homing behaviours of transplanted bears (Miller and Ballard 1982), extended directional movements are rare (McLellan and Hovey 2001). Explanations for long- range bear movements include seeking out quality hab- itat, spacing behaviour resulting from social interac- tions with other bears, the abundance and distribution of food (e.g., following migrating caribou herds), in- breeding avoidance, and maximizing reproductive fit- ness, or dispersal immediately following separation from their mother (Rogers 1987; Pasitschniak-Arts and Messier 2000). All these factors likely contributed to the movement patterns of subadult males in the Canadian central Arctic. McLoughlin et al. (1999) also pointed out that Barren-ground Grizzly Bears have larger scale movements when compared to other griz- zly populations principally due to low primary pro- ductivity in the tundra environment. The spatial behaviour of subadult male grizzlies increases their probability of coming in contact with humans even when sites of human activity (e.g., ex- 242 ploration and hunting camps, industrial developments, and communities) are of considerable distance from the central home range of an individual. Subadult bears in particular do not have the life-experience of mature bears, are often imbued with more curiosity, and thus are highly susceptible to human activity. Management of bears in the central Arctic should focus on main- taining low levels of human-caused mortality of bears, with the realization that communities, hunting camps, and mining/exploration camps may impact bears from more than just the general vicinity. Also, the wide- spread movements of subadult bears are a compelling argument for a comprehensive and consistent bear conflict avoidance program throughout the central Arctic. Developments that might not be considered in optimum Grizzly Bear habitat should have the tools or resources to effectively manage potential problem bears. Acknowledgments Funding for our research was supplied by the Government of the NWT Department of Resources, Wildlife, and Economic Development, University of Saskatchewan, West Kitikmeot/Slave Society, Indian and Northern Affairs Canada, BHP Diamonds, Rescan Environmental Services, Axys Environmental Consult- ing Ltd., Diavik Diamond Project, Monopros Limited, Echo Bay Mines Ltd., Nunavut Wildlife Management Board, the Federal Department of Indian Affairs and Northern Development, the Northern Scientific Train- ing Program, the Natural Sciences and Engineering Research Council of Canada, and the Polar Conti- nental Shelf Project (number 01298). Literature Cited Costello, C. M., R. M. Inman, K. L. Higgins, D. E. Jones, H. B. Quigley, and S. L. Simek. 2001. Comparison of an expandable radiocollar and an eartag transmitter for moni- toring juvenile black bears. Western Black Bear Work- shop 7: 17. Gau, R. J., R. Case, D. F. Penner, and P. D. McLoughlin. 2002. Feeding patterns of barren-ground grizzly bears in the central Canadian Arctic. Arctic 55: 339-344. Government of the Northwest Territories. 2000. NWT species 2000: general status ranks of wild species in the Northwest Territories. Government of the NWT, Yellow- knife, Northwest Territories. 50 pages. Jonkel, C. J. 1987. Brown bear. Pages 457-473 in Wild Fur- bearer Management and Conservation in North America. Edited by M. Novak. Ontario Ministry of Natural Re- sources, Toronto, Ontario. Lefranc, M. N., M. B. Moss, K. A. Patnode, and W. C. Sugg. 1987. Grizzly bear compendium. Interagency Griz- zly Bear Commitee, Bozeman, Montana. 540 pages. THE CANADIAN FIELD-NATURALIST Vol. 118 Mace, R. D., and J. S. Waller. 1997. Spatial and temporal interaction of male and female grizzly bears in northwest- ern Montana. Journal of Wildlife Management 61: 39-52. McLellan, B. N., and F. W. Hovey. 2001. Natal dispersal of grizzly bears. Canadian Journal of Zoology 79: 838-844. McLoughlin, P. D. 2000. The spatial organization and habi- tat selection patterns of barren-ground grizzly bears in Nunavut and the Northwest Territories, Canada. Ph.D. dissertation, University of Saskatchewan, Saskatoon, Sas- katchewan. 151 pages. McLoughlin, P. D., R. L. Case, R. J. Gau, S. Ferguson, and F. Messier. 1999. Annual and seasonal movement patterns of barren-ground grizzly bears in the central Northwest Territories. Ursus 11: 79-86. McLoughlin, P. D., H. D. Cluff, R. J. Gau, R. Mulders, R. L. Case, and F. Messier. 2002. Population delineation of barren-ground grizzly bears in the central Canadian Arctic. Wildlife Society Bulletin 30: 728-737. 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Research Links 8: 4-9. Received 3 February 2002 Accepted 11 November 2004 Severe Chronic Neck Injury Caused by a Snare in a Coyote, Canis latrans PIERRE-YVES Daoust!:3 and PETER H. NICHOLSON! 2 ‘Canadian Cooperative Wildlife Health Centre, Department of Pathology and Microbiology (Daoust), and Class of 2002 (Nicholson), Atlantic Veterinary College, University of Prince Edward Island, 550 University Avenue, Charlottetown, Prince Edward Island CIA 4P3, Canada Present address: College Village Animal Clinic, 2036 E. Northern Lights Boulevard, Anchorage, Alaska, 99508, USA 3Corresponding author: e-mail: daoust @upei.ca Daoust, Pierre-Yves, and Peter H. Nicholson. 2004. Severe chronic neck injury caused by a snare in a Coyote, Canis latrans. Canadian Field-Naturalist 118(2): 243-246. A two-year-old male Coyote, Canis latrans, in poor body condition was found in a moribund state with a snare deeply embedded in the ventral portion of its neck, more than a month after the official end of the trapping season on Prince Edward Island. This snare had presumably malfunctioned, and the cable had cut through the soft tissues of the neck as well as the trachea and had obstructed both jugular veins and both common carotid arteries but had largely spared both vagosympathetic trunks. Cases like this illustrate the need to continue to work on improving the efficiency of trapping methods, through research and trapper education. Key Words: Coyote, Canis latrans, trapping, injury, snare, Prince Edward Island. Animal welfare issues surrounding trapping meth- ods used to capture furbearers have been an ongoing topic of discussion between wildlife managers and animal protection groups for many years (Proulx and Barrett 1991). Because of this and of the economic importance of the fur trade to Canada, this country has taken a leading role in developing national and international standards for the performance of restrain- ing- and killing-type traps. In the section of the Agree- ment on International Humane Trapping Standards (AIHTS) (1997*) dealing with killing type traps such as the commonly used rotating-jaw Conibear trap, the designated time limit to irreversible loss of corneal reflexes in at least 80% of the animals of the target species caught in the trap varies from 45 seconds in Ermine (Mustela erminea) to 120 seconds in Pine Mar- ten (Martes americana) to 300 seconds in 17 other species of North American and European furbearing animals. This Agreement requires that the parties in- volved continue research with a view to lowering the threshold requirements. Nonetheless, it implicitly rec- ognizes that no killing trapping method used to cap- ture wild animals can as yet guarantee a humanely acceptable process in each instance. Manual neck snares (where the animal provides the energy neces- sary to tighten the snare) are widely used by trappers to capture various furbearing animals, and Guthery and Beasom (1978) and Boddicker (1982) discussed the advantages and disadvantages of snares as compared to other trapping devices. Snares are allowed in most Canadian jurisdictions, except the southern regions of some provinces. Being considered devices that are constructed by the trappers themselves, they are not subject to the testing requirements under the AIHTS (1997), but their design must be approved by relevant competent authorities such as provincial Departments of Wildlife. This article describes an unsuccessful cap- ture by snare of a Coyote (Canis latrans), with severe consequences. Case Description On 26 February, 2002, a male Coyote, subsequently determined to be 2 years old (Matson’s Laboratory, Milltown, Montana, USA), was found alive but mori- bund in a field in Kings County (46°08'N, 62°36'W), Prince Edward Island (PEI), and was euthanized by a shot to the head. It was submitted for necropsy to the Diagnostic Laboratory of the Atlantic Veterinary Col- lege, University of PEI. Postmortem examination revealed that the animal had been caught in a snare and had managed to escape. The snare’s cable, made of galvanized steel wire 2 mm in diameter, was equipped with a standard U-shaped lock, similar to an Adams lock. The snare surrounded the animal’s neck, with a portion of cable approximately 20 cm long dangling beyond the neck. The coyote was in poor body con- dition; the stomach was completely empty, and the intestines contained only a small amount of ingesta. However, some fat was still present around the base of the heart, and a sample of diaphyseal bone marrow from one of the femurs contained approximately 54% fat, as measured by comparison of the wet and dry weights of the sample. A large area of the skin along the ventral side of the neck, about 50 cm?, was miss- ing and had been replaced by scar tissue that released a foul odour indicative of suppuration. The portion of the cable along the ventral side of the neck had tran- sected the full diameter of the trachea and was embedded in scar tissue between the trachea and the esophagus (Figure 1). The wall of the severed portion of the trachea was completely healed, although the tissue of repair had extended as a thin circular band 243 244 into the tracheal lumen, reducing its surface area to about one-seventh of its original size (Figure 2). The esophageal lumen was patent. There was no gross evi- dence of suppuration accompanying the scar tissue that surrounded the trachea and the cable. At the time of necropsy, the cable was freely movable within the affected tissues. The architecture of both jugular veins disappeared at the level of the scar tissue. A thin probe inserted into the cranial portion of the common caro- tid artery on both sides of the trachea was blocked from further progression at this level. The outline of the vagosympathetic trunk on both sides was also lost among the scar tissue. No other significant lesion was seen at necropsy. Samples of the affected portion of trachea and surrounding soft tissues were fixed in for- malin, processed routinely for microscopic examina- tion, and stained with hematoxylin and eosin. Microscopically, the scar tissue surrounding the affected portion of trachea consisted of abundant dense fibrous tissue, much of which was birefringent under polarized light. This fibrous tissue enclosed several pockets of necrotic material (including debris of in- flammatory cells) and, in some areas, contained numer- ous cross sections of hair shafts. Serial sections of this tissue, parallel to the length of the trachea, revealed on both sides an artery, about 3 mm in diameter, and a nerve, about 1 mm in diameter. Taking into account the shrinkage that normally occurs in tissues processed for microscopic examination, the diameters of these two structures were compatible with those of the com- mon carotid artery and vagosympathetic trunk, respec- tively, for an animal of this size. Depending on the section examined, the lumen of the common carotid artery on both sides of the trachea was either empty or partly or completely filled with a well organized thrombus. The vagosympathetic trunk on both sides was morphologically normal, except for a few degen- erative lesions suggesting some loss of axons and their myelin sheaths (wallerian degeneration). The tissue within the tracheal lumen also consisted of dense fi- brous tissue that enclosed some necrotic material and numerous cross sections of hair shafts. Discussion Ideally, a snare should cause rapid death by neck strangulation and subsequent asphyxiation. A quick death is more humane and also gives the animal less time to find a way out of the snare. A variety of factors can influence the performance of a snare. Some lock- ing mechanisms tighten less quickly and firmly than others (Phillips 1996), or small kinks in the cable may weaken it and make it easier to break or may prevent full function of the lock. Inclusion of a swivel attach- ing the lock to the cable (not present in this case) can prevent the cable from kinking as the lock slides along it. If the cable is not anchored firmly enough, the animal may be able to pull off the snare before maximum closure has been achieved. If death is not THE CANADIAN FIELD-NATURALIST Vol. 118 rapid and the cable is anchored too low, it may be more easily accessible to the animal’s jaws and be chewed through. Seventeen of 131 (13%) Coyotes caught by one of three types of snares studied by Phillips (1996) managed to escape, one by breaking the lock and 16 by chewing through the cable. In this Coyote, the relative abundance and density of the scar tissue in the neck and, microscopically, its birefringence under polarized light suggested that the failed capture had occurred at least a few weeks prior to death. The trapping season for Coyote and Red Fox (Vulpes vulpes) on PEI officially terminated on Janu- ary 15, more than a month before this animal was found. The reason why it managed to escape capture was not determined, although the portion of cable dan- gling from its neck was of the right length to suggest that the animal had chewed through it rather than pull the whole cable from where it was anchored. Unfortu- nately, the animal’s gums and teeth were not examined for the presence of recent damage that would have sup- ported this suggestion. At the time of escape, sufficient closure of the snare may have already been achieved for the cable to exert a strong pressure on soft tissues of the ventral side of the neck and be forced gradually through these tissues over the following weeks. Grad- ual transection and concurrent repair of the tracheal wall would thus have allowed it to remain continuous. Alternatively, transection of all soft tissues, including the trachea, could have occurred at the time of the animal’s initial struggle in the snare. Such rapid tran- section of the trachea, however, would more likely have resulted in the formation of a gap between its cut ends, thus preventing them from healing together. Moreover, if the trachea had been rapidly transected, the jugular veins would probably also have been cut and the animal would presumably have bled to death. The thicker hair coat and skin on the dorsal side of the neck as compared to the ventral side (Scott et al. 2001) and the tough nuchal ligament on the dorsal side (Evans 1993) may have prevented the cable from moving into the tissues from that side. The common carotid arteries and the vagus nerves are important structures running along both sides of the trachea. In this animal, the lumen of both common carotid arteries was occluded. However, this is not a critical problem, at least in canids. In dogs, both com- mon carotid arteries can be ligated experimentally without causing any clinical signs, implying that col- lateral circulation (e.g., via the vertebral arteries) is able to quickly compensate and transport sufficient blood to the brain (Whisnant et al. 1956). Such col- lateral circulation would have even more time to devel- op if the obstruction were gradual, as is suspected to have occurred in this case. The same reasoning applies to the jugular veins, located much more superficially in the ventral region of the neck, since collateral cir- culation within the venous system can develop even more readily than in the arterial side. Proper function 2004 DAOUST AND NICHOLSON: NECK INJURY IN A COYOTE 245 FiGure |. Right lateral view of the neck of a Coyote. The carcass is lying on its back, with the head to the left. The cable from a snare has cut through the trachea (T) and is now lodged between it and the esophagus (E). The distortion of the wall of the trachea just above the cable as been caused by scar tissue of repair. A metal rod has been inserted in the lumen of the esophagus. The common carotid artery (C) and vagosympathetic trunk (V) are clearly visible to the left of the cable. The lock of the snare is at the bottom of the picture. (Bar = 2 cm.) : of at least one of the vagosympathetic trunks is, how- ever, essential to life. The vagal (parasympathetic) portion of these nerves originates in the brain and supplies efferent and afferent innervation to many vital organs, including the cardiovascular system and digestive tract (Andrews and Lawes 1992; Esler 1992; Guyton and Hall 1996). In particular, vagal stimula- tion protects against excessive stimulation of cardiac function by the sympathetic nervous system during physical activity and stress. In this Coyote, it was not possible to dissect the vagosympathetic trunk on either side of the affected portion of trachea, because both trunks were well embedded in scar tissue at this level. However, careful microscopic examination showed that both trunks were in large part morphologically in- tact. Although their structure had likely been stretched, and their path distorded, by the cable, the gradual pro- cess through which this happened would presumably have provided sufficient time for anatomic and physio- logical compensation. The exact cause of the eventual demise of this coy- ote was not clear. There are undocumented reports by veterinary practitioners of collars being left permanent- ly on growing dogs and eventually becoming buried under the skin, with damage limited to chronic sup- puration of the subcutis (J. B. Miller, Department of Companion Animals, Atlantic Veterinary College, Uni- versity of PEI, personal communication). Similarly, a square of a monofilament gillnet mesh was found al- most totally embedded in the snout of an otherwise healthy Porbeagle (Lamna nasus) that had been caught on the Scotian Shelf (Benz et al. 2001). However, the marked reduction in size of the tracheal lumen in this Coyote may have reduced its stamina and prevented it from foraging efficiently. The esophageal lumen was FiGurRE 2. Cranial view of the trachea of a Coyote at the level where the cable of a snare has cut through this organ. The ventral side of the trachea is at the top. The carti- laginous ring (CR) outlines what should be the normal size of the tracheal lumen. Scar tissue of repair has greatly narrowed this lumen. (Bar = | cm.) 246 patent, but the presence of the cable against its ventral surface may have interfered with the passage of large food boluses. Albeit seemingly superficial, the suppu- rative process and, therefore, probable bacterial in- fection, associated with the soft tissue damage would have further drained the animal’s energy reserves. Although this animal had not yet reached a stage of emaciation, its poor body condition, combined with the energy demands imposed by the winter season, may have been sufficient to bring it to a stage of exhaustion, perhaps combined with hypothermia. The clinical significance, if any, of the mild damage seen microscopically in the vagosympathetic trunks could have been assessed only in the live animal. The efficacy of various trapping devices, including snares, can now be properly tested under standardized laboratory conditions (Proulx and Barrett 1990, 1991). Ultimately, however, the proportion of animals that fail to die rapidly and undergo prolonged suffering depends much on the experience and expertise of the trappers in the field. Regardless of the exact cause of failure of the snare in this case, its outcome emphasizes the need for continued attention to appropriate train- ing of hunters and trappers, in order to ensure that the best trapping devices and capture techniques are used consistently and, as a result, as few animals as pos- sible undergo undue suffering. It also illustrates the extreme resilience of Coyotes. Acknowledgments This article benefited greatly from discussions with Neal Jotham. Documents Cited [marked * in text] Agreement on International Humane Trapping Standards. The European Community, the Government of Canada, and the Government of the Russian Federation. 1997. Department of Foreign Affairs and International Trade, Ottawa, Ontario, Canada. 31 pages. THE CANADIAN FIELD-NATURALIST Vol. 118 Literature Cited Andrews, P. L. R, and I. N. C. Lawes. 1992. A protective role for vagal afferents: an hypothesis. Pages 280-302 in Neuroanatomy and physiology of abdominal vagal afferents. Edited by S. Ritter, R. C. Ritter and C. D. Barnes. CRC Press, Boca Raton, Florida, 316 pages. Benz, G. W., A. Kingman, and J. D. Borucinska. 2001. Gillnet survival and healing by a Porbeagle, Lamna nasus. Canadian Field-Naturalist 115: 506-509. Boddicker, M. L. 1982. Snares for predator control. Pro- ceedings of the Vertebrate Pest Conference 10: 50-54. Esler, M. 1992. The autonomic nervous system and cardiac arrhythmias. Clinical Autonomic Research 2: 133-135. Evans, H. E. 1993. Arthrology. Pages 219-257 in Miller’s anatomy of the dog. Edited by H. E. Evans. W. B. Saunders Company, Toronto, 3" edition, 1113 pages. Guthery, F. S., and S. L. Beasom. 1978. Effectiveness and selectivity of neck snares in predator control. Journal of Wildlife Management 42: 457-459. Guyton, A. C., and J. E. Hall 1996. Textbook of medical physiology. W. B. Saunders Company, Toronto, 9" edition: pages 107-119, 793-802. Phillips, R. L. 1996. Evaluation of 3 types of snares for cap- turing coyotes. Wildlife Society Bulletin 24: 107-110. Proulx, G., and M. W. Barrett. 1990. Assessment of power snares to effectively kill red fox. Wildlife Society Bul- letin 18: 27-30. Proulx, G., and M. W. Barrett. 1991. Ideological conflict between animal rightists and wildlife professionals over trapping wild furbearers. Transactions of the North Ameri- can Wildlife and Natural Resources Conference 56: 387-399. Scott, D. W., W. H. Miller Jr., and C. E. Griffin. 2001. Muller & Kirk’s small animal dermatology. W. B. Saunders Company, Toronto, 6" edition, pages 1-70. Whisnant, J. P., C. H. Millikan, K. G. Wakim, and G. P. Sayre. 1956. Collateral circulation to the brain of the dog following bilateral ligation of the carotid and vertebral arteries. American Journal of Physiology 186: 275-277. Received 7 February 2003 Accepted 26 October 2004 Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Preserve, Alaska Tom S. SMITH*, STEVEN T. PARTRIDGE”, and JOHN W. SCHOEN? 4 United States Geological Survey Alaska Science Center, 1011 E. Tudor Road, Anchorage, Alaska 99503 USA >Alaska Office — National Audubon Society, 308 G. Street, #217, Anchorage, Alaska 99501 USA Smith, Tom S., Steven T. Partridge, and John W. Schoen. 2003. Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Preserve, Alaska. Canadian Field-Naturalist 118(2): 247-250. We describe several encounters between Brown Bears (Ursus arctos) and Gray Wolves (Canis lupus) that were observed at Katmai National Park and Preserve in southwest Alaska. Katmai Brown Bears and Gray Wolves were observed interacting in a variety of behavioral modes that ranged from agonistic to tolerant. These observations provide additional insight regard- ing the behavioral plasticity associated with bear-wolf interactions. Key Words: Brown Bear, Ursus arctos, Gray Wolf, Canis lupus, interactions, Katmai National Park and Preserve, Alaska. Flexibility in mammalian behavior parallels the evo- lution of large brains (Gilbert 1989). Gittleman (1986) presented data showing that bears and wolves have the highest brain:body weight ratios within the order Carnivora, and claimed that this accounts for their behavioral plasticity. The context of the encounter set- ting, as well as the behavior of individual animals, like- ly influences the resulting relationship between bears and wolves at any given time. Alaska Brown [Grizzly] Bears (Ursus arctos) and Gray Wolves (Canis lupus) are sympatric over much of their range and often compete for access to the same food resources (Adams et al. 1995). As sym- patric, apical predators, bears and wolves probably interact frequently, although reports in the scientific literature are scant. The accounts reported illustrate the variable nature of Brown Bear — Gray Wolf inter- actions, and range from mutual tolerance (Lent 1964), to competitive (Adams et al. 1995; Hornbeck and Horejsi 1986; Ballard 1982), to predatory (Ballard 1980; Hayes and Mossop 1987). This note describes additional Brown Bear — Gray Wolf interactions ob- served at Katmai National Park and Preserve on the Alaska Peninsula, approximately 400 km southwest of Anchorage, Alaska. Wolves harass bears On 24 June 2001, while conducting Brown Bear research at Hallo Bay (154° 05’ W, 58° 27’ N) on the Katmai coast, the authors observed a pack of five wolves harass a pair of courting Brown Bears on the edge of extensive salt marsh meadows (Carex spp., Plantago spp., Triglochin spp.). Prior to this observa- tion, we had repeatedly heard wolves “chorus howI- ing” (Harrington and Asa 2003), and noted that the pack was gradually nearing our research field camp situated on the far end of the meadow. Anticipating their arrival, we positioned ourselves where we could observe them when they entered the meadow. Although it was late in the evening (2230 h Alaska Daylight Time), it was still light enough to see so we patiently waited while scanning the meadow. About 400 m south of our position, a pair of Brown Bears foraged on sedges in the meadow near a willow-alder (Salix spp., Alnus spp.) thicket. After we waited 10 minutes, the wolf pack suddenly appeared, emerging from the dense thicket about 25 m north of the two bears. The wolves trotted single file in the direction of the bears. Three of the five wolves had black-gray coats, similar in pat- tern to that of the German Shepherd (C. domesticus) breed of domestic dog; one wolf’s coat was pure white; the other coal black. All five wolves appeared to be mature adults, standing nearly 1 m at the shoulder and similar in stature. As one, the wolves circled south from the meadow’s center toward the bears, which by then were focused on the wolves. The five wolves lined up, head-to-head, with the large male Brown Bear with < 2 m separating the lead wolf and the bear. Suddenly, one wolf rushed the maie bear, which in turn lunged and swatted at the attacking wolf with its foreleg. Leaping forward, duck- ing the bear’s defensive swat, then breaking away, the wolves took turns harassing the male bear. The bear responded to the attack with short lunges and aggres- sive paw swats while slowly backing away. About a minute into the confrontation, two wolves circled the bear and lunge-nipped at his unprotected hindquarters. In defense, the male bear swiftly spun around and swat- ted at the closest attacker, but the wolf dodged the bear’s swinging paw then trotted a few meters away, tail wag- ging high. While the male bear counter-attacked the wolves formerly at his rear, the other three wolves, now at his unprotected hindquarters also began a nipping attack, to which the bear again whirled around, lunged, growled, and aggressively swatted at the wolves. The wolves easily outmaneuvered the large bear’s paw swipes and lunges with quick, evasive maneuvers. The wolves and male bear repeated this attack-count- er attack interaction several times before a group of three wolves split from the pack and rushed the female bear that had been watching from a distance of about 10 meters, near the edge of the thicket. Repeating the same lunge, nip, and run strategy, the two groups of wolves attacked each bear, which rapidly spun around to defend themselves from the lunging-biting wolves. At this point, approximately 10-20 meters separated the two bears as they fended off the wolves. During these aggressive interactions, neither wolves nor bears were observed making physical contact, although the confrontation appeared quite serious. After 5 minutes of sustained confrontation, two wolves abruptly termi- nated their attack and began chasing each other in tight circles a short distance from the other wolves, wag- ging tails held high. Moments later, the remaining three wolves also withdrew, excitedly darted about, wagging tails, and then joined the others. After regrouping about 30 m from the bears, all wolves participated in a series of long, drawn-out “chorus” howls. From the forest to the west, responding howls (possibly from a single wolf) were heard in apparent reply. Moments later, the pack trotted southward into the rapidly dimming twilight, away from the bears and researchers. The entire interaction took approximately 20 minutes from the time the wolves were observed entering the mead- ow to when they disappeared into the dusk. To the best of our knowledge, there had been no food source from which the wolves were attempting to drive the bears. It appeared to us that this wolf pack had encountered and approached these two bears and attempted to test, or harass, them for a short period before breaking off the encounter and moving on. Wolves steal fish from bears In mid-July 2001, a pair of Gray Wolves was ob- served stealing Chum Salmon (Oncorhynchus keta) from Brown Bears at Middle Creek where it empties into Hallo Bay on the Katmai coast. National Park Service (NPS) ranger Stephens Harper first saw the wolves when they trotted past the rangers’ camp at 0830 h, heading to the nearby beach. Bears and wolves frequent the tidal flats because the stream channel is quite shallow at low tide, thus making salmon more vulnerable to predation. Harper walked to the stream mouth a few minutes before noon and observed two wolves and five single adult bears fishing in close proximity to one another. Both wolves were gray with black highlights; one appeared mature, while the other was younger, lankier, and smaller (Figure 1). Fishing at the river mouth were five adult single bears. The younger wolf bedded on a gravel bar some distance from the bears while the other, much closer, visually scanned them, apparently wait- ing for one to catch a fish successfully. Once a bear caught a fish, the older wolf would move in, head low, and attempt to steal it by suddenly lunging and snag- ging a portion away from the bear. In response, the THE CANADIAN FIELD-NATURALIST Vol. 118 bear would protect its quarry by bolting and running after the wolf. Although there was a lot of chasing up and down the stream banks, the wolf easily outran and outmaneuvered these bears. This strategy worked, however, in getting bears to abandon what was left of the fish and the wolf would circle back around and eat these scraps. In turn, the older wolf would trot over to where the other was bedded, regurgitate a portion, and then head back to where the bears were fishing. During the entire hour that Harper watched, the wolf determinedly attempted to snatch fish from bears. Although these wolves attempted to catch salmon for themselves on several occasions, they were unsuccess- ful. One wolf, however, scavenged a Starry Flounder (Platichthys stellatus) lying on the beach that had been exposed at low tide. Wolf displaces a bear from a moose carcass In June 1997, an NPS employee observed a single wolf feeding on a Moose (Alces alces) carcass near a gravel pit adjacent to the Valley of Ten Thousand Smokes Road, approximately 13 km from Brooks Camp. Adjacent to the gravel pit is a wetland-pond complex impounded by a Beaver (Castor canadensis) dam. The following day, NPS employees Jim Gavin and Tom Ferguson observed a wolf and bear simultaneous- ly at the site. The wolf was gray with black highlights, appeared mature, but thin. The bear was a young fe- male, approximately 4 years of age. Initially, only the bear fed on the carcass while the bedded wolf watched from 40 m distant. After a while, the wolf moved in close and began harassing the bear by charging and biting at its hind legs. The wolf’s constant attacks dis- rupted the bear’s feeding as it defended itself. After 1.5 min of intense harassment, the bear took flight into the forest and the wolf usurped the carcass and started feeding. After approximately 10 min of feeding, the satiated wolf trotted off into the forest, reappeared 20 min later, then repeated the process several times, likely caching food (L. Adams, USGS research biol- ogist, personal communication), or transporting it to young at a den nearby. Gavin observed the wolf feed in this manner for three consecutive days. On the third day, while the wolf was absent, the bear returned and resumed control of the Moose carcass. The wolf re- appeared but did not approach any closer than 40 m to the feeding bear. Gavin did not observe the wolf attempt to displace the bear as before, although his observation sessions were only 30 min daily. Bears and wolves fishing together Chum Salmon enter freshwater to spawn in Hallo Bay’s Middle Creek from early July through August. In 2001, Brown Bears and wolves were repeatedly observed fishing side-by-side (approximately 5 m apart) at this location with no apparent interactions or obvious concern toward the other species. Like bears, wolves sat patiently on the gravel bank and scanned the water for incoming salmon. Upon catching sight of an incom- 2004 ing salmon, both bears and wolves would leap into the stream and attempt to capture it. Both bears and wolves appeared indifferent to one another, completely fo- cused on fishing. Similarly, in August 1992, NPS rangers B. Holmes and S. Klenzendorf observed a lone wolf and several bears fishing in close association for 2 h at Margo Creek (155° 03’ W, 58° 29’ N) in the western portion of Katmai National Park. At 1210 h on 13 August, an old female Brown Bear was fishing when a wolf ap- peared suddenly on the riverbank, watched the bear, and then sat down. The wolf was moderate in size and of gray coloration. The wolf wandered in the direction of the bear, entered the creek and retrieved a spawn- ing Red Salmon (Oncorhynchus nerka). As the wolf fed, the bear wandered downstream towards the wolf. Consequently, the wolf abandoned its catch, moved quickly away, picked up another salmon, and then moved into the nearby White Spruce (Picea glauca) forest. At 1245 h, the wolf reappeared from the forest, plunged into the creek and emerged with a live salmon. Carrying the salmon in its mouth, the wolf disappeared again into the heavy cover of the forest. At 1255 h, the wolf reappeared and entered the creek in search of sal- mon. As the wolf waded about searching for fish, the same bear waded downstream until she and the wolf were about 20 m apart. In apparent response, the wolf abruptly ceased fishing, climbed out of the river and sat and watched the bear as it sought fish in that section of stream. Twenty minutes later, the wolf moved up- stream and caught a salmon. As the wolf fed, a large adult male bear suddenly appeared thus causing the wolf to leave the area. The large male bear quickly caught a fish, then left the river, at which time the wolf reappeared. Moments later, the wolf caught a fish then reentered the forest. The observers left at 1402 h, im- pressed with the degree of tolerance the two species appeared to have for one another. Wolf and bears traveling together In June 1989, Aleska Szweda, an NPS employee stationed at Brooks Camp (155° 05’ W, 58°33 ‘ N) in the western portion of Katmai National Park, observed a lone wolf traveling in the company of two Brown Bears. Szweda was driving along the park road, approx- imately 8 km from Brooks Camp, when she noticed what appeared to be two bears in the middle of the roadway, approximately 400 m ahead. The lumbering vehicle’s approach prompted the bears to abandon the roadway suddenly, exposing a wolf that had appar- ently been in their immediate company (<2 m). The wolf too left the road, following the bears into the forest. The following day, five park visitors reported see- ing two bears and a wolf cross the Brooks River at a location known locally as the Oxbow. It was generally believed that these were the same three observed by Szweda the day previous along the park road, 8 km distant. The bears and wolf were clearly traveling as SMITH, PARTRIDGE, and SCHOEN: INTERACTIONS OF BEARS AND WOLVES 249 FIGURE 1. Gray Wolf at Hallo Bay, Katmai National Park and Preserve, Alaska (Kent Fredriksson photograph). a group as they worked their way down the riverbank, probed the riverbed for the shallowest crossing, and then emerged on the other side. It was reported that the trio fed together in an unusual manner; when the bears caught fish they fed only upon the brains and skin, leaving the rest for the wolf which, sitting im- mediately adjacent, would spring forward, snatch and eat the remainder. It is not unusual to see Red Foxes (Vulpes vulpes) or gulls (Larus spp.) attempting to snatch fish scraps from Brown Bears on salmon streams. But bears in these instances are far from cooperative, often lunging at the would-be thief, catching and killing them on occasion (T. Smith, personal observation). However, in this case, the bears tolerated the wolf’s presence and, according to observers, did nothing to discourage its very close (1 m) proximity to them. Acknowledgments We gratefully acknowledge the contributions of Brian Holmes, Ronald Squibb, Tamara Olson, Kerry Sullivan, Jim Gavin, Aleska Szweda, Stephens Harper, and Layne Adams. We also thank Rick Clark and Troy Hamon of Katmai National Park for funding Brown Bear research in the park. Literature Cited Adams, L. G., F. J. Singer, and B. W. Dale. 1995. Caribou calf mortality in Denali National Park, Alaska. Journal of Wildlife Management 59: 584-594. Ballard, W. B. 1980. Brown Bear kills Gray Wolf. Canadian Field-Naturalist 94: 91. 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. Gilbert, B. K. 1989. Behavioral plasticity and bear-human conflicts. Pages 1-8 in Bear-People Conflicts: Proceedings of a Symposium on Management Strategies. Edited by M. Bromley. Yellowknife, Northwest Territories. THE CANADIAN FIELD-NATURALIST Vol. 118 Gittleman, J. L. 1986. Carnivore brain size, behavioral ecol- ogy and phylogeny. Journal of Mammalogy 67: 23-36. Harrington, F. H., and C. S. Asa. 2003. Wolf communica- tion. Pages 66-103 in Wolves: behavior, ecology and con- servation. Edited by L. David Mech and Luigi Boitani. The University of Chicago Press. Chicago, Illinois. Hayes, R. D., and D. H. Mossop. 1987. Interactions of wolves, Canis lupus, and brown bears, Ursus arctos, at a wolf den in the northern Yukon. Canadian Field-Naturalist 101: 603- 604. Hornbeck, G. F., and B. L. Horejsi. 1986. Grizzly bear, Ursus arctos, usurps wolf, Canis lupus, kill. Canadian Field- Naturalist 100: 259-260. Lent, P. C. 1964. Tolerance between grizzlies and wolves. Journal of Mammalogy 45: 304-305. Received 17 February 2003 Accepted 2 December 2003 Wild Turkey, Meleagris gallopavo silvestris, Behavior in Central Ontario During Winter Linu P. Ncuyen,! JOSEF HAMR,” and GLENN H. PARKER! 'Department of Biology, Laurentian University, Ramsey Lake Road, Sudbury, Ontario P3E 2C6 Canada *Northern Environmental Heritage Institute, Cambrian College of Applied Arts and Technology, 1400 Barrydowne Road, Sudbury, Ontario P3A 3V8 Canada 3Present address: Watershed Ecosystem Graduate Program, Trent University, 1600 West Bank Drive, Peterborough, Ontario K9J 7B8 Canada Nguyen, Linh P., Josef Hamr, and Glenn H. Parker. 2004. Wild Turkey, Meleagris gallopavo silvestris, Behavior in Central Ontario During Winter. Canadian Field-Naturalist 118(2): 251-255. Home range size, food habits, and roost site selection are described for the Eastern Wild Turkey (Meleagris gallopavo silvestris) introduced on the Precambrian Shield in central Ontario during the winters 1999 and 2000. Monthly home range size was correlated primarily to snow depth, although it was also likely associated to other factors, including food avail- ability and/or roost site availability. Ferns and allies were used more than available, whereas monocots were used less than expected. Roost site-selection was primarily influenced by tree height. If the Eastern Wild Turkey is to expand its northern range in Ontario, winter food and roost site availability may be the primary determinants for successful introductions. Key Words: Eastern Wild Turkey, Meleagris gallopavo silvestris, home range size, roost site, food availability, Ontario. The Eastern Wild Turkey (Meleagris gallopavo sil- vestris) is a large, highly adaptable gallinaceous bird that is common throughout southern Ontario as a result of introductions in the last 20 years (Bellamy 2001). For northern turkey populations, winter is the most stressful season for satisfying energy demands (Ober- lag et al. 1990). Variations in home range size and re- source selection appear to be governed by snow depth (Porter et al. 1980; Vander Haegen et al. 1989) and, to a lesser extent, ambient temperature (Oberlag et al. 1990). Numerous accounts of turkey winter biology (e.g., Glover and Bailey 1949; Wunz and Hayden 1975; Porter et al. 1980; Kilpatrick et al. 1988; Vander Haegen et al. 1989), refer primarily to studies conducted in the United States. The winter biology of turkeys in these southern habitats may be different from that of birds living in Ontario due to inherent differences in climate and habitat. Hence, we report on home range size, food habits, and roost site use by Eastern Wild Turkeys introduced to the Precambrian Shield in central Ontar- io during the winters 1999 and 2000. It was hypothe- sized that turkey movement and forage availability were inversely related to snow depth. It was also hypothe- sized that the tallest trees provided optimum wild tur- key roost sites due to their sturdiness. Study Area This study was conducted from November to March 1999/2000 and 2000/2001 near Noélville, approximate- ly 60 km southeast of Sudbury, Ontario (46°10'N, 80°25'W). The 169-km/? study area was located within the Great Lakes—St. Lawrence Ecotonal Forest Region (Rowe 1972), characterized by flat to rolling topog- raphy, interrupted by rock outcrops and narrow valleys. The habitat consisted of 20% softwood forests, 37% hardwood forests, 28% abandoned pasture and hay- fields, and 15% residential areas and rock outcrops. Beef farming was the dominant land use, with many fields cultivated for corn silage or pasture grasses. The forested areas were dominated by White Birch (Betula papyrifera) and Trembling Aspen (Populus tremuloides), interspersed with Balsam Fir (Abies balsamea), Eastern White and Red Pine (Pinus .strobus, P. resinosa), Red Oak (Quercus rubra), White Spruce (Picea glauca), Red and Sugar Maple (Acer rubrum, A. saccharum), and Eastern Hemlock (Tsuga canaden- sis). Shrubs included raspberries (Rubus spp.), Bracken Fern (Pteridium aquilinum), blueberries (Vaccinium spp.), Beaked Hazel (Corylus cornuta), and asters (Aster spp.). January, the coldest month of the year, had a mean temperature of —13.9°C and —10.5°C in 2000 and 2001, respectively. Total snowfall was 216.0 cm in 1999- 2000 (10.0% below the 30-year norm) and 328.3 cm in 2000-2001 (22.2% above the 30-year norm). Snow depths exceeded 25 cm for 38 days in 1999-2000 and 111 days in 2000-2001. Methods Capture and Radio-Tracking Wild Turkeys were captured with rocket nets (Hawk- ins et al. 1968) in southern Ontario and upper New York for introduction to the study area in February and March 1999 (10 males and 26 females) and March 2000 (13 females). Female turkeys were fitted with backpack-style, 32.5 g (1% of mean body mass), mortality-mode VHF radio-transmitters (Holohil Sys- tems Ltd., Carp, Ontario). We tracked radio-fitted birds two to four days per week, or until battery failure, from 1999 to 2001. Locations of birds were determined by 251 Diy) triangulation using >3 locations (Heezen and Tester 1967) taken less than 15 minutes apart with a 2-element H antenna and portable receiver-scanner (Model STR- 1000, Lotek Engineering Inc., Newmarket, Ontario). Average telemetry error was 156.9 m + 21.1 SE (n= 40). Home Range Monthly home range sizes were calculated using the 100% minimum convex polygon method (White and Garrott 1990). All spatial analyses were conducted using an ArcView GIS software (Environmental Sys- tems Research Institute, Redlands, California), with the Animal Movement Analysis (Hooge and Eichenlaub 1997) and Spatial Analyst Extensions. Forage Selection Active turkey feeding areas were identified by tracks, and forage plants were identified and collected. Crop contents from dead specimens provided a supplemen- tal source of information. Forage species eaten were recorded regardless of the number of bites using a modified point-quarter method (Jost et al. 1999) during 2000-2001. Point samples of available plants adjacent to turkey tracks in the herb (less than 0.5 m) and shrub (0.5 to 2.0 m) layers were selected randomly and identified. Plants were grouped into five classes: (1) mosses, (2) ferns and allies, (3) conifers and allies, (4) monocots, and (5) dicots. Roost Site Roost sites were found by (1) observing commonly used trees, (2) finding turkey droppings under trees (Hoffman 1968), (3) examining areas where Wild Tur- keys were common after snowfalls, or (4) locating radio-fitted birds on trees before sunset. Locations of roost sites were determined using a Global Positioning System (GARMIN International Inc., Olathe, Kansas). Equal numbers of random trees that may have pro- vided potential roost sites within our study area were generated in ArcView GIS. Random trees were located using a Global Positioning System, and trees that had diameter at breast height (dbh) less than 10.2 cm were discarded (Kilpatrick et al. 1988). Elevation, tree dbh, percent canopy cover, canopy density, distance to habitat edge, distance to open water, and tree height for actual and selected trees were mea- sured. Elevation was measured with an altimeter, tree dbh with a diameter tape, percent canopy cover by averaging readings at each compass directions (N, S, W, E) using a convex spherical densiometer, canopy density by counting tree trunks of dbh =10.2 cm with- in 5 m of actual roost sites, and tree height from esti- mates on a subjectively selected “average” tree. Dis- tances to habitat edge and open water were also deter- mined by plotting roost sites on Ontario Base Maps (OBM) and Forest Resource Inventory (FRI) maps. The used habitat variables either had been previously described in the literature (Tzilkowski 1971; Kilpatrick et al. 1988; Chamberlain et al. 2000), or were hypoth- THE CANADIAN FIELD-NATURALIST Vol. 118 esized correlates based on the winter biology of the species. Statistical Analyses Data were pooled to maintain sufficient sample size, unless otherwise noted (Alldredge and Ratti 1986; Thomas and Taylor 1990). Home range sizes were compared between years using the Mann-Whitney U- test. Monthly home ranges were compared using the Kruskal Wallis test (Zar 1999). Two measurements used to examine the effects of winter on monthly home range size were (1) number of days with snow depths over 25 cm (Porter et al. 1980), and (2) number of days with minimum temperature less than —16.2°C (Oberlag et al. 1990). Spearman rank-order correlation analyses were used to compare mean monthly home range size with these two variables. The number of days with snow depth over 25 cm and the number of days with minimum temperature less than —16.2°C were also compared between years using the Mann- Whitney U-test. Chi-square analysis was used to compare the fre- quencies of plants browsed by turkeys to availability (Neu et al. 1974). When forage selection differed, a Bonferroni Z-test was used to identify plants that were browsed more or less than availability (Byers et al. 1984). A series of one-way analyses of variance (ANOVA) was used to compare each habitat variable associated with roosting and random sites. Pearson cor- relation analyses were performed for the significant habitat variables. In order to minimize artifacts from environmental variations in habitat characteristics, an analysis of covariance (ANCOVA) was used to deter- mine significant habitat variables. All analyses were conducted using Statistical Package for the Social Sciences (SPSS Inc., Chicago, Illinois) with signifi- cance level set at a = 0.10. Results Home Range We obtained 441 locations from 12 radio-fitted turkeys from November 1999 to March 2000 and from November 2000 to January 2001. Mean winter home range size differed between years (U = 142.0, P =0.002), as did monthly home range size (Novem- ber: U = 9.0, P = 0.083; December: U = 8.0, P = 0.060: January: U = 3.0, P = 0.020). In addition, home range size differed among months in the two years (c? = 7.79, df = 4, P = 0.100; Table 1). Monthly home range size was correlated with the number of days with snow depth more than 25 cm (7, = —0.71, n= 59, P = 0.050) and the number of days with minimum temperature less than —16.2°C (r, = 0.75, n= 59, P = 0.031). The number of days with snow depth over 25 cm differed (U = 4.0, P = 0.095) between 1999-2000 (9.60 + 5.91 days, n = 5) and 2000-2001 (22.6 + 4.88 days, n = 5), while the number of days with mini- mum temperature less than —16.2°C did not differ (U = 10.5, P = 0.690) between 1999-2000 (10.00 + 2004 NGUYEN, HAmr, and PARKER: WILD TURKEY BEHAVIOR 255 TABLE |. Winter home range size (ha) of Eastern Wild Turkey hens in Noélville, near Sudbury, Ontario, 1999-2001 (SE = standard error of the mean). 1999-2000 2000-2001 1999-2001 (Pooled Data) n Mean SE n Mean SE n Mean SE Winter 9 249 60 4 58 19 13 204 47 Monthly November 9 399 198 5 90, 54 14 288 132 December 9 145 60 5) 4] 8 14 108 40 January 9 302 130 4 38 10 13 221 95 February 9 75 27 - re - 9 15 27 March 9 323 170 - - - 9 323 170 4.15 days, n = 5) and 2000-2001 (11.40 + 3.63 days, Pine, and White Spruce, were used in proportion to n= 5). In 1999-2000, snow was generally packed or availability. Monocots, including Quackgrass (Elymus crusted with seeps and small streams remaining un- repens) and Timothy (Phleum pratense), were used less frozen. In 2000-2001, deep, powdery snow was com- _ than available, while dicots, including ash (Fraxinus mon, and most seeps were frozen by mid-winter. spp.), aster, Beaked Hazel, burdock, Choke Cherry (Prunus virginiana), Dogwood (Cornus stolonifera), Evening Primrose (Oenothera biennis), Fireweed (Epi- lobium angustifolium), goldenrod, Northern Wild Rai- sin (Viburnum cassinoides), raspberry (Rubus spp.), Soybean, Speckled Alder (Alnus incana), Swamp Thistle (Cirsium muticum), and willow (Salix spp.) were used in proportion to availability. Forage Selection Wild Turkeys foraged primarily on clovers (Trifo- lium spp.), asters (Aster spp.), goldenrods (Solidago spp.), and fertile fronds of the Sensitive Fern (Onoclea sensibilis) in the winter 1999-2000. Turkeys avoided species such as Cattail (Typha latifolia) and Meadow Sweet (Spiraea latifolia). Crops (n = 2) contained grass seeds and Sensitive Fern spore heads in late winter, Roost Site Characteristics confirming observations of turkeys feeding in the field. Trembling Aspen, American Basswood (Tilia ameri- Ostrich Fern (Matteuccia struthiopteris), Trembling cana), Jack Pine (Pinus banksiana), Eastern White Aspen (P. tremuloides) buds, Corn (Zea mays), Smooth Pine, White Spruce, and Eastern White Cedar (Thuja Wild Rose (Rosa blanda), and ragged moss (Brachy- occidentalis) were used as roost sites. Tree heights at thecium spp.) were other winter foods. Burdocks roosting sites averaged 13.9 + 0.8 m, and were higher (Arctium spp.), Sensitive Fern, and Soybeans (Glycine _ than the tree heights (8.8 + 0.7 m) measured at ran- max) comprised 21.4%, 23.0%, and 37.4% of the tur- dom sites (F, 5, = 20.98, P = 0.0001; Table 3). Mean keys’ diet, respectively, in 2000-2001. dbh of trees associated with roost sites, 37.7 + 3.0 cm, Forages used by turkeys differed from availability was significantly larger (F, 5, = 15.69, P = 0.0007) (c? = 40.38, df = 4, P < 0.0001; Table 2). Mosses _ than tree dbhs on random sites (24.1 + 1.7 cm). Roost were used in proportion to availability while ferns and __ site topographical elevations averaged 217.1 + 7.1 m allies, including Bracken (Pteridium aquilinum) and and were significantly higher (F, 5, = 4.49, P = 0.0455) Sensitive Ferns, were selected more than available. than the 188.8 + 11.3 m measured at random sites. Can- Conifer and allies, including Balsam Fir, Eastern White opy cover (73.3 + 8.8%), tree density (849.3 + 127.7 TABLE 2. Chi-square analysis and Bonferroni Confidence Intervals for winter ground forage used by Eastern Wild Turkeys during winter in Noélville, near Sudbury, Ontario (v7 = 40.38, df = 4, P < 0.0001). Forage Forage Forage Proportion Proportion 90% Confidence Class Selected Available Selected Available Interval on Proportion Mosses 0.1 1.0 0.001 0.004 0.000-0.006° Ferns and allies 43.0 32.0 0.230 0.124 0.158-0.302° Conifers and allies 5.0 4.0 0.027 0.016 0.000-0.055° Monocots 0.1 13.0 0.001 0.050 0.000-0.006* Dicots 139.0 208.0 0.743 0.806 0.669-0.817° Total 187.2 258.0 “Proportions greater than the upper confidence limit indicates use less than available. Proportions within the confidence limit indicates use equal to availability. ‘Proportions less than the lower confidence limit indicates selection greater than available. THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 3. Comparison of physiographic characteristics of Eastern Wild Turkey roost (n = 12) and random (n = 12) sites in Noélville, near Sudbury, Ontario. Roost sites Habitat characteristic Mean SE Height (m) 13:9 0.8 Tree dbh (cm) B07 3.0 Canopy coverage (%) 133 8.8 Density (trees/ha) 849.3 127 Elevation (m) PAIN HA Distance to habitat edge (m) 97.5 Del Distance to open water (m) ABS 18.6 trees/ha), distance to clearing (97.5 + 21.1 m), and dis- tance to open water (73.5 + 18.6 m) of roost trees did not differ (all P > 0.10) from those at random sites. Height of roost site was correlated with tree dbh (r= 0.78, n = 24, P = 0.003). There was no correlation between elevation and tree dbh (r = —0.36, n = 24, P = 0.245) nor elevation and tree height (7 = 0.04, n = 24, P = 0.900). Tree height was primarily responsi- ble for the differences between roosting and random sites, regardless of the covariate tree dbh (F, 5, = 3.45, P = 0.077). Tree dbh did not differ between roosting and random sites once the covariate height was statis- tically controlled (F, 5, = 15.87, P = 0.513). Discussion Winter mean home range sizes for turkeys released in northern Ontario (204 + 47 ha) were similar to those of turkeys introduced to Indiana (Miller et al. 1985), and Iowa (Little and Varland 1981), but exceed- ed those reported in Ohio (Clark 1985) and in southern Ontario (Weaver 1989). Although previous studies reported a decline in monthly home range size be- tween November to March (Porter 1977; Miller et al. 1985), this pattern was not observed in 1999-2000. This suggested that movement in late winter (1999- 2000) was not limited by snow depth. On several occa- sions when snow covered local food resources, birds increased flight distance to otherwise unused parts of the home range. Snow depth was inversely related to winter and monthly home range size, confirming data from previous studies in Minnesota (Porter 1977), Indiana (Miller et al. 1985), Pennsylvania (Wunz and Hayden 1975), and southern Ontario (Weaver 1989). Glover and Bailey (1949) described the Wild Turkey as a nomadic feeder with a tendency to sample a wide variety of forages, primarily dictated by their avail- ability. The heavy use of ferns and allies was almost exclusively due to the selection of fertile fronds of the Sensitive Fern found in lowland hardwood sites and along seeps. Decker et al. (1991) reported that Sensitive Ferns were foraged in large quantities by Wild Turkeys because of their high nutritional content (crude protein 18.6%) and/or the high availability of Random sites Mean SE F P 8.8 0.7 20.98 0.0001 24.1 N57 15.69 0.0007 70.4 9.7 0.05 0.8303 764.3 110.9 0.25 0.6206 188.8 ARS 4.49 0.0455 64.8 7-7 1.42 0.2468 122.1 34.7 152 0.2303 this food in concentrated, relatively snow-free patches. Vander Haegen et al. (1989) reported that fields spread with manure were also important sources of winter food in Massachusetts, but the value of manure to win- tering turkeys in central Ontario was questionable, as emaciated birds were observed to ignore manure piles. Glover and Bailey (1949) reported wariness by turkeys of supplemental feeding sites with corn and oats in West Virginia when natural forage (e.g., wild grape, Vitis spp.) was absent. Standing crops, such as Soybean or clover, provid- ed nutritious food in moderate snow conditions during early winter. Soybeans contain protein, but also pro- duce trypsin inhibitors, which lower digestibility and fat absorption (McNaughton and Reece 1980, in Loesch and Kaminski 1989). Post-mortems in winter 2000- 2001 (n = 5) verified that emaciated turkeys were under severe nutritional stress, and had catabolized significant amounts of muscle tissue. Forages that were present in the study area, but may have been underestimated in use, included conifer and hardwood trees. Although primarily ground feeders, Eastern Wild Turkeys are capable of flight, but forage obtained in the tree-canopy layer is difficult to sys- tematically record. Turkeys were observed feeding on Trembling Aspen buds on several occasions; however, the results indicate that prolonged periods of deep snow severely limited food availability for Wild Turkeys in northern Ontario. Roosting sites of turkeys were the tallest and largest trees, usually conifers, which were found at higher elevations. Conifers may reduce wind speed and heat loss (Kilpatrick et al. 1988). Tzilkowski (1971) found that turkey winter roosts were dominated by large decid- uous trees in Pennsylvania, suggesting that tree height and sturdiness were important factors. However, Cham- berlain et al. (2000) suggested that turkeys roost in the nearest suitable habitat at the end of the day. Although roost sites were closer to open water than random sites, this proximity appeared to be due to forage availability. Turkeys used snow to meet winter water requirements in this study. Exum et al. (1985) argued against Wild Turkey dependence on open water in southern Ala- 1 ii 2004 bama during spring and summer; however, Kilpatrick et al. (1988) suggested that winter foods in Rhode Is- land were too low in water content to meet the species’ needs. Roosting sites may have been closer to water because of the accelerated growth of trees associated with moist environments (Kilpatrick et al. 1988). The results of this study suggest that (1) winter food and roost site availability are the primary factors deter- mining successful turkey introductions in northern On- tario, (2) stands of Soybean or Corn are not sufficient to support Wild Turkey populations unless natural foods are available, and (3) snow depth is an important parameter influencing successful reintroduction of this species on the Precambrian Shield. Acknowledgments Financial and logistic support was provided by the West Arm/French River Wild Turkey Chapter, Ontario Federation of Anglers and Hunters, Cambrian College of Applied Arts and Technology, Laurentian University, Ontario Ministry of Natural Resources, Sudbury Dis- trict, and the Ontario Living Legacy Fund. We thank J. Craig and other landowners in Noélville for allow- ing fieldwork on their property, M. N. Hall for the Ontario Base Maps and Forest Resource Inventory data, and G. M. Courtin for meteorological data. Literature Cited Alldredge, J. R., and J. T. Ratti. 1986. Comparison of some Statistical techniques for analysis of resource selection. Journal of Wildlife Management 50: 157-165. Bellamy, K. 2001. Annual Ontario Wild Turkey status reports (1992-2001). Ontario Ministry of Natural Resources, Wild- life Branch, Peterborough, Ontario. Byers, C. R., R. K. Steinhorst, and P. R. Krausman. 1984. Clarification of a technique for analysis of utilization- availability data. Journal of Wildlife Management 48: 1050-1053. Chamberlain, M. J., B. D. Leopold, and L. W. Burger. 2000. Characteristics of roost sites of adult Wild Turkey females. Journal of Wildlife Management 64: 1025-1032. Clark, L. G. 1985. Adjustment by transplanted Wild Turkeys to an Ohio farmland area. Proceedings of the National Wild Turkey Symposium 5: 33-47. Decker, S. R., P. J. Pekins, and W. W. Mautz. 1991. Nutri- tional evaluation of winter foods of Wild Turkeys. Cana- dian Journal of Zoology 69: 2128-2132. Exum, J. H., J. A. McGlincy, D. W. Speake, J. L. Buckner, and F. M. Stanley. 1985. Evidence against dependence upon surface water by turkey hens and poults in southern Alabama. Proceedings of the National Wild Turkey Sym- posium 5: 83-89. Glover, F. A., and R. W. Bailey. 1949. Wild Turkey foods in West Virginia. Journal of Wildlife Management 13: 255- 265. Hawkins, R. E., L. D. Martoglio, and G. G. Montgomery. 1968. Cannon-netting deer. Journal of Wildlife Manage- ment 32: 191-195. 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. NGUYEN, HAmr, and PARKER: WILD TURKEY BEHAVIOR Hoffman, D. M. 1968. Roosting sites and habits of Merriam’s Turkeys in Colorado. Journal of Wildlife Management 32: 859-866. Hooge, P. N., and B. Eichenlaub. 1997. Animal movement extension to ARCVIEW. Version 1.1. Alaska Biological Science Center, United States Geological Survey, Anch- orage. 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. Kilpatrick, H. J., T. P. Husband, and C. A. Pringle. 1988. Winter roost site characteristics of Eastern Wild Turkeys. Journal of Wildlife Management 52: 461-463. Little, T. W., and K. L. Varland. 1981. Reproduction and dispersal of transplanted Wild Turkeys in Iowa. Journal of Wildlife Management 45: 419-427. Loesch, C. R., and R. M. Kaminski. 1989. Winter body- weight patterns of female Mallards fed agricultural seeds. Journal of Wildlife Management 53: 1081-1087. Miller, B. K., P. D. Major, and S. E. Backs. 1985. Move- ments and productivity of transplanted Eastern Wild Tur- keys in west-central Indiana farmland. Proceedings of the National Wild Turkey Symposium 5: 233-244. Neu, C. W., C. R. Byers, and J. M. Peek. 1974. A technique for analysis of utilization-availability data. Journal of Wild- life Management 38: 541-545. Oberlag, D. F., P. J. Pekins, and W. W. Mautz. 1990. Influ- ence of seasonal temperatures on Wild Turkey metabolism. Journal of Wildlife Management 54: 663-667. Porter, W. F. 1977. Home range dynamics of Wild Turkeys in southeastern Minnesota. Journal of Wildlife Manage- ment 41: 434-437, Porter, W. F., R. D. Tangen, G. C. Nelson, and G. C. Hamilton. 1980. Effects of corn food plots on Wild Tur- keys in the upper Mississippi Valley. Journal of Wildlife Management 44: 456-462. Rowe, J. S. 1972. Forest regions of Canada. Publication 1300. Canadian Forestry Service, Department of the Environ- ment, Ottawa. Thomas, D. L., and E. J. Taylor. 1990. Study designs and tests for comparing resource use and availability. Journal of Wildlife Management 54: 322-330. Tzilkowski, W. M. 1971. Winter roost sites of Wild Turkeys in southwest Pennsylvania. Transactions of the Northeast Fish and Wildlife Conference 28: 167-178. Vander Haegen, W. M., M. W. Sayre, and W. E. Dodge. 1989. Winter use of agricultural habitats by Wild Turkeys in Massachusetts. Journal of Wildlife Management 53: 30-33. Weaver, J. E. 1989. On the ecology of Wild Turkeys reintro- duced in southern Ontario. M. Sc. thesis, University of Western Ontario, London. White, G. C., and R. A. Garrott. 1990. Analysis of wild- life radio-tracking data. Academic Press Inc., New York. Wunz, G. A., and A. H. Hayden. 1975. Winter mortality and supplemental feeding of turkeys in Pennsylvania. Proceed- ings of the National Wild Turkey Symposium 3: 61-69. Zar, J. H. 1999. Biostatistical analysis. Fourth edition. Prentice-Hall Inc., Upper Saddle River. Received 24 March 2003 Accepted 15 November 2004 Fifteenth Census of Seabird Populations in the Sanctuaries of the North Shore of the Gulf of St. Lawrence, 1998-1999 JEAN-FRANCOIS RAIL and GILLES CHAPDELAINE Canadian Wildlife Service, P. O. Box 10100, Ste-Foy, Québec G1V 4H5 Canada Rail, Jean-Francois, and Gilles Chapdelaine. 2004. Fifteenth census of seabird populations in the sanctuaries of the North Shore of the Gulf of St. Lawrence, 1998-1999. Canadian Field-Naturalist 118(2): 256-263. For the first time since the tradition began in 1925, the quinquennial census of seabirds in the Migratory Bird Sanctuaries of the North Shore of the Gulf of St. Lawrence was divided between two years in 1998-1999. Trends between 1993 and 1998-1999 were variable across species and sanctuaries. In particular, following the large decreases in both species noted in 1993, the 1998-1999 survey showed that Black-legged Kittiwakes had declined further, while the number of Herring Gulls had stabilized. Alcids were all doing well except for the Atlantic Puffin which showed severe yet unexplained drops at all major colonies. Law enforcement efforts appear reflected in seabird population trends, as well-patrolled sanctuaries such as St. Mary’s Islands seem to do well, whereas many species at the Baie des Loups and Ile a la Brume sanctuaries are far from their historical levels. In addition to a better law enforcement program, research is needed in order to identify other conservation problems that some species may be facing. Pour la premiere fois depuis 1925, l’inventaire quinquennal des oiseaux marins des refuges d’ oiseaux migrateurs de la Céte- Nord du Golfe Saint-Laurent fut mené sur deux ans, soit en 1998 et 1999. Les tendances des populations entre 1993 et 1998- 1999 étaient variables selon |’espéce et le refuge. En particulier, suite a une décroissance notée chez les deux espéces en 1993, Vinventaire de 1998-1999 a démontré que la Mouette tridactyle avait encore diminué, tandis que le nombre de Goélands argentés s’ était stabilisé. Les populations d’ Alcidés ont toutes semblé en bonne santé sauf celle du Macareux moine dont les colonies importantes ont diminué de facon sévére et inexpliquée. Les efforts de protection par l’application de la loi se reflétent visiblement dans les tendances des populations d’ oiseaux de mer, puisque les refuges les mieux surveillés comme celui des Iles Sainte-Marie vont bien, alors que plusieurs espéces aux refuges de Baie des Loups et de l’ile 4 la Brume sont loin de leurs niveaux historiques. En plus d’un meilleur programme d’ application de la loi, la recherche est nécessaire pour identifier les autres problémes de conservation auxquels certaines espéces font face. Key Words: Seabirds, populations, sanctuaries, Gulf of St. Lawrence, Atlantic Puffin, larids, alcids. Apart from its major use in providing updated infor- mation on seabird numbers and distribution through- out the migratory bird sanctuaries on the North Shore of the Gulf of St. Lawrence (Figure 1), another major ap- plication of the results of this traditional survey is in identifying trends. Follow-up censuses have been car- ried out fairly regularly since the first census in 1925 (Lewis 1925, 1931, 1937, 1942; Hewitt 1950; Tener 1951; Lemieux 1956; Moisan 1962; Moisan and Fyfe 1967; Nettleship and Lock 1973; Chapdelaine 1980, 1995; Chapdelaine and Brousseau 1984, 1991), provid- ing the trends over a period of approximately 75 years, including the present survey. This survey also provides one of the ways we monitor the broader marine envi- ronment. In effect, these wide-ranging birds are acting as our sampling agents. As human impact on the seas increases, whether directly by exploitation of resources or marine pollution, or less directly by factors such as global warming, early indications of changes are vital. Clearly any changes in the size of seabird breeding col- onies are of major importance to conservationists to ori- ent any concrete action to be taken or to seabird biol- ogists to learn more about the regulation of seabird numbers and the wide range of factors known to influ- ence the birds. This article presents the current status of the 15 ma- rine bird species breeding in the sanctuaries in 1998- 1999, and compares them with those of the 1993 census. Scientific names are given in Table 1. Methods The census of seabirds in the sanctuaries of the North Shore of the Gulf of St. Lawrence was conducted over two years. All of the technical details of this census (methods, estimates calculations, mapping of the col- onies, weather conditions) are discussed in Rail and Chapdelaine (2002) and summarized below for each family of birds. GaviiDs — We counted all Red-throated Loon nests by systematically walking around the ponds on the islands of each sanctuary. In a few exceptional cases we noted the presence of adults instead of landing on the island. HyDROBATIDS — We conducted a systematic count of active burrows. A burrow was considered active if we were able to reach an adult inside, or detect signs of recent occupation (soil freshly excavated or oily odour characteristic of petrels at the entrance of the burrow). ANATIDS — On Corossol Island, we used a system of quadrats from which we extrapolated and average density (nests/ha) for the entire area deemed suitable 256 RAIL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS _/Me du Corossol —~Betchouane Gaspé Peninsula NEW BRUNSWICK ic Seaiiwiune Soran ogg “Gu, GF Watshishou lle a la Brume Gulf of St. Lawrence f~ = 4 LABRADOR 57°00' W 52°00' N QUEBEC yew Baie de Brador “ Saint-Augustin } ib Gros Mécatina \ t ‘i Ce ony 2 oe lles Sainte-Mari ) logging es Sainte-Marie, lles aux Perroquets Baie des Loups ‘ NEWFOUNDLAND ee FiGurE |. Location of the sanctuaries of the North Shore of the Gulf of St. Lawrence. for Common Eiders (Chapdelaine 1978*). In the Bet- chouane Bird Sanctuary, we conducted a systematic nest count on Calculot Island, and (as in preceding censuses) used a system of line-transects of varying length on Innu Island (see Caughley 1977). In sanc- tuaries featuring many islands (e.g., Watshishou, ile a la Brume, Baie des Loups), we counted all the nests on at least 27% of the land area and extrapolated an average density over the entire area of all the islands. On the Iles aux Perroquets and [les Sainte-Marie sanc- tuaries, extrapolation was done after systematic counts were made on most large islands, covering 57% and 98% of total land area, respectively (but see Rail and Chapdelaine 2002 for details). PHALACROCORACIDS — Systematic counts of all Great and Double-crested Cormorant nests were carried out. At the Corossol Island colony, Double-crested Cormo- rant nests in treetops were counted from an elevated lookout point. Larips — In the large Herring Gull colony of Coros- sol Island, we sampled sub-colonies where the number of nests (Np) and the number of adults (Ni) were determined. Then, using the factor k (k=Np/Ni), we estimated the number of pairs in sectors where we counted only the number of adults present. In the Betchouane Sanctuary, all gull nests were counted sys- tematically, and were attributed to Herring and Great Black-backed gulls according to the observed propor- tions of adults of both species. In the other sanctu- aries all larids were censused with a combination of est counts and adult counts. Proportions of Arctic and Common terns in mixed tern colonies were not always determined, so the results for the two species are com- bined in Table 1. All Black-legged Kittiwake nests attended by adults were counted. ALCIDS — We carried out systematic counts of eggs and active burrows in colonies of Atlantic Puffins, Razorbills and Common Murres that could be accessed with minimal disturbance. Elsewhere we counted indi- viduals at the colony and on adjacent waters from a distance. All colonies with large numbers of Common Murres were carefully avoided because of their sensi- tivity to disturbance. In particular, at the Iles Sainte- Marie Sanctuary, where 87% of breeding Common Murres on the North Shore are found, 98% of the population estimate for the Common Murre results from adult counts. For Atlantic Puffins nesting on the Blacklands Island in the Baie des Loups Sanctuary, we used the factor k method described above for larids. On Ile aux Perroquets in the Baie de Brador Sanctuary, we used evenly spaced grids and line transects to de- limit the area of both the puffin and Razorbill colonies, evaluate mean nest densities, and then extrapolated the populations (see Nettleship 1973* ; Chapdelaine 1978*). The Black Guillemot population was estimated from adult bird counts around the islands, although exceptionally we recorded a pair when a nest was found with no adult in sight. 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YOR] Coane 99 (6 2 (a7 ge 06 0) (Ale (0)! Z (6 DIDIJAIS DIAV) UOOT PayvOI)-poy 66-8661 £661 6661 £661 6661 £661 6661 £661 6661 £661 6661 £661 6661 £661 8661 £661 8661 £661 8661 <66l satoodg [PI0OL, Jopeig BUILD OLIRIAJ-JUTRS sjanbo.lag sdnoq unig noysrysieA ouenoyneq [OSSO1OD, ap oeg SOID Sel] Xne Sol] sap aieg PR OT] Op 9] 258 "66-8661 PUP £66 POUDIME'T ‘1S JO J[ND aup Jo a1oYS YON ay} JO soLMNjouRs pI dy) UI (STENPIAIPUT JO JaquUUNU) spIqvas JO snsUd_ “| ATAV], 2004 RaiL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 259 TABLE 2. Changes in the numbers of seabirds in sanctuaries on the North Shore of the Gulf of St Lawrence, Québec, 1988 to 1998-1999. Years of survey Compound annual growth* rate by period Species 1988 1993 1998-1999 1988-1993 1993-1998** Red-throated Loon 76 66 80 -2.82% 3.21% Leach’s Storm-Petrel 1744 1840 718 1.07% -18.78% Great Cormorant 86 78 340 -1.95% 24.54% Double-crested Cormorant 4558 3472 2830 -5.44% -3.49% Common Eider 8536 14548 13072 10.66% -2.10% Ring-billed Gull 288 104 484 -20.37% 31.04% Herring Gull 16195 4879 4929 -24.00% 0.49% Great Black-backed Gull 1883 2284 2381 3.86% 1.15% Black-legged Kittiwake 8536 6294 3816 -6.09% -10.02% Caspian Tern 15 0 0 -100.00% - Common and Arctic Terns 1350 545 294 -18.14% -12.03% Common Murre 26049 30829 30111 S370 -0.39% Razorbill 7036 8389 13953 35200 8.58% Black Guillemot 521 411 651 -4.74% 8.70 % Atlantic Puffin 35142 46684 29030 5.68% -7.92% log. N (t) - log. N (0) so — 100% t ** Populations in sanctuaries censused in 1999 were back-calculated to 1998 to get the compound annual growth rate for all North Shore sanctuaries after 5 years Results In the eight sanctuaries censused in 1993 and 1998- 1999 (Gros Mecatina excluded), the total number of birds had decreased by 15% or 17 734 individuals (see Table 1 for details). This is mostly due to the dra- matic 38% decline of the Atlantic Puffin (-17 654 indi- viduals), which was by far the most abundant seabird in the North Shore sanctuaries in 1993 (but was out- numbered by the Common Murre in 1999). Leach’s Storm-Petrels, Black-legged Kittiwakes and Common and Arctic Terns also declined at a very rapid rate between 1993 and 1998-1999 (Table 2). Double-crested Cormorants numbers also diminished. The Razorbill is the only abundant species showing a large increase, with the 1998-1999 population estimate exceeding by 5564 individuals the result of 1993 (+66%). This species has made a spectacular comeback since 1972, when less than 3000 individuals were recorded in the sanctuaries (compared to 14 341 in 1998-1999). The small popula- tions of Great Cormorants, Ring-billed Gulls and Black Guillemots also grew rapidly between 1993 and 1998- 1999, as did Red-throated Loons but more slowly. In four species, numbers appeared rather stabilized, the Common Eider and Herring Gull after having showed very sharp trends between 1988 and 1993 (annual growth rates of +11% and -24%, respectively), the Great Black-backed Gull and Common Murre follow- ing less striking but significant rises (annual growth rate of +3.9 and +3.4%, respectively). Finally, no Cas- pian Terns were found for the second quinquennial census in a row. We visited the Corossol Island Sanctuary on 30 and 31 May 1998. Major increases in large gulls (Herring and Great Black-backed gulls) were noted. Populations of alcids also expanded, and in particular the number of Black Guillemots more than doubled compared to 1993. Puffins were recorded for the first time during quinquennial censuses, bringing up to 10 the number of seabird species breeding there. The Common Eider was also more abundant. On the other hand, the colony of Leach’s Storm-Petrels was reduced to less than half between 1993 and 1998, and Black-legged Kittiwakes also declined quite rapidly. Double-crested Cormorants were down 25%, and colony localization had seemed to change a bit, possibly prompted by forest decay caused by their long-term occupation (nests are in tree- tops of conifers) in some areas. On 2 and 3 June 1998, we visited the Betchouane Bird Sanctuary. The most important change in the sea- bird community was the 36% decrease in Common Eider, by far the most abundant bird there. The Razor- bill apparently continued its spectacular comeback, with a 72% population increase between 1993 and 1998. Herring Gulls, Great Black-backed Gulls, Black- legged Kittiwakes and Atlantic Puffins were almost stable or slightly increasing. 260 After a tremendous population explosion following 1982, the population growth of the Common Eider in Watshishou appeared to have slowed down consid- erably between 1993 and 1998. The Black Guillemot population increased threefold during the same period. Double-crested Cormorant and Great Black-backed Gull numbers showed opposite trends; the former spe- cies underwent a 50% increase whereas the latter was reduced by half. Herring Gull population size appeared slightly reduced compared to 1993, but the 1998 result was by far the highest for the species in Watshishou, except for 1993. The 1998 survey confirmed a cata- strophic (-96%) and inexplicable decline of terns in Watshishou since 1988. In the past, this sanctuary, con- sisting of more than 200 islands, always had a fair number of terns (range 128 — 1490 individuals), and was the most important sanctuary on the North Shore in that regard. In 1998, however, despite better cover- age (more islands were visited than in previous cen- suses), we found only 19 breeding pairs distributed in 5 very small colonies. The Watshishou Bird Sanc- tuary was visited on 6, 7 and 8 June 1998. The following year (in 1999), seabirds were cen- sused on 14 June at the Ile 4 la Brume Sanctuary. The Common Eider had decreased slightly since 1993, but was still relatively abundant. Population trends were up for Herring and Ring-billed gulls; however numbers of these two species are rather small and tended to vary a lot in the past. The number of Great Black-backed Gulls dropped further between 1993 and 1999, reaching its lowest level ever in 1999. Terns declined sharply during the same period, and in fact since 1988 their population trends at the Ile a la Brume (-84%) and Watshishou sanctuaries are similar. Status of the very small populations of Red-throated Loon and Razorbill are still precarious, as their numbers remained stable between 1993 and 1999 at two and eight individuals, respectively. Finally, no Caspian Terns were found for the second quinquennial census in a row at Ile a la Brume, the only traditional breed- ing site for this species in the Province of Québec. We visited the Baie des Loups Bird Sanctuary on 18 June. The most abundant bird there, the Atlantic Puffin, had declined by 40% since 1993. The Great Black-backed Gull population also decreased notice- ably (36%). Breeding Leach’s Storm-Petrels apparent- ly deserted the sanctuary, which leaves the Corossol Island Sanctuary as the only active breeding site of that species on the North Shore, and maybe in the whole province. On the brighter side, the Common Eider pop- ulation increased by 50%. Also, the Common Murre and the Razorbill, once abundant in the sanctuary but which had declined steadily up to the very low and worrying levels observed in 1993, bounced back and increased threefold between 1993 and 1999. Red- throated Loon, Herring Gull, terns, and Black Guille- mot numbers remained fairly stable. THE CANADIAN FIELD-NATURALIST Vol. 118 The Iles aux Perroquets Bird Sanctuary was visited on 11 and 21 June 1999. The population of Atlantic Puffin showed a similar drop (-53%) as in the other large colonies on the North Shore. In contrast to the general upward trend observed in these species at the other sanctuaries, a decrease in Common Murres and Razorbills was noted at the Iles aux Perroquets Bird Sanctuary in 1999. The number of Common Eiders declined only slightly. Except for the Black-legged Kittiwakes whose population collapsed (-82%) between 1993 and 1999, numbers of larids were quite stable. With only 11 nests found in 1999, the kittiwake may well disappear as a breeding species at the Iles aux Perroquets in the future. On the positive side, the two smallest seabird populations breeding in the sanctuary in 1993, those of the Red-throated Loon and Black Guillemot, both showed large increases (140% and 164%, respectively). Also worthy of notice is the addi- tion of the Great Cormorant as a breeding species in the sanctuary. The Iles Sainte-Marie Bird Sanctuary (visited on 10, 11, 13, 17 and 19 June) was again found to have the highest seabird diversity (12 breeding species) and abundance (total of 39 316 individuals). Common Murres represented two-thirds of the seabird popula- tion with an estimated 26 156 breeding birds. The number of murres appeared quite stable between 1993 and 1999, after the spectacular and steady recovery of the species following 1972 (when only 4120 birds were counted). Razorbills show the same long-term trend as murres, but were still going up in 1999 (116% rise compared to 1993). The Great Cormorant colony expanded unexpectedly (+318%) to become the largest of this species on the North Shore. Increases were noted in Black-legged Kittiwakes, Black Guillemots and terns as well. Numbers of Double-crested Cormorants, Com- mon Eiders, and Atlantic Puffins were reduced rough- ly by half. Herring Gulls were again found to be declin- ing very sharply, whereas Great Black-backed Gulls showed a moderate decrease. Finally, the number of Red-throated Loons in 1999 was only one pair short of the highest counts which were recorded on the two pre- ceding censuses. A little farther to the east, a few small offshore islands constitute the Gros Mécatina Bird Sanctuary, which was created in 1996. On 23 June 1999, the sanc- tuary was included for the first time in a quinquennial census. Species diversity was impressive for such a small sanctuary, as nine seabird species were found breeding. Numbers of each species were low though, and the most abundant species was the Razorbill with 388 birds observed. According to naturalists such as Coues (1862), these islands were once inhabited by tens of thousands of Common Murres, so the sanctuary has definite potential for seabird conservation if it is well protected. 2004 The Baie de Brador Bird Sanctuary is well-known for its colonies of Atlantic Puffin and Razorbill. On 28 and 29 June 1999 however, we were quite surprised to discover the addition of Black-legged Kittiwakes and terns breeding there for the first time, as well as Common Murres which had not bred there since 1965. Herring Gulls and Great Black-backed Gulls seemed well established as breeders for the second census in a row (they were present in very small num- bers or absent prior to 1993). Also, as in most other sanctuaries, Razorbill numbers grew very fast between 1993 and 1999 (+175%). Nevertheless, all these results were darkened by the finding that Atlantic Puffins had declined by a third since 1993, when the population seemed on its way to get back to historic levels. Because it has always been the stronghold of puffins in our province, the Baie de Brador Bird Sanctuary plays a leading role in the conservation of the species in Québec. Discussion Total number of seabirds recorded in 1998-1999 in the North Shore Bird Sanctuaries dropped 15% (or 17 734 birds, Gros Mécatina excluded). But most of this decrease could be attributed to the Atlantic Puffin whose 38% decline represent a loss of 17551 birds. Population trends in other species were variable, as four species increased rapidly, four species decreased at a fast rate, and the others showed more stable or moderate trends. Numbers of Razorbills and Black Guillemots increased at a very rapid rate in almost every sanctuary (where they are present). The only Great Cormorant colony recorded during the 1993 census was found at the [les Sainte-Marie Bird Sanctuary; by 1999, that colony had grown so much that immigration must be partly involved. Also, three new colonies of this species were found elsewhere in the sanctuaries in 1999. The Ring-billed Gull is the last seabird species which increased at a very fast rate since 1993. How- ever, this species is poorly represented in the North Shore Bird Sanctuaries, where their numbers always fluctuated widely in the past. Instead of suggesting a real population trend, our results may rather reflect colony displacements as the species exhibits poor site fidelity on the North Shore. The Atlantic Puffin was not the only species show- ing a sharp decline between the last two censuses of the North Shore sanctuaries. The case of the Leach’s Storm-Petrel is equally worrying; the species was found breeding in four sanctuaries in 1988, but in 1998- 1999 only the Corossol Island colony was still active, and its number of nests was reduced to less than half compared to 1993. Moreover, this may be the only Leach’s Storm-Petrel colony left in the whole province. Thus the status of the species appears precarious in Québec. The Black-legged Kittiwake was found breed- ing in six sanctuaries in 1998-1999, whereas in the past it had been breeding in no more than four sanctuaries. RAIL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 261 However, over 90% of the kittiwakes in the sanctu- aries are concentrated at the Corossol Island colony, where a 39% drop was observed between 1993 and 1998. In fact the colony decreased by 55% since 1988 and the trend was even faster in the last five years. Observations made on diet and breeding success of larids at Corossol Island in 1996-1998 showed that kittiwake breeding success was severely affected by Herring Gull predation on large chicks, especially when capelin appeared less available. The poor kittiwake productivity measured in 1997-1998 (0.60 and 0.27 young fledged per pair, respectively) means that re- cruitment is likely not enough to compensate for nat- ural mortality, so that further decline of the colony is expected. Terns, after the fashion of Ring-billed Gulls, are poorly represented in the sanctuaries and their population trends partly reflect the unpredictability of their breeding activities. However, their numbers dropped severely for a second quinquennial census in a row, resulting in a 80% decline since 1988. The case of the Watshishou Bird Sanctuary, which has always been the most important for terns, is particularly intrigu- ing as that tern population reached an unprecedented low level in 1998. Among species with near-stable population trends between 1993 and 1998-1999 are the Common Eider and the Common Murre, two species whose numbers finally appeared to level off after spectacular increases were noted in each of the four preceding censuses. Herring Gull population trends on the North Shore (and especially at Corossol Island) have been corre- lated with cod fishery, an industry that provided signif- icant amounts of fish offal available to gulls (Chap- delaine and Rail 1997). After a boom in the early 1980s, cod fishery and Herring Gull populations collapsed concurrently in the late 1980s-early 1990s. But in 1998- 1999 it seemed that after a drastic (-78%) decline be- tween 1988 and 1993, the Herring Gull population was stabilized, perhaps to a more “natural” level. The num- ber of Red-throated Loons increased only slightly be- tween the last two surveys, yet the 1998-1999 total (82 individuals) is the highest ever recorded in the sanctuaries. The Double-crested Cormorant population declined but only at a slow rate after 1993, and in 1998- 1999 the total population estimate for the sanctuaries was still the third highest since 1925. The steady rise of Great Black-backed Gull numbers continued to slow down to be nearly stable between 1993 and 1998. Final- ly, the Caspian Tern did not breed in 1993 and 1999 at the fle 4 la Brume Sanctuary, where it consistently nest- ed between 1925 and 1988. The more time passes by, the more unlikely this species will be back, unless appropriate measures are taken to prevent disturbance of the only breeding site of Caspian Terns in Québec. While some population trends varied regionally, others suggest large-scale factors are involved. In particular, Black-legged Kittiwake and Herring Gull declines since the end of the 1980s are reported not only 262 in North Shore sanctuaries, but also at all large colo- nies around the Gaspé Peninsula and at the Magdalen Islands (Chapdelaine et al. 2003*). Atlantic Puffin num- bers dropped at all major North Shore concentrations between 1993 and 1999. On the other hand, Razorbill colonies continued to expand everywhere in the Gulf and in the Estuary (Chapdelaine et al. 2001), and the small populations of Great Cormorant on the North Shore and around the Gaspé Peninsula grew signifi- cantly since 1988-1990. So far we have presented some evidence that the Herring Gull population had been influenced by cod fishery activities (Chapdelaine and Rail 1997), and that an abundance of small fish such as sandlance (Ammodytes sp.) and Capelin (Mallotus villosus) had a positive effect on alcid breeding per- formance and populations (Chapdelaine and Brousseau 1991, 1996; Rail et al. 1996). But most population trends are left unexplained. The fact that law enforcement efforts have not been distributed uniformly across sanctuaries probably ex- plains why, in sanctuaries such as Ile a la Brume and Baie des Loups, many seabird populations are at low levels and signs of disturbance are omnipresent. Con- versely, the well-patrolled [es Sainte-Marie Bird Sanctuary appears in good shape. Seabird conservation in North Shore bird sanctuaries face many challenges: ecotourism, disturbance, poaching, episodic fox inva- sion in the sanctuaries, seabird bycatch in nearby fish- ing nets, along with ecological factors at all scales. This stresses the need to maintain and improve law enforce- ment and public educational programs, and highlights the importance of research to tackle the factors respon- sible for the observed seabird population trends. Acknowledgments Many thanks to all those who contributed to this. Fieldwork was carried out with the help of our col- leagues Pierre Brousseau and Jocelyn Thibault, Gilles Falardeau with his assistants Linda Burr and Daniel Daigneault, and Canadian Wildlife Service wardens Freddy Strickland and Wilson Evans. Members of Parks Canada in Mingan and assisting volunteers also gave us a big hand, as well as Andrew Rowsell from Fish- eries and Oceans Canada in Blanc-Sablon. Finally, our special thaughts go to M. Gallienne and his family for their hospitality on Corossol Island. Documents Cited (marked * in text) Chapdelaine, G. 1978. Onziéme inventaire des oiseaux colo- niaux des refuges de la c6te nord du Golfe Saint-Laurent et révision globale de la fluctuation des populations depuis 1925 jusqu’a 1977. Service canadien de la faune, région du Québec, Environnement Canada, Sainte-Foy, 166 pages. Chapdelaine, G., P. Brousseau, and J.-F. Rail. 2003. Banque informatisée des oiseaux marins du Québec (BIOMQ). Database. Last update 12 March 2003. Environ- ment Canada, Canadian Wildlife Service, Québec Region. Nettleship, D. N. 1973. Census of seabirds in the sanctuaries of the North Shore of the Gulf of St. Lawrence, summer THE CANADIAN FIELD-NATURALIST Vol. 118 1972. Studies on northern seabirds No 20. Canadian Wild- life Service Report, Ottawa, 160 pages. Literature Cited Caughley, G. 1977. Analysis of vertebrate populations. John Wiley and Sons, London. Chapdelaine, G. 1980. Onziéme inventaire et analyse des fluctuations des populations d’oiseaux marins dans les refuges de la Cote Nord du Golfe Saint-Laurent. Cana- dian Field-Naturalist 94: 34-42. Chapdelaine, G. 1995. Fourteenth census of seabird popu- lations in the sanctuaries of the North Shore of the Gulf of St. Lawrence, 1993. Canadian Field-Naturalist 109: 220-226. Chapdelaine, G., and P. Brousseau. 1984. Douzieme inven- taire des populations d’ oiseaux marins dans les refuges de la Cote-Nord du golfe du Saint-Laurent. Canadian Field- Naturalist 98: 178-183. Chapdelaine, G., and P. Brousseau. 1991. Thirteenth cen- sus of seabird populations in the sanctuaries of the North Shore of the Gulf St. Lawrence, 1982-1988. Canadian Field-Naturalist 105: 60-66. Chapdelaine, G., and P. Brousseau. 1996. Diet of Razor- bill Alca torda chicks and breeding success in the St. Mary’s Islands, Gulf of St. Lawrence, Québec, Canada, 1990-1992. Pages 27-37 in Studies of high-latitude sea- birds. 4. Trophic relationships and energetics of endotherms in cold ocean systems. Edited by W. A. Montevecchi. Canadian Wildlife Service Occasional Paper 91. Ottawa. Chapdelaine, G., A. W. Diamond, R. Elliot, and G. J. Robertson. 2001. Status and population trends of the Razorbill in eastern North America. Canadian Wildlife Service Occasional Paper 105. Ottawa. Chapdelaine, G., and J.-F. Rail. 1997. Relationship between cod fishery activities and the population of herring gulls on the North Shore of the Gulf of St Lawrence, Québec, Canada. ICES Journal of Marine Science 54: 708-713. Coues, E. 1862. Notes on the ornithology of Labrador. Pro- ceedings of the Academy of Natural Sciences of Phila- delphia (1861): 215-257. Hewitt, O. H. 1950. Fifth census of non-passerine birds in the bird sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 64: 73-76. Lemieux, L. 1956. Seventh census of nonpasserine birds in the bird sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 70: 183-185. Lewis, H. F. 1925. The new bird sanctuaries in the Gulf of St. Lawrence. Canadian Field-Naturalist 39: 177-179. Lewis, H. F. 1931. Five years’ progress in the bird sanctu- aries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 45: 73-78. Lewis, H. F. 1937. A decade of progress in the bird sanctu- aries of the North Shore of the Gulf of St. Lawrence. Can- adian Field-Naturalist 51: 51-55. Lewis, H. F. 1942. Fourth census of non-passerine birds in the bird sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 56: 5-8. Moisan, G. 1962. Eighth census of non-passerine birds in the bird sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 76: 78-82. Moisan, G., and R. W. Fyfe. 1967. Ninth census of non- passerine birds in the sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian-Field-Naturalist 81: 67-70. 2004 Nettleship, D. N., and A. R. Lock. 1973. Tenth census of seabirds in the sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 87: 395-402. Rail, J.-F, and G. Chapdelaine. 2002. Quinziéme inven- taire des oiseaux marins dans les refuges de la Cote-Nord : techniques et résultats détaillés. Série de rapports tech- niques No. 392. Service canadien de la faune, région du Québec, Environnement Canada, Sainte-Foy, xvi + 307 pages. Rail, J.-F., G. Chapdelaine, P. Brousseau, and J.-P. L. Savard. 1996. Utilisation des oiseaux marins comme bio- RAIL and CHAPDELAINE: CENSUS OF SEABIRD POPULATIONS 263 indicateurs de l’écosystéme du Saint-Laurent. Série de rap- ports techniques No. 254. Service canadien de la faune, région du Québec, Environnement Canada, Sainte-Foy, ii + 113 pages. Tener, J. S. 1951. Sixth census of non-passerine birds in the bird sanctuaries of the North Shore of the Gulf of St. Lawrence. Canadian Field-Naturalist 65: 65-68. Received 16 June 2003 Accepted 5 July 2004 Notes Observations of Above-Surface Littoral Foraging in Two Sea Ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in Coastal Southwestern British Columbia DEBORAH L. LACROIX!, KENNETH G. WRIGHT?, and DANIEL KENT? 'Centre for Wildlife Ecology, Simon Fraser University, Burnaby, British Columbia VSA 1S6 Canada 26090 Blink Bonnie Road, West Vancouver, British Columbia V7W 1V8 Canada >Vancouver Aquarium Marine Science Centre, Vancouver, British Columbia V6B 3X8 Canada Lacroix, Deborah L., Kenneth G. Wright, and Daniel Kent. 2004. Observation of above-surface littoral foraging in two sea ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in coastal southwestern British Columbia. Canadian Field-Naturalist 118(2): 264-265. Barrow’s Goldeneyes (Bucephala islandica) and Surf Scoters (Melanitta perspicillata) were observed on four separate occa- sions, by three different observers, foraging on Bay Mussels (Mytilus trossulus) above the water surface. This unique foraging behaviour could be attributed to diurnal spring tides and reduced lower intertidal mussel abundance. Key Words: Barrow’s Goldeneye, Bucephala islandica, Surf Scoter, Melanitta perspicillata, foraging behaviour, sea ducks, Bay Mussels, Mytilus trossulus, British Columbia. The sea ducks, Barrow’s Goldeneye (Bucephala islandica) and Surf Scoter (Melanitta perspicillata), winter in abundance along the coast of British Colum- bia where they forage diurnally on marine inverte- brates in bays, harbours, beaches, and inlets (Bellrose 1980; Vermeer 1981, 1982; Vermeer and Bourne 1984; Campbell et al. 1990; Savard et al. 1998; Eadie et al. 2000). In rocky habitats, both sea ducks forage pre- dominantly on Bay Mussels (Mytilus trossulus) (for- merly classified as M. edulis; McDonald and Koehn 1988, McDonald et al. 1991) (Vermeer and Levings 1977; Hirsch 1980; Vermeer 1981, 1982; Vermeer and Bourne 1984; Lacroix 2001). Both sea ducks dive to locate and retrieve mussels. Mussels are pried from the substrate and swallowed whole (Savard et al. 1998; Eadie et al. 2000). Three observers, on four separate occasions, ob- served Surf Scoters and/or Barrow’s Goldeneyes for- aging on Bay Mussels above the water surface in coastal southwestern British Columbia. The first observation, on 22 February 1999, involved juvenile male and fe- male Surf Scoters and Barrow’s Goldeneyes eating exposed mussels at a breakwater on Popham Island. The event was photo-documented by D. Kent. On the second occasion, on 7 March 1999, 1 juvenile male and 7 females, mostly juveniles, Barrow’s Goldeneyes and 3 Surf Scoters, (1 female and 2 immature males), re- moved mussels from a rocky point at Cape Roger Curtis, while they sat on a reef, 1.5 m above the water line. Later that day a flock of approximately 12 Bar- row’s Goldeneyes, mostly juveniles, was seen feeding on mussels while hauled out on a reef on Hermit Island. This above-surface foraging tactic was also noticed later in March when three juvenile Surf Scoters, and six Barrow’s Goldeneyes, mostly juveniles, fed on ex- posed mussels on large boulders. All the observations coincided with low diurnal tides. The tide height ranged between 1.74 to 2.16 m, above Chart Datum (a.c.d.). We are unaware of any previously published or unpub- lished accounts of this foraging behaviour. Our multiple observations suggest that the behaviour occurs frequent- ly but has gone unreported. All of these observations were made during late February and March in Howe Sound and Burrard Inlet, British Columbia, located in the same geograph- ic vicinity (49°19.32’N, 123°09.92’W — 49°21.60’N, 123°29.15’W). These observations share several sim- ilar characteristics including: (1) all ducks were feed- ing on Bay Mussels; (2) the observations were made during low diurnal tides; (3) only small groups, often consisting of mixed-species flocks, were exhibiting this foraging behaviour, and (4) these foraging groups consisted mostly of juveniles. Discussion Bay Mussels are a dominant species in protected coastal rocky intertidal areas (Seed and Suchanek 1992: Ricketts et al. 1995). In our observation area, the vertical distribution of the Bay Mussel ranges from 1.5 to 3.7 m, a.c.d. (Quayle 1978). Although Bay Mus- sels are intertidal, mussel beds are rarely completely exposed during daylight hours in winter as the lowest low tides of the semidiurnal tidal regime occur at night (Thomson 1981). As winter advances into spring, the lowest low tides are diurnal; therefore expose mussel beds during daylight hours (Figure 1). Exposed mussels 264 2004 FIGURE | Percent of daylight hours fully submerged Oct Nov Dec Jan Feb Mar Apr May Time Ficure |. The percent of daylight hours that Bay Mussel beds are completely submerged in coastal British Columbia. Mussel submergence time was estimated by calculat- ing the number of hours the tide was above 3.7 m, a.c.d, using Canadian tide charts (1999), during day- light hours, between sunrise and sunset. may be more vulnerable to diving ducks such as Surf Scoter and Barrow’s Goldeneye as the ducks do not need to expend energy diving to reach the mussels. However, Smeathers and Vincent (1979) found that mussels exposed to air have twice the tensile strength of those submerged. If the exposed mussels have un- dergone some degree of desiccation, they may require more energy to remove than submerged ones, and may therefore, not be as profitable as first postulated. Alternatively, juvenile and sub-adult Surf Scoters and Barrow’s Goldeneyes may be forced to feed on exposed mussels owing to the reduced mussel abundance and distribution from over-winter predation. The lower por- tion of the distribution of Bay Mussels is determined by biological factors, primarily predation from the Ochre Sea Star (Pisaster ochraceus) (Seed and Sucha- nek 1992; Quayle 1978) and sea ducks (Lacroix 2001). Through the winter, the combination of sea star and sea duck predation may eliminate the lower distribution of mussels, hence reducing their overall abundance. It is therefore plausible that the ducks observed were forced to forage on the less profitable prey (1.e., the ex- posed mussels) because there are few or no submerged mussels in the lower portion of their distribution. Acknowledgments We thank R. W. Butler and A. J. Erskine for making valuable suggestions to the manuscript. D. L. Lacroix received funding from a postgraduate NSERC schol- arship and from the Canadian Wildlife Service Geor- gia Basin Initiative. Literature Cited Bellrose, F. C. 1980. Ducks, geese and swans of North Amer- ica. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, and G. W. Kaiser. 1990. The birds of British NOTES 265 Columbia. Volume |. Nonpasserines, introduction and loons through waterfowl. Royal British Columbia Museum, Victoria, British Columbia. 513 pages. Canadian Hydrographic Service, Fisheries and Oceans Canada. 1999. Canadian tide and current tables. Volume 5. Minister of Fisheries and Oceans Canada. 120 pages. Eadie, J. M., J.-P. Savard, and M. L. Mallory. 2000. Barrow’s Goldeneye (Bucephala islandica). Pages 1-32 in The Birds of North America. (548). Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, Pennsylvania and the American Ornithologists’ Union, Washington, D.C. Hirsch, K. V. 1980. Winter ecology of sea ducks in the inland marine waters of Washington. M.Sc. thesis, University of Washington, Seattle, Washington. 92 pages Lacroix, D. L. 2001. Foraging impacts and patterns of win- tering surf scoters feeding on bay mussels in coastal Strait of Georgia, British Columbia. M.Sc. thesis, Simon Fraser University, Burnaby, British Columbia. 126 pages. McDonald, J. H., and R. K. Koehn. 1988. The mussels Mytilus galloprovincialis and M. trossulus on the Pacific coast of North America. Marine Biology 99: 111-118. McDonald, J. H., R. Seed, and R. K. Koehn. 1991. Allo- zymes and morphometric characters of three species of Mytilus in the Northern and Southern Hemispheres. Marine Biology 111: 323-333. Quayle, D. B. 1978. A preliminary report on the possibili- ties of mussel culture in British Columbia. Fisheries and Marine Service Technical Report 815. 37 pages. Ricketts, E. F., J. Calvin, J. W. Hedgpeth, and D. W. Philips. 1995. Between pacific tides. Fifth Edition. Stanford Uni- versity Press, Stanford, California. Savard, J-P. L., D. Bordage, and A. Reed. 1998. Surf Scoter (Melanitta perspicillata). Pages 1-28 in The Birds of North America. (363) Edited by A. Poole and F. Gill. The Aca- demy of Natural Sciences, Philadelphia, Pennsylvania and the American Ornithologists’ Union, Washington, D.C. Seed, R., and T. H. Suchanek. 1992. Population and com- munity ecology of Mytilus. Pages 87-157 in The mussel Mytilus: ecology, physiology, genetics and culture. Edited by E. Gosling. Developments in Aquaculture and Fisher- ies Science 25, Elsevier, Amsterdam. 589 pages. Smeathers, J. B., and J. F. Vincent. 1979. Mechanical prop- erties of mussel byssus threads. Journal of Molluscan Studies 45: 219-230. Thomson, R. E. 1981. Oceanography of the British Columbia coast. Canadian Special Publication Fisheries and Aquatic Science 56. 291 pages. Vermeer, K. 1981. Food and populations of Surf Scoters in British Columbia. Wildfowl] 32: 107-116. Vermeer, K. 1982. Food and distribution of three Bucephala species in British Columbia. Wildfowl 33: 22-30. Vermeer, K., and N. Bourne. 1984. The White-winged Scoter diet in British Columbia waters: resource partitioning with other scoters. Pages 62-73 in Marine birds: their feeding ecology and commercial fisheries relationships. Edited by D. N. Nettleship, G. A. Sanger, and P. F. Springer. Pro- ceedings of the Pacific Seabird Group Symposium, Seattle, Washington, 1982. Vermeer, K., and C. D. Levings. 1977. Populations, biomass and food habits of ducks on the Fraser Delta intertidal area, British Columbia. Wildfowl 28: 49-60. Received 23 July 2001 Accepted 6 August 2004 266 THE CANADIAN FIELD-NATURALIST Vol. 118 New Records of Cyperaceae and Juncaceae from the Yukon Territory Stuart A. HARRIS! AND PETER W. BALL? 'Department of Geography, University of Calgary, Calgary, Alberta T3A 1E4 Canada *Department of Botany, Erindale College, University of Toronto in Mississauga, Mississauga, Ontario LSL 1C6 Canada Harris, Stuart A., and Peter W. Ball. 2004. New records of Cyperaceae and Juncaceae from the Yukon Territory. Canadian Field-Naturalist 118(2): 269-270. Two new species of Cyperaceae are reported, viz., Carex hoodii, and Eleocharis elliptica. Also, range extensions for eight species of Carex, Eriophorum, and Juncus are listed. Key Words: Vascular plants, Yukon Territory, new records, range extensions. In the last ten years, considerable advances have taken place in the knowledge of the vascular flora of the Yukon Territory (Cody, 1996, 2000; Cody et al., 1998, 2000, 2001, 2002, 2003). In spite of this, the flora is still incompletely known, as indicated by the frequent and substantial additions. In 1980, the first author commenced detailed field studies of the distribution of permafrost in the south and central Yukon Territory, together with the nature and dynamics of the associated landforms (Harris 1998, 2004; see also Wall et al. 1987). As part of the work, plants were collected and have been system- atically identified. For the more difficult groups, the material was sent to specialists who have helped in the identification. P.W.B. carried out the task of iden- tifying the Cyperaceae and some of the Juncaceae. The result is a substantial collection of sheets that are in the collection of the first author (UAC #60 000 — 74 000). Comparison of these with the published lists of species and their distributions indicates that there are at least two species not previously described for the Yukon Territory, as well as some range extensions and name changes of varying importance. This paper will describe these changes. The nomenclature used follows that in the recent Flora of North America, volume 23 (Flora of North America Editorial Com- mittee 2002). Species New to the Yukon CYPERACEAE Carex Hoodii Boott — YUKON. Hart River road at 1180 m elevation, east of the Dempster Highway at km 78, about 138°14'W., 64°30'N. (UAC 66735). It occurs on the wet shrub tundra between the clumps of Betula glandulosa and Salix spp., and is unlike any of the other Carex species. This species should be added to the list of rare plants in the Yukon (Douglas et al. 1981). Eleocharis elliptica Kunth. — YUKON. Klondike River bridge, Dempster Highway at 138°44'W., 63°59'N. (UAC 70230). This North American species is widespread in Alberta (Moss 1983, page 158), and should be looked for elsewhere in the southern Yukon Territory. It is characterised by yellow, orange or brown achenes with 12-20 vertical ridges. It should be added to the list of rare plants of the Yukon Territory (Douglas et al. 1981). Range Extensions in the Yukon CYPERACEAE Carex foenea Willdenow — YUKON. Tagish Camp- ground, 134°15'W., 60°19'N. (UAC 70310), and Lucky Lake, east of Watson Lake, 134°30'W., 60°O1'N. (UAC 61656). It is probably present at scattered localities across the south- ern Yukon Territory in wet places in Lodgepole Pine forests. It has previously been found in three localities (Cody, 1994, page 149: Cody et al. 1998, page 301) including Watson Lake. Carex glareosa Wahlenburg — YUKON. Minto Landing in grassy glades at 136°53'W., 62°03'N. (UAC 70268). Douglas et al. (1981), and Cody (1994, page 440; 1996; Cody et al. 1998, page 301) have previously reported it from north of the 69" parallel, but this represents a large extension of its range southwards by some 7° of latitude. Carex lapponica O. F. Lang — YUKON. Sheldon Lake, 131°13'W., 61°38'N. (UAC 70271). Cody et al. (1988, page 301) had earlier collected it in the extreme southeast of the Territory. Carex magellanica Lamark ssp. irrigua (Wahlenburg) Hiitonen — YUKON. Watson Lake at 128°41'W., 60°03'N. (UAC 73050). It was previously collected further west and north (Cody, 1996, page 157) and further east (Cody et al. 1998, page 302). Carex marina Dewey — YUKON. Minto Landing, at 136°53'W., 62°03'N. (UAC 66965), as a rare compo- nent of the grassy open areas in the boreal forest along the river bank, close to the steep, grassy, south-facing slopes of the valley walls. This is presumably an extension of its limited range in adjacent south-central Alaska in the rain shadow of the Wrangel Mountains (Hultén, 1968, page 239). Cody (1996, page 157) only reported it from Herschel Island, the Lower Blow River delta (Cody et al. 1998, page 303), and the Mal- colm River delta. Carex pachystachya Chamisso ex Steudel — YUKON. Macmillan Pass between 1097 m and 1250 m at 130°30'W., 63°21'N. and 130°06'W., 63°23'N. (UAC 61619, 61620), Tombstone Mountain Campground, km 75, Dempster Highway, about 138°14'W., 64°30'N. (UAC 61615), and Dragon Lake, North Canol Road, 131°20'W., 62°33'N. (UAC 70287). 2004 These sites represent a large range extension from the south Yukon (Cody 1996, page 162) to the northern limit of the Boreal Forest. Harris (1998, page 269) previously listed it as being present at km 161.7, Robert Campbell Highway. It’s rarity needs to be verified. JUNCACEAE Juncus stygius Linnaeus ssp. americanus (Buch.) Hultn — YUKON. Thermokarst Mounds, South Fork, Blackstone River, Dempster Highway at 138°22'W., 64°48'N., July 1987 (UAC 70257). Cody (1996, page 190) previously reported it from the Keno Hill area, while Cody et al. (1998, page 305) found it at Coal River in the southeast Yukon. Literature Cited Cody, W. J. 1994. The flora of the Yukon Territory: Additions, range extensions and comments. Canadian Field-Natural- ist 108: 428-476. Cody, W. J. 1996. Flora of the Yukon Territory. NRC Research Press, Ottawa, Ontario, Canada. 643 pages. Cody, W. J. 2000. Flora of the Yukon Territory. 2" Edition, NRC Research 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. Cody, W. J., C. E. Kennedy, and B. Bennett. 2000. New records of vascular plants in the 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 III. Canadian Field-Naturalist 115: 301-322. NOTES 267 Cody, W. J., C. E. Kennedy, B. Bennett, and V. Loewan. 2002. New records of vascular plants in the Yukon Terri- tory [V. Canadian Field-Naturalist 116: 446-474. Cody, W. J., C. E. Kennedy, B. Bennett, and J. Staniforth. 2003. New records of vascular plants in the Yukon Terri- tory [V. Canadian Field-Naturalist 117: 278-301. Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. Brunton. 1981. The rare vascular plants of the Yukon. Syllogeus #28. National Museums of Canada, Ottawa. 61 pages. Flora of North America Editorial Commitee. 2002. Flora of North America 23. Oxford University Press, New York. 608 pages. Harris, S. A. 1998. Effects of vegetation cover on soil heat flux in the southern Yukon Territory. Erdkunde 52: 265-285. Harris, S. A. 2004. Source areas of North Cordilleran en- demic flora: Evidence from Sheep and Outpost Mountains, Kluane National Park, Yukon Territory. Erdkunde 58: 62-81. Hultén, E. 1968. Flora of Alaska and neighbouring Terri- tories. Stanford University Press, Stanford. 1008 pages. Moss, E. H. 1983. Flora of Alberta. 24 Edition, revised by John G. Packer. University of Toronto Press, Toronto. 687 pages. Wahl, H. E., D. B. Fraser, A. C. Harvey, and J. B. Maxwell. 1987. Climate of Yukon. Climatological Studies #40. Atmospherric Environment Service, Environment Canada. 323 pages. Received 21 February 2002 Accepted 21 August 2004 “Ashkui’ Vernal Ice-cover Phenomena and Their Ecological Role in Southern Labrador SHAUNA M. BAILLie!, CORINNE D. WILKERSON? and TINA L. NEWBURY? ‘Biology Department, P.O. Box 5000, St. Francis Xavier University, Antigonish, Nova Scotia B2G 2W5; e-mail:: sbaillie@ stfx.ca * Biology Department, Memorial University of Newfoundland, St. John’s, Newfoundland A1B 3X9; e-mail: r34cdw @mun.ca 3 10 Highland Avenue, Corner Brook, Newfoundland A2H 2Y5; e-mail: tinalnewbury @hotmail.com Baillie, Shauna M., Corinne D. Wilkerson, and L. Tina Newbury. 2004. “Ashkui’” vernal ice-cover phenomena and their ecological role in southern Labrador 2002. Canadian Field-Naturalist 118(2): 267-269. This is the first documented incident of River Otter (Lutra canadensis) feeding on Common Goldeneye (Bucephala clangula) in a little studied region, southern Labrador. Our observations were made during spring staging when waterfowl aggregate at open water sites in frozen lakes and rivers, locally known as ashkui. We suggest that otters and raptors opportunistically forage on staging waterfowl at ashkui. Key Words: River Otter, Lutra canadensis, Common Goldeneye, Bucephala, clangula, Bald Eagle, Haliaeetus leuco- cephalus, predator-prey interactions, staging waterfowl, ashkui, Labrador. Ashkui (singular and plural form) is the Innu name given to sites of open water in river and lake systems within the frozen spring landscape of Labrador (Flet- cher and Breeze 2000*). Migratory waterfowl, includ- ing Common Goldeneye (Bucephala clangula), use ashkui as staging areas enroute to their breeding grounds. These birds arrive in groups of tens to hun- dreds to rest and rebuild energy reserves by feeding on invertebrates, fish, seeds and other plant material at ashkui (Newbury 2002*). Beaver (Castor canadensis), River Otter (Lutra canadensis) and Muskrat (Ondatra zibethicus), have been observed at these sites. Osprey (Pandion haliaetus) and Bald Eagle (Haliaeetus leu- cocephalus) are known to fish at ashkui (Fletcher and Breeze 2000*). The temporal existence, number and distribution of ashkui sites are influenced by a high 268 degree of inter-annual variability in the onset of spring thaw. As the winter snows disintegrate and waterfowl await availability of breeding habitat, they may be sus- ceptible to predation. In 2002, the opening of ashkui occurred two weeks later than in previous years (un- published data, Environment Canada, St. John’s, New- foundland). Our observations were made during a study exam- ining the effects of military low level flying on the behaviour of staging waterfowl at Lac Fourmont in southern Labrador (52°00'N, 60°15'W; Newbury 2004*). While conducting waterfowl behavioural ob- servations, we witnessed a River Otter feeding on a male Common Goldeneye. We also observed a Bald Eagle attack another goldeneye resting on the ice. Study Area Lac Fourmont is situated within the Boreal Forest Region of eastern Canada and is influenced by con- tinental climatic regimes (Lopoukhine et al. 1977*). This region is covered with ice and snow until daily spring thaw temperatures rise to 5.1°C in May (30 year average for Goose Bay, Labrador; Environment Canada Meteorological Service, St. John’s, Newfoundland). Daytime average temperatures were 4.1°C in May 2002 and ranged from 4.0 to 9.0°C for May during the years 1993 to 2002. River width at the observa- tion location was approximately | km, and the ashkui site was 20 m in breadth. Ice breadth from ashkui to shoreline was approximately 50 m. Forest stands were composed of mainly Black Spruce (Picea mariana) and Balsam Fir (Abies balsamea) interspersed with approximately 10 % Paper Birch (Betula papyrifera), Balsam Poplar (Populus balsamifera), and Trembling Aspen (Populus tremuloides). Lake-edge shrubs con- sisted of willows (Salix spp.) and alders (Alnus rugosa and A. crispa). River and bank substrate was composed of coarse-grained sand and pebbles with exposed boul- ders and bedrock of gneiss and mafic intrusions. River basin and landscape were low gradient with shallow sloping banks, wide flood plain and low relief hills. We conducted daily observations on waterfowl be- haviour from blinds using a Swarovski 60x spotting scope from 9:00 to 12:00 and 13:00 to 16:00 between 26 April and 27 May 2002. Also, we conducted thir- teen random 1.5 h watches during dawn and dusk. Observations Fourteen species of waterfowl occupied the Lac Fourmont ashkui. Daily waterfowl numbers at the ashkui ranged from 34 to 376 during this study. A single River Otter was noted at the ashkui during at least five observation days. The number of sightings of mature and immature Bald Eagles, Osprey and Rough- legged Hawk (Buteo lagopus) totaled 7, 2, 1 and 1, respectively, over 8 non-consecutive study days. On 6 May 2002 at 11:35, we observed a River Otter grasping a male Common Goldeneye on the ice, ap- THE CANADIAN FIELD-NATURALIST Vol. 118 proximately 70 m from our observation blind on the south side of the ashkui. Although the capture was not observed, the goldeneye was alive and struggling when the otter was initially observed. The otter sat on its rear haunches with the goldeneye braced in its fore- paws and consumed the head of the duck first before eating the breast muscles and abdominal viscera. After approximately 10 minutes of feeding, the otter slipped into the water and swam downstream without the car- cass remains. We did not observe the fate of the duck carcass, as it was no longer visible on the ice when the otter left the feeding site. The bird flock, during this observation period, comprised mainly 65 Com- mon Goldeneyes, 27 Black Ducks, 113 Canada Geese (Branta canadensis). On 18 May 2002 at 16:03, we observed a Bald Eagle circle the ashkui and plunge downward toward a small group of Common Goldeneyes resting and sleeping on the ice, approximately 150 m from our position. Upon impact, the eagle initially captured a goldeneye. The eagle then struggled with the golden- eye while standing on the ice edge but was unable to maintain its grasp. The goldeneye fell from the ice to the water then became inter-mixed with the flock. This interaction lasted approximately one minute and, we were unable to identify the attacked goldeneye after its escape. The eagle then flew to the other side of the ashkui and rested on the ice. The bird flock, during this observation, comprised mainly 7 Common Gol- deneyes, 29 Black Ducks (Anas rubripes), 67 Canada Geese (Branta canadensis), 11 Greater Scaup (Aythya marila) and 20 Ring-necked Ducks (Aythya collards). Discussion Fish are often the main prey of River Otter and rap- torial predators (Chubbs and Trimper 1998; Lariviére and Walton 1998; Fletcher and Breeze 2000*; Heath et al. 2001; Jedrzejewska et al. 2001). Studies in the Great Lakes and North American boreal ecosystems have shown that the diet of River Otter comprises mostly fish (Lariviere and Walton 1998). However, otters have been known to feed opportunistically on small mammals, molluscs, reptiles, birds and fruits (Lariviére and Walton 1998). In west-central Idaho, otters supplemented their fish diet with invertebrates and reptiles (Melquist and Hornocker 1983). In Great Britain and Ireland, otters consume primarily fish, and lesser amounts of small mammals, medium-sized mam- mals, birds, herpetofauna, earthworms, other inverte- brates and carrion of large mammals (McDonald 2002). Jedrzejewska et al. (2001) showed that otters special- ize on prey taken from water. It is plausible that an otter fishing for fish under water may opportunisti- cally capture diving goldeneyes. Although River Otters were known to occasionally feed on birds (Lariviére and Walton 1998), we believe that this was the first documented case of an otter feeding on a Common Goldeneye. 2004 Instances of Bald Eagles preying on duck species are not rare (Jackman et al. 1999; Heath et al. 2001). _ Although fish dominate their diet, Bald Eagles have been reported to modify foraging behaviour and may forage opportunistically on staging waterfowl in situa- tions where return on such effort makes them econom- ical (Brown et al. 1998; Jackman et al. 1999). The temporal existence, number and distribution of ashkui, in southern Labrador during spring, may influ- ence the spatial and temporal distribution of piscivo- rous predators and migratory waterfowl by limiting the availability of open-water feeding and resting areas. Migratory waterfowl arriving from southern destina- tions must accumulate in greater concentrations when ashkui openings are limited. Though the behaviour and distribution of River Otter were not documented for Labrador, Lariviére and Walton (1998) stated that otters made heavy use of ashkui in winter and “almost exclusively” used open water for locomotion and _ foraging (Madsen and Prang 2001; Ruiz-Olmo et al. 2001). Piscivorous raptors returning to breeding areas must travel to and congregate at available ashkui to _ fish. It was likely that spatial and temporal distribution of ashkui also affected predator-prey interactions. Our observations are examples of opportunistic for- aging events on waterfowl by predators that appear to use ashkui regardless of the presence of waterfowl. Further investigations in Labrador are needed to under- stand better the ecological importance of the tem- poral and spatial distribution of ashkui to mammalian and raptorial predators. Acknowledgments This project was funded in partnership by the In- stitute for Environmental Monitoring and Research (IEMR), the Canadian Wildlife Service (CWS), McGill University and the Department of National Defence (DND), and by the Northern Scientific Training Pro- gram. Many thanks to Sean Sharpe (IEMR), Tony Chubbs (DND) and Roger Titman (McGill University) for invaluable discussion, advice and/or informal manu- script review. Jean Sealy (CWS), Scott Gilliland (CWS) and Dawn Laing (DND) helped us with literature searches. Thanks to Keith Chaulk (CWS), Bruce Whif- fen (Environment Canada) and Sarah Hall (CWS) for providing information on weather and ice conditions. Documents Cited [marked * in text] Fletcher, C., and H. Breeze. 2000. Ashkui sites in the low- level flight training area, Labrador. Report prepared under NOTES 269 contract to Institute for Environmental Monitoring and Research (1.E.M.R.). Gorsebrook Research Institute for Atlantic Canada Studies, Saint Mary’s University, Halifax, Nova Scotia. 22 pages. Lopoukhine, N., N. A. Prout, and H. E. Hirvonen. 1977. The ecological land classification of Labrador. Ecologi- cal Land Classification Series, Number 4. Fisheries and Environment Canada. Halifax, Nova Scotia. Newbury, Tina L. 2002. Effect of low-level flights on the behaviour of spring staging waterfowl in Central Labrador. M.Sc. thesis Proposal. McGill University. Ste. Anne-de- Bellevue, Quebec, Canada. Newbury, Tina L. 2004. Effect of low-level military jet over- flights on the behaviour of spring-staging waterfowl at Lac Fourmont ashkui, Labrador, Canada. M.Sc. thesis. McGill University. Ste. Anne-de-Bellevue, Quebec, Canada. Literature Cited Brown, B. T., L. E. Stevens, and T. A. Yates. 1998. Influ- ences of fluctuating riverflows on Bald Eagle foraging behaviour. Condor 100: 745-748. Chubbs, Tony E., and Perry G. Trimper. 1998 The diet of nesting Ospreys, Pandion haliaetus, in Labrador. Canadian Field-Naturalist 112: 502-505. Heath, Joel P., Geoff Goodyear, and Joe Brazil. 2001. Observation of a Golden Eagle, Aquila chrysaetos, attack on a Harlequin Duck, Histrionicus histrionicus, in northern Labrador. Canadian Field-Naturalist 115: 515-516. Jackman, R. E., W. G. Hunt, J. M. Jenkins and P. J. Detrich. 1999. Prey of nesting bald eagles in northern California. Journal of Raptor Research 332: 87-96. Jedrzejewska B., V. E. Sidorovich, M. M. Pikulik, and W. Jedrejewski. 2001. Feeding habits of the otter’and the American mink in Bialowieza Primeval Forest (Poland) compared to other Eurasian populations. Ecography 242: 165-180. Lariviere. Serge, and Lyle R. Walton. 1998. Lutra cana- densis. Mammalian Species (587) American Society of Mammalogists. Pages 1-8. Madsen, A. B., and A. Prang. 2001 Habitat factors and the presence or absence of Otters Lutra lutra in Denmark. Acta Theriologica 46: 171-179. McDonald, R. A. 2002. Resource partitioning among British and Irish mustelids. Journal of Animal Ecology 71: 185- 200. Melquist, W. E. and M. G. Hornocker. 1983. Ecology of river otters in west central Idaho. Wildlife Monographs 83: 1-60. Ruiz-Almo. J., J. M. Lopez-Martin, and S. Palazon. 2001. The influence of fish abundance on the otter (Lutra lutra) population in Iberian Mediterranean habitats. Journal of Zoology 254: 325-336. Received 28 June 2002 Accepted 19 October 2004 270 THE CANADIAN FIELD-NATURALIST Vol. 118 Sea Otter, Enhydra lutris, Sightings off Haida Gwaii / Queen Charlotte Islands, British Columbia, 1972-2002 KIMBERLY RAUM-SURYAN!3, KENNETH PITCHER!, and RICHARD LAMY? ' Alaska Department of Fish and Game, Division of Wildlife Conservation, 525 W. 67 Avenue, Anchorage, Alaska 99518 USA; e-mail: kraumsuryan @charter.net ? Gwaii Haanas National Park Reserve/Haida Heritage Site, P.O. Box 37, 120 Second Avenue, Queen Charlotte, British Columbia VOT 1SO Canada >Current address: 928 NW Cottage Street, Newport, Oregon 97365 USA Raum-Suryan, Kimberly, Kenneth Pitcher, and Richard Lamy. 2004. Sea Otter, Enhydra lutris, sightings off Haida Gwaii/Queen Charlotte Islands, British Columbia, 1972-2002. Canadian Field-Naturalist 118(2): 270-272. On 27 June 2001 we observed and photographed a Sea Otter (Enhydra lutris) adjacent to a Steller Sea Lion (Eumetopias jubatus) haulout near Sgang Gwaay (Anthony Island), Haida Gwaii / Queen Charlotte Islands. This is one of only eight documented sightings of Sea Otters in these waters during the past 30 years. These sightings may represent the beginning of the expansion of Sea Otters to their former range off Haida Gwaii. Key Words: Sea Otter, Enhydra lutris, Haida Gwaii, Queen Charlotte Islands, British Columbia. The Sea Otter (Enhydra lutris) once ranged in near- shore waters along the North Pacific Rim from Japan to Baja California. The worldwide population of Sea Otters in the early 1700s was estimated to be between 150 000 (Kenyon 1969) and 300 000 animals (John- son 1982). Middens (human food waste mounds) in- dicate that aboriginal people, including the Haida, hunted Sea Otters (Simenstad et al. 1978; Acheson 1998; Sloan 2004). Although this harvest may have caused local reductions of Sea Otter populations near village sites (Simenstad et al. 1978), the species was abundant throughout its range prior to commercial ex- ploitation. Following the advent of commercial hunting in 1741, the worldwide population of Sea Otters de- clined to less than 2000 animals by 1911, limited to remnant groups throughout the range (Kenyon 1969). Many of these remnant groups (including one group located off the Haida Gwaii / Queen Charlotte Islands, British Columbia, Canada) were extirpated, likely due to their small size (Watson et al. 1997). The last Sea Otter recorded from Haida Gwaii was before 1920 (Kenyon 1969) and the last recorded in British Colum- bia, near Vancouver Island (prior to reintroduction), was in 1929 (Cowan and Guiguet 1960; Kenyon 1969). In 1911 an International Treaty provided protection of Sea Otters from additional exploitation. Furthermore, to aid in the recovery of Sea Otters, various translo- cation projects were conducted from the 1950s to early 1970s in the Pacific coastal United States and Canada (Kenyon and Spencer 1960; Kenyon 1969; Burris and McKnight 1973; Bigg and MacAskie 1978; Jameson et al. 1982; Jameson et al. 1986; Riedman and Estes 1990). Since reintroduction, the Sea Otter population in British Columbia has increased 18.6% per year from 70 animals in 1977 to an estimated 2 500 animals in 1998 (Watson 2000). Although Sea Otters continue to expand their range along Vancouver Island and the central coast of British Columbia (Watson 2000; Figure 1), sightings off Haida Gwaii / Queen Charlotte Islands remain rare. On 27 June 2001 we observed and photographed a Sea Otter of unknown sex adjacent to a Steller Sea Lion haulout (52°04.966’ N, 131°13.811° W) just south of Sgang Gwaay (Anthony Island), Haida Gwaii / Queen Charlotte Islands (Figure 1). The Sea Otter was swim- ming on its back eating a sea urchin (Strongylocentrotus sp.). We observed the otter for approximately 30 min- utes as it swam and dove. This observation is one of only eight documented sightings (Edie 1973; Taylor and Gough 1977; Heise et al. 2003) of Sea Otters in waters surrounding Haida Gwaii / Queen Charlotte Islands during the past 30 years (Figure 1). This sighting raises some interesting questions regarding the origin of this individual and whether or not Sea Otters are expanding their range to include the Haida Gwaii / Queen Charlotte Islands once again. Kenyon (1969) suggested that the most significant factor limiting the spread of Sea Otter populations is a tendency of individual Sea Otters to occupy a limited home range. Although Sea Otters are not known to migrate, as populations increase, males generally are the first to explore new areas and their presence some- times indicates imminent range reoccupation (Loughlin 1980; Garshelis et al. 1984; Pitcher 1989). Garshelis and Garshelis (1984) recorded long-distance move- ments of > 100 km for five adult territorial males. They postulated that the Sea Otters might have left an area of limited food resources to travel to an area where food was more abundant. When Sea Otters were extirpated, many of their primary benthic invertebrate prey became larger and more abundant (Estes and Palmisano 1974; Garshelis et al. 1986). When Sea Otters recolonize they initially encounter more abundant and larger prey resources than may be expected to persist, leading to densities of Sea Otters in recovering popu- lations greater than those that can be sustainable over Southeast Alaska NOTES 27) 51 introduced in 1965 International Boundary Haida Gwaii / Queen Charlotte Islands July 1999 August 1990 August 1976 June 2001 July 1972 @ Sea Otter sightings A Sea Otter translocation sites C»> Current Sea Otter range @ August 2002 55 introduced in 1965 Ga British Columbia Wd D : 2 IN s a Q VY Goose Islands Group 89 introduced between 1969 and 197 200 Kilometers 134° 132° 130° 128° 126° FicureE 1. Sightings of Sea Otters around Haida Gwaii / Queen Charlotte Islands, locations of Sea Otter translocations nearest Haida Gwaii, and current Sea Otter distribution in Southeast Alaska and British Columbia (map adapted from Parks Canada Report 038, Living marine legacy of Gwaii Haanas. IV: Marine mammal baseline to 2003 and marine mammal-related management issues throughout the Haida Gwaii region by Heise et al. 2003). time (Bodkin et al. 2000). As prey is consumed, Sea Otter densities should decline and fluctuate at an equil- ibrium density (Estes 1990) through decreased repro- duction, increased mortality, or through emigration (Bodkin et al. 2000). Sea Otter population increases in the Aleutian Islands were achieved largely by range expansion over deep, wide ocean passes (Estes 1990). Unless the Sea Otter we observed is a member of a previously undetected group off Haida Gwaii / Queen Charlotte Islands, this individual would have had to travel at least 175 km across an open expanse of water from the nearest known group of otters along the British Columbia coast (Figure 1). If this individual traveled from southeast 272 Alaska or northern Vancouver Island, the distance would be even greater. If Sea Otter populations have reached or exceeded carrying capacity in certain regions of British Columbia or southeast Alaska (Watson et al. 1997; Watson 2000), this may be the beginning of the expansion of Sea Otters to their former range off Haida Gwaii / Queen Charlotte Islands. This range expansion may be particularly important given the precipitous population decline of Western Alaska Sea Otters dur- ing the past 15 years (Doroff et al. 2003) and the pro- posal to list the population as threatened under the United States Endangered Species Act (Federal Regis- ter: 11 February 2004; Volume 69, Number 28). Acknowledgments We thank the Gwaii Haanas National Park Reserve and Haida Heritage Site for allowing us to conduct Steller Sea Lion surveys (Research Permit 01-01) off Haida Gwaii, thereby allowing us to observe the Sea Otter mentioned. We are grateful to Jane Watson, Norm Sloan, Tom Gelatt, and two anonymous reviewers for their comments regarding this note. Literature Cited Acheson, S. 1998. In the wake of the ya’aats’ xaatgaay [Iron People’ ]: a study of changing settlement strategies among the Kunghit Haida. British Archaelogical Reports Inter- national Series 711. 209 pages. Bigg, M. B., and I. B. MacAskie. 1978. Sea otters reestab- lished in British Columbia. Journal of Mammalogy 59: 874-876. Bodkin, J. L., A. M. Burdin, and D. A. Ryazanov. 2000. Age- and sex-specific mortality and population structure in sea otters. Marine Mammal Science 16: 201-219. Burris, O. E., and D. E. McKnight. 1973. Game transplants in Alaska. Alaska Department of Fish and Game, Game Technical Bulletin (4). Cowan, I. M., and C. J. Guiguet. 1960. The mammals of British Columbia. British Columbia Provincial Museum Handbook (11). Doroff, A. M., J. E. Estes, M. T. Tinker, D. M. Burn, and T. J. Evans. 2003. Sea otter population declines in the Aleutian archipelago. Journal of Mammalogy 84: 55-64. Edie, A. G. 1973. Sea otter sighting at Cape St. James, Bri- tish Columbia. Syesis 6: 265. Estes, J. A. 1990. Growth and equilibrium in sea otter popu- lations. Journal of Animal Ecology 59: 385-401. Estes, J. A., and J. F. Palmisano. 1974. Sea otters: Their role in structuring nearshore communities. Science 185: 1058- 1060. Garshelis, D. L., and J. A. Garshelis. 1984. Movements and management of sea otters in Alaska. Journal of Wildlife Management 48: 665-678. Garshelis, D. L., A. M. Johnson, and J. A. Garshelis. 1984. Social organization of sea otters in Prince William Sound, Alaska. Canadian Journal of Zoology 62: 2648-2658. THE CANADIAN FIELD-NATURALIST Vol. 118 Garshelis, D. L., J. A. Garshelis, and A. T. Kimker. 1986. Sea otter time budgets and prey relationships in Alaska. Journal of Wildlife Management 50: 637-647. Heise, K. A., N. A. Sloan, P. F. Olesiuk, P. M. Bartier, and J. K. B. Ford. 2003. Living marine legacy of Gwaii Haanas IV: Marine mammal baseline to 2003 and marine mammal- related management issues throughout the Haida Gwaii region. Parks Canada Technical Reports in Ecosystem Science Report 038. 150 pages. Jameson, R. J., K. W. Kenyon, A. M. Johnson, and H. M. Wight. 1982. History and status of translocated sea otter populations in North America. Wildlife Society Bulletin 10: 100-107. Jameson, R. J., K. W. Kenyon, S. Jeffries, and G. R. VanBlaricom. 1986. Status of a translocated sea otter population and its habitat in Washington. The Murrelet 67: 84-87. Johnson, A. M. 1982. Status of Alaska sea otter populations and developing conflicts with fisheries. Pages 293-299 in Proceedings of the Forty-seventh North America Wildlife Conference, Alaska. Wildlife Management Institute, Wash- ington, D.C. Kenyon, K. W., and D. L. Spencer. 1960. Sea otter popu- lation and transplant studies in Alaska, 1959. U.S. Fish and Wildlife Service Special Scientific Report — Wildlife 49. 29 pages. Kenyon, K. W. 1969. The sea otter in the eastern Pacific Ocean. North American Fauna 68: 1-352. Loughlin, T. R. 1980. Home range and territoriality of sea otters near Monterey, California. Journal of Wildlife Man- agement 44: 576-582. Pitcher, K. W. 1989. Studies of the southern Alaska sea otter populations: Distribution, abundance, structure, range ex- pansion, and potential conflicts with shellfisheries. United States Fish and Wildlife Service Cooperative Agreement Number 14-16-009-954. Riedman, M. L., and J. A. Estes. 1990. The sea otter (En- hydra lutris): behavior, ecology and natural history. United States Fish and Wildlife Service Biological Report 90. 126 pages. Simenstad, C. A., J. A. Estes, and K. W. Kenyon. 1978. Aleuts, sea otters, and alternate stable-state communities. Science 200: 403-411. Sloan, N. A. 2004. Northern abalone: Using an invertebrate to focus marine conservation ideas and values. Coastal Management 32: 129-143. Taylor, R., and B. Gough. 1977. New sighting of sea otter reported for Queen Charlotte Islands. Syesis 10: 177. Watson, J. C. 2000. The effects of sea otters (Enhydra lutris) on abalone (Haliotis spp.) populations. Pages 123-132 in Workshop on Rebuilding Abalone Stocks in British Col- umbia. Edited by A. Campbell. Canadian Special Publica- tion Fisheries and Aquatic Sciences 130. Watson, J. C., G. M. Ellis, T. G. Smith, and J. K. B. Ford. 1997. Updated status of the sea otter, Enhydra lutris, in Canada. Canadian Field-Naturalist 111: 2: 277-286. Received 6 August 2002 Accepted 26 August 2004 ISABELLE SCHMELZER! and FRANK PHILLIPS” NOTES 273 _ First Record of a Barred Owl, Strix varia, in Labrador 'Department of Environment and Conservation, Wildlife Division, Government of Newfoundland and Labrador. P.O. Box 2007, Corner Brook, Newfoundland A2H 7S1 Canada; e-mail: [sabelleSchmelzer@ gov.nl.ca "Department of Natural Resources, Forestry Division, Government of Newfoundland and Labrador, Goose Bay, Labrador AOP 1CO Canada Schmelzer, Isabelle, and Frank Phillips. 2004. First record of a Barred Owl, Strix varia, for Labrador. Canadian Field-Naturalist 118(2): 273-276. A Barred Owl (Strix varia) was heard calling in central Labrador, Canada in 2001, and a dead owl was found in the same area in 2004. These are the first confirmed records for the Province of Newfoundland and Labrador. Key Words: Strix varia, Barred Owl, range, vagrant, Newfoundland and Labrador. Barred Owls (Strix varia) are common in southern Canadian forests from British Columbia through Nova Scotia (AOU 1998), occurring in mixed evergreen and mature deciduous woodlands, but also into the boreal forest in the northern portion of its range (Erskine 1977; Godfrey 1986; Mazur et al. 1998). There are no confirmed records of Barred Owls for the Province of Newfoundland and Labrador (Todd 1963; ACCDC 2004*). We document two occurrences of a Barred Owl near Happy Valley — Goose Bay, Labrador, three years apart. On the evening of 24 August 2001, at 22:00 hrs, a lone Barred Owl was heard calling in the forest north of Happy- Valley Goose Bay, Labrador, along the banks of the Goose River (53° 24'N, 60° 26'W) by one of the authors (IS) (Figure 1). The owl called continu- ously for several minutes from the same location, approximately 250 m away. The song was made up of the characteristic pattern of 8 hoots, separated into two parts by a pause after 4 hoots, and ending with a characteristic descending pitch on the last hoot (‘hoo- hoo hoohoo, hoohoo hoohooahhw’ ). The calls of this owl may be unfamiliar to residents of Labrador, but Barred Owl calls are very distinctive, and were imme- diately recognizable to one of the authors (IS). The only other owl species in the area with which the calls could possibly be confused is the Great Horned Owl (Bubo virginianus), whose call is of a deeper tone, and consists of five or six hoots, beginning with one em- phatic hoot, followed by several shorter and quieter calls, and ending with two long hoots on the same pitch (‘Hooo! hoohoohoo Hoooo, Hoooo’). The calls of the two owls differ in tone, pattern and rhythm, and are not difficult to tell apart by experienced listeners. Using a vocal imitation of the call, the author at- tempted to elicit a callback. The owl responded by leaving its perch, approaching her, and selecting a new perch within 50 m of her location. It made no other movements, and she was able to elicit vocal respons- es to her calls for nearly an hour. During this period, the owl occasionally varied its call either by repeat- ing only the last section ‘hoohooahhw’, or by short- ening the first section to a rising series of 3 notes followed immediately by the distinctive descending cackle (‘hoohoohoo hoohooahhw’ ). Unfortunately, due to darkness and heavy forest cover, the owl was not sighted. At the time of observation, the air temperature was 10°C, winds were calm, and the skies were clear. Tourists camped at the site reported hearing the same calls during the previous evening. On the evenings of 26, 28, and 31 August, at approx- imately 22:00, IS returned to the area to perform a nocturnal callback survey (using a vocal imitation of the call) to check for the presence of the owl. Roads and paths in forested areas within 10 km of the origi- nal sighting were surveyed, with stops every 0.3 — 0.5 km, and several minutes spent at each’ station calling and listening for a response. No Barred owls were detected during these subsequent surveys. The second observation occurred three years later, in virtually the same location (53° 27'N 60° 18'W, on 16 July 2004 (Figure 1). A dead owl was seen on the side of Route 530, at approximately 08:00 hrs. On closer inspection, the owl carcass showed no obvious sign of injury or illness, and was completely intact, with no sign of scavenging, and no fly larvae. The eyes were present and moist (not sunken, retracted, or absent as in older kills). Given the abundance of foxes (Vulpes vulpes), Ravens (Corvus corax), Gray Jays (Perisoreus canadensis) and other scavengers, it is the opinion of the authors that the owl had been dead for a few hours at most. Field marks on the owl included dark eyes, vertical barring on the belly and short, horizontal bars across the chest, all characteristic of Barred Owls (Figure 2). The owl was also assessed using the criteria of Pyle (1997). The lack of down on the head and contour feathers, flight feathers that were broad and uniform in color and wear, and the presence of squared bars on the primary coverts all suggest the bird was an adult (After Hatch Year or AHY). On dissection, the owl was determined to be a female by the presence of an ovary. No brood patch was present. Barred owls have been observed in Québec north of the St. Lawrence River (Gagnon and Bombardier 1996). The breeding range has been reported to extend 274 Quebec Grebe 1992 0) 150 300 450 Kilometers Ve ay THE CANADIAN FIELD-NATURALIST Vol. 118 FiGure 1. A summary of Barred Owl records in Labrador and nearby Quebec. Further description of the records is given in the text. to 50° North in eastern Canada (David 1980; Godfrey 1986). Our review of the literature, however, revealed several sightings north of this latitude in Québec. For example, a Barred Owl was reported near Lac aux Cédres, 52° 00'N, 67° 07'W, near the Labrador border in 1917 (Todd 1963). It has been reported more recently further west near Nemiscau, 51° 42'N, 76° 15'W, and Lake Evans, 50° 55'N, 77° 00'W, (Grebe 1992). Addi- tionally, Comeau referred to the bird as “tolerably common” near Godbout, 49° 19'N, 67° 18'W, on the North Shore of the Gulf of St. Lawrence (in Merriam 1882), and a specimen was captured at Lac Paterson in Roberval County (50° 11'N, longitude not specified) in 1954 (Godfrey 1957). Nonetheless, the occurrence of the owl in central Labrador is 500 km distant from the nearest previous report, and 700 km outside regions where the species is noted regularly. During the past several decades, the geographic range of the Barred Owl has expanded in western North America, and most recent discussion has focused on its movements into the Pacific Northwest (Levy 1999; Boxall and Stepney 1982; Wright and Hay- ward 1998). However, on the basis of sightings in Quebec north of the St. Lawrence River, Harper (1958) proposed that the range of the Barred Owl was also extending northward in eastern North America. Al- though the Gulf of St. Lawrence and the Strait of Belle Isle present significant barriers to occurrence on the island of Newfoundland, the forests of Quebec and Labrador are continuous and present no obsta- cles to movement. The environs of Happy Valley — Goose Bay, including the Churchill River valley and coastal plain surrounding Lake Melville, belong to the “High Boreal Forest’ ecoregions, (Government of Newfoundland and Labrador 2004*) and consist of mature, dense mixed deciduous/coniferous forest on sandy soil. Predominant tree species are Trembling Aspen (Populus tremuloides) and White Birch (Betula papyrifera), with coniferous species such as Balsam Fir (Abies balsamea), and White and Black spruce (Picea glauca, P. mariana) occurring secondarily. Suitable breeding habitat for Barred Owls is often associated with the presence of snags used as nest cavities (Haney 1997), but also nests of other raptors or corvids (Elderkin 1987). This region features the most favorable climate in all of Labrador, including a growing season of 120-140 days. Conversely, the up- land plateaus of central and western Labrador are 2004 FiGureE 2: Adult female Barred Owl collected near Happy Valley — Goose Bay in July 2004, which showed char- acteristic gray-brown coloration, yellow bill, vertical barring on the belly and horizontal barring on the chest. Flight feathers (right wing shown) are broad and uni- form in colour and wear, suggesting an adult bird that has not yet molted. characterized by a subarctic forest and climate, and border the former ecoregion to the north and west. The elevation, vegetative communities, and severe cli- mate may pose a barrier to an eastward expansion from northern Québec. To the south and east however, broad river valleys and mixedwood boreal forests predomi- nate. In the absence of evidence of breeding, it is un- known whether this record represents a possible range expansion by the Barred Owl, or simply documents the occurrence of a vagrant bird in potentially suit- able habitat on two separate occasions. The fact that in One instance the owl was an adult female, and that she was found within the breeding season, does not NOTES 275 preclude range expansion as a possible explanation. The absence of geographical barriers to movement from northeastern Quebec, particularly from the north shore of the St. Lawrence, into south-central Labrador supports this notion. The establishment of owl surveys in Labrador should help to clarify such questions, and others, in the future. Acknowledgements We extend thanks to John Thomas and Joe Brazil for sharing their knowledge of bird records for Labrador, and for encouraging us to contribute and document our own sightings. Bruce Rodrigues used his banding expertise to sex and age the owl specimen. Chris Earley, Paul Linegar, and Bruce Mactavish provided constructive and useful comments on earlier drafts of the manuscript. Documents Cited (marked * in text) Atlantic Canada Conservation Data Center. 2004. Species Rarity Ranks: Labrador Birds, August 2004. P.O. Box 6416, Sackville, New Brunswick. E4L 1G6 http:// www.accde. com. Government of Newfoundland and Labrador. 2004. Department of Natural Resources-Forestry. Ecoregions of Labrador. http://www.gov.nf.ca/forestry/maps/eco_lab.stm. GREBE. 1992. Compexe Grande-Baleine. Avant-projet, phase II. Compte réndu de I’ étude télémétrique de I’ Aigle royal réalisée dans le complexe Grande Baleine en juin 1992; mission G4. Report for Hydro Québec vice prési- dence environnement Montréal. 63 pages. Literature Cited American Ornithologists’ Union (AOU). 1998. Check-list of North American Birds, 7 edition. American Ornitho- logists’ Union. Allen Press, Inc. Lawrence Kansas. 829 pages. Boxall, P. C., and P. H. R. Stepney. 1982. The distribution and status of the Barred owl (Strix varia) in Alberta (Can- ada). Canadian Field-Naturalist 96: 46-50. David, N. 1980. Etat et distribution des oiseaux du Québec méridional. Cahiers d’ ornithologie Victor-Gaboriault, (3). Club des ornithologues du Québec, Charlesbourg. 213 pages. Elderkin, M. F. 1987. The breeding and feeding ecology of a Barred owl, Strix varia Barton, population in King’s County, Nova Scotia. M.Sc. thesis, Acadia University, Wolfville, Nova Scotia. 203 pages. Erskine, A. J. 1977. Birds in boreal Canada: communities, densities and adaptations. Canadian Wildlife Service. Report Series number 41. 73 pages. Gagnon, C., and M. Bombardier. 1996. Barred Owl 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 des ornithologues, Province of Québec Society for the Protection of Birds, Canadian Wildlife Service, Environment Canada, Québec Region Montréal. 1302 pages. Godfrey, W. E. 1957. Some distributional notes on Canadian birds. Canadian Field-Naturalist 70: 136-138. Godfrey, W. E. 1986. The birds of Canada, revised edition. National Museum of Natural Sciences, Ottawa. 595 pages. 276 Haney, J. C. 1997. Spatial incidence of Barred owl (Strix varia) reproduction in old-growth forest of the Appala- chian Plateau. Journal of Raptor Research 31: 241-252. Harper, F. 1958. Birds of the Ungava Peninsula. Museum of Natural History, University of Kansas. Miscellaneous Publications number 17. 171 pages. Levy, S. 1999. Owl vs. owl. Natural History 108: 28-32. Mazur, K. M., S. D. Frith, and P. C. James. 1998. Barred owl home range and habitat selection in the boreal forest of central Saskatchewan. Auk 115: 746-754. Merriam, C. Hart. 1882. List of birds ascertained to occur within ten miles from Point de Monts, Province of Qué- bec, Canada; based chiefly upon the notes of Napoleon THE CANADIAN FIELD-NATURALIST Vol. 118 A. Comeau. Bulletin of the Nuttall Ornithological Club 8: 244-245. Pyle, P. 1997. Identification guide to North American birds — part 1. Slate Creek Press, Bolinas, California, USA. Todd, W. E. C. 1963. Birds of the Labrador Peninsula and Adjacent Areas. A distributional list. University of Toronto Press, Toronto. 819 pages. Wright, A. L., and G. D. Hayward. 1998. Barred owl range expansion into the central Idaho wilderness. Journal of Raptor Research 32: 77-81. Received 23 January 2002 Accepted 30 August 2004 Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: A New Addition to the Flora of Quebec JACQUES BRISSON Institut de recherche en biologie végétale, Université de Montréal, 4101, Sherbrooke Street East, Montréal, Québec H1X 2B2 Canada Brisson, Jacques. 2004. Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: a new addition to the flora of Quebec. Canadian Field-Naturalist 118(2): 276-277. A population of Forked Three-awned Grass (Aristida basiramea Engelm. ex Vasey; Poaceae) was found for the first time in Quebec, on a sand barren of the Cazaville region (Haut-Saint-Laurent). The only other region where this species is known in Canada is on the southern side of Georgian Bay in Ontario. Key Words: Aristida basiramea, Forked Three-awned Grass, Poaceae, rare plants, Quebec Une population d’Aristida basiramea Engelm. ex Vasey (Poaceae) fut trouvée pour la premiére fois au Québec, dans une lande sableuse de la région de Cazaville (Haut-Saint-Laurent). Le seul autre endroit ot: l’espéce est présente au Canada est sur le coté sud de la Baie Georgienne en Ontario. Mots-clés : Aristida basiramea, Poaceae, plantes rares, Québec In September 2001, during an ecological survey near Cazaville, in the Haut-Saint-Laurent region of south- ern Quebec, a population of Forked Three-awned Grass (Aristida basiramea Engelm. ex Vasey) was found in a dry, sandy grass-field. Other populations were later located in similar habitats nearby. This is the first time the species has been reported for Quebec. In Canada, A. basiramea is known from four extant naturally-occurring sites located in southern Ontario, three of which are in Simcoe County, and one in Muskoka County (Allen 2001). The extent of occurrence in Ontario only totals 16 hectares. There is also one adventive station in northwestern Ontario, at Rainy River (Allen 2001). A few other reports exist, such as one in Norfolk County in Ontario, and a few more in Manitoba. However, the locality of the Norfolk specimen may be the result of a labelling error (Reznicek 1984), while the reports from Manitoba are considered questionable due to the absence of existing specimens (Scoggan 1957; Allen 2001). Thus, the Cazaville area becomes the second area in Canada where the presence of the species is confirmed. The species is listed as rare in Ontario (Reznicek 1984) and is considered threatened in Can- ada (COSEWIC 2003). On the basis of the newly discovered population, A. basiramea should be added to the list of rare vascular plants likely to be designated threatened or vulnerable (Labrecque and Lavoie 2002). The genus Aristida is represented by 250 to 300 spe- cies, 29 of which are native to North America north of Mexico (Flora of North America Editorial Com- mittee 2003). A. basiramea is an annual plant from 30 to 60 cm high characterized by glumes of unequal length and 1-flowered spikelets terminated by three long awns, one in the middle with a twisted base and two shorter straight awns on each side. It is abundant on dry sterile or sandy soil in the midwestern states. At the northeastern periphery of its range in the United States, it forms disjunct populations on dry lands and along sandy roadsides, some populations of which ap- pear to be adventive. The closest reports of A. basira- mea south of the border with Québec were from Platts- burg, Clinton Co., New York (S. J. Smith, 25 July 1965, NYS); Columbia, northern Coos Co., New Hampshire (A. S. Pease, 17 September 1955, NEBC); and Avon, Franklin Co., Maine (A. Haines, 26 October 1990, MAINE). The species is recognized as rare in Maine, Iowa and Colorado (Allen 2001). The Cazaville area (45°03’ N, 74°22’ W) is located in the Mixed Plain Ecozone of the St-Lawrence Low- land Ecoregion (Ecological Stratification Working Group 1995). The area is characterized by a vast sandy plain of littoral origin dating from the last post-gla- cial period. White Pine (Pinus strobus) forests were abundant before European settlement (Brisson and Bouchard 2003), but today, the area is occupied by sandy, grassy fields, sand barrens and secondary forests of Red Maple (Acer rubrum), Trembling Aspen (Popu- lus tremuloides) and Gray Birch (Betula populifolia). Aristida basiramea was found on open, sandy grass- field with Poa compressa, Danthonia spicata, and vari- ous types of lichens. The sandy plain of Cazaville is host to other rare plant species. The northernmost colony of Monarda punctata, a species rare for Quebec and Canada, was recently found nearby (Boudreault and Brisson 1994). As well, Hedeoma hispida, consid- ered rare for Quebec (Labrecque and Lavoie 2002), is also found on the sandy plain. Specimens of A. basiramea were deposited at the Marie-Victorin Herbarium (MT: Brisson Number JBO1- 25) Acknowledgments I thank Stuart Hay (Marie-Victorin Herbarium) and Stephen Darbyshire (Agriculture and Agri-Food Can- ada), who reviewed the manuscript and provided useful information. I am also grateful to Richard Mitchell (Biological Survey, New York State Museum) and Ray Angelo (Harvard University Herbaria, New England Botanical Club) for kindly providing information on A. basiramea in the U.S. NOTES Documents Cited (marked * in text) Allen, G. M. 2001. COSEWIC Status Report on Forked Three-awned Grass. Committee on the Status of Endan- gered Wildlife in Canada. Environment Canada, Ottawa, Ontario. 26 pages. Literature Cited Boudreault, C., and J. Brisson. 1994. Une addition a la flore du Québec: Monarda punctata var. villicaulis (Lamiaceae). Canadian Field-Naturalist 108: 499-500. Brisson, J., and A. Bouchard. 2003. Human activities caused major changes in tree species composition in southern Quebec, Canada. Ecoscience 10: 236-246. COSEWIC. 2003. Canadian Species at Risk, November 2003. Committee on the Status of Endangered Wildlife in Canada. 44 pages. Ecological Stratification Working Group. 1995. A National Ecological Framework for Canada. Agriculture and Agri- Food Canada, Research Branch, and Centre for Land and Biological Resources Research and Environment Canada, State of the Environment Directorate, Ecozone Analysis Branch, Ottawa/Hull. Report and National map at 1: 7 500 000 scale. Flora of North America Editorial Committee. 2003. Flora of North America North of Mexico. Volume 25: Poaceae. (Part 2). New York and Oxford. 814 pages. Labrecque, J., and G. Lavoie. 2002. Les plantes vasculaires menacées ou vulnérables du Québec. Gouvernement du Québec, Ministére de l’environnement. Direction du patri- moine écologique et du développement durable, Québec, 200 pages. Reznicek, A. A. 1984. Poaceae — Aristida basiramea . One page in Atlas of the rare vascular plants of Ontario. Edited by G. W. Argus and D. J. White. 1982-1987. National Museum of Natural Sciences. Looseleaf. Scoggan, H. J. 1957. Flora of Manitoba. National Museum of Canada, Bulletin 140. 619 pages. Received 30 November 2001 Accepted 21 June 2004 278 THE CANADIAN FIELD-NATURALIST Vol. 118 Evidence for the Use of Vocalization to Coordinate the Killing of a White-Tailed Deer, Odocoileus virginianus, by Coyotes, Canis latrans ERICH M. Muntz! and BRENT R. PATTERSON? Department of Natural Resources, Wildlife Division, 136 Exhibition Street, Kentville, Nova Scotia B4N 4E5 Canada ‘Current address: Parks Canada Agency, Cape Breton Highlands National Park of Canada, Ingonish Beach, Nova Scotia BOC 1L0 Canada *Current address: Ontario Ministry of Natural Resources, Wildlife Research and Development Section, 300 Water Street, 3" Floor North, Peterborough, Ontario K9J 8M5 Canada; e-mail: brent.patterson@mnr.gov.on.ca. Author to whom correspondence should be addressed. Muntz, Erich M., and Brent R. Patterson. 2004. Evidence for the use of vocalization to coordinate the killing of a White-Tailed deer, Odocoileus virginianus, by Coyotes, Canis latrans. Canadian Field-Naturalist 118(2): 278-280. Among the social canids, howling is largely accepted as playing a role in territory maintenance. However, its role in commu- nication within packs, such as announcing departures from den and rendezvous sites and coordinating reunions or movements, remains largely speculative. We report an observation where a radio-collared adult male Coyote (Canis latrans) and his mate seemed to summon two other Coyotes (presumed to be their offspring) from ~700 m away to join in the successful pursuit of an adult male White-tailed Deer (Odocoileus virginianus). Our observation suggests that Coyotes can use vocalization as an effective means of coordinating social activities such as the hunting of large prey. Key Words: Eastern Coyote, Canis latrans, predation, vocalization, social organization, Nova Scotia. Auditory communication serves an important role in the social ecology of the Canidae (Harrington and Mech 1978, 1979; Theberge and Falls 1967; Gese and Ruff 1998). Howling is a means of long-distance communication that apparently can be heard at dis- tances of > 6 km (McCarley 1975; Harrington and Mech 1979). Among Coyotes (Canis latrans) and Wolves (Canis lupus), howling is largely accepted as playing a role in territory maintenance (Joslin 1967, Harrington and Mech 1978, 1979; Gese et al. 1988). Intrapack communicatory roles, such as announcing departures from den and rendezvous sites, and coordinating re- unions or movements remain largely speculative (The- berge and Falls 1967; Mech 1970; Harrington and Mech 1978). Bender et al. (1996) suggested that howl- ing among Coyotes might serve to reunify packs to facilitate the hunting of ungulates. Herein we report a case where a breeding pair of Coyotes in pursuit of a large White-tailed Deer (Odocoileus virginianus) was able to entice two other Coyotes to join the pursuit from ~700 m away. Observation We interpreted the details of the chase while snow- tracking a radio-collared Coyote and his mate on 27 February 1994 as part of a study of the effects of the distribution and abundance of Snowshoe Hares, Lepus americanus, and White-tailed Deer on the life history of Coyotes in Nova Scotia (Patterson and Messier 2000, 2001; Patterson et al. 1998). The actual event probably occurred during the previous night. Radio-collared Coyote AM3 and his mate were traveling west on a snow-covered (10-15 cm) secondary road | km east of Kejimkujik National Park (44°20'N, 65°15'W) when they abruptly veered due north. Lengthened strides indicated that their pace increased, and after 40 m the pair jumped three deer, of which two headed west and the other east. Both Coyotes chased the deer that ran to the east. After 280 m the two Coyotes split up, with the larger radio-collared male chasing the deer down a steep bank into a bowl- shaped depression. AM3 then swung wide to the left of the deer, apparently in an attempt to steer the deer into the path of the other Coyote that had remained on the rim of the depression. The second Coyote cir- cled the depression and ran down the opposite side in front of the deer. We interpreted this as an effort to prevent the deer from leaving the depression. Within 3 m of the second Coyote’s resumption of the chase, both Coyotes attacked the deer and drew hair but no blood. The deer escaped and again attempted to run up the side of the depression when AM3 swung to the right and turned the deer back down into the depres- sion, where the second Coyote was waiting and again resumed the chase. The Coyotes attacked the deer again at 383 m where a larger area of snow was trampled down than during the first attack. There was more hair strewn about, but still no blood. At this point two more Coyotes became involved in the chase. At 390 m more hair was detected as well as blood, indicating another attack. The deer broke away once more only to be attacked again at 397 m. After escaping yet again the deer made a long run across the middle of the depression. Considerable amounts of blood and hair strewn about a 20 x 30 m packed-down area at 590 m indicated a more serious struggle. The deer made one final escape and was pulled down by the front end at 630 m, with tracks indicating that the deer was drag- 2004 ging at least two of the Coyotes, which appear to have been hanging off its sides. After dragging the Coyotes ~ for 40 m the deer was killed at the 670 m mark. Examination of the carcass revealed that the deer was a large 3.5 year old buck that showed no obvious debilitations and was apparently in good health (>80% femur marrow fat content and other visible body fat reserves). We believe that the deer would likely have escaped if it could have got out of the depression. The two Coyotes initially involved in the chase appeared to have trouble drawing blood from the deer until the other two Coyotes joined them. Backtracking later revealed that the two Coyotes that joined the chase in progress had been traveling in another direction when they abruptly turned and trotted 690 m in a direct line to join the chase. Forest cover was dense, precluding any possibility that these Coyotes observed the chase prior to joining in. AM3 and his mate were typically accompanied by two of their young of the year when traveling in winter 1994 (Patterson and Messier 2001). We believe that the two Coyotes that joined the chase “in progress” were probably these same juveniles that had been temporarily disassociated from their parents. We speculate that they must have heard AM3 or his mate howling or yipping and were able to determine that it would be to their benefit to join the breeding pair promptly. Discussion Although we can not verify that howling was used to draw the other two Coyotes to the scene of the chase, we can think of only one other means by which two Coyotes ~700 m away in forested cover may have been able to so directly and rapidly locate the scene of the chase. White-tailed Deer can snort loudly when alarmed, but snorts are generally only given when a deer per- ceives danger but does not feel directly threatened (Hirth and McCullough 1977; Marchinton and Hirth 1984). Furthermore, snorts are more likely to be given by maternal family groups than by bucks (Hirth and McCullough 1977). Thus it seems unlikely that snort- ing by the buck alerted the other two Coyotes to the chase. Among forest-dwelling eastern Coyotes increased reproductive fitness and inclusive fitness for juveniles before dispersal seem to be the ultimate factors influ- encing group living (Messier and Barrette 1982; Pat- terson and Messier 2001). Increased efficiency at using large prey appears to be a secondary benefit (Gese et al. 1988; Messier and Barrette 1982; Patterson and Messier 2001). Coyotes in Nova Scotia were more suc- cessful at killing deer when thick snow cover impeded deer movements (Patterson and Messier 2000). There was only 10-15 cm of snow on the ground during the event described here and our observations suggest that in this particular incidence the snow cover may have been a hindrance to the Coyotes, thus benefiting the NOTES 279 deer. It was clear that AM3 and his mate were having difficulty subduing the deer on their own. We believe that they would not likely have been able to make phys- ical contact with the deer if it had not entered the depression. Although Patterson and Messier (2000) did not detect a consistent increase in deer killing rates for groups of 2-5 Coyotes, groups of >4 Coyotes killed proportionately more deer (Patterson 1999). We suspect that larger group sizes may be more advan- tageous in hunting large prey in the absence of other contributing factors such as thick snowcover or glare ice. The proximate mechanism is likely an increase in the probability of at least one member of the group making physical contact with the deer and slowing it down enough for other group members to assist in dispatching it. Our observation supports this idea and suggests that Coyotes can use howling as an effective means of coordinating social activities such as the hunting of large prey. Acknowledgments This work was supported by Parks Canada, the Nova Scotia Department of Natural Resources and Acadia University. We appreciate comments by D. O. Joly on an earlier draft of the manuscript. Literature Cited Bender, D. J., E. M. Bayne, and R. M. Brigham. 1996. Lunar condition influences coyote (Canis latrans) howling. American Midland Naturalist 136: 413-417. Gese, E. M., and R. L. Ruff. 1998. Howling by coyotes (Canis latrans): variation among social classes, seasons, and pack sizes. Canadian Journal of Zoology 76: 1037- 1043. Gese, E. M., O. J. Rongstad, and W. R. Mytton. 1988. Relationship between coyote group size and diet in south- eastern Colorado. Journal of Wildlife Management 52: 647-653. Harrington, F. H., and L. D. Mech. 1978. Howling at two Minnesota wolf pack summer homesites. Canadian Journal of Zoology 56: 2024-2028. Harrington, F. H., and L. D. Mech. 1979. Wolf howling and its role in territory maintenance. Behaviour 68: 207-249. Hirth, D. H., and D. R. McCullough. 1977. Evolution of alarm signals in ungulates with special reference to white- tailed deer. American Naturalist 111: 31-42. Joslin, P. W. B. 1967. Movements and homesites of timber wolves in Algonquin Park. American Zoologist 7: 279-288. Marchinton, R. L., and D. H. Hirth. 1984. Behavior. Pages 129-168 in White-tailed deer: Ecology and Management. Edited by L. K. Halls. Stackpole Books, Harrisburg, Pennsylvania. McCarley, H. 1975. Long distance vocalizations of coyotes (Canis latrans). Journal of Mammalogy 56: 847-856. Mech, L. D. 1970. The wolf: the ecology and behavior of an endangered species. Natural History Press, New York. 384 pages. Messier, F., and C. Barrette. 1982. The social system of the coyote (Canis latrans) in a forested habitat. Canadian Jour- nal of Zoology 60: 1743-1753. 280 Patterson, B. R. 1999. The effects of prey distribution and abundance on eastern coyote life history and predation on white-tailed deer. Ph.D. thesis, University of Saskat- chewan, Saskatoon, 204 pages. Patterson, B. R., L. K. Benjamin, and F. Messier. 1998. Prey Switching and feeding habits of Eastern Coyotes in Relation to Snowshoe Hare and White-Tailed Deer Densi- ties. Canadian Journal of Zoology 76: 1885-1897. Patterson, B. R., and F. Messier. 2000. Factors Influencing killing rates of White-Tailed Deer by Coyotes in Eastern Canada. Journal of Wildlife Management 64: 721-732. THE CANADIAN FIELD-NATURALIST Vol. 118 Patterson, B. R., and F. Messier. 2001. Social organization and space use of Coyotes in Eastern Canada relative to prey distribution and abundance. Journal of Mammalogy 82: 463-477. Theberge, J. B., and J. B. Falls. 1967. Howling as a means of communication in timber wolves. American Zoologist 7: 331 -338. Received 24 September 2002 Accepted 11 October 2004 Book Reviews ZOOLOGY Fish of Alberta By Amanda Joynt and Michael G. Sullivan. 2003. Lone Pine Publishing, Edmonton, Alberta. 176 pages. $18.95. ISBN 1-55105-191-5. This book is a popular account of the 54 established and 11 more rare fishes of Alberta. It consists of Acknowledgements, Foreword (by J. S. Nelson, the expert on fishes of Alberta), Reference Guide, Intro- duction, Keys to the Fishes, Species Accounts, Size Comparisons, Other Alberta Fishes (the rarer species), Glossary, Checklist, Selected References, Further Infor- mation, Index of Scientific Names, Index of Com- mon Names and About the Authors. These are mostly self-explanatory. The Reference Guide is a colour pictorial summary of each of the 54 species while Size Comparisons show the fish in relation to a loonie or a page in the book. The Glossary, Selected Refer- ences and Further Information sections are short but give some additional sources and explications of terms. Schreckstoff is misspelled and the term “alarm sub- stance” is more user-friendly. Laterally compressed is a tautonym, as compressed means flattened from side to side. For a more complete listing of terms used in ichthyology see www.briancoad.com. The ichthyofauna includes 16 families with min- nows (Cyprinidae) having 17 species, 3 introduced; trouts (Salmonidae) 16 species, 6 introduced; and suck- ers (Catostomidae) 7 species. Other families are lam- preys, sturgeons, mooneyes, bullhead catfishes, pikes, trout-perches, cods, livebearers (2 species introduced to the Banff hot spring marshes), sticklebacks (1 introduced), sculpins, sunfishes (introduced), perches and cichlids (1 introduced to the Banff marshes). The Introduction has information about identifying fishes, the lives of fishes, adaptations to life in water, the underwater world, drainage basins of Alberta, top Alberta fishwatching sites and fishwatching (“not yet a popular sport’), history of fishes and conservation issues. Key words are in bold text and some, but not all of these, have illustrations. One illustration, of Bull Trout and Walleye larvae used to explain reproductive strategies, is confusing in that these two distinctive species have the same larva illustrated. The Species Accounts comprise two pages for each established species, in landscape orientation. This requires some twisting for reading introductory and other sections that have a standard orientation. None- theless, the landscape arrangement enables the fish illustrations to spread across the broadest reach of the page. Each Account has an introductory section, Viewing Tips (localities and habitats where the fish can be seen), Feeding, Spawning, Other Names, Did You Know? (an anecdote or interesting factoid), ID (identification characters), Similar Species, Status, Habitat, Length and Weight, the colour illustration and a colour, shaded distribution map. Generally, the book is a good, popular treatment of this fish fauna written in a readable style that is not as dry as more scientific accounts. Some errors occur, per- haps as a result of this style or perhaps from the more pedantic viewpoint of the reviewer; e.g., in the Long- nose Sucker account pharyngeal teeth are said to be in the mouth which could mislead the reader into thinking these throat teeth are readily visible. The definition of “fish” and “fishes” is given on page 10 in the Introduction, fish being a single individual or more than one individual of a single species while fishes refers to more than one species. Curiously, the book is wrongly titled. The Reference Guide has a col- our code for groups of species but since these colours are various shades of green and blue, they do not lend themselves to a ready means of locating species groups. The fish illustrations are positioned with the head to the right, not a problem for most users, but somewhat disconcerting to those familiar with most fish books where traditionally the head faces left. The colour illus- trations lack the details that can be included on line drawings, so certain critical features are not always visible. The illustrations have streaks of white meant as highlights but in some specimens look like a colour pattern. Scientific names are given for all species and are ac- curate and, although Stizostedion may now be correctly Sander, the decision to leave it as the more familiar North American genus is probably apposite for infor- mation retrieval. Conversely, the Arctic Lamprey is given its newer name, Lampetra camtschatica, which is less well known than the older version, L. japonica, which is not mentioned and would inhibit search for information in other books. Common names are stan- dard and include the Northern Pikeminnow, although its former name Northern Squawfish was not so much an “unpopular” designation as a derogatory one. Despite the minor criticisms outlined above, this book is a good introductory guide to Alberta’s fishes. Alberta seems well served in this respect with Nelson and Paetz’s earlier book (1992). The older work pro- vides a more detailed account of the fish fauna and may be more suited to the serious student of Alberta’s 281 fishes — there is an extensive list of references (24 pages compared to only 11 references in Joynt and Sullivan). The book reviewed here has accounts for 65 species while Nelson and Paetz have a more complete analysis of 59 species and a further 29 species recorded as rare or doubtful for Alberta. Sharks By Andrea and Antonella Ferrari. 2002. Firefly Books, Tor- onto, Ontario. 256 pages. $24.95. ISBN 1-55209-629-7. Sharks are the most popular of fishes for publica- tions as people seem to have a fascination for organ- isms that can eat you (although more sharks are killed by people than the reverse). This is one of many that have appeared on this topic. Despite its title, the book covers the appearance and behaviour of 120 species of sharks and rays. Coverage is therefore not complete as sharks and rays number over 1000 species (as the Foreword states), the aim be- ing to give an overview of the main groups by select- ing typical and unusual or fascinating species. Four species of rabbitfishes (out of about 31 species in the Chimaeriformes) are also mentioned. The book has an Introduction of 77 pages with descriptions of anatomy and biology. Three pages are devoted, deservedly, to explaining the ampullae of Lorenzini, an important feature of shark anatomy and biology, first discovered by an Italian scientist in the seventeenth century and, surely by coincidence, this book has Italian authors and is a translation from an Italian version. This is followed by how to avoid sharks, how to assist them (by not eating them or buying shark teeth), marine organisms more dangerous than sharks (mostly venomous sea snakes, cone shells, sea urchins, jellyfishes and fishes, but also fish that bite divers en- thusiastically), and personal accounts of shark encoun- ters. The Entries section describes the sharks, rays and rabbitfishes (153 pages), the Appendices (17 pages) give a Classification of Sharks (but not rays or rabbit- fishes), an Index, Bibliography and Websites (each a page long), and Photographic Credits. The websites “Catalog of Fishes” and “FishBase” are not quoted, although these give entry to much of the names, biol- ogy and literature on sharks and rays world-wide for the more serious student. There are pictorial keys to In Search of the Golden Frog By Martha Crump. 2002. University of Chicago Press, 5801 Ellis Avenue, Chicago, Illinois 60637 USA. xiv + 298 pages, Cloth US$27. Any young person thinking of taking a degree in one of the biological sciences, particularly if they plan to become a field biologist, should read this book. The author, currently Adjunct Professor of Biology at Northern Arizona University, chronicles her life from THE CANADIAN FIELD-NATURALIST Vol. 118 Literature Cited Nelson, J. S., and M. J. Paetz. 1992. The Fishes of Alberta. The University of Alberta Press, Edmonton and the University of Calgary Press. 2"4 Edition, 437 pages. BRIAN W. COAD Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada the shark and ray orders but not any for families or species so the book cannot be used, nor is it meant, as a field guide. The Entries section describes each selected species, allotting half to two pages per species. There is an annoying symbol system at the top of each account to indicate when the shark is active and its danger level with respect to humans. This information could easily have been included in the text. A distribution map is given but at | x 2 cm can only convey a general impression of where these sharks and rays are found. The map for the Bluntnose Shark is in- accurate, for example, showing it in Canadian Arctic waters when it was first caught at its northern limit in Nova Scotian waters in 1989 (Gilhen and Coad 1991). The common and scientific names are given but the author and date of the scientific name is always in parentheses (here and in the Classification of Sharks). The authors (or text editor?) seem not to be aware that parentheses are only used when the species is placed in a genus other than the one it was originally described under. The text comprises the Family to which the shark or ray belongs, Range, Habitat, Size, and Habits. All parts of the book are richly illustrated with colour photographs, over 450 in all. Despite the comments above, this book is a good general introduction to sharks and rays. It is lavishly illustrated, as all such books must be, of a convenient size, and with a reasonable price. Literature Cited Gilhen, John, and Brian W. Coad. 1991. The bluntnose sixgill shark Hexanchus griseus (Bonnatere, 1788), new to the fish fauna of At- lantic Canada. Proceedings of the Nova Scotian Institute of Science, 39 (1989): 75-77. BRIAN W. COAD Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 Canada her postgraduate work in 1968 through various projects to 1998. The book concentrates on the field programs that are the real love of Professor Crump’s life. Her main interests are reptiles and amphibians, but her passion is frogs. Taking advantage of an opportunity offered by her discerning professor, she joins a team going to the Amazon section of Ecuador. From this exposure which 2004 reinforces her desire to work in the field, she returns | repeatedly to work in South America. Her narrative _ includes her excitement at being a scientist and mak- ing discoveries to contribute to her profession. She also chronicles some of the problems she encounters. These do not include frightening encounters with large or dangerous animals. The hazards fall into two cate- gories. The small pests, mosquitoes, chiggers, chigoes, bot flies and ants, causes plenty of discomfort. But it is the large pest, humans, that causes the greatest levels of concern. She notes the ordinary people are by and large, friendly and helpful. It is the bandits, police, _ military, industrialists and rioting natives that disrupt her life. As I started to read this book I became very dis- concerted. In the past few years I have looked for amphibians and reptiles in tropical jungles on many _ occasions. My success rate is very low. The author travels to the same locations in South America and immediately starts finding many individuals of multi- ple species. My estimation of my frog finding prowess dropped page by page. By the time the author reaches the 1990s, however, her success rate has plummeted. By 1996 she says “we hug and screech with excite- ment” on finding one frog. I realize that my lack of success is due less to my incompetence than to the Conversations with an Eagle By Brenda Cox. 2002. Greystone Books, Douglas & McIntyre Publishing Group, 2323 Quebec Street, Vancouver, Brit- ish Columbia VST 487. 261 pages, paperback. This is a story of dedication to an eagle rather than conversations with one. Brenda Cox was a volunteer worker at a raptor rehabilitation centre south of Van- couver (Orphaned Wild Life — OWL), and became enthralled with raptors of all kinds. Her special inter- est was a Bald Eagle, which was three months old when it arrived at the Centre, and her book describes the trials and tribulations of her efforts to train Ichabod so that she could be used in the OWL education pro- gramme. In the course of the story there are passages de- scribing the methods used to rehabilitate owls, hawks and eagles. When the eagle was still young its behav- iour was typically fearful, but the bird became more and more aggressive towards Cox, such that she had to stand against a pillar armed with a household mop to protect herself if she was attacked. In spite of the danger, she persisted in training the bird to come to her gloved arm, and eventually it obeyed three com- mands: Up, Wait and Off. But it was always uncer- tain whether the bird would cooperate or not on any given day. She enlisted the help of expert falconers in the Vancouver area. They were doubtful whether a Book REVIEWS rapid loss of frogs in my lifetime. This is even more depressing. When I go to Monteverde next spring I have little hope of seeing the fabulous Golden Frog. On 7 April 1987 this author saw “over one hundred dazzling bright golden toads“ at one small pool at Monteverde. The author also deserves another accolade for, while she 1s pursuing her scientific research, she is also be- ing a parent to two children. I was delighted with the way she involved them in her work and gave them opportunities to broaden their knowledge. This may have made her life harder and certainly caused her concerns, but I am sure these children will benefit. So if you want to find out about the life of a field researcher read this entertaining book. Follow the trials of first reaching the field locations using often unreli- able local transport. Join in meals that are good, bad, or bizarre. Meet local people, many of whom care greatly about their environment. Find out how a sci- entist collects data and uses it to create a new under- standing of our planet. Roy JOHN 2193 Emard Crescent, Ottawa, Ontario, K1J 6K5 Canada Bald Eagle could be reliably trained, though they helped and advised her. Bald Eagles have seldom been trained successfully, while Golden Eagles have been used by falconers in many parts of the world. Cox was able to train the eagle such that she could take her from her cage to a perch in a field, using jesses and a stout leash attached to her waist, but there were increasing dangerous attacks which inflicted talon wounds on Cox’s head, arms and feet. After several years, the OWL management decided that they would never be able to use the bird in their programme and asked Cox to remove it from the facil- ity. In its new quarters, shortly afterwards the bird developed a lung disease and was euthanized. Cox’s dedication to her volunteer job is remarkable since, in the course of the seven years, she did not have steady employment or reliable housing or transportation. Since Ichabod’s death, she has now found a steady career: as a conductor with British Columbia Rail, and watches the wild raptors along the railway lines. Her story would appeal to someone interested in raptor rehabilitation methods and the lore of falconry. JANE ATKINSON 255 Malcolm Circle, Dorval, Quebec H9S 1T6 Canada 284 THE CANADIAN FIELD-NATURALIST Vol. 118 North American Owls, Biology and Natural History, Second Edition Paul A. Johnsgard. 2002. Smithsonian Institution Press, Wash- ington. 298 pages, 42 colour plates, 69 multi-figured black- and-white maps and sketches, 10 tables, 2 graphs. $46.25 Canadian, hardcover. US$27.95. Although Dr. Johnsgard’s long history of produc- ing books about birds is well known, his entry to the “owl-lover’s camp” is more recent. The first edition of this book came out in 1988 and immediately became the standard reference book for owls of the United States and Canada. For the second edition, 14 years later, Johnsgard has added twelve species of Mexican owls and nearly doubled the number of references (although, contrary to convention, many of the recent references are not mentioned by name in the text!). Extremely few changes have been made to the excel- lent introductory chapters, but this leaves them less up- to-date than the individual species accounts. Often he cites the authors of the Birds of North America account, rather than the original observers who merit the credit; an example is Gottfred and Gottfred’s new informa- tion concerning courtship and copulation of the Great Horned Owl, published in Blue Jay in 1996. The strengths of the first edition have been main- tained: the introductory chapters and species accounts are neatly arranged; maps of each species’ range are detailed; Johnsgard’s pen-and-ink sketches of aspects of behaviour are delightful; the colour plates and pho- tographs are superb. Johnsgard writes well, but in a few occasions new material has been interjected clum- sily, marring the previous smooth flow of information in the first edition. Some compromises were necessary. To make room for the twelve Mexican species, detailed descriptions of plumage have been omitted in this second edition. This does not excuse three sloppy errors. Johnsgard tells of the Pygmy Owl range extending altitudinally to 37 000 meters in Mexico, whereas the quoted source gave a credible 3700 m. A paper by Jack Holt on Great Horned Owls in the Cincinnati region is credited to Denver Holt. Thirdly, when Johnsgard withdrew his overdrawn sketch of the “false eye-spots” on the back of the Northern Pygmy Owl’s head from the first edi- tion and substituted a more realistic sketch of the Fer- ruginous Pygmy Owl in the second edition, he failed to change the figure numbers in the text to comply with this change. These minor caveats aside, this masterful second edition will be welcomed by owl enthusiasts around the world, and should be purchased even by those own- ing the first edition. It is written more for the scientist than the amateur, but, by identifying deficiencies in our current knowledge, this book should offer possi- bilities for future study by graduate students and others who love our night-time friends. Johnsgard’s exquisite sketches, 10 coloured paint- ings of owls by Louis Agassiz Fuertes, and 31 fine col- our photographs, including nine of Mexican owls, make this an unusually attractive book. Buy it! MARTEN J. STOFFEL Box 183, RR#4, Saskatoon, Saskatchewan S7K 3J7 Canada The Mountain White-Crowned Sparrow: Migration and Reproduction at High Altitude By Martin L. Morton. 2002. Studies in Avian Biology Number 24. Cooper Ornithological Society, Camarillo, California. 236 pages, 8 colour plates, pencil illustrations heading each chapter. U.S.$27. The northern limit of the distribution of the Moun- tain White-Crowned Sparrow (Zonotrichia leucophrys oriantha) extends slightly into southern British Colum- bia, Alberta and Saskatchewan. Its main breeding range is in mountainous regions of the western U.S. Here in Waterton Lakes National Park we’re at the contact zone between oriantha and Z. |. gambelii. Morton’s 25-year study of oriantha in the Tioga Pass area of California’s Sierra Nevada ranks with P. J. Greenwood’s lengthy study of the Great Tit (Parus major) in Europe and G. E. Woofenden’s work with the Florida Scrub Jay (Aphelocoma coerulescens) in showing the value of continuous long-term field inves- tigations in advancing a broad spectrum of ideas and hypotheses in avian biology. Morton’s focus is differ- ent, however, in emphasizing physiology more than behaviour or ecology. In 1968, Morton recognized that there were significant gaps in knowledge of mi- gratory passerines on their summering grounds, espe- cially in mountains where large variations in environ- mental conditions occur. As my interests lie more with natural history, life history data (age at maturity, number, size and sex ratio of offspring, dispersal and survival rates) and ecolog- ical factors than with physiology, I found Morton’s first three chapters on Migration Arrival, Social System and Behavior, and Demography of most interest (al- though a bit disappointing at only one-quarter of the book’s pages). The dynamics of arrival of oriantha varied greatly from year-to-year depending upon the amount of remaining snowpack and the frequency of spring storms, as well as by age and sex. By trapping and colour-marking individuals, the researchers found that older males (age 2+ years) generally arrive ear- liest, followed by one-year-old males, older females and one-year-old females. Throughout the monograph, Morton does an excellent job of analysing observed data in terms of costs and benefits, and ecological fact- 2004 ors, in this case, suggesting that older, experienced birds knew the migration route and recognized the _ breeding area once they reached it, even it if was snow- covered. Morton’s efforts over many reproductive seasons made it possible to measure mate fidelity, age of mates, frequency of polygamous pairings, agressive behav- iours, and the functions of vocalizations, in addition to the usual study of territory establishment, pairing, and between-year breeding dispersal. Although I expected the chapters on Gonadal Con- dition, and Body Size and Body Condition, to be of less personal interest, Morton’s explanations of the connections between physiology and behaviour, and | description of the role of environmental cues in annual _ cycles made these sections much more interesting than anticipated. Environmental factors are either ultimate (e.g., availability of an adequate food supply, predation pressure, weather patterns) or proximate (e.g., photo- | period, ambient temperature) in their effects on the timing of reproduction. Nest history (chapters 7, 8, 9 and 10 on Nests and Eggs, Nestlings and Fledglings, Nest Failure and Book REVIEWS Reproductive Success, respectively) provided reliable information on physiological and behavioural respons- es of breeding birds to environmental variation, often to the level of individuals because of Morton’s use of marked birds. And the researchers determined that the snowpack, because of its effects on nesting sched- ules and nest locations, was a stronger environmental factor on reproductive success than sub-freezing tem- peratures or summer storms. Oriantha exhibited plas- ticity in responding to snow conditions by abandon- ing ground-nesting and building their nests in elevated sites when there was more snow, as opposed to Her- mit Thrushes (Catharus guttatus) or Dark-eyed Juncos (Junco hyemalis), which nested on the ground no matter what the environmental conditions. Morton’s study shows that challenges posed by en- vironmental variation often can be met with existing behavioural and physiological responses; adaptation occurs through flexibility rather than through acqui- sition of new abilities or mechanisms. CynpI M. SMITH Box 5, Waterton Park, Alberta TOK 2MO Canada Seabird Bycatch: Trends, Roadblocks, and Solutions Edited by Edward F. Melvin and Julia K. Parrish. 2001. Uni- versity of Alaska Sea Grant AK-SG-01-01, Fairbanks, Alaska. viii + 206 pages. U.S.$20.00. The incidental catch of various water birds and other aquatic organisms (e.g., turtles, sea otters, porpoises, seals) in fishing nets has long been of concern to naturalists for conservation reasons and to the fishing industry for economic reasons. However, until recent- ly this issue has been on the “back burner” compared with more pressing and widespread environmental problems. The advent of intensified fishing efforts, huge factory ships, and collapsing fish stocks have elevated this bycatch into both a major international environmental issue and a serious economic problem. This book, essentially the proceedings of a sympo- sium, presents a recent update on the extent of the problem in marine waters, world-wide, evaluations of several proposed solutions as they apply to specific fisheries and problems that have yet to be overcome. The editors begin the book with a short preface that outlines the history and extent of the problem, espe- cially since it was identified as a major conservation issue in the early 1970s. The first full chapter is a syn- thesis, also by the editors, of a symposium organized by the Pacific Seabird Group in Blaine, Washington, in 1999. The synthesis summarizes the complexity of the problem, “roadblocks” in the way of solving it and guidelines to possible solutions. The remainder of the book consists of nine peer-reviewed, scientific papers (eight of which were presented at the symposium) and seven abstracts. Two of the papers were reprinted from recent (1999 and 2000) scientific journals. One paper, by John Cooper, John Croxall and Kim Rivera outlines the complexities of international efforts to re- duce bycatch through research, local and international regulations and cooperation among scientists, the fish- ing industry and conservation groups. An afterword by Craig Harrison graphically illustrates the complexity of the problem by outlining the political and practical difficulties of coordinating efforts to reduce bycatch in the sockeye salmon fishery in the “shared waters” of British Columbia and Washington, where two fed- eral, one provincial and one state government share jurisdiction with “21 Native American tribes” [a total that applies only to Washington and omits the first nations of British Columbia]. The rest of the papers and abstracts document the extent of the bycatch in specific fisheries, results of various experiments to reduce bycatch, and attempts to measure the impact of specific fisheries on specific populations of birds. As some seabirds spend non- breeding times thousands of kilometers from their breeding sites, measuring the effects of a specific fish- ery on a particular breeding population is very diffi- cult, especially as many seabirds are long-lived and don’t breed every year. Moreover, fishing in a given area may affect some species much more profoundly than their close relatives (for example, effects of by- catch on Black-footed Albatross populations are much more significant than on Laysan Albatrosses) and may affect different age groups differently (for example, juvenile Black-footed Albatrosses are caught at a high- er rate than adults). Research presented in this vol- ume from the Atlantic, Pacific and Southern (Antarc- 286 tic) oceans shows that modifications to fishing gear (weights, design, visual and acoustic alerts), timing of fishing (daily and seasonally) and other techniques can reduce bycatch substantially without substantially reducing the take of target species, but that these mod- ifications have different effects on the rate of bycatch of different species and sometimes of a given species in different areas, different times of the day or different seasons. Banding continues to be helpful in sorting out the origins of birds caught in specific fisheries, while higher tech tracking devices and research on genetic markers are also starting to improve our knowl- edge of the complexities of movements by these highly mobile species. Albatrosses receive the greatest amount of attention, but data on various other “tubenoses,” alcids, larids and some marine mammals are also pre- sented. Canadian content figures prominently in this book. A 1972 paper by Canadian C. Eric Tull and co-authors on the magnitude of Thick-billed Murre mortality in Greenland fisheries is credited (page v) in drawing attention to the significance of fisheries in seabird population declines, and international legal efforts to reduce bycatch in longlines stem from a resolution at a 1996 meeting in Montreal of the World Conserva- tion Congress of the World Conservation Union (page 11). Data from British Columbia are included in a genetics (DNA) study of the breeding locations of Common Murres entangled in fishing nets in Wash- ington’s San Juan Islands and Puget Sound (pages 125-127) and in an abstract (pages 191-192) on sea- bird avoidance experiments in northern Pacific waters. Other papers and abstracts on observations and re- search in Alaskan and Washington waters involve populations of birds that spend portions of their lives in British Columbian waters. Research from Atlantic coast provinces and British Columbia are cited fre- The Life of Mammals By David Attenborough. 2002. Princeton University Press, New Jersey. 320 pages, U.S.$29.95. As the latest edition in the BBC “Life” series com- bining television and print media, the book on mam- mals is nicely illustrated with a good selection of colour photographs that closely follows the engaging text by David Attenborough as he presents interesting stories on these fascinating animals. The first chapter, “A Winning Design’, starts off with the ability of mammals to adapt to different environments on earth, including the harsh arctic conditions where lemmings live year round. After describing some basic charac- ters of mammals, such as hair and the production of milk, there is a general introduction to the origin and evolution of this group of warm-blooded organisms. The chapter finishes off with two early mammalian radiations that cover the egg-laying monotremes and THE CANADIAN FIELD-NATURALIST Vol. 118 quently. This book provides a useful compilation of recent developments on a complex issue, with plenty of in- formation on legal (national and international)/poli- tical aspects, recent research on seabird movements, recent research on genetics of seabird populations, and practical tests of various techniques under a variety of conditions in a variety of waters. The literature cited sections provide numerous additional references for those requiring further information. The book should be on the library shelves of any researchers, govern- ment agencies and non-government conservation and scientific organizations studying seabird biology and/ or the conservation of seabirds. Errors appear to be few — the reference to Wilson et al. (1985) cited on page 116 is not listed in that chapter’s literature cited section, which includes two references that don’t appear to have been cited. However, since most of tne chapters are written as scientific papers, some may be too tech- nical for some readers. The review by Cooper et al. of legal and quasilegal aspects of the issue provides a valuable reference compendium, but is so riddled with acronyms that I found myself wishing for a glossary. I also hope that the seven abstracts at the end of the book are expanded into full papers somewhere. As valuable a contribution as the book is in itself, this volume will undoubtedly also stimulate more research that will require another update before long. Such an update could usefully also include papers or chapters on aquatic bird bycatch of fisheries on inland waters, such as those of the Great Lakes, several large prairie province lakes and similar lakes on other continents. MARTIN K. MCNICHOLL 4735 Canada Way, Burnaby, British Columbia V5G 1L3 Canada the marsupials, which give birth to under-developed young. The remaining nine chapters deal with the placen- tal eutherian mammals but instead of continuing to describe them by scientific groups the format switches to artificial categories such as diet and habitat. This unnatural classification seems awkward at times with bats, which people can readily identify with, split into two separate chapters, “Insect Hunters” (curiously including vampire bats) and “Life in the Trees” (al- though some insect-eating bats also live in trees). There was a missed opportunity to educate readers in scien- tific classification and evolution while still entertaining them with a plethora of amazing natural history stories. Primates get star billing in the book with the last two and a half chapters devoted to this charismatic order of mammals. Although the higher-level taxonomy (or common names employed) is not current, it begins 2004 _ with prosimians as an early branch of the primates. The latest view is that this is not a natural group be- cause, for example, tarsiers are more closely related _to monkeys and apes, as alluded to but nonetheless still included in the chapter with lemurs. The other primates are grouped to cover the new and old world _ monkeys, and ending with the gibbons and great apes, including a branch for humans. The last half chapter concentrates on both physical and social anthropology from the first evidence of bipedal locomotion to cul- tivation and civilization. The book is definitely aimed at a general but knowl- _ edgeable audience with an interest in nature and mam- _mals. I am sure, however, that practising biologists will still find a few facts new to them in their non- specialist group because the background research is _ relatively good. It was nice to see some recent scien- tific hypotheses on mammalian evolution making it into the book such as the close relationship between BOOK REVIEWS whales and hippopotamuses. But some other emerging ideas based primarily on molecular data did not, in- cluding the association of bats with carnivores, ungu- lates and whales, as opposed to insectivores, or tree shrews, flying lemurs and primates. My criticisms are mostly biologically oriented be- cause the book is attractively presented with most photographs of good quality and information well writ- ten. But there should be more books that combine current scientific research with an explanation of the deeper implications or processes involved for wider distribution to the general public looking for mean- ingful substance beyond the usual cursory facts. BurRTON K. Lim Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, 100 Queen’s Park, Toronto, Ontario M5S 2C6 Canada _ Geographic Variation in Size and Shape of Savannah Sparrows (Passerculus sandwichensis) _ By James D. Rising. 2001. Studies in Avian Biology Number 23. Cooper Ornithological Society, Camarillo, California. 65 pages. U.S.$7. The newer field guides, such as The Sibley Guide to Birds, occasionally describe and illustrate subspecies of birds, usually based on differences in plumage or _ soft parts. The “bander’s bible”, Peter Pyle’s Identi- _ fication Guide to North American Birds, includes wing and tail measurements to help banders identify birds to the subspecies level — for example, he dis- cusses 14 subspecies of the Savannah Sparrow. But have you ever wondered why these differences exist in the natural world? Rising does and asks two questions in this regard: Why do features such as body size, wing length, or bill size and shape differ across a species’ range? and, if these differences reflect adaptations to the different environments to which the species is exposed, what are the selective factors that have caused them? One of the classic explanations for geographic vari- ation in size is Bergmann’s Rule, which holds that individuals of a species (vertebrates only) from colder areas are generally larger-bodied than individuals from warmer areas. Allen’s Rule takes this one step fur- ther, stating that within such species, individuals from colder areas will have smaller appendages relative to their body size than individuals from warmer areas. Rising asked his questions of the Savannah Spar- row, which is one of the most wide-spread songbirds in North America. He describes and quantifies geogra- phic variation in the species throughout its breeding range, from Alaska to the Maritimes to central Mexico, and relates trends in phenotypic variation to environ- mental variation. He found some clinal variation in size of Savannah Sparrows, with birds from the northeast being slightly larger than those from the west, and birds in cool, moist areas were larger than those where it is hot and dry. But the species overall did not seem to follow Bergmann’s Rule; rather, measures of summer tem- perature and precipitation explained well the patterns of size variation. His more significant find, though, was that birds were larger on islands than on main- land sites, whether in the Aleutian Islands, Alaska, or on Sable Island, Nova Scotia. He speculates that the long, cool, moist summers on these islands results in a predictable and fairly rich food supply. This, com- bined with the rather long breeding season, allows multiple broods and perhaps enhanced competition for high quality territories. This competition for either food or territories might select for larger body size. Rising ends with some taxonomic comments, com- ing out on the side of the “lumpers’’. He sees no virtue in naming subspecies where the only way they can be reliably separated is by locality. He suggests recog- nizing only two subspecies of non-saltmarsh Savannah Sparrows, P. s. sandwichensis (large size) and P. s. princeps (large and pallid), whereas the nine saltmarsh subspecies seem to be clearly separable by morpho- logical characters. This is in contrast to the 17 sub- species currently recognized by the American Orni- thologists’ Union. Rising’s work relies on Principal Components and Discriminant Functions analyses, which, although I found it a little heavy going, was well presented sequen- tially and a good example of the use of these techniques for morphometric comparisons. It will be interesting to watch in the coming years how Rising’s thorough morphological studies interact with genetic analyses in assessing variation and the sub- species of Savannah Sparrows. CyNnbDI M. SMITH Box 5, Waterton Park, Alberta TOK 2MO Canada 288 THE CANADIAN FIELD-NATURALIST Vol. 118 Warblers of the Great Lakes and Eastern North America By Chris Early. 2003. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto, Ontario, M2H 3K1. 131 + pages. Cloth $24.95: paper $16.95 Sparrows and Finches of the Great Lakes and Eastern North America By Chris Early. 2003. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto, Ontario, M2H 3K1. 128 + pages. Cloth $24.95: paper $.16.95 These books are an expansion and revision of two earlier books by Chris Early (Warblers of Ontario and Sparrows and Finches of Ontario). They both follow the successful format of the earlier books. Each spe- cies is shown in two to five photographs (Warblers) or two to seven photographs (Sparrows). The actual number of photographs for each species depends on how variable the bird’s plumage can be. A short intro- ductory note is followed by descriptions of the key plumage characteristics and other relevant details. A 5 x 3 cm map of America shows the summer and winter ranges. “Warblers” covers 37 species plus one hybrid in some detail. An additional seven vagrant species and one race (the Yellow-rumped “Audubon’s” Warbler) are covered by brief comments and a single accom- panying photograph. “Sparrows” includes 25 species covered in full and 20 vagrant species. In addition to the North American sparrows (buntings) this volume includes longspurs, finches, grosbeaks, crossbills, snow bunting and house sparrow. There are two features used by this author that will make this book particularly valuable to beginners. The first is called a cheat sheet and lists all the birds in groups according to a key characteristic — such as an unstreaked breast. The birds are further grouped by a second characteristic such as a breast spot. The second, and even better innovation, is a series of com- Birds of the Yukon Territory Edited by P. H. Sinclair, W. A. Nixon, C. D. Eckert and N. L. Hughes. University of British Columbia Press, Van- couver, British Columbia. My appetite for a long-overdue return trip to the Yukon has been incredibly whetted! This book is not just an atlas, but also a coffee table book, a very use- ful bird reference, and a source necessary for a planning a trip to the Yukon. This book has stunning photographs throughout, and not just of birds, but of dramatic scenery (i.e., habitat shots) that will make birders and non-birders alike think, “I must go there.” Prior to the species accounts, there are chapters covering the environment, aboriginal use of birds, conservation, history and a month-by-month overview of birdlife. There is also a short chapter outlining the parison tables. These consist of a set of photographs with birds in similar poses. The page is arranged to show the most similar birds close together. For example, there are three Common Redpoll photographs imme- diately above three equivalent Hoary Redpoll photo- graphs. Similarly, all the rufous-capped sparrows are on one page. I think the arrangement of the warbler comparison table is very clever. Here the Mourning Warbler photograph is adjacent to a Connecticut War- bler photograph. In turn, the Connecticut is above a photograph of a Nashville Warbler, which in turn is above a Northern Parula. All four birds share similar combinations of colour and cause confusion to the novice. This arrangement allows for easy compari- son, either up and down or across the rows. For the warblers there are tables showing the spring and fall plumages. These features are on fold-out sheets at the end of the book, making them quickly available in the field. At the front of the book is a seasonal distribution list for Point Pelee National Park. Using line thickness the author also indicates the abundance of each species. I find such distribution lists to be extremely useful. These then are two small, portable books that deal with birds that many find difficult to identify. The text is well written. The photographs are both beautiful and illustrative. Any one who needs help with the species covered will find these books a tremendous help. Per- haps we can encourage the author to write a third book on shorebirds. Roy JOHN 2193 Emard Crescent, Ontario, K1J 6K5 Canada data management for the more than 160 000 records that were used. Each species account spans one to three pages. All species are illustrated by a line drawing and at least one, sometimes several, colour photographs; frequent- ly, there is a very good habitat shot as well. Each account also has a distribution map, and a histogram outlining the number of database records for each week of the year. The base layers of the maps show major roads and rivers as well as ecozones; I would have liked to see the dominant population centers marked as well though. The text is thorough and reads well. All the expect- ed sections are there (distribution, nesting, habitat...), and, where appropriate, a section on aboriginal use of 2004 the bird is included. Several appendices and a 600+ reference list end the book. The items that bothered me were few, and mostly minor. Noteworthy was the absence of birding essays, a feature I first saw in Birds of Delaware, and thought added spark to the thick reference book. Page numbers were located in the most irritating choice possible, along the inner margin of each page, thus not permit- ting a rapid flip-through while searching through this 600 page book. The final, and dare I say incompre- hensible feature of the book was the uninteresting and out of focus (soft focus?) cover shot of the Territorial bird, the Common Raven. People do judge a book by BOTANY BooK REVIEWS 289 its cover, rightly or wrongly, and in a book with hun- dreds of stellar shots, I remain baffled as to why that particular shot was chosen to be most people’s first impression of the book. Overall though, this is a very, very good book, and a must-have for people interested in the birdlife of the North. RANDY LAUFF Department of Biology, St. Francis Xavier University, Anti- gonish, Nova Scotia B2G 2W5 Literature Cited Hess, G. K., R. L. West, M. V. Barnhill I, and L. M. Fleming, 1999, Birds of Delaware. University of Pittsburgh Press. 635 pages. Wild Flowers of the Yukon, Alaska & Northwestern Canada, 2™ Edition By John G. Trelawny. 2003. Harbour Publishing Co. Ltd., P.O. Box 219, Madeira Park, British Columbia VON 2HO Canada. 224 pages. $24.95. The first edition of this wild flower book was pub- lished in 1983 and I reviewed it in 1984. The layout in this new volume is much better. The type is larger and more spaced on a purely white paper and is wrapped around the absolutely beautiful colour pho- tographs. Many of the pictures of the 332 species treated are the same as those in the first edition but the more modern reproductions and paper have made them shine. The descriptive text for each species has been only slightly revised. The surrounding text includes Acknowledgments, an easy-to-read map, an Introduc- tion, an Illustrated Glossary, a Key to Species using Flower Colour and Shape, Photo Credits, Bibliogra- phy, an Index, and a list of Additional Field Guides from Harbour Publishing. This is a most interesting and delightful book for anyone living in, visiting or planning to visit this wonderful area. WILLIAM J. Coby Biodiversity, Program on Environmental Health, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada. Wild Flowers of Field & Slope in the Pacific Northwest By Lewis J. Clark. 2002. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia VON 2HO. $9.95. Wild Flowers of Forest & Woodland in the Pacific Northwest By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia VON 2H0. $12.95 Wild Flowers of the Mountains in the Pacific Northwest By Lewis J. Clark. 2003. Harbour Publishing, P.O. Box 219, Madeira Park, British Columbia VON 2HO0. $12.95. These three books contain absolutely beautiful col- our photographs. The photographs are numbered sequentially and each one has a marker [x0.5] to in- dicate its size. Each is accompanied by a printed para- graph with the common and scientific names, a de- tailed description, together with the habitat and range, and a number to indicate its sequence in the book. There are 108 in the first book, 100 in the second and 106 in the third. There is a four- or five-page interesting introduction at the front of each book and an index, glossary and a list of Additional Field Guides available from Har- bour Publishing at the end. There are most interest- ing pictures on the front covers of each book. On the back covers there is a note about the author, Dr. Lewis J. Clark, together with a map of the Pacific Northwest on which there are shaded areas depicting where the wild flowers can be found. All are elegant. WILLIAM J. Copy Biodiversity, Program on Environmental Health, Agriculture and Agri-Food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Canada 290 MISCELLANEOUS The Canoe: A Living Tradition By John Jennings. Firefly Books, Toronto. 272 pages. $59.95. There could hardly be a more poetic way to ex- plore the natural world than by canoe. While not all observers of natural history are canoeists, most canoe- ists have respect for and curiosity about nature. In- deed, a loon’s call or the slap of a beaver’s tail, accom- panied by the quiet burbling of a canoe’s wake, has served as an introduction to the contemplation of nature for countless North Americans. The Canoe: A Living Tradition is, in a sense, a nat- ural history of traditional North American watercraft. Included in this beautifully illustrated, coffee table- sized book are chapters dedicated to the history and construction of birch bark and dugout canoes, as well as skin kayaks and umiaks (large, open vessels from the Canadian Arctic). This book also discusses the influence of canoes on the fur trade and the settlement of northern North America, as well as the develop- ment of recreational canoes and the preservation of canoe history in more recent times. A respected expert writes each chapter on the type of craft or historical context being discussed. This makes for inconsistent writing at times, but also gives the book an authority that would not have been achieved by a single author. I found that the technical details about the dimen- sions (i.e., width and length, etc.) of canoes a bit repet- itive. On the other hand, the documentation of these measurements may be valuable in the future. As John Jennings notes in the first chapter, the original canoe designs of several First Nations in North America have been erased without a trace, a poignant reminder of the terrible loss of traditional knowledge that contin- ues to this day. While the book brilliantly covers the historical context of canoeing and canoe building (over the span of 230 pages), it pays scant attention to modern uses of the canoe (2 pages). After all, as the title sug- gests, canoeing and canoe building is a “living tradi- THE CANADIAN FIELD-NATURALIST Vol. 118 tion’. Canoeing is still one of the most elegant ways to explore nature and there are many fine canoe builders in North America currently manufacturing modern and traditional canoes from a wide range of natural and synthetic materials. It seems as though a chapter could have been devoted to recent developments in the design and construction of modern canoes. The book gives a good overview of how birch bark and dugout canoes are constructed, including useful photographic illustrations; however, before you run out and start assembling materials, take heed: the information provided is insufficient for the “do it your- self’ canoe builder. A noticeable shortcoming is the omission of information about the many resources available to those interested in building traditional birch bark and dugout canoes, as well as skin kayaks. For instance, while the book contains a photograph of a canoe built by César Nawashish, it fails to refer- ence the excellent 1971 National Film Board film “César’s Bark Canoe”, which documents Nawashish’s completion of a bark canoe with exquisite detail. Other contemporary builders of traditional craft have also produced thorough “how to” publications on building birch bark canoes and skin kayaks including David Gidmark, Robert Morris and Wolfgang Brink, which have not been referenced in this book. While Adney and Chapelle’s Bark Canoes and Skin Boats of North America will remain the bible of traditional canoe and kayak design, The Canoe: A Living Tradition is broader (literally and figuratively), more colourful and more accessible. It is simply irre- sistible for the canoe enthusiast. Its shortcomings are few and its design and production qualities are excel- lent. In summary, it is a valuable addition to the annals of canoe culture. Congratulations to John Jennings and the Canadian Canoe Museum for their fine work. PATRICK WILLISTON 41 Nielson Road, Smithers, British Columbia VOJ 2N2 Canada A Passion for Wildlife: The History of the Canadian Wildlife Service By J. A. Burnett. UBC Press, Vancouver, British Columbia. Hardcover. 331 pages, $27.95 paper, $85.00 cloth. Anyone who is prejudiced against “civil servants” should read this book. The Canadian Wildlife Service, from its inception, has contained a cadre of dedicated scientists who work long hours, share their knowledge widely, and on occasion risk their lives for the sake of the environment. Burnett, an insider, tells the story of this valuable and generally cost-effective organiza- tion, beginning in 1916 with the five men who were involved in drafting and implementing, from the Canadian side, the Migratory Birds Convention Act. In 1918, Hoyes Lloyd was appointed as Supervisor of Wild Life Protection in Canada. Lloyd, in turn, appointed three Federal Migratory Bird Officers. The Dominion Wildlife Service (now the Canadian Wild- life Service or CWS) was founded on | November 1947. Burnett follows a roughly chronological sequence, with 67 photographs. His writing skills introduce us to many interesting scientists and their achievements, without the inevitable plethora of names becoming objectionable. The footnotes list the main publica- tions that resulted from extensive research. He covers objectively the lowest point in CWS history, the short but destructive reign of Suzanne Blais-Grenier as federal Minister of the Environment. 2004 This excellent history will be of interest to most ornithologists, mammalogists and environmentalists, and will be a useful reference in every major library. However, I must admit to being flabbergasted by the publication of this book by a reputable university press, without any mention, let alone acknowledge- ment, of the previous publication, with the identical title, and with 99 per cent of the text which filled one issue of The Canadian Field-Naturalist 113:1-214, January-March 1999. Readers of this journal do not need to buy the book; they already have a soft-cover version on their shelves. What does the new version offer as compared to the unmentioned and unacknowledged version? An appropriate foreword by Janet Foster and a few new photographs have been added, but most are unchanged except that their quality has been improved and some have increased in size. Additions have been made to the following sub-chapters: “Seabirds,” a paragraph about Tony Gaston’s work on the Ancient Murrelet; “Shorebirds,” two paragraphs on the Latin American cooperation in shorebird banding and mention of the Kees Vermeer Award now presented annually at Simon Fraser University; “Habitat Management,’ a paragraph on the purchase of Westham Island in the Fraser River delta; “Consolidation,” two new paragraphs on wildlife policy changes; “Peregrine Falcon,” Richard Fyfe’s investiture into the Order of Canada on 26 April 2000; “Species Protection,’ a paragraph on the delays and rocky road of Bill C-5, the Canada Species at Risk Act, which finally passed on 11 June 2002; “A Work in Progress,” four sentences of reminiscences Birding on Borrowed Time By Phoebe Snetsinger. 2003. ABA Inc. Box 6599, Colorado Springs, Colorado. 307 pages. Paper US$19.19. I never met Phoebe Snetsinger. In fact I have only a vague recollection of hearing her name. I was sur- prised at her sudden prominence after she died. Now I have read her book and I understand. I expected someone who saw over 8000 species to be the typical “combat “ birder; a person who relishes getting the tick on her checklist above everything else. Phoebe Snetsinger was not a combat birder; she was a true birdwatcher. She insisted on really seeing and recording the birds she counted. She did not in- clude those flits that zip into view for an instant and are identified by the local guide. I have seen combat birders joyfully tick off such birds on their list and immediately move rapidly on in search of their next “lifer.” Indeed the author did not seem to be truly competitive until after her 7000" bird species and in line for the Guinness Book of Records. She ceased being competitive after achieving the world record at her 8000" species of bird. Her book is the account of her travels in accumu- lating this incredible total, and the joy she had in see- BOOK REVIEWS 29] by Rob Butler. The title of Chapter 5 has been im- proved from “Policy Implementation” to “Emergence of Environment Canada.” A few small editorial im- provements are evident, such as a change from “at the same time that” to “even as.” Otherwise the two ver- sions are almost word for word and paragraph for para- graph in the text and word for word in the index. Sadly, an opportunity to index each annual wildlife conference was not seized. What has been deleted? A short biography of the author, Sandy Burnett; a short afterword by editor, Francis Cook, explaining the contributions to editing of the Canadian Field-Naturalist made by Harrison F. Lewis, C. H. D. Clarke, and A. J. Erskine; and a useful 31-page bibliography of major publications by CWS personnel, compiled by Erskine. The Canadian Field-Naturalist exists to record and spread information of value, and contains a clear statement that copyright for this particular paper re- sides with “Her Majesty the Queen in Right of Canada” as stated in 113(1). However, I remain surprised that the author and the publisher appear, consciously or unconsciously, to have chosen to withhold information about this material having previously appeared in the very scientific journal: in which this review is now appearing. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Canada 3 Copies of CFN 113(1) are available for $10.00 from the Business Manager. ing all of these birds. She was a focused, energetic and hardworking individual who took her birdwatching extremely seriously. She studied her target species long before trying to see them. She kept copious notes on what she saw. I agree with this approach and try to use it myself. I do not understand why people pay so much money and effort to go to a prime nature locale, yet do not do their homework. Spurred into action by the threat of cancer and the potential to die young, Phoebe Snetsinger began a life of traveling for birds. In the process she made numer- ous friends in over 60 countries. I estimate she did ten to a dozen trips a year and spent about U.S.$50 000/ year in today’s currency. Overall, she must have made hundreds of trips. A neat bonus is the group of 29 black-and-white and 16 colour illustrations by Douglas Pratt. They por- tray the key milestone birds of Phoebe’s impressive odyssey. They are well done and make a nice coun- terpoint to the text. This is certainly a book for all birders. However, it is not simply a long list of birds seen, but a rampag- ing account of Phoebe’s obsession. This lady survived 297 gang rape, robbery, accidents, family crises, mechani- cal breakdowns, bad service and bad food against a background threat of cancer. Nothing seemed to slow her down. It is a tale of the fortitude of the human spirit. Indeed, I think a revision of this with the birding- related material severely edited (after all, which non- birder will realize the importance of a Red-shouldered Vanga — a small, House Sparrow-sized grey, black- and-white bird with bright chestnut shoulders — and New TITLES Zoology +The American Bison. By D. Lott. 2004. University of California Press. 229 pages, $40 U.S. Paper. + Annotated Bibliography of the Quaternary Vertebrates of Northern North America. 2003. Edited by C. R. Harring- ton. University of Toronto Press. xxii plus 539 pages, not illustrated. $150 Cloth, $75. Paper. Annotated Checklist of the Birds of Chile. By Manuel Marin. Lynx Edicions, Barcelona, Spain. US$15.75. Behavior and Ecology of Pacific Salmon and Trout. By T. Quinn. UBC Press, 2029 West Mall, Vancouver, British Columbia V6T 1Z2 Canada. 400 pages, $85. Big Cat Diary: Leopard. By Jonathan Scott and Angela Scott. Harper Collins. 128 pages, $42. * The Bird Almanac. By D. Bird. 2004. Key Porter Books, 70 The Esplanade, Toronto, Ontario, Canada. xvii + 460 pages, not illustrated, $24.95. Paper. Birding in Venezuela. By Mary Lou Goodwin. Lynx Edi- cions, Barcelona, Spain. US$ 5.60. Birds of South Asia: The Ripley Guide. Volumes I and II. By Pamela C. Rasmussen and John C. Anderton. Lynx Edi- cions, Barcelona, Spain. US$95. Bowerbirds — Ptilonorhychidae. Edited by Clifford Frith. Oxford University Press, 70 Wynford Drive, Don Mills, Ontario M3C 1J9 Canada. 416 pages. $209 Cloth. + British Columbia - A Natural History. By R. and S. Cannings. 2004. Greystone Books, Douglas and McIntyre, Vancouver, British Columbia, Canada. ix + 341 pages. $39.95 Paper. + The Buffalo Wolf. 2003. By Lu Carbyn. Smithsonian books, Washington, D.C., USA. 248 pages. Paper. Curassows and Related Birds. By Jean Delacour and Dean Amadon. Update Chapter by Josep del Hoyo and Anna Motis. Illustrated by Albert Earl Gilbert. Lynx Edicions, Barcelona, Spain. US$75. 7 Ecology and Conservation of Birds in the Salton Sink: an Endangered Ecosystem. By W. David Shuford and Kathy C. Molina. 2004. Studies in Avian Biology Number 27. Cooper Ornithological Society. THE CANADIAN FIELD-NATURALIST Vol. 118 its ilk?) would make it ideal for one of those inspira- tional stories we find in Reader’s Digest. Having final- ly seen a Red-shouldered Vanga, Phoebe was asleep in the tour bus when the bus crashed and she was killed. So she did not die of cancer, but binoculars in hand, on a birding trip. I never did meet this lady — pity! Roy JOHN 2193 Emard Crescent, Ottawa, Ontario K1J 6K5 Canada * A Field Guide to the Bird’s Nests and Eggs of Alaska Coastal Tundra. By T. Bowman. 2004. Alaska Sea Grant College Program. + A Guide to Hawk Watching in NA. By D. Heintzleman. 2004. The Globe Pequot Press. + Mammal Tracks and Signs. By Mark Elbroch. 2004. Stackpole Books. + Key Marine Habitat Sites for Migratory Birds in Nuna- vut and the Northwest Territories. 2004. By M. Mallory and A. Fontaine. 2004. Canadian Wildlife Service, Ottawa, Ontario K1A OH3. Paper. 7 Orca — Visions of the Killer Whale. By P. Knudtson. 2004. Greystone Books, Vancouver, British Columbia. xvii +110 pages. $19.95. Paper. Readers Digest Birds of Canada. Edited by R. Greenberg et al. 2004. Dorling Kindersly India Ltd., c/o Tourmaline Editions, 662 King Street West, Suite 304, Toronto, Ontario MSV 1M7. Reptiles and Amphibians of the Amazon: An Ecotourist’s Guide. By R. D. Bartlett and P. Bartlett Florida University Presses, USA. 291 pages. $57 Canadian. Paper. Wild Down Under: The Natural History of Australasia. By Neil Nightingale, Jeni Clevers, Neil Pearson and Mary Summerill. BBC Video. 240 pages, $50 Canadian. Cloth. Botany The Wild Flowers of Britain and Ireland — The Complete Guide to the British and Irish Flora. By Marjorie Blamey, Richard Fitter and Alastair Fitter. A & C Black. 512 pages, $42 Canadian. Paper. Environment *Bull’s Eye — Unraveling the medical mystery of Lyme Disease. By Jonathon Edlow. 2004. Yale University Press, Box 209040, New Haven, Connecticut. + Canoeing, Kayaking & Hiking Temagami. By Hap Wil- son. 2004. The Boston Mills Press (Firefly Books), 132 Main Street, Erin, Ontario. 112 pages, $24.95. + Chicken Soup for the Fisherman’s Soul. By J. Canfield, M. Hansen, K. & Dahlynn McKowen. 2004. Health Com- munications Inc. Paper. 2004 Dictionary of Ecology. By Michael Allaby. 2004. Oxford University Press, 70 Wynford Drive, Don Mills, Ontario M3C 1J9. 448 pages, $29.95. + The Importance of Species. Edited by P. Karliva and S. Levin. 2004. Princeton University Press, 41 William Street, Princeton, New Jersey. xvi + 427 pages, not illustrated, Paper. + Lewis and Clarke on the Great Plains. By Paul Johns- gard. 2003. University of Nebraska Press. xiv + 143 pages, $14.95 U.S. + Missinaibi. By Hap Wilson. 2004. The Boston Mills Press (Firefly Books), 132 Main Street, Erin, Ontario. 134 pages, $24.95. + A place between the tides: A naturalist’s reflections on the salt marsh. By Harry Thurston. 2004. Greystone Books. Paper. $22.95 CAAD, $15.00 U.S. + Politics of the Wild - Canada & Endangered Species. Edited by K. Beazley and R. Boardman. 2004. Oxford Uni- versity Press, 70 Wynford Drive, Don Mills, Ontario. x + 254 pages, not illustrated, $29.95 Paper. Population Ecology: First Principles. By John H. Vander- meer and Deborah E. Goldberg. Princeton UP, USA. 280 pages, $123 Cdn, Cloth. A Primer of Ecological Genetics. By J. Conner and D. Hartt. 2004. Sinauer Associates, Box 407, Sunderland, Massachu- setts. 207 pages, $34.95 U.S., Paper. BOOK REVIEWS 293 * Resource and Environmental Management in Canada. Edited by B. Mitchell. 2004. Oxford University Press, 70 Wynford Drive, Don Mills, Ontario. x + 600 pages, $29.95 Paper. + Rivers of the Upper Ottawa Valley. By Hap Wilson. 2004. The Boston Mills Press (Firefly Books), 132 Main Street, Erin, Ontario. 112 pages, $24.95. + The Selbourne Pioneer. By T. Dodswell. 2004. Ashgate Publising Company, Suite 420, 101 Cherry St., Burlington, Vermont, USA. xvii + 238 pages, not illustrated. Paper. + Shaped by the West Wind — Nature and History in Georgian Bay. By C. Campbell. 2004. UBC Press, 2029 West Mall, Vancouver, British Columbia, V6T 172 Canada. 256 pages, $85 Cdn. The Smithsonian Atlas of the Amazon. By Michael Gould- ing, Ronaldo Barthem and Efrem Ferreira. Smithsonian Books, Washington, D.C., USA. 255 pages, $75 Cdn. Cloth. + Teaching in Eden. 2003. By J. Janovy Jr. Routledge, 29 West 35th St., New York, New York 10001 USA. 187 pages, not illustrated, $22.95 U.S., Paper. + Wilderness Rivers of Manitoba. By Hap Wilson and S. Aykroyd. 2004. The Boston Mills Press (Firefly Books), 132 Main Street, Erin, Ontario, Canada. 112 pages, $24.95. + Available * Assigned News and Comment Update on Quebec Amphibian and Reptile Atlas Project: Rana-Saura 6(2) The newsletter Rana-Saura resumes with Volume 6, Num- ber 2 issued for April 2004. The previous issue was published in March 2000. Due to translation expense no English ver- sion is currently available of the new issue (of 383 participants, only 23 are anglophones). Contents are: Et ¢a continue! — Atlas des amphibiens et des reptiles du Québec — Suivi des populations d’amphibiens du Québec — La présence de la Rai- nette faux-grillon boreale [Pseudarcris maculata] enfin con- firmée [Jean-Francois Desroches] — Un nouvelle reptile pour Québec! [Jean-Fran¢ois Desroches] — Le tout nouveau guide d’identification des amphibiens et des reptiles du Québec! Rana-Saura is edited by David Rodrigue, for the Atlas of amphibians and reptiles of Quebec (continuing in 2004 in its 17" year; since 2002 the entries have increased from 22 000 to nearly 45 000) and the Survey of populations of amphibians of Quebec (in 2004 there will be 45 participants conducting 42 road surveys). These projects are coordinated by the Eco- musuem of the St. Lawrence Valley Natural History Society and financially aided by La Sociéte de la Faune et des Parcs du Québec. It is distributed by St. Lawrence Valley Natural History Society, 21-125 chemin Sainte-Marie, Sainte-Anne-de-Belle- vue, Quebec H9X 3Y7 Canada. Phone (514) 457-9449, extension 105: Fax (514) 457-0769; e-mail: ecomus @total.net. Froglog: Newsletter of the Declining Amphibian Populations Task Force (62) Number 62, April 2004. Contents: Southern African Frog Atlas Project completed! (Marius Burger and J. A. Harrison) — Mark Twain’s frog not croaked after all! (Robert Stack) — Decline of the Kihansi Spray Toad, Nectophrynoides asper- ginis from the Udzungwa Mountains, Tanzania (Che Weldon & Louis H. du Preez) — The Amphibian Fauna of Nagaland, India: Species and Habitats (Meren Ao & Sabitry Bordoloi) — Seed Grant News — Froglog Shorts. Froglog is the bi-monthly newsletter of the Declining Am- phibian 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 Arizona State 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 Zoo- logical Institute, P. O. Box 39, Royal Oak, Michigan 48068- 0039, USA. Publication of issue 62 was also supported by Peace Frogs www.peacefrogs.com and by RANA and the U.S. National Science Foundation grant DEB-0130273. Amphipacifica: Journal of Aquatic Systematic Biology 3(4) Volume 3, Number 4, dated 30 March 2004, is 113 pages and contains the talitroidean amphipod family Najnidae in the North Pacific region: systematics and distributional ecol- ogy (E. L. Bousfield and Pierre Marcoux) and The amphi- pod family Pleustidae (mainly subfamilies Mesopleustinae. Neopleustinae, and Stenopleustinae) from the Pacific coast of North America: systematics and distributional ecology Marine Turtle Newsletter (104) April 2004. 36 pages: ARTICLES: Nesting of Hawksbill Turtles in Paraiba-Brazil: Avoiding Light Pollution Effects — Stranding of Small Juvenile Leatherback Turtle in West- ern Australia — Post-nesting Movements of Green Turtles Tagged in the Turtle Islands, Tawi-Tawi, Philippines — Notes from Preliminary Market Surveys in Morocco — NOTES: Recoveries of Two Post-hatching Loggerhead Turtles in the Northern Adriatic Sea — Green Turtle Nesting on the Gulf of Oman Coastline of the Islamic Republic of Iran — Confirmed Nesting of the Loggerhead Turtle in Corsica — Marine Tur- tles in Iran: Results from 2002 — Sightings of the Leatherback Turtle off the Southern Coast of Rio de Janeiro, Brazil — (E. A. Hendrycks and E. L. Bousfield). Amphipacifica (ISSN 1189-9905) is published quarterly by Amphipacifica Research Publications, Edward L. Bousfield, Managing Editor, 1710-1275 Richmond Road, Ottawa, On- tario K2B 8E3 Canada. Annual subscription rates are $40.00 US or $50.00 Canadian funds. Back numbers are available at $10 each. MEETING REPORTS — IUCN/SSC MTSG News — BOOK REVIEWS — ANNOUNCEMENTS NEWS & LEGAL BRIEFS — RECENT PUBLI- CATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, School of Biological Sciences, University of Exeter, Exeter EX4 4PS United Kingdom; e-mail MTN @seaturtle. org; Fax +44 1392 263700. Subscriptions and donations to- wards the production of both the MTN and its Spanish edi- tion NTM [Noticiero de Tortugas Marinas] should be made online at www.seaturtle.org/mtn or c/o SEATURTLE.ORG 11400 Classical Lane, Silver Spring, Maryland 20901 USA. 294 2004 THE CANADIAN FIELD-NATURALIST 295 Erratum 118(1): 80 Sloan, N. A., and P. M. Garier. 2004. Introduced marine species in the Haida Gwaii (Queen Charlotte Islands) Region, British Columbia. Canadian Field-Naturalist 118(1): 77-84. Langara Island 7 Parry Passage Cryptosula pallasiana He Rose Limnodrilus monothecus | Masset Spit / Delkatla / ; / ® Pelonaia corrugata / Inlet Heh Ve ~ Trechus obtusus Limnodrilus monothecus om Cryptosula pallasiana . a a x \ xe Kumdis Bay \ Yakoun Bay Graham Rennell Sound o Island : Hecate Skidegate he Inlet A Moresby ‘ Island Cartwright Sound Tasu Sound Bowerbankia gracilis Hotspring Ciona intestinalis Island Legend STUNG a a Sargassum muticum Dolomite "Burnaby" Narrows O Mya arenaria Qo oe A Harriet © Ampithoe vallida @ Other invertebrate sightings as noted B Harbour pl ceeeen eels at 5a Bowerbankia gracilis ~~ — nt Houston Stewart Channel Schizoporella unicornis ® Tubularia crocea/ 131°W FicureE |. Map of Haida Gwaii showing locations mentioned in the text and collection sites of introduced marine plant and invertebrate species, based on data from Sloan and Bartier (2000) and Sloan et al. (2001). Editor’s Report for Volume 117 (2003) Mailing dates for issues in volume 117 were: (1) 30 September 2003, (2) 23 March 2004, (3) 12 July 2004, (4) 16 September 2004. A summary of membership and subscriber totals 2003 is given in Table 1. The number of articles and notes in volume 117 is summarized in Table 2 by topic; totals for Book Reviews and New Titles are given in Table 3, and the distribution of content by page totals per issue in Table 4. Council continued contribution of 40% of membership dues for publication. All of sub- scriptions (both individual and institutional) also go toward publication. As well, the Council allocated 80% of the annual interest from the Manning Fund and other capital funds to The Canadian Field-Naturalist. The Manning fund portion is specifically to offset the publication cost of northern papers where author and institutional contributions were insufficient to cover page charges. Emil Holst arranged setting and printing at St. Joseph Print Group, Ottawa, for 117(1), with special thanks due Cameron Fraser and Willy Silfwerbrand, and at Gilmore Printers, Ottawa, for 117 (2,3,4) with special thanks to Wendy Cotie and Ally Reckzin. Leslie Cody prepared the Index, Elizabeth Morton read the galleys. Bill Cody as Business Manager handled all reprint requests and charges and oversaw and proofed the compilation of the Index. Wilson Eedy continued as Book-Review Editor until his death in June, and Roy John replaced him in August 2004. Manuscripts (excluding book reviews, notices, and reports) submitted to The Canadian Field-Naturalist totalled 84 in 2003, down 16 from 100 in 2002. The following reviewed for papers submitted in 2003: Asso- ciate Editors: (number of manuscripts reviewed in par- entheses): R. Anderson, Canadian Museum of Nature, Ottawa (5): C. D. Bird, Erskine, Alberta (14); B. W. TABLE 2. Number of articles and notes published in The Cana- dian Field-Naturalist Volume 117 (2003) by major field of study. Subject Articles Notes Total Mammals 19 * 14 33 Birds Lie 6 23 Amphibians + reptiles 4 1 BD) Fish 3 D. 5 Invertebrates 6 1 6 Plants** 16 3 19 Tributes 2 0 2 Totals 67 27 93 *one article counted twice, one in each group: **includes one feature article. Coad, Canadian Museum of Nature, Ottawa (10); R. R. Campbell, St. Albert, Ontario (6); P. M. Catling, Agriculture and Agri-food Canada, Ottawa (11); A. J. Erskine, CWS, Sackville, New Brunswick (13); D. F. McAlpine, New Brunswick Museum, Saint John, N.B. (8); D. W. Nagorsen, Victoria, British Columbia (11); W. O. Pruitt, Jr., University of Manitoba, Winnipeg (21); Others: (one review each except as marked): P. Achuff, Waterton National Park, Alberta; Y. Alarie, Laurentian University, Sudbury, Ontario; N. Alfonso, Canadian Museum of Nature, Ottawa, Ontario; J. F Bain, Uni- versity of Lethbridge, Alberta; R. Bird, Cascadia Re- search, Olympia, Washington; K. Bellamy, OMNR, Peterborough; J. R. Bider, Ecomuseum, Ste-Anne-de- Bellevue, Quebec (3); E. L. Bousfield, Ottawa, On- tario; J. P. Bogart, University of Guelph, Ontario; I. Brodo, Canadian Museum of Nature, Ottawa, Ontario (2); R. J. Brooks, University of Guelph, Ontario; J. Calambokidis, Cascadia Research, Olympia, Washing- ton; S. E. Campana, Bedford Institute of Oceanography, TABLE |. The 2003 circulation of The Canadian Field-Naturalist (2002 in parenthesis). 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 E, (OMI) 36) G2) 7 (8) 838 (951) Subscriptions Individuals 163 (164) 60 (65) 6 (5) 229° (234) Institutions lesa — is(GuZAl)) 241 (252) 307762) 428 (455) Totals 1005 (1246) 337 (349) 43 (45) 1495 (1640) 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. 296 2003 TABLE 3. Number of reviews and new titles published in Book Review section of The Canadian Field-Naturalist Volume 117 by topic. Reviews New Titles Zoology 40 30 Botany 11 23 Environment 22 4] Miscellaneous 12 18 Young Naturalists 0 18 Totals 85 130 Dartmouth, Nova Scotia; L. Carbyn, Canadian Wild- life Service, Edmonton, Alberta (4); A. Ceska, Victoria, British Columbia; A. H. Clarke, Jr., Portland, Texas; W. J. Cody, Agriculture and Agri-food Canada, Ottawa; V. Craig, EcoLogic Research, Gabriola Island, British Columbia; M. Crete, Ministere de 1’ Environnement et de la Faune, Quebec; E. J. Crossman, Royal Ontario Museum, Toronto, Ontario; S. J. Darbyshire, Agriculture and Agri-food Canada, Ottawa; R. Day, Canadian Museum of Nature, Ottawa; C. Ely, U. S. Fish and Wildlife Service, Anchorage, Alaska; M. Engstrom, Royal Ontario Museum, Toronto; B. Erickson, Salon. Ohio; M. Fairbarns, Conservation Data Centre, B. C. Government, Victoria; M. Gosselin, Canadian Museum of Nature, Ottawa; T. Goward, Clearwater, British Columbia; W. A. Fuller, Athabasca University, Alberta; D. Galbraith, Royal Botanical Gardens, Hamilton, Ontario; Marco Festa-Bianchet, University of Sher- brooke, Quebec; P. T. Gregory, University of Victoria, British Columbia (2); D. Gummer, Provincial Museum of Alberta, Edmonton (2); E. Haber, National Botani- cal Services, Ottawa; F. Harrington, Mt. St. Vincent University, Halifax, Nova Scotia; R. Harrington, Cana- dian Museum of Nature, Ottawa; D. Henry. Parks Canada, Haines Junction, Yukon Territory; S. Herrero, University of Calgary, Alberta; G. V. Hilderbrand, Alaska Department of Fish and Game, Anchorage; R. P. Hodge, Gig Harbor, Washington; C. S. Houston, Saskatoon, Saskatchewan (2); E. Holm, Royal Ontario Museum, Toronto; G. Jarrell, University of Alaska Museum, Fairbanks, Alaska; D. Johnson, Canadian Forest Service, Edmonton, Alberta; N. Jotham, NRJ Technical Services, Trade & Animal Welfare, Ottawa: R. B. King, Northern Illinois University, DeKalb, Illi- nois; J. L. Koprowski, Wildlife and Fisheries Resources, University of Arizona, Tucson; S. Lariviere, Delta Waterfowl Foundation, Portage La Prairie, Manitoba; D. Larson, Memorial University of Newfoundland, St. John’s; L. E. Licht, York University, North York, Ontario; J. Lien, Memorial University of Newfound- land, St. John’s; S. Lindgren, University of Northern British Columbia, Prince George; G. L. Mackie, Uni- versity of Guelph, Ontario; F. F. Mallory, Laurentian University, Sudbury, Ontario; A. Martel, Canadian EDITOR’S REPORT 297 TABLE 4. Number of pages per section published in The Canadian Field-Naturalist Volume 117 (2003) by issue. (1) (2) (3) (4) Total Articles WAI SB) 109 119 490 Notes 12 16 15 17 60 Tributes 1] 0 16 (0) Dil, Book Reviews’ 13 25 26 18 82 CFN/OFNC Reports x 0 2 8 212. News and Comment 3 2 | 2 8 Index 0) 0 0 30 30 Advice to Contributors 0 0 | 0 | Totals 166 180 176 188 710 *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). Museum of Nature, Ottawa; John Maunder, New- foundland Museum, St. John’s, Newfoundland; M. K. McNicholl, Burnaby, British Columbia; G. R. Miche- ner, University of Lethbridge, Alberta; G. Mulligan, Agriculture and Agri-food Canada, Ottawa; E. Nol, Trent University, Peterbourgh; P. Paquet, Meacham, Saskatchewan; K. Prior, Canadian Wildlife Service, Ottawa; G. Proulx, Alpha Research & Management Ltd., Sherwood Park, Alberta; C. Renaud, Canadian Museum of Nature, Ottawa, Ontario; B. Riddell, Paci- fic Biological Station, Nainimo, British Columbia; R. A. Ring, University of Victoria, British Columbia; James Schaefer, Trent University. Peterborough, On- tario; W. B. Schofield, University of British Columbia, Vancouver; F. W. Schueler, Oxford Station, Ontario; B. Slough, Whitehorse, Yukon Territory; I. Stirling, Canadian Wildlife Service, Edmonton, Alberta (2); J. Straley, University of Alaska Southeast, Sitka, Alaska; D. Strickland, Dwight, Ontario; E. S. Telfer, Edmon- ton, Alberta; I. Thompson, Canadian Forest Service, Sault Ste. Marie, Ontario; C. Traynor, Ottawa, Ontario; N. Vachon, Sociéte de la faune et des parcs du Québec, Longueuil; P. Weigl, Wake Forest University, Winston- Salem, North Carolina; G. Wobeser, Western College Veterinary Medicine, University of Saskatchewan, Sas- katoon; J. N. Womble, University of Alaska-Fairbanks, Juneau, Alaska; P. M. Youngman, Ottawa, Ontario (2). I am also indebted to the President of the Ottawa Field-Naturalists’ Club Gary McNulty and the Club Council for continuing support of the journal; Chair- man 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 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 read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, 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. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit, to the Editor, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. All authors should have read and approved it. Institutional or contract approval for the publication of the data must have been obtained by the authors. Also authors are ex- pected to have complied with all pertinent legislation regard- ing the study, disturbance, or collection of animals, plants or minerals. The place where voucher specimens have been de- posited, and their catalogue numbers, should be given. Lati- tude and longitude should be included for all individual local- ities where collections or observations have been made. Print the manuscript on standard-size paper, doublespace throughout, leave generous margins to allow for copy mark- ing, and number each page. For Articles and Notes provide a citation bibliographic strip, an abstract and a list of key words. Generally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manuscripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in the text or in a sepa- rate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “British” or “American” spellings are acceptable in English but should be consistent within one manuscript. 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If grant or institutional funds are not available, club members and subscribers may apply for a waiver of charges for the first five pages. Limited joumal funds are available to help offset publi- cation charges to authors with minimal financial resources. Requests for financial assistance should be made to the Busi- ness Manager when the manuscript is accepted. Reprints An order form for the purchase of repents will accompany the galley proofs sent to the authors. FRANCIS R. CooK, Editor | RR 3 North Augusta, Ontario KOG TRO Canada | 298 TABLE OF CONTENTS (concluded) Volume 118 Number 2 Notes Observations of-above surface littoral foraging in two sea ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in coastal southwestern British Columbia DEBORAH L. LACROIX, KENNETH G. WRIGHT, and DANIEL KEN1 New records of Cyperaceae and Juncaceae from the Yukon Territory STUART A. HARRIS and PETER W. BALL “Ashkui”, vernal ice-cover phenomena and their ecological role in southern Labrador SHAUNA M. BAILLIE, CORINNE D. WILKERSON, and TINA L. NEWBURY Sea Otter, Enydra lutris, sightings off Haida Gwaii/Queen Charlotte Islands, Canada, 1972-2002 KIMBERLY RAUM-SURYAN, KENNETH PITCHER, and RICHARD LAMY iFirst record of a Barred Owl, Strix varia, in Labrador ISABELLE SCHMELZER and FRANK PHILLIPS Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: a new addition to the flora of Quebec JACQUES BRISSON Evidence for the use of vocalization to coordinate the killing of a White-tailed Deer, Odocoileus virginianus, by Coyotes, Canis latrans ERICH M. MUNTZ and BRENT R. PATTERSON Book Reviews ZOoLoGy: Fish of Alberta — Sharks — In Search of the Golden Frog — Conversations with an Eagle — North American Owls, Biology and Natural History, Second Edition — The Mountain White-Crowned Sparrow: Migration and Reproduction at High Altitude — Seabird Bycatch: Trends, Roadblocks, and Solutions — The Life of Mammals — Geographic Variation in Size and Shape of Savannah Sparrows (Passerculus sandwichensis) — Warblers of the Great Lakes and Eastern North America — Sparrows and Finches of the Great Lakes and Eastern North America — Birds of the Yukon Territory Botany: Wild Flowers of the Yukon, Alaska & Northwestern Canada, 2™ edition — Wild Flowers of Field | & Slope in the Pacific Northwest — Wild Flowers of Forest & Woodland in the Pacific Northwest — Wild Flowers of the Mountains in the Pacific Northwest MISCELLANEOUS: The Canoe: A Living Tradition — A Passion for Wildlife: The History of the Canadian Wildlife Service — Birding on Borrowed Time New TITLES News and Comment Update on Quebec Amphibian and Reptile Atlas Project: Rana-Saura 6(2) — Froglog: Newsletter of the Declining Amphibian Populations Task Force (62) — Amphipacifica: Journal of Aquatic Systematic Biology 3(4) — Marine Turtle Newsletter (104) — Erratum: Sloan and Bartiers 118(1): 80 | Editor’s Report Volume 117 (2003) FRANCIS R. COOK | Advice for Contributors Mailing date of the previous issue 118(1): 15 December 2004 2004 264 266 THE CANADIAN FIELD-NATURALIST Volume 118 Number 2 2004 Articles Morphology and population characteristics of Vancouver Island Cougars, Puma concolor vancouverensis STEVEN F. WILSON, APRYL HAHN, AARON GLADDERS, | KAREN M. L. Gou, and Davip M. SHACKLETON 159 Conservation evaluation of Lemmon’s Holy Fern, Polystichum lemmonii, a threatened fern in Canada GEORGE W. DOUGLAS 164 Conservation evaluation of the Pacific population of Tall Woolly-heads, Psilocarphus elatior, an endangered herb in Canada GEORGE W. DOUGLAS and JEANNE M. ILLINGWORTH 169 Conservation evaluation of Howell’s Triteleia, Triteleia howellii, an endangered lily in Canada GEORGE W. DOUGLAS and JENIFER L. PENNY 174 Conservation evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada TYLER W. SMITH, GEORGE W. DOUGLAS, and ALLAN G. HARRIS 175) Immobilizatin of clover-trapped White-tailed Deer, Odocoileus virginianus, with medetomidine and ketamine, and antagonism with atipamezole JOSHUA J. MILLSPAUGH, BRIAN E. WASHBURN, TAMARA M. MEYER, JEFF BERINGER, and LONNIE P. HANSEN 185 Predation on two Mule Deer, Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the southern Canadian Rocky Mountains DorTE PosziG, CLAYTON D. Apps, and ALAN DIBB 191 Effect of fire intensity and depth of burn on Lowbush Blueberry, Vaccinium angustifolium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, production in eastern Ontario Luc C. DUCHESNE and SUSANNE WETZEL 195 Spring dispersal patterns of Red-winged Blackbirds, Agelaius phoeniceus, staging in eastern South Dakota H. J. HomMaAN, G. M. Linz, R. M. ENGEMAN, and L. B. PENRY 201 1f Changes in loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) populations in the Lower Matanuska-Susita Valley, Alaska TAMARA K. MILLS and BRAD A. ANDRES 210 The Barred Owl, Strix varia, in Alberta: distribution and status LisA TAKATS PRIESTLEY 215 Use of Eelgrass, Zostera marina, wrack by three species of ladybird beetles (Coleoptera: Coccinellidae) in Prince Edward Island DaviD J. GARBARY, SARAH FRASER, CARRIE FERGUSON, and RANDOLPH F. LAUFF 225 Predicting the effects of Cerulean Warbler, Dendroica cerulea, management on eastern Ontario bird species JASON JONES, WILLIAM J. MCLEISH, and RALEIGH J. ROBERTSON Ds Parasite prevalence in Dark-eyed Juncos, Junco hyemalis, breeding at different elevations i HEATHER BEARS 235 iF Movements of subadult male Grizzly Bears, Ursus arctos, in the central Canadian Arctic ROBERT J. GAU, PHILIP D. MCLOUGHLIN, RAY CASE, H. DEAN CLUFF, ROBERT MULDERS, and FRANCOIS MESSIER 239 Severe chronic neck injury caused by a snare in a Coyote, Canis latrans PIERRE- Y VES DaAousT and PETER H. NICHOLSON 243 Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Preserve, Alaska Tom S. SMITH, STEVEN T. PARTRIDGE, and JOHN W. SCHOEN 247 Wild Turkey, Meleagris gallopavo silvestris, behavior in central Ontario during winter LINH P. NGUYEN, JOSEPH HAMR, and GLENN H. PARKER 25 (continued on inside back cover, ISSN 0008-3550 | The CANADIAN — FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 118, Number 3 July-September 2004 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 information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining 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 2004 Council President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor Francis R. Cook Cendrine Huemer To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, 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 www.ofnc.ca 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 IRO; (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 KIZ 1A2; (613) 759-1374 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6K5, e-mail: roy.john@pwegsc.ge.ca 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 except Book Reviews should be addressed to the Editor and sent by postal mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, Book-review Editor. 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Date of this issue: July-September 2004 (September 2005). Cover: International zone on rugged rock headland of the Queen Charlotte Islands, British Columbia, each dominant species cre- ating a distinct zone. The white lichen is mainly Coccotrema maritimum and the black one is almost entirely Verru cana maura. Below the Verru cana is a zone bare of both lichens and marine algae, and, still lower, the algae take over. Figure 64 (page 77). Lichens of North America. By I. M. Brodo, S. D. Sharnoff, and S. Sharnoff. 2001. Yale University Press, New Haven, Connecticutt. See article by Brodo and Sloan on lichen zonatum pages 405-424. | | | | | Volume 118, Number 3 The Canadian Field-Naturalist vA. July—September 2004 Population Dynamics of Deer Mice, Peromyscus maniculatus, and Yellow-pine Chipmunks, Tamias amoenus, in Old Field and Orchard Habitats THOMAS P. SULLIVAN!, DRUSCILLA S. SULLIVAN?, and EUGENE J. HOGUE? 'Agroecology Program, Faculty of Land and Food Systems, University of British Columbia, Vancouver, British Columbia V6T 1Z4 Canada (corresponding author). ? Applied Mammal Research Institute, 11010 Mitchell Avenue, Summerland, British Columbia VOH 1Z8 Canada 3 Pacific Agri-Food Research Centre, Agriculture and Agri-Food Canada, 4200 Highway 97, Summerland, British Columbia V1X 7S3 Canada Sullivan, Thomas P., Druscilla S. Sullivan, and Eugene J. Hogue. 2004. Population dynamics of Deer Mice, Peromyscus mani- culatus, and Yellow-pine Chipmunks, Tamias amoenus, in old field and orchard habitats. Canadian Field-Naturalist 118(3): 299-308. There are often several rodent species included in the small mammal communities in orchard agro-ecosystems. This study was designed to test the hypothesis that the population levels of Deer Mice (Peromyscus maniculatus) and Yellow-pine Chipmunks (Tamias amoenus) would be enhanced in old field compared with orchard habitats. Rodent populations were intensively live- trapped in replicate old field and orchard sites over a four-year period at Summerland, British Columbia, Canada. Deer Mouse populations were, on average, significantly higher (2.5 — 3.4 times) in the old field than orchard sites in summer and winter periods. Mean numbers/ha of Deer Mice ranged from 12.1 to 60.4 in old field sites and from 3.3 to 19.9 in orchard sites. Breeding seasons in orchards were significantly longer than those in old field sites, in terms of proportion of reproductive male Deer Mice. Recruitment of new animals and early juvenile survival of Deer Mice were similar in orchard and old field sites. Populations of Yellow-pine Chipmunks ranged in mean abundance/ha from 5.6 — 19.0 in old field sites and from 1.9 — 17.5 on one orchard site, with no difference in mean abundance in 2 of 4 years of the study. Recruitment and mean survival of Yellow-pine Chipmunks also followed this pattern. This study is the first detailed comparison of the population dynamics of these rodent species in old field and orchard habitats. These species should be able to maintain their population levels and help contribute to a diversity of small mammals in this agrarian landscape. Key Words: Deer Mouse, Peromyscus maniculatus, Yellow-pine Chipmunk, Tamias amoenus, abundance, demography, old field, orchard, population dynamics, recruitment, reproduction, British Columbia. Voles of the genus Microtus are often the major rodent species in perennial grasslands and agro-eco- systems. In addition to grasses and herbs, these micro- tines feed on vascular tissues of tree and other crop plants primarily during winter months (Byers 1985; Lewis and O’Brien 1990). Other major rodent species include the Deer Mouse (Peromyscus maniculatus) and Yellow-pine Chipmunk (TZamias amoenus), which may occur in tree fruit orchards and adjacent habitats in the inland regions of the Pacific Northwest of North America. Neither of these associated species is known to damage orchard trees by their feeding activity as they are granivorous or insectivorous in their feeding habits (Baker 1968; Webster and Jones 1982; Sutton 1992). Deer Mice or White-footed Mice (P. leucopus) com- monly occur with Microtus in old fields and other perennial grassland habitats (Tamarin 1977; Krebs 1979; Dueser et al. 1981; Sullivan and Krebs 1981; Schweiger et al. 2000; Manson et al. 2001; Pearson et al. 2001) and in orchard agro-ecosystems (Sullivan et al. 1998). Few studies of voles in orchards have report- ed on the population dynamics of other species in the small mammal community. Sullivan et al. (1998) dis- cussed the population responses of all species to vege- tation management in apple (Malus domestica) orchards and Sullivan et al. (2000) reported changes in species diversity of these communities in orchards and old fields. Orchard habitats usually have frequent mowing and vegetation management treatments during summer months, and hence conditions for small mammals are more changeable than in old fields. Thus, this study was designed to test the hypothesis that the popula- tion dynamics of Deer Mice and Yellow-pine Chip- munks would be enhanced in old field compared with orchard habitats. To evaluate this hypothesis, we pro- vide a detailed analysis of the population dynamics of these species in the two habitats. 299 300 Materials and Methods Study area and experimental design This study was conducted at the Pacific Agri-Food Research Centre in the Okanagan Valley, Summerland, British Columbia, Canada. The experimental design consisted of two replicate “old field” and two repli- cate orchard habitats. The old field habitats were abandoned (225 years) hay fields composed of Crest- ed Wheatgrass (Agropyron cristatum), Quack Grass (Agropyron repens), Downy Brome (Bromus tecto- rum), Diffuse Knapweed (Centaurea diffusa), with some minor herbaceous species such as Yellow Salsify (Tragopogon dubius), Great Mullein (Verbascum thapsus), American Vetch (Vicia americana), Prickly Lettuce (Lactuca serriola), and Tall Tumble-mustard (Sisymbrium altissimum). These old field sites were each 2 to 3 ha in area within a mosaic of sagebrush (Artemisia tridentata), Ponderosa Pine (Pinus pon- derosa) forest, and orchard habitats. These old fields had resident populations of Deer Mice and Yellow- pine Chipmunks. Other species included the Montane Vole (Microtus montanus), Great Basin Pocket Mouse (Perognathus parvus), Western Harvest Mouse (Reith- rodontomys megalotis), and a few Long-tailed Voles (M. longicaudus). The orchards were: (A) a five-year-old apple orchard unit, and (B) a 10-year-old apple orchard combined with a 15-year-old pear (Pyrus sp.) orchard as one unit. Both 1.2-ha orchards were located within a 90- ha mosaic of tree fruits and vineyards. Thus, our ex- perimental design had two true replicates of old field sites and two replicates of orchard sites. Each pair of old field and orchard sites was spatially segregated to enhance statistical independence (Hurlbert 1984). A third replicate pair would have strengthened the study but was not possible within the operational setting of the Research Centre. Common grass species on the orchard sites included Orchard Grass (Dactylis glomerata), Quack Grass, bluegrass (Poa spp.), Smooth Brome (Bromus inermis), and Crested Wheatgrass. These orchards were mowed five or six times each summer. Rodenticides were applied 3-4 times each winter in poison-bait feeder stations (Radvanyi 1974) for voles (Mouse Bait IT® — zinc phosphide and Ramik Brown® — diphacinone). Rodenticides were not present in the old field sites. Deer Mouse and Yellow-pine Chipmunk populations All animals were live-trapped on 1-ha grids with 49 (7 x 7) trap stations located at 14.3-m intervals with one or two Longworth live-traps at each station. One of the old-field grids was an irregular shaped rectangle of 1 ha with the same 49 stations. The four grids (2 orchard and 2 old field) were live-trapped at 3-week (spring, summer, and fall) and at 4- to 6-week (winter) intervals from June 1982 to April 1986. Traps were baited with whole oats, peanut butter, and carrot; coarse brown cotton was supplied as bed- THE CANADIAN FIELD-NATURALIST Vol. 118 ding. Traps were set on day 1, checked on the morning and afternoon of day 2 and morning of day 3, and then locked open between trapping periods. All animals captured were ear-tagged with serially numbered tags, breeding condition noted, weighed on Pesola spring balances, and point of capture recorded. The duration of the breeding season was noted by palpation of male testes and the condition of mammaries of the females (Krebs et al. 1969). A pregnancy was considered suc- cessful if a female was lactating during the period following the estimated time of birth of a litter. Ani- mals were released on the grids immediately after processing. Seasons were defined as summer (April to Septem- ber) and winter (October to March) periods. Thus, there were four summer and four winter periods from 1982 to 1986. We used mass at sexual maturity to infer age classes of animals. Body mass was used as an index of age. The percentage of sexually mature animals was used to determine the mass limitations for juveniles and adults assuming that juveniles were seldom, if ever, sexually mature, and that at least 50% of the adults were sexually mature in their lowest mass class. Deer Mice (juvenile = 1 — 20 g, adult > 21 g) and Yellow-pine Chipmunks (juvenile = 1 — 44 g, adult 2 45 g) were classified as juvenile or adult by body mass. Juveniles were considered to be young animals recruited during the study. Recruits were defined as new animals that entered the population through repro- duction and immigration. All handling of animals was in accordance with the principles of the Animal Care Committee, University of British Columbia. Demographic parameters Population densities were estimated by the Jolly- Seber model for reasons indicated by Jolly and Dick- son (1983). The Jolly-Seber (J-S) model provides the best.estimates of population size for mark and recap- ture data when trappability values are generally < 70% (Hilborn et al. 1976). However, when population size falls very low and no marked animals are recaptured, the J-S estimate becomes unreliable and impossible to calculate (Krebs et al. 1986). For these sample weeks, a minimum number of animals known to be alive (MNA) (Krebs 1966) value was substituted for a bio- logically unreasonable J-S estimate. Measurements of recruitment, number of lactating females, and early juvenile survival were derived from the sample of animals captured in each trapping ses- sion and then summed for summer periods. Early juve- nile survival is an index relating recruitment of young into the trappable population to the number of lactat- ing females (Krebs 1966). A modified version of this index is number of juvenile animals at week ft divided by the number of lactating females caught in week t — 3. Mean survival rates (28-day) for summer and winter periods were estimated from the Jolly-Seber model. Mean body mass of combined males and females 2004 SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 30] Peromyscus maniculatus Number of animals/na NO (Se) A on oO) (eo) (ep) (ep) (oe) (=) = (eo) --Old field 1 Old field 2 Orchard 1 Orchard 2 J SNJIMMJSNJIMMJISNJIMM JSNJIMM 1982 1983 1984 1985 1986 FiGurRE |. Population densities (Jolly-Seber) per ha of Deer Mice in replicate old field and orchard sites during the study. Shaded bars indicate winter periods. Months of year are represented by J = July; S = September; N = November; J January; M = March; M = May. was used as an index of condition within populations of Deer Mice and Yellow-pine Chipmunks during sum- mer and winter periods. Statistical analysis Mean trappability, mean abundance, mean number of recruits, mean Jolly-Seber survival rates, and mean body mass were evaluated by 95% confidence inter- vals (CI) for Deer Mice and Yellow-pine Chipmunks in old field and orchard sites during summer and winter periods, as per the recommendations of Gerard et al. (1998) and Johnson (1999). Proportion of adult males and adult females breeding was analyzed by a Chi- square 2 x 2 contingency table (Zar 1984) for each of the four summer periods. These datasets of proportion of animals breeding often include animals captured more than once, and hence they are not completely independent. Thus, the Chi-square analyses provide only an indication of the degree of difference between datasets. In all analyses, the level of significance was E1053 Results Deer Mouse populations Totals of 751 and 459 individual Deer Mice were captured on the two old field and two orchard sites, respectively. Mean J-S trappability estimates for Deer Mice ranged from 62.6% to 73.8% in summer and from 46.6% to 76.6% in winter in the old field sites. These estimates in the orchard sites ranged from 48.0% to 81.7% in summer and from 39.0% to 80.8% in winter. Deer Mouse populations were, on average, general- ly higher (2.5 — 3.4 times) in the old field than orchard sites in summer and winter periods (Figure 1). There were some exceptions to this pattern in summers 1982 and 1985 and winter 1985-1986 when mean abun- dance of Deer Mice was similar, at least in one old field—orchard site comparison (Table 1). Numbers of Deer Mice reached annual autumn peaks ranging from 30.8 to 69.3 animals/ha in the old field-1 site and from 57.2 to 67.5 animals/ha in the old field-2 site. 302 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Mean abundance per ha + 95% confidence intervals for Deer Mice during summer and winter periods in replicate old field and orchard sites. Sample size (n = number of trapping periods) in parentheses. Old field Orchard Period 1 2 1 2 Summer 1982 ye LES BikD 15.2 12.6 (7) 95% CI 11.2 — 23.8 22, 7.7 —22.7 9.3 — 16.0 Winter 1982-83 NG 25.8 49.0 16.8 6.0 (5) 95% CI 21.9 — 29.7 43.0 —55.] 10.6 — 23.0 2.6 —9.4 Summer 1983 Ve 41.9 55.6 15:5 19.9 (9) 95% CI 36.2 — 47.6 50.1 — 61.2 8.3 — 22.8 14.4 — 25.4 Winter 1983-84 xr 45.7 B92 10.6 9.0 (8) 95% CI 39.8 — 51.6 29.5 — 48.8 8.7 — 12.5 2.3 — 15.7 Summer 1984 1 ba 60.4 33.8 19.7 aL (8) 95% Cl 54.5 — 66.3 24.0 — 43.6 16.1 — 23.3 14.3 —21.0 Winter 1984-85 x2 34.3 SES ite 333 (6) 95% CI 22.4 — 46.1 5.1-21.8 4.7-17.5 2.2-4.4 Summer 1985 V2 36.3 12.1 125 6.9 (9) 95% Cl 31.9 — 40.7 7.9 — 16.2 5.4 — 19.6 3.7— 10.1 Winter 1985-86 yf 41.1 22:2 19.6 4.4 (7) 95% CI 36.9 — 45.3 6.2 — 38.2 10.5 — 28.7 1.8—7.0 These old field populations also had increased recruit- In terms of mean abundance, Deer Mouse numbers ment in early summer (April — May) in some years _ in old field sites ranged from a low of 12.1/ha to a high which was likely related to the decline in breeding — of 60.4/ha during the four-year study (Table 1). Mean activity by mid-summer (see Figure 2). abundance of Deer Mice on the two orchard sites Peromyscus maniculatus Old field 1 +2 80 fe)) oO & jo) NO fo) Percentage of adult males breeding ! JSNJIMMJISNJIMMJISNJIMMJISNJIMM 1982 1984 1985 1986 FiGurE 2. Percentage of adult males in reproductive condition for Deer Mice for pooled data in old field and orchard sites during breeding seasons each year. Shaded bars indicate winter periods. Months of year are represented by J = July: S = September; N = November; J = January; M = March; M = May. 2004 ranged from a low density of 3.3/ha to a high of 19.9/ha. Orchard Deer Mice had annual increases in recruitment in early summer with generally higher populations (1.5 times) in summer than winter (Figure 1, Table 1). Demographic parameters Adult male Deer Mice commenced breeding in January-February each year and continued up to July in the old field sites and through the summer and autumn months in the orchard sites (Figure 2). This longer breeding season in orchard than old field sites was significantly different in terms of proportion of reproductive males in 3 of 4 years (Table 2). Propor- tion of breeding female Deer Mice tended to follow this pattern but there was a significant difference between sites in 1984 only (Table 2). The pattern of recruitment in terms of mean num- bers of new Deer Mice was generally similar in old field and orchard sites in both summer and winter periods (Table 3). There was only one period (summer SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 303 1985) when an old field — orchard comparison of mean values did not have overlapping 95% CI (Table 3). Thus, productivity within orchard populations of Deer Mice appeared to approach that of old field popula- tions, at least with respect to breeding potential and production of young. Over the four breeding seasons, early juvenile sur- vival was similar with overall mean values of 3.64 and 2.12 young Deer Mice captured per lactating female in old field and orchard sites, respectively. Mean total survival of Deer Mice was consistently higher in the two old field sites and orchard-1 site than in the orch- ard-2 site, except in summer 1983 when survival of Deer Mice was significantly lower in the orchard than old field sites (Figure 3). Mean body mass of Deer Mice was the same or higher in orchard than old field sites during summer and winter periods (Figure 4). Only during winter 1983-1984 did body mass of mice in orchard-2 appear lower than that in the old field sites. TABLE 2. Proportion of adult male and female Deer Mice in reproductive condition during breeding seasons each year for pooled data in replicate old field and orchard sites and results of Chi-square (y)analysis. Sample size (number of mice) in parentheses. Significant P in bold face. Old field Orchard Analysis Year ve PB. Males 1982 0.62 (84) 0.79 (28) 2.60 0.11 1983 0.67 (367) 0.95 (114) 34.87 <0.01 1984 0.67 (253) 0.81 (86) 6.58 0.01 1985 0.79 (150) 0.97 (39) qo <0.01 Females 1982 0.26 (69) 0.38 (42) eg 0.19 1983 0.30 (209) 0.30 (106) 0.00 1.00 1984 0.40 (182) 0.64 (96) 13.82 <0.01 1985 0.47 (103) 0.61 (49) 2.84 0.09 TABLE 3. Mean number of recruits (X) + 95% confidence intervals for Deer Mice during summer and winter periods in replicate old field and orchard sites. Sample size (n = number of trapping periods) in parentheses. Old field Orchard Period 1 2 1 2 Summer 1982 X Syl Sel 4.6 5.6 (7) 95% Cl 2.6 —7.7 6.2 — 20.1 1:27 8:0 2.2 -8.9 Winter 1982-1983 X 4.0 6.6 2.8 2) (5) 95% CI 1.7-6.3 2.6 — 10.6 0.0 —5.6 1.2—3.2 Summer 1983 X 6.7 10.0 49 5.6 (9) 95% CI 2.0— 11.3 SO llsy(9) DAVIS) 2.2 —-8.9 Winter 1983-1984 x BS) 3.5 1.9 3.8 (8) 95% CI 1.5-9.5 169-4 0.8 — 2.9 1.9-5.6 Summer 1984 X 9.9 SS) DS) 5.8 (8) 95% CI 7.3 —12.4 2.5 —8.5 2.5—8.5 3.2 — 8.3 Winter 1984-1985 X 3d 1S De), 0.7 (6) 95% CI 0.6 — 6.7 -0.6 — 3.6 -1.0 —5.3 One? Summer 1985 X 7.8 3.4 3.9 2.0 (9) 95% Cl 4.3—11.2 2.1-—4.8 1.9-—5.9 1.1-—2.9 Winter 1985-1986 X 8.6 3.9 7.0 Dal (7) 95% CI 1.5 — 15.6 0.5 -— 7.3 eS P27/ 0.2 —4.1 304 THE CANADIAN FIELD-NATURALIST Vol. 118 F @ i O Orchard 1 Peromyscus maniculatus orice” mn @ Old field 2 O Orchard 2 | O 2 c LJ 2 06 " © ne) 1) of > = 04 if ? 2 S 14) oO => 0.2 0 Summer Winter Summer Winter Summer Winter Summer Winter 1982 1982-83 1983 1983-84 1984 1984-85 1985 1985-86 FiGurE 3. Mean Jolly-Seber survival (per 28 days) + 95% confidence intervals for Deer Mice in replicate old field and orchard sites during summer and winter periods in the study. Yellow-pine Chipmunk populations Totals of 137 and 79 individual Yellow-pine Chip- munks were captured in the two old field and two orchard sites, respectively. Only three individual chip- munks were captured in the orchard-2 site during the study. Mean J-S trappability estimates ranged from 30.8% to 63.8% in the old field sites, and from 37.1% to 58.3% in the orchard sites. Populations of Yellow-pine Chipmunks ranged in mean abundance/ha from 5.6 to 19.0 in old field sites and from 1.9 to 17.5 in the orchard-1 site (Table 4). Mean abundance of Yellow-pine Chipmunks was sig- nificantly (non-overlapping 95% confidence intervals) lower in the orchard-1 than old field sites in 1982 and 1985 (Table 4). In terms of population changes over the study, Yellow-pine Chipmunk abundance/ha ranged from 1.0 — 27.0 in old field sites and from 1.0 — 46.3 in the orchard-1 site (Figure 5). Mean number of Yellow-pine Chipmunk recruits was similar between the old field and orchard-1 sites (Table 4). Mean (+ SE) Jolly-Seber survival rates averaged 0.78 + 0.05 (summer) and 0.96 + 0.02 (win- ter) in the old field-1 site and 0.71 + 0.05 (summer) and 0.91 + 0.06 (winter) in the orchard-1 site. Discussion Deer Mouse and Yellow-pine Chipmunk populations The pattern of abundance of Deer Mice in our old field sites was somewhat different from that recorded in other old field and perennial grassland studies of Peromyscus and Microtus. Grant (1972) and Baker (1968) suggested that the Deer Mouse suffers from competition from microtine rodents in perennial grass- lands, and is forced to live in woodlands. However, Deer Mouse densities in our studies reached annual peaks ranging from 30.8 to 69.3 animals/ha in the pres- ence of relatively high numbers of Montane Voles (Sullivan et al. 2003). Experimental studies by Grant (1971) and Redfield et al. (1977) concluded that Micro- tus Outcompete Peromyscus in grassland habitats; how- ever, explanations for this process were not given. Conversely, other studies concluded that competitive interactions among microtine rodents and Deer Mice were unimportant (Gilbert and Krebs 1984; Galindo 2004 SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 305 ® Old field 1 O Orchard 1 Old field 2 O Orchard 2 Peromyscus maniculatus 30 35 43 S 86 21 my CD oA 57 a 97 141} J 169 ,, 13 68 478 = t 10440) ze 44 36 > mA 96 90 3 72 158 2 + ' 220 87 = if 355 ; = { = 2) 15 Summer Winter Summer Winter Summer Winter Summer Winter 1982 1982-83 1983 1983-84 1984 1984-85 1985 1985-86 FiGuRE 4. Mean body mass (g) + 95% confidence intervals for Deer Mice in replicate old field and orchard sites during summer and winter periods in the study. TABLE 4. Mean abundance (X) and mean number of recruits per ha (X) (+ 95% confidence intervals) for Yellow-pine Chipmunks during non-hibernation periods in replicate old field and orchard sites. Sample size (x = number of trapping periods) in parentheses. Attribute and Old field Orchard period 1 2 1 2 Abundance 1982 X 5.6 7.9 1.9 - (7) 95% CI 3.6 — 7.7 3.8 — 12:0 0.7 — 3.0 - 1983 X 8.1 11.8 9.3 0.1 (13) 95% Cl 5.3 — 10.9 9.8 — 13.8 2.2 — 16.4 = 1984 X 19.0 8.9 TES) 0.2 (11) 95% CI 15.2 — 22.9 6.2 — 11.6 8.7 — 26.2 - 1985 X 14.5 9.8 3.6 - (10) 95% Cl 12.1 — 16.9 6.2 — 13.4 1.9-—5.4 — Recruits 1982 X 1.1 Da 0.6 - (7) 95% CI 0.5 — 1.8 0.1-—5.3 -0.2 — 1.3 — 1983 X eS) 12 2.3 0.1 (13) 95% Cl 0.1 — 3.0 0.0 — 2.4 0.3 — 4.3 - 1984 x De) 0.7 3.4 0.2 (11) 95% CI E2339 -0.1-1.5 0.6 —6.1 ~ 1985 X 1S 2.1 0.4 - (10) 95% CI 0.7 — 2.3 0.5 — 3.7 -0.3-1.1 = 306 THE CANADIAN FIELD-NATURALIST Vol. 118 Tamias amoenus 50 N () as (o) (~) oO Number of animals/ha =x oO --Old field 1 -# Old field 2 < Orchard 1 J S NJ MM:J SON JOM.M J°S.N JOM Mod -S NLS MiGM 1982 1983 1984 1985 1986 FiGuRE 5. Population densities (Jolly-Seber) per ha of Yellow-pine Chipmunks in two old field sites and one orchard site during the study. Shaded bars indicate winter periods. Months of year are represented by J = July; S = September; N = November; J = January; M = March; M = May. and Krebs 1985). Sullivan and Krebs (1981) reported persistence of Deer Mice with Microtus spp. in a grass- land habitat. Our densities of Deer Mice were as high as reported in other studies of perennial grassland hab- itats and orchards (Sullivan and Krebs 1981; Sullivan et al. 1998). This study is the first detailed investigation of the population dynamics of non-target rodent species in orchards treated with rodenticides for vole control. Populations of Deer Mice in orchard sites showed con- sistent annual changes in abundance averaging 15.0 animals/ha in summer and 10.1 animals/ha in winter periods. These values were similar to those reported for Montane Voles in these same orchards at 16.5 ani- mals/ha in summer and 21.1 animals/ha in winter (Sul- livan et al. 2003). Clearly, Deer Mice seemed produc- tive in the orchard environment, having longer breed- ing seasons than in the old field sites, and comparable survival of young animals. Winter breeding in Deer Mice has been reported, particularly after substantial mast crops in forests (Wolff 1996) or in supplemental food studies (Taitt 1981). Food resources appeared sufficient to support breeding mice during winter peri- ods and at body masses comparable to old field popu- lations during most periods. These orchards received regular inputs of fertilizer and irrigation, and hence likely produced relatively “rich” habitats in terms of plant production and invertebrate biomass. Mean overall survival of Deer Mice in orchards was poor, in at least one of the sites, relative to old field populations. A lack of vegetative cover may have contributed to greater predation by raptors, weasels (Mustela spp.), and Coyotes (Canis latrans). Our orchards were situated in a mosaic of different varieties and age classes of orchards and vineyards with adjacent natural habitats of sagebrush and Pon- derosa Pine (Pinus ponderosa). As discussed by Sul- livan et al. (1998) for their study in this same mosaic, it was possible that our relatively high numbers of Deer Mice (and Yellow-pine Chipmunks) in orchards may have represented transients or animals who lived in nearby sage-pine areas. However, a comprehensive analysis of transient and resident animals suggested strongly that these rodents very likely lived in the actual 2004 orchard units (Sullivan et al. 1998). There was no rea- son to assume that our animal populations behaved differently, and hence our population estimates were considered accurate. The prediction of the hypothesis that population dynamics of Deer Mice and Yellow-pine Chipmunks would be enhanced in old field compared with orch- ard habitats is partially accepted. Abundance, recruit- ment, and overall survival of Deer Mice were higher in old field than orchard sites. However, proportion of breeding animals and length of breeding seasons were greater in the orchard sites. Early juvenile sur- vival and body mass were similar between sites for Deer Mice. Similarly, the general lack of differences in abundance (except in 1982 and 1985) and recruits for Yellow-pine Chipmunks between sites (albeit for one replicate only) also contradicted our prediction. These rodent species should be able to maintain their population levels in association with Microtus in these habitats. Traditional methods of vole control (rodenticides) seem to have had little effect on these two non-target species in the orchard sites. The higher abundance of Deer Mice, and sometimes Yellow-pine Chipmunks, in the old field than orchard sites was more likely a function of habitat quality than expo- sure to rodenticides. Consequently, Deer Mice and Yellow-pine Chipmunks may assist integrated pest management in orchards because both species per- sisted in this managed habitat and their consumption of seeds and invertebrates could provide some degree of assistance in reducing weed and insect pests. In addition, as prey species, they may attract a greater number of predators to these sites. A total of six small mammal species was recorded in these orchards, which suggests that these agro-ecosystems and adja- cent natural lands help contribute to a diversity of habitats in this agrarian landscape. Acknowledgments We thank the Research Branch of Agriculture Can- ada and the Applied Mammal Research Institute for financial support; the study area and facilities provided by the Summerland Research Station; and field assis- tants W. Kaiser, J. Gareau, B. Jones, J. Krebs, and C. Robinson. Literature Cited Baker, R. H. 1968. Habitats and distribution. Pages 98-126 in Biology of Peromyscus (Rodentia). Edited by J. A. King. Special Publication Number 2. American Society of Mam- malogists, Boston, Massachusetts. Byers, R. E. 1985. Management and control. Pages 621-646 in Biology of New World Microtus. Edited by R. H. Tam- arin. Special Publication Number 8. American Society of Mammalogists, Boston, Massachusetts. Dueser, R. D., M. L. Wilson, and R. K. Rose. 1981. Attri- butes of dispersing meadow voles in open-grid populations. Acta Theriologica 26: 139-162. SULLIVAN, SULLIVAN, AND HOGUE: POPULATION DYNAMICS OF DEER MICE AND CHIPMUNKS 307 Galindo, C., and C. J. Krebs. 1985. Habitat use and abun- dance of deer mice: interactions with meadow voles and red-backed voles. Canadian Journal of Zoology 63: 1870- 1879. Gilbert, B. S., and C. J. Krebs. 1984. Competition between Peromyscus maniculatus and other small rodents in the boreal forest of the southern Yukon Territory. Acta Zoo- logica Fennica 172: 51-56. Grant, P. R. 1971. Experimental studies of competitive inter- action in a two species system. II. Microtus and Pero- myscus species in enclosures. Journal of Animal Ecology 40: 323-350. Grant, P. R. 1972. Interspecific competition among rodents. Annual Review of Ecology and Systematics 3: 79-106. Gerard, P. D., D. R. Smith, and G. Weerakkody. 1998. Limits of retrospective power analysis. Journal of Wildlife Management 62: 801-807. Gerard, P. D., D. R. Smith, and G. Weerakkody. 1998. Limits of retrospective power analysis. Journal of Wildlife Management 62: 801-807. Hillborn, R., J. A. Redfield, and C. J. Krebs. 1976. On the reliability of enumeration for mark and recapture census of voles. Canadian Journal of Zoology 54: 1019-1024. Johnson, D. H. 1999. The insignificance of statistical signi- ficance testing. Journal of Wildlife Management 63: 763- 772. Jolly, G. M., and J. M. Dickson. 1983. The problem of un- equal catchability and mark-recapture estimation of small mammal populations. Canadian Journal of Zoology 61: 922-927. Krebs, C. J. 1966. Demographic changes in fluctuating pop- ulations of Microtus californicus. Ecological Monographs 36: 239-273. Krebs, C. J. 1979. Dispersal, spacing behaviour, and genetics in relation to population fluctuations in the vole Microtus townsendii. Fortschritte der Zoologie 25: 61-77. Krebs, C. J., B. S. Gilbert, S. Boutin, A. R. E. Sinclair, and J. N. M. Smith. 1986. Population biology of snow- shoe hares. I. Demography of food-supplemented popula- tions in the Southern Yukon, 1976-1984. Journal of Animal Ecology 55: 963-982. Lewis, S. R., and J. M. O’Brien. 1990. Survey of rodent and rabbit damage to alfalfa hay in Nevada. Pages 116-119 in Proceedings of the 14" Vertebrate Pest Conference. Edited by L. R. Davis and R. E. Marsh. University of California, Davis, California. Manson, R. H., R. S. Ostfeld, and C. D. Canham. 2001. Long-term effects of rodent herbivores on tree invasion dynamics along forest-field edges. Ecology 82: 3320-3329. Pearson, D. E., Y. K. Ortega, K. S. McKelvey, and L. F. Ruggiero. 2001. Small mammal communities and habitat selection in northern Rocky Mountain bunchgrass: Impli- cations for exotic plant invasions. Northwest Science 75: 107-117. Redfield, J. A., C. J. Krebs, and M. J. Taitt. 1977. Com- petition between Peromyscus maniculatus and Microtus townsendii in grasslands of coastal British Columbia. Journal of Animal Ecology 46: 607-616. Schweiger, E. W., J. E. Diffendorfer, R. D. Holt, R. Pier- otti, and M. S. Gaines. 2000. The interaction of habitat fragmentation, plant, and small mammal succession in an old field. Ecological Monographs 70: 383-400. Sullivan, T. P., D. S. Sullivan, and E. J. Hogue. 2000. Impact of orchard vegetation management on small mammal pop- ulation dynamics and species diversity. Pages 398-403 in 308 Proceedings of the Nineteenth Vertebrate Pest Confer- ence. Edited by T. P. Salmon, and A. C. Crabb. University of California, Davis, California. Sullivan, T. P., D. S. Sullivan, E. J. Hogue, R. A. Lauten- schlager, and R. G. Wagner. 1998. Population dynamics of small mammals in relation to vegetation management in orchard agroecosystems: compensatory responses in abundance and biomass. Crop Protection 17: 1-11. Sullivan, T. P., D. S. Sullivan, and E. J. Hogue. 2003. Demography of montane voles in old field and orchard habitats in southern British Columbia. Northwest Science 77: 228-236. Sullivan, T. P., and C. J. Krebs. 1981. An irruption of deer mice after logging of coastal coniferous forest. Canadian Journal of Forest Research 11: 586-592. Sutton, D. A. 1992. Tamias amoenus. Mammalian species, Number 390. American Society of Mammalogists, Provo, Utah, USA. THE CANADIAN FIELD-NATURALIST Vol. 118 Taitt, M. J. 1981. The effect of extra food on small rodent populations. I. Deer mice (Peromyscus maniculatus). Journal of Animal Ecology 50: 111-124. Tamarin, R. H. 1977. Demography of the beach vole (Micro- tus breweri) and the meadow vole (M. pennsylvanicus) in southeastern Massachusetts. Ecology 58: 1310-1321. Webster, W. D., and J. K., Jones, Jr. 1982. Reithrodontomys megalotis. Mammalian species, Number 167, American Society of Mammalogists, Provo, Utah, USA. Wolff, J. O. 1996. Population fluctuations of mast-eating rodents are correlated with production of acorns. Journal of Mammalogy 77: 850-856. Zar, J. H. 1984. Biostatistical Analysis. Prentice-Hall, Engle- wood Cliffs, New Jersey, USA. Received 11 August 2002 Accepted 24 December 2004 Density and Survival of Lady Beetles (Coccinellidae) in Overwintering Sites in Manitoba* W. J. TURNOCK and I. L. WISE Cereal Research Centre, Agriculture and Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, Canada R3T 2M9 Turnock, W. J., and I. L. Wise. 2004. Density and survival of Lady Beetles (Coccinellidae) in overwintering sites in Manitoba. Canadian Field-Naturalist 118(3): 309-317. The densities of lady beetles, Coccinellidae, overwintering as adults (adults per m7) in leaf litter collected in late October for two years in a beach-ridge forest on the south shore of Lake Manitoba were 56.4 for the Thirteen-spotted Lady Beetle, Hippodamia tredecimpunctata (Say), 38.3 for the Seven-spotted Lady Beetle, Coccinella septempunctata (L.), 7.7 for the Transverse Lady Beetle, Coccinella transversoguttata richardsonii Brown, 1.6 for the Convergent Lady Beetle, Hippodamia convergens Guerin, and 0.6 for the Parenthesis Lady Beetle, Hippodamia parenthesis (Say). The mean overwintering survival for these species was 0.254, 0.036, 0.023, 0.0, and 0.0, respectively. The density of overwintering coccinellids was highest near the margins of the forest, particularly on the beach side, where beetles from shore appear to have entered the forest. The mean density over 3 years (2.9 per m7) of all coccinellid species in November in the litter under a remnant grove of riverbank forest in Winnipeg, was lower than in the beach-ridge forest (104.8 per m?), but their survival (0.460) was higher than in the beach-ridge forest (0.154). More species of coccinellids were found in the samples from the riverbank forest than from the beach-ridge forest. Key Words: Lady Beetles, Coccinellidae, leaf litter, beach-ridge forest, overwintering sites, Lake Manitoba, Red River. The overwintering sites of north-temperate zone coc- cinellids include leaf litter, grass tufts, bark crevices, stone piles, and rock clefts (Hodek and Honék 1996). Adults of some species migrate long distances to hiber- nate at high elevations, and even those which move only locally seem to be attracted to even slightly higher elevations and vertical silhouettes. In Bohemia and Moravia, Coccinella septempunctata L. (C7) prefers elevated places, but also hibernates in refuges among cultivated areas and does not migrate long distances (Hodek 1960). In southern Manitoba, where much of the land is flat and cultivated, it is generally believed that lady beetles overwinter in the leaf litter under trees in field shelterbelts, farmsteads and urban plantings, along the shores of streams and lakes, and in “bluffs” (groves of trees on uncultivated land). The species oc- currence, density, and survival of coccinellids in most of these overwintering sites have not been studied. Large numbers of pre-hibernating coccinellids are washed ashore in some years on Manitoba lakes, usu- ally in late August to early October (Lee 1980; Turnock et al. 2003). These beetles usually fly away soon after, to complete their search for hibernating sites, and do not often walk into the adjacent beach-ridge forest. Once, on 1 November 1978, beetles (mainly H. con- vergens) were washed ashore, but subsequent cold weather forced them to remain near the beach, where few survived the winter (Turnock and Turnock 1979). In this paper, the density, distribution, species com- position, and overwintering survival of coccinellids in the litter under a beach-ridge forest are compared with the density and survival of coccinellids in the litter under a riverbank forest. Methods Beach-Ridge Forest. The abundance and species composition of coccinellids overwintering in the sur- face litter and soil under a beach-ridge forest on the south shore of Lake Manitoba at the Delta Marsh Field Station (University of Manitoba), 50°11'N, 98°23'W, about 120 km WNW of Winnipeg, were recorded. The shore is a sand beach with a forested barrier-beach ridge formed by the reworking of alluvial sand depos- its. This beach ridge, about 60 m wide, supports a mature deciduous forest of Manitoba Maple (Acer negundo L.), Green Ash (Fraxinus pennsylvanica Marsh.), Plains Cottonwood (Populus deltoides Marsh.), and Peachleaf Willow (Salix amagdaloides Anderss.) (Kenkel 1986). The ridge has its highest elevation 5- 10 m from the top of the foreshore and slopes down about 50 m to the marsh. A road about 10 m wide along the south edge of the beach ridge separates the forest from the marsh. Samples were taken along five tran- sects, 20 m apart, perpendicular to the beach. Six sam- ples were taken along each transect at 1, 6, 11, 21, 31, and 41 m from the top of the foreshore, where the beach ended and the beach-ridge forest began. A sample of the leaf litter and soil above the sandy par- ent material, 0.25 m? and 5-10 cm deep, was collect- ed at each of the 30 locations. Samples were collected after beetle flight had ceased, on 22 October 1992 and 27 October 1993. Each sample was bagged, sorted by *Contribution Number 1832 from the Cereal Research Centre, Agriculture and Agri-Food Canada, 195 Dafoe Road, Winnipeg, Manitoba, Canada R3T 2M9. 309 310 hand to remove the beetles, and the species and num- ber of coccinellids were recorded. The numbers and species of coccinellids emerging in the spring were determined from cone traps, each covering a surface area of 0.1028 m* (Turnock et al. 1987). One trap was set into the litter at each of the 30 locations, near the place where a litter sample had been collected the previous autumn. The traps were set out at each location in late April or May, depend- ing on the disappearance of snow, in 1993, 1994, and 1995. The traps at the apex of the cones were emptied at regular intervals, and the coccinellids identified to species and counted. Riverbank Forest. The overwintering of coccinellids in a remnant grove of riverbank forest along the Red River at the University of Manitoba, Winnipeg, Mani- toba, was determined from records of the insects found in litter samples and emergence traps used to study the overwintering of flea beetles (Chrysomelidae) (Tur- nock et al. 1987). The grove of trees, about 1.3 ha in area, included Bur Oak (Quercus macrocarpa Michx.), American Elm (Ul/mus americana), Manitoba Maple, Green Ash, Black Ash (Fraxinus nigra Marsh.), and Basswood (Tilia americana L.). In this grove, 50 litter samples, each 0.25 m7, <10 cm deep, were collected in the autumn of the years 1980-1982, and in the springs of 1980, 1981, and 1983. The samples were taken at 10 m intervals along five transects running north to south through the grove. Spring emergence was sam- pled by 100 cone traps, each covering 0.1028 m/, in 1980 and 1981. Two traps were located at 10 m inter- vals along each transect, one on each side of the site of a litter sample, as described by Turnock et al. (1987). For coccinellids, only the total number collected in each sampling period was recorded, except in the litter samples of 3 November 1982, where the species were identified but their numbers were not recorded. Results Beach-Ridge Forest. The dominant species in all collections were H. tredecimpunctata, C. septempunc- tata, and Coccinella transversoguttata. In the autumn litter samples in 1992 (n = 653) and 1993 (n = 918) coccinellid relative abundances were: 0.54/0.54 (H. tredecimpunctata); 0.36/0.37 (C. septempunctata); 0.08/ 0.07 (C. transversoguttata); 0.01/0.02 H. convergens; and 0.004/0.001 H. parenthesis (Say). In the spring emergence samples, the relative abundances of each species in 1993/94/95 were: H. tredecimpunctata — 0.95/0.78/0.92; C. septempunctata — 0.04/0.19/0.06; and C. transversoguttata — 0.02/0/0. Neither H. conver- gens nor H. parenthesis were found in the spring sam- ples. The distribution of numbers per sample of H. tredecimpunctata and C. septempunctata was skewed in both the autumn litter samples and the spring emer- gence samples, with most samples containing 0, 1, or 2 beetles (Figure 1). THE CANADIAN FIELD-NATURALIST Vol. 118 The mean density per m* of coccinellids in the autumn was 87.1 in 1992 and 122.4 in 1993, and 8.5 and 11.7 in the following springs. The density of coc- cinellids in the autumn was generally higher on the margins of the forest, but it varied among species (Figure 2). The numbers of H. tredecimpunctata were highest near the beach margin in 1993, but they were higher near the marsh margin in 1992. The density of both C. septempunctata and C. transversoguttata was higher near the beach margin in both years, with only a slight increase along the marsh margin. These patterns suggest that most of the C. septempunctata and C. transversoguttata found in the litter near the beach were from aggregations that had washed ashore (Turnock et al. 2003). Hippodamia tredecimpunctata seems mainly to have come from the beach in 1993, but in 1992 more appeared to have come across the marsh to the edge of the forest. The proportion of living beetles in the autumn litter samples differed little between years, 81.3% in 1992 (n = 653) and 81.4 % (n= 913) in 1993 (Table 1). The mean proportion of living beetles, by species for both years, was: H. tredecimpunctata = 0.963 (n = 847); C. septempunctata = 0.729 (n = 575); C. transversogut- tata = 0.250 (n = 116); H. convergens = 0.458 (n = 24); and H. parenthesis = 0.500 (n = 4). Survival of all spe- cies was lowest adjacent to the beach and increased toward the edge of the marsh (Figure 3). The decreased survival in the forest edge near the beach was prob- ably caused by the burial of overwintering beetles by sand blown off the beach by strong winds. Coccinellids emerged from the leaf litter during May and early June in traps throughout the beach-ridge. Most of the beetles (98%, n = 187) were of two spe- cies, H. tredecimpunctata and C. septempunctata. The mean density per m? of emerging coccinellids in 1993 to 1995 was: 17.5, 9.1, 27.9 for H. tredecimpunctata; 0.65, 2.3, 1.9 for C. septempunctata; and 0.32, 0, 0 for C. transversoguttata. The pattern of higher densities near the margins of the forest was less prominent in the spring than in the autumn (Figures 2 and 4). The density of emerging beetles was <20 per m? except at distances from the beach of | m in 1995, 31 m in 1994, and 41 m in 1993. The mean overwintering survival (density spring/ density autumn) in the beach-ridge forest for H. tre- decimpunctata was 0.370 in the winter of 1992-1993, and 0.139 for 1993-1994 (Table 1). The survival ratios over the same two winters for C. septempunctata were (0.021 and 0.051, and for all coccinellids, 0.212 and 0.096. The survival of H. tredecimpunctata was quite variable with position within the forest (Figure 5). Overwintering survival of all species was very low at 1 m from the beach margin of the forest, where drifting sand often covered the leaf litter. However, the over- wintering survival did not significantly differ among years, density of coccinellids in the autumn, or distance 2004 TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 31] Coccinella septempunctata Mmm 1992 Gas%] 1993 Frequency 60 80 100 Hippodamia tredecimpunctata Frequency 100 Coccinella transversoguttata Frequency 80 100 0 20 40 60 Beetles per sample FIGURE 1. Frequency distribution of numbers of coccinellids, Hippodamia tredecimpunctata, Coccinella septempunctata, and Coccinella transversoguttata, per sample of litter (0.25 m? x 5-10 cm deep) from the floor of the beach-ridge forest at the Delta Marsh Field Station, Manitoba, in late October of 1992 and 1993. n = 60 for each species. SH THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 1. The mean numbers (N) per m? in autumn litter samples and spring emergence samples, the proportion of living beetles in the autumn samples, and the overwintering survival (density spring emergents/density in autumn), for Coccinella septempunctata (C7), Hippodamia tredecimpunctata (H13), C. transversoguttata (CT), H. convergens (HC), and H. parenthesis (HP), and all coccinellids in the beach-ridge forest at the Delta Marsh Field Station. Species 1992 1993 Autumn Spring H13 N/m? 47.3 IES Survival 0.95 0.37 N 355 54 (Oy N/m? Si1k3 0.65 Survival 0.72 0.02 N 235 2 CT N/m? 6.9 0.32 Survival 0.37 0.05 N 52 1 HC N/m2 101 0 Survival 0.38 0 N 8 0 HP N/m? 0.4 0 Survival 0.67 0 N 3 0 All N/m? 87.1 18.5 Survival 0.81 0.21 N 653 57 1993 1994 1995 Autumn Spring Spring 65.6 9.1 27.9 0.97 0.14 ~ 492 28 86 45.3 ep) 129 0.73 0.05 - 340 q 6 8.5 0 0 0.16 0 - 64 0 0 25k 0 0 0.5 0 - 16 0 0 0.13 0 0 1 0 - 0 0 122.4 11.7 30.5 0.81 0.1 — 913 36* 94% *Includes one specimen of Anistosticta bitriangularis in 1994 and two specimens of Calvia quatuordecimguttata in 1995. from the beach (analysis of variance, proc glm, SAS Inst. Inc. ). The high survival of H. tre- decimpunctata at several locations likely reflects the placing of the emergence traps over local aggrega- tions of beetles in the litter, as does a single record of high survival of C. septempunctata. Riverbank Forest. The litter samples collected on 3 November 1982 contained nine species of coccinel- lids: Stethorus punctum (LeConte), Scymnus (Pullus) brullei Muls., Hyperaspis benedetti (Say), Chilocorus stigma Say, H. tredecimpunctata, Anatis labiculata (Say), Adalia bipunctata L., C. transversoguttata, and Psyllobora virginimaculata (Say). The mean density of all coccinellid species in the autumn litter collec- tions of 1980-1982 (n = 110) was 2.9 per m? (2.0- 4.3), vs. 4.4 (1.5- 10.4) in the spring litter collections (n = 172) of 1980, 1981, and 1983. Overwintering survival, calculated by dividing the number of living beetles in the spring litter samples by the total num- ber of beetles in the autumn samples, was 76% for the winter of 1980-1981 and 43% for the winter of 1982- 1983. Survival during the winter of 1980-1981 was lower, 49%, when the spring density, 0.97 per m*, was based on captures in the emergence traps (n = 10). Discussion and Conclusions The studies of coccinellid density and survival in three overwintering sites in southern Manitoba differ in location, years, and sampling design. However, the results of some comparisons may be helpful to future investigators of site selection and survival by overwin- tering coccinellids. In the autumn, more overwintering lady beetles were found in a beach-ridge forest on the shores of Lake Manitoba in 1992 and 1993 than in a riverbank forest in Winnipeg in 1980 to 1981. The difference in the density between the two sites may have been related to differences in the populations of lady beetles in the years sampled. Another factor could be the availability of overwintering sites in the two areas. The beach-ridge forest was located in an intensively- farmed area with few treed areas, whereas the river- bank forest was near suburban Winnipeg, where trees were more abundant. Beetles washed ashore on the beach contributed to the number overwintering in the beach-ridge forest in 1993, but in 1992 more beetles seem to have entered the forest from the side away from the beach. The beach-ridge forest, despite higher initial den- sities, had fewer species and fewer beetles emerging in the spring than in the riverbank forest, because of lower overwintering survival. Overwintering survival by beetles may be related to their need for moisture (Hodek and Honék 1996). The litter under the river- bank forest was thicker and lay upon a clay soil, thus providing a moister environment than the thinner litter on a sandy soil under the beach-ridge forest. The limited sampling by Turnock and Turnock (1979) on the shores of Lake Manitoba, following a very late flight (1 November) and aggregation of beetles, indicated few beetles survived the winter (2% of 1002 per m? trapped in the beach debris). Even this level of survival is surprising, because beach debris usually is pounded and buried by high water, large 2004 TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES Beetles per m? Beetles per m? Beetles per m? 30 40 50 0 10 20 Distance from forest edge (m) FiGurE 2. Mean density per m? of Hippodamia tredecimpunctata, Coccinella septempunctata, Coccinella transversoguttata, and all coccinellids, in forest litter samples collected at various distances from the forest edge next to the beach in late October 1992 and 1993 at the Delta Marsh Field Station. 314 THE CANADIAN FIELD-NATURALIST Vol. 118 Proportion alive Proportion alive All species Proportion alive 0 10 20 30 40 50 Distance from forest edge (m) FicuRE 3. Proportion of live individuals of Hippodamia tredecimpunctata (H13), Coccinella septempunctata ( (C7). Coccinella transversoguttata (CT), and all coccinellids, in litter samples taken at different distances from forest edge next to the beach in late October of 1992 and 1993. TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 2004 —e— H13 -O::- C7 N —-w— All species = —. = (oy [ou 0) a) x) ® faa) N E = (or) Q. o v zr) ® a N i= = ® a 72) & © ® a Distance from forest edge (m) FiGure 4. Mean density per m? of Hippodamia tredecimpunctata (H13), Coccinella septempunctata (C7), and all coccinel- lids, in emergence trap samples collected at different distances from the forest edge next to the beach. The samples were collected from emergence cones placed over the litter in May of 1993, 1994, and 1995. 316 Survival THE CANADIAN FIELD-NATURALIST Vol. 118 H13-1992/3 H13-1993/4 C7-1992/3 C7-1993/4 Distance from forest edge (m) FiGurE 5. Mean overwintering survival ratios for Hippodamia tredecimpunctata (H13), and Coccinella septempunctata (C7), for the winters of 1992-1993 and 1993-1994 in forest litter on the beach ridge at different distances from the forest edge next to the beach at the Delta Marsh Field Station. waves, and ice chunks from violent autumn and spring storms. In this study the top of the foreshore, where a covering of aspen leaves had accumulated on the sand, had 74 beetles per m* and overwintering survival was 53%. This location is not likely to be a suitable over- wintering site in most years because of high water and waves. The grassy backshore, with little litter, had 28 beetles per m’, of which 18% survived the winter. The foreshore of the sandy beaches and even the shal- low litter in the sparse beach-ridge forest of Trembling Aspen, Populus tremuloides Michx., do not appear to be selected by coccinellids for overwintering. Excep- tions can occur when beetles washed ashore after un- usually late autumn flights aggregate on the beach when subsequent temperatures are too cool to allow flight, but warm enough to allow limited walking up the beach to the backshore. In these cases the beach-ridge forest, even if unsuitable, is mainly a “last-chance” overwin- tering site for these beetles. Four species of coccinellids, H. tredecimpunctata, C. septempunctata, C. transversoguttata, and H. con- vergens (in order of relative abundance) were found in both years in autumn litter samples in the beach- ridge forest, but only the first two species were found in the spring emergence samples in all three years. More H. tredecimpunctata (14 and 38%) survived over winter than C. septempunctata (3 and 5%) or C. trans- versoguttata (5 and 0%). This level of survival of C. septempunctata is much lower than the 97% survival of C. septempunctata that had been covered with litter in experimental cages at Edmonton, Alberta (Ryan and Acorn 1999). Although in Europe C. septempunctata does not migrate long distances to hibernate (Hodek 1960), large flights of this species occur in both autumn and spring, as shown by the numbers of this species that are found in the aggregations washed ashore on the shores of the lakes (Turnock et al. 2003). Southern Manitoba is mostly cultivated and has a flat terrain, so potential overwintering sites are limited to leaf litter under trees. These habitats probably are both attractive and suitable for the hibernation of coc- cinellids, but without information on the preferred over- wintering sites in this area, the true level of overwinter- ing survival by native and introduced species cannot be compared. 2004 Acknowledgments The statistical advice by S. Woods, technical assis- tance of R. J. Bilodeau, and the help given by the staff of the Delta Marsh Field Station (University of Mani- toba) in providing facilities is gratefully acknowledged. This is Publication Number 305 from the Delta Marsh Field Station (University of Manitoba). Literature Cited Hodek, I. 1960. Hibernation-bionomics in Coccinellidae. Acta Societis Entomolicae Cechosloveniae 57(1): 14-17. [In Czech with English “Discussion and Conclusions” }. Hodek, I., and A. Honék. 1996. Ecology of Coccinellidae. Series Entomological. Volume 54. Kluwer Academic Pub- lishers. Dordrecht. 464 pages. Kenkel, N. C. 1986. Vegetation structure and dynamics of the barrier-beach ridge at Delta, Lake Manitoba. Annual Report of the University of Manitoba Delta Marsh Field Station 21: 61-83. TURNOCK and WISE: DENSITY AND SURVIVAL OF LADY BEETLES 317 Lee, R. E. 1980. Aggregations of lady beetles on the shores of lakes. American Midland Naturalist 104; 295-304. Ryan, S. E., and J. H. Acorn. 1999. Overwintering survival of the seven-spot ladybug (Coccinella septempunctata) in Edmonton. Blue Jay 57: 97-100. Turnock, W. J., and R. W. Turnock. 1979. Aggregations of lady beetles (Coleoptera: Coccinellidae) on the shores of Lake Manitoba. Manitoba Entomologist 13: 21-22. Turnock, W. J., R. J. Lamb, and R. J. Bilodeau. 1987. Abun- dance, winter survival, and spring emergence of flea beetles (Coleoptera: Chrysomelidae) in a Manitoba grove. The Canadian Entomologist 119: 419-426. Turnock, W. J., I. L. Wise, and F. O. Matheson. 2003. Abun- dance of native coccinellines (Coleoptera: Coccinellidae) before and after the appearance of Coccinella septem- punctata. The Canadian Entomologist 135: 391-404. Received 9 August 2002 Accepted 24 December 2004 Description of Age-O0 Round Goby, Neogobius melanostomus Pallas (Gobiidae), and Ecotone Utilisation in St. Clair Lowland Waters, Ontario JOHN K. LESLIE and CHARLES A. TIMMINS Great Lakes Laboratory for Fisheries and Aquatic Sciences, Department of Fisheries and Oceans, 867 Lakeshore Road, Burling- ton, Ontario L7R 4A6 Canada Leslie, John K., and Charles A. Timmins. 2004. Description of age-O Round Goby, Neogobius melanostomus Pallas (Gobiidae), and ecotone utilisation in St. Clair lowland waters, Ontario. Canadian Field-Naturalist 118(3): 318-325. Early developmental stages and ecotone utilisation of the non-indigenous Round Goby, Neogobius melanostomus (Pallas, 1811), are described and illustrated. Fish (5-40 mm) were collected in coarse gravel, rocks and debris in the St. Clair River/Lake system, Ontario, in 1994-2000. The Round Goby hatches at about 5 mm with black eyes, flexed urostyle, and developed fins and digestive system. Distinguishing characters include large head, dorsolateral eyes, large fan-shaped pectoral fins, two dorsal fins, fused thoracic pelvic fins and a distinct black spot on the posterior of the spinous dorsal fin. Modal counts for preanal, postanal, and total myomeres were 12, 19, and 31, respectively. Key Words: Round Goby, Neogobius melanostomus, St. Clair aquatic ecosystem, age-0, morphometry, habitat, Ontario. Gobiidae, the largest family of marine fishes, has been found in fresh waters of all continents (Nelson 1984) except Antarctica. None of 68 species endemic to North America is native to the Great Lakes, where two introduced species, the Round Goby, Neogobius melanostomus (Pallas, 1811) and the Tubenose Goby, Proterorhinus marmoratus (Pallas, 1811), recently be- came established. These fishes originate in the Ponto- Caspian region (Caspian, Azov and Black seas and Sea of Marmara). International ships’ ballast was assumed the most probable means by which Round Goby eggs and/or postembryonic stages were transferred from the Black Sea to the Great Lakes (Crossman et al. 1992; Jude et al. 1992). Since its arrival in the mid- late 1980s the relatively innocuous Tubenose Goby has not proliferated beyond the St. Clair-western Lake Erie ecosystem (Leslie et al. 2002). In contrast, the aggressive, eurytopic Round Goby has diffused extensively and is now one of the most abundant fish in the Great Lakes basin. Apparently, repetitive spawn- ing and habitat requirements of this fish conflict with our native species (Dubs and Corkum 1996), some of which they may supplant. The Round Goby is a guarding cavity spawner with a life span of 4-5 years (Miller 1986). Spawning sea- son is typically from spring to late summer. Rocks, gravel, solid objects with crevices and submersed plants are used for egg deposition (Miller 1986; Skora 1997*). Whereas literature abounds on the ecology of adults, information on postembryonic fish is rare. Eggs and early developmental stages in southern Russia and Ukraine have been described by Moskal’kova (1967), Kalinina (1976), and Koblickaja (1981). This report presents information on aspects of taxonomy and eco- tone utilisation of age-O0 Round Gobies in the Great Lakes and considers possible effects the species may have on the aquatic community in general and native fishes in particular. Study Area Eggs and age-0 fish were collected at numerous locations at the shore of the St. Clair River, Ontario, various sites along the southeastern shore of Lake St. Clair, and in Duck Creek (42°17'N, 82°35'W), a low gradient stream tributary to Lake St. Clair (Figure 1). Duck Creek is a disturbed system draining a small area in one of the most highly cultivated agricultural regions in Canada. It is approximately 9 km long and 10-15 m wide near its mouth, where most specimens were collected. Mid-stream depth ranges from 1.2 to 1.6 m. In April to November 1995, mean conductivity was 487 uS/cm and Secchi disc depth 0.3 + 0.1 m, reflecting high suspended particulate load due to run- off from cropland. Substrate at the sampling site is mainly alluvium, with clay and scattered debris of human origin at the base of 1-4 m high stream banks. However, age-O Round Gobies were found only on a small gravelled area at the base of a railway bridge. Duck Creek has a large diverse, albeit transient, fish assemblage of at least 41 species (Leslie and Timmins 1998) dominated by Gizzard Shad Dorosoma cepedi- anum, Bluntnose Minnow Pimephales notatus, Spottail Shiner Notropis hudsonius, and Bluegill Lepomis macrochirus. Submersed plants near the collection site include Pondweed Potamogeton spp., Eelgrass Vallis- neria americana, Canada Waterweed Elodea cana- densis, Eurasian Milfoil Myriophyllum spicatum, and Curly Pondweed Potamogeton crispus. Arrowhead Sagittaria sp., Sedge Carex sp., Cattail Typha spp., and Bulrush Scirpus spp. were abundant emergent species. 318 2004 Michigan Lake Huron ichig 20 kn Lake St. Clair © Duck Creek FiGure 1. Main sites where age-O Neogobius melanostomus were collected in the St. Clair River-Lake ecosystem in 1994-2000: LGS (Lambton Generating Station) and Duck Creek. Round Goby eggs and age-0 developmental stages as well as numerous co-occurring fishes were found at various locations at the shore of the St. Clair River, mainly in dilute condenser cooling water discharge of Lambton Generating Station (LGS) (42°47'N, 82°22'W). Random fish collections took place at LGS (Figure 1) each year between April and November 1994-2000 over a shoreline stabilised with debris, limestone boulders and rocks 0.3-1.0 m in longest dimension. This formation extends 4 m from shore on a natural substrate of sand. Most common fish species collected with age-O Round Gobies in the St. Clair River include age-O0 White Sucker Catostomus com- mersoni, Alewife Alosa pseudoharengus, Brook Silverside Labidesthes sicculus, Rainbow Darter Ethe- ostoma caeruleum, lowa Darter Etheostoma exile, Johnny Darter Etheostoma nigrum, Spottail Shiner, and Emerald Shiner Notropis atherinoides. Of these species, guarding speleophils Rainbow Darter, Iowa Darter, and Johnny Darter utilise common ecotones and probably compete with gobies for food and shelter. Methods A bulging larval fish beach seine (4 m long, 1 m wide, mesh 0.4 mm) was necessary for capture of recently hatched gobies, which only emerged from pro- tective crevices when the bottom of the seine dis- LESLIE and TIMMINS: AGE-O ROUND GoBy IN ONTARIO 319 turbed the substrate. Two flat seines, 3 m long, | m wide (3 mm mesh), and 6 m long, | m wide (6 mm mesh) were used to sample young and adult gobies and co-occurring species. All seine hauls took place repeatedly in contact with gravel at the base of a rail- way bridge over Duck Creek. In the St. Clair River, clusters of eggs attached to rocks or human artefacts such as discarded sewer pipe were preserved intact with 5-10% formalin and enumerated in the laboratory. Water temperature, spe- cific conductivity, water transparency, substrate char- acteristics, and co-occurring fishes were recorded at all sampling locations except four that were devoid of cover and fishes. Fish were preserved in a 12:1 solution of 80% ethanol and glycerin, respectively, and stored at room temperature (22-25°C). A dissecting microscope equipped with an ocular micrometer was used for mea- surement and illustration of fish. Measurements <5 mm were accurate to +0.1 mm, and larger characters to +0.2 mm. Several specimens were cleared to determine vertebra number. Fish used in this study are stored at the Royal Ontario Museum, Toronto, Ontario. In general, terms used to describe fishes follow Trautman (1981). Base length of median fins is defined as the distance from the anterior margin of the first spine or soft ray to the posterior margin at the base of the last spine or ray. Snout length is the distance from the anterior margin of the premaxillary to the-anterior margin of the eye. Interorbital width is the least dis- tance between orbits. Body depth factor (BDF) and caudal depth factor (CDF) indicate general body form and swimming ability (Webb and Weihs 1986). Body depth factor is defined as mean TL/mean body depth, whereas CDF = mean caudal peduncle depth/ mean body depth. A high BDF and CDF denotes an elongate species whose depth is more or less the same along its length; e.g., clupeid or osmerid. Low values are found in centrarchids, where manoeuvrability is enhanced by a deep body and large pectoral fins. Fish volume, determined in each size class = mean total length x (mean body depth)’. Results Reproductive and nursery habitat Environmental conditions for fish reproduction and rearing differed in Duck Creek and the St. Clair River according to water quality and physical characteris- tics. On most dates, water temperature at the shore of the river was about 3°C lower and suspended particu- late load always lower, than in Duck Creek. Whether collected at 4° or 30°C, Round Gobies were recorded in the same habitat from April to November. However, their occurrence was contingent on availability of cover. For example, gobies were not found at LGS in May 1999 when low water level completely exposed rock and rubble habitat. Recently hatched fish were rarely found along featureless, sandy shores, where 320 za Do OI THE CANADIAN FIELD-NATURALIST Vol. 118 SS ES eee a ae ~ a] s FIGURE 2. Neogobius melanostomus at 7.5 mm; upper: dorsal, middle: lateral, lower: ventral view. Length bar = 1 mm. juvenile and adult gobies were occasionally caught. Small boat launches with corrugated concrete ramps were consistent sources of larger (>30 mm) specimens of age-0 gobies. Presence of small age-0 fish in late spring and early autumn indicates a possible prolonged spawning peri- od in Duck Creek, accommodated by water tempera- ture >10°C from early May to mid-October. Earliest evidence of recently hatched fish (5-13 mm TL) on 1 June 1995 suggests spawning takes place in April or May. However, in early May 1998, ten gobies (22-28 mm) were caught near LGS at 10°C above ambient (4°C). These fish may have hatched in late winter in dilute heated power plant effluent. Since small gobies (14-23 mm) were found each year in September, spawning at LGS may extend to late summer. In Duck Creek, Round Gobies endure continuous turbid water, enriched conditions, occasional dense blooms of blue-green algae (Anabeana sp; Cyanophy- ta) and temperature at least 30°C. According to size in autumn, gobies attain a mean length of about 33- 39 mm at the end of first year growth. For example, in Duck Creek fish were 32.8 mm (28-36 mm) in early October 1995, whereas at LGS, they were 37.6 mm (range 31-49 mm) in November 1994 and 39.4 mm (30-45 mm) in November 1995. Reproduction In mid-July 1999, numerous clusters of eggs were found attached to the underside of rocks at a depth of 0.3 m in Duck Creek. A sample (N = 121) of eggs in a cluster of roughly 200 on a rectangular patch of about 30 cm? averaged 3.0 mm long and 1.8 mm wide with 2004 LESLIE and TIMMINS: AGE-O ROUND GoBY IN ONTARIO 32] FIGURE 3. Neogobius melanostomus at 25 mm. Length bar = 2 mm. respective modal values of 3.1 mm and 1.8 mm (range: 2.6-3.3 mm and 1.6-2.0 mm). Eggs are oblong with rounded base and blunt apex, with a fibre-like basal pedestal or stalk 0.4-0.5 mm long. Yolk is light orange and occupies nearly 75% of the capsule. These eggs incubated in situ in turbid, almost lentic water at 28- 30°C. Similarly, egg clusters were found in early June 2000 at 14°C in the St. Clair River beneath limestone rocks and human artefacts. A cluster of approximately 300 eggs formed an oval patch (estimated area = 40 cm?) under a rock at a depth of 0.2-0.3 m in slow flowing water. Eggs (n = 78) were 3.5 mm long and 2.0 mm wide (modal values, 3.7 mm and 2.1 mm). Morphology Yolk, which is usually retained in the gut of fish 4.5 to 6.5 mm long, may persist to 11 mm. At 5-8 mm, minute teeth are present on the jaws. Origin of D1 is at the fourth myomere or fifth vertebra, whereas D2 originates at the seventh or eighth myomere (elev- enth or twelfth vertebra). Scales first appear on mid- caudal peduncle at approximately 8-9 mm and body scalation may be complete at 12-15 mm. The upper lip is protractile and jaw articulation is below the anterior margin of the pupil. The anterior tubular nostril protrudes, whilst the minute posterior nostril is flush. At about 13 mm the cranium is de- pressed, cheeks begin to enlarge, and sub-orbital canals and papillae are barely visible. At all sizes, the head is bluntly arrow-shaped in ventral profile (Figure 2). All spines and fin rays are formed, although principal rays are incomplete in the caudal fin. Pectoral fin rays are joined distally by membrane. A small genital papilla is first obvious at about 23 mm. Origin of D2 is on a vertical line with the anus and separated from D1 by a flap of tissue. Full complement of branched caudal rays is attained at about 20 mm. At 22-25 mm, the pectoral fins are slightly flared ven- trally and depressed D2 and anal fins extend equally to mid-peduncle (Figure 3). Two short, stout spines support a transverse membrane on the pelvic fin, the whole forming a suctorial disc. Fish >30 mm have a moderately rounded snout and large protruding dorsolateral eyes above bulging cheeks. Sub-orbital papillae occur in transverse rows (Figure 4). The tip of the large fleshy upper lip is anterior to the lower, and the opening of the terminal mouth lies below the ventral margin of the eye. The centre of the pupil lies along a horizontal line with the origin of the pectoral fin. Gill openings are narrow. Ctenoid scales cover the body, whereas cycloid scales occupy the anterior nape and gill covers. In outline, Ficure 4. Lateral view of male Neogobius melanostomus at 38 mm. Length bar = 5 mm. B22 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Morphometrics for age 0 Neogobius melanostomus collected in the St. Clair ecosystem, Ontario. Lengths are percentage (with range) of mean total length (TL) in each size class. Size 5.0-8.9 9.0-12.9 13.0-16.9 class n= 20 m=9 n=2 Mean TL Ussies A 10.4 + 1.2 13301 Standard 79.0 79.3 78.1 73-86 78-83 77-719 Preanal 45.8 43.8 41.5 43-52 42-45 41-42 Predorsal 33.0 Silen 30.6 30-42 30-33 30-32 Prepelvic Pes A| PAE | 27.6 18-30 24-31 27-28 Body depth 17.8 16.8 1GS)59) 14-21 15-18 14-17 Peduncle US 8.0 8.3 6-9 7-9 0 Head 26.2 26.3 27.6 21-30 24-29 27-28 Eye? 30.6 Bilal 27.6 25-37 26-35 27-28 Volume (mm?*) 12.3 30.1 5i5).7/ BDF 5.6 6.1 6.5 CDF 0.42 0.49 0.54 Pectoral fin 18.6 19.7 D5 10-22 17-22 21-22 Pelvic fin 1525 Ne, 19.3 9-21 13-19 18-20 Dorsal | base 7.0 7.8 6.5 5-11 6-10 6-7 Dorsal 2 base 24.3 235 23.4 19-30 20-29 23-24 Anal base 20.5 19.7 18.5 18-24 16-23 17-20 *% head length D1 is slightly rounded, D2 slopes slightly, and the anal fin is uniform (Figure 4). Morphometry Relative changes in lengths are generally small as fish grow (Table 1). Head length (24-28% TL) is 1.3 times its width and 1.4 times its depth. Snout length increased from 15% HL at 7 mm to 20% HL at 35 mm. Interorbital width, variable among and within size classes, is 10-22% HL. Pectoral fin rays are 19-24% TL whereas D2 base is 23-26% TL. Longest spines and base of D1 are about equal but less than the longest ray of D2. On fish >35 mm, the pectoral fin base is approximately 55% greatest body depth. Body depth and caudal depth factors are generally constant (Table 1). Meristics Respective modal counts for preanal, postanal, and total myomeres are 12 (range, 11-13), 19 (17-20), and 31 (29-32). Modal spine and fin ray complements are D1: VI, D2: 1 15 (13-16), anal: I 13 (11-13), pectoral: 18 (16-19), pelvic(s): 1 5 (conjoined), principal caudal: 17.0-20.9 21.0-24.9 25.0-32.9 33.0-39.9 n=9 n= n=33 N23, 18.5+0.9 23.5+0.9 2837 = 21 35.3+ 1.8 80.3 79.8 79.5 80.3 79-83 78-83 77-81 79-82 44.0 44.7 44.0 43.4 43-46 41-48 42-51 39-46 30.1 29.7 29.2 28.2 29-32 28-32 27-31 24-29 DAs 27.0 26.6 26.6 26-28 24-32 25-29 25-34 gh) 16.9 17.6 ei 14-20 15-19 15-20 16-19 9.0 8.9 8.9 8.8 9-10 8-9 9-10 8-10 25.5 253 24.8 24.0 25-27 24-27 23-27 23-26 33.0 32.4 B22 30.3 31-35 28-35 28-35 26-35 189 35 ii 2 1401 5.8 6.0 Si 5.6 0.53 0.52 0.51 0.50 23.6 24.0 DES) 23.6 22-25 22-25 19-27 21-26 203 19.9 20.1 19.8 20-22 18-21 18-22 17-23 8.8 8.5 9.0 9.2 8-9 7-12 8-11 7-11 pip) 24.7 25.9 DS) 24-28 20-29 24-27 24-27 19.6 19.2 20.6 20.4 17-22 17-22 17-23 19-23 13 (11-13). There are 33 modal (32-33) vertebrae and 49 (48-52) ctenoid scales in mid-lateral series. Pigmentation On free embryos, pigmentation consists of black eyes and several black spots on posterior spines of D1. However, on many specimens <9 mm, the char- acteristic black spot on D1 is absent. Speckled mel- anophores on snout and dorsum of head, a dark crescent on the occiput and a small patch on the cheek are typical at 10-12 mm. Ventrally, an elongate stellate melanophore is situated mid-gut. About five faint dorsolateral clusters are expressed on the body. There is a small group of stellate melanophores on frontal and parietal areas, a ventral patch on the pectoral fin base and a subcutaneous series on the intestine. Mel- anophores develop at the base of each anal fin ray, and a narrow mid-ventral series prevails from anal fin base to caudal fin base (Figure 2). At 14 mm, striation extends from maxilla to orbit and a black spot is pronounced between fifth and sixth spines on D1. Several faint mid-lateral blotches extend the length of the body. 2004 LESLIE and TIMMINS: AGE-O ROUND GoByY IN ONTARIO 323 FiGuRE 5. Cottus bairdi at 10.5 mm. Length bar = | mm. At 22-25 mm a large brown patch is obvious on the dorsal peduncle of the pectoral fin. First four rays of D1 are lightly pigmented, and patches appear on the base of interradials. A faint stripe exists mid-fin on D1 (Figure 3). Posterior margins of body scales are darkly outlined and rays at the base of the caudal fin are lightly pigmented. A small basicaudal spot persists throughout development. Fish >30 mm have a small stripe on the preopercle, and about seven irregular patches of brown pigment on the side of the body, mainly below the mid-line. Otherwise, lips and dorsum of head are covered with small round spots, whilst the throat is essentially de- void of pigment. Pelvic and caudal fins are lightly spotted, whereas D2 has two pale longitudinal stripes. Pigmentation patterns are intensified over head, body, and fins (Figure 4). Overall colouration on individuals varies from dark brown to grey-brown. Description of age-O Mottled Sculpin Mottled Sculpin larvae (11.4 + 1.1 mm; n = 5) have the following characters: preanal length 41% TL, head 25%, eye 29% head length; head length and width equal; predorsal length 29% TL, prepelvic length 24%. Branchiostegals 3,4. Except for pelvic fin sep- aration, overall body characters are similar to gobies (Figure 5). Pigmentation features black eyes and approximately six lateral saddle marks on the body. Melanophores are absent on the ventral body surface and fins. Meristics for this, and superficially similar, species are given in Table 2. Discussion The Round Goby is an eurythermal fish adapted to slightly brackish environments in the Caspian and Black seas (Miller 1986). In the Great Lakes basin it is ubiquitous in the littoral zone, including ecotones intolerable to many of our native fishes. Eggs incu- bate in almost static water in Duck Creek, a stream burdened with silt, and at times, blooms of blue- green algae at high temperatures. Concurrently, eggs may develop in the St. Clair River in water of high quality and clarity at temperatures several degrees lower than in small turbid streams in the same catch- ment. The reproductive strategy includes parental guarding of eggs and free embryos. In the St. Clair aquatic ecosystem, age-O0 Round Gobies appear to remain natal stream and shore habitats throughout their first year of growth. Solid objects, unlike sand and other unstable substrates, provide surfaces essential for attachment and refuge for small gobies in flowing water. Reproduction is prolonged and extensive but varies geographically. In their native range, gobies migrate to spawning habitat at 9-26°C and reproduce from April-September in intervals of 3 or 4 weeks (Charle- bois et al. 1997). Similar data for the Great Lakes are lacking, although MacInnis and Corkum (2000) suggest a possible spawning period from April to November in the upper Detroit River. In power plant cooling water discharges, the reproductive period may extend from winter to late summer or beyond, since the endo- crine system is active all year (Charlebois et al. 1997) and water temperatures are constantly >10°C. TABLE 2. Comparison of meristics of several Great Lakes age-0 fishes superficially similar in appearance. Species D1 D2 Pelvic Pectoral Anal Preanal _Postanal fin fin fin fin myomeres myomeres References Etheostoma exile VII-X LO=12 5 12-14 II, 7-8 Heufelder 1982 Etheostoma nigrum VIl-XI 10-14 5 11-12 EWVealO els Bil Heufelder 1982 Etheostoma caeruleum x 12 6 13 ind) 16-17 18-19 Cooper 1979 Cottus bairdi VIL-IX 16-19 I, 3-4 13-17 13-14 9-12 19-21 Heufelder 1982 VIL-VIN =15-17_ I, 3-4 13-15 13 9-10 19-21 This study Cottus cognatus VII-IX 16-19 1, 3-4 12-16 10-14 9-12 19 Heufelder 1982 Proterorhinus marmoratus VI-VII 16-19 1,5 14-15 | is) 11 20 Leslie et al. 2002 Neogobius melanostomus VI 1S =1G) les 18 iatles} 12 19 This study 324 In the study area, batches of about 200-300 eggs represent an absolute fecundity of 800-1200, assuming four spawning episodes. MacInnis and Corkum (2000) estimated an absolute fecundity of 252-1818 eggs based on three spawning episodes from late May to the end of July in the upper Detroit River. Miller (1986) report- ed fecundity of gobies 7-13 cm long as 328-5221 in the Black-Caspian-Azov seas, whilst Kuczyfiski (1995) calculated 2700-3000 eggs in the Gulf of Gdansk. In general, fecundity is lower and eggs slightly smaller (3.7-3.9 mm long, 2.0-2.2 mm wide) than in the Black and Baltic seas, possibly because the spawning popu- lation is younger and fish are smaller (MacInnis and Corkum 2000). Moskal’kova (1967) reared embryos that emerged from the egg at 5.5-5.7 mm and remained in the nest 4-9 days. Gobies do not have a true larval stage (Miller 1986) and hatch with most characteristics of juveniles. However, in the present study, smallest gobies (4.5- 5.0 mm) feature a large yolk sac and incomplete com- plement of fin rays. These fish may have extruded from eggs during the collection process and perhaps are not representative of free embryos. Otherwise, smallest specimens found in coarse gravel were clearly un- guarded, free-living fish. The Round Goby is a cryptic, benthic fish, devoid of air bladder and planktonic stage. As such, early developmental stages are found in and among firm structures. Gobies have smaller caudal and body depth factors than larvae capable of high thrust and acceler- ation, such as Anchovy (Eugraulidae), Alewife (Clu- peidae) or percids (C. A. Timmins, unpublished, 2000). The Round Goby may not be a strong or fast swimmer, but its large head, robust body, and large pectoral fins enhance its manoeuvrability. Age-0 Round and Tubenose gobies did not co-occur in any given habitat in the St. Clair ecosystem, al- though they partitioned habitat in a small area near the mouth of Duck Creek (Leslie and Timmins 1998). Usually, small age-O Round and Tubenose gobies may be separated on the basis of head and body pigmen- tation, which is much lighter in the former. Whereas the Round Goby has a conspicuous black spot on the posterior spines of D1, the Tubenose Goby has an oblique stripe on anterior spines. Further, tubular nos- trils that extend beyond the upper lip are peculiar to the Tubenose Goby. However, most mensural char- acters overlap for these gobies. Thus, small (<6 mm) unpigmented fish may not be separable. Fishes with similar habitat requirements and mor- phology, such as cottids, darters, and gobies sometimes co-occur. Recently hatched Mottled Sculpins are pre- cocious and appear similar to gobies (Figure 5). However, all fins are developed in recently hatched Round Gobies, whereas cottids (and darters) have a complete fin fold and separate pelvic fins, which are expressed some weeks post-hatch. In addition, pre- anal and total myomere counts of cottids and darters THE CANADIAN FIELD-NATURALIST Vol. 118 do not overlap with those of gobies. Fin spine and ray counts and pigmentation patterns also differ among these taxa. The Round Goby has survival advantage over many competitors. It reproduces repeatedly from spring to autumn, the critical life stage is not determined by specific seasonal type or quantity of food, its embryos hatch with almost all external structures elaborated, the young are guarded in nests and it persists in natal habitats. Certainly, few native fishes of the Great Lakes basin are as morphologically and physiologically suit- ed to thrive in comparable environmental conditions. Therefore, geographic range expansion and prolifer- ation of the species seems likely in the Great Lakes Basin. Acknowledgments R. E. Dermott, Department of Fisheries and Oceans, Burlington, Ontario, kindly provided specimens of mottled sculpin. We thank two anonymous reviewers for comments that improved the manuscript. Documents Cited (marked * in text) Skora, K. E. 1997. Neogobius melanostomus. In Baltic Research Network on ecology of marine invasions and introductions. Edited by S. Olenin and D. Daunys. Internet: http: www.ku.lt/nemo/mainmeno.htm. Literature Cited Charlebois, P. M., J. E. Marsden, R. G. Goettel, R. K. Wolfe, D. J. Jude, and S. Rudnicka. 1997. The round goby, Neogobius melanostomus (Pallas) a review of Euro- pean and North American literature. Ilinois-Indiana Sea Grant Program and Illinois Natural History Survey. Special Publication Number 20. Cooper, J. E. 1979. Description of eggs and larvae of Fan- tail (Etheostoma flabellum) and Rainbow (E. caeruleum) darters from Lake Erie tributaries. Transactions of Amer- ican Fisheries Society 108: 446-456. Crossman, E. J., E. Holm, R. Cholmondeley, and K. Tuininga. 1992. First record for Canada of the Rudd, Scardinius erythrophthalmus, and notes on the introduced Round Goby, Neogobius melanostomus. Canadian Field- Naturalist 106: 206-209. Dubbs, D. O. L., and L. D. Corkum. 1996. Behavioural interactions between Round Gobies (Neogobius melanos- tomus) and Mottled Sculpin (Cottus bairdi). Journal of Great Lakes Research 22: 838-844. Heufelder, G. R. 1982. Family Cottidae, sculpins. Identi- fication of larval fishes of the Great Lakes basin with emphasis on the Lake Michigan drainage. Pages 1-774 in Great Lakes Fisheries Commission Special Publication 82- 83. Edited by N. A. Auer. Ann Arbor, Michigan. Jude, D. J., R. H. Reider, and G. R. Smith. 1992. Estab- lishment of Gobtidae in the Great Lakes Basin. Canadian Journal of Fisheries and Aquatic Sciences 49: 416-421. Kalinina, E. M. 1976. [Abstract] Reproduction and early development of gobies from the Azov and Black seas. Kiev. Nauk Dumka, Russia. Koblickaja, A. F. 1981. [Abstract]. Opredielitiel molodii priesnowodnyh rby Moskva. Legkaja i pischevaja pro- myslennost 208. [cited in Charlebois et al. 1997]. 2004 Kuczynski, J. 1995. Babka kragtia Neogobius melanostumus (Pallas 1811) — emigrant z Basenu Pontokaspijskiego w zatoce Gdanskiej. Bulletin Sea Fisheries Institute, Gdynia 2(135): 68-71. Leslie, J. K., and C. A. Timmins. 1998. Fish reproduction and distribution in a small tributary of Lake St. Clair. Can- adian Technical Report of Fisheries and Aquatic Sciences 2253): Leslie, J. K., C. A. Timmins, and R. G. Bonnell. 2002. Postembryonic development of the tubenose goby Protero- rhinus marmoratus Pallas (Gobiidae) in Lake St. Clair and the St. Clair River. Archivum fiir Hydrobiologie 154: 341-352. MacInnis, A. J., and L. D. Corkum. 2000. Fecundity and reproductive season of the Round Goby Neogobius melan- ostomus in the upper Detroit River. Transaction of American Fisheries Society 129: 136-144. Miller, P. J. 1986. Gobiidae. Pages 1019-1095 in Fishes of the North-eastern Mediterranean. Edited by P. J. P. White- LESLIE and TIMMINS: AGE-O ROUND GOBY IN ONTARIO 325 head, M.-L. Bauchot, J.-C. Hureau, J. Nielsen, E. Torto- nese. Volume 3: UNESCO, Paris. Moskal’kova, K. I. 1967. Morpho-ecological specifics of development of the Round Goby, Neogobius melanostomus (Pallas). Pages 48-75 in Morphological and ecological analysis of fish development. Nauka, Moskow. [In Rus- sian]. Nelson, J. S. 1984. Fishes of the World. Second edition. John Wiley and Sons. Toronto. 523 pages. Trautman, M. B. 1981. The fishes of Ohio. Ohio State Uni- versity Press. Columbus, Ohio. 782 pages. Webb, P. W., and Weihs, D. 1986. Functional locomotor morphology of early life history stages of fish. Transactions of American Fisheries Society 115: 115-127. Received 17 February 2003 Accepted 15 November 2004 A Systematic Analysis of the Alpine Saxifrage Complex (Saxifragaceae) in the Canadian Arctic Islands Using Morphology and Chloroplast DNA Data CAROLINE HEALY! and LYNN J. GILLESPIE Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4, Canada ' Present address: Department of Biology, University of Ottawa, P.O. Box 450, Station A, Ottawa, Ontario KIN 6N5 Canada Healy, Caroline, and Lynn J. Gillespie. 2004. A systematic analysis of the alpine saxifrage complex (Saxifragaceae) in the Canadian Arctic Islands using morphology and chloroplast DNA data. Canadian Field Naturalist 118(3): 326-340. The Saxifraga nivalis complex displays significant ecological, morphological and cytological variation. Most European studies suggest that the S$. nivalis complex comprises two distinct species: Saxifraga nivalis sensu stricto and Saxifraga tenuis. However, the presence of intermediate morphotypes, inconsistencies in chromosomal counts and variability in morphological keys and descriptions have led to different taxonomic interpretations of the complex in North America. This study investi- gated the systematics of Canadian Arctic Island members of this complex from 157 specimens using 23 morphological characters. Principal component analysis of the morphological data revealed two adjacent clusters, corresponding to the two taxa and consistent with a close morphological similarity and the presence of hybrids. A preliminary restriction site analysis of five non-coding regions of the chloroplast genome, trnH-trnK, trnT-trnF, trnF-trnV, trnV-rbcL and rbcL-ORF106, was conducted using 21 restriction endonucleases. This analysis indicated a length difference between the trnT-tvnF region of S. nivalis and that of S. tenuis, but no difference in restriction sites for any of the assayed regions. These results confirm that in the Canadian Arctic, the S. nivalis complex consists of two closely related, largely sympatric species, with notable morphological variability, and possible hybrids. Key Words: Alpine Saxifrage, Saxifraga nivalis, Saxifraga tenuis, Canadian Arctic, systematics, morphology, chloroplast DNA restriction site analysis. Saxifraga L., the largest genus in the Saxifragaceae, consists of nearly 400 herbaceous, mostly perennial species (Webb 1993; Soltis et al. 1993, 1996). Saxi- fraga nivalis L. sensu lato, commonly known as the Alpine Saxifrage, inhabits arctic and alpine environ- ments throughout the northern hemisphere. The S. nivalis complex is dispersed throughout northern and alpine Europe, Siberia, arctic and subarctic North America and south in the alpine zone to Arizona and the Gaspé, Quebec (Britton and Brown 1913; Krause and Beamish 1973; Scoggan 1978; Webb 1993). Members of the S. nivalis complex display notable morphological, cytological and ecological variation, which has led to different taxonomic interpretations and classifications. Linnaeus (1753) first described S. nivalis, a name that means snow saxifrage, as these plants are often found growing in snow patch plant communities (Brit- ton and Brown 1913). Later, Wahlenberg (1812) recog- nized S. nivalis var. tenuis Wahlenb. for plants with a notably smaller stature, fewer flowers, smaller fruits and recurved stigmas. Smith (in Lindman 1918) elevat- ed Wahlenberg’s variety to Saxifraga tenuis (Wahlenb.) Harry Sm. However, the taxonomy of the S. nivalis complex remained problematic and the acceptance of S. tenuis as a species was controversial. For instance, Bocher (1938) recognized S. nivalis and S. tenuis as distinct species, but also suspected the presence of races and microspecies within the S. nivalis complex. Polunin (1940) found the two taxa difficult to separate morpho- logically with intermediates and suggested the need for breeding and cytological studies before distinguishing them at the species level. Although accepted as a dis- tinct species in Scandinavia and Greenland, based on samples from Alaska and the Yukon, Hultén (1945) concurred at the time with Polunin in treating the taxon as S. nivalis var. tenuis because of its minimal phenotypic distinction from S. nivalis and its low fre- quency of occurrence within the area of distribution of S. nivalis. However, in the Canadian Arctic, Porsild (1957, 1964) and Porsild and Cody (1980) recognized S. nivalis and S. tenuis as distinct species. Upon fur- ther examination of specimens from Alaska and neigh- boring regions, Hultén (1968) also recognized the two taxa at the species level. More recently Scoggan (1978) and Aiken et al. (1998) treat the two taxa as varieties of S. nivalis. Taxonomic and nomenclatural ambiguity also stemmed from the description of additional varieties and microspecies, as well as the re-emergence of Haworth’s (1812) suggestion to revise Linnaeus’ concept of the genus Saxifraga (reviewed in Spongberg 1972). In 1905, Small divided the genus Saxifraga into 13 genera in North America (in Small and Rydberg 1905; reviewed in Soltis et al. 1996). Small treated S. nivalis as Micranthes nivalis (L.) Small, S. nivalis var. tenuis as Micranthes tenuis (Wahlenb.) Small, and distin- guished an additional species in the Canadian eastern Arctic, Micranthes kumlienii Small. However, subse- quent authors did not recognize the genus Micranthes, 326 2004 and M. kumlienii was treated as belonging to S. nival- is (Polunin 1940; Aiken et al. 2000*). Fernald (1917) distinguished Saxifraga nivalis var. labradorica Fern., a variety localized in Labrador, and suggested that this plant corresponded to Small’s concept of Micranthes tenuis. Subsequently, this variety was treated as a syn- onym of S. nivalis var. tenuis (Scoggan 1978) or as a luxuriant form of the latter (Polunin 1940). In the Gaspé Peninsula, Fernald (1917, 1950) identified another member of the S. nivalis complex, Saxifraga gaspensis Fern. Subsequently, this species was included under S. nivalis var. tenuis (Scoggan 1950, 1978), treated as S. nivalis var. gaspensis (Fern.) Boivin (Boivin 1966), and then considered as small S. nivalis s.s. (Blondeau 1989a). Most recently, Gervais et al. (1995) followed Fernald in treating the Gaspé population as a distinct species. In Alaska, Hultén (1968) considered S. nivalis as highly variable, recognizing S. nivalis var. tenuis, and describing S. nivalis var. rufopilosa Hultén for plants characterized by “densely pubescent, rufous- haired leaves and red petals” (page 579) found in Alaska and the Yukon. Porsild (1957, map page 188) appeared to treat S. rufopilosa under S. tenuis ac- cording to his map of that species (see also Porsild and Cody 1980, map page 403; Cody 1996). Later, Porsild (1975) recognized the latter as the distinct species, S. rufopilosa (Hult.) A. E. Porsild, and suggested that it was more closely related to S. tenuis than S. nivalis. Cody (1996) also recognized S. rufopilosa as a distinct species and considered S. tenuis as not present in the Yukon. In 1978, Scoggan’s concept of S. nivalis var. tenuis encompassed S. tenuis, S. gaspensis, S. nivalis var. labradorica and S. nivalis var. rufopilosa. Based on cytology and morphology, Love (1983) and Webb (1964, 1993) discerned the species S. nivalis (2n = 60) and S. tenuis (2n = 20) in Europe. The taxo- nomic ambiguity regarding the S. nivalis complex in North America (Aiken et al. 2000*) appears to stem from the following factors: (1) several morphological characters used to distinguish the species in Europe do not hold up in North America; (2) taxa can be found growing together; and (3) intermediate morphotypes, as well as variation in chromosomal counts suggest hybridization. Various cytological studies of material from North America have demonstrated cytological diversity within the S. nivalis complex with chromo- somal counts of 2n = 20, 40, c. 56, 58, 60 or c. 60. While most support the above European counts for S. nivalis and S. tenuis (Mosquin and Hayley 1966; Hed- berg 1967; Johnson and Packer 1968; Packer and McPherson 1974; Love and Love 1975), several studies found intermediate chromosome counts. Krause and Beamish (1973) provide counts for plants of the S. nivalis-tenuis complex from the Yukon (2n = 20, 40), British Columbia (60) and Idaho (20), which they were unable to identify to species. Intermediate chromosome counts have also been found in the Russian Arctic in a plant described as S. nivalis x S. tenuis (Devyatov et HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 327 al. 1997). In the Gaspé Peninsula S. gaspensis is con- sidered to be a putative endemic hybrid based on chro- mosome number (2n = 40) and apparent intermediate morphology (Gervais et al. 1995). Further confusion results from the variance and discrepancies among the morphological keys used in the identification of this complex and related species, such as S. foliolosa R. Br., which are occasionally misidentified as members of the S. nivalis complex. Consequently, accurate and consistent identification of North American plants belonging to this complex can be problematic. The aim of this study was to investigate the distinc- tiveness of S. nivalis and S. tenuis in the Canadian Arctic Islands based on morphological and molecular differences, and to help reassess the taxonomic status of S. tenuis. Material and Methods Morphological Analysis Specimens Representative specimens from the Canadian Arc- tic were selected from the Canadian National Herb- arium, Canadian Museum of Nature (CAN), and the Agriculture and Agri-foods Canada Herbarium, Cen- tral Experimental Farm, Ottawa (DAO). Morphological data were gathered from observations and measure- ments on 157 herbarium sheets (listed in Appendix I), including two chromosome count vouchers (S. nivalis, 2n = 60, Calder et al. DAO 24148: S.. tenuis, 2n = 20, Calder et al. DAO 24149). One plant per sheet was sampled, except when sheets comprised mixed collections or included DNA vouchers, and then multiple plants per sheet were sampled. A total of 200 plants were sampled, 122 S. nivalis and 78 S. tenuis. More specimens of S. nivalis were available for study than of S. tenuis. Where identifications of exam- ined specimens were judged questionable, specimens were re-identified based on a combination of published keys and preliminary results. Additionally, identifica- tions of many S. tenuis specimens at CAN were sub- sequently confirmed for the Panarctic Flora Project (E. Reidar, 2001, 2002). Morphological characters Characters used in descriptions and keys of the S. nivalis complex (Simmons 1906; Polunin 1940; Porsild 1957; Love 1983; Blondeau 1989a, 1989b; Webb 1993; Aiken et al. 1998, 2000*) were reduced to a set of 30 characters based on ease of observation or meas- urement on dried specimens as well as their taxonom- ic significance. Of these, 23 were selected for statis- tical analysis (Table 1). Petal length was excluded from all analyses due to difficulties in measuring petals that were often folded and partly included within the calyx. The qualitative characters, stem colour, petal shape, prominence of leafy bract on stem, leaf blade shape, leaf blade base shape and apex shape, were excluded due to variation within a plant or difficulties in determining discrete states. 328 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Morphological characters used in analyses of the Saxifraga nivalis complex in the Canadian Arctic. Quantitative characters: Hair length on leaf lower surface (mm): UNDHRLGTH Hair length along leaf margin (mm): MRGHRLGTH SOC eE ES ON Calipers) Ei Plant height (cm): HEIGHT; length from basal rosette to top of inflorescence Stem width (mm): STWIDTH; measured midway along lower half of stem Stem hair density (0, glabrous to 4, dense): STHRDENSITY Stem hair length (mm): STHRLENGTH; measured midway along lower half of stem. Leaf blade length (cm): LFLENGTH; measurement made on the largest leaf Leaf blade width (em): LFWIDTH; measurement made on the largest leaf Number of teeth on leaf: TEETH; count made on the largest leaf 10. Tooth width (mm): TTHWIDTH; measurement made on largest tooth on the largest leaf 11. Tooth length (mm): TTHLENGTH; measurement made on largest tooth on the largest leaf 12. Petal width (mm): PTWIDTH; measurement made on the largest petal Qualitative characters: Stem hair texture (short fine; long coarse) Tooth apex shape (rounded; pointed) Hair colour on leaf lower surface (rust; white) Hair colour on leaf margin (rust; white) Inflorescence density (open; tight cluster) 10. Style shape (curved; straight or essentially so) SEN SION a eS 11. Petal colour (pink or essentially so; white or essentially so) The software package DELTA (Dallwitz et al. 1993) and the character set for the “Flora of the Canadian Arctic Archipelago” (Aiken et al. 2002*) were used to generate preliminary taxon descriptions. These descrip- tions were then modified to read more easily, to ex- clude irrelevant characters, and to include important diagnostic characters not included in the Flora char- acter set. Statistical analysis All statistical analyses were performed with SYSTAT version 8.0. For each quantitative trait, the means for S. tenuis and S. nivalis specimens were compared by performing t-tests. Separate variance values were cal- culated for each taxon. T-tests were conducted on both raw and log-transformed data. A principal component analysis (PCA) using the 12 quantitative characters in- dicated in Table 1 was conducted to visualize possible distinctiveness between S. nivalis and S. tenuis. The character MRGHLGTH was excluded from some PCA analyses due to a large amount of missing data in S. tenuis, which sometimes lacked marginal hairs. The PCA analysis excluded all specimens with missing values. A discriminant function analysis was also per- formed on the quantitative data. For the 11 selected qualitative characters (Table 1) the percentages of spec- imens having each character state was determined for each taxon. To determine if there was any geographical pattern to the morphological variation in S. tenuis, principal component analysis was performed only on the S. tenuis quantitative data (excluding MRGHLGTH). This analysis also included five specimens from Alas- Stem hair colour (rust-coloured; mixed rust and white; white) Regularity of teeth (even in size; variable in size smaller towards leaf blade apex) Hair presence on leaf lower surface (moderate to dense; sparse; absent) Hair presence on leaf margin (moderate to dense; sparse; absent) ka (Calder 6107, 6511, 6249 DAO) and northern British Colombia (Calder & Kukkonen 28151 DAO, Taylor et al. 1204 DAO) not included in the larger analysis. Individuals were coded as occurring in one of three regions: high arctic (67), eastern arctic (11) or western montane (5). Chloroplast DNA analysis Plant material and DNA extraction A total of 15 plants, for which silica gel dried mate- rial was available, were selected for molecular analysis: seven individuals of S. nivalis (Gillespie 6165, 6675-1, 6826a-1, 6826a-2, 6977-2, 6984-5; Aiken 98-055-3), seven S. tenuis (Gillespie 6728-1, 6807-1, 6807-3, 6825-2, 6870-2, 6877-1, 6881-3), and one S. foliolosa (Gillespie 6841) (Appendix 1). The latter species was selected to provide perspective on the chloroplast DNA (cpDNA) data, based on its close phylogenetic relationship to the S. nivalis complex (Soltis et al. 1996). DNA was extracted following Doyle and Doyle’s (1990) CTAB total DNA extraction procedure as modified by Gillespie et al. (1997). Restriction site analysis DNA extracts were PCR amplified using five pairs of universal primers for non-coding regions of chloro- plast DNA. The five regions are: tmmH-tmmK (Demesure et al. 1995), trnT-trnF (Taberlet et al. 1991), trnF-trnV (Dumoulin-Lapegue et al. 1997), trnV-rbcL (Dumou- lin-Lapegue et al. 1997), and rbcL-ORF106 (Arnold et al. 1991). Amplification reaction mix and programs follow Gillespie and Boles (2001). 2004 HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 329 TABLE 2. Descriptive statistics and t-test comparing the means of S. nivalis and S. tenuis at the 95% confidence level for 12 quantitative morphological characters. Range Difference Character Taxon N Mean (+SD) Min Max in means t value df p S. nivalis 122 OFZ) CE3'3i7) af AsO) Plant height (cm) 4.66 13.68 176.2 0.000 S. tenuis 78 4.40 (+1.40) 1.8 8.5 S. nivalis 122 1.37 (+0.35) 0.7 DES Stem width (mm) 0.82 23.06 178.6 0.000 S. tenuis 78 0.56 (+0.15) 0.3 1.0 S. nivalis 122 3.2 (+ 0.70) 1 4 Stem hair density 2.0 24.94 205.0 0.000 S. tenuis 78 1.2 (+ 0.36) 1 3} S. nivalis 1:22 3.2 (+ 0.70) 1 4 Stem hair length (mm) 0.67 35.45 188.5 0.000 S. tenuis Tal 0.19 (+0.09) 0.1 0.5 S. nivalis 119 0.96 (+0.34) 0.4 Dil Leaf blade width (cm) 0.51 14.81 161.2 0.000 S. tenuis 77-044 (+0.13) 0:25 570!8 S. nivalis 119 1.38 (+ 0.50) 0.6 3.6 Leaf blade length (cm) 0.69 13.51 166.5 0.000 S. tenuis TL 0.67 (+ 0.18) 0.4 IES Hair length on lower S. nivalis 120 0.38 (+0.13) 0.2 12, leaf surface (mm) 0.10 725) 97:08 70/000 S. tenuis 78 0.27 (+0.06) 0.1 0.5 Hair length on S. nivalis 121 0.19 (+ 0.07) 0.1 0.4 leaf margin (mm) 0.07 8.47 167.4 0.000 S. tenuis 55% 0.12 (+ 0.04) 0.1 0.2 Number of teeth S. nivalis 118 10.3 (+1.74) 7 14 on leaf blade 2.8 16.63 167.8 0.000 S. tenuis 78 7.5 (40.73) 6 9 S. nivalis 118 1.17 (+ 0.42) 0.5 3h) Tooth width (mm) 0.37 7.86 195.6 0.000 S. tenuis 78 0.80 (+ 0.24) 0.5 1.6 S. nivalis 118 1.52 (+0.48) 0.8 B82. Tooth length (mm) 0.62 11.84 190.5 0.000 S. tenuis 78 0.90 (+0.26) 0.4 1.8 S. nivalis 120 1.12 (+0.24) 0.7 1.8 Petal width (mm) 0.40 14.16 202.0 0.000 S. tenuis al 0.72 (+0.16) 0.5 il * Marginal hair was not present in all S. tenuis PCR products were single digested with twenty one Apal, BamHI, Bgil, BglII, Dral, EcoRI, EcoRV, Hincll, restriction endonucleases according to manufacturer’s PstI, XhoI. To separate restriction fragments, 8 uL of recommendations. These included nine 4-base pair the reaction mixture and 2 uL of buffer-dye mix were (bp) cutting restriction enzymes: Alul, Haelll, Hhal, run on 1.1% agarose gels with ethidium bromide at Mspl, Rsal, Ddel, Hinfl, Sau96], Taql; two 5bp cutting 38 V for 2-3hrs. The restriction fragment patterns were restriction enzymes: BstOl, SinI; and ten 6bp enzymes: visualized and photographed under ultraviolet light. 330 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 3. Distribution of qualitative morphological character states in Saxifraga nivalis and S. tenuis in the Canadian Arctic. Numbers represent percentage of individuals having a particular character state. Character Character state Stem hair colour rust mixed white Stem hair texture short fine long coarse Tooth apex shape rounded pointed Regularity of teeth even-sized variable Hairs presence on leaf lower surface moderate-dense sparse absent Hair colour on leaf lower surface rust white Hair presence on leaf margin moderate-dense sparse absent Hair colour on leaf margin rust white Inflorescence density open tight cluster Style shape curved straight Petal colour pink white Lengths of total amplified product and restriction frag- ments were estimated by comparison with known 100 bp ladder DNA markers. Results Morphological Analysis To determine whether the means of the examined quantitative characters were different between S. nivalis and S. tenuis, t-tests were performed (Table 2). The analysis revealed a significant difference in means be- tween the two taxa for all characters examined. Overall, plants belonging to S. nivalis are more robust than S. tenuis as implied by higher means in both plant height and stem width observed in S. nivalis. Similarly, leaves of S. nivalis are on average longer, wider and have longer, wider and more numerous marginal teeth than S. tenuis. Also, hairs on the stems are longer and denser and hairs on the lower surface and margin of leaves are on average longer in S. nivalis. Petal width was sig- nificantly greater in S. nivalis than in S. tenuis. How- ever, the two taxa have notably different variances, with S. nivalis having a higher variance than S. tenuis for all quantitative characters examined. Considering the very low probability values and the robustness of t-tests for samples of this size, the differences in means can be considered as meaningful. Analyses were also performed on log-transformed data, with no significant differences. The frequency of occurrence of states of qualitative characters differed considerably between the two taxa S. nivalis S. tenuis 3} 74.4 0 25.6 98.7 0 1.6 100 98.4 0 DNS 100 97.5 0 18.3 87.2 81.7 12.8 91.7 84.6 es 15.4 0.8 0 40.0 79.5 60.0 20.5 91.7 10.3 8.3 60.3 0 29.5 20.7 60.0 79.3 40.0 1.8 97.1 98.2 2.9 25.9 84.7 74.1 15}3) 14.9 81.8 85.1 18.2 for most characters (Table 3). The characters, stem hair colour and texture, tooth apex shape and inflorescence density, differed significantly between the two taxa, with less than a five percent overlap in states. Several characters, regularity of teeth, petal colour and style shape, exhibited an overlap in states of 10-26% between the two taxa. In contrast, one character, hair presence on lower leaf surface, displayed very similar frequen- cies of states between S. nivalis and S. tenuis, with hairs present in over 95% of the specimens examined in both taxa, although density was sometimes more sparse in S. tenuis. The remaining characters displayed considerable overlap in states between the taxa. The principal component analysis of quantitative characters (excluding MRGHLGTH) revealed two clusters, positioned adjacent to each other with little overlap (Figure 1). Factor loading for the analysis was STWIDTH, LFWIDTH, LFLENGTH, STHRLENGTH, STHRDENSITY, HEIGHT, TEETH, PIWIDTH and TTHLENGTH for factor 1, and TTHWIDTH and UNDHRLGTH for factor 2, with 61% and 10% of total variance explained by factors | and 2, respectively. As noted above, there is considerable variation among S. nivalis plants for most examined characters. This can be observed on the graph through the large oblique spread of S. nivalis compared to S. tenuis. The PCA including the character MRGHLGTH produced very similar results (although displayed fewer individuals). The discriminant function analysis performed on the same data distinguished between the two taxa 100% 2004 5 4 3 2) ORO Q 1 On S Boo GS ‘OP 2 © O <= LL 9) ES -4 5 5 -1 0 { : : FACTOR(1) FIGURE |. Principal component analysis on the quantitative morphological data set for the Saxifraga nivalis com- plex in the Canadian Arctic. Circles represent individ- uals of S. nivalis, crosses represent S. tenuis. of the time, with one dimension accounting for the variation. The PCA analysis examining geographical patterns to morphological variation in S. tenuis revealed very little geographical structure, with eastern arctic and high arctic plants almost completely overlapping. HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 33] Western montane plants separated from the majority of arctic plants on a combination of both factors (with LFWIDTH, LFLENGTH, and TEETH, loading high on factor one, HEIGHT on factor two), but overlapped considerably with several eastern arctic plants. A dis- criminant function analysis performed on the three geographic groups produced similar results (correctly placing the western montane plants with 100% reli- ability, high arctic plants with 86% reliability, and eastern arctic plants with only 42% reliability). Chloroplast DNA analysis The trnH-trnK amplicons were estimated at 1600 base pairs (bp) for S. nivalis and S. tenuis and 1700 bp for S. foliolosa. The three largest regions, trnF-trnV, trnV-rbcL and rbcL-ORF106, were approximately 3000 bp in length in all three taxa. Interestingly, the tnT-trnF region was estimated to be 1800 bp, 1850 bp and 2000 bp in length for S. nivalis, S. tenuis and S. foliolosa, respectively. Sixteen out of 105 enzyme restriction assays showed restriction site differences between S. foliolosa and the S. nivalis complex, including three site differences in the trnH-trnK region, three in the rbcL-ORF106 region, three in the trnV-rbcL region, five in the trnF- trnV region and two in the trnT-trnF region (Table 4). No restriction site differences were detected bet- ween S. nivalis and S. tenuis. However, digestion of the trnT-trnF region with the restriction enzyme Alul showed clearly the length difference between S. nivalis and S. tenuis (Figure 2). Table 4. Data matrix of restriction site characters for S. nivalis (N), S. tenuis (T) and S. foliolosa (F). “0” represents absence of restriction site; “1” represents presence of restriction site. trnH-trnK rbcL-ORF106 NTF INGE Alu I eal ie 1b kl Apal 000 000 BamHI 000 eeleall Bell 000 000 Bel Il 000 Ave let BstO I 001 Ethel Dral 001 lO EcoRI 000 Ikea EcoR V 000 ae al Hae It Hee al 001 Hha lI 000 000 Hinf I lbw eal Msp I ha al hiked j sO lal al apna iL hal 000 000 000 Write al Pal il ilies al ie ihel oak il 000 001 000 000 trnT-trnF NTF trnF-trnV INiglge trnV-rbcL NTE DR RP eB RrP ORR RRP rR OOOO COOCCrF ORB Re RP ROR RFP RRP rR OCOCOCOCOCCOOF CORR RRB BS OR RP RB Re RO OrR KR OOF OF eee Re RK Or Fer COOCOCcoocoocoorcoor [St ee a) ees) (5) oa ea) (Fea) fos OPER ROORHEH OHM OH COOCOOH SPSS LOLS shore Oise SO oro Sc See =O Ores Orr Ore SO Ora Ore SS OororcjooroqooqococrreKScocorf Oo Oo Nw THE CANADIAN FIELD-NATURALIST Vol. 118 MNNTTTTNNT F FiGuRE 2. Digestion of the chloroplast DNA region trnT-trnF with restriction enzyme Alul for Saxifraga nivalis (N), S. tenuis (T) and S. foliolosa (F). The first lane contains a 100 bp DNA size marker (M). Note absence of the upper band on S. nivalis. Discussion The principal component analysis confirms that the S. nivalis complex in the Canadian Arctic is represent- ed by two principal morphotypes, appearing as adjacent clusters on the PCA graph. The several plants occu- pying an intermediate position between the two clus- ters displayed a somewhat intermediate morphology, but nevertheless could readily be assigned to one of the two taxa. For example, several specimens of S. nivalis from Prince Patrick Island (e.g., Gillespie & Consaul 6869-4, 6926a-3) were very small with slender flowering stems, characters typical of S. tenuis, but otherwise had the morphological characteristics of S. nivalis. Although character states or ranges overlapped for the majority of characters examined, differences between the two taxa were statistically significant for all quantitative and for three qualitative characters. Morphological variability A significant amount of variance was observed for most characters measured. Since the S. nivalis complex is considered an environmental indicator (Aiken et al. 1998, 2000*), the variability observed may be due to phenotypic plasticity and variation in environment. For instance, S. nivalis plants growing in harsher habitats are usually shorter and less robust (e.g., the very short plant shown in Figure 3) than those grow- ing in more nourishing environments. Also plants of S. nivalis may be taller with more slender stems in shady microhabitats as a result of etiolation. Another contributor to variance is the time of collection. Since stems elongate in the fruiting stage, particularly in S. nivalis, plants collected later in the season are more likely to be taller than their younger counterparts. However, most examined collections of S. nivalis and S. tenuis made at the same time and site displayed significant height differences on average between the two taxa. Leaf characters, such as leaf size and teeth number and size, showed much variation both within and between the two taxa. Although S. tenuis is usu- ally found in wetter habitats than S. nivalis, the latter species in particular occupies a broad range of habitats and the two species sometimes co-occur in mixed populations. Some of the morphological variation in the complex, particularly within S. nivalis, may reflect this variation in habitat (Blondeau and Cayouette 2002; Aiken et al. 2000*). Such high intraspecific variation is apparently typical of Saxifrages, particularly in arctic and alpine environments, and several studies have demonstrated that such variation may not be of taxo- nomic significance (Soltis et al. 1996; Brysting et al. 1996). The coloration of petals also varied within each taxon, but season and environment can be a large con- HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX pws ki, y ~ FIGURE 3. Saxifraga nivalis complex in the Canadian Arctic. A. S. tenuis (Gillespie & Consaul 6978-2), habit. B. S. nivalis (Gillespie and Consaul 6977-1), habit. C. Comparison of basal rosette of leaves of S. tenuis (on left, Gillespie & Consaul 6978-2) and S. nivalis (on right, Gillespie and Consaul 6977-1). A and B are the same scale. 334 tributor to petal coloration. Plants growing in more open, sunnier environments tend to have more pink or dark red anthocyanin pigment throughout the plant, including the petals. White petals may also sometimes turn pinkish with age. In contrast, little variation was observed within each taxon for the texture and color- ation of stem hair, and these characters were signifi- cantly different between taxa. Among traits traditionally used for identification of the S. nivalis complex, many displayed considerable intrataxon variance, with ranges often overlapping be- tween taxa. Our results also indicate a greater degree of variability within S. nivalis than in S. tenuis in the Canadian Arctic. This pattern of variation results in a somewhat obscure morphological boundary between the two species and has led to difficulties in construct- ing good identification keys. Characters and taxonomic identification Despite the considerable intraspecific variation, most commonly used identification characters were significantly different between the two taxa. Indeed, plants of S. nivalis were overall larger and more robust than S. tenuis, and the latter often had a more reddish tinge on the petals and stem hair, as previously found by numerous authors (Simmons 1906; Polunin 1940; Porsild 1957; Boécher et al. 1968; Léve 1983; Blon- deau 1989a; Webb 1993). Flowering stem width was an excellent identifica- tion character apart from a few individuals, with S. tenuis characterized by a width of 1 mm or less, con- sistent with Webb’s (1993) use of it as a key character. Consistently used by various botanists (e.g., Webb 1993), stem hair characters were among the most use- ful for identification. Saxifraga nivalis usually has a much more conspicuous, longer, denser, coarse crisped white vestiture, while S. tenuis has a less conspicuous, shorter, finer, usually glandular pubescence. Canadian Arctic S. tenuis was found to always have stem hairs, thus not conforming to descriptions of the taxon in Greenland and Svalbard as having mostly glabrous stems (Bécher et al. 1968, 1978; Ro@nning 1996). Generally, S. nivalis plants were taller than S. tenuis, but due to its variance, height was not useful for iden- tification. Saxifraga nivalis had more numerous teeth on the leaves than S. tenuis, and the latter had rounder ob- tuse teeth versus more pointed, as suggested in the literature (e.g., B6cher et al. 1968), The width and length of S. tenuis teeth were also found to be less than those observed on S. nivalis, but, this is not sur- prising considering that S. nivalis had on average longer and wider leaves. Interestingly, S$. nivalis had a notably less regular dentition than S. tenuis in regards to the size of the teeth. The former generally had small teeth at the base with teeth becoming larger towards the apex, whereas S. tenuis had more evenly sized teeth. Although size of the largest leaf differed significantly between the two taxa, leaf size characters THE CANADIAN FIELD-NATURALIST Vol. 118 make poor diagnostic characters due to overlap and age related variation. Also, leaf shape varied tremen- dously and thus should not be considered a good discriminating character. However, the leaf apices of S. tenuis did tend to be more rounded than those of S. nivalis. Inflorescence compactness provides a useful identi- fication character, with S. nivalis having one or more compact head-like clusters and S. tenuis a more open inflorescence (as described in Porsild 1957). Flower pedicels, especially of the lowermost flowers, are longer in S. tenuis than in S. nivalis. Since pedicels of the latter may elongate somewhat in fruit, this difference may be obscured later in the season. Although not con- stant, petals tend to be whiter, wider and ovate in S. nivalis and pink, narrower, more elongated and rounder at the apex in S. tenuis. BOcher (1938), Bocher et al. (1968), and more recently Love (1983) and Rgnning (1996) state that S. nivalis has straight or slightly spreading styles, while S. tenuis has styles that curve strongly downwards. This was not always the case in Canadian Arctic specimens we examined. The way specimens were pressed and the time of collection might be partly responsible for the variation in this character within the two species. Other characters showed variation that was not use- ful in separating the two taxa. While S. nivalis usually had hairs on leaf margins, this character was also found to a lesser degree in S. tenuis. Most specimens examined had a purplish or reddish stem, thus not specific to S. tenuis. Also, the flowering stems of both taxa had on occasion a reduced leaf (bract) or two near the inflorescence, although usually more prominent in S. tenuis. The presence of rust-coloured hairs on the lower surface of the leaves of S. tenuis has been commonly used to distinguish it from S. nivalis in the Canadian Arctic, and was used as a key character by several authors (Porsild 1957; Scoggan 1978; Porsild and Cody 1980). However, this study reveals that hairs are present on the leaf undersurface in over 99% of the plants examined of both species in the Canadian Arctic, and that hairs may be rust-coloured in both species, about 42% of S. nivalis plants examined and 74% of S. tenuis plants. This error may have resulted from Porsild’s inclusion of S. rufopilosa within his concept of S. tenuis (distribution maps of S. tenuis in Porsild (1957) and Porsild and Cody (1980) appear to have included S. rufopilosa). Saxifraga rufopilosa, a species now considered to be endemic to unglaciated areas of the Yukon and Alaska, is characterized by a mat of long crinkly rusty hairs on the leaf undersurface (Cody 1996), much longer and denser than in either S. tenuis or S. nivalis. Whatever its origin, use of rust-coloured hair presence as a key diagnostic character in identify- ing Canadian Arctic material of the S. nivalis complex resulted in many misidentified specimens and much confusion as to the identity of S. tenuis. 2004. HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 335 Sy e Specimen data (CAN, DAO) ‘\ == Adjacent species range from S Porsild and Cody (1980) we e Specimen data (CAN, DAO) -\se=, Adjacent species range from OB Porsild and Cody (1980) Ficure 4. Distribution of the Saxifraga nivalis complex in the Canadian Arctic. (above) S. nivalis. (below) S. tenuis. Shading indicates approximate distribution outside of the Arctic Islands (from Porsild and Cody 1980, and CAN and DAO herbarium records). 336 The many herbarium collections having both S. nivalis and S. tenuis plants on the same sheet and under the same number (e.g., Bruggeman 358, Savile 4562, Macdonald 150, Calder et al. 24149) shows just how problematic identification of this complex in the Cana- dian Arctic has been. About 20% of herbarium speci- mens identified as S. tenuis at both CAN and DAO were in fact mixed collections including both S. tenuis and S. nivalis plants, or less often including small plants of S. foliolosa or S. hieracifolia Waldst. & Kit. Pre- vious identification problems of the S. nivalis complex in the Canadian Arctic may also have been due to misinterpretation of key characters. For example, S. nivalis with several flower heads could be keyed out under lax (open) inflorescence. Although S. nivalis and S. tenuis can be readily dis- tinguished in the Canadian Arctic, a small percentage of specimens had one or a few characteristics of the other species. For example, some high arctic S. nivalis had very slender stems or fruit with strongly recurved styles. While this may be part of the natural variation within S. nivalis, it may also suggest a low level of hybridization. Some of the overlap in quantitative char- acter states shown in our morphological analyses may also be the result of hybridization. Recent cytological studies have suggested the existence of S. nivalis-S. tenuis hybrids (2n = 40) in regions of the Russian Arctic and the Yukon where S. nivalis and S. tenuis are found growing together (Krause and Beamish 1973; Devyatov et al. 1997). The population in the Gaspé Peninsula has the same intermediate chromosome num- ber and may represent a stabilized relictual hybrid in an area whether neither parent now occurs (Gervais et al. 1995). In addition Polunin (1959) suggested that hybridization may also occur between the S. nivalis complex and other closely related species, such as S. hieracifolia. Geographical variation in Saxifraga tenuis Savile (1961) proposed that North American S. tenuis may be divided into three disjunct and morphologi- cally distinct “populations.” The first is a high arctic population composed of plants identical to material from the north of Greenland and hypothesized to have spread from there following the Pleistocene. The second population is apparently found mostly along Hudson Straight in southern Baffin Island, northern- most Quebec and Southampton Island, but also scattered northward to northeast Baffin Island. Savile treated plants in the Gaspé Peninsula of Quebec, con- sidered by some as a separate species S. gaspensis, as part of this population. Plants of this population appar- ently tend to be larger and less purplish in colour than in the high arctic. The third highly variable population, found in Alaska and the Yukon, is usually character- ized by a highly purplish pigmentation and whitish scape hairs, and was suggested to be intermediate between S. nivalis and high arctic S. tenuis. THE CANADIAN FIELD-NATURALIST Vol. 118 Our analysis indicates that eastern and high arctic plants of S. tenuis cannot be distinguished based on examined quantitative characters, while western mon- tane plants (based on our small sample size) may be differentiated from most arctic plants based on larger size. Since Savile (1961) used mostly qualitative char- acters to distinguish the three groups, further analyses based on both qualitative and quantitative characters and including more western montane and eastern arctic specimens should be made to further test his hypoth- esis. Chloroplast DNA analysis The five non-coding cpDNA regions of S. nivalis and S. tenuis were identical for restriction sites as- sayed. Considering the conservative nature of cpDNA evolution, it is not surprising to observe such similar- ity, with closely related species often having identical cpDNA restriction site profiles (Olmstead and Palmer 1994). However, S. nivalis and S. tenuis displayed an estimated 50 bp length difference in the trnT-trnF region. This difference was observed both among plants from the same site and from different locations. This suggests that the size variation in the t7T-trnF region is not background variation among populations or a phenomenon limited to a particular population, but represents a molecular difference between S. nivalis and S. tenuis, either a deletion event in S. nivalis or an insertion event in S. tenuis. Sequencing of the tnT-trnF region would prove useful in specifying the nature of this size difference. The RFLP analysis re- vealed molecular differences for 16 of 105 restriction enzyme assays between S. foliolosa and the other two species. These results are similar to a study of Saxifraga based on the cpDNA matK sequences (Soltis et al. 1996; plus sequences in Genbank), in which S. nivalis and S. tenuis would appear to be more closely related to each other than to S. foliolosa. Based on comparison of the Genbank sequences, the single plant of S. fenuis differed from S. nivalis in four nucleotide substitu- tions, and from S. foliolosa in 79 substitutions. Implications Our results suggest that S. nivalis and S. tenuis should be recognized as distinct taxa, and are most appropriately recognized at the species level following Porsild (1957), Porsild and Cody (1980), and Webb (1993). The two taxa are distinguishable at both the molecular and cytological level and have morpho- logical differences that correspond to these differences. Although no morphological characters give 100% sep- aration, many quantitative and qualitative characters show statistically significant differences between the two taxa. The two taxa overlap in geographic distribu- tion in the Canadian Arctic, with the range of the less common S. tenuis included within that of S. nivalis. Although ranges of habitat preferences differ, there is considerable overlap, with the two taxa sometimes oc- curring in mixed populations. Lack of geographical 2004 separation and considerable habitat overlap make recognition at the subspecific level inappropriate, while the morphological distinctiveness of the two taxa would make recognition at the varietal level inappropriate. Saxifraga nivalis complex in the Canadian Arctic: key to species A. Inflorescence one to several dense head-like clusters of numerous flowers; petals white or essentially so (sometimes becoming pink with age); flowering stem (0.5-) 1-2.5 mm wide, moderately to densely hairy, with conspicuous long coarse white hairs; plant usu- ally robust in appearance S. nivalis B. Inflorescence an open cyme of fewer flowers, flowers on distinct pedicels; petals pink or less often white; flowering stem 0.3-1 mm wide, usually sparsely hairy, with short fine hairs that are usually inconspicuous; plant delicate in appearance S. tenuis Saxifraga nivalis L. Plants perennial herbs, (2) 5—17 cm high, with a basal ros- ette of leaves. Leaves simple, alternate, all basal in a rosette, evergreen; petiole usually distinct, 0.2-2.5 mm long. Leaf blade ovate, obovate, orbicular or spatulate, 6-36 mm long (mean 14 mm), 4-21 mm wide (mean 10 mm), slightly leath- ery; green above, green or reddish-purple beneath, base obtuse and usually abruptly narrowed at petiole; apex obtuse, rounded or rarely acute; appearing single-veined or with inconspic- uous lateral veins; upper surface glabrous; lower surface sparsely to densely hairy; hairs 0.2-1.2 mm long, rust-coloured, whitish or translucent; margins coarsely serrate or crenate, usually with rust-coloured or whitish hairs; teeth 7-14, usually increasing in size towards blade apex, tips usually pointed. Flowering stem erect, (0.5) 1-2.5 mm wide, green to purple, moderately to densely hairy, with hairs conspicuous, (0.5) 0.7-1.2 mm long, crisped (crinkly and irregularly wavy), white or translucent with tips and crosswalls white or sometimes rust-coloured or purplish, the tips sometimes glandular, usually leafless or occasionally with reduced leaves (bracts) close to the inflorescence. Inflorescence a dense head-like cluster of numerous flowers, sometimes with 1-2 smaller lateral clusters on short to long peduncles; flowers on very short pedicels, subtended by bracts. Flowers: sepals 5, free, green or purple, glabrous; petals 5, ovate or elliptic, 0.7-1.8 mm wide, equal to or longer than the sepals, white or cream-coloured, some- times with reddish apex, mostly obtuse at apex; stamens 10; gynoecium partly inferior; carpels 2, partly fused; styles 2, free; placentation axile; ovules numerous. Fruit a capsule, spherical in lower half, with straight or slightly divergent free carpels above, 34 mm long, 4.5—5.5 mm wide, glabrous, dry, dehiscent; styles straight or slightly divergent, persistent; sepals persistent in fruit. Seeds numerous, 0.5—1 mm long, yellowish brown, with surface verrucose. Figure 3b, c; map 4a. Saxifraga tenuis (Wahlenb.) Harry Sm. Plants perennial herbs, 2-9 cm high, with a basal rosette of leaves. Leaves simple, alternate, all basal in a rosette, ever- green; petiole usually indistinct, ~0.5—2.5 mm long. Leaf blade narrowly ovate, obovate or spatulate, 4-15 mm long (mean 7 mm), 2-8 mm wide (mean 5 mm); slightly leathery, green to reddish-purple above, reddish-purple beneath; base obtuse or broadly attenuate and decurrent onto petiole; apex obtuse, rounded or rarely acute; appearing single-veined or with inconspicuous lateral veins; upper surface glabrous; lower HEALY AND GILLESPIE: ALPINE SAXIFRAGE COMPLEX 337 surface usually sparsely hairy, with hairs 0.1-0.4 mm long, rust-coloured, whitish or translucent; margins coarsely serrate or crenate, with sparse rust-coloured or whitish hairs or some- times glabrous; teeth 6-9, usually evenly sized, tips usually rounded. Flowering stem erect, 0.3-1 mm wide, usually dark purple, sparsely hairy, with hairs mostly inconspicuous, 0.1- 0.3 (0.5) mm long, fine, crinkled or straight, rust-coloured to translucent, often with purplish crosswalls and tips, the tips usually glandular; sometimes with a reduced leaf (bract) on the upper part of the stem. Inflorescence an open head of few flowers; flowers subtended by bracts; lower-most flowers often long-pedicellate and each subtended by a conspicuous bract. Flowers: sepals 5, free, green or purple, glabrous; petals 5, ovate or elliptic, 0.5-1.1 mm wide, equal to or longer than the sepals, usually pink, sometimes white, often with reddish apex, often rounded at apex; stamens 10; gynoecium partly inferior; carpels 2, partly fused; styles 2, free; placentation axile; ovules numerous. Fruit a capsule, spherical in lower half, with diverging free carpels above, 3-4 mm long, 4.5—5.5 mm wide, glabrous, dry, dehiscent; styles usually strongly re-curved at maturity, persistent; sepals persistent in fruit. Seeds num- erous, 0.5—1 mm long, yellowish brown, with surface verru- cose. Figure 3a, c; map 4b. Acknowledgments We thank Kristina Makkay for assistance with the Statistical analysis and figures; Laurie Consaul for sug- gestions in statistical analysis and assistance with map- ping; Michelle Leblanc for assistance with mapping: Susan Aiken for interesting discussions; Gisele Mitrow and William J. Cody for assistance at the DAO herb- arium. Also, a special thank-you to Marcel Blondeau from the Université Laval for sharing his knowledge and insights on the Saxifraga nivalis complex. We acknowledge and thank Polar Continental Shelf Project, Nunavut Research Institute and Aurora Research Institute for their logistic support in the field. This paper is based on a fourth year honours thesis project by C. H. submitted to the Biology Department, Uni- versity of Ottawa. Documents Cited (marked * in text). Aiken, S. G., M. J. Dallwitz, L. L. Consaul, C. L McJannet, L. J. Gillespie, R. L. Boles, G. W. Argus, J. M. Gillett, P. J. Scott, R. Elven, M. C. LeBlanc, and A. E. Zamluk. 2000. 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Axel Heiberg Island: 79°28'N 87°40'W, Beschel 10979 (CAN); Bukken River, 80°31.57'N 92°21.83'W, Gillespie et al. 6613 (CAN); Crusoe Glacier, 97°27'N 91°12'W, Kuc 476-3 (CAN); Crusoe River, 79°24'N 90°S0'W, Kuc s.n. (CAN 331187); Diana Lake, 79°30°N 88°30’ W, Porsild 16699-2 (CAN); Expedition Fjord, 79°20'N 92°W, Kuc s.n. (CAN 331189, CAN 331189); Mesa Brook, 79°37'N 95°W, Beschel 13129 (CAN); Mokka Fiord, 79°45'N 87°W, Parker 73061E (CAN). Baffin Island: 63°24'N 64°35'W, McLaren 91 (CAN); 66°40'N 70°W, Soper s.n. (CAN 65150); Admiralty Inlet, 73°13'N 84°W, Malte 118993 (CAN); Apex, 63°43'N 68°27'W, Gillespie & Soreng 6785 (CAN); Beekman Peninsula, 63°24'N 64°35'W, McLaren 9] (CAN); Burwash Bay, 65°59'N 71°18'W, Jacobs & Maus s.n. (CAN 517697); Frobisher Bay, 63°45'N 68°31'W, Gillett 19018 (CAN); Dorset Island, 64°12'N 76°32'W, Hainaud & Norman 5698 (CAN); Foxe Peninsula, 66°23'N 83°13'W, Manning 265 (CAN); Foxe Peninsula, 69°52'N 76°54'W, Manning 231 (CAN); Frobisher Bay, 62°57'N 66°03'W, Aiken et al. 86-260 (CAN); Frobisher Bay, 63°44'N 68°27'W, Aiken et al. 86-493 (CAN); Frobisher Bay, 62°57'N 66°03'W, Calder et al. s.n. (DAO 24148); Frobisher Bay, 62°57'N 66°03'W, Wynne-Edwards 7353 (CAN); Pond Inlet, 72°50'N 76°40'W, Dutilly 1224a (CAN); Stising Valley, Weber 94 (CAN); Taverner Bay, 67°12'N 72°25'W, Manning 44, 68, 77 (CAN). Bathurst Island: Polar Bear Pass, 75°43'N 98°23'W, Aiken & Maus 92-022 (CAN); Goodsir Inlet, 75°45'N 97°45'W, Lamothe 69-25 (CAN); Polar Bear Pass, 75°43.8'N 98°25.45'W, Gillespie & Consaul 6984 (CAN) [DNA voucher, individuals 1-5 examined]. Bylot Island: 72°53'N 76°W, Wilcox 125631 (CAN). Cameron Island: 76°19.5'N 104.5°W, Aiken 92-039 (CAN). Cornwallis Island: Resolute, 74°40'N 94°50'W, Aiken & MacCormac 98-055 (CAN 581909, plants 1-5; CAN 582554); Resolute, 74°40'N 95°04'W, Aiken et al. 93-072 (CAN). Devon Island: Dundas Harbour, 74°31.3'N 82°33.5'W, Gillespie et al. 6675-1 (CAN); Truelove Lowland, 75°40'N 84°40'W Kerik s.n. (CAN 409363). Digges Island: 62°32'N 77°45'W, Gaston 25, 29 (CAN); 62°35'N 77°50'W, Gaston 17 (CAN). Ellef Ringnes Island: Isachsen, 78°47'N 103°50'W, Macdonald 242 (CAN). Ellesmere Island: 79°34'N 84°44'W, Edlund & Roncato- Spencer 264 (CAN); Alexandra Fiord, 78°53'N 75°45'W, Gillespie & Vogel 6165 (CAN); Alexandra Fiord, 78°53'N 75°50'W, Gillett & Shchepanek 18108 (CAN); Craig Harbour, 76°20'N 81°30'W, Dutilly 1267 (CAN): Hazen, 68°30'N 72°45'W, Soper 8194 (CAN); Hazen, 81°49'N 71°20'W, Gillespie & Vogel 6238 (CAN); Sawtooth Range, 79°43.54'N 83°09.44'W, Gillespie et al. 6643 (CAN); Slidre Fjord, 80°N 86°15'W, Tener 95 (CAN): Tanquary, 78°53'N 75°45'W, Gillespie & Vogel 5982 (CAN). King William Island: Gjoa Haven, Cooper 28 (CAN), 73 (CAN), 226 (CAN); Gjoa Haven, 68°38'N 95°53'W, Larson 25 (CAN); Victory Point, Cooper 164 (CAN), 122 (CAN). Lougheed Island: 77°30'N 105°38'W, Edlund 2091 (CAN). Meighen Island: 79°55'N 99°30'W, Kuc (CAN 331171). Pelly Bay: 68°53'N 89°51'W, Campbell s.n. (CAN 282778). Prince Charles Island: 67°10'N 76°43'W, Baldwin 1919 (CAN). Victoria Island: 68°53'N 105°W, Edlund & Argus 12730-1 (CAN). NORTHWEST TERRITORIES. Banks Island: 73°N 117°W, Aiken 99-203 (CAN); 73°24'N 117°0'W, Porsild 17711 (CAN); 74°31'N 121°07'W, Manning & Macpherson 164 (CAN); Bernard River, 73°22'N 121°47'W, Maher & Maclean 75 (CAN); Egg River, 72°27'N 124°36'W, Lambert s.n. (CAN 535943); Lambton, 71°05'N 123°09'W, Porsild 17579 (CAN); Shoran Lake, 71°51'N 113°23'W, MaclInnes s.n. (CAN 535570, 535598, 535733). Fitzwilliam Owen Island: 77°07'N 113°47'W, Kuc s.n. (CAN 331114). Melville Island: 75°56'N 114°48'W, Edlund & Aiken 148 (CAN); Canrobert Hills 75°47.30'N 115°56'W, Edlund 179 (CAN); Ibbett Bay, 75°54'N 114°30'W, Edlund 43 (CAN); Kitson River, 76°02'N 113°05'W, Edlund 18 (CAN); Marie Bay 76°12'N 114°55'W, Edlund 305 (CAN); McCormick Inlet, 75°46.2'N 112°31.64'W, Gillespie & Consaul 6977 (CAN, plants 1, 2), 6964a (CAN); Shellabear Point, 71°51'N 113°23'W, Edlund 115 (CAN); Sherard Bay, 76°08'N 108°07'W, Dugal s.n. (CAN 535598); Winter Harbour, 74°17'N 110°42'W, Kuc s.n. (CAN 400104); Winter Harbour, 74°17'N 110°42'W, Tener & Harington 149 (CAN). Prince Patrick Island: Green Bay. 76°33.67'N 118°53.09'W, Gillespie & Consaul 6869 (CAN, plants 1-6); Intrepid Inlet, 76°22.01'N 118°35.45'W, Gillespie 340 THE CANADIAN FIELD-NATURALIST Vol. 118 & Consaul 6876 (CAN); Mould Bay 76°12N 119°25'W, MacDonald 146 (CAN); Mould Bay, 76°14.50'N 119°21'W, Gillespie & Consaul 6806 (CAN, plants 1-5); Mould Bay, 76°14'N 118°57'W, Gillespie & Consaul 6827 (CAN, plants 1, 2), 6826 (CAN, plants al-a5); Mould Bay, 76°21.50'N 119°30.50'W, Gillespie & Consaul 6884 (CAN, plants 1, 2), 6984 (CAN, plants 1-5), 6806 (CAN, plants 1-5). Victoria Island: Minto, 71°34'N 115°21'W, Edlund 594 (CAN). Saxifraga tenuis NUNAVUT. Axel Heiberg Island: Bastion Ridge, 79°21'N 90°48'W, Beschel 12920 (CAN); Mesa Brook, 79°35'N 95°30'W, Beschel 13129 (CAN 295548): 80°17'N 88°27'W, Scotter & Zoltai 45133 (DAO), 4518/1 (DAO). Baffin Island: Resolution Island, 61°20'N 64°55'W, Wynne-Edwards 7225 (CAN); Erik Harbour, 72°40'N 76°30'W, Calder s.n. (DAO 30869, 30871); Ogac Lake, 62°51.7'N 67°21'W, McLaren s.n. (CAN 274125); Frobisher Bay, 63°45'N, 68°32'W, Senn 3658 (DAO); Blackhead Island, 64°59'N 66°19'W, Soper s.n. (CAN 125598); Savage Islands, 61°49.15'N 65°42.62'W, Gillespie et al. 6728 (CAN), 6732 (CAN); Inugsuin Fjord, 69°50'N 69°08'W, Beschel 3649A (CAN). Bathurst Island: 76°36'N 100°04'W, Blake 24a (DAO). Bylot Island: 73.03'N 80.07'W, Scotter & Zoltai 6751A (DAO). Ellef Ringnes Island: Isachsen, 78°49'N 103°37'W, Savile 4275 (DAO), 4219 (DAO), 4224 (DAO), 4285 (DAO), 4355 (DAO), 4293 (DAO); Isachsen, 78°47'N 103°30'W, MacDonald 251 (CAN), 252 (CAN); Christopher Peninsula, 78°59'N 101°35'W, Savile 4194 (DAO); 78°45'N 102°50'W, Savile 4148 (DAO). Ellesmere Island: Eureka, 79°59'N 85°50'W, Scotter & Zoltai 45294-B (DAO); Eureka, 80°10'N, 85°39'W, Bruggemann S05 (DAO); Fosheim Peninsula, 79°34'N 84°44'W, Edlund & Roncato- Spencer 262 (CAN 535348, 533280); Fosheim Peninsula, 79°58'N 84°26'W, Edlund & Roncato-Spencer 275 (CAN); Hazen, 81°49'N 71°21'W, Savile 4485 (DAO), 4752 (DAO), 4655 (DAO); Hazen, 81°45'N 68°30'W, Powell 338 (CAN); Hazen, 82°N 70°W, Soper 8273 (CAN); Makinsen Inlet, 76°41'N 81°37'W, Blake 7 (DAO); Mount Pullen, 82°25'N 62°17'W, MacDonald 46 (CAN); 81°45'N 62°11'W, Powell 520 (CAN); Mt. Pullen, 82°26'N 62°11'W, Bruggemann 215 (DAO); Tanquary Fjord, 79°12'N 83°29'W, Brassard 3306 (CAN); Van Hauen Pass, 81°07'N 86°55'W, Brassard 3047 (CAN); Vesle Fjord, 79°12'N 83°29'W, Edlund & Roncato-Spencer 282 (CAN). Somerset Island: 72°56'N 94°57'W, Zoltai 741044 (DAO). NORTHWEST TERRITORIES. Melville Island: 74°58'N 115°02'W, Parker 44f (DAO); Bailey Point, 74°58'N,115°02'W, Parker 48b (DAO); McCormick Inlet, 75°46.2'N 112°31.64'W, Gillespie & Consaul 6978 (CAN, plants 1, 2), 6964b (CAN). Prince Patrick Island: Green Bay, 76°33.67'N 118°53.09'W, Gillespie & Consaul 6870 (CAN, plant 1); Intrepid Inlet, 75°46.9'N 112°22.09'W, Gillespie & Consaul 6877 (CAN, plant 1); Mould Bay, 76°21.5'N 119°30.5'W, Gillespie & Consaul 6881 (CAN, plants 1-6); Mould Bay, 76°14'N 118°57'W, Gillespie & Consaul 6825 (CAN, plants 1-3); Mould Bay, 76°14.5'N 119°21'W, Gillespie & Consaul 6807 (CAN, plants 1-5). Victoria Island: Ulusartok Peninsula, 70°44'N 117°44'W, Edlund 875 (CAN). ALASKA. Kenai Peninsula. 60°49'N 149°33'W, Calder 6511, 6249, 6107 (DAO). BRITISH COLUMBIA. Haines Junction, Calder & Kukkonen 28151 (DAO); Haines, Taylor et al. 1204 (DAO). Mixed collections - Saxifraga tenuis and Saxifraga nivalis on same sheet NUNAVUT. Baffin Island: Apex, 63°43'N 68°27'W, Calder et al. s.n. (DAO 24149); Home Bay, 68°45'N 67°10'W, Smith VP-53-61 (CAN). Ellesmere Island: Tanquary Fjord, 79°12'N 83°29'W, Brassard 3317 (CAN); Hazen, 81°49'N 71°21'W, Savile 4562 (DAO). NORTHWEST TERRITORIES. Victoria Island: near Holman, 70°43'N 117°43'W, Edlund 719 (CAN). Prince Patrick Island: Mould Bay, 76°12'N 119°25'W, MacDonald 150 (CAN, S. tenuis: 1, 3, 4, 7, 8, 9; S. nivalis: 2, 5, 6, 10). _ The Influence of Prey Availability and Vegetation Characteristics on | Scent Station Visitation Rates of Coyotes, Canis latrans, in a Heterogeneous Environment LynpDA A. RANDA! and JOHN A. YUNGER2 Department of Biological Sciences, Northern [Illinois University, DeKalb, Illinois 60115 USA 'Present address: Natural and Applied Sciences Division, College of DuPage, 425 Fawell Boulevard, Glen Ellyn, Illinois 60137 USA Present address: Environmental Biology Program, Governors State University, University Park, Illinois 60466 USA Randa, Lynda A., and John A. Yunger. 2004. The influence of prey availability and vegetation characteristics on scent station visitation rates of Coyotes, Canis latrans, in a heterogeneous environment. Canadian Field-Naturalist 118(3): 341-353. We investigated the effects of local prey fluctuations and habitat variables on the scent station visitation rates of the Coyote (Canis latrans) in northern Illinois within a heterogeneous environment. Availability of small mammalian prey was assessed by monthly mark-recapture sampling and visual counts conducted along three, 192-m transects in each of seven habitats that ranged from grassland to wooded sites. Habitat metrics, which included foliage density, ground cover, and canopy cover, were also collected for the same seven habitats. Visitation rates of Coyotes were determined from scent station lines parallel to the small mammal trapping transects. A multiple regression analysis indicated that Coyote visitation rates across the study site were influenced positively by vole (Microtus spp.) abundance and negatively by canopy cover. When Coyote visitation rates were regressed on vole abundance for only the habitats in which voles occurred, the relationship was not significant. This may be attributed to the general avoidance of wooded areas by Coyotes. Coyotes did, however, respond to experimentally- induced abundant patches of Peromyscus. These findings suggest Coyotes selectively use grassland habitats within a hetero- geneous environment and may modify their use according to prey availability. Key Words: Coyote, Canis latrans, habitat use, heterogeneous environment, prey availability, scent station, Illinois. The Coyote, Canis latrans, is among the most adapt- able of North American predators. Across an expan- sive range, the species occupies a variety of habitats and consumes a diversity of foods (Bekoff 1978). As such, the Coyote has been the focus of various local and regional studies of distribution, activity patterns, and diet selection. In particular, spatial and temporal activity patterns have been explicitly described (e.g., Andelt and Gipson 1979; Holzman et al. 1992), yet there has often been only circumstantial evidence as to causal factors (e.g., Person and Hirth 1991). Under- standing potential determinants of Coyote spatial dis- tribution and activity patterns requires an understanding of factors such as diet composition and specific use of available habitat types. Predators exhibiting flexible behavior patterns, such as Coyotes, provide an opportunity for studying possi- ble rapid responses to prey availability. Previous studies have shown that diets (Andelt et al. 1987: Brillhart and Kaufman 1995; Todd et al. 1981) and temporal activities of Coyotes (Shivik and Crabtree 1995) can vary seasonally based upon prey availability. Because space use and movements of vertebrate predators are closely coupled to foraging behavior, Coyotes could also alter their spatial use patterns in response to chang- es in prey availability. However, vertebrate predators frequently move over large spatial scales, whereas their smaller prey tend to be restricted to specific habitats or patches within a habitat. The study of prey in only one or a limited number of locations used by the pred- ators (Hamlin et al. 1984; Jaksic et al. 1993; Korsch- gen and Stuart 1972) may preclude accurate inferences on the foraging behavior of vertebrate predators. Thus, it is important to relate predator foraging behavior among habitats to prey abundances within habitats of heterogeneous landscapes (Dunk and Cooper 1994; Korpimaki 1994). In addition to prey, variations in vegetation structure, such as foliage density and height, among different habitats can also influence the movements and activities of vertebrate predators (Clark et al. 1993; Lamberson et al. 1994), especially in fragmented landscapes. Habitat differences may constrain predator foraging in certain areas through predators selectively using or avoiding particular habitat types (DeJong 1995; Small- wood 1995). As a result, predators may not respond to changes in prey availability within certain habitats. However, vegetation measures used to quantify pred- ator habitat use may also be related to or reflect prey usage (Anthony et al. 1981). Such inter-relatedness must be considered when making inferences of pred- ator behavior. We investigated the relationships of spatial and tem- poral distribution of prey and vegetation characteristics to the habitat visitation of Coyotes across a hetero- geneous environment. We incorporated experimental manipulations of habitat and prey, along with natural perturbations, to elicit predator behavioral responses 341 342 within a relatively short time scale. Data collected on prey abundance and habitat variables were used to ad- dress whether scent station visitation rates of Coyotes vary among habitats and within a particular habitat in response to changes in prey availability. Study Areas The study site was located in northern Illinois at Fermi National Accelerator Laboratory (Fermilab) in Batavia, Illinois (41°50'N, 88°15'W). Fermilab encom- passes approximately 3200 ha and is characterized as a heterogeneous area, marked by distinct habitat bound- aries. The areas surrounding Fermilab consisted of small open spaces of mainly agricultural land inter- spersed with light industry and residential neighbor- hoods, providing potential predator access into or out of the study site. The study was conducted at seven different locations within Fermilab, representing five habitat types: (1) oldfield, dominated by Queen Anne’s Lace (Daucus carota) and Tall Goldenrod, (2) tallgrass prairie, dominated by Big Bluestem (Andropogon ger- ardii) and Indian Grass (Sorgastrum nutans); (3) brome grass field, dominated by Smooth Brome (Bromus inermis); (4) shrubby oldfield, a heterogeneous mix of dense shrubs and shrub-grassland mixture dominated by Tall Goldenrod (Solidago altissima) and Gray Dog- wood (Cornus racemosa) with small stands of Trem- bling Aspen (Populus tremuloides); and (5) oak wood- land, dominated by mature oaks (Quercus spp.), one woodland (Oak Woods I) characterized by less dense canopy (primarily Q. macrocarpa and Q. rubra) than the other (Oak Woods II; predominantly Q. alba and Q. rubra; Figure 1). These different habitats were cho- sen because they represented the heterogeneity of the study site and included all prey species potentially found in Coyote diets at Fermilab. A concurrent inves- tigation indicated that three prey, voles (primarily Meadow Voles, Microtus pennsylvanicus, and rarely Prairie Voles, M. ochrogaster), Eastern Cottontails (Sylvilagus floridanus), and mice (Peromyscus spp.), constituted 39.4%, 31.3%, and 8.4% of the Coyote diets, respectively, which was at least two times great- er than the proportion of any other prey item (Randa 1996). Methods Availability of prey From March 1994 through May 1995, small mam- mal species compositions and abundances were esti- mated by mark-recapture live-trapping on three, 192 m transects located within each of the seven habitats (Figure 1). The transects were parallel and spaced from 70 to 85 m to help maximize their independence in relation to small mammal movements. At least 50 m was maintained from each transect to habitat bound- aries to avoid edge effects. Seventeen trap stations were spaced at 12-m intervals along each transect, totaling 51 trap stations per habitat. One 23 x 9 x 7.5 cm THE CANADIAN FIELD-NATURALIST Vol. 118 Sherman live-trap, baited with a peanut butter and oats mixture, was placed at each station. All habitats were trapped for two nights on a monthly basis. Traps were checked early morning and late afternoon (ca. every 12 hr). Captured individuals were marked with a uniquely numbered eartag (Monel Number 1, Na- tional Band and Tag Co., Newport, Kentucky). Nocturnal visual counts, using a high-powered spot- light (Woolf et al. 1993), were conducted for Eastern Cottontails in each of the seven habitats. The counts were conducted twice during each of the 15 small mammal trap sessions by sighting with binoculars, at one end and midway, along the small mammal trap- ping transects. Animals within ca. 25 m of the tran- sect were counted. For the visual counts, the trapping transects were lengthened to 250 m, delineated by flag- ging at opposing ends, to allow for the larger spatial movements of rabbits. Care was taken not to double count animals potentially moving between transects. For the experimental studies, fire was used as a means of manipulating prey. Tallgrass Prairie I was burned between the October and November 1994 small mammal surveys, a period of time when small mammal abundances peak in northern Illinois (Yunger 2002). This site was adjacent to Oak Woods I (Figure 1), which concurrently experienced a high acorn production year. Both habitats contained two closely related species of small mammals (Peromyscus mani- culatus, the Deer Mouse, in the prairie and P. leu- copus, the White-footed Mouse, in the woods), which were similar in body size and morphology. Numerical responses, caused by immigration of Deer Mice to prairie burns, have been documented previously (Cook 1959; Kaufman et al. 1983, 1988). Similarly, acorns have been shown to be an important local food source for mice (Peromyscus spp: Ostfeld et al. 1996; Wolff 1996). Consequently, we predicted a localized increase of P. maniculatus in Tallgrass Prairie I and P. leucopus in Oakwoods I. The proximity of these two habitats provided the opportunity to compare changes in Coy- ote habitat visitation following the manipulations of prey abundance. Prey also was manipulated in Tallgrass Prairie II through food supplementation. From October 1994 through March 1995, 10 kg of commercial rodent chow was hand broadcasted at weekly intervals on two of four (i.e., two treatment and two control), 0.60-ha square plots. As part of a separate study on the popu- lation dynamics of small mammals (Yunger 2002), prey numbers were determined by monthly trapping on a 6 X 6 grid with 12-m spacing in each plot. Predator spatial and temporal visitation responses to resulting changes in prey due to the food supplemen- tation experiment were also compared to the natural increase in acorns of Oak Woods I and to the prey response to experimental burning of Tallgrass Prairie I. Visitation rates of Coyotes 2004 RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES es me mee n e le hu | = A ( (|) Ibo! oO a senses “amas om sow 6 ame a : 1 | ear Bee oo Bea Pha 1 Oldfield Wea 2 Tallgrass Prairie I mane 3 Oak Woods I 4 Shrubby Oldfield 5 Oak Woods II 6 Brome Grass Field 10) 500 m Ace N ——— ee ransect 7 Tallgrass Prairie II FIGURE 1. Map of Fermilab showing the seven different study locations (habitats) and approximate locations of scent station transects in each habitat. Dashed line represents boundary of Fermilab. Roads, buildings, and open water (dark shaded areas) are included to further depict heterogeneity of the study site. 344 Coyote visitation rates were monitored through the use of scent stations. These were constructed by exca- vating a 50-cm diameter by 10-cm deep circular depres- sion, rimmed up to the ground surface with a 10-cm wide strip of aluminum flashing to inhibit invasive growth from surrounding vegetation, and filled with fine-grained sand. The resulting stations were level with the ground surface and fairly inconspicuous except for the lack of vegetation. Scent stations were located along 175-m transects, parallel to the small mammal trapping transects, in each of the seven loca- tions. Transects were spaced ca. 100 m apart while maintaining at least 50 m from a transect to the habi- tat edge. Eight scent stations were spaced every 25 m along each transect in the seven habitats, for a total of 24 scent stations per location. At the start of each monitoring period, all scent stations were baited by placing a cotton swab dipped in a liquid commercial predator lure (Cronk’s Predator 500, Wisscasset, Maine) upright in the middle of the scent station and the sand was smoothed. Stations were simultaneously monitored for a minimum of 2 nights (without precipitation), | to 3 times monthly (mean of 2.1 times per month); sand was not smoothed or otherwise disturbed by the authors during each monitoring period. Monthly scent station use was calculated for each habitat based upon visitation rates, or the proportion of scent stations exhibiting Coyote tracks in a given habitat, and divid- ed by the total number of operative nights to account for any variation in sampling duration. Within-habitat Coyote visitation rates were evaluated through the small mammal food supplementation experiment conducted in Tallgrass Prairie II. In Jan- uary 1995, one scent station was placed at each corner of the four plots previously described for this habitat. These 16 stations were monitored from February 1995 through May 1995 to compare scent station visitation rates of Coyotes to prey numbers in supplemented and non-supplemented plots. Vegetation analysis Vertical foliage density, percent ground cover, and canopy cover were used to describe habitat structure for each of the seven habitats. Vegetation measurements were collected at 24 stations per habitat, along alter- nating stations of the small mammal trapping transects. Foliage density was measured using a modified profile board technique (MacArthur and MacArthur 1961). Percent cover of three, 30 x 50-cm profile boards, divided into 15, 10 x 10-cm cells, was measured at 0.5 m, 1.0 m and 1.5 m from ground to mid-level of the board. A cell was considered covered if 50% or more of the cell, viewed from a 5-m distance, was ob- structed by vegetation. The proportion of cells covered determined percent cover of an individual board. All readings were taken facing north and there was no apparent patterning or trending in vegetation struc- ture and diversity among each of the four cardinal directions. To arrive at a single foliage density value THE CANADIAN FIELD-NATURALIST Vol. 118 for each habitat, the proportion of vegetation cover was averaged across the three vertical profile boards for each sampling station, then the mean of the result- ing 24 values was calculated. Percent ground cover was assessed at each sam- pling station by the amount of vegetation (herbaceous, woody, and leaf litter) occupying a 1-m? quadrat against visible patches of ground (Brower et al. 1990). Canopy cover was estimated by the presence or absence (scored as | or 0, respectively) of vegetation viewed through a vertical ocular tube (James and Shugart 1970) at each sampling station. Average ground cover and canopy cover for each habitat was obtained by taking the mean of the 24 values of each metric. The vegetation analysis was conducted once in March 1994 and once in August 1994, representing vegetation structure characteristic of the non-growing and growing seasons, respectively. Two months of the year, May and November, represent transitions between the growing and non-growing seasons and hence, char- acteristic vegetation density. For this study, vegetation variables for May were described by the August sur- vey (i.e., high foliage density), and vegetation variables for November were described by the March survey (i.e., low foliage density). Data analysis Analyses of small mammal prey were based upon minimum number known alive (MNKA). This esti- mate was generated for each individual transect per month using Package C. M. R. (Le Boulengé 1987). The estimates then were averaged for the three tran- sects to yield monthly small mammal species compo- sition and abundance for each habitat. To examine whether Coyotes shifted visitation rates in response to prey, it was necessary to determine whether spatial and temporal differences in prey abun- dances existed. Analysis of covariance (ANCOVA), conducted with SAS PROC GLM (SAS Institute Inc. 1990), was used to compare abundances of the most important prey species identified in Fermilab Coyote diets (Randa 1996). Specifically, analyses were con- ducted to test for significant fluctuations in prey abun- dances over time among each of the seven habitats. ANCOVA can be used to test for an interaction or heterogeneity of slopes (Littell et al. 1991) and was chosen because sampling units (i.e., habitats) were not replicated in space. Temporal changes of prey popu- lations were indicated by crossing the categorical vari- able (1.e., prey abundances across habitats or prey abun- dances per habitat) with the continuous covariate (time). Traditional model building techniques were used (Box et al. 1978) in which non-significant, higher- order interactions were removed from the model. If no significant sources of variation were detected, the full model was reported. If significant interaction of variables occurred (i.e., heterogeneity of slopes), sig- nificance of main effects were inferred from the plot- ted data. Statistical inferences were based on type III 2004 sum-of-squares and significance accepted at a = 0.05. Multiple regression (SAS PROC REG; SAS Insti- tute Inc. 1990) was used to evaluate the effect prey availability and vegetation structure had on Coyote visitation rates. Six independent variables were used in the model: the abundances of the three main prey genera, (1) Microtus, (2) Peromyscus, and (3) Sylvi- lagus, and the three vegetation variables, (4) vertical foliage density, (5) percent ground cover, and (6) can- opy cover. Each observation in the multiple regres- sion represented a single measure of the independent variable in one of the seven habitats during one of 15 months. Thus, there were 105 observations for each independent variable, or a total of 630 data points for the regression analysis. Data were logarithmic-trans- formed prior to analysis to help satisfy assumption of normality of the residuals. Collinearity diagnostics, such as tolerance values and variance inflation factors, indicated a lack of colinearity between the regres- sors, hence all independent variables were retained in the model. Simple linear regression, performed with SAS PROC REG (SAS Institute Inc. 1990), was used to compare Coyote visitation rates against mice abundances in each of the two adjacent habitats, Tallgrass Prairie I and Oak Woods I. We performed a separate analysis for the response of Coyotes to prey increase following manipulation of the prey’s food supply. Visitation rates of Coyotes around the experimental plots in Tallgrass Prairie II were analyzed using a one-way repeated measures analysis of variance ((mANOVA), with SAS PROC GLM (SAS Institute Inc. 1990). Inferences were based on Huynh-Feldt adjusted P-values for the rmANOVA. Results Among-habitat prey fluctuations There was variation in spatial-temporal abundances of prey species (Figure 2) with significant species (R? = 0.658, Fro. 264) = 6.21, P = 0.002), time (Fry 264) = 21.29, P < 0.001), and time? (Fy, 264) = 14.17, P < 0.001) effects, indicating that overall numbers of the three prey differed significantly. The significant time X species (Fy 264) = 18.29, P < 0.001) and time* x spe- cies (Fi, 964; = 11.74, P < 0.001) interactions indi- cated that the rate at which prey species numbers changed also differed significantly. This was primar- ily due to the substantial increase in mice numbers (Figure 2). Furthermore, the significant time x habi- tat x species (Fry. 264) = 4.51, P < 0.001) and time? x habitat x species (Fi, 364; = 4-84, P < 0.001) inter- actions indicated that the differential rates in changes of numbers of species also changed among the dif- ferent habitats. For example, the highest abundances of mice and voles occurred in November 1994 in the habitats Oak Woods I and Tallgrass Prairie I, res- pectively. The highest Sylvilagus abundance was docu- mented in October 1994 in the Oldfield, with nearly equally high numbers recorded in July 1994 in the RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 345 two prairie sites, probably following the weaning period of young rabbits in northern Illinois (Hoffmeister 1989). These abundance peaks are reflected in the quadratic function of the change in prey numbers over time. Among-habitat Coyote visitation rates Scent station visitation rates fluctuated widely among habitats and over time; no signs of Coyotes were found in Oak Woods II and the majority of tracks were re- corded in the grassland habitats (Figure 3). Overall, the multiple regression of Coyote scent station visita- tion rates across Fermilab with the three main prey (Figure 4) and the three habitat structure variables (Figure 5) was significant (R* = 0.241, Fy, og) = 4.40, P = (0.003; Table 1). The unexplained variance in the model may be attributed to the observed variation in Coyote visitation (Figure 3) and 0 values recorded for prey abundances. Based upon results of the multiple regression, we conducted a separate analysis to discern whether the visitation rates in grasslands by Coyotes was due to prey (Microtus) or avoidance of wooded areas, from which voles were absent. Regression of scent station visitation rates with vole abundance, ex- cluding data from the two oak woods, was not signi- ficant (7? = 0.050, Fy, 73; = 3.75, P = 0.057). In comparing Coyote visitation in the two adjacent habitats, Tallgrass Prairie I and Oak Woods I, scent station visitation rates, when averaged over the study period, were over three times greater in the prairie (0.035 + 0.011 (mean + SE ); range 0.000—0.0645), than the woods (0.008, + 0.003, range 0.000—0.0179; tog) = 2-486, P = 0.026). However, mice abundance was two times greater in the oak woods (4.7 individ- uals per transect + 1.4, range 0.0—19.3) than the prairie (2.3 individuals per transect + 0.6, range 0.3—8.0; fig) = 1.637, P = 0.113). Within-habitat Coyote visitation rates The fire in Tallgrass Prairie I did not result in a substantial increase in prey (Figure 2). However, the perturbation in Oak Woods I was followed by a con- siderable increase in mice abundance, enabling a within-habitat comparison to Coyote activity. Despite the high prey abundance, Coyote visitation rates were not significantly related to mice abundance (7° = 0.158, Fry, 13) = 1.03, P = 0.389) in Oak Woods I. Food supplementation in Tallgrass Prairie II resulted in an approximately three-fold increase in numbers of mice compared to the non-supplemented plots by February 1995 (Figure 6). Numbers subsequently dec- lined until densities converged on the supplemented and non-supplemented plots in May 1995. Coyotes responded to the peak in mice densities with mean visitation around the food supplemented plots over five times greater than on non-supplemented plots in Feb- ruary 1995 (Figure 6). Coyote visitation around these plots in the ensuing three months reflected the trend in prey availability. The result was a significant food effect (Fi, >, = 25.60, P = 0.037), a significant time effect (Fi; 6 = 9-79, P = 0.013), and a significant 346 THE CANADIAN FIELD-NATURALIST Vol. 118 @—® Tallgrass Prairie I &—L Oak Woods II 4— A Oldfield O--O Shrubby Oldfield v--v Brome Grass Field C—O Tallgrass Prairie II @— 8 Oak Woods I Peromyscus Microtus Mean Number of Individuals per 200-m Transect Months FiGuRE 2. Abundances of the three main prey, Peromyscus, Microtus, and Sylvilagus, in each of the seven habitats over time. 2004 0.35 0.30 O25 0.20 0.15 0.10 0.05 Mean Scent Station Visitation Rate (proportion stations visited / operative days) 0.00 RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 347 Tallgrass Prairie I Tallgrass Prairie II Oakwoods I Oakwoods II Oldfield Shrubby Oldfield Brome Grass Field Months FicureE 3. Habitat visitation of Coyotes, based upon monthly average scent station visitation rates, over time among the seven habitats. visitation X time interaction (F/; ) = 10.32, P = 0.012). Vegetation characteristics of habitats Of the three vegetation metrics collected, percent ground cover and canopy cover estimates were fairly consistent between the non-growing and growing seasons for each of the seven habitats sampled (Table 2). Vertical foliage density was greater in the growing season than the non-growing season for all habitats, reflecting the increase in above-ground plant biomass. Tallgrass Prairie I had the greatest difference in ground cover and foliage density between these seasons. This was due to the almost complete absence of litter and standing vegetation during the March survey, except for small clumps of tall grasses, a result of the previous fall’s fire. There were similar trends of vegetation met- rics among similar habitats. Vertical foliage density was greatest in the prairies compared to other habitats during the growing season, but was relatively low for the woods. The habitats dominated by grasses, the prairies and brome grass field, exhibited > 97% ground cover, exceeding estimates of other habitats. Canopy cover, as expected, was highest in the woods, with Oakwoods II having a denser overstory than Oak- woods I. The presence of canopy in the brome grass field was attributed to a few isolated trees located near the middle of the field, which was not expansive enough to markedly influence Coyote visitation. Discussion Coyote habitat visitation, measured through scent station visitation rates, fluctuated greatly over time, showing no clear temporal pattern among habitats. However, overall visitation indicated an extensive use of grasslands and avoidance of woodlands. Through- out the duration of the study, there was no evidence of Coyote tracks or scat in Oak Woods II, a mature woodlot with few gaps in its overstory, even though its entire western side bordered a tallgrass prairie, a habitat where Coyote scat and tracks in snow were observed. Oak Woods I also experienced relatively little Coyote visitation during most of the study. Although scent station transects were 50 m from habitat edges, all Coyote tracks were observed at the western end of the transects, which were closest to a grassy area near the woods. The shrubby oldfield experienced slightly more visitation, overall, than the woods, but tracks were found primarily at scent sta- tions within open grassy areas and in the western por- 348 THE CANADIAN FIELD-NATURALIST Vol. 118 Peromyscus Sylvilagus Mean Coyote Visitation Rate (proportion stations visited / operative days) 0.0 0.5 1.0 IS 2.0 De) 3.0 Mean Number of Prey per 200-m Transect Ficure 4. Partial regression plots for activity of Coyotes vs. abundances of each of the three main prey species. Each point represents a single observation recorded during | of 15 months in one of the seven habitats. ) 2004 RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 349 vertical foliage density canopy cover ground cover Mean Coyote Vistation Rate (proportion stations visited / operative nights) Means of Vegetation Variables | Ficure 5. Partial regression plots for activity of Coyotes vs. each of the three vegetation variables. Each point represents a single observation recorded during | of 15 months in one of the seven habitats. 350 GY WSS WN 4 ‘ Lf Ny NS SS fee KK Mean Number of Peromyscus per Plot Yj Y j L Y Months THE CANADIAN FIELD-NATURALIST —@— Peromyscus density, +F —O©Q-—. Peromyscus density, -F WZ, Visitation rate, +F [___] Visitation rate, -F Vol. 118 Mean Coyote Visitation Rate (proportion stations visited / operative nights) FIGURE 6. Comparison of activity of Coyotes and abundance of mice on food-supplemented (+F) and non-supplemented (-F) plots in Tallgrass Prairie II. tion of this habitat which bordered a tallgrass prairie. These observations were supported by the significant negative relationship of canopy cover to Coyote visita- tion rates in the multiple regression analysis. Hence, Coyotes may move a limited distance into woods but tend to avoid core areas of heavily wooded habitats. This behavior was corroborated by a two-year radio telemetry study of Coyotes in northeastern Illinois (Roth et al. 1999°). The multiple regression analysis also indicated scent station visitation rates by Coyotes were affected by abundance of voles, which was not found in the oak woods studied. However, a simple regression of Coy- TABLE |. Parameter estimates for independent variables used in the multiple regression analysis of Fermilab Coyote activities. Variable df Regression coefficient Jz Intercept 0.0261 0.040 Peromyscus 1 -0.0082 0.192 Microtus 1 0.1192 0.006 Sylvilagus -0.0169 0.359 Foliage Density 1 -0.0441 0.295 Canopy Cover 1 -0.0632 0.004 Ground Cover 1 0.0082 0.871 ote visitation rates on vole abundance, excluding data from the two woods, did not reveal a significant rela- tionship. Thus, it appeared that factors other than prey, such as vegetation structure, strongly influenced Coy- ote habitat visitation at Fermilab. In addition, overall visitation rates were not related to the abundance of Eastern Cottontails, which comprised the greatest biomass of prey in the Coyote diets (Randa 1996). This could possibly be attributed to the association of cottontail rabbits along habitat edges (Althoff et al. 1997; Mankin and Warner 1999; Smith and Livaitis 2000). Although some of the scent station transects were located within habitat areas where rabbits were found, Coyotes could have spent more time foraging for rabbits along these edges where transects were not located. The selective use or visitation of grasslands by the Coyotes in this investigation may be associated with their historical range. Prior to European settlement, Coyotes occupied the southwest and central plains of the U. S. including grassland of southern Canada. Clearing of forests and extermination of the Grey Wolf (Canis lupus) facilitated the north- and eastward expan- sion of Coyotes. Coyotes moved northward along the Great Lakes, and colonized the northeastern U.S. ) 2004. RANDA and YUNGER: SCENT STATION VISITATION RATES OF COYOTES 35] TABLE 2. Summary of vegetation characteristics for the seven different habitats sampled twice during 1995 at Fermilab; NG = non-growing season sampling, G = growing season sampling. Mean Vertical Mean Percent Mean Canopy Foliage Density* Ground Cover Cover? Habitat NG G NG G NG G Tallgrass Prairie I 0.031 0.703 352, 98.0 0.000 0.000 Tallgrass Prairie II 0.320 0.761 OED 97.5 0.000 0.000 Oakwoods I 0.196 0.377 94.7 89.7 0.750 0.792 Oakwoods II 0.132 0.172 66.4 52.4 1.000 1.000 Oldfield 0.328 0.459 89.4 89.2 0.000 0.000 Shrubby Oldfield 0.248 0.587 96.7 89.5 0.389 0.340 Brome Grass Field 0.180 0.346 100.0 99.7 0.042 0.021 “Based upon the average proportion of three profile boards obscured by vegetation for 24 sampling stations per habitat. >Based upon presence/absence scores (1/0 respectively) for 24 sampling stations per habitat. (Richens and Hugie 1974) and the Midwest (Moore and Parker 1992) by the 1940s. Most Coyotes still in- habit deserts and grasslands, and those in regions with forests tend to use open, non-forested areas in a great- er proportion than available (Todd et al. 1981; Toweill and Anthony 1988; Cypher 1991; Holzman et al. 1992; Murray et al. 1994; Kamler and Gipson 2000). Tracks of Coyotes observed in the snow at our study site indicated that grasslands, such as the brome grass field and tallgrass prairie, were occasionally traversed by Coyote tracks following a relatively straight path, reflecting non-foraging behavior. Hence, these open areas may not only serve as preferential foraging sites but as areas facilitating movement between habitats. A preferential use of grasslands may explain why chipmunks and tree squirrels, which were moderately common in the woodlands, were rarely detected in Coyote diets (Randa 1996). Selective use of grasslands may have restricted the ability of Coyotes to respond spatially and temporally to localized patches of abundant prey within a habi- tat. In Fall 1994, when mice densities in Oak Woods I were more than twice the prey abundances at any other location, Coyote visitation did not increase sig- nificantly in that habitat. This discrepancy of prey abun- dance and coyote response in woodland corresponds with findings in southeastern Quebec (Richer et al. 2002). However, Coyotes demonstrated an ability to closely track prey within a grassland (Tallgrass Prairie II) as their visitation rates were correlated with ex- perimentally manipulated abundant patches of mice. The high level and subsequent decline in Coyote visi- tation rates detected on scent stations around food- manipulated plots paralleled the change in mice densities. Hence, once mice decreased following a peak in densities on these plots, Coyote visitation rates likewise diminished. It was unlikely that Coyotes were responding to a novel item in the environment (such as the odor of the food) because supplementation had been initiated four months prior to the scent station monitoring. Because voles were not detected in this habitat during the experiment, a similar test of within- habitat Coyote response to abundant patches of this commonly consumed prey was not feasible. Yet, as with mice, Coyotes could have responded to isolated patches of voles within habitats throughout Fermilab, given that voles were still a major component of Coy- ote diets despite their low availability. One limitation of the scent station transects was the difficulty in identifying Coyote responses to possible small, isolated patches of prey. Also, due to the large areas over which Coyotes may move, there was not sufficient separation to avoid visitation by the same individual between scent station transects (Diefenbach et al. 1994; Roughton and Sweeny 1982). However, by intensively sampling each habitat, these transects were effective in discerning overall trends (i.e., among- habitat comparisons) in Coyote visitation rates and rel- ative habitat use; they were not intended to indicate finer-scale or localized responses (e.g., patch-level foraging) within the study site. The results we obtained from both the experimen- tal plots and scent station transects demonstrated that Coyotes exhibit variable responses to prey availability, depending upon habitat type. Reacting relatively quick- ly to a local increase in prey, as seen around the ex- perimental plots in prairie, would enhance Coyote foraging efficiency. However, preferential use of open areas, as suggested by the transect data, may con- strain foraging efficiency during periods of very low prey densities when alternative prey in woodlands is under-utilized. Hence, supporting Coyote populations near urbanized areas, as in northern Illinois, requires open land expanses such as grasslands to facilitate movements across large spatial extents within the land- scape and to provide an adequate prey base. Acknowledgments P. L. Meserve lent support and guidance throughout the course of this investigation. S. Scheiner and C. von Ende provided statistical advice. D. Cooper shared in much of the field activity. J. Mendelson, E. Miller, 392 C. Petersen, B. Satterfield, and several anonymous reviewers provided helpful comments on earlier drafts of this manuscript. R. Walton oversaw Fermilab activ- ities associated with this project and helped obtain funding. The National Environmental Research Park of the U.S. Department of Energy (MO-RS023), and the Department of Biological Sciences, Northern Illinois University provided funding for this study. Documents Cited (marked * in text) Roth, C., E. Miller, and J. Yunger. 1999. 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Journal of Mam- malogy 83: 267-279. Received 27 February 2003 Accepted 14 October 2004 Presence of Cavities in Snags Retained in Forest Cutblocks: Do Management Policies Promote Species Retention? Ki T. Everett!:? AND KEN A. OTTER? ‘Forestry Program, University of Northern British Columbia, Prince George, British Columbia V2N 4Z9 Canada "Ecosystem Science & Management Program, University of Northern British Columbia, Prince George, British Columbia V2N 4Z9 Canada 3Current Address: Department of Biology, University of Victoria, Victoria, British Columbia V8W 2Y2 Canada Everett, Kim T., and Ken A. Otter. 2004. Presence of cavities in snags retained in forest cutblocks: do management policies promote species retention? Canadian Field-Naturalist 118(3): 354-359. Tree cavities, which are frequently excavated by primary cavity nesters, are typically used by a number of avian and non-avian species and are thus important components in maintaining biodiversity in forest ecosystems. One way to provide these habitat opportunities in harvested areas is through the retention of snags. In this study, we assessed the habitat and snag characteristics that promote cavity excavation, using the presence of cavities to infer activity of primary cavity excavators. Snags retained closer to the forest/cutblock edge contained a greater density of cavities than trees further from edge. However, the proportion of cavities found within cutblocks declined at a more rapid rate with distance from edge than did those in adja- cent forested stands. There was also a tendency for cavities to occur more frequently in trees that were at the advanced stages of decay. The results of our study suggest management for snags in harvest areas should include the retention of snags closer to the forest edge combined with incorporating trees showing signs of advanced decay. Key Words: tree cavities, snags, forest edge, birds, mammals, reptiles, British Columbia. Snags (standing dead trees) provide specialized hab- itat for many species, which in turn play an integral role in maintaining ecosystem functions (Machmer and Steeger 1995*). Approximately 16% of the indigenous birds, mammals, and reptiles in British Columbia de- pend to some extent on snags or cavities for repro- duction, feeding, and shelter (Backhouse and Lousier 1991*). These dependent species then create habitat opportunities for other wildlife species, facilitate seed dispersal, increase soil aeration, accelerate organic decomposition, and can even aid in controlling forest pests (Machmer and Steeger 1995*). Although snags play an important role in the ecosystem, their removal has traditionally occurred in past forest management (ostensibly to reduce hazards — lightning attraction, falling snags, etc. — and foci of infection for healthy trees). By retaining trees of varying ages, diameters, and decay classes in cutblocks, the vertical structure of vegetation, total percent vegetation, and percent decid- uous cover increase. These changes can lead to greater species richness and diversity in both plant and mam- mal communities (Davis et al. 1999*). Population den- sities of cavity-nesting species are often limited by potential nest sites (Runde and Capen 1987), and inten- sive forest management that removes snags reduces their densities in managed stands compared to natural stands (Haapanen 1965; Pojar 1995). Primary cavity nesters (those which excavate their own cavities, such as woodpeckers and some chicka- dees: Thomas et al. 1979) appear to prefer stands with a high number of snags (Zarnowitz and Manuwal 1985; Raphael and White 1984). As most primary cavity- nesting birds rarely occupy nests that they themselves have not excavated (Short 1979), suitable trees, in- cluding snags, must be available for excavation for nests or roosts. Snags offer softer excavating substrates as well as important foraging sites for these species. Conversion of natural forests into intensively managed stands which lack snags may, therefore, have long-term negative effects on the entire cavity-nesting populations, as secondary cavity-nesting species rely on primary excavators for cavity creation (Peterson and Grubb 1983). Bodies that govern forestry practices recognize the importance of snags, and advocate the retention of a variety of tree species at various stages of decay in managed forests (e.g., Machmer and Steeger 1995*). In an effort to reduce the impact of large openings created by cutblocks, some forest companies retain windfirm deciduous trees and advanced regeneration that fall below utilization standards to aid in main- taining structural diversity in cutblocks. This limits the distance wildlife must travel through the cutblock without vegetative cover. The traditional paradigm has been that efforts such as these will be sufficient to maintain wildlife and biodiversity throughout managed forests. One aspect of snag retention policy that has been overlooked, however, is the behavioral characteristics of primary cavity-nesting species that may deter mem- bers of this guild from occupying trees left specifically for their use. Many forest birds avoid areas outside forested cover (Hegner 1985; Desrochers and Hannon 354 2004 1997), as open spaces leave these species susceptible to hawk and falcon predation (Corral 1989). Several studies have shown that many forest dwelling species are reluctant to cross gaps between fragmented forest- ed areas (Lens and Dhondt 1994; Desrochers and Hannon 1997). Desrochers and Hannon (1997) found that forest birds were three times less likely to cross gaps 70 metres wide, and eight times less likely to cross gaps 100 metres wide compared to similar dis- tances within the forest. Machtans et al. (1996) also found that movement rates of forest birds across cut- blocks were significantly lower than movement rates within the forest. Instead, forest birds preferred to travel much greater distances through corridors, rather than crossing narrow gaps (Desrochers and Hannon 1997). The avoidance of open spaces indicated that forest birds — with the majority of those studied belonging to the cavity-nesting guild — have fewer habitat oppor- tunities as the landscape becomes increasingly frag- mented. Although retained snags may be scattered throughout cutblocks to provide nest sites, the gap between the forest edge and the snag may be too sub- stantial for birds to cross. In turn, the number of cavity- nesting birds may decline with increased distance into cutblocks as compared to the surrounding forest. In this study, we documented the presence of cavi- ties in retained snags within forest cutblocks as a mea- sure of use by cavity-excavating species, and compared this to similar trees found within the surrounding for- est. If primary cavity nesting species avoid openings in forest cover, we predict fewer cavities as one travels further from the forest edge into cutblocks. By assess- ing attributes of cavity-possessing trees, we attempt to identify which tree characteristics and tree species cavity nesting birds tend to utilize. Our aim is to provide well-substantiated recommendations towards the placement and creation of retained snags within cutblocks that maximize potential usage by cavity nesting species. Methods Study Area This study was carried out from 30 September to 5 November 2000, approximately 30 kilometers west of Prince George, British Columbia (53°36'35"N, 122°57'29"W). All samples were located within the Sub-Boreal Spruce biogeoclimatic zone, which covers much of the central interior of British Columbia (MacKinnon et al. 1992). The study area was primarily composed of coniferous and mixedwood forests, with species including Lodgepole Pine (Pinus contorta var. latifolia), hybrid White Spruce (Picea glauca x engelmannii), Sub-alpine Fir (Abies lasiocarpa), Doug- las-fir (Pseudotsuga menziesii var. glauca), Trembling Aspen (Populus tremuloides), Black Cottonwood (Pop- ulus balsamifera ssp. trichocarpa) and Paper Birch (Betula papyrifera). Harvested areas are typically re- planted with Lodgepole Pine, with a small component of hybrid White Spruce on moister sites. EVERETT AND OTTER: PRESENCE OF CAVITIES IN SNAGS 355 Sampling Procedure We randomly selected eight cutblocks for the survey from a Forest Development Map. The cutblocks ranged in age from 3 to 10 years post-harvest, and from 30 to 60 hectares in size. A line-transect method of sampling was used. In this method, all retained snags falling with- in 5 metres of each side of the transect were measured. We ran transects on a bearing perpendicular to the for- est edge, with a random point of commencement along the edge being selected from the Forest Development map. Measurements were carried out until the centre- line of the cutblock was reached and at least 20 snags were encountered. In one case, the cutblock contained only 17 retained snags. Due to the minimal number of retention trees present, the line-transect method would have been relatively ineffective; therefore; all retention trees in the cutblock were measured. Snags were defined as any dead, standing tree (Smith et al. 1997: 489). We recorded all cavities that were either actively excavated or were natural cavities of similar size, which displayed signs of active use (e.g., modification to the cavity, presence of feathers in entrance, etc.). Cavities were only recorded if they had a minimum diameter of 3.5 cm, as the cavity had to be large enough for use by a small bird or mammal. However, as our study was looking for evidence of use of the trees rather than specifically focusing on nesting, we did not distinguish between cavities used for roost- ing versus those where nesting had occurred. For each snag encountered along the transect, we recorded sev- eral attributes: tree species, height, diameter at breast height (DBH, measured at 1.3 m from the base of the tree), decay class, number of cavities, distance from the nearest timber edge, percent overhead cover, and the number of snags and live trees within an 11.28 m radius around the trees. Only trees greater than 10 cen- timeters DBH were recorded, as this reduced the chance of incorporating regeneration into the study. Decay class was determined using a six-step hardwood classi- fication scheme combined with a corresponding nine- step coniferous classification scheme (Ministry of Forests 2000*). Similar transects were placed in the adjacent forest- ed stands running perpendicular to the edge. In order to measure a similar number of trees over the same dis- tance in both habitat types, every third tree classified as decay class 2 or greater was measured. In order to determine the initial tree for measurement along each transect, a coin toss was completed to decide if the first or second tree should be measured. For each tree encountered along transects in forested stands, the same attributes were measured as those collected for cutblock sampling. Statistical Analysis A multiple regression was used to determine if the habitat (forest or cutblock) and distance from the for- est edge influenced the proportion of cavities found (SYSTAT 9.01). To compare the distribution of tree 356 types across the two habitats, we employed a Chi- square and used a Mann-Whitney U test to compare the frequency of decay classes of trees between cut- blocks and forested stands. To determine which characteristics tended to predict the presence of cavities in wildlife trees, we conduct- ed a Principal Component Analysis (PCA) on trees in the forested stands. As the forest stands constitute the natural habitat of many of the primary cavity-nesting species, determining the characteristics of cavity-bear- ing trees in these areas may give insight into which types of wildlife trees are important to retain in cutblocks. The following characteristics of the focal tree and the immediate area were subjected to PCA: height, diameter, and decay class of the focal tree, along with the total number of trees, number of live trees, number of snags, and percent overhead cover within the 11.28 metre radius plot. A linear regression was used to compare the number of tree cavities against the PCA factor scores. The PCA was carried out in SYSTAT 9.01 using Varimax rotation. Two retention trees measured had extremely high numbers of cavities and were identified during initial analysis as significant outliers in the data set (Durbin- Watson D Statistic = 1.984, P < 0.01). These were excluded from subsequent analysis. Results In total, 185 trees were measured at our eight sam- pling sites; 94 trees within cutblocks and 91 trees within forested stands. We found 46 cavity-bearing trees, con- taining a total of 67 cavities. Of the 46 trees containing cavities, 29 were deciduous while the remaining 17 were conifers. A greater proportion of deciduous trees contained cavities compared to coniferous species. Species of willow had the highest proportion of cavities, with two of the three trees observed containing cavities. THE CANADIAN FIELD-NATURALIST Vol. 118 Trembling aspen was found to have the second highest frequency of cavities with 17 of the 62 trees observed containing cavities. A breakdown of the number of cavities recorded by species is given in Table 1. The species composition of trees differed between the two habitats (x? = 76.6, df = 6, p< 0.01 — Figure 1). Snags in cutblocks were primarily deciduous species; although these trees were also present in forested stands, the latter stands had a proportionately higher coniferous component. Distance From Edge and Cavity Abundance Distance from forest edge and stand type (cutblock or forest) both had a significant effect on the number of cavities found (multiple regression: 17=0.051, Fo 19) = 4-857, p = 0.009). The further a tree was located from the forest-cutblock edge, the lower the frequency of cavities (p = 0.027). There was also a sig- nificant effect between the stand types, as more cavities were found in the forest than were found in the cut- block (p = 0.020). In the forest stands, the decline in number of cavities found with distance from the edge was not as dramatic as that seen in cutblocks (Figure 2a and b). Stand Characteristics Associated with Snags The principal component analysis identified two factors that accounted for 56.2% of the total variance among retained snags (Table 2). Principal component one (PC 1) accounted for 31.2% of the variance, with strong positive weightings for a high number of total trees and a high number of live trees within an 11.3 m radius plot. As a result, PC 1 can be considered a meas- ure of the density of trees surrounding the snag. Principal component two (PC 2) accounted for 25.0% of the total variance, with a strong positive weight- ing for decay class and a strong negative weighting for tree height. PC 2 indicates trees at the latter stages of the decay process, as tree height frequently decreases TABLE |. A breakdown of the number of cavities by tree species. Tree Species Number of trees observed Trembling Aspen Populus tremuloides 62 Lodgepole Pine Pinus contorta var. latifolia 52 Paper Birch Betula papyrifera Sy) Hybrid Spruce Picea glauca X engelmannii 24 Willow Salix sp. 3 Black Cottonwood Populus balsamifera ssp. trichocarpa 8 Douglas Fir Pseudotsuga menziesii glauca 4 Total (Average) 185 Total number of trees Percentage of trees Total number with cavities with cavities of cavities 17 27.4 32 13 25.0 14 8 25.0 10 4 16.7 5 2 66.7 3 2 25.0 3) 0 0 0 46 (24.9) 67 2004 70 | Cutblock ” Forest a) o Qa 2) S38 ae ES 25 CJ Q n x o ” Populus balsamifera Populus tremuloides Betula papyrifera Pseudotsuga menziesii Pinus contorta Picea glauca Tree Species FiGure 1. Distribution of sampled snags within cutblocks versus surrounding forest stands. as the tree reaches the higher decay classes (Towers et al. 1992). Response of Cavity Nesters to Stand Characteristics There was no significant relationship between the number of cavities found and PC | (regression: r* = 0.003, Fo 99) = 0.281, p = 0.60). There was, however, a significant effect between the number of cavities being found and PC 2 (regression: 1? = 0.092, F,, 99) = 9.13, p = 0.003) indicating that more cavities occurred in trees with a higher amount of decay. On average, the snags retained in clearcuts had less decay than the comparison cohort of trees in the forested areas (MWU: U = 2918.5, n= 93, 91, p<0.01). Table 2. Principal component analysis of stand characteris- tics in forest stands. The first two Principal Components ac- count for greater than 50% of the total variance among stand attributes; therefore only these two factors were considered in subsequent analysis. Variables that were within 10% of the highest loading were considered to be strong weightings, and are shown in bold. Variable Real PE2 (clustering of (decay retention trees) stage) Tree Diameter -0.407 -0.395 Tree Height 0.272 -0.714 Decay Class -0.353 0.738 Total Number of Trees 0.947 0.212 Number of Live Trees 0.956 0.115 Number of Snags 0.000 0.392 Percent Overhead Cover -0.029 0.573 Percent of Total Variance explained by each Factor SIRS 25.02 EVERETT AND OTTER: PRESENCE OF CAVITIES IN SNAGS a. Forest n ut = > Département de biologie et Centre d’études nordiques, Pavillon Vachon, Université Laval, Sainte-Foy, Québec G1K 7P4 Canada > Département de biologie et Centre d’ études nordiques, Université du Québec a Rimouski, 300 Allée des Ursulines, Rimouski, Québec GS5L 3A1 Canada + Present address: Environment Canada, Region de Québec, Service Canadien de la faune, 1141 Route de LEglise, C.P. 10100, Ste-Foy, Québec G1 V 4H5 Canada Daigle, Claude, Michel Créte, Louis Lesage, Jean-Pierre Ouellet, and Jean Huot. 2004. Summer diet of two White-tailed Deer, Odocoileus virginianus, populations living at low and high density in southern Québec. Canadian Field-Naturalist 118(3): 360-367. We investigated summer diets of two White-tailed Deer (Odocoileus virginianus) populations through rumen content analyses. Samples from 93 deer were collected in a low density, LD (1 deer/km?) and a high density, HD (14 deer/km?) area of south- ern Québec during the growing seasons of 1997 and 1998. Availability of preferred forage in forests was greater in LD than in HD, whereas agriculture covered a larger proportion of the area in HD than LD. Rumen composition differed between the two populations. Deer from HD consumed less forbs and leaves of shrubs and trees than did LD deer, whereas they con- sumed more fruits, grasses and farm crops. The rarity in HD rumens of food items common in LD, as well as in many parts of the White-tailed Deer range (i.e., Lilliacae), indicated that deer could not compensate for the rarity of preferred forest forage by increasing foraging time and had to feed on cultivated crops. Rumen contents of LD deer had a higher level of cell solubles and lignin, which reflected their greater reliance on quality forage growing in forests. Feeding habits and forage quality can explain why deer body size decreased in HD between the 1970s and 1990s whereas LD deer remained large. Key Words: White-tailed Deer, Odocoileus virginianus, diet, density, forest, agriculture, plant, Québec, Canada Nous avons examiné le régime alimentaire estival de deux populations de cerf de Virginie (Odocoileus virginianus) par analyse du contenu des rumens. Des échantillons ont été prélevés sur 93 cerfs provenant d’un secteur a faible densité de cerfs, LD, (1 cerf/km?) et d’un secteur a haute densité, HD, (14 cerfs/km?) du sud du Québec durant la saison de croissance des végé- taux de 1997 et 1998. La disponibilité des aliments forestiers préférés des cerfs était plus grande dans LD que dans HD alors que la proportion du territoire agricole était supérieure dans HD. La composition des rumens des cerfs de chacune des régions différait. Les cerfs de HD ont consommé de moins grandes proportions de plantes herbacées et de feuilles d’arbustes et d’ar- bres que ceux du secteur LD, mais de plus grandes proportions de fruits, de graminées et de plantes agricoles. La rareté dans les rumens du secteur HD d’aliments communs dans ceux du secteur LD et dans plusieurs autres régions de l’aire de répar- tition du cerf (e.g., Lilliacae), indique que les cerfs du secteur HD ne pouvaient compenser pour la rareté de leurs aliments préférés par un accroissement de la durée de la quéte alimentaire, et qu’ils devaient consommer des plantes agricoles. Les cerfs de LD ont mangé des aliments contenant plus de solubles cellulaires et de lignine que ceux de HD, ce qui refléte une consommation de plantes forestieres de bonne qualité. Le régime alimentaire et la qualité de la nourriture peuvent expliquer pourquoi la taille des cerfs de HD a diminué entre les années 1970 et 1990 alors que celle des cerfs de LD est demeurée grande. White-tailed Deer, Odocoileus virginianus, are selec- tive feeders and although they ingest a wide variety of plants, only a few taxa make up large proportions of their diet (Johnson et al. 1995; Healy 1971; Korschgen et al. 1980; McCaffery et al. 1974). Plant phenology influences their diet; forbs and leaves of deciduous woody plants are common food items in spring and summer, and fruits become major components of the autumn diet. White-tailed Deer also consume mush- rooms in natural ecosystems (Crawford 1982; Johnson et al. 1995; Korschgen et al. 1980; Short 1971; Skinner and Telfer 1974), whereas farm crops can be major food sources in agricultural landscapes (Nixon et al. 1991). During winter, White-tailed Deer switch to twigs of both deciduous and evergreen woody plants and, where snow cover allows access, they also use grasses and farm crops left after harvest (Johnson et al. 1995; McCaffery et al. 1974; McCullough 1985; Nixon et al. 1991; Skinner and Telfer 1974). Forage competition results in consumption of food of reduced quality or availability, which lengthens ingestion and digestion. At high density, forage compe- tition can modify feeding habits of ungulates, in partic- ular for small, selective species (Kie and Bowyer 1999) and reduce body size in cervids (Ashley et al. 1998; Créte et al. 1993; Hjeljord and Histol 1999; Lesage et al. 2001). In mid-latitude deer, forage competition during the growing season likely determines adult body 360 2004 size because growth and replenishment of body reserves occur during this period of the year (Hjeljord and Histol 1999; Lesage et al. 2001; Boucher 2004). On the south shore of the St. Lawrence River in | Québec, the carrying capacity of the deer range de- creases northward because suitable forest stands for wintering become increasingly rare with increasing | winter severity (Boucher 2004). Density averaged ~1 | deer/ km? in northernmost hunting zones of Québec | whereas it locally exceeded 30 deer/km? in southern zones after populations erupted due to mild winters and a conservative hunting regime. Forage competi- _ tion in winter regulates deer numbers in the absence _ of Grey Wolves (Canis lupus) and under conservative | hunting regime (Dumont et al. 2000), but agriculture can help deer to survive winter by providing some food _ during the dormant season (Rouleau et al. 2002a). In recent decades, browsing pressure exhibited a pro- _ nounced gradient on the south shore of the St. Law- rence River, which resulted in a scarcity of preferred summer forage in southern vs. northern zones (Rou- | leau et al. 2002a). Vegetation sampling throughout | the deer range in Québec showed a general pattern of | negative relationship between deer density on one hand, and preferred forest forage and deer size on the _ other (Boucher 2004). Deer living at low density tend- ed to avoid cultivated fields (Lesage et al. 2002) and reached the largest size among Québec deer (Lesage | etal. 2001). We studied the summer feeding habits of White- | tailed Deer in two areas which differed markedly with respect to deer density, forage availability, impor- _ tance of agriculture, and winter severity. The objective _ of this study was to compare composition and quality _ of summer diets of these two White-tailed Deer pop- | ulations. We anticipated that preferred forage of deer | living at low density would be less abundant in rumens of deer living at high density and that the low density population would consume forage of higher quality due to relaxed forage competition during summer. Study Area Samples were collected in a low-density area (LD) and a high-density area (HD) (Figure 1). The LD study area is located in a transition zone between northern hardwood forests and boreal forests (Rowe 1972*). Among trees, Balsam Fir (Abies balsamea), White Spruce (Picea glauca), Eastern White Cedar (Thuja occidentalis), Trembling Aspen (Populus tremuloides), and Yellow Birch (Betula alleghaniensis) dominate (Lamoureux 1994*). Commercial logging has been intensive in this region and the spruce budworm (Choristoneura fumiferana) adversely impacted fir and spruce stands in the late 1970s. Agriculture rep- resents about 10% of the land use in the study area and hay; alfalfa (Medicago sativa), cultivated and up- graded pastures, oat (Avena sativa), and barley (Hor- deum vulgare) cover most cultivated fields. The grow- DAIGLE, CRETE, LESAGE, OUELLET, AND HUoT: SUMMER DIET OF DEER 36] ing season averages 160 days (Wilson 1971*) and snow cover often exceeds 50 cm for more than SO days. Deer density was estimated through double-count aerial surveys (Potvin et al. 1992) and averaged 1.1 deer/km? in summer 1998 (Lamoureux and Pelletier 2000*). Northern hardwood forests cover most of the HD study area. Commercial logging has affected mainly coniferous stands on small areas due to land owner- ship and forest composition (Gosselin 1994*). Most common tree species include Sugar Maple (Acer sac- charum), Red Maple (Acer rubrum), Yellow Birch, American Beech (Fagus grandifolia), White Ash (Frax- inus americana), Balsam Fir, Hemlock (Tsuga cana- densis), Eastern White Cedar, and White Spruce. Forests have been cleared for agriculture throughout the region. They have progressively grown back on abandoned farms; and introduced plants such as apple trees (Malus pumila) are common in today’s forests. Agriculture now covers about 23% of the area. Hay, corn (Zea mays), uncultivated pastures, cultivated and upgraded pastures, and alfalfa dominate cultivat- ed fields. The growing season averages 190 days and snow cover rarely exceeds 50 cm. Deer density aver- aged 14.1 deer/km? in summer 1996 (Dicaire 1999*); locally it may exceed 30 deer/km?. Methods Forage composition — We collected rumen samples from road kills during the growing season (13 May — 2 November 1997 and 1998). Rumen samples were generally collected within 3 days after collision (maxi- mum 5 days), provided that identifiable forage frag- ments remained. Rumen content was hand mixed and two one-litre samples were extracted. The first litre was frozen for forage identification and the other pre- served in 4% formalin for chemical analysis. At the laboratory, frozen samples were thawed and washed through a 7.9 mm and a 4.0 mm mesh size sieve (Créte et al. 1990; Gauthier et al. 1989). About 50 ml of the particles kept by the 4.0 mm sieve were spread into a 20 x 50 cm sampling tray (Chamrad and Box 1964) and covered with 1 cm of water. One hundred particles were systematically selected by point sampling (Créte et al. 1990) and identified through macroscopic and microscopic (10 — 25x) examination to family, genus or species whenever possible. Reference plant samples served for comparison during the identification process. Samples were grouped by taxon and their volume measured by water displacement (+0.25 mL). Plants were grouped into eight categories for statistical analy- ses: graminoids (Graminae and Cyperacae), farm crops, forbs (without flowers or fruits), leaves of shrubs and trees, wild flowers, wild fruits, mushrooms, and un- known. Forage quality — We reasoned that rumens of deer consuming poor quality forage would contain fewer nutrients with rapid assimilation and more nutrients with slow digestibility than counterparts feeding on 362 better quality forage. Poor quality forage could also include more secondary compounds slowing or in- hibiting digestion. We measured cell solubles, lignin, and phenolics as proxy for rapid digestibility, slow digestibility and defensive compounds. We also deter- mined nitrogen (N) content because it proved to be a good indicator of summer diet quality in Caribou (Rangifer tarandus) (Créte et al. 1990). We used the detergent method (Gauthier et al. 1989; Goering and Van Soest 1970) to measure cell solubles (1-NDF) and acid detergent lignin (ADL) (Mould and Robbins 1982). Cell solubles are highly digestible whereas lignin is largely not (Robbins 1993: 251, 294). Total nitrogen content (N) was measured with an automat- ed Macro-Kjeldahl analyzer and total phenolic con- tent was determined by a calorimetric method using the Folin-Ciocalteu reagent (Marigo 1973; Sauvesry et al. 1991). Proteins are also highly digestible (Rob- bins 1993: 294) whereas phenolics includes 8000 sub- stances, of which tannins are known for reducing pro- tein digestion (Robbins 1993: 300). Some samples could not be analysed due to inadequate quantities. All analyses were duplicated or triplicated whenever a difference of more than 5% was found between the first two measurements. The average of the two clos- est estimates served as results. Forage availability — We estimated forage avail- ability during a companion study in the same study areas (Rouleau et al. 2002a). We measured forage bio- mass between the ground and 1.5 m in height at sites used by radio-collared deer in the two areas through- out the growing season with the help of 11 linear regression models. Vertical and lateral plant coverage over 2-m transects served as dependant variables in regression models to predict plant biomass by species (Rouleau et al. 2002b). Data analysis — In a first step, we determined whether sex (for deer >1 year old) and age (fawns vs. >1 year old) influenced forage composition or quality as summer progressed using MANOVA. We restricted the analysis to the HD area because our samples from deer with known sex and age were too small in the LD area. Analyses were performed on sex and age with respect to months, including sex by month, and age by month interactions. We used ANOVA for contrasting forage composi- tion and quality between the two study areas. We pooled months two by two for this analysis to ensure adequate sample size for each period in both regions. ANOVA were followed by pairwise protected LSD tests (PROC GLM, PDIFF option, SAS Institute Inc. 1988). For all analyses, we ensured that residuals were normally distributed (Shapiro-Wilk test) and homog- enous (visual inspection of the plot). For forage com- position, we carried out ANOVA on ranks because we could not normalise residuals. We estimated means and their standard error with the LSMEANS statement (SAS Institute Inc. 1988). THE CANADIAN FIELD-NATURALIST Vol. 118 Results Rumen samples were collected from 93 deer (HD 68 / LD 25): 64 adults, 11 fawns, and 18 of unknown age. On a volumetric basis, 80% of the particles examined could be identified to 68 food items. Males and females showed no overall differences in the food items they selected in the HD area when considering forage categories (F = 0.52; df = 6, 38; P = 0.79) or diet quality (F = 0.21; df = 5, 22; P = 0.95), without sex X age interaction for composition (F = 0.77; df = 24, 134; P = 0.76) and quality (F = 0.69; df = 20, 74; P = 0.83). Neither did age influence forage com- position (F = 0.26; df = 6, 49; P = 0.95) or quality (F = 2.27; df = 5, 28; P = 0.08), nor did it interact with month on a composition (F = 2.31; df = 6, 53; P = 0.05) or quality (F = 1.43; df = 5, 28; P = 0.24) basis. Given these results, we did not take into account age and sex in subsequent analyses. Diet composition — Rumen contents differed between the two study areas (Figure 2). In spring, rumens con- tained no farm crops in LD but 13% in HD (P = 0.03). Grasses (P = 0.08) and flowers (P = 0.07) also tended to be more important food items in HD than in LD. Conversely, forbs (P = 0.04) and leaves of shrubs and trees (P = 0.12) accounted for a larger volume in LD samples. Diets during July-August were more-com- parable; the only significant difference was for forbs: 32% in LD vs. 7% in HD (P = 0.01). Rumens collected during September and October contained more mush- rooms (P < 0.01) and leaves of shrubs and trees (P = 0.08) in LD, whereas those collected in HD includ- ed more fruits (P < 0.01) and grasses (P = 0.08). Differences between the two study areas were more striking when considering taxa for the whole growing season (Table 1). For this comparison, we eliminated fruits, mostly apples, because they were almost restrict- ed to HD and their more three-dimensional shape contrasting with that of most other two-dimensional plant tissues resulted in exceptionally large volume. Excluding fruits, 19 food items each accounted for more than 1% of the rumen volume in LD, summing up to 95 % of volume, compared with 20 food items making up 87 % of rumen content in HD. Forest forbs, such as Yellow Clintonia (Clintonia borealis) and Wild- Lily-of-the-Valley (Maianthemum canadense) were common food items in LD but rare in rumens collect- ed in HD. They were replaced in HD by forbs typical of openings and by farm crops, such as clover (77i- folium sp.) and alfalfa. Mountain Maple (Acer spica- tum) and Mountain Ash (Sorbus americana), the most common shrub leaves in rumens collected in LD, were replaced by Red Maple, Choke Cherry (Prunus vir- giniana), and Sugar Maple in HD; the latter species were also consumed by LD deer, but to a much lesser extent. Finally, apples, the most common item in HD rumens, were found in only one LD sample. Diet quality — Nitrogen (F = 0.91; df = 1, 52; P = 0.34) and phenolic content (F = 0.13; df = 1, 52; P = 0.71) 2004 DAIGLE, CRETE, LESAGE, OUELLET, AND Huot: SUMMER DIET OF DEER 363 | Saint-Jean) | Lac 70° 68° 66° 64° ) Septlles aaa | | fe na | ae et nee lle “ieee, 48° | | White-Tailed Deer rer! distribution Af |__| Study areas Ficure |. White-tailed Deer distribution in Québec and location of the two study areas, low density (LD) and high density (HD). did not differ between the two areas, with nitrogen remaining stable as the growing season progressed, | and phenolic content declining in September-October (Table 2). However, LD rumens contained more cell solubles (F = 5.66; df = 1, 69; P = 0.02) and lignin (F = 4.68; df = 1, 69; P = 0.03) than those from LD. Discussion Diet composition — Forage availability is reflected in diet composition. Although climate differed between LD and HD, the two regions shared most plant taxa and natural forage species had similar digestibility in both areas throughout the growing season (Lesage et al. 2000). However, biomass of preferred forage species found at sites used by deer was much larger in LD than HD (Rouleau et al. 2002a). Food items which domi- nated in LD diets remained almost absent from HD rumens. For instance Liliacae, such as Yellow Clintonia and Wild-Lily-of-the- Valley, which White-tailed Deer normally consume in the Northeast (Crawford 1982; Korschgen et al. 1980; McCaffery et al. 1974; Skinner and Telfer 1974; Waller and Alverson 1997) occupied a marginal volume in HD rumens. Liliacae are partic- ularly vulnerable to deer browsing (Créte et al. 2001; Waller and Alverson 1997) and can be used as indi- cators to estimate the impact of White-tailed Deer on plant communities (Balgooyen and Waller 1995). Wild- Lily-of-the-Valley was common in HD in the early 1980s (Guérard and Legris 1984*). At the time of our study, its biomass was 20 times lower in HD than in LD (Rouleau et al. 2002a) and occurred as a marginal food item in the diet of HD deer (Table 1). Although not abundant on a volumetric basis, Wild-Lily-of-the- Valley occurred in 16% of the rumens collected in HD, showing that deer were still looking for this species even though its availability was largely reduced. Moun- tain Maple, which is known to be affected by deer (Balgooyen and Waller 1995), was a common species in HD in the past (Guérard and Legris 1984*), but seemed almost extirpated at the time of our study (Rouleau et al. 2002a). In LD, where natural forage is abundant (Rouleau et al. 2002a), Lesage et al. (2002) found that deer avoided cultivated fields. Our results support their findings and indicate that White-tailed Deer prefer to consume forest forages when they are readily available during the growing season. In HD, where most preferred food items had become scarce (Rouleau et al. 2002a; Boucher 2004), deer switched to other natural forage still persisting (e.g., Red Maple, Choke Cherry and Sugar Maple) and relied on farm 364 THE CANADIAN FIELD-NATURALIST Vol. 118 May-June GLD gm HD Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms July-August Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms September-October Farm crops Forbs Flowers Fruits Grasses Leaves Mushrooms FIGURE 2. Composition (% volume + SE) of rumen samples collected in a low-density LD (n = 25) and a high-density HD (n = 68) area of southern Québec during the growing seasons of 1997 and 1998. crops for a large part of their summer diet. Comparison Fruits, mostly apples (90%), were a major compo- of forage availability and consumption (Table 1) sug- __ nent of the fall diet in HD. Our results support the idea gests that Red Maple, Choke Cherry, and Sugar Maple that fruits are important food items where and when suffered high browsing pressure in HD. they are available (McCaffery et al. 1974; Short 1971; 2004 DAIGLE, CRETE, LESAGE, OUELLET, AND HUoT: SUMMER DIET OF DEER 365 | TABLE |. Most common food items (volume >1%) found in White-tailed Deer rumens from a low density LD (n=23) and a high density HD area (n=63) of southern Québec during the growing seasons of 1997 and 1998. Skinner and Telfer 1974). Apple trees and oak are absent from forests in LD. We assumed that apples found in one LD sample came from baits set by hunters. Baiting was also common in HD, but we think that apples ingested by deer in this region primarily origi- nated from uncultivated apple trees growing in young forests based on the fact that most specimens found in rumens were smaller than commercial apples. Mushrooms were a common food item in both study areas. Like apples, but to a lesser extent, their shape and size may have exaggerated their relative importance on a volumetric basis. Their frequent occurrence in rumens indicates that they represented a common food item for White-tailed Deer. Mushrooms were consumed mostly in spring in HD and in fall in LD, which may be explained by different seasonal availability or by the possibility that White-tailed Deer feed on them when preferred foods are rare at both ends of the growing season. Many studies reported large amounts of mush- | % volume % occurence Availability (g/m) | Food items Category LD HD LD HD LD HD Leaves, stems, and grain | Acer spicatum Leaf 26.9 0.6 30.4 4.8 19.0 <0.) Fungi Mushroom 14.7 16.4 26.1 20.6 <0. 1 <0.) | Botrychium sp. Forb a2, 0 4.3 0 | Clintonia borealis Forb 6.2 0.1 34.8 1.6 Dei 0.1 | Maianthemum canadense Forb 5.9 Ie) 47.8 15.9 el 0.2 | Sorbus americana Leat 4.4 0 26.1 0 <0.1 | Fraxinus nigra Leaf 4.0 0 4.3 0 | Epilobium angustifolium Forb 3.6 0 PAVE 0 0.8 Prunus virginiana Leaf 3.6 8.9 Paved 28.6 <0.1 <0.1 Prunus pennsylvanica Leaf pos) 0.9 26.1 4.8 <0.1 | Trillium sp. Forb PRS} 0.2 13.0 6.3 <0.1 <0.1 Aralia nudicaulis Forb 23 2.1 8.7 6.3 2.3 <0.1 » Acer rubrum Leaf 1.9 10.3 PSH 41.3 2.6 0.1 | Taraxacum officinale Forb 19 2D 13.0 Vas7) 0.9 0.2 Populus tremuloides Leaf od Ie 13.0 95 4.3 0.1 | Graminae Grass ites) 8.4 43.5 76.2 2.0 | Acer saccharum Leaf 13 6.6 13.0 14.3 1.6 <0.1 Taxus canadensis Leaf 1.3 <0.1 4.3 1.6 | Salix sp. Leaf Les 0.5 8.7 4.8 <0.1 | Solidago sp. Forb 0 1.2 0 4.8 | Fragaria sp. Forb 0 1.5 0 19.0 0.1 _ Lotus corniculatus Farm crop 0 1.6 0 1.6 | Prunus serotina Leaf 0 1) 0 Hilal 0.3 | Sonchus sp. Forb 0.2 1.9 4.3 322 <0.1 | Rubus sp. Leaf 0.7 23 8.7 27.0 | Potemogeton sp. Forb 0 3.0 0 32 | Malus pumila Leaf 0 32 0 6.3 | Zea mays Farm crop 0 3.6 0 7.9 Medicago sativa Farm crop 0 3.8 0 7.9 Trifolium sp. Farm crop 0.1 4.8 4.3 20.6 Fruits Malus pumila Fruit 100 90.1 4.3 36.5 <0.1 <0.1 Quercus sp. Fruit 0 3.6 0 1.6 Fragaria sp. Farm crop 0 Sal 0 1.6 Crataegus sp. Fruit 0 22 0 6.3 rooms in White-tailed Deer diet (Crawford 1982; John- son et al. 1995; Korschgen et al. 1980; McCaffery et al. 1974), but nobody has yet considered the potential impact of deer herbivory on mushrooms. Diet quality - LD deer had access to a greater bio- mass of preferred forage (Rouleau et al. 2002a) and consumed more forbs and leaves of shrubs and trees than HD deer, which compensated by ingesting farm crops, grasses and fruits. Different forage intake result- ed in a diet containing more cell solubles and lignin in LD than in HD deer. Rumen contents did not differ with respect to phenolics, which may have reflected the complex nature of the chemical group (Robbins 1993: 253). Protein content followed a similar trend as that for cell solubles, although the difference between LD and HD was small and non-significant. Between May and September, cell solubles were 8% greater in LD than HD rumens. Cell solubles, which are composed of sugars, protein, non-protein nitrogen, lipid, organic 366 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 2. Mean (SE; n) content (%) of cell solubles, lignin, phenolics, and nitrogen in White-tailed Deer rumen samples collected in a high-density (HD) and a low-density (LD) study area of southern Québec during the growing seasons of 1997 and 1998. May-June HD LD Cell solubles 32.4 40.7 (1.4; 21) (2.0; 14) Lignin 222 23.3 (1.0; 21) (1.2; 14) Phenolics 3i5) 3.6 (0.1; 16) (0.2; 13) Nitrogen Te 7.0 (0.3; 16) (0.3; 13) acid and soluble minerals (Iason and Van Wieren 1999), are highly and rapidly digestible, whereas cell walls (1- cell solubles) are only partly digestible and necessitate some retention in the rumen for microbes to degrade them (Iason and Van Wieren 1999). LD does benefit- ed therefore from a richer diet than HD counterparts during the end of gestation and peak lactation, a peri- od with elevated energy demand in cervids (Mauget et al. 1997). In late summer, the high consumption of apples, which contain little fiber (Robbins 1993), can explain the improved quality of rumen contents in HD. The higher concentration of lignin in LD than in HD rumens likely came from the greater consumption of farm crops and grasses in the latter area, which have a lower level of lignin than natural forages do (Rob- bins et al. 1987). The difference in forage quality likely reduced the growth and condition of White-tailed Deer in HD, where strong competition for forage has prevailed. Deer density in HD increased by 10 fold since the late 1970s, whereas it remained relatively low and stable in LD (Lesage et al. 2001). Deer had the same size in both study areas before 1987 (Potvin 1989), but their body mass progressively decreased in HD (Lesage et al. 2001; Potvin 1994*). At the time of our study, the eviscerated carcass mass of fully grown male deer aver- aged 80 vs. 116 kg in HD and LD, respectively (Lesage et al. 2001). Additional measurements throughout the deer range in Québec revealed that deer size began to decline when forage competition intensified in sum- mer; 1.e., forage availability <10 000 kg/deer (Boucher 2004). Our results concur with the hypothesis that rural deer living at high density cannot fully compensate for the rarity of natural forage in woodlots during summer by consuming cultivated plants (Rouleau et al 2002a). All of these studies suggest that deer populations have reached excessive densities in HD. Acknowledgments We thank R. Bélanger, J. Boisvert, B. Deschamps, and M. Desrochers from Transports Québec, who kind- ly gave us access to deer carcasses. J. Bachand, C. Bélanger, M. Bélanger, N. Bergeron and E. Jeanson helped in collecting the rumen samples. We are grateful to G. Picard for conducting the chemical analysis, to July-August September-October HD LD HD LD 34.1 42.7 40.7 3919 (1.8; 11) (2.1; 4) @i5223) @a72) 21.9 23.4 17.8 24.0 (1.4; 11) (2.3; 4) (1.0; 23) (6322) 3.4 Bi) 2.4 Di (0.2; 7) (0.3; 4) (0.1; 16) (0.4; 2) 6.8 6.5 6.9 7.0 (0.4; 7) (0.6; 4) (0.3; 16) (0.8; 2) N. Desrosiers, and E. Normand for their assistance in the laboratory, and to C. 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Wildlife feeding and nutrition, .2"¢ edi- tion. Academic Press, San Diego. 352 pages. Robbins, C. T., S. Mole, A. E. Hagerman, and T. A. Hanley. 1987. Role of tannins in defending plants against ruminants: reduction in dry matter digestion? Ecology 68: 1606-1615. Rouleau, I., M. Créte, and J.-P. Ouellet. 2002a. Contrasting the summer ecology of White-tailed Deer inhabiting a for- est and a rural landscape. Ecoscience 9: 459-469. Rouleau, I., M. Créte, G. Daigle, and P. Etchevary. 2002b. Rapid estimation of plant biomass used as forage or cover by White-tailed Deer, Odocoileus virginianus, and Snow- shoe Hare, Lepus americanus, in mixed and coniferous for- est of southeastern Québec. The Canadian Field-Naturalist 116: 523-528. SAS Institute Inc. 1988. SAS/STAT User’s guide. SAS Insti- tute Inc. Cary, North Carolina Sauvesty, A. F., F. Page, and J. Huot. 1991. A simple method for extracting plant phenolic compounds. Canadian Jour- nal of Forest Research 22: 654-659. Short, H. L. 1971. Forage digestibility and diet of deer on southern upland range. Journal of Wildlife Management 35: 698-706. Skinner, W. R., and E. S. Telfer. 1974. Spring, summer. and fall foods of deer in New Brunswick. Journal of Wildlife Management 38: 210-214. Waller, D. M., and W. S. Alverson. 1997. The white-tailed deer: a keystone herbivore. Wildlife Society Bulletin 25: 217-226. Received 31 March 2003 Accepted 1 November 2004 Winter Habitat Use by Wolves, Canis lupus, in Relation to Forest Harvesting in West-central Alberta GERALD W. Kuzyk!, JEFF KNETEMAN?, and FIONA K. A. SCHMIEGELOW! ' Department of Renewable Resources, University of Alberta, Edmonton, Alberta, T9H 4N1 Canada > Fish and Wildlife Division, Alberta Sustainable Resource Development, Hinton, Alberta, T7V 2E6 Canada > Corresponding author: gkuzyk @ualberta.ca Kuzyk, Gerald W., Jeff Kneteman, and Fiona K.A. Schmiegelow. 2004. Winter habitat use by Wolves, Canis lupus, in relation to forest harvesting in west-central Alberta. Canadian Field-Naturalist 118(3): 368-375. Forested landscapes in west-central Alberta are facing increased pressures from forest harvesting and other land-use activities, which may alter the movements and distribution of Wolves and ungulates. Information on habitat use by Wolves in logged forests is scarce, potentially limiting effective land-use planning in the boreal forest. Nine Wolves, from four Wolf packs, were fitted with GPS radiocollars in the Rocky Mountain foothills, near Grande Cache, Alberta (2000-2001). We found Wolves did not use the landscape randomly, but rather exhibited a significant preference for non-forested natural habitats (shrubs, water), relative to their availability. Within forest habitats, Wolves used cutblocks proportionately more than unharvested forest and non-forested anthropogenic habitats (pipelines, clearings); however, selection of forest cutblocks was not statistically significant. We found no evidence that Wolves preferred or avoided forest cutblock edges. Wolf pack territories contained various levels of timber harvesting, but most areas were still in the early stages of harvest. Nevertheless, these areas have been allocated for large-scale harvesting. Understanding the potential responses of Wolves to rapidly changing landscape mosaics poses a significant challenge to researchers and managers, but such information is important to informing future land-management and conservation strategies for boreal forest Wolf-prey systems. Key Words: Wolf, Canis lupus, Caribou, Rangifer tarandus, forestry, habitat, Moose, Alces alces, predation, Alberta. Much of the world’s boreal forest is undergoing in- creased demands from resource extraction industries, where related activities such as forest harvesting may alter habitat use by large carnivores (McLellan and Hovey 2001; White et al. 2001). Wolves (Canis lupus) are a common predator of ungulates in Canada’s for- ests, yet there are few data on how their use of habitat might be affected by forest harvesting (Jedrzejewska et al. 1994; Kohira and Rexstad 1997; Kunkel and Pletscher 2000). Logging of forests can change the spa- tial dynamics of Wolves and their prey, resulting in conflict between resource development and wildlife management (Hervieux et al. 1996). For example, in Alberta, Woodland Caribou (Rangifer tarandus caribou) are classed as a threatened species (Edmonds 1998), and Wolf predation is considered a primary reason for their decline (Edmonds 1988; McLoughlin et al. 2003). It is therefore important to understand how Wolves respond to forest harvesting, as habitat use by Wolves may change in response to logging activities, and could affect predation risk to Caribou and other ungulates. Forest harvesting can cause habitat fragmentation and alter predator-prey systems. Predators may follow habitat edges due to ease of travel (Bider 1968). As well, when patch size decreases, predator numbers may increase due to increased prey density and diversity (Gates and Gysel 1978; Yahner 1988). For example, Red Fox (Vulpes vulpes) and Coyote (Canis latrans) densities can increase with more landscape fragmenta- tion, and habitat edges are favored for hunting (Oehler and Litvaitis 1996). In a Wolf-prey system in Canada’s boreal forest, forestry activities have the potential to alter the preda- tion risk to ungulates from Wolves in three ways. First, differential spatial and temporal habitat selection sep- arates Woodland Caribou and Moose (Alces alces) distribution, reducing the risk of predation to Caribou from Wolves (Bergerud and Elliot 1986). Caribou sel- ection for higher elevations reduces the chance of en- counter by Wolves hunting Moose, a primary prey spe- cies, in lower elevations riparian areas (Seip 1992). If the spatial separation of Caribou and Moose is altered by logging roads and forest cutblocks, it has been ar- gued that Wolves will have increased access to, and greater encounter rates with Caribou, resulting in a Cari- bou decline (Bergerud 1988; Seip 1992). In northeast Alberta, linear developments (roads, seismic lines, trails) were found to affect the spatial separation be- tween Caribou and Moose, where linear corridors en- hanced Wolf travel efficiency (James 1999) and Cari- bou mortalities caused by Wolves were found closer to linear corridors than expected by chance (James and Stuart-Smith 2000). Second, Caribou select for older forests (Szkorupa 2002) and distribute at low densities (Bergerud 1988). Caribou spatial overlap with Wolves is reduced and, correspondingly, so too is the risk of detection and pre- dation. Reducing the amount and patch size of older forest by timber harvest may temporarily increase Caribou densities (Bergerud 1988), and Moose, deer (Odocoileus spp.) and Elk (Cervus elaphus) may use these remaining patches for cover. Higher densities of 368 2004 Caribou in more constricted areas, and wider distribu- tion of Wolves in response to primary prey distribution, may increase the chances of Caribou being detected by Wolves. Third, Moose, Elk and deer are attracted to recently logged areas that support high quality regenerating forage (Peek et al. 1976; Tomm et al. 1981; Stelfox et al. 2001). This represents a concentrated prey base for Wolves, which may influence how Wolves use land- scapes. If Wolves frequent forest cutblocks searching for Moose, Elk and deer, and if cutblocks occur near preferred Caribou habitats, this may increase predation risk to Caribou. All three outcomes potentially affect- ing Caribou depend on information about Wolf behav- ior and use of changing landscapes. Forest harvesting in west-central Alberta was large- ly initiated in the late 1960s and has accelerated in recent years. Energy sector activities (oil and gas ex- ploration and development) are also altering these landscapes, resulting in cumulative land use impacts (Hervieux et al. 1996). For decisions concerning long term wildlife conservation, resource managers and land-use planners require new information about how Wolves use habitat in logged forests and under chang- ing landscape conditions. ALBERTA Study area shown indetai 400 Kilomaers 7 gs Wolf locations (GPS) © Wolf pack territory ». Town of Grande Cache — Major Rivers Zones Data Source: Alberta Environment Natural Resources, Govemment of Canada 2000; DEM published with permission of Alberta Sustainable Resource Development. Natural Resource Canada and the Spatial Data Warehouse. Weyerhaeuser Company. KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 369 We used Global Positioning Systems (GPS) radio- collar technology to examine winter habitat use of Wolves in west-central Alberta. We chose to examine fine-scale Wolf movements that correspond with John- son’s (1980) third order habitat selection: movements of animals within their home range. We examined two questions. First, do Wolves use forest cutblocks prefer- entially over other habitat types? Second, do Wolves prefer cutblock edges? We predicted that Wolves would prefer forest cutblocks over other habitats, due to the expected increase in ungulate densities in regenerating forests. We also predicted Wolves would prefer forest cutblock edges relative to areas farther away from them, due to ungulate use of cutblock edges for feeding and proximity to cover (Stelfox et al. 2001*). Study Area The study area is approximately 5000 square kilo- meters, located in the foothills of west-central Alberta, near the town of Grande Cache (54°N 119'W) (Fig- ure 1). The area is classed into subalpine and boreal natural subregions (Beckingham and Archibald 1996), and contains several main rivers and a dendritic pat- tern of creeks; lakes are scarce. Elevations range from 1300-1800 meters, and the climate is subarctic, with Map compied by S. Shirkoffand G. Kuzyk. Unwersty of Alberta FiGure 1. Distribution of four Wolf packs monitored in winters of 2000 and 2001 in west-central Alberta. 370 short wet summers and long cold winters. Temperatures average 16°C in July and -13.5°C in December (Beck- ingham and Archibald 1996). The forests are primarily Lodgepole Pine (Pinus contorta) and some White Spruce (Picea glauca). The wetland complexes support mostly Black Spruce (Picea mariana) and some Tam- arack (Larix laricina). Some south facing slopes sup- port Trembling Aspen (Populus tremuloides) and wil- low (Salix spp.). This area supports a high diversity of large mam- mals: Woodland Caribou, Moose, Elk, Mule Deer (Odocoileus hemionus), White-tailed Deer (Odocoileus virginianus), Bighorn Sheep (Ovis canadensis), Moun- tain Goats (Oreamnos americanus) and Wild Horses (Equus cabalus). Wolves, Coyotes (Canis latrans), Grizzly Bears (Ursus arctos), Black Bears (Ursus americanus) and Cougars (Felis concolor) also exist throughout the study area. Major land use activities include forest harvesting, oil and gas exploration and development, coal mining, commercial trapping, and public uses such as hunting, fishing, hiking, horse packing and camping. Access is primarily on roads created for resource extraction, pipelines and seismic lines. Further descriptions of the study area can be found in Edmonds (1988) and Smith et al. (2000). Wolf Location Data Nine Wolves in four packs were captured and fitted with GPS radiocollars in winters of 2000 and 2001 (Table 1). Three packs were located in areas with a migratory mountain Caribou population and one pack with a sedentary boreal Caribou population. All Wolf handling was approved by the Faculty of Agriculture, Forestry and Home Economics Animal Care Policy (Number 96-99D), subject to the protocols of the Canadian Council of Animal Welfare. Wolf captures were accomplished by helicopter darting (Ballard et al. 1991) or netgunning, then physically restraining the Wolf with restraining forks, and hand-injecting 1-2 mls of telazol at 200 mg/ml (Kuzyk 2002a). Wolves were fitted with store aboard GPS radiocollars (Lotek En- gineering Sytems, Newmarket, Ontario). In the winter of 2001, the Prairie Creek and Cutbank packs each had two members with GPS radiocollars. To avoid pseudo- THE CANADIAN FIELD-NATURALIST Vol. 118 replication (Hurlburt 1984), location data and asso- ciated patterns of habitat use from these individuals were averaged for their respective packs (Table 1). As this study was designed to understand Wolf hab- itat use in winter, the following criteria were used to select Wolf location data: (1) Location data from 31 January to 25 April 25 in 2000 and 2001 were used for analyses. These dates were used for two reasons: first, most Caribou in the study area (the migra- tory mountain ecotype) leave the forested foothills in late winter and spring to calve in the nearby mountains (Edmonds 1988); second, a spring cutoff time also has ecological rele- vance to Wolves. In spring near whelping time, Wolves are thought to change their hunting patterns by switching from hunting as a pack and preying on ungulates, to hunting alone or in small units in search of smaller prey, with their activities centering on the den and pups (Mech 1970). There- fore a single GPS collared Wolf would no longer represent the behavior of their pack, and would not meet our design criteria. (2) To provide consistency in GPS collar programming, six-hour locations were chosen (4 per day) as the minimum common sampling unit for analysis. Wolves are sporadic in their movements, and may travel at rates of about 8 km/hr while hunting (Mech 1966), or relatively short distances when near a killsite (Kuzyk 2002a). When near a killsite, they sel- dom rest in one location for periods longer than six hours (Mech 1970). (3) Wolf locations outside calculated pack territories were not used. These Wolves were assumed to be dispersing from their natal territory and thus behaving differently from their pack (Gese and Mech 1991). Data were differentially corrected using N4Win Version 2.40 program (Lotek Engineering Inc. 2000) and were assumed accurate within 14 meters, 95% of the time (Lotek Engineering Inc. 2000). Wolf locations with Dilution of Precision (DOP) values greater than 15 were removed from the analysis (<2 % of total loca- tions). High DOP values and radiocollar malfunctions made for unequal locations per Wolf pack (range 152 to 279) over the duration of this study (Table 1). Habitat Classification and GIS Methods Wolf location data were imported into ArcView Version 3.1 (Environmental Systems Research Insti- tute Inc., Redlands, California). Current, digital forest inventory coverages were obtained from Weyerhauser Canada Limited, Canadian Forest-Products and Al- TABLE |. Wolf packs with associated number of GPS locations and area of habitat use (territory size) in west-central Alberta in winters of 2000 and 2001. Wolf Pack Wolf Year Dates Number of Locations Area (km?) Cutbank W5 2000 24 January — 16 March 185 714 Prairie Creek W9 2000 28 January — 25 April 252 286 Simonette W13 2000 31 January — 25 April 279 786 *Prairie Creek W5 and W9 2001 18 February — 25 April 231 182 *Cutbank W19andW21 = 2001 15 February — 25 April 258 448 Lynx Creek W22 2001 15 February — 25 April 247 1848 Simonette W30 2001 17 February — 4 April 152 398 * Locations were averaged for two collared Wolves which belonged to the same pack. 2004 KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 37] TABLE 2. Description of habitat categories used in compositional analysis for Wolf packs in west-central Alberta during late winters of 2000 and 2001. Habitat 2 Unharvested forest all harvestable forest burn Habitat | Forest cutblocks all harvested forest Habitat 3 Habitat 4 Non-forest (natural) Non-forest (anthropogenic) herbaceous grassland clearing sand, flooded land right-of-way closed and open shrub industrial infrastructure coniferous scrub pipelines deciduous scrub brush, windfall open and treed muskeg water geophysical perennial forest crops* *Perennial forest crops are denoted as anthropogenic by the forest companies and account for < 0.6km?* of one Wolf pack’s territory. (Simonette pack — total territory size is 786 km7). TABLE 3. The percentage of use (GPS locations) and availability (area in km?) of five habitat categories for Wolf pack territories in west-central Alberta during late winters of 2000 and 2001. Cutblock Forest Non-forest* Non-forest* Non-forest Shrubs Water Anthropogenic Wolf Pack Year Used_1 Avail_1 Used_2 Avail_2 Used_3 Avail_3 Used_4 Avail_4 Used_5 Avail_5 Cutbank 2000 34.1 28.9 57.8 64.6 7.0 A: 1.1 0.3 0.0 0.8 Prairie Creek 2000 Lieve Aes) 77.8 80.2 6.7 4.8 LED, 1.0 2.4 Hes) Simonette 2000 PN SYS) 61.6 78.8 Gs 3.0 DES 0.6 TD 1.8 Prairie Creek 2001 6.3 5.9 81.2 87.1 9.1 4.1 fe 1.1 M7 1.8 Cutbank 2001 Asay SD) 35/9 60.0 20.2 3.4 0.4 0.3 0.4 0.8 Lynx Creek 2001 0.8 5.4 1S} 85.9 202 8.5 0.8 0.1 0.8 0.2 Simonette 2001 16:4: - 1420 68.4 78.3 113) 5) 0.7 0.5 1.3 Le, Total Mean 19.1 16.9 65.7 76.4 1220 5.0 le, 0.6 2.0 1D, SE S27, 4.3 6.0 3.9 23, 0.7 0.3 0.1 0.9 0.2 *Non-forest natural is subdivided into shrub and water categories for descriptive purposes only. berta Government Phase 3. The minimum mapping unit for these coverages was | ha for forest polygons. However, the resolution of line features, such as roads, was much greater, as these were spatially referenced from high-resolution remotely sensed data (~5 m res- olution), or from GPS readings taken in the field. Thus, the resolution of our animal location data was com- mensurate with the resolution of the landscape cover- ages used, for purposes of evaluating coarse-scale habi- tat use. Minimum convex polygons (MCPs) of Wolf territories were initially calculated with an animal movement extension in ArcView (Hooge and Eichen- laub 1997). MCPs were considered an appropriate method for delineating general territory boundaries in order to evaluate coarse-scale habitat use. Due to a small portion of the GIS coverages missing within each Wolf pack territory, territory sizes for subsequent analyses were adjusted by summing all the areas within the MCPs for which we had GIS coverages (Table 1). As Wolves live in a defined territory (Mech 1970), each territory was classified into four habitat categories to reflect coarse scale patterns of use. These categories were: (1) forest cutblocks, (2) unharvested forest, (3) non-forest natural (shrubs and water) and (4) non- forest anthropogenic (pipelines, wellsites) (Table 2). The area of non-forest natural was divided into “shrub” and “water” classes for descriptive purposes (Table 3), but the data were pooled for analysis. As the focus of this study was to determine habitat use of Wolves in managed landscapes, a further analysis was con- ducted to determine Wolf use of forest cutblock edges. Forest cutblocks were buffered using specified dis- tances starting from the edge of the forest-cutblock and proceeding into the forest. We did not use locations inside cutblocks in this analysis. Buffer distances were consistent with those studying Caribou avoidance of linear features (Dyer et al. 2001; Oberg 2001), starting from the edge of the cutblock to 100 m, 101-250 m, 251-500 m, 501-1000 m and >1000 m. The category of >1000 m was also chosen as the farthest distance for comparison to Smith et al. (2000), who found that Caribou in west-central Alberta may avoid cutblocks by about 1200 m. Statistical Analysis Compositional analysis (Aebischer et al. 1993) was conducted by integrating Wolf GPS location data and forest inventory data within a GIS (ArcView 3.1) to B72 determine if there was a preference in Wolf use of habi- tat or buffer categories. Aebischer et al. (1993) suggest a minimum of six radiotagged animals are required to perform compositional analysis, and replication across years is acceptable. Therefore, our sample of seven Wolves over two winters was adequate for this test. Compositional analysis compares the amount of “used habitat” to the amount of “available habitat” and tests whether habitats are preferred or avoided more than expected by random (Johnson 1980). The number of Wolf locations in each habitat or buffer category rep- resented used habitat. The available habitat was the total area of each habitat or buffer category (Table 3). If there was no use of a habitat category, 0% use was replaced with 0.001%, as this represented a value lower than the smallest recorded nonzero percentage (Aebis- cher et al. 1993). A chi-square test was used to determine if Wolf use of habitat or buffer categories was significantly non- random, then each habitat category was ranked in terms of its use. To determine if any habitats were selected over others, a difference for each pair-wise comparison was calculated using log ratios. This compared each habitat category within each Wolf pack territory. The means and standard errors for each comparison were calculated across all Wolf packs, and the pair-wise differences were tested for significance using a t-test (Aebischer et al. 1993). Alpha level for all tests was set at 0.05. Results Wolf Habitat Use Territory size for the four Wolf packs ranged from 182 — 1848 km? (Table 1). The availability of each of the four habitat categories varied markedly: the per- centage of forest averaged 76.40% (SE 3.90) for all packs, and thus was the most dominant habitat, where- as non-forest anthropogenic cover averaged only 1.23% (SE 0.25) of available habitat across Wolf territories (Table 3). Wolves showed a significant deviation from random use of the four habitat types (v7 = 7.815, df = 3, p = 0.036) selecting non-forest natural (shrubs-water) habitats over both forest (t = -4.281, df = 6, p = 0.005) and cutblocks (t = -2.92, df = 6, p = 0.027), in relation to their availability (Table 4). No other pair-wise com- parisons were significant. However, when ranked in preference by habitat type, forest cutblocks were pref- THE CANADIAN FIELD-NATURALIST Vol. 118 erred more than both forest and non-forest anthropo- genic areas. Wolf Response to Forest Cutblock Edges The available areas for all distance buffers less than 1000 m were similar, with variation due mostly to dissolving buffers for adjacent cutblocks. Wolf use of distance buffers did not deviate significantly from ran- dom (x7 = 2.349, df = 3, p = 0.503). When buffer dis- tances were compared using compositional analysis, no significant difference was found between distance categories related to forest cutblock edges. When ranked, the 501-1000 m buffer distance was the most preferred, followed by the 0-100m buffer, with the least preferred being the buffer >1000 m. Discussion Wolves have been described as habitat generalists (Mech 1970; Mladenoff et al. 1995). On a coarse spa- tial scale, Wolves inhabit large tracts of forest (Mech 1995) and may prefer mixed wood forests over either homogenous coniferous or deciduous forests (Mladen- off et al. 1995; Krizan 1997). Wolves may use forests altered by logging, as these areas provide good deer habitat, and thus support an important prey base for Wolves (Mladenoff and Sickley 1998). In this study, GPS radiocollar technology allowed for a more re- fined examination of Wolf habitat preferences, show- ing Wolves in our study area do not use the landscape randomly. In general, Wolves preferred habitats with young vegetation, in both non-forest natural habitats and forest cutblocks. This is consistent with increased ungulate abundance in areas of young vegetation (Peek et al. 1976; Stelfox et al. 2001*), which attract Wolves (Bergerud 1988). However, increased road access into these areas may also allow humans to alter activity patterns of Wolves (Theuerkauf et al. 2003), or affect Wolf numbers by direct or indirect killing (Mech 1995, Mladenoff and Sickley 1998). In this study, the least used habitat by Wolves was non-forest anthro- pogenic (pipelines, right-of-ways), possibly to avoid human contact. Two radio-collared Wolves were known to have been shot during this study, and several other collars were lost to unknown factors (Kuzyk 2002a). We found little support for our first prediction that Wolves select forest cutblocks. Wolves did use cut- blocks proportionately more than their availability and were ranked above forest or anthropogenic features, TABLE 4. Results from compositional analysis (p values in parenthesis; + denotes row > column and — column > row) for comparing four habitat categories for four Wolf packs in west-central Alberta during late winters of 2000 and 2001. 1 2) Cutblock Forest 1 Cutblock +(0.947) 2 Forest - 3 Non-for_natural + + 4 Non-for_anthro. - - * denotes significance at (p<0.05). 3 4 Non-for_natural Non-for_anthropogenic -(0.027) * +(0.902) -(0.005) * +(0.903) +(0.177) — 2004 | but differences were not significant. Similarly, Wolves _ in Ontario were found to use cutblocks in proportion to their occurrence (Krizan 1997). We had the advan- tage of GPS radiocollars which allowed for a large collection of location data, compared with the traditi- onal VHE collars used by Krizan (1997). However, we acknowledge that our analyses still lacked statistical power, due to the relatively small sample of Wolves radiocollared. The amount of harvested forest differed substan- | tially between Wolf packs and may have accounted | for variation in Wolf use of forest cutblocks. The Lynx Creek pack had only 5% of its territory as forest cut- blocks, whereas 36% of the Cutbank pack’s territory consisted of recent cutblocks. This seven-fold differ- ence in the amount of harvested forest between packs may have influenced habitat preferences. Kohira and Rexstad (1997) found no evidence that Wolf diets differed between logged and unlogged areas in the coastal rainforests of Alaska. About 6% of that total study area was logged, with the amount of area logged ranging from 1-26% between Wolf pack territories. This differs from our study area, where about 17%, or approximately three times as much area has been logged. In southeast British Columbia, researchers also did not find evidence that forest harvesting increased the vulnerability of Moose to predation by Wolves, where about 13% of the area was logged (Kunkel and Pletscher 2000). Wolves in our study did show a significant prefer- ence for non-forested natural habitats (shrubs/water) over cutblocks and forest, which might be explained by a number of selection criteria. Wolves prefer to rest in open areas, and may travel several kilometers to reach such preferred sites (Mech 1970). Wolves in this study area were observed on numerous occasions resting in open meadows, muskegs, hillsides and beav- er ponds, often when they were near killsites (Kuzyk 2002a). The shrubs in these habitats have open crowns, which allows both penetration of sunlight and structure for protection from the wind, thus providing Wolves cover while resting. The shrubs in this non-forest habitat type also pro- vide forage and cover for ungulates. During this study, Wolves made deer kills in shrubby willow areas, and Moose, deer and Elk kills in or near riparian areas (Kuzyk 2002a). Elk are primarily grazers, and may be attracted to these shrub patches due to the increased availability of grasses. Bjorge and Gunson (1989), in a nearby Wolf study, noted that Elk, especially Elk calves, were a preferred prey for Wolves in winter. During the limited kill rate work associated with this study (Kuz- yk 2002a), only one cow Elk kill was documented. In Jasper National Park, Wolves hunt deer while mov- ing to pockets of Elk (Carbyn 1974; Weaver 1994). It is possible that shrubby areas do represent reliable patches of prey, and the Wolves investigate them for prey regularly. KUZYK, KNETEMAN, and SCHMIEGELOW: HABITAT USE BY WOLVES 373, Water was also included in this preferred habitat class. It is common for Wolves to use frozen waterways as travel routes where snow is most compacted and ice makes travel easy. In winter Wolves probably use creeks and rivers to travel among ungulate winter ranges. Also there is an increased chance of encounter- ing Moose that use riparian areas in winter (Hayes et al. 2000) and Wolves are known to frequently kill ungulates on iced surfaces (Mech 1991). Our second prediction of Wolf preference for for- est cutblock edges was also not supported. There was no significant difference in Wolf preference for any buffer distance categories, nor was the 0-100 m buffer class ranked highest. Habitat was not controlled for in the buffer categories, which may have confounded the analysis. The behaviors of Wolves, such as feeding at killsites, resting and hunting may also be diluting the effect of any preference or avoidance of cutblock edges. Wolves hunt a diversity of prey, and chase dis- tance varies with each prey type (Paquet 1989). For example, the average chase distance for a Moose is 883 m (Paquet 1989). Depending on where Wolves lo- cate them, Moose could choose to run to the nearest forest to avoid attacking Wolves (Stephens and Peter- son 1984), or remain stationary and aggressive (Mech 1966), or stationary and non-aggressive (Kuzyk 2002b), and still avoid attack by Wolves. These results, and those related to broader habitat selection questions, suggest that consideration of behaviors associated with different habitat types is an important component of interpretation in Wolf habitat use studies. Forest harvesting alters both the amount and spatial distribution of habitat types. We measured habitat use by Wolves directly, and found that Wolf use of land- scapes was not random. We suggest that patterns of habitat use may be influenced by the relative avail- ability of different habitat types, specifically natural shrubs and waterways, and to a lesser degree, by re- cent forest cutblocks. Nevertheless, our results clearly show that consideration of shrub/waterway habitats is an important criterion for land-use decisions regard- ing Wolves. In our study area, Caribou prefer forests greater than 80 years old, especially those stands aged 120-160 years (Szkorupa 2002), and have been found to avoid forest cutblocks by 1200 m (Smith et al. 2000). As most of the winter range of these Caribou has been allocated for timber extraction, areas of older forest will become increasingly small and isolated. If the forest continues to be harvested at present rates, all Wolf packs we studied will have a substantial amount of logged area within their territories within a relatively short time. As Moose, deer and Elk are the primary prey of Wolves in this study area, informa- tion is required on how these ungulates are respond- ing to the changing landscape mosaic, as this may ultimately determine how Wolves use the landscape. Understanding the dynamic relationship between pred- ator and prey in a system undergoing rapid change 374 poses an enormous challenge. Future research on Wolf habitat use should concentrate on increasing sample sizes, refining habitat classifications, and linking be- havior with patterns of habitat use. Acknowledgments Funding for this research was provided by the West- Central Alberta Caribou Standing Committee, the Al- berta Sport, Recreation, Parks and Wildlife Foundation, and a University of Alberta Challenge Grant in Bio- diversity (supported by the Alberta Conservation Asso- ciation). A University of Alberta Graduate Research Assistantship and Margaret Brown Award in Environ- mental Studies and Wildlife Resources also provided financial assistance to G. Kuzyk. Additional financial support and digital forest inventory data were kindly provided by Weyerhaeuser Canada Ltd. and Cana- dian Forest Products. S. Shirkoff provided GIS sup- port. 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Identifying the bird communities of a habitat could contribute to conservation efforts and provide benchmarks for ecosystem studies. Garry Oak (Quercus garryana) ecosystems in British Columbia are among the most endangered in Canada and warrant conservation. Four bird communities were determined by analyzing an extensive sample of Garry Oak habitat bird data. These communities were defined objectively by aggregations of the bird species themselves from across the various sites and areas. Characteristic species of these communities include American Goldfinch (Carduelis tristis), Spotted Towhee (Pipilo maculatus), Bewick’s Wren (Thryomanes bewickii), Rufous Hummingbird (Selasphorus rufus) and Chestnut-backed Chickadee (Poecile rufescens) in community 1; House Wren (Troglodytes aedon), Olive-sided Flycatcher (Contopus cooper), Purple Finch (Carpodacus purpureus), White-crowned Sparrow (Zonotrichia leucophrys), and Pine Siskin (Carduelis pinus) in community 2; Western Tanager (Piranga ludoviciana), Yellow-rumped Warbler (Dendroica coronata), American Goldfinch, Pine Siskin, Pacific-slope Flycatcher (Empidonax difficilis), Cassin’s Vireo (Vireo cassinii) and Chipping Sparrow (Spizella passerina) in community 3; and Northwestern Crow (Corvus caurinus) and European Starling (Sturnus vulgaris) in commu- nity 4. Differences between the communities are suggested from the life history traits of the species, including a community consisting mostly of insectivores when on breeding territory (number 1), one with species foraging primarily in shrubs and trees (community 3), and another with tree-nesting ground gleaners (number 4). One community (number 3) had analogues in two widely disparate areas: oak-associated in north-central New Mexico, and aspen (Populus tremuloides)-related in northcentral British Columbia; otherwise communities reported in the literature were generally not directly comparable. Key Words: bird communities, Garry Oak, Quercus garryana, multivariate classification, British Columbia. Garry Oak, Quercus garryana habitat is unique within Canada, and related biogeographically to Cal- ifornia (Erickson 1996). Garry Oak ecosystems are at their northern margin in British Columbia, and are among the most endangered ecosystems in Canada (Erickson 1993*, 2000a). Native stands have been reduced by urban development and are threatened with invasion by alien species. Bird communities of Garry Oak habitat in the Pacific Northwest have not yet been fully investigated. Defining these communi- ties is a basic step toward conservation, and a requisite to identifying critical habitat requirements. Predictive abilities and assessments for preservation and man- agement of the Garry Oak habitat could be strength- ened by further understanding these communities. Garry Oak occurs in an area of southwestern British Columbia with a distinctive modified mediterranean climate in the strong rain-shadow of the Olympic and Vancouver Island mountains. Mild, wet winters, vari- able precipitation and regular summer drought are typ- ical. The result on the landscape is a mosaic of park- lands with spring forb meadows and oak clumps; mossy bluffs often with shrub oaks; open grassy savannahs; and woodlands sometimes mixed with Douglas-fir (Pseudotsuga menziesii). As well as being diverse and productive for plant growth, this landscape is attractive for human habitation and agricultural development. The consequence has been habitat loss and endanger- ment for Garry Oak ecosystems. There is a scientific and conservation interest in knowing more about the elements of these ecosystems, including their bird communities. Garry Oak habitat has been selected as a focus of Partners in Flight for its importance to migrating and wintering birds in the Pacific Northwest (de Groot et al. 2000). Their work includes restoration of habitat within the British Columbia range of Garry Oak (Figure1). The purpose of this paper is to identify the bird com- munities of Garry Oak habitat, make comparisons of the life history traits of constituent species, and com- pare these communities with the literature on other habitats. Methods I used Detrended Correspondance Analysis (DCA, PCORD 3.0, McCune and Mefford 1997*) (Hill and Gauch 1980; Peet et al. 1988); interpretive graphing; and a quantitative similarity index (Motyka’s modifi- cation of the Sorenson index, Brower et al. 1990): to determine bird communities. I took records of bird species occurrence by ear and by sight while sampling representative ecological plots over 120 Garry Oak areas (Figure 2). This was part of an ecological reconnaissance in 1993 and 1994, cov- 376 2004 in British Columbia | Ficure 1. Garry Oak range in British Columbia. This map is courtesy of British Columbia Ministry of Water, Land and Air Protection, December 2003. ering an area bounded by East Sooke, Gonzales Hill and Courtenay on Vancouver Island; and East Point, Saturna Island, and Helliwell Point, Hornby Island on the Gulf Islands (Erickson 1996). These locations are between approximately 48°N, 123°W and 49°30'N, 125°W. The Garry Oak stands sampled were primarily mature, but their canopy varied from shrub-like with exposure, to large and open on deep soils. Elevations ranged from sea level to about 550 m. Plots were approximately 200 m?, consisted of relatively uniform vegetation and topography, and varied in size accord- ing to plant community boundaries. Species presence was recorded in an observation effort of approximately 90 minutes per plot. A total of 1243 records were taken ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT SEH on 286 plots (Erickson 2000b). Sampling was primarily in spring, during May and June. This time interval has migratory bird influx, spring vegetation growth, territory establishment, nesting, summer plant growth cessation and bird dispersal. The data set had been previously checked with spe- cies accumulation curves (McCune and Mefford 1997; Erickson 2000b, unpublished data; Smith et al. 2002) to determine the adequacy of the sample. Occurrence by plot of all species with a frequency 5% for each year was entered into a data set; 23 species in 1993, and 29 species in 1994. This qualifying criterion reduced the number of plots to 135 in 1993 and 127 in 1994, for a total of 262 in the analysis. The DCA method uses chi-squared distances to simultaneously ordinate, in this case, bird species data against plots, and vice-versa. Axis solutions account for, and therefore represent the most variation in the data set. Bird species are sepa- rated and referenced by their scores relative to the de- trended and re-scaled axes. Graphing the axis combi- nations provides a view of the reference coordinates in multivariate space. DCA is among the most widely used analytical methods in ecology (Peet et al. 1988). It has been applied in bird community work in oak woodland and other settings (Huff and Raley 1991; Pojar 1995; Garcia et al. 1998; Abernethy et al. 2001*; Parody et al. 2001). In the interpretive graphing method I framed pro- portional circles on Axis | vs. 2, and Axis 1 vs. 3, on the DCA output graphs in order to outline potential bird groups. The circles represent four potential bird groups for each year’s data, a number which was deter- mined from species accumulation curve results. These Helliwell Point Sea ) Nanaimo Vancouver Island $ So [Sime eames | re; <1 UU ¢ = 5 oe. | S eS oe Cone E : : Se Georgia Strait Ne Tee xX Sea 2e NS SS x ; SS East Point se = Gonzales Hill +etoria Notes: 1 to 10 plots per circle. Note lateral map distortion \ East Sooke Sse RS ES] FiGure 2. Garry Oak bird community sampling on Vancouver Island, British Columbia. 378 TABLE |. Characteristic species classification. Species classification Symbol and similarity value d (10) c (7) n (3) Distinct Companion Non differential results indicated that groups of approximately 35 plots could potentially cover >60% of the total species. The circles were centred on the graph coordinates from the median of the top-ranking species. Species rank was judged from the scores against each axis and the dominant axis combination (1 and 2) in the DCA result (Table 1, 2). These groups quantitatively represent the co-occurrence of species in different plots from the data, and they served as the first stage in identifying bird communities. Species were classified according to their distinctness to a group from the graphs and numer- ic values assigned (Table 1). Quantitative similarity index comparisons were then completed and the groups aggregated into communities based on the results. Results Species scores against the DCA axes are shown in Tables 2, 3 and 4. Four bird communities of Garry Oak habitat were identified by the analysis (Figures 3-6). Combinations of characteristic species, includ- ing most of the twenty top-ranking species in overall frequency, helped define the communities (Table 5). Characteristic species included American Goldfinch (Carduelis tristis), Spotted! Towhee (Pipilo macula- tus), Bewick’s Wren (Thryomanes bewickii), Rufous Hummingbird (Selasphorus rufus) and Chestnut-backed Chickadee (Poecile rufescens) (community 1); House Wren (Zroglodytes aedon), Olive-sided Flycatcher (Con- topus cooperi), Purple Finch (Carpodacus purpureus), White-crowned Sparrow (Zonotrichia leucophrys), and Pine Siskin (Carduelis pinus) (community 2); West- ern Tanager (Piranga ludoviciana), Yellow-rumped Warbler (Dendroica coronata), American Goldfinch, Pine Siskin, Pacific-coast Flycatcher (Empidonax dif- ficilis), Cassin’s' Vireo (Vireo cassinii) and Chipping Sparrow (Spizella passerine) (community 3- 1993 only); and Northwestern Crow (Corvus caurinus) and Euro- pean Starling (Sturnus vulgaris) (community 4- 1994 only). Three groups had resulted in each of the two years, which were then reduced to the four communities. Two of these occurred across the years and two were unique to a single year. Communities were defined by the combination of their characteristic species. Simi- larity index values were quite low for the two across- year communities: 0.36 for community | and 0.43 for community 2. The two within-year communities had no similarity to each other (0 index value). THE CANADIAN FIELD-NATURALIST Vol. 118 Association with a group on both axis combinations unique to a group on both axis combinations unique to the group on one axis combination distinct on one axis combination TABLE 2. Garry Oak bird species in 1993 with the highest scores against DCA axes, with median species designated*. Species Score Axis |: Northwestern Crow 522 Song Sparrow 499 California Quail* 388 Spotted Towhee* 388 Cedar Waxwing 388 Bewick’s Wren 378 Axis 2: Cedar Waxwing 495 Pine Siskin 446 Bewick’s Wren 400 Western Tanager* 388 Cassin’s Vireo* 384 Yellow-rumped Warbler 384 California Quail Sl Song Sparrow 345 Axis 3: Chestnut-backed Chickadee 428 Rufous Hummingbird 336 Northern Flicker* 318 Pacific slope Flycatcher 294 Cedar Waxwing 2S) TABLE 3. Garry Oak bird species in 1993 and 1994 with the highest scores against DCA axis combination | and 2, with median species designated*. Year and Species Score Score 1993 axis | axis 2 California Quail 388 371 White-crowned Sparrow 183 250 Olive-sided Flycatcher* 119 230 American Goldfinch 302 342 House Wren 69 135 1994 Brown Creeper 222 230 Rufous Hummingbird 406 275 Brown-headed Cowbird* 373 262 Northern Flicker* 258 426 Pacific-slope Flycatcher 216 162 Cassin’s Vireo 254 224 Note: The former common names (Common Flicker, Rufous- sided Towhee, Common Starling, and Solitary Vireo) were used in the data and Figures for Northern Flicker, Spotted Towhee, European Starling and Cassin’s Vireo. ' The former common names, (Common Flicker, Rufous-sided Towhee, Common Starling, and Solitary Vireo) were used in the data and Figures 1-4 for Northern Flicker, Spotted Towhee, European Starling and Cassin’s Vireo. 2004 TABLE 4. Garry Oak bird species in 1994 with the highest scores against DCA axes, with median species designated*. Species Score Axis 1: Song Sparrow 583 Northwestern Crow* 460 European Starling* 420 Rufous Hummingbird 406 Axis 2: Common Flicker 426 White-crowned Sparrow 40] Song Sparrow* 294 Chipping Sparrow 283 Rufous Hummingbird PAIS Axis 3: European starling 506 Violet-green Swallow 396 Olive-sided Flycatcher* 378 House Wren* 358 White-crowned Sparrow 327 Spotted Towhee 326 Note: The former common names (Common Flicker, Rufous- sided Towhee, Common Starling, and Solitary Vireo) were used in the data and Figures for Northern Flicker, Spotted Towhee, European Starling and Cassin’s Vireo. The overall frequency (Erickson 2000b) of these characteristic species gives an indication of the abun- dance of the communities. Community 1 was most fre- quent, with the #3-rank Spotted Towhee; the #4 Chest- nut-backed Chickadee; #6 American Goldfinch; #14 Bewick’s Wren and #16-rank Rufous Hummingbird. 4PiSi Unngmed group Community 3 SomMRWa ele Axis | ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT 379 Community 2 had the #5-rank White-crowned Sparrow; #10 House Wren; #12 Pine Siskin; #13 Olive-sided Flycatcher; and the #19-rank Purple Finch. Commu- nity 3 had the #6-rank, American Goldfinch; #7 Pacific-coast Flycatcher; #11 Chipping Sparrow; #12 Pine Siskin; #18 Yellow-rumped Warbler; #21 Cassin’s Vireo; and #24-rank Western Tanager. Com- munity 4 had the #15 rank Northwestern Crow and the #26 rank European Starling. Discussion Sampling and analysis Communities are differentiated here by their visual distinctness in the multivariate space defined by bird frequency and composition, and by differences defined using thresholds in quantitative index of similarity com- parisons. Both measures use quantity along with com- position, which ensures that the communities arising from the data are actually different from each other. Although eigenvalues can measure the overall strength of a multivariate relationship, such as the multivariate coefficient of variation for each axis in an ordination, they are not a test statistic in Detrended Correspondance Analysis.This method uses detrending and rescaling to avoid the spurious, secondary arch effect of previous techniques, but this in turn prevents the use of eigen- values. Although the differences by year were signifi- cant for many species in previous t-test comparisons (Erickson, unpublished data), these tests are not used here, as they are not appropriate for analysis of non- experimental survey data (e.g., Hurlbert 1984). Sampling occurred on discrete but much smaller (0.02 ha) plots in this study than in many others. How- se ogee nl 2 OCWa Community 4 4NWCr FiGurE 3. Garry Oak bird communities from DCA analysis of 1993 data, Axis 1 vs. 2. Large square represents the multi- variate space formed from the data centred on the DCA axes. Circles are objectively defined groups or communities. Small square is a subjectively placed community. Bird species coordinates are marked by a cross and designated by the first letters of the common name (former names as noted in the text). Other data points are the plot coordinates. 380 THE CANADIAN FIELD-NATURALIST Vol. 118 Cc ity 4 Unnamed group saa Community 3 | ” PSEI aI sar SowRWa FiGure 4. Garry Oak bird communities from DCA analysis of 1993 data, Axis | vs. 3. Large square represents the multi- variate space formed from the data centred on the DCA axes. Circles are objectively defined groups or communities. Small square is a subjectively placed community. Bird species coordinates are marked by a cross and designated by the first letters of the common name (former names as noted in the text). Other data points are the plot coordinates. oF {ChSp Vie f Community 3 GS ; Fh OWA / ie een ies | w Axis 2 Conur a: 4WCSp Community | 94 group 4 DSp Ficure 5. Garry Oak bird communities from DCA analysis of 1994 data, Axis 1 vs. 2. See explanation for Figure 4. ever, the number of plots was correspondingly larger (262 plots) and records were taken over a much longer observation period (90 minutes per plot) than in many studies. Consequently, the results do not represent ex- treme low values in comparison to studies using other methods in oak woodlands. In my species accumula- tion curves, about one half the species were covered by 15 randomly selected plots, and three-quarters of the species by 45 plots (Erickson 2000a). The average number of species detections per plot (3.7, 1993; 5.0, 1994: Erickson 2000a) is from the same order of magnitude for two count periods (3 species, 5 species per 0.28 ha plot) in Jalisco, Mexico, oak woodland; but lower than two other count peri- 2004 Community 2 / SEI -HoWr | 4GSw chSp \ aQn \ 4RBNu Axis 3 | ; ” deel \ PSFI Axis | ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT \ REC'Bp 4PuFi th.QCWa? NRG oe eee SvaTBCh |: Community3—\, Community 1 CeWa 381 CoSt Community 4 I it eet AS 94 group 4 BHCo \ 4RuHu, FIGURE 6. Garry Oak bird communities from DCA analysis of 1994 data, Axis 1 vs. 3. See explanation for Figure 4. _ ods (10, 12 species: Corcuera and Butterfield 1999). _ My averages were much lower than those recorded on multiple plot, large-area studies: 21 to 25 late-spring _ species on five, 70 to 130 ha, stands in Oregon (Ander- _ son 1970), and 9 to 29 breeding species from 40 ha oak census plots in Pennsylvania (Probst 1980). My total number of species detections (66) is higher than in a number of studies in oak woodland (e.g., 38 spe- cies in Dedon et al. 1984; 20 to 62 species in Leidolf et al. 2000; 43 to 51 species in Stone no date*; 50 and 58 species in each of two seasons, five forest types, Corcuera and Butterfield 1999) but is lower than in some other comparable results (e.g., 77 species in Gar- cia et al. 1998). My total would be increased by an additional 12 species which I recorded as “out of plot’, “out of habitat’, or “overhead”. However, much of this difference is irrelevant, in that my interest in defining bird communities was at the plot (not area) level, and was focused on frequently occurring species which are easily detected within my plot parameters, not on less common species used to round out a full species list. The graphical location method is an objective one, in its centering technique and the proportional alloca- tion of multivariate space to each community. Unbi- ased shapes, such as squares or circles, were used to define multivariate species aggregations and overlap in composition was accepted. Other methods include subjective ones (e.g., Pojar 1995), and other objective approaches, such as the use of a “fuzzy-clustering partition coefficient” (Abernethy et al. 2001*). These methods tend to result in elliptic forms, irregular bound- aries and volumes of multivariate space, and no overlap in composition. The elliptical forms assume a trend in dimensionality that does not actually occur with the DCA methodology, as it has demonstrated potential to produce evenly distibuted sample points in multi- variate space (Peet et al. 1988). Communities ; Of the six groups first resulting from the DCA analy- sis, two from each year were similar enough (similarity index > 0.33) to combine. One group from 1994 was dropped because it had less than my conceptual thresh- old of three species. Two within-year communities were not differentiated by the results of the other year, but it was possible to represent them subjectively. To do so, I placed the square frame (Figures 3-6) on the graphs of the other year, and in doing so, included species and multivariate space not already covered. Through this process, Dark-eyed Junco (Junco hyemalis) was added to community 3 as a distinct species in 1994, although in 1993 it was limited to one of the two one-axis com- binations. The two top-ranking species in frequency, Orange-crowned Warbler (Vermivora celata) and Amer- ican Robin (Turdus migratorius) did not contribute to the characteristic combination of species for the com- munities. American Robin did qualify as a character- istic species, but was not used because it was in three out of four communities. Consideration was given to dropping another group which occurred only in 1994, as it had only two dis- tinct species, Northwestern Crow and European Star- ling. However, it was kept (as Community 4) because these two species are potentially important as ecolog- ical indicators. They may both signify a disturbance zone and represent a source of disturbance themselves, via nest predation and nest cavity competition. In addition, this community can be characterized by the absence of other species. There may be an inversely 382 THE CANADIAN FIELD-NATURALIST TABLE 5. Bird communities and characteristic species of Garry Oak habitat. Species Group 94-3 Community | American Goldfinch Spotted Towhee Bewick’s Wren Rufous Hummingbird Chestnut-backed Chickadee Community 2 House Wren Olive-sided Flycatcher Purple Finch White-crowned Sparrow Pine Siskin Community 3 Western Tanager Yellow-rumped Warbler Pacific-slope Flycatcher Cassin’s Vireo Chipping Sparrow Dark-eyed Junco* Community 4 European Starling Northwestern Crow Other species: Song Sparrow n California Quail d Violet-green Swallow Orange-crowned Warbler n Townsend’s Warbler Brown Creeper Cedar Waxwing n Swainson’s Thrush n Common Flicker American Robin n n Vol. 118 93-3 94-2 93-2 94-1 Cc (¢ (e n n n nN d d Cc d (e Cc n Cc n n n n (© d d d n Cc (Gj c c n d d (& n n n n n n n Nn (© Cc Notes: Communities are shown in the boxes. See Table | for species classification codes. The addition of Dark-eyed Junco to community 3 is partly subjectively derived. proportional relationship to the presence of North- western Crow and European Starling across various sites. In this study, the lack of importance of another disturbance indicator, Brown-headed Cowbird (Molo- thrus ater), contrasts with the high numbers of cowbird nestlings on two Garry Oak sites west of Victoria, British Columbia (Shepard 2000). However, the present results reflect only detections of adults and fledged birds in a reconnaissance investigation. Cross-year similarity index values were relatively low, approximately one-half of my expected threshold value of 0.66. Therefore I considered using the concept of assemblages to denote a weaker level of association. However, both the terms community (e.g., Huff and Raley 1991; Pojar 1995; Garcia et al. 1998; Abernethy et al. 2001*; Parody et al. 2001) and assemblage (e.g., Manuwal 1986; Corcuera and Butterfield 1999; Hagar and Stern 2001) receive only general use in the litera- ture, and the term community is referred to more wide- ly. The year differences I encountered could relate to increases in bird frequency and dominance shifts in 1994, possibly linked to yearly weather (Haila et al. 1993; Erickson 2000b). The communities may differ in the life history traits of their characteristic species, as shown in Table 6 (Leidolf et al. 2000; Ehrlich et al. 1988), and there- fore in their general habitat and guild use. Species of community | have mixed nest locations and most are insectivorous when on breedng territory. Those of com- munity 2 have an equal dominance of granivores with insectivores. Most species of community 3 forage in shrubs and trees. Community 4 species are tree-nest- 2004 Community and species Community 1: Nest Location TABLE 6. Life history traits for characteristic species of Garry Oak bird communities. Foraging Layer ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT American Goldfinch SH Spotted Towhee GR/ SH Bewick’s Wren TR Rufous Hummingbird VN/TR Chestnut-backed Chickadee SN American Robin SH/ TR Community 2: House Wren TR Olive-sided Flycatcher TR Purple Finch TR American Robin SH/ TR White-crowned Sparrow SH Pine Siskin TR Community 3: Western Tanager TR Yellow-rumped Warbler TR American Goldfinch SH American Robin SH/ TR Pine Siskin TR Pacific-slope Flycatcher TR Cassin’s Vireo TR Chipping Sparrow SH/ TR Dark-eyed Junco GR Community 4: Northwestern Crow TR European Starling TR Foraging Method Food Type HB/ SH/ TR FG GV GR GG IN/ OM GR GG IN HB HG NE SH/ TR BG IN GR/ SH/ TR GG/FG IN/ FR GR GG IN AIR HA IN GR GG GV GR/ SH/ TR GG/ FG IN/ FR GR GG GV HB/ SH/ TR FG GV SH/ TR FG IN SH/ TR FG IN HB/ SH/ TR FG GV GR/ SH/ TR GG/FG IN/ FR HB/ SH/ TR FG GV AIR HA IN SH/ TR FG IN GR GG IN/ GV GR GG GV GR GG OM GR GG IN Note: Nest location: TR-tree, VN-vine; GR-ground, SH-shrub, SN-snag. Foraging Layer: AIR-air, GR-ground, HB-herb, SH- shrub, TR-tree. Foraging Method: BG-bark glean, FG-foliage glean, GG-ground glean, HA-hawk, HG-hover and glean. Food Type: FR-frugivore; GV-granivore; IN-insectivore; NE-nectarivore, OM-omnivore. ing ground gleaners. Shared traits are as follows: tree nesting insectivores (community 2,3) and ground glean- ing (1,2,4). The comparison may suggest general habi- tat and guild relationships for the communities. This could include open savanna settings (community 2,4) and shrubby woodlands or shrub oak (community 1,3). More work is needed on habitat relationships though. Comparison with other studies In this study, bird communities have been delineat- ed within the Garry Oak habitat at a detailed level by aggregations of the bird species themselves. This ap- proach is not typical in the literature. There are indica- tions of different bird communities in the discussion of Hagar and Stern (2001) for Garry Oak woodlands, including Chipping Sparrow as a species of semi-open woodland and Orange-crowned Warbler as shrub- associated. Dedon et al. (1984) employed habitat suit- ability class lists, possibly comparable to more detailed communities, in California Black Oak (Quercus kel- loggii) habitat. One shrub community was identified to subdivide Gambel Oak (Quercus gambellii) wood- lands (Stone no date*) in Colorado. Artman et al. (2001) found in Ohio that differences in bird commu- nities within mixed oak forests could be related to three moisture index classes. Pojar (1995) described separate bird communities within Trembling Aspen forests of north-central British Columbia. In most oak woodland bird studies, bird communities have been identified at a more general level. Broad habitat groupings have formed bird communities, guilds have been separated by shared traits, or com- munities have been grouped by successional stages (e.g., Anderson 1970; Smith 1977; Probst 1980; Davis et al. 2000; Hagar and Stern 2001). Both Leidolf et al (2000) and Stone (no date*) described bird commu- nities associated with four different Gambel Oak habi- tats (i.e., oak woodland, submontane shrub, Ponderosa Pine (Pinus ponderosa)/oak woodland, and mixed conifer/oak woodland), and locations. Garcia et al. (1998) and Corcuera and Butterfield (1999) referred to separate bird communities in oak woodlands, mixed oaks (Quercus rugosa, Q. candicans, Q. obtusata, Q. laurina) and other groupings for north-central Michoa- can, Mexico. Some comparisons and comments can be made about the composition of communities from the present study in relation to these others. Shepard (2000) out- lined the importance of House Wren (found in com- munity 2) for two Garry Oak sites west of Victoria, 384 British Columbia. However, those sites had steady num- bers of Red-breasted Nuthatch (Sitta canadensis) and Brown Creeper (Certhia americana), species which were unimportant in the present study. Most of the frequent birds from my study were present in Black Oak habitat in northern California, but no more than one or two species were listed by habitat suitability class (Dedon et al. 1984). Five of the seven species of community 3 were resident or breeding species in the ponderosa pine/Gambel Oak forest in north-central New Mexico (Leidolf et al 2000). The two remaining species are not found in the range of Gambel Oak. However, the shrub subdivision in Gambel Oak bird communities (Stone no date*) is not similar to the four communities except for the presence of Spotted Towhee. A community from Trembling Aspen forests in north-central British Columbia had similarities (4/7 species) to community 3 (Pojar 1995). Although this was labelled a group of conifer-related species, three of four species were associated with Gambel Oak and Trembling Aspen in southwestern Colorado (Stone no date*). In addition, Shepard (2000) mentioned high densities of Pacific-slope Flycatcher, also a species of community 3; and the importance of Chesnut-backed Chickadee (found in community 1), for two Garry Oak sites in British Columbia. These species have similarly been thought of as conifer (Douglas-fir)-related (Huff and Raley 1991). Neither the other two Trembling Aspen bird communities (Pojar 1995), nor the ones described for Douglas-fir habitat (Shepard 2000; Huff and Raley 1991) had any substantial similarity to the four communities in the present study. Summary Testing of wildlife habitat relationship models based on broad habitats (e.g., oak woodland, old-growth Douglas-fir forest) has given results which questioned their basic utility (e.g., Dedon et al. 1984; Laymon 1989). This suggests that other methods could be examined. An alternative approach, which empirical- ly defines spatial and temporal bird aggregations as bird communities, is presented here, and may war- rant further investigation. The four communities identified in this work could serve as benchmarks in Garry Oak ecosystem studies. Like any classification, they should be tested with fur- ther field work and analysis, particularly for differen- tiation by habitat features. For example, Erickson and Campbell (2001) related correlations involving mois- ture regime, oak diameters, tree form complexity, total wildlife habitat features, and, for one year, average number of bird species per plot. Monitoring is recom- mended as a first step in maintaining characteristic Garry Oak bird communities, a focus which should parallel growing concern for this endangered habitat and contribute to increasing conservation efforts. THE CANADIAN FIELD-NATURALIST Vol. 118 Acknowledgments The field work for this study was completed in an M.Sc. program at the University of Victoria, toward which British Columbia Ministry of Forests approved a part-time education leave. Both Canadian Wildlife Service and a Canadian Wildlife Service employee’s fund provided grants which assisted in the first year of the study. I thank A. J. Erskine for a helpful review. Documents Cited (marked * in text) Abernethy, V. J., D. I. McCracken, A. Adam, I. Downie, G. N. Foster, R. W. Furness, K. J. Murphy, I. Ribera, A. Waterhouse, and W. L. Wilson 2001. Functional analysis of plant-invertebrate-bird biodiversity on Scottish agricultural land. Website posting (Accessed 13 January 2005): http://www. gla.ac.uk/ibls/DEEB/kjm/divpap.html. Erickson, W. R. 1993. Garry oak ecosystems at risk (brochure). Wildlife Branch, British Columbia Ministry of Environ- ment, Lands and Parks, Victoria. Website posting (Accessed 13 January, 2005): http://wlapwww.gov.be.ca/wld/list.htm. McCune, B., and M. J. Mefford. 1997. PCORD: Multivariate analysis of ecological data, version 3.0. MjM Software Design. Gleneden Beach, Oregon. Stone, E. no date. Avian communities of Gambel oak habitats in southwestern Colorado. Website posting (Accessed 10 Februrary 2003): www.colorado.edu/epob/epob4630estone/ ES/ResearchStuff/Oak/Oak. html Literature Cited Anderson, S. H. 1970. The avifaunal composition of Oregeon white oak stands. Condor 72: 417-423. Artman, V. L., E. K. Sutherland, and J. F. Downhower. 2001. Prescribed burning to restore mixed-oak communities in southern Ohio: Effects on breeding-bird populations. Conservation Biology 15: 1423-1434. 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So a mighty oak can grow: A bird’s eye view of the Oak Woodlands Restora- tion project. Bird Conservation 14: 6-7. Website posting (Accessed 25 November 2004): http://www.partnersinflight. org/pubs/birdcons/14pgs6-7.pdf. Ehrlich, P. R., D. S. Dobkin, and D. Wheaye. 1988. The birder’s handbook: A field guide to the natural history of North American birds. Simon and Schuster, Toronto. 2004 Erickson, W. R. 1996. Classification and interpretation of Garry oak (Quercus garryana) plant communities and ecosystems in southwestern British Columbia. M.Sc. the- sis. University of Victoria, Victoria, British Columbia. Erickson, W. R. 2000a. Garry oak communities in Canada: Classification, characterization and conservation. Interna- tional Oaks 10: 40-54. Erickson, W. R. 2000b. Birds of the Garry oak habitat in British Columbia. British Columbia Birds 10: 5-12. Erickson, W. R., and D. C. Campbell. 2001. Aspects of the form, ecological function and aesthetics of Garry oak and its environment in southwestern British Columbia, Canada. International Oaks 12: 39-47. Garcia, S., D. M. Finch, and L. D. Chavez. 1998. Patterns of forest use and endemism in resident bird communities of north-central Michoacan, Mexico. Forest Ecology and Management 110: 151-171. Hagar, J. C., and M. A. Stern. 2001. Avifauna in oak wood- lands of the Willamette Valley, Oregon. Northwest Natu- ralist 82: 12-25. Haila Y., I. Hanski, and S. Ravio. 1993. Turnover of breeding birds in small forest fragments: The sampling colonization hypothesis corroborated. Ecology 74: 714-725. Hill, M. O., and H. G. Gauch. 1980. Detrended correspon- dance analysis: an improved ordination technique. Vege- tatio 42: 47-58. Huff, M. H., and C. M. Raley. 1991. Regional patterns of diurnal breeding bird communities in Oregon and Wash- ington. Pages 117-121 in Wildlife and vegetation of unman- aged Douglas-fir forests. Technically coordinated by L. F. Ruggiero, K. B. Aubry, A. B. Carey and M. H. Huff. United States Department of Agriculture Forest Service General Technical Report PNW-GTR-285. Portland, Oregon. Hurlbert, S. H. 1984. Pseudo-replication and the design of ecological field experiments. Ecological Monographs 54: 187-211. Laymon, S. A. 1989. A test of the California Wildlife-Habitat Relationship system for breeding birds in Valley-Foothill riparian habitat. Pages 307-313 in Proceedings of the Cali- fornia riparian systems conference. United States Depart- ment of Agriculture Forest Service General Technical Report PSW-119. Davis, California. ERICKSON: BIRD COMMUNITIES OF GARRY OAK HABITAT Leidolf, A., M. L. Wolfe, and R. L Pendleton. 2000. Bird communities of Gambel oak: A descriptive analysis. United States Department of Agriculture Forest Service General Technical Report RMRS-GTR-48. Fort Collins, Colorado. Manuwal, D. A. 1986. Characteristics of bird assemblages along linear riparian zones in western Montana. Murrelet 67: 10-18. Parody, J. M., F. J. Cuthbert, and E. H. Decker. 2001. The effect of 50 years of landscape change on species richness and community composition. Global Ecology and Biogeog- raphy 10: 305-313. Peet, R. K., R. G. Knox, J. S. Case, and R. B. Allen. 1988. Putting things in order: The advantage of Detrended Cor- respondance Analysis. American Naturalist 131: 924-934. Pojar, R. A. 1995. Breeding bird communities in aspen forests of the Sub-Boreal Spruce (dk Subzone) in the Prince Rupert Forest Region. British Columbia Ministry of Forests, Land Management Handbook 33. Victoria. Probst, J. R. 1980. Oak forest bird communities. Pages 80- 88 in Management of North Central and Northeastern for- ests for nongame birds. Workshop Proceedings, July 23- 25, Compiled by R. M., DeGraaf and K. E. Evans. United States Department of Agriculture Forest Service General Technical Report NC-51. Central Forest Experimental Station. Minneapolis Minnesota. Shepard, M. G. 2000. Breeding bird censuses in Garry oak forests, with notes on habitat use, metapopulations, and consequences of forest fragmentation. Pages 277-278 in Proceedings of a conference on the biology and manage- ment of species and habitats at risk, Feb. 15-19, Kamloops, British Columbia. Edited by L. M. Darling. British Colum- bia Ministry of Environment, Lands and Parks, Victoria. Smith, K. G. 1977. Distribution of summer birds along a forest moisture gradient in an Ozark watershed. Ecology 58: 810-819. Smith, W., A. Solow, and S. Chu. 2002. An index of the con- tribution of spatial community structure to the species- accumulation curve. Ecology 81: 3233-3236. Received 17 April 2003 Accepted 24 January 2005 Invertebrate Diversity under Artificial Cover in Relation to Boreal Forest Habitat Characteristics STEVEN H. FERGUSON! and DANIELLE K. A. BERUBE? 'Fisheries and Oceans Canada, Freshwater Institute, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Canada > Department of Biology, Lakehead University, 955 Oliver Road, Thunder Bay, Ontario P7B 5E1 Canada Ferguson, Steven H., and Danielle K. A. Berube. 2004. Invertebrate diversity under artificial cover in relation to boreal forest habitat characteristics. Canadian Field-Naturalist 118(3): 386-392 We investigated invertebrate diversity in boreal forests using an experimental design that consisted of counting soil inverte- brates under artificial cover. The aim was to assess the utility of using soil invertebrate diversity as a measure of ecosystem health. The study area was grouped into five habitats: upland hardwood, lowland hardwood, conifer, shrub, and conifer- grass. Simpson’s and Shannon’s indices of invertebrate diversity were negatively correlated with percent herbaceous cover. Number of recognizable taxonomic units (RTU richness) was negatively correlated with percent litter cover. The number of individual invertebrates was positively correlated with soil moisture and negatively correlated with percent conifer cover. Invertebrate diversity varied among habitat types, with conifer forests (spruce, fir, pine) having the highest diversity and regenerating conifer-grass forests having the lowest diversity, suggesting that successional stages affect diversity. The most productive sites, upland and lowland hardwood habitats, had the highest abundance of soil invertebrates, although interme- diate diversity compared to the other five habitats. The results are consistent with the view that diversity increases and then decreases with productivity and disturbance over succession (ca. 50-100 yr). Hence, maintenance of soil invertebrate diversity in managed boreal forests requires the provision of a varied landscape with a mosaic of disturbance regimes. Key Words: arthropods, biodiversity, conifer, earthworms, indices of diversity, moisture, Ontario. Concerns about the effects of the widespread loss of biodiversity have prompted many recent studies investigating the relationship between biodiversity and ecosystem function (Symstad et al. 2000). Inver- tebrates are important in the functioning of nearly all environments, with changes in species composition potentially reflecting changes in the ecosystem (Majer 1990; Madden and Fox 1997). Hence invertebrates are increasingly being viewed as reliable indicators to assess human impacts on the general level of distur- bance of an ecosystem (Majer 1983; Greenslade 1984; Andersen 1990). Biodiversity includes all levels of natural variation and thus diversity indices provide a relative measure of variation within a community (Til- man and Pacala 1993). Monitoring diversity across spatial and temporal scales allows for measurement of system complexity, functionality, and stability. Knowl- edge of diversity helps in understanding changes in ecosystem complexity before and after disturbance. Information on the habitat characteristics that influence diversity at various levels and knowledge of habitat changes resulting from human disturbance are required for management and conservation (Madden and Fox 19977): Forest managers can assess diversity changes asso- ciated with human disturbances that include various forestry practices through an understanding of the relationship between animal diversity and forest struc- ture to determine ecosystem changes (Noss 2000). Animal diversity includes soil invertebrates common- ly found under logs and rocks in managed forests (Kolstrom and Lumatjarvi 1999). Downed wood is im- portant for organisms in providing shelter and moisture, and in preventing light penetration. The use of soil fauna diversity has the potential to act as a surrogate of forest biodiversity. A number of forest characteristics have been shown to relate to soil invertebrate diversity, including understorey vegetation and litter (Bird et al. 2000), plant functional diversity (Siemann et al. 1998), coarse woody debris (Marra and Edmonds 1998), conifer species composition (Lattin 1993), forest suc- cession (Paquin and Corderre 1997), soil moisture (MacKay et al. 1986) and structural complexity (Fer- guson 2001). To further this research, we used a method of sur- veying invertebrate diversity under artificial cover and relate indices of invertebrate diversity to measured for- est characteristics. The study design consisted of (1) surveys of soil invertebrates (springtails, beetles, cen- tipedes, slugs, earthworms and isopods) found under sand-filled cardboard boxes placed on the forest floor; and (2) surveys of the sampled forest characteristics (snags, logs, soil moisture, overstorey, understorey, and ground cover). Forest characteristics were used to identify habitats within a boreal forest landscape, and diversity of soil invertebrates was measured. The goal was to relate forest habitat characteristics to soil invertebrate diversity to assess their utility as surro- gate measures of ecosystem changes associated with forest management practices. 386 2004. FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 327 RS 1.0 cb) O S is) —~ jp <2) © 0) 0.5 iS rs) =) LL Forest types Ficure 1. Location of study area in northwestern Ontario and dendrogram of 20 plots (0) located within a 1 km? stand of boreal forest grouped using cluster analysis based on forest characteristics (e.g., Overstorey. understorey, ground cover, moisture, coarse-woody debris). 388 Study Area The study area (Figure 1) was located in northwest- ern Ontario, Canada, and consisted of a 1 km? area of boreal forest located along the McIntyre River within Lakehead University’s natural forest (48°22'N, 89°19'W). The mixed boreal forest consisted of Jack Pine (Pinus banksiana Lamb.), Black Spruce (Picea mariana (Mill.) B.S.P.), Balsam Fir (Abies balsamea (L.) Mill.), White Spruce (P. glauca (Moench) Voss), White Birch (Betula papyrifera Marsh.), and Trem- bling Aspen (Populus tremuloides Michx.). The study site lies within the Boreal Ecosystem that consists of rolling rocky uplands with coarse well-drained soils (Rowe 1972). The climate is humid continental with a mean minimum January air temperature of -15°C and a mean maximum daily air temperature for July of 18°C (Environment Canada 2001"). Mean annual precipitation is approximately 700 mm, including a mean winter snowfall of 196 cm (Environment Cana- da 2001"). For the study period (May to September), mean monthly daily air temperature (1961-1990 nor- mals) varied from 9.0°C in May to 17.7°C in July (annual mean = 2.4°C) and precipitation varied from 69.3 mm in May to 88.5 mm in August (annual mean = 58.6 mm) (Environment Canada 2001"). Methods and Materials Study design and sample plots The sampling method was designed to assess soil invertebrate diversity under a standardized collection technique that optimized sampling replication, effort, and coarse taxonomic resolution. Coarse woody debris occurred in various shapes, sizes, and material. The use of cardboard boxes standardized the collection and reduced this variability while sampling a more diverse fauna than other methods such as pitfall traps. We decided to use a coarse taxonomic resolution (Bolger et al. 2000), thereby allowing inexperienced observers with minimal training to obtain reasonable survey counts efficiently. A trade-off associated with group- ing invertebrates occurs between ease of surveys by observers with minimal taxonomic experience and the loss of more detailed guild and life history infor- mation related to individual species. Twenty plots, each consisting of three adjacent boxes, were randomly distributed within the forest (minimum distance between plots was 5 m; Figure 1). Each box consisted of two 2-liter milk cartons with a plastic coating of red and white color. These were fitted one inside the other to create a solid box and filled with sand (approximately 2 kg) to create a footprint-sized depression 21 by 9.5 cm. Boxes depressed the leaf litter an average of 1.3 cm (Ferguson 2000), creating a microhabitat of increased humidity and decreased temperature similar to that beneath a log or rock rest- ing on the forest floor. Boxes were overturned and the numbers of all soil invertebrates (>1 mm) were visually counted. Boxes were lifted individually without disturbing adjacent THE CANADIAN FIELD-NATURALIST Vol. 118 boxes or the underlying litter. Twenty weekly surveys were conducted from 9 May to 24 September 2000 between the hours of 1100 to 1700. The survey num- bers reflect a relative abundance of soil invertebrates. Observer bias was consistent across the landscape. Recognizable Taxonomic Units (RTU; Bolger et al. 2000; Ferguson 2004) were used to group all macroin- vertebrates (>3 mm) observed under boxes based on differences in size and feeding habits (Eisenbeis and Wichard 1987; Brock et al., 1994) and included spring- tails (Collembola), spiders (Araneae), ants (Formicidae), ant larvae, centipedes (Chilopoda), Diplura, adult flies (Diptera), phytophagous mites (Acarina: Oribatida), bugs (Hemiptera and Homoptera), Pseudoscorpionida, wasps (non-ant Hymenoptera), moth and butterfly larvae (Lepidoptera), Gastropoda (snails and slugs), Isopoda (woodlouse; Tracheoniscus rathkei), beetles (Coleoptera and Staphylinidae species — adults and larvae), milli- pedes (Diplopoda), and earthworms (Oligochaeta). Forest characteristics measured Site-specific habitat variables were measured 21- 23 August 2000 using 5 x 5 m quadrats centered at three box plots. One quadrat was located at the centre of each plot. Four other quadrats were located 10 m from the centre in cardinal compass directions. A total of 100 quadrats were sampled for the following habitat characteristics: percent overstorey cover (>5 m), per- cent understorey cover (saplings and shrubs 0.5-5 m), percent herbaceous cover, percent litter cover, percent grass cover, percent moss cover, percent fern cover, number of snags (dead standing trees with dbh > 5 cm), number of decaying logs (> 5 cm diameter), percent cover by conifers, and a relative measure of soil mois- ture (1.e., xeric=1, mesic=2, hydric=3). Values for the five quadrats were averaged for each plot. Measures of invertebrate diversity Diversity of soil invertebrates was calculated for each plot across surveys using RTU richness (num- ber of RTU at each site), Shannon-Wiener, and Simp- son’s Indices of Diversity (Ludwig and Reynolds 1988). Shannon’s entropy (H) is a measure of species diversity used in relation to relative frequencies (probabilities) of the different species i of the sample and was calculated as: H=-% [(n/n) In(n/n)] , from i=1 ton (1) where n, is the number of individuals belonging to the ith RTU in the sample and 7 is the total number of individuals in the sample. H = 0 (minimum value) when the sample contains only a single RTU, where- as diversity H increases with the number of RTUs. H is maximum when all RTUs are equally distributed in the sample. The Shannon-Wiener index of diversity is sensitive to changes in the rare species in a com- munity sample (Pielou 1966). Invertebrate diversity was also measured by Simpson’s index of diversity (Simpson 1949), which is sensitive to changes in the more abundant species and was measured as: A= 1- Yn (n, - 1)/n(n-1) (2) 2004 Simpson’s index varies from 0 to 1, and gives the probability that two individuals drawn at random from a population belong to the same RTU. If the number approaches () then individuals belong to the same RTU and the diversity of the community sample is low. These diversity measures were calculated using the minimum number of invertebrates observed for the RTU in a given sampling period and were represent- ed by an average value for each survey. Statistical analysis We performed a cluster analysis of the 11 forest measures to group the 20 sampling sites into forest habitats. Classification of habitats types was conducted by the average-linkage clustering method using a Euclid- ean distance similarity index (Romesburg 1984). All variables were standardized between 0 and 1. Many (8 of 17) forest and soil invertebrate variables were not normally distributed (Wilk’s statistic) and transformations (e.g., log, arc-sine) failed to normalize all variables. Therefore, we used nonparametric analy- ses by ranking nonparametric data before correlation analyses (Conover and Iman 1981). Measures of diver- sity (Simpson’s and Shannon’s), RTU richness, and number of individuals were normally distributed and did not require transformations. We report untrans- formed means in the Figures and Tables in the Results section to simplify presentations. Analyses were performed to determine the relation- ship between (1) soil invertebrate diversity indices and forest characteristics, and (2) soil invertebrate numbers for each of the abundant RTU (springtails, beetles, centipedes, slugs, earthworms, and isopods) and for- est characteristics. We tested for significant effects of forest characteristics on dependent measures (e.g., diversity) with Spearman’s correlations and partial correlation analyses (i.e., multiple regression of ranked data without replacement). ANOVA was used to com- pare forest characteristics, indices of diversity, and soil invertebrate abundance relative to forest types, fol- FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 389 lowed by a Tukey multiple range test if the ANOVA was significant. Spearman correlations for nonpara metric data and Pearson’s correlations for parametric data compared RTU abundance with forest character- istics. Sample units were the 3-box groups (n = 20) sampled every week (n = 20). All statistical analyses were done using SAS (SAS Institute Inc., Cary, North Carolina 1987) statistical software for microcomput- ers: Results Both indices of soil invertebrate diversity (Shan- non’s and Simpson’s) were negatively related to per- cent herbaceous cover (R? = 0.32, F = 5.89, P = 0.01, n= 20 and R? = 0.35, F = 6.64, P = 0.01 respectively; Table 1, Figures 2A, and 2B). RTU richness (number of RTU) was negatively correlated with percent litter cover, which explained 48.6% of the variance in the model (F' = 17.0, P= 0.001, n = 20; Table 1, Figure 2C). Diversity indices were also related to litter cover as herbaceous cover and litter cover covaried (r = -0.94). Soil moisture (69.5% of variation explained) and per- cent conifer cover (9%; Figure 2D) best explained the number of individual invertebrates. Greater numbers of soil invertebrates occurred in wet habitats and with less conifer cover (Table 1). The forest measures were grouped into five forest types defined as upland hardwood, lowland hard- wood (mesic), shrub, conifer, and conifer-grass (Fig- ure 1). The upland hardwood forest consisted prima- rily of Trembling Aspen, with little understorey (47% cover) and the most abundant herbaceous cover (90%; Table 2). The soil invertebrate community in the upland hardwood forest consisted of numerous individuals (mean = 24/plot), moderate RTU diversity, and large numbers of isopods (Table 2). The lowland hardwood habitat consisted of mixed hardwoods in a mesic site that included an intermittent stream with a deep humus layer. The soil invertebrate community TABLE 1. Seven multiple regression results used to determine the significant effects of 11 forest characteristics on (1) Shannon’s index of diversity; (2) Simpson’s index of diversity; (3) RTU richness (where RTU = recognizable taxonomic units); (4) Number of individual invertebrates, (5) springtail abundance; (6) earthworm abundance; (7) isopod abundance. Of the 11 explanatory variables included (soil moisture, % overstorey, % understorey; % herb cover; % litter; % grass: % moss; % ferns: number of snags; number of logs; and % conifer cover), only those with significant (P < 0.05) relationships are shown. Dependent Independent Coefficient Partials Model variable variable direction R? R? P (1) Shannon’s diversity index % herb cover negative 0.318 0.318 0.008 (2) Simpson’s diversity index % herb cover negative 0.349 0.349 0.006 (3) RTU richness % litter cover negative 0.486 0.486 0.001 (4) Number of RTU Soil moisture positive 0.695 0.695 0.0001 % conifer cover negative 0.087 0.782 0.02 (5) Springtail abundance % conifer cover negative 0.224 0.224 0.04 (6) Earthworm abundance Conifer cover negative 0.657 0.657 0.0001 Number of logs positive 0.100 0.757 0.02 (7) Isopod abundance % conifer cover negative 0.624 0.624 0.0001 390 Shannon's index 0 25 50 75 Percent herb cover Species richness 100 0 25 50 75 Percent litter cover THE CANADIAN FIELD-NATURALIST Vol. 118 © Ce) Simpson's index Percent herb cover 800 600 400 Number of individuals 0 25 50 75 Percent conifer cover FIGURE 2. Bivariate relationships between two indices of diversity (Simpson’s and Shannon’s) of soil invertebrates and major explanatory habitat variables: (A) Shannon’s index of diversity, and percent herbaceous cover (y = -0.307x + 2.29, 7 = 0.318); (B) Simpson’s index of diversity and percent herbaceous cover (y = -0.0810x + 0.883, 7 = 0.349); (C) RTU richness (number of Recognizable Taxonomic Units) and percent litter cover (y = -0.0692x + 18.6, 7° = 0.486): and (D) number of individual invertebrates and percent conifer cover (y = -9.38x + 712, r? = 0.695). living in this forest consisted of moderate diversity, the greatest number of individuals (mean = 37/plot), and the most springtails and earthworms. The shrub habitat had the highest understorey cover (78%), low- est overstorey (34%), and moderate conifer and litter cover. The soil community in the shrub habitat was moderately diverse and consisted of a moderate num- ber of individuals (mean = 16/plot). The conifer for- est was the most extensive habitat and was composed of a mixed softwood/ hardwood (56/44%) forest with high overstorey cover (66%), high understorey cover (66%), low herb cover (33%), and high litter cover (64%). The conifer forest supported the highest soil invertebrate diversity although few RTU (low RTU richness) and few individuals (mean = 10/plot; Table 2). The conifer-grass forest type was represented by a relatively low overstorey (55%), intermediate under- storey (56%), high conifer cover (53%) with the most grass (13%) and more herb than litter cover (66/22%). The soil invertebrate community in the conifer-grass habitat was the least diverse with the lowest RTU richness and fewest individuals per plot (n = 9). The abundance of three out of the six most numerous RTU were significantly related to forest characteristics 2004 FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 39 | TABLE 2. Comparison of forest and soil invertebrate characteristics for five forest types. Mean invertebrate abundances (numbers per m7) are given for six of the more common invertebrates. Means with the same letters do not differ significantly according to Tukey’s multiple range test. Upland hardwood Variable F IP (n= 6) Forest characteristics Snags 6.0 0.005 ab 13.3 Logs 10.5 0.0003 ab 7.0 Soil moisture 26.7 0.0001 b 0.6 Overstorey Sel 0.008 ab 52.8 Understorey 4.4 0.01 b 46.5 Herb 28.3 0.0001 a 89.5 Litter 26.0 0.0001 c 2.8 Grass 2967]: 0.0001 10), 117/ Conifer 19.0 0.0001 b 21.8 Indices of diversity Shannon’s 5.64 0.006 a2.15 Simpson’s 37 0.002 ab 0.191 RTU richness S/3 0.005 a 19.3 Number of individuals 9.11 0.0006 ab 475 Soil invertebrates Springtails 3.64 0.03 ab 67.8 Beetles 0.6 0.70 16.5 Centipedes 2 0.34 US) Slugs 0.8 0.52 4.0 Earthworms 8.8 0.002 a 54.5 Isopods ae) 0.005 a 127.3 (Table 1). Springtail abundance increased with greater litter cover (22.4% of variation explained; F = 5.2, P =0.04). Earthworm abundance was negatively cor- related with percent conifer cover (65.7% explained variance) and positively correlated with number of logs (10%; F = 34.4, P < 0.001). Isopod abundance was negatively related to percent conifer cover (62.4% of variation explained; F = 29.9, P < 0.001; Table 1). In contrast, beetles, centipedes, and slugs were relatively evenly distributed across forest types and their abun- dance was not significantly related to forest charac- teristics (not shown in Table 1). Discussion Our findings are consistent with the view that diver- sity is greater in the most common habitat within a varied landscape, such as a mixed-wood boreal forest (<80 y) and is often characterised by intermediate productivity and disturbance. Simpson’s and Shannon’s Indices of Diversity explained similar amounts of vari- ation in the pattern of diversity of soil invertebrates with forest characteristics. Both found percent herba- ceous cover the most important environmental factor explaining the pattern of soil invertebrate diversity among forest habitats. Also, both indices showed simi- lar correlations, in magnitude and direction, such that Forest types Lowland Shrub Conifer Conifer- hardwood grass (n = 2) (n=4) (n=6) (n= 2) a 16.0 b 4.3 ab 7.5 b 5.5 a 15.0 o23 be 2.7 c 2.0 a 1.4 c 0.0 c 0.1 c 0.0 a 69.5 b 34.3 a 65.7 ab 54.5 ab 54.5 a 77.8 ab 66.3 ab 56.0 ab 68.5 be 56.8 c 32.8 b 65.5 cb 14.0 b 31.8 a 64.2 cb 21.5 b 0.5 b 3.5 bee, a 13.0 b 26.0 ab 36.0 a 55.8 a 53.0 ab 1.99 ads Lif; a 2.19 b 1.90 b 0.155 b 0.147 b 0.142 a 0.228 ab 17.0 ab 16.3 ab 14.5 b 14.0 a 723 be 313 c 190 c 179 a 120.5 ab 65.0 b 46.2 ab 76.5 10.0 eS, 16.5 4.5 10.5 12.0 6.5 20.0 6.0 3:3 6.3 6.0 a 116.5 ab 32.3 b 21.0 b 14.0 ab 96.5 ab 64.5 b 25.3 b 14.5 as diversity increased the percentage of herbaceous cover decreased. In contrast to our findings, soil inver- tebrates from 27 orders did not differ significantly among five different landscapes (with the exception of earthworms) (Kalisz and Powell 2000). We found a negative relation of soil invertebrate diversity with increasing composition of conifer veg- etation. Few herbs and more litter are present under the conifer cover due to greater soil acidity (Kimmins 1997). Numbers of invertebrates were greater under herbaceous cover but diversity was reduced. Other studies have found invertebrates occurring in greater numbers under deciduous and herbaceous cover rela- tive to conifer litter (Wallwork 1983; Paquin and Coderre 1997; Hammond 1997; Marra and Edmonds 1998). Within varied forest landscapes, the greatest diversity of soil invertebrates likely occurs for the most common (in time and space) successional community. In northern boreal forests this is likely for older conifer stands with greater overstorey and reduced ground veg- etation understorey. Removal of conifers by natural disturbance or forest harvesting likely results in lower initial diversity with increasing diversity over time as other plants colonize the habitat (i.e., succession). We found greater invertebrate density under herbaceous cover associated with hardwood forest, but diversity 392 was lower relative to the conifer forest. More research is required to explain these differences. There are some limitations to the sampling method- ology used in this study. Taxonomic levels lower than the one used here to identify invertebrates would cer- tainly provide more detailed results though sampling efficiency would be reduced. Sampling was done around mid-day, which limits sampling to invertebrate groups that are active during the day (Eisenbeis and Wichard 1987). Results are limited to the invertebrates visually observed (i.e., >1 mm which may exclude arthropods such as small mites) found under boxes whereas smaller individuals and groups found in deep- er soil are under-represented. Another potential con- cern is that ants, due to their clumped distribution and significant effects on community structure, may obscure patterns among other invertebrate groups (Madden and Fox 1997). Diversity indices respond to both changes in species richness estimates and changes in species evenness. For example, the mesic lowland hardwood site record- ed few RTU but high total density. Among those com- mon taxa associated with the mesic lowland sites are springtails, earthworms, and isopods, which might be expected to be more abundant in moist soils. As a result, one problem with interpretation of diversity results is the possible relationship between measures of richness and diversity relative to measures of density. Both Shannon’s and Simpson’s indices are expected to be lowest when richness and evenness are low and to increase with richness and evenness. Shannon’s index is more sensitive to richness and is impacted by the inclusion of rare taxa, while Simpson’s index is more sensitive to evenness of the more common taxa. The result is usually that the two are generally correlated, although the relative ranking of sites may differ some- what. Among the five forest types reported in Table 2, the two hardwood sites had the highest numbers of individuals. Both Shannon’s and Simpson’s indices are reported as moderate for these sites with higher esti- mates for both in the upland sites. The two conifer- grass sites yielded the lowest number of individuals. The Shannon’s estimate is the lowest of the five sites while the Simpson’s is the highest. Between the other two sites, which recorded somewhat higher numbers of individuals, the Shannon’s estimates are the two highest while the Simpson’s estimates are the two low- est. The net effect is that across all five habitat types the correlation coefficient between the Shannon and Simpson estimates for the forest stands arranged along a moisture gradient had a negative slope, although not significant, contrary to expectations. However, both indices were negatively correlated with % herb cover and positively correlated with each other. Greater repli- cates are required in various forest stands along envi- ronmental gradients to account for forest characteris- tics that covary. Although increasing plant diversity significantly increases invertebrate diversity, local herbivore diversi- THE CANADIAN FIELD-NATURALIST Vol. 118 ty is also maintained by a diversity of parasites (e.g., flies and nematodes) and predators (Siemann et al. 1998; Ferguson 2001). A community-level prediction is that invertebrate diversity increases with increasing plant species diversity, as many invertebrates forage on the leaves and litter of herbaceous plants (Symstad et al. 2000). However, if an entire functional group of plants, such as conifers, is absent from a habitat, then a landscape-level decrease in soil invertebrate diver- sity would occur, as many taxa are associated with the acidic soil and fungal hyphae characteristic of conif- erous forest floors (Lattin 1993; Butterfield 1999). Alternatively, diversity is hypothesized to increase and decrease across the two dimensions of productivity and disturbance, respectively (Kondoh 2001). Our results conform to this view, as conifer cover had the highest diversity at the late successional stage but the lowest measure of diversity at the early conifer-grass stage. In contrast, more constant mesic hardwood habitats had high productivity and low disturbance, and showed intermediate indices of invertebrate diversity. Appar- ently, the pattern of invertebrate community diversity varied among habitat types according to productivity and disturbance gradients. The decrease in soil invertebrate diversity with increasing grass cover may be due to the low nutri- tive value of grass and the microhabitat conditions of grassy areas. Low moisture, associated with dry grassy areas, has been found to have an adverse effect on soil invertebrate abundance due to a reduced oxygen level that degrades soil composition, increases erosion, and mobilizes carbohydrates and nutrients (Marra and Edmonds 1998). The majority of soil invertebrates are best suited to moderate moisture levels, as some spe- cies have little or no exoskeleton (e.g., earthworms) to protect against high and low soil moisture (Schaefer 1995; Ferguson 2004). Soil invertebrate diversity was not significantly related to many of the forest characteristics. Still, the considerable diversity of invertebrates ensures that some species are adapted to many of the diverse con- ditions. For example, soil invertebrates did not show a relationship to the number of downed logs. This lack of a relationship with logs may have been relat- ed to the experimental technique of providing a simi- lar microhabitat using boxes. Many soil invertebrates are known to depend on cover provided by coarse woody debris, such as downed logs, due to their pro- vision of nutrients and protection from predators and stability of microclimatic conditions (Lattin 1993; Ferguson and Joly 2002). Slugs showed few relation- ships with forest characteristics and were found in a diversity of habitats. Slugs forage on larvae of bee- tles and flies and on cellulose and other plant poly- saccharides (Port and Port 1986). Earthworms bur- row in moist rich soil and feed on decaying organic matter from fallen leaves and vegetation (Edwards and Bohlen 1996). Earthworms were found to be positively associated with herbaceous cover, nega- 2004 tively associated with conifer cover, and positively associated with soil moisture. The major food items of centipedes are earthworms and small arthropods (Formanowicz and Bradley 1987). Although cen- tipedes have been found to prefer moist habitats (Corey and Stout 1992), such as under logs, no sig- nificant relationships were found with the forest characteristics measured in this study. Springtails are abundant in soil, and consume decomposing plant material and fungal hyphae (Hop- kin 1997). Their abundance was negatively associat- ed with litter cover, perhaps because it is living annu- al plants that provide the overwinter dead material that fungal hyphae provide as springtail food. The non-significant correlation (P = 0.06) of springtail abundance with herbaceous cover and the negative association with conifer composition contradicts other studies (Butterfield 1999; Paquin and Coderre 1997). A possible explanation for these differences is the differing sampling methodologies used whereby surface-dwelling springtails were sampled here, in contrast to sub-surface sampling from other studies. Beetles showed little dependence on any of the measured forest characteristics in this study, in con- trast to another study that found habitat dependencies (Fournier and Loreau 2001). Beetles are adapted to a wide range of environments partly due to their exoskeleton enabling life in a variety of moisture conditions (Eisenbeis and Wichard 1987). Isopods occur in greater abundance under stones and in damp environments (Sutton 1980), which is confirmed in this study by the positive association with relative soil moisture. Also, isopods were positively associat- ed with herbaceous versus conifer cover, which is related to intolerance to acidic conditions of conifer soils and the greater food availability in deciduous and herbaceous forest cover (David et al. 2001). These findings differ with previous research that found isopod abundance unrelated to forest attributes (Bolger et al. 2000). Forest management, with the goal of preserving forest biodiversity that includes soil invertebrates, needs to consider the requirements of individual taxa in boreal ecosystems by providing varied landscapes within a mosaic of forest stands. For example, cli- mate change may profoundly influence boreal forest ecosystems and their management, via increased temperature and altered precipitation regimes (e.g., fires, etc.; Parker et al. 2000). In the boreal forest area studied here, conifer habitat had the highest diversity, although abundance and RTU richness were lower. However, our results need to be guarded- ly interpreted as they are based upon observations of a partly artificial system and our invertebrate diversi- ty results are not sufficiently comprehensive to draw management conclusions. Still, monitoring soil invertebrate diversity can provide a means to assess changes in forest environments with climate warm- FERGUSON and BERUBE: INVERTEBRATE DIVERSITY 393 ing as well as forest management practices that include silvicultural interventions to maintain forest health. Acknowledgments Students in the “Fish and Wildlife Practice” 4" year course offered at Lakehead University participated in data analysis. Financial support was provided by post- doctoral (SHF) and postgraduate (DKAB) funding from NSERC (Natural Sciences and Engineering Research Council) and Bowater Pulp and Paper Inc., Thunder Bay Woodlands Division. Documents Cited (marked * in text) Environment Canada. 2001. 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Observations of habitat use by Polar Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, and northern Bering Seas. Canadian Field-Naturalist 118(3): 395-399. A total of 1 112 Polar Bears (Ursus maritimus) at 482 sightings were recorded during aerial surveys in the Beaufort, Chukchi, and northern Bering Seas conducted primarily during September and October from 1979-1999. Of these bears, 784 were observed offshore at 400 sightings. The surveys were conducted by the Naval Ocean Systems Center and Minerals Manage- ment Service; they were designed to monitor the fall Bowhead Whale (Balaena mysticetus) migration. Over the 20-year period, 1 096 620 kilometers of surveys were flown. The majority of the offshore Polar Bears, 595 bears at 290 sightings, and most of the kill sites and polar bear tracks were recorded in 80-100% ice cover. The number of bears per kilometer increased substantially in >24% ice cover, with the highest number observed in 80-100% ice cover. This habitat use proba- bly is related to the availability of seals, their primary prey. There were 328 bears (83 sightings) recorded on land, and most of them were associated with whale carcasses and bowhead whale subsistence harvest sites along the Alaskan Beaufort Sea coast. Key Words: Polar Bear, Ursus maritimus, habitat, Beaufort Sea, Chukchi Sea, Northern Bering Sea, Alaska. The preference of Polar Bears for extensive ice habitats is well documented (Stirling et al. 1993; Stir- ling and Derocher 1993; Amstrup 1995). Polar Bears need an ice platform to hunt successfully for seals, their primary prey (Stirling, Andriashek, and Calvert 1993). The bears select ice habitats that increase ac- cessibility (availability) of seals. Ferguson et al. (2000) compared ice-habitat use by Polar Bears of the Cana- dian Arctic Archipelago and Baffin Bay. They found ice-habitat use to be similar for the two regions. How- ever, Baffin Bay bears had more limited access to Ringed Seals (Phoca hispida) because of the complete absence of ice cover during the summer season. The Canadian Arctic Archipelago had more consistent sea- sonal ice cover, which provided the bears with reli- able access to the seals. Although the Baffin Bay region had a much higher density of Ringed Seals (1.4-2.1 seals/square kilometer: Finley et al. 1983; Stirling and Oritsland 1995) than the Arctic Archipelago region (0.28-0.97seals/square kilometer: Kingsley, Stirling, and Calvert 1985), both regions had very similar Polar Bear densities. Ferguson et al. (2000) concluded that Polar Bear abundance is not always proportional to prey density because of differences in Polar Bear access to seals (i.e., availability of prey). During September and October (open-water season), the Alaskan Beaufort Sea has great variability in ice coverage from year to year (Figures la and 1b). Like Baffin Bay, the Arctic coast of Alaska during light ice years is free of ice during the open-water season, when the pack ice is more than 100 miles from the coast (see Figure la). During the absence of ice in coastal Beau- fort Sea habitats, bears that occur along the coast either scavenge on animal carcasses that wash ashore, feed on the remains of whale carcasses at subsistence whale harvest sites, or fast until shorefast ice forms and seals become available (United States Department of the Interior, Fish and Wildlife Service 1995). During heavy ice years, the pack ice remains on the continental shelf; ice floes are common in coastal waters, and shorefast ice remains or forms early dur- ing the fall (Figure 1b). These varying ice conditions affected the movements and distribution of Polar Bears in the Beaufort and Chukchi seas (Amstrup 2000). Methods and Materials From 1979-1999, the Naval Ocean Systems Center under contract with the Bureau of Land Management (1979-1982) and Minerals Management Service (1982- 1999) under the Bowhead Whale Aerial Survey Project has conducted aerial surveys in the Beaufort, Chukchi, and Bering seas (Ljungblad et al. 1986; Moore and Clarke 1992; Treacy 2000) to monitor the fall migration (late August through October) of endangered whales (Treacy 2000). Information on Polar Bears and sea-ice coverage was acquired mostly from these aerial surveys. Surveys were flown primarily at 1500 feet (458 meters), weather permitting, or at a minimum of 1000 feet (305 meters) to avoid disturbing marine mammals. The aircraft used was a de Havilland Twin Otter Series 300 with bubble windows on both sides of the plane. Earlier flights (1979-1991) over the northern Bering, Chukchi, and western Beaufort seas used a Grumman Turbo Goose model G21G (Ljungblad et al. 1986; Moore and Clarke 1992). Information also was used from aerial whale sur- veys conducted during the spring and summer (April through August) in the northern Bering Sea as well 395 396 Arctic North Slope of Alaska THE CANADIAN FIELD-NATURALIST Maximum Retreat of the Pack Ice on 2 October 1998 Vol. 118 “600 Miles Ficure la. Light Ice Year Information derived from the National Naval Ice Center (http://www.natice.noaa.gov/westarct1.htm). Maximum Retreat of the Pack Ice on 18 September 1983 Beaufort Sea Arctic North Slope of Alaska Maximum Retreat of the Pack Ice on September 18, 1983 ak. Ficure 1b. Heavy Ice Year information derived from the National Naval Ice Center (http://www.natice.noaa.gov/westarct1.htm). as in the Chukchi and Beaufort seas (Ljungblad et al. 1986; Moore and Clarke 1992). Incidental sightings of Polar Bears and their behavior were recorded along north-south transects and off transects along the re- mainder of the flight tracks (Figure 2). Information was recorded on percent ice coverage within about 1- 2 kilometers of the aircraft at each position update (once about every 5 minutes) and at each sighting loca- | 1 2004 ioe. Chukchi Sea Alaska Bering Sea & HANSEN: HABITAT USE BY POLAR BEARS 397 Polar Bear Sightings Survey Track FiGure 2. Study are with survey tracks and Polar Bear sightings. tion. Position updates, sighting locations, and ice- coverage data were recorded on an onboard comput- er system connected to a Global Positioning System receiver in the aircraft. These records included infor- mation on numbers of Polar Bears per sighting and ice cover within about 1-2 kilometers of the aircraft. Polar Bear tracks and kill sites also were recorded dur- ing September and October from 1987 through 1999. Survey effort in the Beaufort Sea focused primarily along the coast, continental shelf, and shelf break, while surveys in the northern Bering and eastern Chukchi seas extended from the coast to the International Date Line (Figure 2). The Polar Bear sightings, kill sites, locations of bear tracks, and ice-coverage data were converted into database files and analyzed with Arcview Geo- graphic Information System Program 3.2a (devel- oped by ESRI in Redland, Calif.). Tables and graphs were created in Microsoft Word and Microsoft Excel (Windows 2000) to compare bear numbers and sight- ings with ice coverage (Table | and Figure 3) and kill sites and bear tracks with ice cover (Figures 4 and 5). The number of bear sightings and the survey effort (kilometers flown) were plotted by ice category to com- pare ice-cover use (Figure 2). This analysis is provid- ing useful information on habitat use by Polar Bears occurring in Alaskan waters. Results and Discussion Table 1 shows the number of Polar Bears and num- ber of sightings recorded offshore by ice cover and on the mainland or on barrier islands. Figure 4 shows the number of Polar Bear kill sites and ice coverage recorded at the kill locations. Figure 5 shows the num- bers of Polar Bear tracks recorded and ice coverage at their locations. Over the 20-year period and the 1 096 620 kilome- ters of surveys flown, 1112 Polar Bears at 482 sightings were recorded in the Beaufort, Chukchi, and Bering 398 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE |. Number of Polar Bears and Number of Sightings (Recorded Per Year) by Ice Coverage and Number Recorded on Land. Year 80-100% Ice 50-79% 25-49% Ice Ice Ice 1979 15 (8) 1980 29 (2) ayy) iy, <6) 1981 S005) Sii(Q) Dale) 1982 16.3) 4 (4) ives) 1983 165 365) 14 (11) Brn) 1984 23)a- 1 CLS) (2) Sy) 1985 5 (3) ihe (Cb) 1986 9 (6) 3y2@) 1987 4 (2) i eo Gh) 1988 69 (43) 1G) PEO) 1989 155) 0) a= 9A) One®) 1990 52 (40) Perth) 1991 DD ran 3) Aa (2) Gd) 1992 LOS Gib) Sie G2) 1993 Sh aC) 1994 26°, (12) PO) 32) 1995 Disi(eld) 1996 9 (7) ta) 1997 53 (8) 1) ED) 1998 15 (6) 9 (4) 1b Gb) 1999 2 (2) 1b) Totals: 5957 1(290) 91 (52) 45 (23) Polar Bear Sightings/Kilometer @ bear/km 79-50% 49-25% — 24-1% 0% Ranges of Ice Coverage 100-80% Ficure 3. Polar Bear sightings per kilometer recorded from April 1979 through October 1999 in the Beaufort, Chukchi and Bering Seas. Ice Coverages at Kill Sites 1 { 77) o = 2) S - ) _ o Q = 3 rs 100% 99-95% 94-90% 89-80% 79-70% 69-50% 49-40% Percent Ice Coverage 39-1% FiGuRE 4. Number of kill sites and ice coveage recorded at each site. 1-24% Ice No Ice Total Number of On Land Bears and Sightings 51) 20 (10) (29) (20) 1 (1) 22 (19) 93 (49) 3-7(2) et) S50 1@2) 6 (4) (10) 356) 72 (46) (25) 53 (41) (18) (48) Sp (@) (16) 2 @) 10 (8) LL @) 65 (11) 217 (57) 247 (73) 33 (19) 39 (25) 328 (83) 1112 (482) 67 (5) 8 (5) S13) 28 17) 25, (7) seas from 1979-1999 (Table 1). Of these bears, 784 were observed offshore at 400 sightings. Figure 2 shows the number of Polar Bears per kilometer of survey effort over ranges of ice coverage. The num- ber of bears per kilometer increased substantially in >24% ice cover, with the highest number observed in 80-100% ice cover. These observations suggest that Polar Bears may use 80-100% ice cover more often than lower percentages of ice cover. This habitat use probably is related to the availability of seals, their pri- mary prey. Table | lists the number of bears and sight- ings seen offshore in other ice concentrations. Thirty- five kill sites and 769 Polar Bear track sightings were recorded, mostly in 80-100% ice cover (Figures 3 and 4, respectively). Tracks with zero ice cover were record- ed on land or barrier islands. A total of 328 bears (at 83 sightings) were recorded on the mainland or on barrier islands (Table 1). Most of these bears were associated with whale carcasses and bowhead whale subsistence harvest sites along the Beaufort Sea coast. The large numbers of bears were recorded on land in 1992 (67 bears) and 1998 (217 bears), when repeat sightings were made of sever- al bears feeding on whale carcasses along the Beau- fort Sea coast (Table 1). Studies of radio- and satellite-tagged Polar Bears have provided useful information on the movements and habitat use of individual Polar Bears (Arthur et al. 1996; Ferguson et al. 1998; Amstrup et al. 2001; Durner et al. 2004). These studies recorded Polar Bear locations and compared them with satellite-derived, 2004 Number of Polar Bear Tracks nm a ao nm o n x rs) £ (= ° - o a = Ss z 100 99-95 94-90 89-80 79-70 69-60 59-50 49-40 39-30 29-20 19-1% 0 Percent Ice Coverage FiGurE 5. Number of Polar Bear tracks and ice coverage recorded at each site. digitized maps of ice types and ice concentrations in _ the general areas where the bears were located. These remote-sensing techniques provide general informa- tion on habitat use by Polar Bears throughout the year. Aerial surveys in this study recorded ice types and ice concentrations within 1-2 kilometers of the aircraft along the flight track and at the specific loca- tions where Polar Bears were observed. Both tech- niques record information on habitat use. Aerial sur- veys provide real-time information on habitat use at the actual locations where the bears were observed, _ while remote-sensing techniques provide valuable information on bear movements and general informa- tion on habitat use over time. Many aerial surveys conducted specifically for Polar Bears are flown at lower altitudes than the 1500 foot surveys conducted in this study. Aerial surveys that are conducted at very low altitudes (200-300 feet) are very likely to disturb Polar Bears and other wild- life. Our surveys were conducted at 1500 feet to avoid disturbing marine mammals. Even at 1500 feet some _ of the bears observed reacted to the aircraft by run- ning away when the aircraft past overhead. Polar Bears are quite visible on the ice at 1500 feet because their tracks are very detectable. Other marine mammals of comparable size to Polar Bears, such as Beluga Whales, are easily detected at 1500 feet. _ Acknowledgments This study was funded by the U.S. Department of the Interior, Minerals Management Service, as part of its Alaska Environmental Studies Program. As the Principal Investigator, I would like to thank Stephen Treacy, Project Manager for the Minerals Manage- ment Service Bowhead Whale Aerial Survey Project, for providing the survey data on Polar Bears and ice coverage along the survey tracks from 1979 through 2000, and for his comments on the draft manuscript. I also would like to thank Elinore M. Anker, who edited the manuscript. Literature Cited Amstrup, S. C. 1995. Movements, distribution, and population dynamics of Polar Bears in the Beaufort Sea. Ph.D. thesis. HANSEN: HABITAT USE BY POLAR BEARS 399 University of Alaska, Fairbanks, Fairbanks, Alaska. 299 pages. Amstrup, S. C. 2000. Polar Bear. Pages 133-157 in Chapter 7 in The natural history of an Arctic oil field development and the biota. Edited by J.C. Truett and S. R. Johnson. Academic Press, San Diego, California. Amstrup, S. C., G. M. Durner, T. L. McDonald, D. M. Mulcahy, and G. W. Garner. 2001. Comparing movement patterns of satellite-tagged male and female Polar Bears. Canadian Journal of Zoology 79: 2147-2158. Arthur, S. M., B. F. J. Manly, L. L. McDonald, and G. W. Garner. 1996. Assessing habitat selection when avail- ability changes. Ecology 77: 215-227. Durner, G. M., S. C. Amstrup, R. Neilson, and T. McDon- ald. 2004. The use of sea ice habitat by female Polar Bears in the Beaufort Sea. OCS Study MMS 2004-014. USDOL, MMS, Alaska OCS Region, Anchorage, Alaska. 41 pages. Ferguson, S. H., M. K. Taylor, and F. Messier. 2000. Influ- ence of sea ice dynamics on habitat selection by Polar Bears. Ecology 81: 761-772. Ferguson. S. H., M. K. Taylor, E. W. Born, and F. Messier. 1998. Fractals, sea-ice landscape and special patterns of Polar Bears. Journal of Biogeography 25: 1081-1092. Finley, K. J., G. W. Miller, R. A. Davis, and W. R. Koski. 1983. A distinctive large breeding population of Ringed Seals (Phoca hispida) inhabiting the Baffin Bay pack ice. Arctic 36: 162-173. Kingsley, M. C. S., I. Stirling, and W. Calvert. 1985. The distribution and abundance of seals in the Canadian high Arctic, 1980-1985. Canadian Journal of Fisheries and Aquatic Science 42: 1189-1210. Ljungblad, D. K., S. E. Moore, J. T. Clarke, and J. C. Ben- nett. 1986. Aerial surveys of endangered whales in the north- em Bering, eastern Chukchi, and Alaskan Beaufort Seas, 1985: with a seven year review, 1979-85. OCS Study, MMS 86-0002. USDOI, MMS, Alaska OCS Region, Anchorage, Alaska. 142 pages. Moore, S. E., and J. T. Clarke. 1992. Distribution, abundance, and behavior of endangered whales in the Alaskan Chukchi and western Beaufort Seas, 1991: with a review 1982-91. OCS Study, MMS 92-0029. USDOI, MMS, Alaska OCS Region, Anchorage, Alaska. 126 pages. Stirling, I., and A. E. Derocher. 1993. Possible impacts of climatic warming on Polar Bears. Arctic 46: 240-245. Stirling, I., D. Andriashek, and W. Calvert. 1993. Habitat preferences of Polar Bears in western Canadian Arctic in late winter and spring. Polar Record 29(168): 13-24. Stirling, I., and N. A. Oritsland. 1995. Relationships between estimates of Ringed Seal (Phoca hispida) and Polar Bear (Ursus maritimus) populations in the Canadian Arctic. Canadian Journal of Fisheries and Aquatic Sciences 52: 2594-2612. Treacy, S. D. 2000. Aerial surveys of endangered whales in the Beaufort Sea fall 1998, and 1999. OCS Study, MMS 2000-066. USDOI, MMS, Alaska OCS Region, Anchor- age, Alaska. 135 pages. USDOI, Fish and Wildlife Service. 1995. Habitat conser- vation strategy for Polar Bears in Alaska. USDOI, Fish and Wildlife Service, Alaska Region, Anchorage, Alaska. 119 pages. Received 23 April 2003 Accepted 16 August 2004 New Records for the Arctic Shrew, Sorex arcticus and the Newly Recognized Maritime Shrew, Sorex maritimensis Nett D. Perry!, DONALD T. STEWART?, ELIZABETH M. MADDEN®, and THOMAS J. MAIER* 'Department of Wildlife and Fisheries Science, Texas A&M University, 210 Nagle Hall, College Station, Texas, 77843-2258 USA. Corresponding author. “Department of Biology, Acadia University, Wolfville, Nova Scotia, B4P 2R6 Canada ‘U.S. Fish and Wildlife Service, Medicine Lake National Wildlife Refuge, 223 North Shore Road, Medicine Lake, Montana 59247 USA USDA Forest Service, Northeastern Research Station, University of Massachusetts, Amherst, Massachusetts 01003-9285 USA Perry, Neil D., Donald T. Stewart, Elizabeth M. Madden, and Thomas J. Maier. 2004. New records for the Arctic Shrew, Sorex arcticus, and the newly recognized Maritime Shrew, Sorex maritimensis. Canadian Field-Naturalist 118(3): 400-404. We report the first record for the Arctic Shrew (Sorex arcticus) in the state of Montana, USA. We also report range extensions for the closely related Maritime Shrew (Sorex maritimensis) in New Brunswick and Nova Scotia, Canada. These collections augment our limited knowledge of the ranges and habitat associations of these rarely collected shrews, and highlight the need for a careful assessment of the status of S. maritimensis in Canada. Key Words: Arctic Shrew, Sorex arcticus, Maritime Shrew, Sorex maritimensis, range, state record, Montana, New Brunswick, Nova Scotia, Quebec. Approximately 38 species of shrews (genus Sorex) are currently recognized in North America (Hall 1981; Jones et al. 1986; George 1988; Wolsan and Hutterer 1998; Fumagalli et al. 1999). Most North American shrews belong to the subgenus Otisorex, which is pri- marily restricted to this continent. Until recently, the only two recognized members of the subgenus Sorex found in North America were the Tundra Shrew (Sorex tundrensis) and the Arctic Shrew (Sorex arcticus) (van Zyll de Jong 1983a), the latter species documented throughout much of the boreal forest region of North America. Recent molecular work, however, has support- ed the recognition of the Maritime Shrew (Sorex mari- timensis, previously S$. arcticus maritimensis) as a distinct species (Stewart et al. 2002). The range of the Maritime Shrew is limited to the eastern portion of New Brunswick and Nova Scotia. Sorex arcticus and S. maritimensis are infrequently observed or collected. Both species exhibit a preference for grass-sedge meadows and wetland edges (Wrigley et al. 1979; van Zyll de Jong 1983b; Kirkland and Schmidt 1996). Although there are limited studies which include estimates of density (Buckner 1966), both spe- cies appear to exist at lower population densities than other common mammal species of the boreal region (e.g., Masked Shrew [Sorex cinereus] and Meadow Vole [Microtus pennsylvanicus]), although they may be locally common in appropriate habitat (Wrigley et al. 1979). Sorex maritimensis, because of its limited range and restriction to areas of grass-sedges, is con- sidered rare (van Zyll de Jong 1983b). This may be a result of competitive exclusion by similar sized Smoky Shrews (Sorex fumeus) which exhibit a preference for less mesic, wooded habitats (van Zyll de Jong 1983a). Herein we report the first record of S. arcticus in Montana and range extensions of S. maritimensis in New Brunswick and Nova Scotia, and provide addi- tional data on habitat associations in these locations. We also report a collection of S. arcticus in Sept Hes, Quebec, one of only six records within the province (Peterson 1966; van Zyll de Jong 1983b). In the follow- ing sections, we detail trapping methods and results for each collection. Methods Montana, USA. Between 22 and 27 July 2001 we conducted small mammal baseline surveys on wet meadow habitats at Medicine Lake National Wildlife Refuge, Sheridan County, northeast Montana (48°30'N, 104°20'W; Figure 1). The refuge is located in the gla- cially influenced prairie pothole region, noted for its gentle rolling plains with occasional shallow depres- sions—host to vast wetlands and seasonally flooded meadows. We set two 150 m transects, each with 10 trap sta- tions set 15 m apart for five consecutive nights. Each station included three different Victor® snap-traps: one mouse trap, one museum special trap, and one rat trap (300 trap nights). Nearby pitfall traps consisted of two 5-m fences (aluminum flashing) with 5-L paint buckets (dry, not baited) at either end (40 trap nights; each night a bucket was open was considered a trap night). The transects were located in the Lake Creek flood plain, an area seasonally inundated with water during spring run-off and early summer rains (Stuart and Kantrud 1971). The plant community was herba- ceous, dominated by sedges (Carex spp.), grasses (Agro- pyron spp. and Spartina spp.), and rushes (Juncus 400 2004 PERRY, STEWART, MADDEN, and MAIER: ARCTIC AND MARINE SHREWS 40] FicurE 1. Suggested range for Sorex arcticus. Includes extensions presented herein and ranges suggested by Peterson (1966) and van Zyll de Jong (1983b). Insets: A + Indicates location of Montana S. arcticus collections, Medicine Lake National Wildlife Refuge. @ Indicates location of nearest known record, Lostwood National Wildlife Refuge, North Dakota. B. o Indicates the location of the Quebec S. arcticus collection, other points (@) indicate location of historical collections in Quebec. spp.), with a variety of wetland forbs interspersed. Iden- tical survey efforts were conducted in two additional habitat types on the refuge: native prairie and planted perennial grasslands. Total combined effort for all three habitat types was 900 snap trap nights and 120 pitfall nights. Quebec, Canada. One hundred pitfall traps (800 trap nights) were set at each of two sites, 17-21 July 1990 (Figure 1). Sites were 8 km north and 8 km east of Sépt Iles, Quebec (50°12'N, 66°23'W) in coniferous wood- land near the edge of a marsh and in an old grassy field next to a road, respectively. The coniferous woodland/ marsh was characterized by White Spruce (Picea glauca), Balsam Fir (Abies balsamea) and sedges (Carex spp.). New Brunswick, Canada. A total of 175 non-baited pitfall traps (525 trap nights) were set 3 km southeast of St. George, near L’Etete, New Brunswick (45°8'N, 66°50'W), 3-7 August 1990 (Figure 2). Traps were set in thickets of predominantly alder (Alnus sp.) with some mixed conifer growth and some grasses. A small brook (0.5 m wide) transected the site, maintaining relatively moist soils. Nova Scotia, Canada. [Method information is not available] Results Montana, USA. Six S. arcticus, the first confirma- tion of this species in Montana (Foresman 2001), were collected in wet meadows at Medicine Lake National Wildlife Refuge (Figure 1). The nearest known previous collection was at Lostwood National Wildlife Refuge, Burke and Mountrail counties, North Dakota, approx- imately 190 km to the east (R. Murphy, personal com- munication). 402 THE CANADIAN FIELD-NATURALIST Vol. 118 200 Kilometers _ FiGureE 2. Suggested range of Sorex maritimensis, including collections described herein. A indicates the location of the St. George collection, New Brunswick. B indicates location of the Belle Isle collection, Nova Scotia. Historic locations, indicated by solid dots, were taken from van Zyll de Jong (1983b). We found no extralimital collections since that publication. These specimens were captured using Victor® Mu- seum Specials (n = 5) and Victor® rat traps (n = 1) baited with a mixture of peanut butter, oatmeal, flour, and black sunflower seeds. No S. arcticus were col- lected in either pitfall traps or smaller Victor® mouse- traps. Although identical survey efforts were conduct- ed in native prairie and planted perennial grasslands, S. arcticus were collected only in wet meadow habi- tats. Other species collected from wet meadow sites included: Meadow Vole (n = 19), Deer Mouse (Pero- myscus maniculatus, n = 1), and Masked Shrew (n = 8). All specimens were confirmed as S. arcticus; how- ever, they were collected on the periphery of the S. a. arcticus and S. a. laricorum ranges, and identification to subspecies is difficult. These specimens are cata- logued at the Philip L. Wright Zoological Museum at the University of Montana, Missoula (catalog numbers UMZM 18554 — 18559). Quebec, Canada. Two Arctic Shrews (S. a. arcticus; Royal Ontario Museum [ROM] catalogue numbers 110254 and 110255) were collected by D.T-.S. near Sept Iles, Quebec (50°12'N, 66°23'W), 17-21 July 1990 (Figure 1). This is the sixth reported collection of S. arcticus in this province (Peterson 1966; van Zyll de Jong 1983b). Both specimens were collected from the grassy site east of Sept Iles. Other species collected during this effort were Masked Shrew (n = 7), Pygmy Shrew (Sorex hoyi; n = 1), and Meadow Vole (n = 1). This record is as far north along the immediate coast of the St. Lawrence River as S. arcticus have previously been reported. The last specimen in this general area, near Moisie River, just north of Sept Iles, was trapped in 1937 (van Zyll de Jong 1983b). Though little trapping has been conducted in this region, van Zyll de Jong (1983b) speculates that the species is distributed further north in Quebec and Labrador throughout the boreal forest, which includes extensive marshy habitats. New Brunswick, Canada. Three Maritime Shrews (originally identified as S. a. maritimensis; ROM cata- logue numbers 110314, 110315, 110331) were col- lected 3 km southeast of St. George, near L’Etete, New Brunswick (45°8'N, 66°50'W), 3-7 August 1990 (Fig- ure 2). This collection extends the known range of S. maritimensis ca. 100 km south. Other species collected from this effort were Masked Shrew (n = 21) and Northern Short-tailed Shrew (Blarina brevicauda; e—1))' Nova Scotia, Canada. On 4 October 1992, a single specimen of Sorex maritimensis (originally identified as §. a. maritimensis) was collected at Belle Isle, Nova Scotia (Tom Herman, personal communication). The specimen, collected in a marshy area dominated by the sedge Scirpus cyperinus, represents a provin- cial range extension of this species by ca.100 km (Figure 2). 2004 Discussion Sorex arcticus. The range of the Arctic Shrew is strongly associated with the boreal forest region of North America. Southward range expansions of four other boreal species, Masked Shrew, Meadow Vole, Meadow Jumping Mouse (Zapus hudsonius), and the Least Weasel (Mustela nivalis), have been correlated with contemporary cool, mesic climate patterns in the Great Plains region (Frey 1992). Jannett and Huber (1994) speculate that a recent southward extension of S. arcticus in Minnesota is associated with these cool- ing climate patterns. Indeed, this southwestward exten- sion of S. arcticus into Montana could be correlated with this phenomenon. Nevertheless, few small mammal surveys have been conducted in northeastern Montana (D. Flath, person- al communication); it is possible this has prevented earlier detection in the state. Similar wet meadow habitat exists approximately 40 km to the south as the Big Muddy Creek feeds into the Missouri River. Thus, it is plausible that the range of S. arcticus ex- tends further into Montana. All Montana S. arcticus specimens were sexually inactive young-of-year. Clough (1963) observed that over-wintering S. arcticus captured between February and July were reproductively active. Also, the char- acteristic tri-colored pelage for adults of this species was indistinct in these specimens, further supporting our designation of these shrews as young-of-year, products of early season breeding by the previous year’s cohort (Clough 1963; Baird et al. 1983). These specimens were collected in two groups of three, each group approximately 3 km apart and separated by a small perennial stream. Given their inactive re- productive status and that individuals from each group were collected no greater than 15 m apart, group mem- bers might have been litter-mates. As such, a viable population of S. arcticus likely exists at Medicine Lake National Wildlife Refuge. Sorex maritimensis. The Maritime Shrew was pre- viously considered restricted to the north and east of the St. John River system, with the nearest previous collection for this species near Saint John, New Bruns- wick (Peterson 1966; van Zyll de Jong 1983b). The collection of S$. maritimensis near L’ Etete, New Bruns- wick, brings this species within 30 km of the Maine border. Maine and New Brunswick are separated in this area by the St. Croix River which could be a bar- rier to shrew dispersal. There is suitable habitat for S. maritimensis on the U.S. side of the border and it is possible they will be found there; however, to date there is no record of S. maritimensis in Maine (J. Albright, R. Boone, and L. Master, personal communications). S. maritimensis is currently recognized as one of only four mammals endemic to Canada (the others are the Varying Lemmings /Dicrostonyx hudsonius D. richard- soni], the Gaspé Shrew [Sorex gaspensis], and the Vancouver Marmot [Marmota vancouverensis]). PERRY, STEWART, MADDEN, and MAIER: ARCTIC AND MARINE SHREWS 403 The Nova Scotia and New Brunswick collections of the newly recognized S. maritimensis imply a greater range, extending inland and likely including the entire- ty of mainland Nova Scotia. Stewart et al. (2002) speculate that recent glacial encroachment (ca. 20 000 y) may have isolated this species on the coastal flood plain of Nova Scotia. These recent collections may either provide evidence that S. maritimensis is re- colonizing former range, following the reestablish- ment of the boreal forest after the last ice age, or may reflect the paucity of efforts to collect S. maritimensis within suitable habitats in these provinces. These col- lections also support a hypothesis that this species is limited to moist grasslands and bogs associated with the boreal forest, competitively excluded from habitats occupied by the Smokey Shrew, a closely related wood- land associate. This limited distribution and restriction to fragile wet meadow habitats suggests this species may warrant conservation concern. These collections offer some data regarding the range and niche characteristics of these shrews. Given that S. arcticus and S$. maritimensis have been the focus of very few research efforts, more research targeting specific life history traits are necessary to better under- stand the habitat associations and range restrictions of these shrew species. Acknowledgments We particularly thank Tom Herman for providing data on his specimen of S. maritimensis collected in Nova Scotia and Kerry Foresman for facilitating the processing and accession of Montana specimens into the University of Montana mammal collection. We also thank the staff at Medicine Lake National Wildlife Refuge for funding and support of small mammal col- lections on the refuge. We thank D. Flath and R. Mur- phy for insight into the status of S. arcticus in Mon- tana and North Dakota. DTS thanks S. Hindocha and H. Russell for assistance in the field; and J. Albright, R. Boone, and L. Master for comments on the status of S. maritimensis in Maine. The research program of DTS is supported by an NSERC Discovery grant. 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Received 12 May 2003 Accepted 21 December 2004 Lichen Zonation on Coastal Rocks in Gwaii Haanas National Park Reserve, Haida Gwaii (Queen Charlotte Islands), British Columbia IRWIN M. Bropo! and Norm A. SLOAN2 ‘Canadian Museum of Nature, P. O. Box 3443 Station D, Ottawa, Ontario K1P 6P4 Canada > Parks Canada, Gwaii Haanas National Park Reserve and Haida Heritage Site, P. O. Box 37, Queen Charlotte City, British Columbia VOT 1SO Canada Brodo, Irwin M., and Norm A. Sloan. 2004. Lichen zonation on coastal rocks in Gwaii Haanas National Park Reserve. Haida Gwaii (Queen Charlotte Islands), British Columbia. Canadian Field-Naturalist 118(3): 405-424. The occurrence of 43 marine lichen species on intertidal rocky shores of southern Haida Gwaii (Queen Charlotte Islands), British Columbia is described and related particularly to elevation on the shore (duration of seawater immersion) and exposure to waves. In the area of Gwaii Haanas National Park Reserve and Haida Heritage Site on Moresby Island, rock- dwelling marine lichens are distributed in zones much as they are elsewhere in the world, although some species found abundantly only in Haida Gwaii give the local shores a unique appearance. In common with other areas, there is a conspicuous black band of Verrucaria species (in this case, nine species plus other black lichens) at the upper edge of the intertidal zone. A conspicuous white band of Coccotrema maritimum above the black band is a unique feature of this flora. The unusually large percentage of endemic, near-endemic or disjunct lichen species and their phytogeography suggest that at least the headland rocks along the west coast were refugia during the last glacial maximum. Verrucaria striatula and V. sandstedei are reported for the first time from British Columbia. Key Words: Maritime lichens, zonation, Haida Gwaii, Pacific Northwest, British Columbia. Lichens form a dominant, yet often overlooked, component of the maritime vegetation along rocky sea- shores. They create distinct bands along such shores from polar to tropical seas with maximum abundance, and the most studies, in north-temperate regions (Flet- cher 1980; Little and Kitching 1996). The species res- ponsible for these bands in the northeast Pacific (1.e., the Pacific Northwest of North America; see the cover photo, and Plate 64 in Brodo et. al. 2001: 77), however, have rarely been investigated. The shoreline lichen vegetation of Gwaii Haanas National Park Reserve and Haida Heritage Site can be understood in the context of the total lichen flora of Haida Gwaii, British Columbia. Five field trips by the senior author to Haida Gwaii since 1967 have revealed a diverse lichen flora of about 580 species with more than 25 new to science (Brodo 1995). Many of the spe- cies are rare worldwide, and more than a dozen are en- demic to Haida Gwaii or are known only from a few nearby localities in southeast Alaska and Vancouver Island. Further, many species are disjuncts, having their nearest additional locality thousands of kilometres dis- tant, often in western Europe, Asia or the southern hemisphere (Brodo 1992). With the establishment of Gwaii Haanas as a Na- tional Park Reserve, there is a need to develop a base- line inventory of lichen species for various uses, for example, to facilitate the conservation of lichen bio- diversity (Hunter and Webb 2002), to track human and natural perturbations, to assist in nature interpretation, and to aid studies of shoreline ecology. Lichens are well-known indicators of environmental health in use world-wide (Nimis et al. 2002). Regionally, they have been used for air quality monitoring in the Tongass National Forest in southeast Alaska (Geiser et el. 1994*). In addition, lichen studies have been made to evaluate the effects of oil spills and post-spill cleaning on marine ecosystems (Lallemant and Van Haluwyn 1981; Flet- cher and Crump 2002). A knowledge of maritime lich- ens is clearly relevant to the prospect of oil and gas development in the Hecate Strait separating Haida Gwaii from the British Columbia mainland (Anony- mous 2002*). It is important to know if the shoreline lichens of Haida Gwaii differ from those found elsewhere on the west coast of North America. The only other detailed study of Pacific Northwest marine lichen zonation was done by Ryan (1988a, 1988b) in Washington State, al- though some observations on shoreline lichens from southern British Columbia were presented by Noble (1982). We report here on the distribution of lichen species in the upper rocky intertidal zone landward into the salt spray zone in the Gwaii Haanas area. Because splash patterns and exposure affect establishment and growth of shoreline lichens, we examined rocky shores with a range of exposures to wave action from “protected” to “very exposed” to reveal species occurrence patterns. Finally, we compared the rock-dwelling marine lichen flora and zonation of southern Haida Gwaii with the flora and patterns seen in other regions. Study Area Haida Gwaii, known also as the Queen Charlotte Islands, is an archipelago of about 138 islands in the hypermaritime zone of the Pacific Northwest, lying 405 406 about 80 km off the mainland British Columbia coast between Vancouver Island and southeast Alaska (Fig- ure |). Its geography, climate and vegetation were des- cribed in some detail in Brodo (1995). The two largest land masses of Haida Gwaii are Graham Island to the north and Moresby Island to the south. The lower elevations of Graham Island (the eastern two-thirds) and Moresby Island (the northern third) have been disturbed by intensive logging activ- ities, but the southern two-thirds of Moresby Island were set aside in 1988 as a National Park Reserve called Gwaii Haanas lying roughly between 52° and 53°N longitude (Figure 2) covering ca. 1470 km? of land and ca. 1700 km of shoreline (Sloan and Bartier 2000). The lichen studies described here mainly centre on the extreme southern end of the Park Reserve, at the tip of Moresby Island and on Kunghit Island (Fig- ures 2 and 3). Methods To characterize our sample locations, we used Gwaii Haanas’ shoreline classification system (Harper et al. 1994*). This is in the park’s geographic information sys- tem (GIS) and is based largely upon substrate type, texture and exposure to wave action. This system was first developed in Gwaii Haanas and has since become the standard physical shoreline classification system of British Columbia (Howes et al. 1994*) and Washington State [http://www2.wadnr.gov/nearshore/index.asp]. The system has a biological zoning component in which visible lichens are coarsely grouped into a “Verrucaria band” (Searing et al. 1995). The British Columbia scheme has been used in broad-scale region- al studies of intertidal species diversity (Zacharias and Roff 2001). In Harper et al. (1994*), lichens are mentioned as forming conspicuous shoreline bands. There are refer- ences to the “Verrucaria zone” and the “white lichen zone,” but the diversity of species is overlooked. For example, it is not made clear that the Verrucaria zone consists of nine black Verrucaria species and other dark lichens (Brodo and Santesson 1997), and that the white zone is caused by the dominance of Coccotrema maritimum, a species described from Haida Gwaii, whose closest relatives are in Asia and the southern hemisphere. It is rare in the Pacific Northwest outside Haida Gwaii and adjacent coastal regions (Brodo 1973). Other lichens of intertidal rocks also proved to be new to science and generally restricted to Haida Gwaii and nearby coastal localities. These include Caloplaca litoricola, Fuscidea thomsonii, Verrucaria schofieldii, V. epimaura, and Porina pacifica (Brodo 1984: Brodo and Wirth 1998; Brodo and Santesson 1997; Brodo 2004). Several other novelties remain to be formally described. Fourteen locations within Gwaii Haanas and one on Limestone Island were sampled (Figures 2, 3). At each location, one to three transects were laid out for a total of 18 transects. A 12 m tape measure was deployed THE CANADIAN FIELD-NATURALIST Vol. 117 conforming closely to the rock surface and perpendi- cular to the high tide line. Transects usually began at the upper limit of the rockweed (Fucus gardneri) and barnacle (Balanus and Semibalanus spp.) zone (the “FB zone,” approximating the mean high water level [MHW)]) and ran landward to the point at which terres- trial vascular plants or mosses began. This distance was divided into eight or nine equal sections to give approximately nine to ten quadrat points. Because ex- posed, wave-splashed beaches and shallow slopes pro- duced longer transects than those on protected, rela- tively steep shores, the increments of height above the FB zone from quadrat to quadrat were not uniform, nor were the distances between quadrats. At each quadrat point, a 10 x 10 cm grid on trans- parent plastic with a hundred 1-cm? squares, was cen- tered on the transect tape. All lichen species under the grid were recorded together with their coverage in percent (each square representing 1% coverage). For smooth rocks, the coverage readings were fairly accu- rate, but with increasing surface roughness, the esti- mates were correspondingly approximate. After the quadrats were sampled and the lichen data recorded, the vertical height of each quadrat above the starting point in the FB zone was recorded. Results The 18 transects were sorted and analyzed accord- ing to degree of shoreline exposure from Protected to Very Exposed, based on Harper et al. (1994*), as listed in Table 1. Two locations had multiple transects, there were no transects for localities 6 or 10, and transect 11 had incomplete data and was excluded. Limestone transects are indicated (L) and all other transects were over mixed volcanic and fine-grained sedimentary rocks characteristic of the region’s shoreline (Sutherland Brown and Yorath 1989). The transects in Gwaii Haanas yielded 43 lichen species. Sixty-seven rock-dwelling species occurring on maritime rocks are currently known from Haida Gwaii as a whole. An annotated list of all shoreline species in Haida Gwaii is presented as an Appendix. This updates the list in Sloan and Bartier (2000; Ap- pendix B, Part 2) compiled by the senior author based on specimens at the National Herbarium of Canada (CANL), Ottawa. The zones indicated in Sloan and Bartier (2000) were assigned tentatively, based on label data, and have been corrected in this study. Two of the lichens from our transects (Verrucaria striatula and V. sandstedei) are new lichens for Haida Gwaii (and British Columbia), and ten others were not includ- ed in the Sloan and Bartier (2000) list because they were not regarded to be maritime. These are: Adelole- cia kolaensis, Gyalecta jenensis, Opegrapha gyro- carpa, Parmelia saxatilis, Placopsis lambii, Porpidia contraponenda, P. speirea and P. thomsonii. Many rela- tively common maritime and marine lichens found elsewhere in Haida Gwaii were not seen in the Gwaii Haanas transects. These include Caloplaca inconspecta, 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 407 Langara 1327 'W Island Dixon Entrance Canada Masset e f , British A Columbia | + ak Graham Island Hecate Queen Charlotte : Citys. Skidegate Strait ® Sandspit Louise Island Moresby Island GD Gwaii Haanas Burnaby A Island National Park Reserve and Haida Heritage Site / proposed National Marine Conservation Area SGaang Gwaii Kunghit Island Queen 100 Km Cape Charlotte St. James Sound FicurE 1. Location of Gwaii Haanas within Haida Gwaii (Queen Charlotte Islands) including some of the larger islands. 408 132°W Louise Island va Moresby Island Gwaii Haanas National Park Reserve and Haida Heritage Site and proposed National Marine Conservation Area THE CANADIAN FIELD-NATURALIST Vol. 117 131°W Ron 18 Limestone Island Lyell Hecate Island eunah Strait Burnaby Island Inset 2 17 Kunghit Treat Bay Island GOON 50 Kilometres 132°W 131°W FIGURE 2. Map of the Gwaii Haanas area showing Limestone Island (location 18) to the north and Treat Bay (location 17) to the south of the insets (see Figure 3 for insets). Catillaria calybeia, Rhizocarpon hensseniae and Rino- dina gennari. At least one lichen (Porina pacifica) was found growing abundantly just outside the quadrats on a protected shore. The frequency of occurrence (%) of each species in each transect is listed according to shore exposure class in Table 2. Verrucaria maura occurred in all tran- sects on all shores. Among the eight most abundant lichens (frequencies exceeding 50% in all transects), four are endemic or nearly endemic to Haida Gwaii: Caloplaca litoricola, Coccotrema maritimum, Verru- caria epimaura and V. schofieldii; and one is endemic to the coastal Pacific Northwest (Herteliana alasken- sis). The semi-exposed shores had the most species and the most exposed shores yielded the fewest species. The results of the transect observations, expressed as metres above the FB (Fucus/barnacle) zone, are given in Tables 3 and 4 organized according to the degree of 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS ; Ee ; F bY e TS phelican Foint Tnset pre wie, eas Skincuttle han i 4 ; ee : Hecate : \ # &. a Ka Strait ns Gamenten 8 7 - Ingraham Point Bay ray ¢ 5 Kilometres Houston Stewart Channel Islands 4 Kilometres SSS Ficure 3. Inset maps from Figure 2 showing lichen sampling locations 1-16 within Gwaii Haanas (see Table 1). 409 410 THE CANADIAN FIELD-NATURALIST Vol. 117 TABLE 1. Locations of shore lichen transects in the Gwaii Haanas area grouped according to the shore exposure classification of Harper et al. (1994). Exposure Class Location Name Transect No.* Latitude (°N) Longitude (°W) Protected Carpenter Bay 9 131.1660 52.2312 SGaang Gwaii 13 13022511 S2a1015 Limestone Island 18(L>) 131.6170 52.9086 Semi-protected Ellen Island 1 131.0918 52.1544 Orion Point 5) 131.0231 52.1229 Semi-exposed Gordon Islands 2-1, 2-2, 2-3 131.1443 52.0967 Orion Point + 131.0217 52.1241 Pelican Point 14 (L’) 131.2615 52.3472 Pelican Point 15 131.2618 52.3458 Houston-Stewart Channel 16-1, 16-2 131.1261 21258 Treat Bay ity) 131.0080 52.0659 Exposed Ingraham Point 7 131.0325 32.232) Ingraham Point 8 131.0331 52.2319 Very exposed Gordon Islands 3 131.1493 52.0965 SGaang Gwaii 1 131.2330 52.1021 4 Two locations (2 and 16) had multiple transects, locations 6 and 10 had no transects, the transect at location 11 was in- complete and excluded. >L = limestone rock exposure of each shore. These data are graphed Figures in 4 and 5, with the ordination based on average mini- mum values (the base of the thick black line). Because the values for all shore exposures except “Very Ex- posed” appear to largely overlap with respect to the height above the FB zone, the data are pooled in Table 3 and Figure 4; Table 4 and Figure 5 shows the data only for Very Exposed shores. The data for Quadrat Position (Table 5), being relative, are not greatly affect- ed by shore exposure, so all shores are considered together. The results of all transects with 8-10 quadrats per transect are shown in Figure 6. (Transects having only 6 or 7 quadrats cannot be compared with those having 8-10 transects.) Translating the continuum of species position into a zonal scheme with names that correspond with previ- ous North American or European studies proved to be challenging. The graphs presented in Figures 4 and 5 based on metres above the upper limit of the FB zone (Tables 3 and 4) were compared to Figure 6 showing the species according to their quadrat placement along the transects (Table 5) with the aim of finding natural breaks in the continua. This was only partially suc- cessful. Thus, in Table 6, which presents a summary of the zones, their position and their dominant lichens, a few species are listed twice, in different zonal cate- gories, according to height above the Fucus-barnacle limit versus their relative quadrat position. Our zonal terminology basically follows the British System, es- pecially Fletcher (1973a, 1973b) who, in agreement with Lewis (1961), argued that the zones are best described and defined on biological criteria, not on physical (i.e., tidal) criteria. Lewis (1961) recommend- ed adopting “Littoral Fringe” in place of “Supralit- toral Fringe” used by Stephenson and Stephenson (1949) because it better describes the community that inhabits the upper fringe of the intertidal zone, washed by most tides in relatively protected shorelines. We replace Mesic, Submesic and Xeric subdivisions of the Supralittoral Zone with the more easily understood terms Lower, Middle and Upper, respectively. In addi- tion, we find that the term “xeric,” even taken in its relative sense, is not an accurate description of a shore- line habitat. Ryan (1988b) used a numbering system for his zones, but they basically followed the British scheme following the distribution of shoreline organ- isms. O’Clair et al. (1996) used a simplified system based on wave exposure, and this was adopted by Sloan and Bartier (2000). Some zonal synonymy, discussed in detail by Ryan (1988b), is given in the first column of Table 6. In the Gwaii Haanas area, Verrucaria mucosa and Pyrenocollema halodytes are the most abundant spe- cies at the upper edge of the intertidal zone; i.e., the lowest point on our transects, where rockweed or bar- nacles occur (Lower Littoral Fringe; quadrat 1). They grow there together with some of the smaller and rarer species of Verrucaria such as V. sandstedei and V. stri- atula. These are generally too small, thin, and scattered to create a black zone and, therefore, are hard to find. The black zone begins with the appearance of the blacker, thicker species of Verrucaria: V. erichsenii, V. maura, V. epimaura and V. schofieldii, all of which can have broad coverage (Table 7) in the Upper Lit- toral Fringe (quadrats 2-4). Although V. maura con- tinues to be important well up on the shoreline, the community changes its character and color with the appearance of the gray-brown, gray, and orange species such as Herteliana alaskensis, Arthonia phaeobaea, and the near-endemic but abundant Haida Gwaii lichen, Caloplaca litoricola, in the Lower Supralittoral Zone (quadrats 5-7). Also in this zone is Adelolecia kolaensis and the cyanophilic lichen Spilonema revertens. Above this is a poorly defined orange zone (Middle Supra- EE qC“eer_ 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 41] TABLE 2. Percentage occurrence of shoreline lichen species at locations according to shore exposure classification of Harper et al. (1994) in the Gwaii Haanas area. P = protected; SP = semi-protected; SE = semi-exposed; E = exposed; VE = very exposed. % occurrence according to shore exposure class % Lichen species P (3) SP (2) Verrucaria maura 100 100 Spilonema revertens 67 100 Caloplaca litoricola 33 100 Coccotrema maritimum 67 50 Verrucaria epimaura 0 100 Herteliana alaskensis 67 100 Pyrenocollema halodytes 67 0 Verrucaria schofieldii 67 100 Arthonia phaeobaea 33 50 Verrucaria amphibia 100 0 Lecidella stigmatea 67 100 Rhizocarpon geminatum 33 50 Xanthoria candelaria 0 50 Amandinea coniops 0 100 Physcia caesia 33 50 Porpidia speirea 0 Verrucaria erichsenii 0 Verrucaria mucosa 67 Caloplaca flavogranulosa 33 Collema fecundum 33 Fuscidea thomsonii Adelolecia kolaensis Verrucaria degelii Verrucaria sandstedei Lecanora muralis Ochrolechia subplicans Parmelia saxatilis Placopsis lambii Porpidia thomsonii Rhizocarpon haidensis ined. Aspicilia caesiocinerea Caloplaca citrina Caloplaca rosei Caloplaca verruculifera Candelariella sp. Fuscopannaria maritima Gyalecta jenensis Lecanora sp. #2 Lecanora sp. #1 Lecanora straminea Opegrapha gyrocarpa Opegrapha sp. Porina pacifica Porpidia carlottiana Porpidia contraponenda Rhizocarpon hochstetteri s. str Verrucaria sp. #5 Verrucaria striatula i>) Sees Nn Nn Nn Nn SESS nO SIS OSS 2 OS 2 SEO OES TOO So. OrO rr Orr Oo Orr WwW SrOL ISTO Dru iOrO LOO OS OOS Os OOS O20 FSGS nn nN ies) nn Oo — \o i) Total lichen species littoral Zone; quadrats 7-8) with Caloplaca flavogran- ulosa, C. rosei (both west coast endemics) and the widespread Xanthoria candelaria, as well as the in- conspicuous crust, Rhizocarpon “haidensis.” The Upper Supralittoral Zone (quadrats 8 — 9) almost at the upper edge of the beach is well marked by the dom- inance of the pinkish white lichen, Coccotrema mari- (number of transects) occurrence in SE (9) E (2) VE (2) all transects 100 100 100 100 56 100 50 67 56 100 50 61 67 100 0 61 67 50 100 61 33 100 50 56 56 50 100 56 56 0 0 50 33 100 50 44 33 50 0 39 0 0 100 33 22, 0 50 28 22 0 100 28 22 0 0 22 22 0 0 22 44 0 0 22 33 0 50 22 yp) 0 0 22 22, 0 0 17 11 50 0 17 22 50 0 17 0 50 0 17 33 0 0 17 33 0 0 17 0 0 50 11 11 50 0 11 11 50 0 11 22 0 0 11 11 50 0 11 22 0 0 11 0) 0 0) 6 11 0 0 6 0 0 0 6 0 0 50 6 0) 0 0) 6 11 0 0 6 11 0 0 6 0 0 0 6 0 0 0 6 0 0 50 6 0 0 0 6 11 0 0 6 0 0 0 6 0 0 0 6 11 0 0 6 0 0 0 6 11 0 0 6 0 0 50 6 33 15 15 timum, which forms a conspicuous white band on the coastal rocks of Haida Gwaii, especially on the Pacific side, although it is less common outside of the archi- pelago. Accompanying the Coccotrema are such wide- spread lichens as Placopsis lambii, various species of Rhizocarpon, and rarer west coast species such as Och- rolechia subplicans. Higher on the shore and influ- 412 THE CANADIAN FIELD-NATURALIST Vol. 117 TABLE 3. Summary data for Protected to Exposed shores, metres above Fucus/barnacle zone (16 quadrats). Species # Extreme quadrants minimum Adelolecia kolaensis 3 135 Arthonia phaeobaea 7 0.55 Aspicilia caesiocinerea 1 2.05 Caloplaca citrina 1 1.05 Caloplaca flavogranulosa 3 1.24 Caloplaca litoricola 10 0.55 Caloplaca rosei 1 2.05 Candelariella sp. 1 2.65, Coccotrema maritimum 11 1.10 Collema fecundum 3} 0.80 Fuscidea thomsonii 3 2.20 Fuscopannaria maritima 1 2.40 Gyalecta jenensis 1 0.90 Herteliana (Bacidia) alaskensis 9 0.60 Lecanora muralis 2 2.65 Lecanora sp. no.1 1 0.55 Lecanora sp. no.2 1 0.90 Lecidella stigmatea 4 0.60 Ochrolechia subplicans 2 1.80 Opegrapha gyrocarpa 1 1.83 Opegrapha sp. 1 1.80 Parmelia saxatilis 2 2.00 Physcia caesia 4 1.00 Placopsis lambii p) 1.85 Porina pacifica if 2.70 Porpidia carlottiana 1 1.50 Porpidia contraponenda 1 1.60 Porpidia speirea + 1.10 Porpidia thomsonii 2 2.00 Pyrenocollema halodytes 8 -0.20 Rhizocarpon geminatum 4 1.70 Rhizocarpon haidensis y) E25) Rhizocarpon hochstetteri s. str 1 1.30 Spilonema revertens 11 0.60 Verrucaria amphibia 7 0.20 Verrucaria degelii 3 0.30 Verrucaria epimaura 9 0.25 Verrucaria erichsenii 3 0.00 Verrucaria maura 16 -0.20 Verrucaria mucosa 4 -0.20 Verrucaria sandstedei 3 0.00 Verrucaria schofieldii 9 -0.15 Verrucaria sp. #5 1 0.90 Xanthoria candelaria 3 1.40 enced only by intermittent salt spray, is the Terrestrial Zone (quadrats 9-10), characterized by widespread non- maritime but salt-tolerant species such as Parmelia saxatilis as well a Haida Gwaii endemic, Fuscidea thomsonii, and a few lichens restricted to the Pacific Northwest, such as Fuscopannaria maritima and Por- pidia carlottiana. The bird rock community (ornitho- coprophilous lichens), including Lecanora straminea, L. muralis, and Caloplaca verruculifera, also occurs in this zone. In comparing the graph showing zonation on Pro- tected to Exposed shores (Figure 4) with the graph deal- ing with Very Exposed shores (Figure 5), it is imme- diately apparent that on very exposed shorelines, most Average Average Extreme minimum (m) maximum (m) maximum (m) 1.43 2.03 BES 1.04 1.34 2.40 2.05 3.15 3.15 1.05 1.05 1.05 1.85 1.85 2.20 1.07 1.43 Balls 2.05 2.65 2.65 2.65 2.65 2.65 2.05 239) 3.60 1.40 1.50 1.90 2.73 D213. 3.60 2.40 2.40 2.40 0.90 0.90 0.90 1.08 1.20 2.45 2.65 2.65 2.65 0.55 0.55 0.55 0.90 0.90 0.90 25 1.30 2.05 2.10 2.40 2.40 1.83 1.83 1.83 1.80 2.30 2.30 2.20 2.20 2.40 1.11 1.99 Sul 1.93 Pil} 2.40 2.70 3.60 3.60 1.50 1.50 1.50 1.60 1.60 1.60 2.28 2.44 3.60 2.05 2.05 2.10 0.00 0.00 0.35 2S 2.61 Bulld 2.08 DSS 2.90 1.30 1.30 1.30 1.65 2.21 3.60 0.44 0.61 0.69 0.32 0.32 0.35 0.45 0.81 DDS 0.35 0.87 0.90 0.45 1.93 3.60 -0.05 0.08 0.20 0.20 0.33 0.35 0.62 0.78 1.50 0.90 1.05 1.05 1.97 233 3.65 of the lichen species are much higher on the shore, the zones themselves are much broader (reflecting the height of wave splash), and not all species are in the same relative quadrat position. For example, the cyano- bacterial lichen, Spilonema revertens, which mainly occurs in the central section of the transects (Middle Suppralittoral) on Protected to Exposed shores, is found at the top of the transects (Terrestrial) on Very Exposed shores. Verrucaria maura begins to invade the rock over a metre above the Fucus/barnacle limit, and other shoreline lichens are proportionally higher on the shore. The five species at the top of the transects are terrestrial, but salt-tolerant, lichens, not really mari- time. Also unique to the Very Exposed shores are the 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 413 TABLE 4. Summary data for Very Exposed shores, metres above Fucus/barnacle zone (2 transects). Species # Extreme quadrants minimum Arthonia phaeobaea | 2.05 Caloplaca litoricola | 2.05 Caloplaca verruculifera I 3.65 Herteliana alaskensis I 3.65 Lecanora muralis | 6.70 Lecanora straminea | 3.65 Lecidella stigmatea 2, Poet fe) Pyrenocollema halodytes 2 0.00 Rhizocarpon geminatum 1 6.70 Spilonema revertens | 5.70 Verrucaria epimaura 2 1-75 Verrucaria erichsenii ] 0.70 Verrucaria maura 2 1.10 Verrucaria schofieldii I 1.10 Verrucaria striatula | 0.00 Xanthoria candelaria 2 3.65 large areas of uncolonized rock (bare of lichens as well as barnacles or marine algae). The relative zonal position of some lichen species on protected shores differs when comparing relative quadrat position and actual distance above the FB zone (compare Figures 4 and 6). This may be due to the ability of these species to occupy broader zones than others, coupled with the method of averaging mini- mum and maximum values to draw the charts. The species included in more than one zonal category are marked with an asterisk (*) or exclamation (!) in Table 6. Species richness (diversity) appears to be greatest on Semi-exposed shores (33 species) but, since this category of exposure has the largest number of tran- sects (9), the significance of this number is difficult to evaluate. If one combines the lists in Table 2 for Pro- tected and Semi-protected shores (5 transects) and those of Exposed and Very Exposed shores (4 tran- sects), the numbers of species are closer: P+SP: 29 species; SE: 33 species; E+VE: 23 species. Interest- ingly, the Very Exposed shores, with only 16 species, include 3 species found in no other exposure type, Verrucaria striatula, Caloplaca verruculifera and Lecanora straminea. One indication of species importance along the rocky shores is the ability of each species to form large patches, 1.e., with high coverage values. Table 7 sum- marizes the coverage classes of all the species that appeared in quadrats in this study. Some species, and even species groups, are clearly able to cover larger areas than others. Almost all the black Verrucaria spe- cies, for example, are able to cover over 50% of the 100 cm? quadrats, which is why they form a conspic- uous black belt on the shore. Similarly, Coccotrema maritimum forms large patches as does Caloplaca litoricola and Ochrolechia subplicans. Some species never dominate their zones, however, always occur- ring in small patches even if they are frequently en- Extreme maximum (m) Average minimum (m) Average maximum (m) 2.05 2.05 2.05 2.05 3.65 3.65 3.65 3.65 3.65 3.65 3.65 3.65 6.70 6.70 6.70 3.65 3.65 3.65 4.73 4.83 6.70 0.00 0.00 0.00 6.70 6.70 6.70 5.70 6.70 6.70 2:23 2.38 2.70 0.70 1.70 1.70 1.40 4.83 6.70 1.10 1.10 1.10 0.00 0.00 0.00 5.18 5.18 6.70 countered. Examples are Spilonema revertens, Pyreno- collema halodytes, Herteliana alaskensis and Arthonia phaeobaea. Rock type generally has a major influence on spe- cies composition in lichen communities, but some spe- cies are more tolerant of varying rock type than others. Verrucaria mucosa was especially abundant on lime- stone although present on siliceous rock as well. Some lichens found only, or most commonly, on the lime- stone transects were Verrucaria amphibia, V. species number 5, Caloplaca citrina and Gyalecta jenensis. Pyrenocollema halodytes occurs on limestone rocks as well as on barnacles (both rich in calcium carbonate). On the other hand, many common lichens; e.g., Cocco- trema maritimum, Caloplaca litoricola, Verrucaria epimaura, Herteliana alaskensis and Collema fecun- dum, were found only on siliceous rock. Species appar- ently oblivious to rock chemistry include Verrucaria maura and V. schofieldii. Discussion Marine and maritime lichens distribute themselves on rocky shorelines in response to many factors, with the dominant influences being wave exposure and, particularly, immersion duration (Southward 1958; Fletcher 1973a; Taylor 1982). Fletcher (1973a) said that lichens such as Verrucaria mucosa and V. striatula, which grow among the barnacles in the upper inter- tidal zone, require 52% submergence per year, the highest amount for any lichen. There is experimental evidence (Ramkaer 1977) that spore germination and hyphal growth at young stages of development are influenced by salt concentrations, and that the ability of different lichens to become established in shoreline habitats is positively correlated with their position on shoreline rocks. Those lowest on the shore (longest immersion duration) are the species that have best germination and hyphal growth under moderate or even high concentrations of seawater. Mature thalli 414 THE CANADIAN FIELD-NATURALIST Table 5. Summary data for relative quadrat position for all transects with 8-10 quadrats (15 transects). Species # Extreme quadrants minimum Adelolecia kolaensis 3 4.00 Amandinea coniops 4 6.00 Arthonia phaeobaea 7 4.00 Aspicilia caesiocinerea 1 6.00 Caloplaca flavogranulosa 3 6.00 Caloplaca litoricola 10 4.00 Caloplaca rosei 1 7.00 Caloplaca verruculifera 1 9.00 Candelariella sp. 1 7.00 Coccotrema maritimum 9 6.00 Collema fecundum 3 5.00 Fuscidea thomsonii 3 9.00 Fuscopannaria maritima 1 9.00 Herteliana alaskensis 10 3.00 Lecanora muralis 2 7.00 Lecanora straminea 1 9.00 Lecanora sp. no.1 1 4.00 Lecanora sp. no.2 1 3.00 Lecidella stigmatea 6 4.00 Ochrolechia subplicans 2 7.00 Opegrapha gyrocarpa 1 8.00 Parmelia saxatilis 2 6.00 Physcia caesia 3 6.00 Placopsis lambii 2 8.00 Porina pacifica 1 7.00 Porpidia carlottiana 1 9.00 Porpidia speirea 3 6.00 Porpidia thomsonii 2 8.00 Pyrenocollema halodytes 9 1.00 Rhizocarpon geminatum 5 5.00 Rhizocarpon haidensis 2 7.00 Rhizocarpon hochstetteri s. str 1 8.00 Spilonema revertens 11 5.00 Verrucaria amphibia =) 1.00 Verrucaria degelii 2 1.00 Verrucaria epimaura 10 2.00 Verrucaria erichsenii 4 1.00 Verrucaria maura 15 1.00 Verrucaria mucosa 2 1.00 Verrucaria sandstedei 3 1.00 Verrucaria schofieldii 8 1.00 Verrucaria striatula 1 1.00 Xanthoria candelaria 5 6.00 might be affected by osmotic problems, acidity (sea water has a relatively high pH), and other seawater minerals. Other factors that influence zonation include degree of wetness and seepage of acidic fresh water from the land above the shore (Fletcher 1973b; Ryan 1988b) and excess heating or freezing (Southward 1958). Verrucaria maura occurs far up on exposed shores in depressions or shaded nooks indicating a requirement for moisture. This was noted by European workers, including Fletcher (1973b), who nevertheless regards V. maura as one of the more drought-tolerant of the marine Verrucaria species. Fletcher (1973a) found that V. maura actually has no requirement for submergence in sea water at all. It can survive with Vol. 117 Average Average Extreme minimum (m) maximum (m) maximum (m) 5.67 7.00 8.00 6.50 6.50 7.00 6.00 6.57 9.00 6.00 8.00 8.00 7.33 N33 8.00 4.80 6.30 9.00 7.00 7.00 7.00 9.00 9.00 9.00 7.00 7.00 7.00 8.00 9.00 10.00 6.00 6.67 7.00 9.33 9.33 10.00 9.00 9.00 9.00 4.80 5.30 9.00 7.50 7.50 8.00 9.00 9.00 9.00 4.00 4.00 4.00 3.00 3.00 3.00 6.17 6.67 8.00 8.00 9.00 9.00 8.00 8.00 8.00 8.50 9.00 9.00 6.00 7.00 8.00 8.00 8.50 9.00 7.00 8.00 8.00 9.00 9.00 9.00 8.00 8.67 10.00 8.00 8.00 8.00 1.00 1.00 1.00 7.80 8.00 9.00 7.00 9.50 10.00 8.00 8.00 8.00 6.27 7.73 10.00 2.40 2.60 4.00 1.00 1.00 1.00 2.90 3.90 5.00 DIDS 4.50 7.00 2.47 UAL 10.00 1.00 1.50 2.00 1.00 1.67 3.00 325 4.00 7.00 1.00 1.00 1.00 7.20 7.60 8.00 only sea spray. Competition with algae and grazing animals (such as limpets and snails) is regarded to be a significant factor by some observers (Southward 1958; Ryan 1988b) and minimal by others (Fletcher 1973a). We did not see much evidence of grazing on the lichens of Gwaii Haanas. As noted by Fletcher (1980), the lichen bands can be characterized by life form as well as species. In the Littoral Fringe, all lichens are anatomically unstratified (i.e., without a differentiated cortex, algal layer and medulla) and almost all open by an ostiole rather than having a discoid apothecium. An exception in Gwaii Haanas is Herteliana alaskensis, a lichen with bia- torine apothecia found in the Lower Supralittoral Zone a 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 415 TABLE 6. Zones formed by lichens on maritime rocks in Gwaii Hanaas. The names of equivalent zones in O’ Clair et al. (1996) and Ryan (1988b) are also given. For species listed twice under “Characteristic lichens,” an asterisk (*) indicates the lichen’s zonal position according to its occurrence in the quadrat series (see Figure 6); an exclamation (!) indicates its zonal position according to its height above the Fucus-barnacle (FB) zone (see Figure 4). P to E = protected to exposed shores; VE = very exposed shores. Quad series m above Zone Type Quadrat FB zone Terrestrial (Halophilic) 9-10 2.3-3.6 O’Clair: upper salt spray Ryan: Zones 2A- 2B Upper Supralittoral Zone 8-9 2.1-2.6 O’Clair: mid salt spray Ryan: Zone 2C-2D Middle Supralittoral Zone 7-8 1.4-2.3 O’Clair: low salt spray Ryan: Zone 2D Lower Supralittoral Zone 5-7 0.8-1.5 O’ Clair: splash zone Ryan: Zone 2E Littoral Fringe, Upper 2-4 0.3-0.8 O’Clair : high intertidal (p.p.) Ryan: Zone 3-2E Littoral Fringe, Lower 1-2 -0.3-0.3 O’Clair: high intertidal (p.p.) Ryan: Zone 4A-3 occasionally in the company of black species of Ver- rucaria. Lichens with apothecia, also having somewhat stratified crustose thalli, first appear in the Lower Su- pralittoral Zone. Higher still, in the Middle and Upper Supralittoral Zones, the stratified foliose lichens appear. The biological reasons for this phenomenon are fairly easy to surmise. Lichens that are submerged frequently have no need for a loosely organized medulla with air Characteristic lichens PtoE VE Parmelia saxatilis Fuscidea thomsonii Fuscopannaria maritima Rhizocarpon geminatum! Lecanora muralis! Porina pacifica Caloplaca verruculifera* Lecanora straminea* Porpidea speirea! Rhizocarpon geminatum Lecanora muralis Spilonema revertens Coccotrema maratimum Placopsis lambii Ochrolechia subplicans Xanthoria candelaria! Porpidia thomsonii Aspicilia caesiocinerea Rhizocarpon “haidensis”! Caloplaca rosei! Porpidea speirea* Rhizocarpon geminatum* Lecidella stigmatea Xanthoria candelaria Caloplaca flavogranulosa Caloplaca rosei* Rhizocarpon “haidensis” * Lecanora muralis* Xanthoria candelaria* Adelolecia kolaensis! Spilonema revertens! Caloplaca verruculifera Herteliana alaskensis Lecanora straminea Herteliana alaskensis Caloplaca litoricola Arthonia phaeobaea Physcia caesia Collema fecundum Adelolecia kolaensis* Spilonema revertens* Caloplaca litoricola Arthonia phaeobaea Verrucaria maura V.epimaura Verrucaria maura Verrucaria epimaura Verrucaria erichsenii Verrucaria schofieldii Verrucaria amphibia Verrucaria erichseniti V. schofieldii Pyrenocollema halodytes Verrucaria striatula Verrucaria mucosa Verrucaria sandstedei Pyrenocollema halodytes Verrucaria striatula spaces for gas exchange, and their spores are better distributed by oozing out of an ostiole than being shot into the air as would occur from an apothecial disk. Stratified apothecial lichens are adapted for a basically terrestrial life style. The shift in relative position in the zonation by Spilonema revertens on Very Exposed (where it is Terrestrial) as compared to less exposed shorelines Vol. 117 Metres above Fucus/barnacles THE CANADIAN FIELD-NATURALIST 416 4.00 3.50 - 3.00 2.50 2.00 1.50 1.00 0.50 0.00 -0.50 2 2 O@ Sf & © oN DS 3 V © O © Ww @ Oo HS So © © @ oe & dS @ a > & > © © & / Oo & os os e) os ee © is & SES Oe” ss CLE SX o* SELES ES é es yy Sos eS LE COE EFC FEF BOE GF SEF OB EE ESE SF EEF EB E EP EE EES ES oF oF F EEF ESE @ as fs a” -® eo” SS @ 2S ee & S s Ra eS S oe @ SEOe PF oe & & 5 & Pes < Ss & & > we @ & os & eS PS eS SF FS SS SF PD GS FP OS SF SF So Pie Fe S ee & & P° QPS LY EF oF F : Oe 7 0 or 2 oo “<. RS ws Rs Fo oe ae es Fe & 3 & Fee ss ror & QP x eS RO is) Ko s e ¢ x ES eg Z S SF WANA RO w CN SC SS Se ie oe Ke ak es) 0 & ee PF ¥ of SS es Co we < ge x < « e € The thick line is the range of average values. The thin lines represent the total range (extreme maximum and FicureE 4. Zonation of maritime lichens on Protected to Exposed shores, ordered by metres above the Fucus-barnacle zone. minimum) values. 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 417 ecies etres above Fucus/barnacles mn Sp Metres above Fucus/barnacl “% Ge) % @ y ° A I) 7) rN on fo) Ni 00 °% ° ° r=) ° ° ° ° ° ro) Pe ro) ro) ro) } ro) rs) ro) } } FiGuRE 5. Zonation of maritime lichens on Very Exposed shores, ordered by metres above the Fucus-barnacle zone. The thick line is the range of average values. The thin lines represent the total range (extreme maximum and minimum) values. 418 THE CANADIAN FIELD-NATURALIST Vola? Species 4 Quadrant %, %e ke es y ° Nn h a oo S =a j=) S S beh ro) ros) ro) ) ) 8 S FiGcuRE 6. Zonation of maritime lichens on Protected to Very Exposed shores, ordered by relative quadrat position. The thick line is the range of average values. The thin lines represent the total range (extreme maximum and minimum) values. 2003 BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS 419 TABLE 7. The number of quadrats sampled in which lichen species are represented, according to percentage coverage category. Coverage category (%) Lichen species 1-5 6-10 11-20 Adelolecia kolaensis 4 | 2 Amandinea coniops 3 Arthonia phaeobaea 5 | | Aspicilia caesiocinerea | ] Caloplaca citrina Caloplaca flavogranulosa 1 Caloplaca litoricola 6 4 3 Caloplaca rosei 2 Caloplaca verruculifera 1 Candelariella sp. | Coccotrema maritimum 1 4 2 Collema fecundum Fuscidea thomsonii 3 Fuscopannaria maritima 1 Gyalecta jenensis 1 Herteliana alaskensis 7 4 3 Lecanora muralis 2 Lecanora straminea 1 Lecanora sp. #1 Lecanora sp. #2 Lecidella stigmatea 4 Ochrolechia subplicans 1 Opegrapha gyrocarpa Opegrapha sp. Parmelia saxatilis Physcia caesia 1 Placopsis lambii 2 1 Porina pacifica Porpidia carlottiana Porpidia contraponenda Porpidia speirea 4 Porpidia thomsonii 1 Pyrenocollema halodytes 10 3 2 — Sas a — i) BRNNRK eK — Rhizocarpon geminatum Rhizocarpon haidensis ined. Rhizocarpon hochstetteri s. str 1 Spilonema revertens 17 Verrucaria amphibia 2 Verrucaria degelii Verrucaria epimaura 2 Verrucaria erichsenti 1 Verrucaria maura 11 Verrucaria mucosa Verrucaria sandstedei Verrucaria schofieldii Z 2 2 Verrucaria striatula Verrucaria sp. #5 Xanthoria candelaria + i [id ) SN Oe aN — me (where it is Middle Supralittoral) may be due, in part, to the species’ sensitivity to excessive inundation by tidal splash. More likely, it is due to its inability to physically withstand the pounding of waves on highly exposed headland rocks. Spilonema is a microfruticose lichen and is attached to the rock relatively loosely compared to its crustose neighbours. A similar shift in the distribution of the fruticose Lichina pygmaea occurs on European shores (Fletcher 1973a). Spilo- 21-30 31-40 41-50 51-60 61-70 71-80 81-90 91-100 w See nNWe oo tS — iT) NCO nema revertens is not a particularly maritime lichen, however, being frequently found in inland areas (Brodo et al. 2001). The species diversity on protected and semiprotect- ed Gwaii Haanas shores is not significantly lower than that of more exposed shores. Chu et al. (2000) found the opposite on Hong Kong shores. They hypothesized that wave-exposed shores have more microhabitats to be exploited than do quiet bays. Our data do not sup- 420 port Chu’s observations, but our data suffer from un- even sampling in the various exposure categories and may not be as reliable. Light influences species composition especially on the more terrestrial shoreline habitats (Fletcher 1973a). For example, Caloplaca and Xanthoria species are almost entirely restricted to sunny rocks in Gwaii Haanas. Fletcher (1973a) found that most lichens in the Littoral Fringe and even the Lower Supralittoral Zone are largely indifferent to light conditions, but Ryan (1988b) reported that Verrucaria mucosa only occurs in shaded habitats. This species was found in both unshaded and shaded sites in Gwaii Haanas, although it was more abundant on shaded limestone. Since exposure to sun affects moisture availability and retention, some correlation of species distribution ac- cording to shade conditions is to be expected. Shaded shores are almost always also protected shores, so it is difficult to separate the two factors in explaining differences in lichen vegetation. Shoreline rocks are frequently subject to manuring by sea birds, increasing the nitrogen load (urea and ammonia) and raising the pH of the rock surface, and this affects the composition of the saxicolous lichen communities, especially in the salt spray zone. Orni- thocoprophilous lichens that prefer such habitats in- clude Lecanora straminea, Caloplaca verruculifera, Physcia caesia and Xanthoria candelaria, all present on the shores of Gwaii Haanas wherever the conditions are suitable. The relative position of the marine and maritime species in Gwaii Haanas agree closely with observa- tions made by others, in so far as there are species in common, e.g., Verrucaria mucosa, V. striatula, Pyreno- collema halodytes occur in the upper edges of the in- tertidal zone; V. maura appears higher and has a broad distribution; Xanthoria candelaria, Physcia caesia and Caloplaca verruculifera are associated with bird rocks; etc. (Eliasson 1965; Ryan 1988b; Sheard and Ferry 1967; Sheard 1968; Fletcher 1973a; Fletcher 1973b). The west coast endemics, and especially the mari- time lichens closely restricted to Haida Gwaii, com- prise 17 of the 67 species listed in the Appendix. It is useful to examine these species to see what their eco- logical and taxonomic equivalents would be elsewhere in the world. We will begin with ecological relation- ships. Caloplaca litoricola, a Haida Gwaii specialty, is in the Supralittoral Caloplaca zone as one would expect, but, since it is grey rather than yellow, it does not help to create an orange zone as other species of Caloplaca do in Europe or even eastern North America. On the other hand, the Pacific Northwest endemics C. rosei and C. inconspecta, and especially Caloplaca flavo- granulosa, are the ecological equivalents of the similar European C. marina (also in western North America from California to Vancouver Island, but not reaching Haida Gwaii; Arup 1992), which is part of the “orange zone” of European shores. Instead of the fruticose THE CANADIAN FIELD-NATURALIST Vol. 117 cyanophilic lichen, Lichina pygmaea, common in the British Isles (and rare on the American east coast), we have Spilonema revertens, a microfruticose cyano- philic lichen in Gwaii Haanas, although it tends to be higher in the transects than Lichina would appear. In- stead of species like the orange foliose lichen, Xan- thoria parietina (common in both Europe and along the east coast of North America) in the Middle to Upper Supralittoral Zone, Haida Gwaii has the white crustose lichen Coccotrema maritimum, resulting in a conspicuous white zone replacing the orange zone seen in Europe. We have no ecological equivalent for their maritime saxicolous fruticose lichen, Ramalina siliquosa, but instead of Fuscidea cyathoides, Haida Gwaii has the narrowly endemic F: thomsonii in the Terrestrial zone (Brodo and Wirth 1998). The distribution of geographically or ecologically separated, closely related taxa (vicariants) with respect to these endemics or near-endemics is especially in- teresting. The categories of relationships and the taxa involved are summarized below, in most cases, with references to discussions already published. High Mountain connections The following species either have close relatives in the arctic-alpine flora, or have populations/subspecies that are found only on maritime and alpine rocks, not in between. Caloplaca litoricola: C. exsecuta (Ny1.) Dalla Torre & Sarnth. (Brodo 1984) Ochrolechia subplicans: subsp. hultenii 1s on shore- line rocks; subsp. subplicans grows in the alpine zone from Haida Gwaii to Alaska (Brodo 1988). Porpidia carlottiana: populations on Haida Gwaii either on shore or in mountains; close to P. glauco- phaea (Korber) Hertel & Knoph in Hertel, an ocean- ic species (Gowan 1989). Southern Hemisphere connections These species are closely related to species common in the southern hemisphere, either in South America or Australasia. Coccotrema maritimum: C. cucurbitula common on trees in Chile, etc. as well as in other parts of the Paci- fic Basin, e.g., Japan (Brodo 1973) Collema fecundum: C. novozelandicum Degel., on calcareous non-maritime and maritime rocks and soil, South Island, New Zealand (Degelius 1974, 1979) Verrucaria epimaura: V. durietzii 1. M. Lamb, mari- time lichen from the Aukland Islands (Brodo and Santesson 1997) Kohlmeyera complicatula: the same or related spe- cies are On maritime rocks in western South America. (Brodo 1976) Related to widespread maritime species These species are most closely related to other mari- time lichens. Lecanora species number 1: L. actophila, L. heli- copsis This undescribed Lecanora with a thick, white, xanthone-containing thallus will be discussed in a pub- 2003 lication in preparation. It is clearly related to other species in the L. dispersa aggregate such as those listed above. L. poliophaea (undescribed subspecies): subsp. poliophaea. The Haida Gwaii populations of L. poliophaea differ in some respects from European populations and are probably deserve taxonomic recognition. This will be discussed in the publication in preparation mentioned above. Verrucaria. schofieldii: V. erichsenii, a widespread supralittoral species (Brodo and Santesson 1997). Related to widespread non-maritime species The following species have their closest relatives as Temperate or Boreal, non-maritime lichens. Fuscidea thomsonii: F. intercincta (Nyl.) Poelt, F: atlantica (H. Magn) P. James & Poelt, both are Euro- pean species found in regions with an oceanic climate (Brodo and Wirth 1998; Oberhollenzer and Wirth 1984). Fuscopannaria maritima: F. thiersii P. M. J@rg., a Pacific Northwest species on iron-rich moist rocks (Jérgensen 2000); and F: leucostictoides Ohlsson, a western American corticolous species (Jorgensen 1978). Porina pacifica: Porina chlorotica (Ach.) Miill. Arg., widespread northern species on shaded rock faces (Brodo 2004). Rhizocarpon maritimum ined: R. cinereovirens is a Temperate species on shaded and exposed rocks, B. C., Black Hills, Long Island, N.Y. (The species, which will be described in a forthcoming paper, is closely related to R. cinereovirens (Fryday 2002: 463). These phytogeographic relationships of the endemic or near-endemic maritime lichens of Haida Gwaii and their vicariant taxa suggest survival in the area during the Pleistocene glaciations. Relationships with montane species may indicate, as has been suggested (Clague 1989; Heusser 1989), that during the several glacial advances of the Pleistocene, some coastal areas and the highest mountain peaks of Haida Gwaii remained exposed. They would, therefore, have been available for colonization by lichens. It is possible that species that evolved at sea level from montane ancestors were highly restricted to coastal rocks and never were able to reinvade the intervening elevations although the highly exposed and harsh seacoast environment has much in common with alpine habitats and often shares the same or related species. It is probably significant that the closest relative of Coccotrema maritimum, a rock- dwelling species, is C. cucubitula, primarily a corti- colous species throughout its range. A glacial relic would, of course, have to survive on non-arboreal sub- strates. Schofield (1989) notes that all the Haida Gwaii endemics among the bryoflora occur on rock or soil. Many organisms are distributed around the Pacific Rim, or are disjunct from regions thousands of kilo- metres away. Plants and animals showing these pat- terns usually are related at the genus level or above, but these relationships show up at the species level in lichens and sometimes bryophytes (Schofield 1989). BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS The fact that the land mass of Haida Gwaii represents a crustal plate fragment (terrane) that originated off the coast of Peru and and started its northward migration approximately 230 million years ago (Sutherland- Brown and Yorath 1989) provides a possible way for the species to have been transported (assuming the spe- cies had evolved by that time), although long distance dispersal is another means. Similar patterns are known from bryophytes; e.g., the moss Polytrichadelphus lyellii Mitt, which is known from British Columbia to California. The other 20 species of the genus are from South America, New Zealand and Australia (Persson, H. 1968. The possible migration routes of the species with southern hemisphere affinities are discussed by Brodo (1973). Maritime lichens on Haida Gwaii that are disjunct with western Europe or even eastern North America probably reveal ancient, probably Tertiary, distribu- tion patterns that were at first continuous but became dissected by climatic and geological events over the past 10-20 million years leaving the Haida Gwaii populations clinging to their outposts in microclimat- ically suitable habitats. Equivalent patterns among the mosses are common (Schofield 1984). Conclusions Rock-dwelling marine lichens of Gwaii Haanas are distributed in zones much as they are elsewhere in the world, although some species found abundantly only in Haida Gwaii create a unique appearance of these zones, such as the white Coccotrema zone, as compared with other regions. The unusually large percentage of en- demic, near-endemic or disjunct species suggests that at least the headland rocks along the west coast were refugia during the last glacial maximum. Acknowledgments We thank Gwaii Haanas (Parks Canada) for fund- ing the summer 2000 fieldwork and Doug Burles and Lee Edenshaw for field assistance. Dr. Fenja Brodo made numerous helpful suggestions on early drafts and assisted with some quadrat studies when not execut- ing her own Gwaii Haanas inventory project. IMB, now retired from the Canadian Museum of Nature, thanks the Museum for appointing him as a Research Associate and providing him with space and facilities for his lichen research. We dedicate this paper to the memory of Bruce Ryan, who first studied the maritime zonation of lichens in the Pacific Northwest and did it so well. Documents Cited (marked * in text) Anonymous. 2002. Final Report—Offshore Oil and Gas Task Force. (http://www.em.gov.bc.ca/Oil& Gas/offshore/Off shoreOilGasReport/) Geiser, L. H., C. C. Derr, and K. L. Dillman. 1994. Air quality monitoring on the Tongass National Forest. Meth- ods and baselines using lichens. USDA-Forest Service, Alaska Region, Report R10-TB-46. 98 pages. 422 Harper, J. R., W. C. Austin, M. M. Morris, P. D. Reimer, and R. Reitmeier. 1994. Ecological Classification of Gwaii Haanas — Biophysical Inventory of Coastal Resources. Report prepared for Parks Canada, Calgary, Alberta by Coastal & Ocean Resources Ltd., Sidney, British Colum- bia. 115 pages. Howes, D., J. R. Harper, and E. Owens. 1994. B.C. physical shore-zone mapping system. British Columbia Resource Inventory Committee Publication 8. 71 pages. Literature Cited Arup, U. 1992. Caloplaca marina and C. rosei, two difficult species in North America. Bryologist 95: 148-160. Brodo, I. M. 1973. The lichen genus Coccotrema in North America. Bryologist 76: 260-270. Brodo, I. M. 1976. 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Kitching. 1996. The biology of rocky shores. Oxford University Press, Oxford. 240 pages. Nimis, P. L., C. Scheidegger, and P. A. Wolseley. Editors. 2002. Monitoring with lichens — lichen monitoring. NATO Science Series IV, Earth and Environmental Sciences 7. Kluwer Academic Publishers, Dordrecht, Netherlands. Noble, W. 1982. The lichens of the coastal Douglas-fir dry subzone of British Columbia. Ph.D. thesis, University of British Columbia, Vancouver. Oberhollenzer, H., and V. Wirth. 1984. Beitrige zur Revision der Flechtengattung Fuscidea. Beiheft zur Nova Hedwigia 79: 537-595. O’ Clair, R. M., S. C. Lindstrom, and I. M. Brodo. 1996. Southeast Alaska’s Rocky Shores. Seaweeds and Lichens. Plant Press, Auke Bay, Alaska. 149 pages. Persson, H. 1968. Bryophytes from the Aleutian Islands, Alaska, collected mainly by Hansford T. Shacklette. Svensk Botanisk Tidskrift 62: 369-387. 2003 Ramkaer, K. 1977. The influence of salinity on the estab- lishing phase of rocky shore lichens. Botanisk Tidskrift AZ W9=-123.. Ryan, B. D. 1988a. Marine and maritime lichens on serpen- tine rock on Fildago Island, Washington. Bryologist 91: 186-190. Ryan, B. D. 1988b. Zonation of lichens on a rocky seashore on Fildago Island, Washington. Bryologist 91: 167-180. Schofield, W. B. 1984. Bryogeography of the Pacific Coast of North America. Journal of the Hattori Botanical Labora- tory 55: 35-43. Schofield, W. B. 1989. Structure and affinities of the bryo- flora of the Queen Charlotte Islands. Pages 109-119 in The outer shores. Edited by G. G. E. Scudder and N. Gessler. Queen Charlotte Islands Museum Press, Skidegate, British Columbia. Searing, G. G., H. R. Frith, and D. Howes. 1995. B.C. biological shore-zone mapping system. British Columbia Resource Inventory Committee Publication 9: 1-46. Sheard, J. W. 1968. The zonation of lichens on three rocky shores of Inishowen, Co. Donegal. Proceedings of the Royal Irish Academy 66, Section B: 101-112, plate VI. Sheard, J. W., and B. W. Ferry. 1967. The lichen flora of the Isle of May. Transactions of the Botanical Society of Ediburgh 40: 268-282. BRODO AND SLOAN: LICHEN ZONATION ON COASTAL ROCKS Sloan, N. A., and P. M. Bartier. 2000. Living marine Jega- cy of Gwaii Haanas I: Marine plant baseline to 1999 and plant-related management issues. Parks Canada Technical Reports in Ecosystem Science 27: 1-104. Southward, A. J. 1958. The zonation of plants and animals on rocky sea shores. Biological Review 33: 137-177. Stephenson, T. A., and A. Stephenson. 1949. The universal features of zonation between tide marks on rocky coasts. Journal of Ecology 37: 289-305. Sutherland Brown, A., and C. J. Yorath. 1989. Geology and non-renewable resources of the Queen Charlotte Islands. Pages 3-26 in The outer shores. Edited by G. G. E. Scudder and N. Gessler. Queen Charlotte Islands Museum Press, Skidegate, British Columbia. Taylor, R. M. 1982. Marine flora and fauna of the north- eastern United States. Lichens (Ascomycetes) of the inter- tidal region. National Oceanic and Atmospheric Adminis- tration Technical Report, National Marine Fisheries Service Circular 446. Department of Commerce, Washington, D.C. Zacharias, M. A., and J. C. Roff. 2001. Explanations of pat- terns of intertidal diversity at regional scales. Journal of Biogeography 28: 1-13. Received 30 June 2003 Accepted 17 February 2004 424 THE CANADIAN FIELD-NATURALIST Volany Appendix: Lichens of Haida Gwaii Found on Maritime Rocks Note: Many of the lichens listed below are non-maritime lichens that, on occasion, are found growing in the maritime zone thereby demonstrating a tolerance for salt spray. Species not found in Gwaii Haanas National Park Reserve are marked with an asterisk (*). Those new to British Columbia are in boldface. Voucher specimens of all these records can be found in the Canadian Museum of Nature (CANL). Adelolecia koalensis (Ny1.) Hertel & Rambold (Syn. Lecidea conferenda Nyl.) This was first reported for British Col- umbia in Brodo (1995). *Amandinea coniops (Wahlenb. in Ach.) M. Choisy ex Scheid. & H. Mayrh. Arthonia phaeobaea (Norman) Norman Aspicilia caesiocinerea (Nyl. ex Malbr.) Arn. *A spicilia supertegens Arn. This is not normally a maritime species. The single voucher specimen requires further study. *Bacidina inundata (Fr.) Vézda. This species was also found in streams in Haida Gwaii. *Caloplaca chlorina (Flotow) H. Olivier. The voucher speci- men was verified by both T. Tonsberg and C. M. Wetmore. Caloplaca citrina (Hoffm.) Th. Fr. Caloplaca flavogranulosa Arup *Caloplaca inconspecta Arup Caloplaca litoricola Brodo Caloplaca rosei Hasse Caloplaca verruculifera (Vainio) Zahlbr. *Catillaria chalybeia (Borrer) Massal. *Cliostomum griffithii (Sm.) Coppins Coccotrema maritimum Brodo Collema fecundum Degel. *Ephebe lanata (L.) Vainio *Fuscidea mollis (Wahlenb.) V. Wirth & Vézda. The voucher specimen is not typical of the species, so this record is questionable. Fuscidea thomsonii Brodo & V. Wirth Fuscopannaria maritima (P. M. Jorg.) P. M. Jorg. Gyalecta jenensis (Batsch) Zahlbr. Herteliana alaskensis (Nyl.) Zahlbr. *Kohlmeyera complicatula (Nyl.) Schatz Lecanora muralis (Schreber) Rabenh. *Lecanora poliophaea (Wahlenb.in Ach.) Ach. s. lat. Lecanora straminea Wahlenb. ex Ach. *Lecanora tenera (Nyl.) Cromb. Lecanora sp. number 1. This is a xanthone-containing. mari- time, saxicolous member of the Lecanora dispersa group and almost certainly is undescribed. Lecanora sp. number 5. This lichen is similar in many respects with L. contractula Ny\l. *Lecidella scabra (Taylor) Hertel & Leuckert Lecidella stigmatea (Ach.) Hertel & Leuckert f. stigmatea Ochrolechia subplicans (Ny1.) Brodo subsp. hultenii (Erichs.) Brodo Opegrapha gyrocarpa Flotow Opegrapha sp. This is a sorediate species containing roccel- lic acid, with 3-septate spores and lirelliform ascomata unlike those of O. gyrocarpa. It may be undescribed. Parmelia saxatilis (L.) Ach. Physcia caesia (Hoffm.) Fiirnr. Placopsis lambii Hertel & Wirth. There is a single record of P. gelida (L.) Lindsay s. str. on a rock in a tidal marsh, but otherwise the species is only known on Haida Gwaii from inland localities, usually on roadsides and fresh rock exposures. *Polychidium muscicola (Sw.) S. F. Gray Porina pacifica Brodo. This species, which superficially resembles P. chlorotica (Ach.) Miill. Arg., was only recent- ly described (Brodo 2004) Porpidia carlottiana Gowan Porpidia contraponenda (Arnold) Knoph & Hertel Porpidia speirea (Ach.) Kremp. Porpidia thomsonii Gowan *Punctelia stictica (Duby) Krog Pyrenocollema halodytes (Nyl.) R. C. Harris s. lat. We have in- cluded specimens with sunken perithecia as well as those with sessile perithecia under this name if the perithecia were under 0.3 mm in diameter. The much larger P. sub- litorale (Leighton) R. C. Harris ex Fletcher in Coppins, P. James & D. Hawksw. has not yet been found in Haida Gwail. Rhizocarpon geminatum Korber Rhizocarpon “haidensis” Brodo, ined. This species is closely related to R. cinereovirens (Miill. Arg.) Vainio (Fryday 2002). Its description is being prepared for publication. *Rhizocarpon hensseniae Brodo Rhizocarpon hochstetteri (K6rber) Vainio s. str. A detailed discussion of this species is provided by Fryday (2002). *Rinodina gennarii Bag}. Spilonema revertens Ny. *Tephromela atra (Hudson) Hafellner in Kalb *Tylothallia biformigera P. James & R. Kilias in R. Kilias Verrucaria amphibia Clemente Verrucaria degelii R. Sant. Verrucaria epimaura Brodo Verrucaria erichsenii Zsch. *Verrucaria halizoa Leighton Verrucaria maura Wahlenb. in Ach. Verrucaria mucosa Wahlenb. in Ach. Verrucaria sandstedei de Lesd. This species, like V. striatula listed below, was reported from coastal rocks in Fildago Island, Washington, by Ryan (1988a, b). Verrucaria schofieldii Brodo *Verrucaria Silicicola Fink Verrucaria striatula Wahlenb. See comment under V. sand- stedel. Verrucaria sp. number 5 (dry rock at edge of beach) Xanthoria candelaria (L.) Th. Fr. Predaceous Water Beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Collected Along the Horton and Thelon Rivers in the Arctic Central Barrens of Canada HELENA V. SHAVERDO! and DONNA J. GIBERSON? ‘Department of Entomology, 213 Animal Science/Entomology Building, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada "Department of Biology, University of Prince Edward Island, Charlottetown, Prince Edward Island C1A 4P3 Canada Shaverdo, Helena V., and Donna J. Giberson. Predaceous water beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) col- lected along the Horton and Thelon rivers in the Arctic Central Barrens of Canada. Canadian Field-Naturalist 118(3): 425-433. Predaceous water beetles were collected during expeditions along two northern Canadian rivers during 2000 and 2002. Twelve species of Dytiscidae (including 11 named species and one additional genus identified from a larva) and one species of Gyrinidae are recorded from 20 sites along the Horton and Thelon rivers in the Central Barrens area of the Canadian Arctic. These records represent an extension of the distributions of four species to the northeast in the Northwest Territories (NWT), and two species to the northwest in Nunavut (NU). Oreodytes sanmarkii is reported for NWT and for the mainland of NU for the first time. //ybius erichsoni, Hydroporus geniculatus, and Gyrinus opacus are reported for NU for the first time. Five species were recorded for the first time from the Southern Arctic ecozone, and one from the Taiga Shield ecozone. The majority of specimens were collected in habitats that were consistent with those previously known for each species. Key Words: Insecta, Coleoptera, Adephaga, Dytiscidae, Gyrinidae, predaceous water beetles, faunistics, river, Northwest Territories, Nunavut, Arctic Central Barrens. The Central Barrens area of the Canadian Arctic is a vast and very sparsely populated area dominated by arctic tundra, scattered clumps of taiga vegetation, large lakes, and rivers. The inaccessibility of the region has meant that the area has been poorly collected for all insect groups (Currie et al. 2000), including preda- ceous water beetles. Only three surveys have been con- ducted on aquatic Adephaga in the Yukon, Alaska, and the eastern Canadian Arctic (Alarie and Maire 1991; Larson 1991, 1997), though data on this group of Coleoptera in arctic and subarctic regions can also be found in Larson and Roughley (1991), Roughley (1991), Anderson (1997), and Larson et al. (2000). Recently a series of expeditions were mounted to ex- plore the aquatic insect fauna along large river systems on the treeless barrens of the mainland Canadian Arc- tic zone (Currie et al. 2000, 2002). The objective of this paper is to report on the predaceous water beetles collected from habitats in and near the Horton River Northwest Territories (NWT) and the Thelon River, NWT and Nunavut (NU), and present faunistic and habitat data from the collections. Methods Two expeditions were made by DJG and colleagues from various institutions to northern Canadian rivers during 2000 and 2002 (Figure 1). Expedition mem- bers flew in to upstream sections of the Horton and Thelon rivers in NWT and travelled downriver by canoe, stopping frequently to sample habitats along the linear corridor of each river. Aquatic sampling was carried out using a D-frame kick net with a 200 um mesh. In flowing water habitats, the net was placed into the stream and the substrate disturbed by kicking or by moving rocks by hand, and the dislodged insects were carried into the net by the current. In areas with little or no flow, the substrate was disturbed as above, and then the net was drawn through the water by hand to catch any dislodged insects. Insects were sorted alive from the substrate materials on site, and preserved in 80% ethanol. Predaceous water beetles were not specifically targeted in the study, but those that were collected (61 adults and 6 larvae) were exam- ined by HVS. Collection records were compared to the maps and distribution table in Larson et al. (2000) to determine whether any of them represented range ex- tensions among the provinces and ecozones. The doc- ument A National Ecological Framework for Canada (Ecological Stratification Working Group 1995) was consulted for the classification of Canada’s ecological regions. Collecting Permits were obtained from the Aurora Research Institute (Inuvik, NWT) and the Nunavut Research Institute (Iqaluit, NU) to carry out the research. Voucher specimens have been deposited in the collections of the Wallace Museum (University of Manitoba) and the Royal Ontario Museum, and in the private collection of D. J. G. Study Site and Sampling Regime The Horton River begins west of Horton Lake, which is located north of Great Bear Lake in the Northwest Terriories of Canada (Figure 1). The river, which has 425 426 THE CANADIAN FIELD-NATURALIST Vol. 118 Horton River 68°. 127° Tanptalase 123° 121° Thelon River Thelon Game" * Sanctuary \ ynaeunn _souloyuse| ISOMYYON 105°W 100°W FiGureE 1. Location of study area (A), including the Horton River (B) and Thelon River (C) showing sample sites. The fine dashed line indicates the watershed boundary upstream of the lowest sampled site for both watersheds. The grey border on the Thelon Watershed map indicates the boundaries of the Thelon Game Sanctuary. Site co-ordinates are given in Table 1. a drainage basin area of approximately 26 680 km? and a total length of = 800 km, runs entirely in the Southern Arctic Ecozone. Expedition members col- lected over a 700 km stretch of the river from Horton Lake to the Arctic Ocean. The river flows over lime- stone-dominated substrates in its upper reaches, where it is characterized by relatively high pH (>8.0) and spe- cific conductance (>150 uS/cm). In its lower reaches, the river flows through the Smoking Hills, a region containing lignite and magnesium deposits which spon- taneously ignite when exposed to air (for example, during bank slumping during summer). These result in “smoke holes” as smoke pours from places where the coal is burning underground, and the water drain- ing these sites is profoundly affected by the geology. Tributaries and ponds in this region were highly acidic (some with pH values < 2.5), and acid runoff in this area resulted in a decline in river pH to approximately neutral values. The river is clear and swift-flowing for most of its length, but becomes muddy and meander- ing in its lower portion. Sampling was carried out in 37 sites on or near the river between 17 July and 8 Au- gust 2000. Predaceous water beetles were collected from 5 of the 37 sites along the river, corresponding to sampling sites 8, 11, 14, 18, and 23 of the larger Hor- ton River survey; all were located in the middle sec- tion, which was clear and swift-moving (Figures 1b, 2; see Table 1 for descriptions of each site). The Thelon River begins near Lynx Lake, to the east of Great Slave Lake (Figure 1). The river flows = 900 km from Lynx Lake to Baker Lake (which exits via Chesterfield Inlet to northern Hudson Bay to the east) and drains a total area of = 240 000 km?. The upper portions of the Thelon River are in the Taiga Shield ecozone, then the river flows into the Southern Arctic ecozone. Expedition members flew into the junction of the Hanbury and Thelon rivers, and col- lected along a 300 km stretch of the Thelon River located within the bounds of the Thelon Game Sanc- tuary (Figure Ic). The part of the river covered in this survey (i.e., upstream of Beverly Lake, NU) drains an area of 71 470 km. The river corridor upstream of the study area consists of large numbers of lakes, water- falls, and rapids, but in the study area itself, the river meanders over an ancient glacial lake bottom. At this point, the river is wide and slow moving and dominat- ed by sandy or sand/silt-embedded cobble substrates. Surrounding relief is very low except in a few spots, 2004 Horton Site 8 one SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC Horton Site 11 FIGURE 2. Study sites along the Horton River where predaceous water beetles were collected, July/August 2000 (compare to Figure 1B for site locations). where low hills, boulders, and more rapid flow can be found. Most of the river flows through open tundra, but a small stretch in the upper third of the study area (= 65 km) is located in a remnant spruce and larch forest known as the “Thelon Oasis”. The pH and spe- cific conductance values were quite low, especially in comparison with the Horton River, with pH values gen- erally around 6.0, and specific conductance values = 10 uS/cm for main river sites. pH and conductivity values for tributaries and tundra pools were more variable, however. Sampling was carried out at 34 sites on or near the river between between 29 June and 11 July 2002. Predaceous water beetles were collected from 15 of the 34 sites, representing a combination of river and pool sites corresponding to sites 1, 3, 4, Gy TS OMI lor slSeil9) 22-26::27.) 50and 32-oftthe larger Thelon River survey (Figures Ic, 3; see Table 1 for descriptions of each site). Results and Discussion Oreodytes laevis (Kirby) and O. sanmarkii (Sahlberg) were the two most common species found in our north- em rivers survey, and were also the only species found in both rivers (Table 1). Oreodytes laevis accounted for 26 of the 61 adults collected (> 40%) and was found in eight of the 20 sites where water beetles were found. 428 THE CANADIAN FIELD-NATURALIST Vol. 118 _ Thelon Site 7 FiGuRE 3a. Study sites along the upper reaches of the Thelon River where predaceous water beetles were collected, July/ August 2000 (compare to Figure 1C for site locations). 2004 SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 429 Thelon Site 18 i Thelon Site 19 yA Thelon Site 30 FIGURE 3b. Study sites along the lower reaches of the Thelon River where predaceous water beetles were collected, July/August 2000 (compare to Figure Ic for site locations). 430 TABLE |. Distribution and abundance of predaceous water beetles collected along the Horton and Thelon Rivers. All species were collected in the main branch of the rivers except where specifically indicated. All species were in the family Dytisci- THE CANADIAN FIELD-NATURALIST dae except Gyrinis opacus which was in the family Gyrinidae. Site Coordinates / Date Site Details Horton River, NWT 8 67°59°24.5"N 123°13°55.5°W 20 July 2000 68°09°29"N 123°22°44.9"W 22 July 2000 68°23°31.5°N 123°38°03.4°W 23 July 2000 68 39°3.18°N 124°08°53.0"W 25 July 2000 68°44’ 19.9"N 124°59°0.8°W 28 July 2000 Thelon River, NWT 1 13 16 18 63 38°04.8"N 104°32°18.6"W 29 June 2002 63 42°04.4”"N 104°25’26.6" W 30 June 2002 63 48’22°N 104°18’21.1"W 30 June 2002 63°49’58.4"N 104°06°20.9"W 1 July 2002 63°55°02.7"N 103°56’59.5°W 1 July 2002 64°02’ 13.2”N 103°51°41.8°W 2 July 2002 64°17°01.8"°N 103°33°22.4”"W 3 July 2002 64°19" 18.6”N 103°19715.7°W 4 July 2002 64°17°56.2”N 102°43°20.6"W 5 July 2002 cobble dominated riffle and shore section, where river slowed and flowed through a broad valley; pH: 8.3; conductivity: 150 wS/cm; temp.: 15°C slow, weedy shoreline area along the main channel of the river (fill in from logbook); pH: 8.2; conductivity: 180 uS/cm; temp.: 10.5°C backwater zone and a riffle, from a portion of the river in a narrow valley surrounded by high limestone cliffs; pH: 8.2; conductivity: 180 uS/cm; temp.: 12°C the river is a series of very shallow flats and riffles; temp.: 12°C canyon/white-water area of the river, deep bedrock pools and fast riffles; at camp spot no obvious riffles though water moving very fast; sampled from shore; temp.: 17°C relatively slow and wave washed section near shore, in silt/sand embedded cobble; pH: 6.2; conductivity: 10 uS/cm; temp.: 11°C rocky shore with fairly strong current, and in backwater pools behind large boulders; pH: 6.2; conductivity: 10 wS/cm; temp.: 11°C slow section near shore, with cobble substrate embedded in sand and silt; pH: 6.2; conductivity: 10 uS/cm; temp.: 10.5°C submerged grasses in slow current near shore, 10 m upstream of a tributary entering from the south; pH: 6.2; conductivity: 40 uS/cm; temp.: 14°C cobble and silt substrate near shore where main river narrowed and became swift flowing, even near shore; pH: 6.2; conductivity: 10 uS/cm; temp.: 11°C at Hornby Point; well embedded cobble in fast, shallow riffle near shore; pH: 6.1; conductivity: 10 uS/cm; temp.: 12°C cobble substrate in fast water in main channel, plus slower water near shore; pH: 6.3; conductivity: 30 uS/cm; temp.: 13°C unnamed tributary entering Thelon River from forest to the north; riffle with large clean cobble substrate; pH: 7.6; conductivity: 40 uS/cm; temp.: 16°C river wide and lake-like; wave-washed area, in cobble substrate well embedded in sand; pH: 6.3; conductivity: 10 uS/cm; temp.: 13.5°C Species Oreodytes laevis (Kirby) Oreodytes sanmarkii Sahlberg Oreodytes sanmarkii Oreodytes laevis Oreodytes sanmarkii Oreodytes sanmarkii Oreodytes sanmarkii Oreodytes laevis Oreodytes sanmarkii Oreodytes laevis Oreodytes laevis Hygrotus novemlineatus (Stephens) Oreodytes laevis Stictotarsus griseostriatus (DeGeer) Oreodytes laevis Stictotarsus griseostriatus Oreodytes sanmarkii Oreodytes laevis Vol. 118 Specimens | 1° ie) lo 2c¢c lo ilxe} 2c¢c oil © 2004 SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 43] TABLE | (continued). Distribution and abundance of predaceous water beetles collected along the Horton and Thelon Rivers. All species were collected in the main branch of the rivers except where specifically indicated. All species were in the fami- ly Dytiscidae except Gyrinis opacus which was in the family Gyrinidae. Site 19 64°10°46.1°N 102°36°58.1"W 6 July 2002 23 64°11°05.4"N 102°19°06.9° W 7 July 2002 Thelon River, NU 26 64°19713.2”°N 101°50’11.1°W 8 July 2002 27 64°19’ 13.2”N 101°50°11.1"W 8 July 2002 30 64°26°15°N 101°43°55.1"W 9 July 2002 64°32°25.2”N 101°24°48.9°W 10 July 2002 Coordinates / Date Site Details small brown-water tundra pools above the river on north, pools draining to each other and seeping to the river; overhanging vegetation, undercut banks, stony bottom; pH: 6.0; conductivity: 20 uS/cm; temp.: 14°C unnamed tributary seep entering the river from the north and small brown-water tundra pools surrounded by willows, shorelines with sphagnum and grass; pH: 5.7; conductivity: 50 wS/cm; temp.: 9°C unnamed tributary entering the river from the north; cobble substrate in riffle area of the stream; pH: 6.5; conductivity: 10 uS/cm; temp.: 11°C small brown-water tundra pond beside site 26; much submerged and decaying vegetation (grass); pH: 6.0; conductivity: 10 uS/cm; temp.: 15°C large tundra pond on southern bank above the river, with overhanging willows and soft bottom substrates; pH: 6.0; conductivity: 10 uS/cm; temp.: 12°C unnamed small tributary entering the river from the south - small pools and seeps between the pools; overhanging vegetation; pH: 6.2; conductivity: 20 wS/cm; temp.: 14°C Species Specimens Hydroporus appalachius 2d¢0 Sherman DIRE) Agabus confines 499 (Gyllenhal) il bake Ilybius erichsoni 1E9 (Gemminger and Harold) Agabus sp. DES Oreodytes sanmarkii 1° Hydroporus geniculatus lo Thomson Ilybius erichsoni lo 2IE3e Stictotarsus griseostriatus lo Agabus arcticus ls (Paykull) 6 29 Rhantus sp. WLR Gyrinus opacus Sahlberg le Hydroporus morio 3.99 Aubé Agabus thomsoni IN Gr Ale. (Sahlberg) Note: * L3 — larva of third instar; L2 — larva of second instar. Eleven specimens of Oreodytes sanmarkii were also found at eight sites. Stictotarsus griseostriatus (De- Geer) specimens were collected in three of the sites, but there was only a single specimen in each collection. The remaining species were all restricted to a single site each. Although several specimens of individual species were often found, for example, the sites with Agabus confinis (Gyllenhal) and A. arcticus (Paykull) (Table 1). These collections are range extensions among the provinces and ecozones of Canada for most of the species collected, and represent a “filling in of gaps” in the distributions of many northern species. Because of logistic difficulties in sampling these interior regions of the arctic barrens, many previously reported species distributions have appeared disjunctive, or restricted to coastal areas (e.g., Larson et al. 2000). For example, the known ranges of Hydroporus appalachius Sher- man, Oreodytes laevis, and Agabus confinis are extend- ed to the northeast within the Northwest Territories, and those of Agabus arcticus and A. thomsoni Salh- berg are extended to the northwest within Nunavut. Oreodytes laevis and Hydroporus appalachius are widespread in the boreal low arctic zone of the Nearc- tic, so these records represent an extension into the Tundra zone. With respect to ecozone distribution, O. laevis, H. appalachius, and A. confinis are recorded for the first time from the Southern Arctic ecozone, and A. confinis is now known from all ecozones of Canada except the prairies. [/ybius erichsonii (Gem- minger and Harold) is reported for the first time for Nunavut, and its known range is extended to the north- east within the Northwest Territories, and into the Southern Arctic Ecozone. Ilybius erichsonii has now been reported from all the ecozones of Canada. Hydro- porus geniculatus Thomson is a Holarctic species which occurs in the boreal and alpine zones in north- western North America. In Canada, it is distributed throughout most of British Columbia and western Alberta, and north into the Yukon Territory and Alaska (Larson et al. 2000), and is reported for the first time in this study in Nunavut and in the Southern Arctic ecozone. The present record is an extension of its known 432 distribution far to the northeast, and therefore the range for this species appears disjunctive. However, H. geniculatus belongs to the poorly studied nigellus- group, and revision of the group is needed to clarify its taxonomic position and distribution. We report the presence of Oreodytes sanmarkii in the Northwest Territories and in the Taiga Shield ecozone for the first time, and the collections along the Thelon River also represent the first record of the species on the Nunavut mainland. This Holarctic species was previously known in Canada from a few sites in northern Yukon and a site on Southampton Island (NU) (Larson et al. 2000). Hygrotus novemlineatus (Stephens), Hydroporus morio (Aubé), and Stictotarsus griseostriatus have been reported in the Canadian Barrens to the east and west of the sampled areas, but these collections indicate that their distributions are more continuous across the north than previous distribution maps might indicate. Gyrinus opacus (Gyrinidae) is reported for Nunavut for the first time, and collections on the Thelon River may represent the most northeasterly record for the species (Oygur and Wolfe 1991). However, this record is in need of confirmation since it is based on study of a single female specimen. Most of the species were collected in habitats that matched those that were previously reported for each species (e.g., Larson et al. 2000). However, this study has produced some new and additional data on habi- tat distributions for several species. Oreodytes laevis and O. sanmarkii have been reported in rivers, streams, and along exposed shorelines of cold lakes with little or no submerged vegetation, and with mineral substrates. These two species dominated in these types of habi- tat along the Thelon and Horton rivers. According to Alarie and Maire (1991; as H. hudsonicus) and Larson et al. (2000), Hygrotus novemlineatus is circumpolar in the low arctic region and has been collected in Canada (near Churchill Manitoba, and in Quebec) in brackish splash pools (rock-pools) along the coast. However, in our survey it was collected among dense submerged grass in slow water current along the shore of the Thelon River. The species may have relatively wide habitat preferences, since Nilsson and Holmen (1995) reported it in Fennoscandia from similar habitats to the Thelon site, as well as from sandy oligotrophic lake margins and sparsely vegetated ponds. In the southern parts of its range, Hydroporus appalachius is usually found in habitats where there is some water movement, for example in streams, springs, and the margins of small lakes (Larson et al. 2000). In Labrador, the species was collected from a protected shoreline of a large lake (Larson et al. 2000), and it has been found in inland pools in the Quebec subarctic region (Alar- ie and Maire 1991). The habitat in the more northern parts of its range has not previously been described, and in this survey, it was found in small tundra pools that drained into each other before spilling into the Thelon River. THE CANADIAN FIELD-NATURALIST Vol. 118 Acknowledgments We are grateful to Rob Roughley and Lisa Purcell for their advice and comments on the manuscript. Financial support for the collections was provided by an NSERC operating grant, a UPEI Senate Research Grant, and a Nature Discovery Grant (National Muse- ums of Canada) to D. J. Giberson. Lisa Purcell helped to collect the specimens. Support for the study of the material was provided by NSERC through a 2001 NATO Science Fellowship to H. V. Shaverdo and an NSERC operating grant to Rob Roughley. Voucher specimens have been deposited in the J. B. Wallace Museum of the University of Manitoba, the Royal Ontario Museum, and in the private collection of H. V. S. Collecting permits were provided by the terri- torial governments through the Aurora Research Insti- tute in the Northwest Territories and the Nunavut Research Institute in Nunavut. Literature Cited Alarie, Y., and A. Maire. 1991. The dytiscid fauna of Québec Subarctic (Coleoptera: Dytiscidae). Coleopterists Bulletin 45: 350-357. Anderson, R. S. 1997. An Overview of the Beetles (Coleop- tera) of the Yukon. Pages 405-444 in Insects of the Yukon. Edited by H.V. Danks and J.A. Downes. Biological Sur- vey of Canada. Monograph series (2), Biological Survey of Canada (Terrestrial Arthropods). Ottawa. Ontario. Currie, D. C., D. J. Giberson, and B. V. Brown. 2000. Insects of Keewatin and Mackenzie. Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) 19: 48-51. Currie, D. C., D. J. Giberson, and B. V. Brown. 2002. Insect biodiversity in the Thelon Wildlife Sanctuary. Newsletter of the Biological Survey of Canada (Terrestrial Arthro- pods) 21: 59-64. Ecological Stratification Working Group. 1995. A National Ecological Framework for Canada. Agriculture and Agri- Food Canada, Research Branch, Centre for Land and Bio- logical Resources Research, and Environment Canada, State of the Environment Directorate, Ecozone Analysis Branch, Ottawa/Hull. Report and national map at 1:7 500 000 scale. Larson, D. J. 1991. A new species of Potamonectes Zim- mermann (Coleoptera: Dytiscidae) from Labrador. Coleop- terists Bulletin 45: 280-284. Larson, D. J. 1997. Dytiscid Water Beetles (Coleoptera: Dytiscidae) of the Yukon. Pages 491-522 in Insects of the Yukon. Edited by H. V. Danks and J. A. Downes. Biological Survey of Canada. Monograph series (2), Biological Sur- vey of Canada (Terrestrial Arthropods). Ottawa. Ontario. Larson, D. J., and R. E. Roughley. 1991. Family Dytiscidae. Pages 62-72 in Checklist of beetles of Canada and Alaska. Edited by Y. Bousquet. Agriculture Canada Publication 1861/E. Larson, D. J., Y. Alarie, and R. E. Roughley. 2000. Preda- ceous Diving Beetles (Coleoptera: Dytiscidae) of the Nearc- tic Region, with emphasis on the fauna of Canada and Alaska. NRC Research Press, Ottawa, Ontario, Canada. 982 pages. Nilsson, A. N., and M. Holmen. 1995. The aquatic Adephaga (Coleoptera) of Fennoscandia and Denmark. II. Dytiscidae. Fauna Entomologica Scandinavica 32: 1-192. 2004 SHAVERDO and GIBERSON: PREDACEOUS WATER BEETLES IN ARCTIC 433 Oygur, S., and G. W. Wolfe. 1991. Classification, distribution, Roughley, R. EK. 1991. Family Gyrinidae. Pages 72-73 in and phylogeny of North American (north of Mexico) spe- Checklist of beetles of Canada and Alaska. Edited by Y. cies of Gyrinus Miiller (Coleoptera: Gyrinidae). Bulletin Bousquet. Agriculture Canada Publication 1861/E. of the American Museum of Natural History 207: 1-97. Received 2 June 2003 Accepted 14 July 2004 Notes Collapsing Burrow Causes Death of a Eurasian Beaver, Castor fiber Liat R. THOMSEN!, FIONA SHARPE2, and FRANK ROSELL? ' Department of Zoology, University of Aarhus, Denmark 2 School of Biology, University of St Andrews, Fife, Scotland 3 Faculty of Arts and Sciences, Department of Environmental and Health Studies, Telemark University College, N-3800 Bo i Telemark, Norway Thomsen, Liat R., Fiona Sharpe, and Frank Rosell. 2004. Collapsing burrow causes death of a Eurasian Beaver, Castor fiber. Canadian Field-Naturalist 118(3): 434-435. The death of a Eurasian Beaver Castor fiber caused by a collapsing burrow in southeast Norway is reported. Two days of heavy rainfall had presumably caused the burrow to collapse, suffocating the animal. Key Words: Beaver, Castor fiber, collapsing burrow, southeast Norway. Several causes of death have been reported for Eurasian Beavers (Castor fiber) (Rosell et al. 1996; Nolet et al. 1997), but to our knowledge this is the first report of a beaver killed due to a collapsing bur- row. The beaver was an adult solitary male (21.5 kg and 5 years old; age was determined by examining annual cementum and dentine layers of the first molar (van Nostrand and Stephenson 1964)). His mate died two months earlier, probably due to old age (18 years old). The animal was one of several radio tagged beavers that were followed during a field study in spring and summer 2000 in Telemark County, south- east Norway (59°25'N, 09°03'E) (see Campbell 2000). On the evening of 14 July 2000 it was noted that the pulse interval of the radio signal had increased. The signal is inversely related to the body temperature of the animal (Alterra 1999), thus a higher pulse interval indicated that the beaver had died. The dead beaver was located in the main part of a partly collapsed bur- row (assessed to be relatively new), which was exca- vated four days later in order to find and retrieve the carcass (Figure 1). Beavers dig burrows where banks are sufficiently high (Wilsson 1971; Zurowski 1992). A burrow usu- ally consists of a single living chamber, a water basin, and a tunnel with exit below the water level (Wilsson 1971). The beaver was lying in the chamber (1 m long and 70 cm wide) of the collapsed burrow, facing to- wards the exit. The chamber was situated 210 cm from the water’s edge, 80 cm below the surface of the riverbank, and 60 cm above the current water level. Apparently the ceiling of the chamber had collapsed on top of the beaver causing the death of the animal, presumably by suffocation. There was loose sand along the flanks of and underneath the body of the dead beaver. The last 10 cm of the beaver’s tail was bent downwards into the sand and the right hind foot was crouched as if the beaver had been attempting to dig with it. The beaver had no external injuries and seemed to be in good condition. of a partly collapsed burrow. The burrow was dug in sandy soil. In the week prior to the discovery of the beaver there had been two days with heavy rainfall in the area with 30.8 mm and 24.6 mm of rain, respectively (data from the Norwegian Meteorological Institute). We conclude that the combination of heavy rainfall and a sandy soil had caused the burrow to collapse, therefore causing the death of the beaver. How preva- lent this cause of death is in beavers is unknown. How- ever, we expect that more field studies using radio telemetry, in areas where beavers dig burrows, could clarify this issue. 434 2004 Acknowledgments We thank Frode Bergan for help with excavating the beaver and Inger Hanssen-Bauer for providing the weather data. The study was financially supported by Telemark University College. The experiments comply with the current Norwegian law, the country in which they were performed. Literature Cited Alterra. 1999. Alterra (IBN/DLO) P.O. Box 23. 6700 AA Wageningen. Netherlands. Campbell, R. D. 2000. Territoriality in the European Beaver, Castor fiber. MSc thesis, School of Biological Sciences, University of East Anglia, Norwich, England. Nolet, B. A., S. Broekhuizen, G. M. Dorrestein, and K. M. Rienks. 1997. Infectious diseases as main factors of NOTES 435 mortality to beavers Castor fiber after translocation to the Netherlands. Journal of Zoology, London 241: 35-42. Rosell, F., H. Parker, and N. B. Kile. 1996. Causes of mor- tality in beaver (Castor fiber & canadensis). Fauna 49: 34-46 [In Norwegian with English summary]. van Nostrand, F. C., and A. B. Stephenson. 1964. Age deter- mination for beavers by tooth development. Journal of Wildlife Management 28: 430-434. Wilsson, L. 1971. Observations and experiments on the ethol- ogy of the European beaver (Castor fiber L.). Viltrevy 8: _ E266: Zurowski, W. 1992. Building activity of beavers. Acta Theriologica 37: 403-411. Received 26 November 2002 Accepted | November 2004 Frequency of Tail Breakage of the Northern Watersnake, Nerodia sipedon sipedon KENNETH D. BOWEN! :2 'Department of Biology, Central Michigan University, Mt. Pleasant, MI 48859 USA Present Address: 709 Ringold Street, Boone, Iowa 50036 USA Bowen, Kenneth D. 2004. Frequency of tail breakage in the Northern Watersnake, Nerodia sipedon sipedon. Canadian Field-Naturalist 118(3): 435-437. I noted the frequency of broken tails of Northern Watersnakes, Nerodia sipedon sipedon, in the Beaver Archipelago of North- eastern Lake Michigan. Overall, 10% (22 of 220) of captured snakes had broken tails. This value is similar to published values for closely related snakes but is smaller than those reported for another Nerodia sipedon sipedon population. Unlike some previously published studies, the frequency of injured tails was not greater for females or lesser for first-year snakes. The mech- anism behind the injury frequency observed here and the reason for differences between this and other studies are unknown. Key Words: Beaver Archipelago, Nerodia sipedon sipedon, Northern Watersnake, tail breakage, Michigan. The frequency of tail injuries in a lizard or snake population was at one time considered to be a useful index of the predation pressure on that population be- cause tail autotomy and breakage (Mendelson 1992) are thought to be important defense mechanisms (re- viewed in Arnold 1988). More recently, theoretical and empirical tests of this hypothesis have led to the asser- tion that loss or injury of the tail may instead be related to predator inefficiency or alternative sources of mor- tality and that careful investigation must be undertaken to determine the mechanism behind observed injury frequencies (Schoener 1979; Medel et al. 1988). While such data must indeed be interpreted with caution, re- porting the frequency of tail injury in study populations can be useful when combined with demographic data (Arnold 1988), for example in forming hypotheses re- garding sex or size-based differences in anti-predator mechanisms (Fitch 2003). Several researchers have reported the frequency of tail injuries in populations of snakes. Fitch (1999) found that the frequency of tail breakage increased with the age/size of individuals in a Kansas population of Nero- dia s. sipedon, the Northern Watersnake. More specif- ically, the frequency of female tail breakage ranged from 21.2 to 44.5%, and the frequency of male tail breakage ranged from 6.5 to 25% in increasing body size categories. Willis et al. (1982) reported that fe- male Eastern Garter Snakes (Thamnophis sirtalis) and Northern Ribbon Snakes (Thamnophis sauritus) had a higher incidence of tail loss than males (13% versus 6%, and 12% versus 7%, respectively) and that tail loss was more prevalent in larger size classes. However, nei- ther trend was statistically significant in populations of Butler’s Garter Snake (Thamnophis butleri). Fitch (2003) found that tail breakage was more common for female T. sirtalis (16.7%) than for males (10.3%) and more common for large snakes (13.7%) than for first- year snakes (2.42%). In general, females and larger snakes appear more likely to have broken tails. I studied the frequency of broken tails of the North- ern Watersnake in the Beaver Archipelago of north- eastern Lake Michigan during the summers of 2000 and 2001. I captured snakes from Beaver Island (45°41.26'N, 85°30.34'W), Garden Island (45°48.28'N, 85°29.41'W), High Island (45°43.88'N, 85°39.54'W), and Hog Island (45°48.39'N, 85°22.15'W). Upon cap- 436 THE CANADIAN FIELD-NATURALIST TABLE 1. Chi-square contingency table for comparison of the frequency of tail breakage among male, female, and first-year Nerodia sipedon sipedon in the Beaver Archipelago of Lake Michigan. The calculated x? was 0.23 (critical x7 = 5.99, df =2). Class Number of Number of injured snakes uninjured snakes Total % Injured Male 7 55 62 AES Female 13 120 133 9.8 First-year 2, 23 25 8.0 Totals 22 198 220 10.0 ture, I brought snakes to the laboratory where I then measured snout-vent length (SVL) and noted the tail condition and sex of each individual. I released all snakes at the point of capture as soon as possible after processing. I pooled snakes from all islands and calculated the overall percentage of snakes with tail breaks. I also compared the frequency of broken tails among males, females, and unsexed first-year snakes, and among three capture locations (Beaver Island Lake Michigan shoreline, Beaver Island interior lake, and Garden Island Lake Michigan shoreline) within sexes using Chi-square contingency table analysis. I hypoth- esized that females would have a higher frequency of tail breakage than males and that first-year snakes would have a lower frequency of tail breakage than both males and females. Two hundred and twenty snakes were captured from the four islands. Beaver Island produced 153 snakes, Garden Island 55, High Island 1, and Hog Island 11 snakes. Captured snakes ranged in size from 149 to 860 mm SVL with a mean of 478.2 mm and a stan- dard deviation of 164 mm. In all, 10% (22) of the snakes had broken tails. There was no statistically sig- nificant difference in the frequency of tail breakage among males, females, and first-year snakes (x7 = 0.23, critical y? = 5.99, df = 2; Table 1). Within males, there was no statistically significant difference in the fre- quency of tail breakage among capture sites (x? = 0.33, critical y7 = 5.99, df =2; Table 2). However, females captured on the Lake Michigan shoreline of Garden Island had a higher frequency of tail breakage than those captured elsewhere (x7 = 8.9, critical y? = 5.99, dfi=2; Table3): My data do not closely match those of any pre- vious studies. The overall frequency of tail breakage is similar to that reported by Willis et al. (1982). How- ever, unlike the populations of N. s. sipedon and T. sirtalis studied by Fitch (1999; 2003), and the popu- lations of T. sirtalis and T. sauritus studied by Willis et al. (1982), the frequency of broken tails is not great- er for females in the Beaver Archipelago overall (al- though it is greater if only Garden Island is consid- ered). Furthermore, first-year snakes in the Beaver Archipelago do not appear to have a lower frequency of tail breakage than larger snakes. The 7. butleri pop- ulations of Willis et al. (1982) appear to be similar to my snakes in the lack of common patterns of tail break- age. The sample size (287) for T. butleri in that study is similar to my own, while the sample sizes of the 7. sirtalis, T. sauritus and N. s. sipedon studies are con- siderably larger. This suggests that the ability to detect patterns in the frequency of tail injuries in snake pop- ulations may be dependent upon sample size. TABLE 2. Chi-square contingency table for comparison of the frequency of tail breakage of male Nerodia sipedon sipedon among capture sites in the Beaver Archipelago of Lake Michigan. The calculated x? was 0.33 (critical y° = 5.99, df =2). Capture Number of site Beaver Island Lake Michigan shoreline Beaver Island interior lakes Garden Island Lake Michigan shoreline Totals mee injured snakes Number of uninjured snakes Total % Injured ils) 17 11.8 20 22 9.1 18 21 14.3 53 60 Hike 7/ TABLE 3. Chi-square contingency table for comparison of the frequency of tail breakage of female Nerodia sipedon sipedon among capture sites in the Beaver Archipelago of Lake Michigan. The calculated x7 was 8.9 (critical y* = 5.99, df =2). Capture site Number of injured snakes Beaver Island Lake Michigan shoreline Beaver Island interior lakes Garden Island Lake Michigan shoreline Totals = wlnony sp Number of uninjured snakes Total % Injured 60 64 6.2 34 36 SD) 21 28 25.0 TS 128 10.1 2004 Fitch (2003) suggested that the frequency of tail breakage should be expected to vary among different study sites based on the presence of predators and availability of escape cover. Such factors may explain the differences in overall frequency of tail breakage between this and other studies and between islands in the Beaver Archipelago within females. Fitch (1999) also suggested that NV. s. sipedon individuals with great- er mass are more likely to have broken tails because they whirl repeatedly in an attempt to escape when grasped by a predator. Greater mass makes tail break- age more likely when this whirling motion is employed. The smaller mass of first-year snakes, the smaller mass of male N. s. sipedon in comparison to females of the same age (Fitch 1999), and the fact that broken tails may potentially damage the hemipenes of males all sug- gest that these groups should have lower frequencies of tail breakage than large snakes and females, res- pectively, regardless of locality. These patterns were not clear in this study or in the populations of 7: butleri studied by Willis et al. (1982). Sample size may be the critical factor in the dis- crepancy between studies. The two studies that did not show common patterns in tail breakage had rela- tively small sample sizes. A large sample size may be necessary for two reasons: (1) the patterns are real but the categorical nature of the data (break or no break) requires that large samples be collected for a pattern to be detectable, or (2) the patterns are not real but stem from the fact that any difference, no matter how small, can be statistically significant if the sample size is large enough (Johnson 1999). The general similarity of patterns of tail breakage across studies with large sample sizes suggests that the first reason may be the case. However, discrepancies among studies underscore the need for more research in this area. Researchers should continue to record and report the frequency of tail breakage in populations of lizards and snakes, but more carefully designed studies that attempt to determine the direct causes and consequences of tail breakage are needed (Medel et al. 1988). Until such studies are carried out, the meaning and utility of tail breakage data will remain speculative. NOTES 437 Acknowledgements This research was done as part of a M.S. thesis at Central Michigan University. I thank my Thesis com- mittee: James C. Gillingham, Michael J. Hamas, and David L. Clark. I also thank John W. Rowe for help and advice in the field and laboratory. E. S. Bowen, M. F. Haussmann, and two anonymous reviewers pro- vided valuable comments on earlier drafts. Financial support was provided by the Central Michigan Uni- versity Department of Biology, the Central Michigan University College of Graduate Studies, and the Central Michigan University Biological Station on Beaver Island. Animals were collected under MDNR Cultural and Scientific Collector’s Permit CA 341 is- sued to J. C. Gillingham. The experimental protocol was approved by the Central Michigan University In- stitutional Animal Use and Care Committee (approval number 02-06). Literature Cited Arnold, E. N. 1988. Caudal autotomy as a defense. Pages 235- 273 in Biology of the Reptilia. Volume 16, Ecology B. Edited by C. Gans and R. B. Huey. Alan R. Liss Inc., New York. Fitch, H. S. 1999. A Kansas snake community: composition and changes over 50 years. Krieger Publishing Company, Malabar, Florida xi + 165 pages. Fitch, H. S. 2003. Tail loss in Garter Snakes. Herpetological Review 34: 212-213. Johnson, D. H. 1999. The insignificance of statistical signifi- cance testing. Journal of Wildife Management 63: 763- TLE: Medel, R. G., J. E. Jimenez, S. F. Fox, and F. M. Jaksic. 1988. Experimental evidence that high population fre- quencies of lizard tail autotomy indicate inefficient preda- tion. Oikos 53: 321-324. Mendelson, J. R. 1992. Frequency of tail breakage in Conio- phanes fissidens (Serpentes: Colubridae). Herpetologica 48: 448-455. Schoener, T. W. 1979. Inferring the properties of predation and other injury-producing agents from injury frequencies. Ecology 60: 1110-1115. Willis, L., S. T. Threlkeld, and C. C. Carpenter. 1982. Tail loss patterns in Thamnophis (Reptilia: Colubridae) and the probable fate of injured individuals. Copeia 1982: 98-101. Received 21 February 2003 Accepted 4 November 2004 438 THE CANADIAN FIELD-NATURALIST Vol. 118 The Heather Vole, Genus Phenacomys, in Alaska S. O. MACDoNALD!:3, AMY M. RUNCK?, and JosEPH A. COOK>*” 1P.O. Box 58, Gila, New Mexico 88038-0058 USA Biological Sciences Department, Idaho State University, Pocatello, Idaho 83209-8007 USA 3Museum of Southwestern Biology, University of New Mexico, Albuquerque, New Mexico 87131 USA; Corresponding author, e-mail cookjose @unm.edu MacDonald, S. O., Amy M. Runck, and Joseph A. Cook. 2004, The heather vole genus Phenacomys, in Alaska. Canadian Field-Naturalist 118(3): 438-440. Four specimens of heather vole (genus Phenacomys) collected in the coastal mountains of Southeast Alaska document the first Recent records of this vole in Alaska. Alternative hypotheses on the relationship of these newly-discovered populations to extant and historical populations are outlined, and additional studies proposed. Key Words: Heather vole, Phenacomys sp., Southeast Alaska. Since publication of a synopsis of the land mammal fauna of Southeast Alaska (MacDonald and Cook 1996), new field and laboratory studies of the mam- mals of this region have resulted in new perspectives on the zoogeography of the North Pacific Coast (Con- roy et al. 1999; Cook and MacDonald 2001; Cook et al. 2001). In our 1996 publication, we speculated that several mammal species found on the eastern side of the coastal mountains in Canada may occur along the Alaska coast. One species, the heather vole (Phena- comys intermedius), had previously been recorded within 2 km of the southern Alaska border in the sub- alpine near Salmon Glacier in British Columbia (ap- proximately 56.17° N, 130.03° W), and Nagorsen and Jones (1981) reported its occurrence in the Chilkat Pass north of Haines and Southeast Alaska. Subsequent trapping efforts in Alaskan mountains near Salmon Glacier and elsewhere along the mainland coast result- ed in the capture of four specimens of this vole (Table 1) from two widely separate localities (Figure 1). These specimens, deposited at the University of Alaska Museum (UAM), constitute the first documented rec- ords of extant populations of heather vole in Alaska. On 11 August 1996, we captured a single heather vole (UAM 42371) above timberline in the glacially- isolated Chilkat Range at the northern end of South- east Alaska near Excursion Inlet, Glacier Bay National Park (58.41° N, 135.43° W; elevation approximately 900 m). Also recorded in this alpine area were Sorex monticolus and melanistic Marmota caligata. In 1999, three heather voles were collected from the far southern end of mainland Southeast Alaska, approximately 400 km south of the 1996 Glacier Bay record. On 23 September 1999, two male (UAM 64239, UAM 64240) and one female (UAM 50000) heather voles were found in subalpine habitat along the Titan Trail in the Reverdy Mountains, approximately 10 km north of Hyder, Alaska (approximately 55.83° N, 130.03° W; elevations between 1128-1220 m). Synap- tomys borealis was also sampled at this site. The distribution of the heather vole is poorly known throughout western North America (Figure 1) and geo- graphic variation has been little studied (for overviews see McAllister and Hoffmann 1988; Nagorsen 1990; Musser and Carleton 1993). Some authorities (Ander- son 1942; Cowan and Guiguet 1965; Peterson 1966; Hallett 1999) recognized the distinctiveness of western populations (P. intermedius of westcentral British Columbia south to California and New Mexico) and separated them from P. ungava in the north and east (southern Yukon Territory and northern British Colum- bia eastward to Newfoundland) (McAllister and Hoff- mann 1988; George 1999); others considered all to comprise a single species (Crowe 1943; Youngman 1975; Honacki et al. 1982; Nagorsen 2004). Hall (1981) speculated that intermedius may be a composite of two or three allopatric species (see McAllister and Hoff- mann 1988), but the lack of specimens from too few areas has prevented meaningful progress in resolving the problem. Reluctance of this species to enter traps may be the primary factor limiting sample size and study materials (McAllister and Hoffmann 1988). The taxonomic affinities of newly-discovered popu- lations of heather vole in Southeast Alaska are not immediately evident. Based on the map provided by Hall (1981: 787, as adjusted by McAllister and Hoff- mann 1988 and Nagorsen 2004), Alaska’s southern- most heather voles could share close kinship with the P. i. intermedius group of “western” heather voles occurring in the coastal mountain ranges of adjacent British Columbia. Heather voles from northern South- east Alaska, on the other hand, might possibly have connection with the “eastern” populations (as P. unga- va mackenzii) in northwestern British Columbia and southern Yukon Territory. Both of these scenarios pre- sume relatively recent colonization from inland pop- ulations following the (ongoing) retreat of the last major glaciation from the region. A similar scenario has been proposed for American Marten (Martes amer- icana) (Stone et al. 2002) and other mammals (Cook 438 2004 NOTES 439 TABLE |. Summary of specimen data from collection notes and records at the University of Alaska Museum (UAM). Alaska Frozen Tissue Number refers to the cryogenic collection administered by UAM. Measurements and abbreviations are as follows: total length-tail length-hind foot-ear from notch (in millimeters)=mass (in grams); t = testes (length and width in mm), plsc = number of placental scars on right and left uterine horn; F = female, M = male. Alaska Frozen Tissue Number UAM Catalog Number Sex (Reproduction) Measurements 42371 17262 50000 30176 64239 30178 64240 30179 et al. 2001). An alternative hypothesis, however, 1s that one or more of these populations are remnants of a former coastal population whose origin in the region preceded the last glacial advance (MacDonald and Cook 1996; Fleming and Cook 2002). The discovery of late Pleistocene fossil remains of Phenacomys in limestone cave deposits in the Alex- ander Archipelago of Southeast Alaska has added his- torical dimension to these hypotheses. Preliminary Chilkat River Gulf of Alaska LOCALITY KEY @ Chilkat Range @ Reverdy Mts. On Your Knees Cave fossil site M (t=2X3) 120-31-19-11=19 F (plsc: 4R, OL) 156-37-18-14=33.9 M (t=3X2) 129-32-17-13=22.8 M (t=3X2) 115-26-16-10=14.6 analyses of thousands of rodent teeth recovered from On Your Knees Cave at the extreme northwest corner of Prince of Wales Island in Southeast Alaska (Figure 1) (Heaton 1995, 2004*; Heaton and Grady 2003) in- dicate that Heather Voles had been present in the vicin- ity of this site during the Middle Wisconsin up to the start of, but not during or following, the last glacial maximum (between about 24 000 to 13 000 years ago) (Heaton 2004*; Heaton and Grady 2003). Other % CANADA Sc , Za 4p Stikine River Unuk River Ficure 1: Localities of specimens of Phenacomys in Southeastern Alaska from Recent and Late Pleistocene sites. Inset range map modified according to McAllister and Hoffman (1988) following subspecific designations suggested by George (1999) and Hallett (1999). 440 rodent remains discovered in these same pre-glacial deposits were Brown Lemming, Lemmus trimucro- natus, a species thought absent from the region but recently discovered by UAM in the mountains near Haines, and the Hoary Marmot, Marmota caligata, a species now restricted to mainland localities. How the new records of Phenacomys from South- east Alaska relate to extant and historical populations has yet to be determined. Analyses of samples using modern molecular and morphological techniques could help clarify these relationships and provide a test of alternate hypotheses on the origin and dynamics of the region’s biota. Clearly, more samples from more localities are needed. An expanded effort to excavate fossils throughout the region, along with a continua- tion of survey efforts to further define the current distribution of Phenacomys and other mammals that share a deep history in the region (e.g., Lemmus, Marmota), as well as provide adequate samples for assessment, are needed. The alpine zone of Southeast Alaska is virtually unexplored and future inventories need to target sites above treeline along the mainland as well as on the more mountainous islands in the Alexander Archipelago. Acknowledgments A number of individuals assisted with field work that led to these new records including C. T. Seaton, Able Duffy, Joni Reese, Mike Brown, Guillermo D’Eha, and John Chythlook. Brandy Jacobsen, Gordon Jarrell, and Dusty McDonald of the University of Alaska Mu- seum provided specimen information. Thanks also to David Nagorsen for reviewing an early draft of this paper. This research was funded by the United States Fish and Wildlife Service, United States Department of Agriculture Forest Service, Alaska Cooperative Fish and Wildlife Research Unit, and the National Science Foundation (DEB0196095 & DEB9981915). Documents Cited [marked * in text citations] Heaton, T. H. 2004. Ice Age paleontology of Southeast Al- aska. http://www.usd.edu/esci/alaska/index.html. 3 Decem- ber 2004. Literature Cited Anderson, R. M. 1942. Canadian voles of the genus Phena- comys with description of two new Canadian subspecies. Canadian Field-Naturalist 56: 56-60. Conroy, C. J., J. R. Demboski, and J. A. Cook. 1999. Mammalian biogeography of the Alexander Archipelago of Alaska: a north temperate nested fauna. Journal of Biogeography 26: 343-352. Cook, J. A., and S. O. MacDonald. 2001. Should endemism be a focus of conservation efforts along the North Pacific Coast of North America? Biological Conservation 97: 207-213. Cook, J. A., A. L. Bidlack, C. J. Conroy, J. R. Demboski, M. A. Fleming, A. M. Runck, K. D. Stone, and S. O. MacDonald. 2001. A phylogeographic perspective on endemism in the Alexander Archipelago of southeast Alaska. Biological Conservation 97: 215-227. THE CANADIAN FIELD-NATURALIST Vol. 118 Cowan, I. M., and C. J. Guiguet. 1965. The mammals of British Columbia. British Columbia Provincial Museum, Handbook No. 11, 24 edition: 1-414. Crowe, P. E. 1943. Notes on some mammals of the southern Canadian Rocky Mountains. Bulletin of the American Museum of Natural History 80: 391-410. Fleming, M. A., and J. A. Cook. 2002. Phylogeography of endemic ermine (Mustela erminea) in southeast Alaska. Molecular Ecology 11: 795-808. George, S. B. 1999. Eastern heather vole, Phenacomys un- gava. Pages 618-619 in The Smithsonian book of North American mammals. Edited by D. E. Wilson and S. Ruff. Smithsonian Institution Press, Washington, D.C., in asso- ciation with the American Society of Mammalogists. 750 pages. Hall, E. R. 1981. The mammals of North America. Wiley- Interscience, New York. 1181 pages. Hallett, J. G. 1999. Western heather vole, Phenacomys inter- medius. Pages 617-618 in The Smithsonian book of North American mammals. Edited by D. E. Wilson and S. Ruff. Smithsonian Institution Press, Washington, D.C., in associa- tion with the American Society of Mammalogists. 750 pages. Heaton, T. H. 1995. Middle Wisconsin bear and rodent re- mains discovered on Prince of Wales Island, Alaska. Cur- rent Research in the Pleistocene 12: 92-94. Heaton, T. H., and F. Grady. 2003. The Late Wisconsin verte- brate history of Prince of Wales Island, Southeast Alaska. Pages 17-53 in Ice Cave Faunas of North America. Edited by B. W. Schubert, J. I. Mead, and R. W. Graham. Indiana University Press. 400 pages. Honacki, J. H., K. E. Kinman, and J. W. Koeppl. Editors. 1982. Mammal species of the world: a taxonomic and geo- graphic reference. Allen Press, Inc. and The Association of Systematics Collections, Lawrence, Kansas. 694 pages. MacDonald, S. O., and J. A. Cook. 1996. The land mammal fauna of southeast Alaska. Canadian Field-Naturalist 110: 571-599. McAllister, J. A., and R. S. Hoffmann. 1988. Phenacomys intermedius. Mammalian Species 305: 1-8. Musser, G. G., and M. D. Carleton. 1993. Family Muridae. Pages 501-756 in Mammal species of the world: a taxo- nomic and geographic reference. Edited by D. E. Wilson and D. M. Reeder. Second Edition. Smithsonian Institution Press, Washington. 1206 pages. Nagorsen, D. W. 1990. The mammals of British Columbia: a taxonomic catalogue. Memoir Number 4, Royal British Columbia Museum. 140 pages. Nagorsen, D. W. 2004. The rodents and lagomorphs of Brit- ish Columbia. Royal British Columbia Museum Hand- book, Volume 4. University of British Columbia Press, Vancouver. 352 pages. Nagorsen, D. W., and D. M. Jones. 1981. First records of the Tundra Shrew (Sorex tundrensis) in British Columbia. Canadian Field-Naturalist 95: 93-94. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press, Toronto. 465 pages. Stone, K., R. Flynn, and J. Cook. 2002. Post-glacial coloni- zation of northwestern North America by the forest associ- ated American marten (Martes americana). Molecular Ecology 11: 2049-2064. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museum of Natural History, Publications in Zoology 10: 1-192. Received 26 November 2002 Accepted 30 December 2004 2004 NOTES 44] Extension de l’aire de distribution connue de la Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du Québec JEAN-FRANCOIS DESROCHES! et ISABELLE PICARD ' Collége de Sherbrooke, Département des Techniques d’écologie appliquée, 475 du Parc, Sherbrooke, Québec JIK 4K] Canada Desroches, Jean-Francois, et Isabelle Picard. 2004. Extension de l’aire de distribution connue de Ja Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du Québec. Canadian Field-Naturalist 118(3) : 441-442. La Musaraigne fuligineuse (Sorex fumeus) figure sur la liste des especes de la faune vertébrée susceptibles d’étre désignées menacées ou vulnérables au Québec. Le 7 juin 1999, nous avons récolté un individu mort en bordure de la riviére Saint-Nicolas, a Godbout, dans la région de la Céte Nord. Cette observation constitue une extension de l’aire de répartition connue de |’ espéce d’ environ 200 km vers le nord-est. Mots-clés : Musaraigne fuligineuse, Sorex fumeus, aire de distribution, Québec. La Musaraigne fuligineuse (Sorex fumeus) est pré- sente dans le nord-est de 1’ Amérique du Nord, des Great Smokies des Carolines au sud jusqu’au centre de l’Ontario et aux Maritimes vers le nord (Banfield 1977). Au Québec, la mention la plus nordique de Pespéce se situe a Val-Jalbert, aux abords du lac Saint- Jean (van Zyll de Jong 1983). Sa répartition connue vers le nord-est, sur la rive nord du fleuve Saint-Laurent, ne dépasse guére la riviére Saguenay (van Zyll de Jong 1983; Banfield 1977). Cette musaraigne figure sur la liste des espéces de vertébrés susceptibles d’étre désig- nées menacées ou vulnérables au Québec (Beaulieu 1992). Les chercheurs contactés en 1989 n’avaient pas rapporté de capture récente au Québec (Beaulieu 1992) mais depuis, elle a été capturée a divers endroits notam- ment dans le sud de la province (G. Lupien, commu- nication personnelle). Le 7 juin 1999, nous avons trouvé un spécimen de la Musaraigne fuligineuse mort, probablement noyé, en bordure de la riviére Saint-Nicolas, a environ 100 métres au sud de la route 138 a Godbout (49°19'10"N, 67°47'24"0), dans la région de la Cote Nord. Lhabitat a cet endroit est une aulnaie, bordée par une sapiniere a Bouleau blanc. Aucun autre micromammifere n’y a été observé. Le spécimen de Musaraigne fuligineuse a été récolté pour identification. Les mesures ont été prises sur le spécimen a l’aide d’un vernier a cou- lisse, par J-F. D. et S. Houde, et elles correspondent a celles de la Musaraigne fuligineuse (tableau 1). Au Québec, la Musaraigne fuligineuse peut étre con- fondue avec la Musaraigne cendrée (Sorex cinereus), ayant toutes deux de quatre a cing dents unicuspides visibles, la limite postérieure du foramen infraorbi- taire antérieure ou prés de la limite entre les molaires 1 et 2, le trou mentonnier situé a la hauteur du proto- conide de la moiaire 1, et la mandibule dépourvue de foramen postmandibulaire (Maisonneuve et al. 1997). Elle s’en distingue toutefois par certaines mesures du crane et du corps (tableau 1). Le spécimen a malheu- reusement été perdu quelques jours plus tard, jeté par erreur avec d’autres micromammiferes morts, mais les mesures prises confirment son identification. Cette observation constitue une extension d’aire de — Vespece de 200 km vers le nord-est, soit a partir de V’embouchure de la riviére Saguenay (van Zyll de Jong 1983). La Musaraigne cendrée (Sorex cinereus), une espece plus commune au Québec (observation per- sonnelle), a une distribution qui s’étend bien plus au TABLEAU |. Mesures comparées du spécimen de musaraigne trouvé sur la Cote Nord, de la Musaraigne fuligineuse et de la Musaraigne cendrée. Longueur Largeur du Longueur Longueur de Espéce du crane (mm) maxillaire (mm) totale (mm) la queue (mm) Spécimen de la Céte Nord WET 4.8 111,4 47,8 Musaraigne fuligineuse 17,5 a 18,8? 4,8 45,4? 1104127! 40 452! SIS? >4,6° 104 a 1252 42 a 54? Musaraigne cendrée 15,2 a 16,92 3.8 44,32 924110! 37 a 46! <173 <4,6° Ta le52 28 a 507 ' Banfield 1977 > van Zyll de Jong 1983 3 Maisonneuve et al. 1997 441 442 nord (van Zyll de Jong 1983; Banfield 1977). La dé- couverte de la Musaraigne fuligineuse 4 Godbout in- dique que la distribution de certaines espéces de petits mammiféres est encore mal connue. En ce sens, il est permis de croire que de futures découvertes seront faites et permettront de mieux connaitre les habitats fréquentés par ces espéces et de préciser leur aire de distribution dans le nord de la province. II est a noter que cette mention de la Musaraigne fuligineuse figure dans |’Atlas des micromammiféres du Québec, sans documentation publié en 2002 (Desrosiers et al. 2002). Remerciements Nous tenons a remercier Mme Stéphanie Houde, technicienne de la faune, pour la vérification de l’iden- tification du spécimen de méme que M. Gilles Lupien, technicien de la faune a la Société de la Faune et des Parcs du Québec, pour ses commentaires sur les cap- tures de l’espéce au Québec. Nous remercions égale- ment le comité ZIP de l’estuaire du Saint-Laurent, qui a permis la réalisation de |’inventaire duquel la présente découverte de la Musaraigne fuligineuse a été faite. THE CANADIAN FIELD-NATURALIST Vol. 118 Littérature citée Banfield, A. W. F. 1977. Les Mammiféres du Canada. Les Presses de l’ Université Laval et University of Toronto Press. Publié par le Musée national des Sciences naturelles, Musées nationaux du Canada. Ottawa, Canada. 406 pages. Beaulieu, H. 1992. Liste des espéces de la faune vertébrée susceptibles d’étre désignées menacées ou vulnérables au Québec. Ministére du Loisir, de la Chasse et de la Péche. Québec. 107 pages. Desrosiers, N., R. Morin, et J. Jutras. 2002. Atlas des micromammiféres du Québec. Société de la faune et des parcs du Québec et Fondation de la Faune du Québec. 92 pages. Maisonneuve, C., R. Mc Nicoll, S. St-Onge, et A. Des- rosiers. 1997. Clé d’identification des micromammiferes du Québec. Ministére de 1’ Environnement et de la Faune, Québec. 19 pages. van Zyll de Jong, C. G. 1983. Traité des Mammiféres du Canada, volume | : Les Marsupiaux et les Insectivores. Mu- sée national des Sciences naturelles, Musées nationaux du Canada. Ottawa, Canada. 217 pages. Recu 28 octobre 2001 Acceptée 15 janvier 2005 Gulls, Larus spp., Foraging at Pink Salmon, Oncorhynchus gorbuscha, Spawning Runs Mary FE. WILLSON 5230 Terrace Place, Juneau, Alaska 99801 USA; e-mail mwillson@ gci.net Willson, Mary F. 2004. Gulls, (Larus spp.), foraging at Pink Salmon, Oncorhynchus gorbuscha, spawning runs. Canadian Field-Naturalist 118(3): 442-443. Small and immature gulls foraged more often on drifting salmon eggs than did large and mature gulls, and large and mature gulls foraged more often on salmon carcasses, at streams in Southeast Alaska. These differences may be related to body size via physical strength and dominance status, as well as foraging experience. Key Words: Gulls, Larus spp., Pink Salmon, Oncorhynchus gorbuscha, Southeast Alaska. Salmon spawning runs attract many foraging birds and mammals (Willson and Halupka 1995) and are clearly an important food resource for wildlife in late summer and fall. Gulls (Larus spp.) are among the most numerous predators and scavengers at salmon runs, sometimes occurring in the thousands (Isleib and Kes- sel 1973). Here I report how four species of gull ex- ploited runs of Pink Salmon (Oncorhynchus gor- buscha) at the mouths of streams in Juneau, Alaska. Study Area and Methods Most of these observations were made at the mouths of Salmon Creek and Sheep Creek (less than 12 km apart), with some supplementary observations at four other small, nearby streams, within the city and bor- ough of Juneau in Southeast Alaska (ca. 58°30'N, 133°30'W). I observed four species of gulls foraging at or near low tides, usually in the morning, from 27 August to 6 September 2002 (>6 hrs of observation). At any one time, many dozens of gulls were present at the two primary study locations. At low tide, these creeks flow over intertidal deltas, often with several shallow channels used by incoming salmon. I scanned these deltas with spotting scope and binoculars, noting the foraging activities of the gulls and counting the individuals engaged in several distinctive foraging ac- tivities. Birds shifted position frequently, using different parts of the deltas; many individuals of the three larger species also changed foraging activity as I watched. Most scans were made at different locations on differ- ent days; if more than one scan occurred on the same day and stream delta, they were separated in time by about 30 min. I recorded foraging observations in three categories: foraging on salmon eggs (drifting eggs are common because later spawners often disturb the grav- els over the nests of earlier spawners, foot-paddling by gulls also stirs up poorly buried eggs), foraging on salmon carcasses (including some extraction of eggs from living or dead fish), and searching for inverte- brates in Fucus mats (where amphipods and isopods were common) and tide pools. 2004 NoTes 443 TABLE |. Foraging activity of four species of gull at salmon spawning runs in southeast Alaska. Species/age class Eggs Salmon Invertebrates N (observations) Bonaparte’s Gull 82% - 18% 353 Mew Gull adults 33% 1% 66% 232 Herring Gull adults* 31% 58% 11% 26 Herring Gull immatures 80% 20% - 15 Glaucous-wing adults* 11% 87% 2% 362 Glaucous-wing immatures 44% 55% 1% 222 ‘Frequency distributions for adults and immatures were significantly different (y’, = 88.5 and 7.1, for the two species respectively; p < 0.05 in both cases). Results Foraging behavior of each species was very con- sistent among the observed streams, but there were marked differences in foraging among species and age classes (Table 1). The small (ca. 190 g, Sibley 2000) Bonaparte’s Gull (L. philadelphia) foraged for salmon eggs chiefly by hovering over the streams and plunge-diving for drifting eggs, but they occasionally sought invertebrates in Fucus mats or tide pools. Most birds were in nonbreeding plumage, and adults were not distinguished from immatures. Mew Gulls (L. canus), of intermediate size (ca. 420 g, Sibley 2000), waded in the shallows, searching for drift eggs, or hunted invertebrates in Fucus mats. They seldom foraged on fish carcasses and then only if no large gulls were nearby. Too few immatures were present for adequate sampling, so all observations per- tain to adults. Both Herring (L. argentatus) and Glaucous-winged (L. glaucescens) gulls (body mass about 1000-1100 g for both species, Sibley 2000) foraged chiefly on car- casses and on eggs, only rarely visiting the Fucus mats on these deltas. Eggs were commonly obtained while wading, occasionally by foot-paddling. Glaucous-wing adults sometimes pulled live salmon from the stream, poking initially at eyes and at the vent area to force egg extrusion. Immatures of both species foraged more often on eggs than did adults, while adults foraged more often on carcasses. Both of these larger gulls often examined very old, bleached and decayed, carcasses that appeared to offer little or nothing edible. Discussion Salmon eggs are rich in lipids (Gende et al. 2001, 2004), offering an easily captured and high-calorie food, and brightly colored, so they are easy to see as they drift downstream. Bonaparte’s Gulls, Mew Gulls, and immatures of the larger species foraged heavily on eggs. The demands of imminent fall migration for the smaller gulls may provide added impetus for rapid energy acquisition; most Bonaparte’s Gulls left the area before the Pink Salmon runs were completely over. The immatures are less experienced foragers than adults, and eggs may be more easily accessed than carcasses. Carcasses are probably less accessible to smaller gulls because the skin is tough and hard to breach and also because the larger gulls (and ravens and eagles) can easily dominate the smaller birds. Glaucous-winged Gulls selectively killed and eviscer- ated female Sockeye Salmon (O. nerka) in shallow streams in westem Alaska (Mossman 1958). The forag- ing differences among gull species here are similar to the observations of Moyle (1966) elsewhere in Alaska. Other species of wildlife also favor salmon eggs. Bears (Ursus spp.) often feed selectively on eggs (and other high-lipid body parts) from live-caught fish (Gende et al. 2001). American Dipper (Cinclus mexi- canus) chicks, juveniles, and adults feed on drift eggs (Obermeyer et al. 1999; K. E. Obermeyer and M. F. Willson, unpublished observations). Foraging: on drift eggs by wildlife has no significant effect on salmon populations, because drift eggs do not survive outside the nesting gravels. Literature Cited Gende, S. M., T. P. Quinn, and M. F. Willson. 2001. Con- sumption choice by bears feeding on salmon. Oecologia 127: 372-382. Gende, S. M., T. P. Quinn, M. F. Willson, R. Heintz, and T. M. Scott. 2004. Magnitude and fate of salmon derived nutrients and energy in a coastal system ecosystem. Journal of Freshwater Ecology 19: 149-160. Isleib, M. E., and B. Kessel. 1973. Birds of the north gulf coast-Prince William Sound region, Alaska. Biological Papers, University of Alaska 14: 1-149. Mossman, A. S. 1958. Selective predation of glaucous-winged gulls upon adult red salmon. Ecology 39: 482-486. Moyle, P. 1966. Feeding behavior of the glaucous-winged gull on an Alaska salmon stream. Wilson Bulletin 78: 175-190. Obermeyer, K. E., A. Hodgson, and M. F. Willson. 1999. American Dipper, Cinclus mexicanus, foraging on salmon eggs. Canadian Field-Naturalist 113: 288-290. Sibley, D. A. 2000. The Sibley guide to birds. Knopf, New York. Willson, M. F., and K. C. Halupka. 1995. Anadromous fish as “keystone” resources in vertebrate communities. Con- servation Biology 9: 489-497. Received 26 February 2003 Accepted 22 January 2004 444 THE CANADIAN FIELD-NATURALIST Vol. 118 Sequential Polyandry in Piping Plover, Charadrius melodus, Nesting in Eastern Canada DIANE L. AMIRAULT!, JONATHAN KIERSTEAD!, PETER MACDONALD? and LARRY MACDONNELL! ! Canadian Wildlife Service, Environment Canada, P.O. Box 6227, Sackville, New Brunswick E4L 1G6 Canada ? Nova Scotia Department of Natural Resources, P.O. Box 99, Tusket, Nova Scotia BOW 3MO Canada Amirault, Diane L., Jonathan Kierstead, Peter MacDonald and Larry MacDonnell. 2004. Sequential polyandry in Piping Plover, Charadrius melodus, nesting in eastern Canada. Canadian Field-Naturalist 118(3): 444-446. On Cape Sable Island, Nova Scotia, we confirmed that a banded female Piping Plover (Charadrius melodus melodus) produced two broods of chicks during the 2000 nesting season, the second on a beach approximately 2 km from the first. The female abandoned her second brood two days after hatching, leaving the male to complete brood rearing. Key Words: Piping Plover, Charadrius melodus, sequential polyandry, Nova Scotia. Piping Plovers (Charadrius melodus) generally have been thought to be monogamous, with males estab- lishing and defending territories and raising only one brood per season (Wilcox 1959; Cairns 1982; Haig and Oring 1988; Haig 1992). Renesting in Piping Plovers is common if a clutch is lost (Haig 1992) and occasionally occurs if a brood of chicks is lost (Mac- Ivor 1990). In Manitoba, Piping Plovers exhibited sequential polyandrous behavior (Oring 1982; Haig 1992) where some breeding adults had multiple nests and mates (Haig and Oring 1988) after predation or storm events caused nest loss. The best example of polyandry in shorebirds has been documented in the Spotted Sandpiper (Actitis macularia) (Oring and Knudson 1973). Within Spotted Sandpipers, females are larger and more aggressive than males and establish and defend nesting territories. Female Spotted Sandpipers lay several clutches of eggs while different males incubate eggs and complete brood-rearing tasks for each. Bottitta et al. (1997) published the first observations of Piping Plovers successfully fledging two broods in one nesting season, at Griswold Point in Old Lyme, Connecticut, and Assateague Island National Seashore, along the Maryland and Virginia coast. During the Bottitta et al. (1997) studies (1989-1994), eight Piping Plover pairs successfully raised two broods each within a reproductive season. Although not all birds involved were banded to enable identification of individuals, mates in most cases were thought to have been re- tained because of plumage characteristics and proximity of second nests to locations of first nests. The potential for Piping Plovers P. m. meliodius to raise a second brood in eastern Canada is thought to be small, owing to the later, and therefore shorter, nest- ing season associated with a more northerly location. With aid of a banding program, we documented the production of a second brood by the same female Piping Plover in southwestern Nova Scotia during the 2000 nesting season. Study Area and Methods Piping Plovers have been closely monitored on two beaches (Daniels Head beach [43°26'N, 65°36'W] and Stoney Island beach [43°28'N, 65°34'W]) on Cape Sable Island in southern Nova Scotia since 1997 (P. MacDonald, unpublished data). Daniels Head beach encompasses a 15-90 m (high-low tide) wide stretch of light-coloured sand extending the full length of the 3.3 km beach, broken by four rocky points and an inlet to a sandy tidal-flat area. Low vegetated (Ammophila breviligulata and other grasses) dunes provide a break between the beach and a brackish inland marsh. Sto- ney Island is a light-coloured sand beach bordered by low, densely vegetated dunes (Ammophila breviligulata, grasses, and shrubs). The 15-75 m (high-low tide) wide beach extends 1.8 km in a crescent shape from a rocky point in the south to an embankment in the north. The beaches are separated by an approximate- ly 2 km long coastal stretch characterized by rocky shoreline, a tidal channel, and headland on which a fish processing plant is located (Boates et al. 1994*). Daniel’s Head beach supported from three to seven nesting pairs of Piping Plovers in the 1997 to 2000 period. Conversely, the single pair at Stoney Island beach was the only successful nesting event since LOOT: We trapped and banded birds on several occasions during May-July 2000. A modified Weller trap was used to capture adult birds incubating eggs (Weller 1957). A trapped bird was immediately removed from the trap, banded with a United States Fish and Wildlife Service numbered metal incoloy band on the lower right leg (for adult birds) and a Canadian Wildlife Service bicoloured plastic band on the lower left leg (specific to province) and released. Results On 24 May 2000, we trapped and banded a female Piping Plover incubating four eggs on Daniels Head beach. Four chicks hatched from this clutch on 30 May. The female was not observed with the brood after 13 2004 June, at which time only one chick remained. This chick was considered fledged on 25 June. The adult male remained with the juvenile and both were ob- served together until 5 July. A nest containing three eggs was discovered on 23 June on Stoney Island beach. Four eggs were confirmed on 25 June. We trapped a female bird on this nest on 29 June and her leg band number confirmed that this individual was the female banded on 24 May at Daniels Head beach. This second nesting event appears to have been com- pleted with a different mate as the male from the first nest remained with the surviving chick until 5 July, well after fledging. The second pair hatched four eggs on 18 July and the female was not observed with the brood (4 chicks still remained) after 20 July. One chick from this brood was considered fledged on 13 August, the last day both the juvenile and adult male were observed. Males from each nest were unbanded, facilitating identification of which adult (male or female) was present with the chicks. Discussion Although Haig and Oring (1988) documented poly- andry in Manitoba, their observations included renest- ing following nest loss but never more than one brood per nesting season. Our observations represent the first documented occurrence of a female Piping Plover relocating to produce a second brood within a nesting season on Canadian nesting grounds. Bottitta et al. (1997) were the first to report Piping Plovers raising more than one brood in a season. However, their oc- currences were documented in the United States and on a single nesting beach. Although closely related species such as Killdeer (Charadrius vociferus) and Snowy Plover (Charadrius alexandrinus) are known to produce two broods, this behaviour is thought to be extremely rare among Piping Plovers (Wilcox 1959; Cairns 1982; Whyte 1985; Haig and Oring 1988; Maclvor 1990; Strauss 1990; Loegering 1992). Thus, most second nest attempts are renests following nest loss. Presumably, the second nest on Stoney Island beach was initiated 18-19 June as Piping Plovers aver- age 6-8 days to complete laying of a full clutch (Haig 1992). The interval between the last sighting of this female with her first brood (13 June) and the calcu- lated initiation of her new clutch is brief (5-6 days). However, Piping Plovers have been known to initiate renesting after a lost clutch in as few as 4 days (Whyte 1985; Haig and Oring 1988; MacIvor 1990; Loegering 1992). Bottitta et al.’s (1997) long-term study documented several examples of two broods being produced with- in a nesting season, all of which were thought to have been completed with the same mates. That is consis- tent with renesting Piping Plovers generally retaining the same mate (Wilcox 1959; Whyte 1985; Haig and Oring 1988; MaclIvor 1990). However, in southern Manitoba, renesting birds often chose a new mate in NOTES 445 cases of high predation and nest loss (Haig and Oring 1988). Our record of intra-year mate switching sug- gests that switching may occur in cases where nest loss has not occurred and may increase an individual’s productivity. Several studies have indicated that it is not uncom- mon for one adult, usually the female, to abandon its brood prior to chicks fledging (Cairns 1977; Whyte 1985; Haig 1987). However, this is the first docu- mented case of a female plover abandoning its brood in order to produce another nest. These observations indicate that a small number of females may leave early to renest and leave further parental care to the male. Previously documented cases of sequential poly- andry in Piping Plovers have been in circumstances associated with high predation rates and catastrophic storm events (Haig and Oring 1988; Maclvor 1990). Our observations demonstrate that sequential poly- andry may also occur where there is potential for increasing productivity. The documentation of a sec- ond brood produced by one female underscores the importance of having early nest attempts succeed. Such success would allow greater chance for a sec- ond nesting event to occur, resulting in greater prod- uctivity. Maclvor (1990) reported that more young fledge from first nest attempts than from renests and from early versus late nests. MacIvor (1990) suggest- ed that increased vulnerability of chicks later in the breeding season (July-August) was due to a greater human presence on beaches then. Haig and Oring (1988) reported greater parental care on first nest at- tempts versus renests; however, no difference in hatch- ing or fledging rates was noted. Therefore, measures to enhance nest success such as placing predator exclo- sures (Bottitta et al. 1997), symbolic fencing, and beach guardian patrols may be particularly important for first nest attempts as a successful early nest may increase reproductive success. The incidence of producing two broods within a nest- ing season is thought to be fairly low, but documenta- tion of one apparent double nesting event by an in- dividual has significant implications for conducting population counts of adult Piping Plovers. This event justifies adherence of any Piping Plover census to a brief window of time, as in the absence of informa- tion such as provided by this study, the second nesting event might have been erroneously recorded as two additional individuals. Counts within a prescribed time frame will decrease the potential for double-counting birds that have relocated to renest or produce a second brood. In past censuses, observers often have con- sidered an adult found on a nest or a single adult with a brood as a nesting pair (i.e., two adult birds), even if only one adult was observed. Our observations sug- gest that a small proportion of late-nesting pairs may actually be successful early nesters on a second nest attempt. Hence, danger in overestimating numbers 446 exists when counting adult Piping Plovers outside a census window as our observations indicate not all birds retain the same mate or beach in a nesting season. Acknowledgments The Canadian Wildlife Service funded the Piping Plover banding and monitoring program during the 2000 season. The Nova Scotia Department of Natural Resources provided in-kind and logistical support for the 2000 seasonal effort. Andrew Boyne and Kevin Davidson provided useful comments on earlier drafts of the manuscript. Documents Cited (marked * in text) Boates, J. S., P. Austin-Smith, G. Dickie, R. Williams, and D. Sam. 1994. Nova Scotia Piping Plover Atlas. Unpub- lished Nova Scotia Department of Natural Resources report. 86 pages. Literature Cited Bottitta, G. E., A. M. Cole, and B. Lapin. 1997. Piping Plov- ers produce two broods. Wilson Bulletin 109: 337-339. Cairns, W. E. 1977. Breeding biology of the piping plover in southern Nova Scotia. M.Sc. thesis, Dalhousie Univer- sity, Halifax. 115 pages. Cairns, W. E. 1982. Biology and behavior of breeding Pip- ing Plovers. Wilson Bulletin 94: 531-545. Haig, S. M. 1987. The population biology and life history patterns of the piping plover. Ph.D. dissertation, University of North Dakota, Grand Forks. 121 pages. Haig, S. M. 1992. Piping Plover. Pages 1-18 in The Birds of North America (2). Edited by A. Poole, P. Stettenheim, THE CANADIAN FIELD-NATURALIST Vol. 118 and F. Gill. The Academy of Natural Sciences, Philadel- phia; The American Ornithologists’ Union, Washington, Die Haig, S. M., and L. W. Oring. 1988. Mate, site, and terri- tory fidelity in Piping Plovers. Auk 105: 268-277. Loegering, J. P. 1992. Piping Plover breeding biology, forag- ing ecology and behavior on Assateague Island National Seashore, Maryland. M.S. thesis, Virginia Polytechnic Insti- tute and State University, Blacksburg, Virginia. Maclvor, L. H. 1990. Population dynamics, breeding ecology, and management of Piping Plovers on outer Cape Cod, Massachusetts. M.S. thesis, University of Massachusetts, Amherst, Massachusetts. Oring, L. W. 1982. Avian mating systems. Pages 1-92 in Avian Biology. Volume VI. Edited by D. S. Farner and J. R. King. Academic Press, Inc., New York. Oring, L. W., and M. L. Knudson. 1973. Monogamy and polyandry in the spotted sandpiper. Living Bird 11: 59-73. Strauss, E. 1990. Reproductive success, life history patterns, and behavioral variation in a population of Piping Plover subjected to human disturbance. Ph.D. dissertation, Tufts University, Boston, Massachusetts. Weller, M. W. 1957. An automatic nest-trap for waterfowl. Journal of Wildlife Management 21: 456-458. Whyte, A. J. 1985. Breeding ecology of the Piping Plover (Charadrius melodus) in central Saskatchewan. M.Sc. thesis, University of Saskatchewan, Saskatoon, Saskatch- ewan. Wilcox, L. 1959. A twenty year banding study of the Piping Plover. Auk 76: 129-152. Received 23 May 2003 Accepted 15 November 2004 American Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, Ascaphus truei Curisty A. MorRISSEY!:? AND ROBERTA J. OLENICK? 'Department of Biological Sciences, Simon Fraser University, 8888 University Drive, Burnaby, British Columbia, V5A 1S6 Canada >Never-Spook-the-Animals Wildlife Photography, 3778 West 13' Avenue, Vancouver, British Columbia, V6R 2S6 Canada >Corresponding author (current address): Canadian Wildlife Service, Environment Canada, 5421 Robertson Road, RR#1, Delta, British Columbia V4K 3N2 Canada; email: christy.morrissey @ec.gc.ca or cmorrissey @alumni.stu.ca Morrissey, Christy A., and Roberta J. Olenick. 2004. American Dipper, Cinclus mexicanus, preys upon larval tailed frogs, Ascaphus truei. Canadian Field-Naturalist 118(3): 446-448. The American Dipper (Cinclus mexicanus) is an aquatic songbird that inhabits fast-flowing mountain streams in western North America. Although dippers are known to feed primarily on aquatic invertebrates, they will also eat juvenile fish and salmon eggs when available. In 2002, while monitoring and photographing nesting activities of the American dipper, we observed and photographed adult dippers capturing Tailed Frog (Ascaphus truei) tadpoles and feeding them to their young. This note is intended to document a rarely observed occurrence and identify interactions between two relatively uncommon species. Key Words: American Dipper, Cinclus mexicanus, Tailed Frog, Ascaphus truei, feeding, Chilliwack River, British Columbia. American Dippers (Cinclus mexicanus) are North America’s only truly aquatic songbirds occupying mountain streams of western North America from Alaska south to Mexico (Kingery 1996). Dippers are known to feed almost exclusively on in stream fauna, diving underwater and probing among the rocks for benthic invertebrates and small fish (salmon and trout) as well as fish eggs (Kingery 1996). To our knowledge, there have been no previous records in the literature of dippers feeding on amphibians. However, while studying and photographing American Dippers in southwestern British Columbia, we observed dippers 2004 NOTES 447 ni eominstiscgagty em sa Creek, British Columbia, in June 2002. Photograph by Roberta Olenick. on several occasions feeding on Tailed Frog (Ascaphus truei) larvae. Field work on the American Dipper was conducted in the Chilliwack River watershed located in the Cas- cade Mountains of southwestern British Columbia, Canada. While observing dipper nests in June and July 2002, we noted at least two different pairs of Ameri- _ can Dippers, attending separate nests on Tamihi Creek, _ a tributary of the Chilliwack River (49.02 N, 121.50 W, Ficure 2: Adult American Dipper delivering Tailed Frog tad- pole to newly fledged dipper on Tamihi Creek, British Columbia in July 2002. Photograph by Roberta Olenick. elevation 545 m) feeding Tailed Frog tadpoles to their young. Roberta Olenick photographed those dippers delivering Tailed Frog tadpoles to nestlings or fledg- lings at each nest site (Figures | and 2). On one occasion, an adult dipper captured a very large Tailed Frog tadpole (estimated length 5-6 cm) and attempted to feed it to one of its fledglings. The fledgling dipper struggled to swallow the large prey item without success. Several times, the adult dipper retrieved the tadpole, knocked it vigorously against a rock and manipulated the large wide head in its bill. Eventually, after several minutes of handling by the adult, the fledgling dipper managed to swallow the large tadpole whole. During routine annual benthic invertebrate collec- tions in April 1999-2002, we infrequently captured Tailed Frog tadpoles in our Surber samplers using kick-sampling procedures. Larval Tailed Frogs have a distinct round mouth, well adapted to use as suction onto the surface of in stream rocks, making them relatively easy to identify. Tailed Frog larvae were found on several tributaries of the Chilliwack River including Tamihi Creek, Borden Creek, and Chipmunk Creek. It is worth noting that all of these creeks have present or past logging activity which is thought to limit the presence and abundance of Tailed Frogs through increased water temperatures and siltation (Nussbaum et al. 1983). No other similar species of amphibians occupy fast flowing stream habitat in our study area. There are few known predators of Ascaphus truei. Tailed Frog tadpoles have been identified as impor- 448 tant prey for larval Pacific Giant (Dicamptodon ensa- tus) and Cope’s salamanders (Dicamptodon copei) and Red-legged Frogs (Rana aurora) (Bury 1968; Jones and Raphael 1998). Observations of garter snakes (Thamnophis spp.) (Karraker 2001) and Cutthroat Trout (Salmo clarki) depredating Tailed Frogs have also been previously reported (Daugherty and Shel- don 1982). We found only one instance in the litera- ture where American Dippers were cited as being a potential predator of Tailed Frogs, in Butler Creek, Montana (Daugherty and Sheldon 1982). However, it was not clear whether those authors actually observed dippers feeding on Tailed Frogs. Given that the Tailed Frog is a provincially blue-listed (vulnerable) species in British Columbia and of special concern in the na- tional COSEWIC listing, documentation of these rare observations is important. Tailed Frogs are endemic to the Pacific Northwest. They range from British Columbia south to Califor- nia, occupying western mountain streams and humid forests throughout a 15-20 year lifespan (Daugherty and Sheldon 1982). Larvae take from | to 4 years to metamorphose into adults in cool fast streams (Bull and Carter 1996). Their distribution directly overlaps that of the American Dipper, which occupies the same habitat and geographic range. Both species are consid- ered sensitive to environmental impacts in mountain- THE CANADIAN FIELD-NATURALIST Vol. 118 ous watersheds from anthropogenic sources (Nuss- baum et al. 1983, Kingery 1996). Given that these two species occupy the same habitat and geographic range, American Dipper predation of Tailed Frogs may be more widespread than previously acknowledged. Literature Cited Bull, E. L., and B. E. Carter. 1996. Tailed Frogs: Distri- bution, ecology, and association with timber harvest in northeastern Oregon. United States Forest Service Re- search Paper 497: 1-12. Bury, R. B. 1968. The distribution of Ascaphus truei in Cali- fornia. Herpetologica 24: 39-46. Daugherty, C. H., and A. L. Sheldon. 1982. Age determi- nation, growth, and life history of a Montana population of the tailed frog. Herpetologica 38: 461-468. Jones, L. and M. Raphael. 1998. Ascaphus truei (Tailed Frog) predation. Herpetological Review 29: 39. Karraker, N. E. 2001. Ascaphus truei predation. Herpeto- logical Review 32: 100. Kingery, H. E. 1996. American Dipper (Cinclus mexicanus). The birds of North America (229): 1-28. Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, and American Ornithologists’ Union, Washington, D.C. Nussbaum, R. A., E. D. Brodie, and R. M. Storm. 1983. Reptiles and amphibians of the Pacific Northwest. Univer- sity Press of Idaho, Moscow, Idaho. Received 28 January 2003 Accepted 6 December 2004 Piping Plover, Charadrius melodus, egg viability after seawater immersion JULIE McKNiGut!, LINDA THOMAS?, and DIANE L. AMIRAULT? ' Canadian Wildlife Service, Environment Canada, Atlantic Region, 45 Alderney Drive, Dartmouth, Nova Scotia B2Y 2N6 Canada ? Prince Edward Island National Park, Parks Canada, 2 Palmers Lane, Charlottetown, Prince Edward Island C1A 5V6 Canada 3Canadian Wildlife Service, Environment Canada, Atlantic Region, 17 Waterfowl Lane, Sackville, New Brunswick E4L 1G6 Canada McKnight, Julie, Linda Thomas and Diane L. Amirault. 2004. Piping Plover, Charadrius melodus, egg viability after seawater immersion. Canadian Field-Naturalist 118(3): 448-450. Four observed nest histories indicate Piping Plover eggs are able to survive immersion in seawater, but little is known regarding their hardiness. As Piping Plover nests are often exposed to tidal flooding, their eggs may be relatively resistant to inundation by seawater. Therefore, we suggest that replacing eggs recently flooded or washed out of-nests is a viable option for the recovery of individual nests. Key Words: Piping Plover, Charadrius melodus melodus, flooding, high tide, hatching success, viability, Prince Edward Island, Nova Scotia. The Atlantic Coast population of Piping Plover (Charadrius melodus melodus) is listed as Endan- gered in Canada and Threatened in the United States (Goossen et al. 2002: U.S. Fish and Wildlife Service 1996). Nesting Atlantic coast Piping Plovers prefer flat coastal beaches with sand and pebble substrate (Boyne and Amirault 1999; Burger 1987; Cairns and McLaren 1980). The male scrapes a shallow depres- sion in the substrate between the mean high water mark and the edge of adjacent dunes or vegetation (Burger 1987; Cairns 1982; Haig 1992). The young hatch after approximately 28 days and leave the nest within hours of hatching (Cairns 1982; Haig 1992). Nesting areas are often flooded by storm-induced overwashes (Cairns and McLaren 1980) and high water levels des- troy many plover nests each season (Sylvester 1991). Four cases in which flooded Piping Plover eggs remained viable were documented during the course of regular monitoring at Prince Edward Island National Park and in southern Nova Scotia. The following nest 2004 histories confirm the viability of flooded eggs. Case 1. On 29 May 1992, a nest at Cavendish Sand- spit in Prince Edward Island National Park (46.5036°N, 63.4408°W) was completed with four eggs. On 5 June, the nest was submerged by a high tide. Both adults were present but not searching for their eggs or vocal- ising excessively. Their eggs were removed from the nest, temporarily replaced with clay eggs and returned to the nest later that day. The shells that had lined the nest were covered by wet sand and some water still remained in the nest. After a brief period, one of the adults returned to the nest and began to incubate and the second adult arrived at the site shortly thereafter. On 24 June, four chicks hatched after 26 days of incubation. The four chicks were considered to have fledged on 14 July. Case 2. On 3 June 1999, a nest at Cavendish Sands- pit (46.5028°N, 63.4322°W) was completed with four eggs. On 17 June, this nest was flooded and the eggs were found approximately | m from the nest in 10 cm of water. Both adults were present and appeared agi- tated. The real eggs were removed and clay eggs were placed in the nest. One severely cracked egg was dis- carded and the three undamaged eggs were returned to the nest once the water receded. One chick hatched on 30 June, and a second was observed on 1 July. The remaining egg was abandoned. Only one chick sur- vived to fledge and it was banded on 12 July. This chick was recaptured as a breeding male on the same beach in 2001. Case 3. On 17 June 1999, a nest with three eggs at Cavendish Sandspit (46.5047°N, 63.4483°W) was flooded without damage to the eggs. The eggs were found under 10 cm of water, more than | m from the nest. They were replaced with clay eggs. An adult was observed incubating the clay eggs approximately 1.5 hours later. The real eggs were then replaced and both adults returned within 30 minutes. Only one adult was observed on subsequent visits and the nest was considered abandoned on 20 June. The eggs were collected on 22 June, candled, found to be viable and placed in an incubator. The eggs were candled again on 30 June, when further growth and movement was observed in all 3 eggs. The eggs were successfully fostered to another Piping Plover pair on 3 July, but were taken by a predator on 4 July. The eggs from this nest were not only flooded, but endured a 24-48 hour lapse in incubation without any apparent decrease in viability. Case 4. A nest with three eggs was discovered on 12 June 2001 at Sebim Beach in Sand Hills Prov- incial Park, Nova Scotia (43.5311°N, 65.5580°W),. The full clutch remained on 25 June, but the high tide had completely overwashed the nest and the sand remained wet. Only two eggs remained on 11 July, and incubation was confirmed on 13 July. One aban- doned egg remained in the nest on 16 July. On 18 NOTES 449 July, a recently hatched chick ( or postal mail to Michael Coyne (online Editor) Marine Tur- tle Newsletter, | Southampton Place, Durham, North Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and Reptile Conservation Network — Reseau Canadien de Conservation des Amphbiens et des Reptiles 8(2) June 2004 Contents: Editor’s Note — Kawartha Turtle Trauma Centre: Now a Recognized Charity — 2993 Digital Frog -CARCNET/ RECCAR Scholarship Winner Kimberley Pearson (Sara L. Ashpole) — A Survey if the Status of the Western Toad (Bufo boreas) in Mount Revelstoke and Glacier National Parks (Jesse Dykstra) — (Other) Upcoming Meetings: 18" Annual Meeting of the Society for Conservation Biology (Danna Schock) — The 9 Annual Meeting of the Canadian Amphibian and Reptile Conservation Network (CARCNET)/ Reseau Canadien de Conservation des Amphibiens et des Canadian Species at Risk May 2004 Issued by the Committee on the Status of Endangered Wild- life in Canada (COSEWIC), the list is 49 pages containing: About COSEWIC: mandate, membership, definitions — Summary tables: Species designated in the “risk” and the Extinct categories and the Not at Risk, and Data Deficient categories (Tables 1-3) — Results of May 2004 COSEWIC meeting (Tables 4-5) — Explanation of status change sym- bols for reassessed species — COSEWIC Assessment Results — Table 6: Species assessed and designated Extinct: Extinct Category — Table 7: Species examined and designated in a “risk category”: Etripated Category, Endangered Category, Threatened Category, Special Concern Category — Table 8, Species assessed and found to be on the Not at Risk cate- gory — Table 9. Species considered and placed in the Data Reptiles RECCAR) Coast Terrace Inn, Edmonton, Alberta 24-27, 2004 — Seasonal and Diurnal Patterns of Calling in Eastern Canadian Amphibians: Compiling the conventional wisdom as a null hypothesis (Frederick W. Schueler with con- tributions from Wayne F. Weller, John Gilhen, Lenny Shirose, and Brian Dalzell — Membership in CARCNET/RECCAR (contact Bruce Pauli, Canadian Wildlife Service, National Wildlife Research Centre, Carleton University, Raven Road, Ottawa, Ontario K1A OH3. Web site: http://www.carcnet.ca/). Deficient category — Record of Status Re-examinations — Record of Name Changes. Listed as of May 2004 (with November 2003 totals in parenthesis) are 12 (12) extinct, 21 (21) extirpated, 169 (153) endangered, 114 (102) threatened, and 140 (143) of special concern. Of the 453 (431) forms in these categories, 66 (64) are mammals, 60 (56) birds, 32 (31) reptiles, 19 (19) amphibians, 79 (807) fishes, 16 (13) arthropods, 21 (18) molluscs, 145 (136) vascular plants, 11 (8) mosses, and 7 (6) lichens. In addition 151 (152) forms have been considered and found not at risk, and 33 (29) to be data deficient. This publication is available from COSEWIC Secretariat, Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. See Web site: http://www.cosewic.gc.ca. 484 2004 ERRATUM 485 Erratum Canadian Field-Naturalist 118(2): Back cover contents omission: Fifteenth census of seabird populations in the sanctuaries of the North Shore of the Gulf of St. Lawrence 1998-1999 JEAN-FRANCOIS RAIL AND GILLES CHAPDELAINE 256 Replacement Figure 2, page 161 for Morphology and population characteristics of Vancouver Island Cougars, Puma concolor vancouverensis, STEVEN F. WILSON, APRYL HAHN, AARON GLADDERS, KAREN M. L. Gon, AND DAviID M. SHACKLETON 4 @ Female 3 & Male 2 Males: y = 1.45Ln(x) - 0.54 u R?= 0.70 O | Ss 6 0 m Females: ® 4 Md @ y = 0.52Ln(x) - 1.09 R? = 0.34 -2 -3 0 2 4 6 8 10 Age (years) FIGURE 2. Relationship between body size (derived from the principal component analysis on morphological measurements, see text) and age for male and female Cougars in two Vancouver Island study areas. Logarithmic trend lines are presented for males (squares m) and females (diamonds ®) pooled by study area. The body size measurement was used because it was less condition-dependent than using total weight alone. Minutes of the 125 Annual Business Meeting of The Ottawa Field-Naturalists’ Club 13 January 2004 Place and time: Canadian Museum of Nature, Ottawa, Ontario, 7:30 p.m. Garry McNulty, President Thirty-five persons attended the meeting. Chairperson: Attendance: 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 7:35 p.m. with some opening remarks from Gary McNulty, the President. 1. Minutes of the Previous Meeting There were no changes to the minutes of the 124" Annual Business Meeting. It was moved by Fenja Brodo/David Hobden that the min- utes be accepted. (Motion Carried) 2. Business Arising from the Minutes Garry McNulty inquired whether we were still receiving the annual reports for the Trinidad and Tobago Naturalists’ Club. Frank Pope reported that we were receiving their annual reports along with their newsletters. The Soiree was revamped last year. Changes were made to the venue and entertainment was included in the course of the evening. Fenja Brodo, the new chair of the Excursions and Lectures Committee, will be working with the Education and Publicity Committee to continue the process of making the Soiree an evening that members will not want to miss. 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 end- ing September 30, 2003, noting that the Club’s net assets had decreased by approximately $60,000. This was due in large part to the contribution of $55,000 to the Alfred Bog Fund. Moved by Frank Pope and seconded by Roy John that the Financial Report be accepted. (Motion Carried) 5. Committee Reports Gary McNulty introduced each of the Committee reports and a representative of the appropriate Committee and asked for questions and comments. He thanked the committee chairs and committee members for their work over the past year. Moved by Mike Murphy/Eleanor Zurbrigg, that the reports be accepted. (Motion Carried) 6. Nomination of the Auditor Moved by Frank Pope, seconded by Bill Cody, that Janet Gehr continue as Auditor for another year. (Motion Carried) 7. Report of the Nominating Committee President Mike Murphy Vice President Gillian Marston Secretary Susan Laurie Bourque Treasurer Frank Pope Past President Gary McNulty Business Manager Bill Cody Editor CFN Francis Cook Editor T&L Karen McLachlan Hamilton Committee Chairs Birds Chris Traynor Computers Conservation Stan Rosenbaum E&P John Cameron E&L Fenja Brodo Finance Louise Schwartz FWG David Hobden Macoun rep Barbara Gaertner Membership Dave Smythe Publications Ron Bedford FON Rep Cendrine Huemer Members at large Kathy Conlan Diane Lepage Barbara Gaertner Christine Lewis Henry Steger Chairs not on Council Awards Ernie Brodo Macoun Rob Lee Nominations Fenja Brodo Retiring from the Council: = Charles Clifford, Roy John, Marcel Gahbauer, Eleanor Zurbrigg New on the Council: Christina Lewis, Henry Steger Moved by Fenja Brodo, seconded by Frank Pope, that the slate of nominations for the 2004 Council be accepted. (Motion Carried) 8. New Business There was no new business. 9. Presentation by David Hobden, ““FWG and its Rise to Fame” David Hobden presented an enjoyable and informative digital photographic presentation created by Henry Steger and 486 2004 David Hobden. The presentation touched on the develop- ment and flora and fauna of the Fletcher Wildlife Garden. The digital format successfully combined prints, slides and digital pictures. It was very well received. MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 487 10. Adjournment Fenja Brodo made an announcement about the upcoming 4"" Annual Great Backdoor Bird Count. Moved by Fenja Brodo/David Hobden that the meeting be adjourned at 9:15 pm. (Motion Carried) SUSAN LAURIE-BOURQUE, Recording Secretary The Ottawa Field-Naturalists’ Club Committee Reports for 2003 Awards Committee The Awards Committee met in January to consider nomi- nations made for the various OFNC Awards. As a result of our deliberations, the following four awards for the year 2002 were presented at the OFNC’s Annual Soirée, which took place 26 April 2003 at St. Basil’s Church in Ottawa. MEMBER OF THE YEAR: Bill Royds: For his many contributions to the Club’s con- servation efforts, especially during this last year as the link between the OFNC and the Greenspace Alliance of Canada’s Capital (GACC). GEORGE MCGEE SERVICE AWARD: David Hobden: For his active participation in the Con- servation Committee and his enthusiastic, productive work on the Fletcher Wildlife Garden Committee. CONSERVATION AWARD — MEMBER: Philip Fry: For establishing and maintaining the “Old Field Garden” near Oxford Mills as a model for habitat restoration and wildflower gardening. CONSERVATION AWARD — NON-MEMBER: Michéle André-St. Cyr: For her relentless work in pre- venting the death of turtles along the highways of southern Ontario by the establishment of “Turtle Crossing” signs, and for her active and successful efforts in educating the public about turtles. I. BRoDO Birds Committee The Birds Committee participated jointly with the Club des Ornithologues de l’Outaouais to run a successful Christmas Bird Count in December 2002. The fall bird count was held again in October 2003 and continues to grow in popularity. We were again active in the Peregrine Falcon Watch at the downtown nest site. One female was successfully fledged. The Bird Record Sub-committee continues to review records of rare birds and is currently compiling a lengthy database of Ottawa bird records. The Bird Record Sub-committee also finished publication of the new Ottawa District Bird Checklist 2002. The checklist is available in many Ottawa nature stores. The third year of surveying for the Ontario Breeding Bird Atlas (Region 24) has been completed. There are plans to add five more under-surveyed squares from an adjacent region for the 2004 season. This years seed-a-thon raised close to $400.00 for the club’s bird feeders. We con- tinue to operate the rare bird alert and the Ottawa bird status line, a recorded telephone message of current bird sightings. Some members of the Committee participated in the Taver- ner Cup competition C. TRAYNOR Computer Management Committee This year, the Committee reviewed and revised their Terms of Reference and the revisions were approved by the Council. It also reviewed the state of the Club’s Membership Data- base System application. Upon a recommendation by the Computer Management Committee, the Council approved the acquisition of Microsoft Access as the Club’s data entry and database management technology. This product provides a relational database engine with VisualBasic language sup- port for applications development. The Membership Database application will be redeveloped using Microsoft Access during 2004. M. MurpHy Conservation Committee Since November 2002 the CC has been assisting local residents calling themselves Larose Defense Fund/Fond de défense Larose with opposition to a proposed zoning change within part of the forest. We expressed support for the preser- vation of francophone heritage at other locations. The Club contributed $1000 towards the Fund’s legal costs at an OMB hearing. In cooperation with the Greenspace Alliance of Canada’s Capital, the CC appeared at City Planning and Development Committee in regard to the 3rd draft of the Official Plan. Some improvements were secured. We helped the FON to hold this workshop at the Museum of Nature in Ottawa. Several Club members reviewed the City’s list of areas to be studied and suggested additional areas. Adrianne Sinclair of the CC was appointed to the Public Advisory Committee. What in the 1970s the OFNC called the South Gloucester Conservation Area gradually became a quarry, but the 177 City-owned acres were known to support uncommon plant species. Despite intervention by the CC and the Greenspace Alliance, the City voted to sell this land to the quarry owner for $1.7M, but imposed requirements for environmental assessment, and assigned the sale revenue to a natural land acquisition fund. S. ROSENBAUM Education And Publicity Committee The committee organized and managed OFNC participa- tion in three major public events: the Ottawa Teachers Fed- eration professional development day, the Carlingwood Wild- life Festival and Health Canada’s celebration of Environment Week. The Wildlife Festival was the largest and most success- ful in attracting public interest. Twenty OFNC volunteers and five Macoun members assisted with this event. A demonstration of bird songs using tapes and slides was provided as a pre-concert activity for the Young Peoples Con- cert Series at the NAC, April 26. The Ottawa Field-Naturalists’ Club presented three Natural History Awards of $50.00 each plus an OFNC membership at the Ottawa Regional Science Fair. The judges were Kathy Conlan and Susan Laurie-Bourque. A French version of the club brochure was prepared and production begun at year’s end. 488 The sales table at the club’s evening presentations at the Canadian Museum of Nature provided revenue of $400. J. CAMERON Excursions And Lectures Committee In 2003 this Committee arranged 38 events and nine monthly meetings including the ABM and the Soirée. Seven general interest excursions were offered to Club members. The Birds Committee ran 12 birding excursions, three bird counts as well as assisted with The Taverner Cup. Two of our monthly meetings were also devoted to birds. There were six botanical excursions or workshops and two mushroom activities. Insects were the subject of one excursion and two of our monthly meetings. Other excursions were for mudpup- pies, astronomy and a museum visit. Workshops were also given on geology and on digital cameras. Two of our monthly meetings were devoted to conservation matters and one intro- duced us to polar environments. The monthly meetings drew about 50 people on average. Excursions and workshops are designed for a smaller number of participants. This year we asked leaders to have partici- pants sign a waiver form. The form is primarily designed to inform participants about liability but it provides a record of attendance. F. BRODO Executive Committee The Executive Committee met in October to review sey- eral items that required responses and/or action plans. As a result of that meeting, action plans have been put in place to address the following issues in 2004: (1) review of electronic publication possibilities (2) digital photo archives (3) how to retain new members (4) system to record names and number of persons parti- cipating in club excursions and activities. The Committee also reviewed the club’s liability insurance, its role in the Taverner Cup, the issue of attracting volunteers and it put in place a succession planning exercise and a sys- tem to review and report on the publications of other nature clubs. These matters require further action in 2004 but should be completed by year end. G. McNuLTy Finance Committee The Finance Committee met three times in calendar year 2003. The following items are highlighted: — a review of existing and proposed honoraria was undertaken. No changes to the current structure were recommended to Council. — the committee recommended to Council that the Alfred Bog Fund be topped up by $4,000 from the de Kiriline Lawrence Fund, bringing the total OFNC contribution to $59,000. — options for replacement accounting software were assessed but no decision was taken. — areview of membership fees for 2004 was carried out, and the Committee recommended to Council that the fees not be raised for the upcoming year. However, given the ex- pected year-end deficit, this issue will need serious con- sideration next year. — a budget for fiscal year 2003-04 was tabled and approved at Council in the fall, showing a small projected deficit. — other items such as the RoweCom bankruptcy, liability and directors’ insurance, and Taverner Cup funding were discussed. L. SCHWARTZ THE CANADIAN FIELD-NATURALIST Vol. 118 Fletcher Wildlife Garden Volunteers contributed over 3000 hours, mainly on Friday or Sunday mornings. Their work is particularly noticeable in the Backyard Garden. The Management Committee met eleven times during the year. The big achievement for the year was the production of an agreement with Agricultureand Agri-Food Canada for use of the site and occupation of the building. Negotiations were completed in the summer and the agreement finally signed in November. It is to run until April 2006. We wish to thank Frank Pope for handling legal and insurance issues and the final signing. During the year we held a public event for International Migratory Bird Day in May and our best plant sale ever in June thanks to Algonquin College and our own volunteers. Our traveling display was seen at Seedy Saturday, The Wildlife Festival, Environment Day at Health Canada, an Invasive Species Workshop at the Museum of Nature and at the OFNC Soiree. This year we had a part-time summer employee, since we did not obtain a grant to support the position as we had in previous years. She dealt with visitors, kept the Interpre- tation Centre open and worked mainly on invasive plant con- trol, supplementing the work of a group of volunteers. The Interpretation Centre was opened by volunteers on Sunday afternoons through the summer. Located there is a small library, including a complete collection of the Cana- dian Field-Naturalist, and two stereo microscopes for examin- ing small specimens. The Centre was used for meetings by OFNC Council and four committees and for workshops. D. HOBDEN Macoun Field Club Committee The committee met just once, to plan the start-up of the Macoun Club year; much of the work of organizing the pro- gram from month to month was done over the telephone and by e-mail. Some committee members arranged for speakers each week and supervised the meetings, while others led field trips twice a month. Some also led the high-school-age members on camping trips at Christmas and in June. Through this program, the children and teenagers of the Macoun Club were introduced to local geology, lichens, plant biology, insects, and wildlife (both Canadian and exotic). Field trips gave members hands-on experience in identifying trees and animal tracks, and allowed them to take part in the Club’s ongoing research on butternut canker disease, lichen bio- diversity, and porcupine populations. The monthly newsletter and the annually produced Little Bear magazine were both prepared by the young people them- selves, while the Club’s web site was maintained jointly with Committee members. Individual Macoun Club members pre- sented natural science projects at the OFNC Soirée in April, and the Club’s executive recounted the group’s activities for the OFNC members assembled there. R. LEE Membership Committee The distribution of memberships for 2003 is shown in the table (below), with the comparable numbers for 2002 in brackets. These statistics do not include the 23 affiliate orga- nizations which receive free copies of the Club’s publications. This year, the Club lost a long time member and valuable contributor to the production of the Canadian Field-Naturalist (CEN), with the death of Dr. Wilson Eedy, member since 1970 and Book-Review Editor of the CFN for many years. D. SMYTHE 2004 MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 489 | ° . . sae . | Distribution of Memberships in The Ottawa Field-Naturalists’ Club CANADIAN FOREIGN | Type Local Other USA Other Total _ Family 317 (340) 28 (29) | (1) | (2) 342 (372) Individual 323 (345) Le C20) 24 (26) 6 (5) 464 (496) Honorary 15 (15) 9 (9) 0 (0) 0 (0) 24 (24) Life 21 (21) 20 (21) 7 (5S) | (1) 49 (48) Sustaining 11 (8) 3 (2) 0 (O) 0) (QO) 1] (10) Total 687 (729) 166 (181) 32. (32) 8 (8) 890 (950) Publications Committee The Publications Committee met twice in 2003. _ Four issues of The Canadian Field-Naturalist were pub- — lished in 2003: Volume 116, #2,3,4 and Volume 117, #1. These four issues contained 698 pages; 47 articles; 20 notes; _ 1 COSEWIC article; 81 book reviews; 144 new titles; 2 _ commemorative tributes; 24 pages of News and Comments; and a 29 page index. The long-time Book Review Editor, Wilson Eedy, died sud- denly in the summer. His position has been taken over by Roy John. All of the Associate Editors will be reappointed for 2004. Six articles qualified for support from the Manning Mem- _ orial Fund for a total of $4044. Steady progress was made in getting the publication sched- ule back on track. The printing of The CFN has been taken from St. Joseph and contracted to Gilmore Printing Services Inc. beginning with Volume 117, #2. It is expected that this will both reduce the printing costs and improve the photo- graphic quality. In response to an author’s complaint, the details of pub- lication charges have been reworded. The new text says, in effect, that only members of The Ottawa Field-Naturalists’ Club and subscribers to The CFN are eligible to apply for waiver of page charges for the first five pages, and then only if institutional funds are unavailable. Volume 37 of Trail & Landscape was published in four issues containing 192 pages with the usual mix of good articles and news of Club events. R. BEDFORD 490 Auditor’s Report THE CANADIAN FIELD-NATURALIST 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, 2003, the statement of changes in net assets, and the statements of operations. These financial statements are the responsibility of the organization’s management. 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 ac- cepted auditing standards. Those standards require that I plan and perform an audit to obtain reasonable as- surance 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 raising activities. These revenues are not readily susceptible to complete audit verification, and accordingly, my veri- fication 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 concern- ing the completeness of the revenues referred to in the preceding paragraph, these financial statements present fairly, in all material respects, the financial position of the OFNC as at September 30, 2003, 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 January 12, 2004 Vol. 118 The Ottawa Field-Naturalists’ Club Balance Sheet September 30, 2003 2003 2002 ASSETS CURRENT Cash (Note 1) $ 2,781 $ 71,087 Investment certificates (Note 1) 43,764 84,790 Marketable securities (Note 2) 246,467 235,758 Accounts receivable 27,690 7,238 Prepaid expenses 1,000 1,000 321,702 399,873 CAPITAL ASSETS (Note 3) - _ Land — Alfred Bog 3,348 3,348 $ 325,050 $403,221 LIABILITIES AND FUND BALANCES CURRENT Accounts payable and accrued liabilities $ 2,000 $ 2,000 Deferred revenue 12,075 13,500 14,075 15,500 Life memberships 12,663 2A NET ASSETS Unrestricted 60,609 80,325 Club reserve 100,000 100,000 Manning principal 100,000 100,000 Manning interest - OFNC 1,434 1,963 - CFN 13,153 13,179 Seedathon 1,618 2,087 Anne Hanes memorial 870 870 de Kiriline-Lawrence 18,798 23,975 Macoun Baillie Birdathon 3 5857/ S27 Alfred Bog 493 52,274 298,312 376,000 $ 325,050 $403,221 2004 MINUTES OF THE 125" ANNUAL BUSINESS MEETING 49] The Ottawa Field-Naturalists’ Club The Ottawa Field-Naturalists’ Club Statement of Operations Canadian Field-Naturalists — Statement Of Operations For the Year Ended September 30, 2003 For the Year Ended September 30, 2003 2003 2002 2003 2002 REVENUE REVENUE Memberships $ 14,615 $ 15,488 Memberships $ 9,743 $ 10,325 Trail and Landscape 193 196 Subscriptions 25,369 28,483 Interest 1,339 1,457 Reprints 9,534 5,608 GST rebate 780 4,532 Publication charges 50,559 16,812 Other 617 760 Interest and exchange 7,940 10,404 17,547 22,433 GST rebate 3,040 3,448 SS a Other 734 339 OPERATING EXPENSES 106,919 75,419 Affiliation fees 680 680 Computer 302 216 EXPENSES Membership Syl 1,437 Publishing $ 80,758 $ 61,570 Office Assistant 1,000 1,000 Reprints 7,307 4,127 Telephone 1,622 1,848 Circulation 11,307 7,653 Insurance 655 655 Editing 4,183 2,652 Audit 1,000 1,000 Office Assistant 5,000 5,000 GST 1,290 136 Honoraria 9,000 9,000 Other 884 2,847 GST rebate 6,508 5,146 Tal RE: Suet 124,615 95,569 CLusB ACTIVITY EXPENSES ExcEss EXPENSES OVER Awards is 471 REVENUE $ (17,696) $ (20,150) Birds 1,696 223 ae Saupe Education and Publicity 73 1322 Excursions and lectures 496 164 Macoun Field Club 851 1,027 Soiree 19 466 Trail and Landscape 8,368 9,088 Fletcher Wildlife Garden (Note 4) 2,623 1,294 Other = _ 14,126 14,055 22,710 25,054 Excess EXPENSES OVER REVENUE $ (5,163) $ (2,621) 492 THE CANADIAN FIELD-NATURALIST The Ottawa Field-Naturalists’ Club Notes to the Financial Statements September 30, 2003 1. CASH Chequing Savings Nesbitt Burns Fletcher Wildlife Garden Investment Certificates: Maturity Maturity Value Date $ 44,385 04/29/04 2. MARKETABLE SECURITIES Investment Certificates: Maturity Maturity Value Date Province of Newfoundland Coupon $ 44,782 10/17/11 Province of Manitoba Coupon 29,847 11/15/04 Ontario Savings Bonds 40,000 06/21/05 CMHC Global Debs 52,000 12/01/06 Province of Ontario Bond 30,000 09/12/07 Province of Newfoundland Bond 20,000 10/07/08 Government of Canada Coupon 30,167 12/01/09 Province of New Brunswick Bond 20,000 06/15/10 3. CAPITAL ASSETS Equipment at a cost of $16,748 is fully amortized. 4. FLETCHER WILDLIFE GARDEN REVENUE Federal government City of Ottawa Taverner Cup Sales GST Donations Other EXPENSES Program Backyard Habitats Interpretation centre Administration Publications GST Library Other 2003 $ (3,777) 1,344 919 4,295 $ 2,781 Yield 1.00% Yield 4.525% 5.119% 6.45% 5.250% 6.125% 6.263% 5.605% 6.231% 243 132 6,346 $ (2,623) Vol. 118 ! 2002 $ 5,589 58,336 919 5,868 $ 70,712 Book Value $ 43,764 Book Value $ 31,421 26,754 41,370 53,539 31,187 20,538 20,590 21,068 $246,467 2002 $ 3,059 1,450 1,354 872 398 292 7,425 3,174 123 1,864 783 1,048 337 303 87 8,719 $ (1,294) 2004 MINUTES OF THE 125™ ANNUAL BUSINESS MEETING 493 The Ottawa Field-Naturalists’ Club Statement of Changes in Net Assets For the Year Ended September 30, 2003 (Note 5) Net Beginning Excess Excess Other Ending Assets Balance Expenses CFN Expenses OFNC Revenue Expenses Balance Unrestricted $ 80,325 $ (17,696) $ (5,163) $ 3,143 a $ 60,609 Club Reserve 100,000 - - - ~ 100,000 Manning Principal 100,000 = - - - 100,000 Manning — OFNC (a) 1,963 ~ - 1,021 1,550 1,434 Manning — CEN (b) 13,179 ~ ~ 4,086 4,112 13,153 Seedathon 2,087 - - 502 971 1,618 Anne Hanes Memorial 870 - - = ~ 870 de Kiriline-Lawrence (c) 23,975 _ - 381 5,558 18,798 Macoun Baillie Birdathon (d) 1.327 — - 10 - 1,337 Alfred Bog (e) 52,274 - - 3,219 55,000 493 $ 376,000 $ (17,696) $ (5,163) $ 12,362 $ 67,191 $ 298,312 STATEMENT OF CHANGES IN NET ASSETS a) Manning OFNC Expenses: Taverner Competition, $1,250; Peregrine Watch, $300; b) Manning CFN Expenses: Vol. 114, No. 2, Douglas Clark, Grizzly Bears in Northern Manitoba $228; Vol. 116, No. 1, Mark Hipfner et al., Common Eiders around Digges Sound $340; Gerald Kuzyk et al., Wolf response to sled dog $160; Kevin White et al., Predation of wolves on wolverines and martens $160; W. A. Fuller, Canada and the buffalo $1,609; Vol. 116, No. 3, John Goold et al., Sperm whale strandings $1,615. c) de Kiriline Lawrence Expenses: Nature Conservancy of Canada — land in Alfred Bog $4,000; Sierra Club, Ottawa Group — Leitrim Wetlands publicity — $1,000; Greenspace Alliance (Unger) — newspaper ad in defence of South Gloucester Conservation Area $558. d) Alfred Bog Expense: Nature Conservancy of Canada, $55,000 proceeds from fund raising campaign for acquisition of land in Alfred Bog. The Ottawa Field-Naturalists’ Club Summary of Significant Accounting Policies September 30, 2003 1. Nature of Business The organization is non-profit and incorporated under the laws of Ontario (1884). The organization promotes the appre- ciation, preservation, and conservation of Canada’s natural heritage. It encourages investigation and publishes the results of the research in all fields of natural history and diffuses information on these fields as widely as possible. It also sup- ports and cooperates with other organizations engaging in preserving, maintaining or restoring environments of high quality for living things. 2. Financial Instruments The organization’s financial instruments consist of cash, accounts receivable, marketable securities, and accounts pay- able. Unless otherwise noted, it is the management’s opinion that the organization is not exposed to significant interest, currency, or credit risks arising from these financial instru- ments. The fair value of these instruments approximate their carrying values. 3. Capital Assets Capital assets acquired after 1989 are expenses. Capital assets acquired prior to 1990 were recorded as assets at cost and amortized on a straight-line basis. These assets have been fully amortized. 4. Statement of Changes in Financial Position A statement of changes in financial position has not been provided as it would not provide additional meaningful infor- mation. 5. Foreign Currency Transactions during the year in U.S. dollars have been converted in the accounts to Canadian dollars at the exchange rate effective at the date of the transaction. All monetary assets in U.S. dollars at year end have been converted to Canadian dollars at the rate effective on Sept. 30, 2003. Gains or losses resulting therefrom are included in revenue or expenses. 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 read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, 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. Potential contributors who are neither members of The Ottawa Field-Naturalists’ Club nor subscribers to The Canadian Field-Naturalist are encour- aged to support the journal by becoming either members or subscribers. Manuscripts Please submit, to the Editor, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. AJ] authors should have read and approved it. Institutional or contract approval for the publication of the data must have been obtained by the authors. Also authors are ex- pected to have complied with all pertinent legislation regard- ing the study, disturbance, or collection of animals, plants or minerals. The place where voucher specimens have been de- posited, and their catalogue numbers, should be given. Lati- tude and longitude should be included for all individual local- ities where collections or observations have been made. Print the manuscript on standard-size paper, doublespace throughout, leave generous margins to allow for copy mark- ing, and number each page. For Articles and Notes provide a citation strip, an abstract and a list of key words. Gener- ally, words should not be abbreviated but use SI symbols for units of measure. The names of authors of scientific names may be omitted except in taxonomic manuscripts or other papers involving nomenclatural problems. “Standard” common names (with initial letters capitalized) should be used at least once for all species of higher animals and plants; all should also be identified by scientific name. The names of journals in the Literature Cited should be written out in full. Unpublished reports and web documents should not be cited here but placed in the text or in a sepa- rate Documents Cited section. List the captions for figures numbered in arabic numerals and typed together on a separate page. Present the tables each titled, numbered consecutively in arabic numerals, and placed on a separate page. Mark in the margin of the text the places for the figures and tables. Check recent issues (particularly Literature Cited) for journal format. Either “British” or “American” spellings are acceptable in English but should be consistent within one manuscript. The Oxford English Dictionary, Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Illustrations Photographs should have a glossy finish and show sharp contrasts. Electronic versions should be high resolution. Photo- graphic reproduction of line drawings, no larger than a standard page, are preferable to large originals. Prepare line drawings with India ink on good quality paper and letter (don’t type) descriptive matter. Write author’s name, title of paper, and figure number on the lower left corner or on the back of each illustration. Reviewing Policy Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who reviews it or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, cho- sen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for pub- lication, and in so doing aims to maintain the scientific quality, content, overall high standards and consistency of style, of the joumal. Special Charges — Please take note Authors must share in the cost of publication by paying $80 for each page, plus $15 for each illustration (any size up to a full page), and up to $80 per page for tables (depending on size). Authors may also be charged for their changes in proofs. Reproduction of color photos is extremely expensive; price quotations may be obtained from the Business Manager. If grant or institutional funds are not available, club members and subscribers may apply for a waiver of charges for the first five pages. Limited joumal funds are available to help offset publi- cation charges to authors with minimal financial resources. Requests for financial assistance should be made to the Busi- ness Manager when the manuscript is accepted. Reprints An order form for the purchase of repents will accompany the galley proofs sent to the authors. FRANCIS R. Cook, Editor RR 3 North Augusta, Ontario KOG IRO Canada 494 ——SaEaS==—_—_=>—=—>= TABLE OF CONTENTS (concluded) Volume 118 Number 3 } ik Sequential polyandry in Piping Plover, Charadrius melodus, nesting in eastern Canada DIANE L. AMIRAULT, JONATHAN KIERSTEAD, PETER MACDONALD, and LARRY MACDONNELL | American Dipper, Cinclus mexicanus, preys upon larval Tailed Frogs, Ascaphus truei CurisTy A. Morrissey and ROBERTA OLENICK Piping Plover, Charadrius melodus, egg viability after seawater immersion JULIE MCKNIGHT, LINDA THOMAS, and DIANE L. AMIRAULT Tribute 1A tribute to Loris Shano Russell, 1904-1998 KEVIN L. SEYMOUR |Book Reviews |ZooLocy: The Uses and Curation of Bird’s Egg Collections: An Examination and Bibliography — Birds of | Nunavut: A Checklist — Birds of the Untamed West: The History of Birdlife in Nebraska, 1750 to 1875 — Birds of Nebraska: Their Distribution and Temporal Occurrence — Canadian Skin and Scales: A Complete Encyclopedia of Canadian Amphibians and Reptiles — Conservation and Ecology of Turtles of the Mid- Atlantic Region: A Symposium — For Love of Insects — The Freshwater Fishes of Manitoba — The History of Ornithology in Virginia — All-Weather Hawk Watcher’s Field Journal —- Self-Portrait with Turtles: A Memoir — Wild Mammals of North America: Biology, Management, and Conservation (Second edition) — Pete Dunne on Bird Watching: The How-to, Where-to and When-to of Birding -|Borany: Alpine Plants of North America: An Encyclopaedia of Mountain Flowers from Rockies to Alaska — Arboretum America, a Philosophy of the Forest — Cape Cod Wiidflowers: A Vanishing Heritage — Lewis Clark’s Field Guide to Wild Flowers of the Sea Coast in the Pacific Northwest — Wild Flowers of Field & Slope in the Pacific Northwest — Wild Flowers of Forest & Woodland in the Pacific Northwest Wild Flowers of the Mountains of the Pacific Northwest ENVIRONMENT: Conserving Living Natural Resources in the Context of a Changing World — Visions of the Land: Science Literature, and the American Environment from the Era of Exploration to the Age of Ecology — Genetic, Demography and Viability of Fragmented Populations — A Primer of Ecological Genetics MISCELLANEOUS: Smithsonian Institution Secretary Charles Doolittle Walcott NEw TITLES |News and Comment Froglog: Newsletter of the Declining Amphibian Populations Task Force (63, 64) — Marine Turtle Newsletter (105) — The Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian Amphibian and Reptile Conservation Network - Reseau Canadien de Conservation des Amphbiens et des Reptiles 8(2) June 2004 — Canadian Species at Risk May 2004 _|Errata: Canadian Field-Naturalist 118(2) Omission from the back cover contents: Rail and Chapdelaine Replacement figure for Wilson et al. page 161 Minutes of the 125 Annual Business Meeting of The Ottawa Field-Naturalists’ Club Tuesday 13 January 2004 | Advise to Contributors to The Canadian Field-Naturalist Mailing date of the previous issue 118(2): 10 May 2005 2004 444 446 448 45] 465 474 477 481 483 484 485 486 494 THE CANADIAN FIELD-NATURALIST Volume 118 Number 3 Articles Population dynamics of Deer Mice, Peromyscus maniculatus, and Yellow-pine Chipmunks, Zamias amoenus, in old field and orchard habitats THOMAS P. SULLIVAN, DRUSCILLA S. SULLIVAN, and EUGENE J. HOGUE Density and survival of lady beetles (Coccinellidae) in overwintering sites in Manitoba W. J. TURNOCK and I. L. WISE Description of age-O Round Goby, Neogobius melanostomus Pallus (Gobiidae) and ecotone utilization in St. Clair lowland waters, Ontario JOHN K. LESLIE and CHARLES A. TIMMINS A systematic analysis of the alpine saxifrage complex (Saxifragaceae) in the Canadian Arctic Islands using morphology and chloroplast DNA data CAROLINE HEALY and LYNN J. GILLESPIE The influence of prey availability and vegetation characteristics on scent station visitation rates of Coyotes in a heterogeneous environment LYNDA A. RANDA and JOHN A. YUNGER Presence of cavities in snags retained in forest cutblocks: Do management policies promote species retention? Kim T. EVERETT and KEN A. OTTER Summer diet of two White-tailed Deer, Odocoileus virginianus, populations living at low and high density in southern Quebec CLAUDE DAIGLE, MICHAEL CRETE, LOUIS LESAGE, JEAN-PIERRE OQUELLET, and JEAN HUOT Winter habitat use by Wolves, Canis lupus, in relation to forest harvesting in west-central Alberta GERALD W. KUZYK, JEFF KNETEMAN, and FIONA K. A. SCHMIEGELOW Bird communities of the Garry Oak habitat in southwestern British Columbia WAYNE R. ERICKSON Invertebrate diversity under artificial cover in relation to boreal forest habitat characteristics STEPHEN H. FERGUSON and DANIELLE K. A. BERUBE Observations of habitat use by Polar Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, and northern Bering Seas DONALD J. HANSEN New records for the Arctic Shrew, Sorex arcticus, and the newly recognized Maritime Shrew, Sorex martimensis NEIL D. PERRY, DONALD T. STEWART, ELIZABETH M. MADDEN, and THOMAS J. MAIER Lichen zonation on coastal rocks on Gwaii Haanas National Park Reserve, Haida Gwaii (Queen Charlotte Islands), British Columbia IRWIN M. BRobo and Norm A. SLOAN Predaceous water beetles (Coleoptera: Adephaga: Dysticidae, Gyrinidae) collected along the Horton and Thelon rivers in the Arctic Central Barrens of Canada HELENA V. SHAVERDO and DONNA J. GIBERSON Notes Collapsing burrow causes death of a Eurasian Beaver, Castor fiber Liat R. THOMSEN, FIONA SHARPE, and FRANK ROSELL Frequency of tail breakage of the Northern Water Snake, Nerodia sipedon sipedon | KENNETH D. BOWEN The Heather Vole, genus Phenacomys, in Alaska S. O. MACDONALD, AMY M. RUNCK, and JOSEPH A. COOK Extension de Vaire de distribution connue de la Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du Quebec JEAN-FRANCOIS DESROCHES and ISABELLE PICARD Gulls, Larus spp., foraging at Pink Salmon, Oncorhynchus gobuscha, spawning runs — MAry F. WILLSON (continued on inside back cover) ISSN 0008-3550 2004 368 376 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada re Ave Volume 118, Number 4 October—December 2004 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 information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining 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 Dan Strickland Francis R. Cook C. Stuart Houston Eugene G. Munroe John B. Theberge Ellaine Dickson George F. Ledingham Robert W. Nero Sheila Thomson 2004 Council President: Mike Murphy Ronald E. Bedford Barbara Gaertner Stanley Rosenbaum Vice-President: Gillian Marston Fenja Brodo Diane Lepage Louise Schwartz Recording Secretary: Susan Laurie-Bourque John Cameron Christina Lewis David Smythe Treasurer: Frank Pope William J. Cody Karen McLachalan Hamilton Henry Steger Past President: Gary McNulty Kathy Conlan David Hobden Chris Traynor Francis R. Cook Cendrine Huemer To communicate with the Club, address postal correspondence to: The Ottawa Field-Naturalists’ Club, 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 www.ofnc.ca 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. PAP Registration Number 9477. We acknowledge the financial support of the Government of Canada through the Publication Assistance Program (PAP) toward our mailing costs. a Canada Editor: Dr. Francis R. Cook, R-R. 3, North Augusta, Ontario KOG IRO; (613) 269-3211; e-mail: cfn@ofnc.ca Copy Editor: Elizabeth Morton Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada KIZ 1A2; (613) 759-1374 Book Review Editor: Roy John, 2193 Emard Crescent, Ottawa, Ontario K1J 6KS5, e-mail: roy.john@pwgsc.ge.ca 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 except Book Reviews should be addressed to the Editor and sent by postal mail. Book-review correspondence should be sent by e-mail or postal mail to Roy John, 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 members (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: April-June 2004 (April 2005). Cover: Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, growing on Jack Pine, Pinus banksiana, Growth in Manitoba. Photograph by Jacques C. Tardif. See article by Epp and Tardif, pages 595-601. MCZ LIBRARY APR 0 3 2006 ARVARD October-December 2004 The Canadian Field-Naturalist Volume 118, Number 4 Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the Northern Gulf of St. Lawrence, Canada LENA Measures!”, BENOIT ROBERGE?, and RICHARD SEARS® ' Fisheries and Oceans, Maurice Lamontagne Institute, 850 Route de la Mer, Mont-Joli, Québec GSH 324 Canada. (Author to whom correspondence should be addressed.) 2 Parks Canada Agency, Mingan Archipelago National Park Reserve of Canada, 1303 de la Digue, Havre-Saint-Pierre, Québec GOG 1P0 Canada 3 Mingan Island Cetacean Study, Inc., 625, rue du Centre, Longue-Pointe-de-Mingan, Québec GOV 1V0 Canada Measures, Lena, Benoit Roberge, and Richard Sears. 2004. Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the Northern Gulf of St. Lawrence, Canada. Canadian Field-Naturalist 118(4): 495-498. A Pygmy Sperm Whale, Kogia breviceps, stranded alive and later died in the Northern Gulf of St. Lawrence, Quebec, Canada on 28 August 2001. This is the northern-most stranding of this species in the western Atlantic. The whale was estimated to be approximately 3 m long and a longitudinal section from one tooth (31 mm long and 5.0 mm in diameter) revealed 3.5 growth layer groups. ~ Key Words: Pygmy Sperm Whale, Kogia breviceps, Gulf of St. Lawrence, Quebec, Canada. A Pygmy Sperm Whale (Kogia breviceps) was found dead 28 August 2001 on the north shore of the Gulf of St. Lawrence. It was observed alive on 27 August by a local resident, Mme Rose-Annette Blais, northwest of St. Charles Island in the Mingan Archi- pelago National Park Reserve. The whale was observed making slow, undulating movements in shallow water close to shore in Trilobites Bay (50°14'07"N; 63°21' 18"W) and was found dead the next morning on a sandy beach in the bay. Personnel from Parks Canada were alerted and arrived on site to examine the carcass. The whale was estimated to be approximately 3 m long (Figure 1). There were various wounds and lacerations on the rostrum, flanks and abdomen (Figure 2). Two teeth (one broken near the root) collected by Jacques A. Thériault and his father, were later submitted for analysis. Fisheries and Oceans Canada and the Min- gan Island Cetacean Study, Inc. were informed of the stranding but before the carcass could be collected for necropsy it had been carried off by a rising tide and was not seen again. Both teeth were deposited in the Maurice Lamontagne Institute collection, Acces- sion Number 11629. The whale was identified as a Pygmy Sperm Whale based on photographs, approximate length (3 m), the low, falcate dorsal fin located posterior to the center of the dorsum (Figures 1, 2) and size of one intact tooth. The tooth was 31 mm long and 5.0 mm in diam- eter at its widest point. In the dentine and cementum layers there were 3.5 growth layer groups (GLG) observed in a longitudinal section of the tooth. It is unknown how many GLGs are laid down per year in Pygmy Sperm Whales. If similar to the Sperm Whale Physeter macrocephalus [1 GLG = 1 year of age (C. Lockyer, personal communication; Perrin and Myrick 1980)], then this animal is almost 4 years old and likely an adult (Handley 1966). Ross (1984) reported 3.5 GLGs in a sexually mature female that stranded with a calf. The Pygmy Sperm Whale can be confused with the Dwarf Sperm Whale (Kogia simus). The latter is smaller (2.7 m maximum length) with a taller dolphin- like dorsal fin and teeth are less than 30 mm long and less than 4.5 mm in diameter (Handley 1966; Leather- wood and Reeves 1983). Pygmy Sperm Whales are rarely observed in eastern Canadian waters and have no COSEWIC (Committee on the Status of Endangered Wildlife in Canada) status (Baird et al. 1996). Seven, probably eight, Pygmy Sperm Whale strandings have been reported prior to the present report (Piers 1923; Sergeant et al. 1970: Nelson et al. 1991; McAlpine et al. 1997; Lucas and Hooker 2000) in Canada over the last century, specif- ically in Halifax and on Sable Island in Nova Scotia, in Blacks Harbour and Saint-John’s, New Brunswick and on the French island of Miquelon near the south- 495 496 THE CANADIAN FIELD-NATURALIST Vol. 118 pau FiGure |. Pygmy Sperm Whale beach-cast in Trilobites Bay on 27 August 2001. Length is approximately 3 m. west coast of Newfoundland. The present report is the northern-most stranding of this species in the western Atlantic and the first within the Gulf of St. Lawrence. The Pygmy Sperm Whale is a small pelagic, mainly deep-water odontocete (occasionally seen motionless in surface waters) found widely offshore on continen- tal slopes throughout tropical and warm temperate waters in the world. Most information on this species has been obtained from beach-cast carcasses and re- cently from live-stranded and rehabilitated animals in- cluding mother-calf pairs or pregnant females (Hiick- stadt and Antezana 2001; Scott et al. 2001; Manire et al. 2002). Stomach analyses indicate that the Pygmy Sperm Whale feeds primarily on cephalopods, crus- taceans (shrimp, crab) and fish (Reeves et al. 2002). In the western Atlantic most strandings have been reported off the southeastern coast of the United States (Caldwell and Caldwell 1989; Odell 1991) and live individuals have been observed beyond the continental shelf and in the Gulf Stream. Recently, a rehabilitated Pygmy Sperm Whale equipped with a time-depth recorder was released in the Gulf Stream off the eastern coast of the United States. It was observed for 5 days and remained east of the continental shelf break (200 m isobath) but west of the eastern edge of the Gulf Stream where sea- surface temperatures were 27.8 — 30.0°C (Scott et al. 2001). Unfortunately, dive depth information was not obtained and it is unclear whether the behaviour of this individual reflects that of the species in general. Odell et al. (1985 as cited in Caldwell and Caldwell 1989) noted that more strandings of Kogia (K. brevi- ceps and K. simus) occur in Florida when the Gulf Stream shifts farther offshore. They suggested that indi- viduals following prey caught in Gulf Stream eddies or rings that suddenly dissipate may become disori- ented and, subsequently, strand. Gulf Stream rings can occur to the north or south of the Gulf Stream but anticyclonic or warm-core rings which are about 1000 m deep and exist on average 4.5 months, drift southwestward to rejoin the Gulf Stream near Cape Hatteras (Richardson 1976; Wiebe 1976). Gulf stream rings have unique physical, chemical and biological characteristics (Wiebe 1976; Joyce and Wiebe 1983; Craddock et al.1992). Although warm-core rings do not ride onto shallow continental shelves, they can push slope water onto the shelf or entrain shelf water into slope waters. Such shelf water entrainment was suggested as providing suitable cephalopod habitat which attracted Sperm Whales in the vicinity of a warm-core ring off Georges Bank (Griffin 1999). The occurrence of a Pygmy Sperm Whale in the Gulf of St. Lawrence, especially in the northern Gulf, seems unusual as water temperatures are relatively cold [sea-surface temperatures are generally 12 — 18°C in August (Vigeant 1987*)]. The minimum temperature 2004 MEASURES, ROBERGE, AND SEARS: STRANDING OF PYGMY SPERM WHALE 497 FiGuRE 2. Pygmy Sperm Whale in shallow water in Trilobites Bay (same specimen as in Figure 1). Note relative position of dorsal fin on trunk. tolerated by Pygmy Sperm Whales is unknown. Annu- al mean air temperatures in most of the northwest At- lantic, especially in the southern Gulf of St. Lawrence, and sea-surface water temperatures throughout eastern Canadian waters in 2001 were generally warmer than normal (Drinkwater et al. 2002a*). In some areas, the upper 30 m in the central and southern Gulf of St. Lawrence show monthly anomalies of 3 - 4°C (Drink- water et al. 2002b*; Gilbert 2002*). The Pygmy Sperm Whale may have swam and drifted northeast follow- ing prey in the Gulf Stream as suggested by Fraser (1974), perhaps using warm-core rings as “stepping- stones” (Peter Wiebe, personal communication) or thermal fronts associated with warm-core rings as observed by Griffin (1999). It may have entered the Gulf of St. Lawrence in warm surface waters on the Cape Breton side of the Cabot Strait when the Cape Breton Current slows in summer (El-Sabh 1977). Dur- ing a DFO research cruise in the Gulf of St. Law- rence in August 2001 squid (/lex illecebrosus) were collected but numbers of specimens collected were not considered exceptional (D. Archambault, DFO, unpublished data). Acknowledgments We thank R-A. Blais and J. A. Thériault for assis- tance in this stranding and providing teeth. We also thank S. Plamondon of Parks Canada who provided photographs and J.-G. Gosselin of Fisheries and Oceans Canada who determined the number of GLGs by cut- ting the tooth. We appreciate that P. Best and J. Mead examined the photographs and concurred with our identification of the species. Documents Cited (marked * in text) Drinkwater, K. F., R. G. Pettipas, and W. M. Petri. 2002a. An overview of meteorological, sea ice and sea-surface temperature conditions off Eastern Canada during 2001. Canadian Science Advisory Secretariat Research Docu- ment 2002/048. Drinkwater, K. F., R. G. Pettipas, and W. M. Petri. 2002b. Physical environmental conditions in the Southern Gulf of St. Lawrence during 2001. Canadian Science Advisory Secretariat Research Document 2002/047. Gilbert, D. 2002. Oceanographic conditions in the Gulf of St. Lawrence in 2001 : physical oceanography. DFO Sci- ence Stock Status Report. 498 Vigeant, G. 1987. Température mensuelle de ]’eau en surface dans l’estuaire et le Golfe du Saint-Laurent. Service de 1’ Environnement Canada. Literature Cited Baird, R. W., D. Nelson, J. Lien, and D. W. Nagorsen. 1996. The status of the pygmy sperm whale, Kogia brevi- ceps, in Canada. Canadian Field-Naturalist 110: 525-532. Caldwell, D. K., and M. C. Caldwell. 1989. Pygmy sperm whale Kogia breviceps (de Blainville, 1838): dwarf sperm whale Kogia simus Owen, 1866. pages 235 — 260 in Hand- book of marine mammals. Edited by S. H. Ridgway and R. Harrison. Academic Press, London. Craddock, J. E., R. H. Backus, and M. A. Daher. 1992. Vertical distribution and species composition of midwater fishes in warm-core Gulf Stream meander/ring 82-H. Deep Sea Research 39: S203-S218. El-Sabh, M. I. 1977. Oceanographic features, currents, and transport in Cabot Strait. Journal of the Fisheries Research Board of Canada 34: 516-528. Fraser, F. C. 1974. Report on Cetacea stranded on the British coasts from 1948 to 1966. British Museum (Natural History) 14. Griffin, R. B. 1999. Sperm whale distributions and commu- nity ecology associated with a warm-core ring off Georges Bank. Marine Mammal Science 15: 33-51. Handley, C. O., Jr. 1966. A synopsis of the genus Kogia (pygmy sperm whales), pages 62 — 69 in Whales, dol- phins and porpoises. Edited by K. S. Norris. University of California Press, Berkeley and Los Angeles, Calif. Hiickstaédt, L., and T. Antezana. 2001. An observation of parturition in a stranded Kogia breviceps. Marine Mammal Science 17: 362-365. Joyce, T., and P. Wiebe. 1983. Warm-core rings of the Gulf Stream. Oceanus 26: 34-44. Leatherwood, S., and R. R. Reeves. 1983. The Sierra Club handbook of whales and dolphins. Sierra Club Books, San Francisco. Lucas, Z. N., and S. H. Hooker. 2000. Cetacean strandings on Sable Island, Nova Scotia, 1970 — 1998. Canadian Field- Naturalist 114: 45-61. Manire, C. A., L. Byrd, H. L. Rhinehart, P. Cunningham- Smith, and D. R. Smith. 2002. Subacute atropine toxicity in a pygmy sperm whale, Kogia breviceps. Journal of Zoo and Wildlife Management 33: 66-72. THE CANADIAN FIELD-NATURALIST Vol. 118 McAlpine, D. F., L. D. Murison, and E. P. Hoberg. 1997. New records for the pygmy sperm whale, Kogia breviceps (Physeteridae) from Atlantic Canada with notes on diet and parasites. Marine Mammal Science 13: 701—704. Nelson, D., A. Desbrosse, J. Lien, P. Ostromm, and R. Seton. 1991. A new stranding record of the pygmy sperm whale, Kogia breviceps, in waters off Eastern Canada. Canadian Field-Naturalist 105: 407-408. Odell, D. K. 1991. A review of the Southeastern United States Marine Mammal Stranding Network: 1978 — 1987. pages 19 — 23 in Marine Mammal Strandings in the United States. Edited by J. E. Reynolds II and D. K. Odell. NOAA Technical Report NMFS 98. Perrin, W. F., and A. C. Myrick, Jr. Editors. 1980. Growth of odontocetes and sirenians: problems in age determina- tion. Reports of the International Whaling Commission Special Issue 3: 1-229. Piers, H. 1923. Accidental occurrence of the pygmy sperm whale (Kogia breviceps) on the coast of Nova Scotia; an extension of its known range, with remarks on the proba- bility of the former presence in these waters of the true sperm whale (Physeter macrocephalus). Proceedings and Transactions of the Nova Scotia Institute of Science 15: 95-114. Reeves, R. R., B. S. Stewart, P. J. Clapham, and J. A. Powell. 2002. National Audubon Society guide to marine mammals of the world. Alfred A. Knopf, New York. Richardson, P. 1976. Gulf Stream rings. Oceanus 19: 65-68. Ross, G. J. B. 1994. The smaller cetaceans of the south east coast of southern Africa. Annals of the Cape Provincial Museums (Natural History) 15: 173-410. Scott, M. D., A. A. Hohn, A. J. Westgate, J. R. Nicolas, B. R. Whitaker, and W. B. Campbell. 2001. A note on the release and tracking of a rehabilitated pygmy sperm whale (Kogia breviceps). Journal of Cetacean Research Man- agement 3: 87- 4. Sergeant, D. E., A. W. Mansfield, and B. Beck. 1970. Inshore records of cetacea for eastern Canada, 1949 — 68. Journal of the Fisheries Research Board of Canada 27: 1903-1915. Wiebe, P. 1976. The biology of cold-core rings. Oceanus 19: 69-76. Received 14 July 2003 Accepted 15 July 2004 Seed Dispersal by Brown Bears, Ursus arctos, in Southeastern Alaska Mary FE. WILLSON! and Scott M. GENDE? '5230 Terrace Place, Juneau, Alaska 99801 USA. National Park Service, Glacier Bay Field Station, 3100 National Park Road, Juneau, Alaska 99801 USA Willson, Mary F., and Scott M. Gende. 2004. Seed dispersal by Brown Bears, Ursus arctos, in southeastern Alaska. Canadian Field-Naturalist 118(4): 499-503. Mammals often consume fleshy fruits and disperse significant quantities of the enclosed seeds. In southeastern Alaska, Brown Bears (Ursus arctos) are among the most important dispersers of seeds for the numerous plant species producing fleshy fruits, because these bears are abundant, often eat large quantities of fruit, and commonly excrete seeds in germinable condition. Scat analyses showed that Brown Bears on Chichagof Island ate increasing quantities of fruit through summer and fall. Scats commonly contained several thousand seeds, often of two or more species. Four kinds of seeds of fleshy- fruited plants that normally grow in forest understory germinated at similar levels when experimentally deposited (in bear scats) in the two most common habitats (forest and muskeg), suggesting that habitat distribution of these plants is not deter- mined simply by germination patterns. Although seed passage through bear digestive tracts and the composition of scats are known to affect germination rates to some degree, the most important role of bears in seed dispersal is probably transport. Key Words: Brown Bears, Ursus arctos, Southeastern Alaska, seed dispersal, fleshy fruits, Rubus, Ribes, Oplopanax, Vaccinium, Streptopus. Coastal rainforest ecosystems in southeastern Alaska include many shrubs and herbs with fleshy fruits, commonly consumed by vertebrates that disperse the enclosed seeds (Willson 1991; Traveset et al. 2004). Potential dispersal agents include corvids, thrushes, warblers, waxwings, Pine Marten (Hickey et al. 1999) and bears (Willson 1993, 1994). Bears are important dispersers of seeds, because they can consume large quantities of fruit (Welch et al. 1997; Farley and Rob- bins 1995) and often range over considerable dis- tances (Patten 1993), distributing seeds far from the source. Furthermore, bears are numerous in south- eastern Alaska, with density reaching several hundred bears per 1000 km — perhaps as many as 700 or even more in some areas (Miller et al. 1997; J. Whitman, personal communication). The abundance of Brown Bears (Ursus arctos) in southeastern Alaska provides an opportunity to examine the consequences of bear frugivory for seed dispersal. The purpose of this paper is to describe the phenol- ogy of Brown Bear frugivory, patterns of scat deposition on a small spatial scale, seed abundance and composi- tion in bear scats, and field tests of seed germination from bear scats. We then integrate this information with previous reports, in order to describe what is known about this ecologically important interaction in southeastern Alaska. Study Area and Methods Field work was conducted on northern Chichagof Island in the Alexander Archipelago, at the southern end of Port Frederick (approximately 58°N, 135°30'W). The only bears on this island are Brown Bears. The rainforest is composed chiefly of Sitka Spruce (Picea sitchensis) and hemlock (Tsuga spp.), with occasional muskegs (bogs) of Sphagnum moss, scattered Lodge- pole Pines (Pinus contorta), and low shrubs. Brown Bear scats were collected from abandoned logging roads, at less than 300 m elevation, in the Neka River valley and near Salt Lake Bay on opposite sides of Port Frederick. During each road survey, all scats were cleared from the road, so that the subsequent survey recorded only scats deposited in the between-survey interval. The roads passed through forests of differing ages, clearcuts, and muskegs. Seed composition of scats The frequency of occurrence of seeds, and the prin- cipal kinds of seeds present, were recorded in June- September 1990-1993 in 4378 scats. A subset of 570 seed-containing scats was collected in August and September 1990 and 1991, for quantitative estimates of seed abundance. Each of these scats was mixed to distribute the seeds throughout, and a subsample aver- aging 24% + 2% (S. E.) of the total scat mass was taken. In the weighed subsample, the seeds were iden- tified to genus and counted; the number of seeds in the entire scat was then estimated by extrapolation. The frequency of seeds in scats through the summer and early fall was examined, using | June as Day 1. June scat collections represent the previous spring months; collections in July, August, and September represent only the preceding 30 days, approximately. Seed germination In order to compare seed germination in two prin- cipal habitats, over 150 of the scats for which total seed content was estimated were placed in two common habitats (muskeg or forest). Scats were “planted” in 499 500 transects in each habitat, in whatever microhabitats were available at about 2 m intervals. Muskeg and for- est transects near Salt Lake Bay were established in late summer, 1990, and in the Neka Valley in 1991. Seedlings were counted in the planted scats annually (in August) for three (Neka) or four (Salt Lake Bay) years. For seedling censuses, each planted scat was covered with a grid, and seedlings were counted in a random selection of the grid squares. The proportion of grid squares sampled usually ranged from 20% to about 50% of the total scat. Emerging seedlings were counted individually when possible, but so many seeds germinated in some squares that it was impossible to count them without destructive sampling, in which case the abundance of seedlings was estimated visually. The total number of emerging seedlings per scat was extrapolated from this subsample. Curious bears and heavy fall rains disturbed some of the planted scats and thus reduced the usable sample size slightly. Because it was impossible to mark individual seedlings in the very dense clusters, the data are simply estimates of annual germination, not of survivorship of seedlings. Mortality of seedlings was very high and only a few survived more than one year. We did not observe rodent and bird predation on seeds in the experimental tran- sects, although seed predation was sometimes evident in scats on the road system. The estimated percent of seeds germinating each year was compared between habitats using a repeated- measures ANOVA on arc-sine transformed data (PROC GLM, SAS Systems Inc., Cary, North Carolina). Because the power of this test was low (two sites, two habitats), our results were re-examined, to see if the outcome was merely a result of the few degrees of freedom in the ANOVA. For this we used a t-test (in which the planted scats were the replicates, so there were more degrees of freedom in the test) to test for differences in seedling germination between habitats for the first and the last years at each site (no seedlings in the censused grid squares survived for the entire length of the experiment). Results Phenology of fruit consumption Scats collected in early summer contained mostly vegetable fiber and occasional deer remains (hair, bone, hoof) with few seeds, as expected, because few fruits are available at that time of year (some overwintered THE CANADIAN FIELD-NATURALIST Vol. 118 ce 100 - A i. ® e (3) @ 0” go 4 ro}) ° e = = 60 = e 5 qo 404 vg e © ® 8 20 + . re) AS 0 & T i T T {1-Jun 30-Jun 29-Jul 27-Aug 25-Sep day Ficure |. Seasonal changes, for four years, in the percent of bear scats containing seeds, Chichagof Island, south- eastern Alaska. R, = 0.79, p < 0.001. Viburnum and Maianthemum; some early Rubus spec- tabilis). The frequency of seed-containing scats in- creased through the summer, from less than 1% in mid-June to over 90% in late August and early Sep- tember (R, = 0.79, p < 0.001, n = 12; Figure 1). Scat distribution at a small spatial scale In addition to widespread deposition of scats on the landscape (Patten 1993), defecations by a moving bear can spread multiple deposits of a single defecation over several meters, which could be ecologically important. A survey of 1163 “scat trails” revealed that 42% were multiple deposits, with 3% of them comprised of over 20 separate droppings spread over as much as 84 m. Seed composition in scats Seeds were identified at least to genus; in some cases, the genus was represented by only one species in this area (Sambucus racemosa, Rubus spectabilis, Oplopanax horridum). In the case of Ribes, the species was most likely bracteosum, which grew commonly in this area. Likewise, Streptopus amplexifolius was far more common in this area than two congeners. Both Vaccinium ovalifolium and V. alaskense were present and were not distinguishable in the scats. The most commonly occurring seeds were Oplopanax, Vaccinium, Ribes, Rubus, and Streptopus (Table 1). Seeds had a high probability of being deposited not only with numerous conspecifics but also with at TABLE 1. Summary of frequency of occurrence of common kinds of seeds in scats on northern Chichagof Island. Data are the number of months that the frequency of occurrence equaled or exceeded the stated value. Kind of seed Vaccinium Ribes Oplopanax Rubus Streptopus Number of months = 20% 5 5) 1 1 3 ia “8 Number of months = 50% 3} 2004 WILLSON and GENDE: SEED DISPERSAL BY BROWN BEARS TABLE 2. Average (S.E.) number of seeds per scat on northern Chichagof Island. N = 570 scats. Year Month Vaccinium Ribes 1990 July 3581 (667) 914 (767) August 4655 (461) 700 (165) September 2534 (468) 1918 (281) 199] July 4375 11(933) 2.02817 (-) August 7193 (760) 364 (109) September 4329 (795) 1302 (293) 4445 1971 x number of seeds/fruit* 47 12 x number of fruits 95 89 *from Traveset et al. 2004. least one other kind of seed: in monthly samples, a median of 60% (range 16-87%) of scats contained at least two kinds of seeds, and sometimes as many as five kinds. Vaccinium and Ribes were consistently the most abundant seeds in scats collected (Table 2). In most months, Rubus was next most abundant, followed by Oplopanax and others. The maximum numbers per scat for each of the genera listed in Table 2 were approximately 37 000 Vaccinium, 14 500 Ribes, 7600 Rubus, 2000 Oplopanax, 1900 Sambucus, and 400 Streptopus. In addition to those represented in Table 2, other kinds of seeds appeared in the scats sporadical- ly (from fleshy fruits, Viburnum, Maianthemum; non- fleshy fruits, Kalmia, Heracleum, Carex, grass). Because the number of seeds per fruit differs greatly among genera (Traveset et al. 2004), the approximate number of consumed fruits of each type per scat differs markedly from the abundances of seeds (Table 2). The average scat represented only approximately 89 — 116 fruits of the most commonly consumed types. Seed germination Germination patterns were examined only for Vac- cinium, Ribes, Rubus, and Oplopanax. A few Vaccinium seeds germinated in the year they were planted (<1% of all seeds estimated to be present); germination levels were generally low (<6%) over the next four years (Table 3). Annual germination of Oplopanax was also rather low (<18%). In contrast, both Ribes and Rubus germinated relatively well (up to 31-34% per year), with the highest levels of germination often in the second year after planting. All these kinds of seeds were clearly capable of dormancy for at least four years. There were no differences in germination percent- ages between habitats for any species (ANOVA; range of p-values: 0.17 — 0.82). The subsequent exploratory t-test also revealed no differences between habitats except for Vaccinium in one site — one statistically sig- nificant outcome in 16 tests, which might happen by chance alone. Variation among scats planted within a habitat was relatively large, but it probably reflected the biological reality of variation on a small spatial scale. 501 Kind of seed Oplopanax Rubus Streptopus Sambucus 325). 1G18) 383 (91) = = 134 (22) 123) (182) 307 (8) 126 (42) 360 = (42) 830 (291) 41 (15) 88 (33) 3K (25) 650 (137) 47 (26) - 228 (40) 583 (120) 81 (20) - 374 (76) 54 = (20) 56 (12) 683 (606) 231 634 5] 110 2 42 22 3 116 15 2 37 Discussion Bears are well-known frugivores, consuming many species of fleshy fruits (Auger et al. 2002; Willson 1993). Fruits can be an important resource for prepar- ing for winter dormancy and lactation (females give birth in winter dens), and berry-crop failures can lead to lower reproductive success (Rogers 1987). Bears can consume many kilograms of fruits in a day (Welch et al. 1997; Farley and Robbins 1995). Although a single bear scat in the present study often contained thou- sands of seeds, this represented relatively few fruits and therefore only a small proportion of possible daily consumption. Composition of the fecal deposit can affect seed germination and seedling growth (Traveset et al. 2001), and the concentration of seeds in scats can lead to post- dispersal seed predation and secondary dispersal via scattering (Bermejo et al. 1998). Germination of seeds passed by captive bears was generally similar to or faster than that of seeds extracted from fruits by researchers, although germination of some species was quite low in all conditions and was sometimes dependent on the substrates on which the seeds hap- pen to land (Traveset and Willson 1997, 1998). But rapid germination is not necessarily beneficial, if seed dormancy is indeed an adaptation for dispersal in time. The most important consistent effects of frugivory by bears on seed dispersal may be transport, rather than shifts of germination behavior of seeds passed through the digestive tract. In a companion study, the probability of seed deposition by Brown Bears on Chichagof Island was greatest at 600-900m from a source (Patten 1993). The greatest distance estimated for seed transport by Brown Bears was over three km (Patten 1993), and even though few seeds are likely to be carried this far from their source, the extended tail of the seed distribution pattern can be dispropor- tionately important for dispersal ecology (Portnoy and Willson 1992) and plant community ecology (Vellend et al. 2003). Little habitat preference was shown by the bears tracked on Chichagof, except for an avoidance of bare-rock areas, so the probability of seed deposi- 502 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 3. Estimated average percent of seeds germinating per year in two common habitats at two sites on Chichagof Island, southeastern Alaska. Site Habitat Year Kind of seed Vaccinium Ribes Oplopanax Rubus Neka Forest 1992 3.9 (0.9) 0.2 (0.2) <0.1 = 22.5 (7.2) n= 30 1993 2.8 (0.5) 30:8. e@li5e7) NS:42) HGLOSI) 28.5 (9.8) 1994 313 (0.9) 14.7 (9.0) 9.5 (4.9) 13.6 (4.8) Muskeg 1992 5.9 (2:2) - Lal (1.0) 20.7 Ga) n= 30 1993 2.9 (1.0) - 10.2 (8.4) 19.7 GS) 1994 1.2 (0.4) — TIEOe., .LO%6) 5.6 68) Salt Lk Bay Forest 1991 2.4 (0.9) 19 x CluES)) 55} A) EOF (933) n=60 1992 3.6 (0.8) 30.8 (16.2) 1.6 (0.8) 34.2 (8.7) 1993 1.8 (0.3) DES (es) iNet (0.5) 19.7 (6.9) 1994 ES (0.2) 1.0 (0.6) 3.0 (1.3) 9.8 (4.8) Muskeg 1991 1.1 (0.5) 0.5 (0.5) 4.6 (2.7) heal (5.1) n=49 1992 3.0 (1.0) 25.6 (10.4) 122, (0.7) 19.6 (6.9) 1993 1.8 (0.7) 6.3 (52) 0.9 (0.7) 10.7 (4.5) 1994 1.6 (0.8) 0.3 (0.2) 0.6 (0.4) 4.3 (3.0) tion was proportional to habitat availability, including logged areas (Patten 1993). Because forest was the common habitat, this meant that the forest fruits, such as most of the species examined here, had a fairly high probability of deposition in the appropriate habitat (Patten 1993). On a smaller scale, defecation by mov- ing bears could increase the probability that seeds from one defecation episode are deposited in more than one habitat or microhabitat, as well as decrease the inten- sity of seedling competition. Germination of several forest and forest-edge species was similar in forest and muskeg, indicating that habi- tat distribution of these shrubs is not determined by germination requirements at the habitat scale. The large variances of the estimates of germination level not only decreased the probability of detecting habitat differences, but they also suggest that microhabitats of seed deposition may be more important in deter- mining germination success than habitats. Mortality of seedlings was very high, in part an apparent result of desiccation during periods of warm, sunny weather. Especially at high densities, intra- and interspecific competition was undoubtedly intense and probably contributed to seedling mortality. However, occasion- ally small groups of seedlings persisted and were still alive when field work ended; similar “bear gardens” were also found in natural settings (Willson 1994). Bears are good seed dispersers: they commonly transport germinable seeds up to hundreds of meters from a source, often in appropriate habitat, simulta- neously with a load of fertilizer that may influence germination and growth. Although scats containing numerous seeds produce conditions of high seedling competition, several factors tend to reduce the impact of competition on the effectiveness of dispersal. Ger- mination may be spread over several years. Relatively rapid passage through the gut and multiple defecations per day within a bear’s home range spread scats over the landscape. Scat trails of moving bears spread out the seeds on a more local scale. Bears may be more reliable dispersal agents for many kinds of seeds than most frugivorous birds in this area, since the coastal region is not a major flyway for migratory forest birds. Acknowledgments. This project was funded by the Pacific Northwest Research Station, U.S. Forest Service. We are grate- ful to J. Zasada for engaging us with bear-fruit inter- actions, to our numerous field assistants, and to A. Traveset for comments on the ms. Literature Cited Auger, J., S. E. Meyer, and H. L. Black. 2002. Are American black bears (Ursus americanus) legitimate seed dispersers for fleshy-fruited shrubs? American Midland Naturalist 147: 352-367. Bermejo, T., A. Traveset, and M. F. Willson. 1998. Post- dispersal seed predation in the temperate rainforest of Southeastern Alaska. Canadian Field-Naturalist 112: 510- 512. Farley, S. D., and C. T. Robbins. 1995. Lactation, hiberna- tion, and mass dynamics of American black and grizzly bears. Canadian Journal of Zoology 73: 2216-2222. Hickey, J. R., R. W. Flynn, S. W. Buskirk, K. G. Gerow, and M. F. Willson. 1999. An evaluation of a mammalian predator, Martes americana, as a disperser of seeds. Oikos 87: 499-508. Miller, S. D., G. C. White, R. A. Sellers, H. V. Reynolds, J. W. Schoen, K. Titus, V. G. Barnes Jr., R. B. Smith, R. R. Nelson, W. B. Ballard, and C. C. Schwartz. 1997. Brown and black bear density estimation in Alaska using radiotelemetry and replicated mark-resight techniques. Wildlife Monographs 133: 1-55. Patten, L. A. 1993. Seed dispersal patterns generated by brown bears (Ursus arctos) in southeastern Alaska. MS thesis, Washington State University, Pullman, Washington. Portnoy, S., and M. F. Willson. 1992. Seed dispersal curves: the behavior of the tail of the distribution. Evolutionary Ecology 7: 25-44. 2004 Rogers, L. L. 1987. Effects of food supply and kinship on social behavior, movements, and population growth of black bears in northeastern Minnesota. Wildlife Mono- graphs 97: 1-72. Traveset, A., and M. F. Willson. 1997. Effect of birds and bears on seed germination of fleshy-fruited plants in tem- perate rainforests of southeastern Alaska. Oikos 80: 89-95. Traveset, A., and M. F. Willson. 1998. Ecology of the fruit- colour polymorphism in Rubus spectabilis. Evolutionary Ecology 12: 331-345. Traveset, A., T. Bermejo, and M. F. Willson. 2001. Effect of manure composition on seedling emergence and growth of two common shrub species of southeastern Alaska. Plant Ecology 155: 29-34. Traveset, A., M. F. Willson, and M. Verdi. 2004. Charac- teristics of fleshy fruits in southeastern Alaska: Phyloge- netic comparison with fruits from Illinois. Ecography 27: 41-48. WILLSON and GENDE: SEED DISPERSAL BY BROWN BEARS Vellend, M., J. A. Myers, S. Gardescu, and P. L. Marks. 2003. Dispersal of Trillium seeds by deer: implications for long-distance migration of forest herbs. Ecology 84: 1067- 1072. Welch, C. A., J. Keay, K. C. Kendall, and C. T. Robbins. 1997. Constraints on frugivory by bears. Ecology 78: 1105- 1119. Willson, M. EF. 1991. Dispersal of seeds by frugivorous animals in temperate forests. Revista Chilena de Historia Natural 64: 537-554. Willson, M. F. 1993. Mammals as seed-dispersal mutualists in North America. Oikos 67: 159-176. Willson, M. F. 1994. Bear gardens. Pacific Discovery 47(1): 8-14. Received 15 July 2003 Accepted 23 October 2004 A Mapping of the Present and Past Forest-types of Prince Edward Island D. G. SoBey! AND W. M. GLEN? ! School of Applied Biological and Chemical Sciences, University of Ulster, Jordanstown, Northern Ireland BT37 OQB United Kingdom. (Research Associate of the Institute of Island Studies, University of Prince Edward Island, Charlottetown, Prince Edward Island) 2 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. 2004. A mapping of the present and past forest-types of Prince Edward Island. Canadian Field-Naturalist 118(4): 504-520. Our aim was to produce maps showing the distribution on Prince Edward Island of five forest-types previously identified from a TWINSPAN analysis of ground flora data collected at 1200 sampling points in a field survey. For this purpose we had available two databases: one on the composition of the tree canopy of 82 957 forest stands, as determined by photo- interpretation of a 1990 aerial, photographic survey of the island; the other on the drainage properties of the same stands from a published soil survey. The tree canopy and drainage criteria for sorting these stands into five stand-types were chosen in the light of the equivalent properties of the TWINSPAN forest-types as evident from the field survey. These criteria were perfected in four trial computer-sortings, followed by the computer-printing of maps showing the distribution of the stand- types. These maps, which were then evaluated by comparing them with the properties of the TWINSPAN forest-types, are the first fine-scale maps of the main forest-types of the island. They reveal that, of the three “primary” forest-types, the upland hardwood forest occurs especially in the central and south-eastern hill-lands, as well as in scattered parcels elsewhere, whereas the Black Spruce forest and the wet species-rich woodland occur primarily in areas of lower elevation in the east and west of the island. The two forest-types resulting from human disturbance, the White Spruce woods and the “disturbed forest”, have a more scattered distribution, with the White Spruce woods being found especially in the central and eastern parts of the island and the disturbed forest in the west and east of the island. A secondary aim was to map the conjectured distribution before European settlement of the three primary forest-types: two maps have been produced, one showing the distribution of upland hardwood forest, the other of the wet forest-types. Key Words: forest classification, forest mapping, forest history, Prince Edward Island. Between 1990 and 1992 the Prince Edward Island Forestry Division carried out a comprehensive Forest Biomass Inventory of the province, a component of which was a field survey in 1991 at 240 randomly- selected forest sites. At each of the sites the percentage cover of the ground flora species in 4-m? plots was recorded at five sampling points. The availability of such ground flora data from such a large number of plots (1200 in total) presented an unprecedented oppor- tunity for the study and analysis of the woodland ground vegetation and forest-types of Prince Edward Island. An earlier paper (Sobey and Glen 2002) presented an analysis of the ground flora data in 1127 of the plots using two multivariate techniques: TWINSPAN — two-way indicator species analysis (a classification technique), and DECORANA -— detrended correspon- dence analysis (an ordination technique). TWINSPAN led to the recognition of five forest-types, each char- acterized by particular tree species and soil drainage properties: (1) a wet species-rich swamp-type wood- land, (2) upland hardwood forest, (3) Black Spruce (Picea mariana) forest, (4) old field White Spruce (Picea glauca) woods, and (5) disturbed conifer-dom- inated forest. It was conjectured that the first three of these forest-types were heavily modified descendants of forest-types occurring at the time of European set- tlement, while the two latter appeared to be largely the product of successional processes associated with the effects of human disturbance and forest clearance. The DECORANA ordination provided further sup- port for the overall importance of soil drainage and human disturbance as the principal factors responsi- ble for the differences between the five forest-types. The aim of this paper is to expand the results of the plant community analysis beyond the 1200 sampling points to include the whole forested area of the island, in the form of maps showing the total distribution of the five forest-types on the island. Since the forest-types were initially segregated and defined by TWINSPAN on the basis of the composition of their ground flora, the ideal mapping approach would have been to use the ground flora composition of all forest stands on the island as the basis for mapping the distribution of the forest-types. However, such data are not available for the whole forested area of the island and are never likely to be. The forest-types thus had to be mapped using data that showed a correlation with the ground flora composition (1.e., data on the composition of the tree canopy) and also for some, data on soil drainage properties. It is fortunate that there were avail- able two comprehensive relevant databases for the 504 2004 whole island: the more important was a database on the tree species composition of all forest stands on the island, based on an aerial photographic survey carried out in mid-summer 1990; the other was the Prince Edward Island Soil Survey (MacDougall et al. 1988) which classified the soils of the island into 44 soil series and mapped their distribution (at a scale of 1:10 000) over the whole island. The soil survey proved useful in the forest mapping because some of the TWINSPAN forest-types had high levels of association with partic- ular soil drainage properties. It should be noted that none of the previous attempts at forest classification and mapping that include Prince Edward Island (Stilgenbauer 1929; Halliday 1937; Rowe 1959; Loucks 1962) aimed to map what was actually present on specific sites, nor do they contain a high level of detail or accuracy. Halliday’s and Rowe’s maps, being part of national studies, show no internal differentiation at all in the forests of the island. Loucks (working at the level of Maritime forests), did sub- divide the island’s forests: he placed them in three “forest districts” but the boundaries of these are very generalized on his map. One comprised the hardwood areas of the central and eastern parts of the island; the other two were coniferous districts: one in the west along the shores of Northumberland Strait, the other running along the length of the northern shore. The map showing the greatest detail (Stilgenbauer 1929) is actually the earliest of the four, but Stilgenbauer’s descriptions of his “forest belts” are brief and quali- tative and the criteria he used for delimiting them are not given. In addition to these mapping studies, there is a useful descriptive summary of the forest-types of the island in Erskine (1960), and in Appendix 1 the stand-types emerging from this study are equated with Erskine’s descriptions. The principal objective of this study was thus, mak- ing use of two comprehensive databases, to map the total distribution on Prince Edward Island of the five forest-types recognized in a TWINSPAN analysis, and to further verify the validity of the maps by comparing the properties of the mapped stands with those of the TWINSPAN forest-types. Such a mapping will not only give us a clear picture of the current actual distribution on the ground of the main forest-types on the island, but should also assist us in understanding the factors responsible for their distributions. A secondary objective (on the basis of any stand and soil relationships that might emerge) was to attempt to extend the mapping back into historical time in order to obtain a picture of the possible distribution of the forest-types before the advent of the large scale forest clearance that began in the eighteenth century with the beginning of Euro- pean settlement on the island. Methods The methodology leading to the description of the forest-types and their mapping makes use of several SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. large independently obtained databases and involves a number of different stages in processing and analysing the data they contain. Figure | shows these databases and stages in diagrammatic form. The end-product, computer-produced maps of the five forest-types, is the result of a computer-sorting of 82 957 demarcated stands of “high forest” (i.e., excluding alder woods, plantations, clear-cut and burned areas) into five forest- types on specific tree canopy and soil drainage criteria. In the description of the various steps in the analysis, the text that follows is linked to the stages shown in Fig- ure | by Roman numerals in square brackets, e.g.: [I]. The databases: 1. The 1991 field survey: ground flora and tree canopy data Of the vast amount of data collected at the 1200 sam- pling points in the 1991 field survey, two data sets, one on the ground flora [II], the other on the tree can- opy [III], are of relevance to this report. We present here only a summary of the methods used to collect these data — full field methods are given in Sobey and Glen (2002). The data were collected between June and Sep- tember 1991 at 1200 ground flora plots located at 240 randomly-selected forest sites. At each sampling point the tree canopy was assessed using a “point sampling” technique involving the use of a “variable- radius plot” centred on the sampling point (Watts 1983). From these field data, the percentage contribu- tion of each tree species to the total woody biomass (to the nearest 10%) in each variable radius plot was cal- culated [III]. The ground flora species were assessed within a circular 4-m? plot, with each species being given a percentage cover value to the nearest 10%, with a minimum value of 5% [II]. The TWINSPAN classification and the recognition of five forest-types The TWINSPAN classification of 1127 of the 1200 ground flora plots (based on the species composition and percentage cover of their ground flora species) resulted in the recognition of 11 ground flora plot groups or community-types [V] (see Sobey and Glen (2002) for a full description of the methodology — note that 73 plots were omitted from the analysis either because no trees were recorded in the area of the sampling point, or because no ground flora species were present in the 4-m* plot). These plot groups were then examined in terms of the species composi- tion of the tree canopy and other properties at the sam- pling sites (especially soil drainage), and as a result five major forest-types were recognized [VII]. (See Sobey and Glen (2002) for summary descriptions of each of these forest-types and maps showing the dis- tribution of their sampling points on the island.) 2. The 1990 aerial survey: the forest stand database Complete aerial photographic coverage of the island was carried out in mid-summer 1990 as part of 506 1990 AERIAL PHOTO SURVEY: 1:17 500-scale photos of the whole Island IV PHOTO-INTERPRETATION: demarcation and description of 82 957 forest stands: = Stand database XI THE CANADIAN FIELD-NATURALIST a Il 1991 FIELD SURVEY: Ground flora data from 4-m plots at 1200 sampling points TWINSPAN CLASSIFICATION: =» Recognition of 11 ground flora community-types VII ANALYSIS of TWINSPAN GROUPS with the aid of tree canopy and soil series data for the 1200 points: =» Recognition of five forest-types VIE SELECTION of TREE CANOPY & SOIL DRAINAGE CRITERIA for classification of 82 957 stands into five forest-types COMPUTER SORTING of the 82 957 forest stands corresponding to the five forest-types, using the selected tree canopy and soil drainage criteria COMPUTER MAPPING of the stands corresponding to each of the island's five 1990 forest-types Vol. 118 1991 FIELD SURVEY: Tree species data from ‘variable-radius’ plots at 1200 sampling points Vi 1988 P.E.I. SOIL SURVEY: Description and mapping of 44 soil series for the Island IX DATABASE of SOIL SERIES & DRAINAGE CLASS for the 1200 sampling points x DATABASE of SOIL DRAINAGE CLASS for 82 957 forest stands FIGURE 1. The stages leading to the mapping of the 1990 forest-types on Prince Edward Island: a flow diagram showing the databases used (in the boxes with heavy borders) and the analyses carried out. the 1990-1992 Prince Edward Island Forest Biomass Inventory [I]. The false-colour infra-red photographs resulting from the survey (scale c. 1:17 500) were analysed by trained photo-interpreters in the Forestry Division with the purpose of dividing the total forest- ed area into “stands” which were then recorded on a map as “polygons” [IV]. For the purpose of the photo-interpretation, a stand was taken to be a group of trees having a relatively uniform visual appearance from the air as evident in the aerial photograph. Stands could comprise a single species or several or many species, but they are generally of a uniform species composition, height and density. For each of the 82 957 stands, up to five tree species, each con- 2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 507 FIGURE 2. The distribution of “high forest”, based on photo-interpretation of the 1990 aerial photographic survey of the island. (High forest excludes Speckled Alder woods, plantations, and clear-cut and burned areas.) tributing greater than 5% crown closure to the canopy, were recorded, with the percentage contribution of each species being estimated to the nearest 10%. The database resulting from the photo-interpretation was computerised, and the stand boundaries were linked to the Geographic Information System (G.LS.) for the province, enabling the direct plotting onto maps of stand boundaries and areas, as well as the extraction of data on their tree species make-up. 3. The 1988 soil survey of Prince Edward Island: a soil series database The most recent major soil survey of the island (MacDougall et al.1988) has resulted in the classifi- cation of the island’s soils into 44 soil types or “series”, and the mapping (scale: 1:10 000) of the distribution of these series over the whole island [VI]. The soil maps have been digitized in the G.I.S. for the island, such that the soil series at each of the sampling points in the 1991 field survey [IX], as well as for each forest stand polygon in the aerial stand database [X], are accessible by computer. For this study it was decided that it would also be useful to group the 44 soil series into six drainage classes: rapidly-drained, well- drained (coarse-textured parent materials), well- drained (medium-textured parent materials), imperfect- ly-drained, poorly-drained, and organic soils. Selection of the tree canopy and soil drainage criteria. The selection of the criteria for dividing the 82 957 stands of high forest (i.e., the area shown in Figure 2) into the five forest-types [VII] (or “stand-types” as they will be termed to distinguish them from the TWINSPAN-defined forest-types), was based on the tree canopy and soil drainage properties of the five TWINSPAN forest-types as evident from the 1991 field survey (Sobey and Glen 2002). Four trial com- puter sortings on selected criteria were carried out in succession; the results of each trial were evaluated and the criteria for each of the five stand-types were then redefined and a new sorting was carried out. Eventually the perfected criteria were chosen (Table 1), and all of the stands in the database were comput- er-sorted on these criteria [XI], followed by the com- puter-printing of maps showing the distribution of each stand-type on the island [XII]. It should be noted that in any of the sortings the order of the sort (i.e., of the selecting and removal of each stand-type) is important, since once a stand has been sorted it is normally no longer available for inclusion in subse- quent stand-types even if it should meet their particu- lar criteria. The following is a more detailed discus- sion of the criteria used for each of the five forest-types, as well as of some of the changes made as a result of the trial sortings: I. Upland hardwood forest. In the TWINSPAN classi- fication Sugar Maple (Acer saccharum) and American Beech (Fagus grandifolia) had been virtually restrict- ed to sampling points belonging to the upland hard- wood forest-type (Sobey and Glen 2002). Yellow 508 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 1. The criteria used in the sorting of 82 957 stands of “high forest” on Prince Edward Island into five stand-types corresponding to the five forest-types recognized in a TWINSPAN classification of 1127 ground flora plots. SELECT AND REMOVE: stands designated as clear-cut, burned, plantation, windfall, or alder. This leaves 82 957 stands of “high forest” First Sort: Excluding any stands containing White Ash, Eastern White Cedar or White Elm, select all stands containing Sugar Maple, American Beech or Eastern Hemlock, plus all stands with Yellow Birch, but for the last species excluding those stands on poorly-drained, imperfectly-drained or organic soils. 1. Upland hardwood forest SECOND Sort: Excluding any stands containing White Ash, Eastern White Cedar or White Elm, select all stands containing Black Spruce with crown closure 2 50%. 2. Black Spruce forest THIRD Sort: Select all stands containing White Elm, White Ash, or Eastern White Cedar, and/or classed as ‘swampy’, plus all stands with a crown closure of total hardwoods = 50% and occurring on poorly-drained or imperfectly-drained soils. 3. Wet rich woodland FourTH Sort: Select all stands containing White Spruce with crown closure = 40%. 4. White Spruce woods RESIDUAL STANDS: Birch (Betula alleghaniensis) had a somewhat wider distribution, also occurring at a rather high level at wet rich woodland sampling points. In the initial trial sortings all stands containing any presence of Sugar Maple, American Beech and Yellow Birch were select- ed; however, for the final sorting it was decided to exclude those stands selected due to the presence of Yellow Birch that occurred on poorly-drained, imper- fectly-drained or organic soils, soils more characteristic of wet rich woodland than of upland hardwood forest. Also, in the final sort, Eastern Hemlock (Tsuga canadensis) was included as a selecting species, and stands containing American Elm (Ulmus americana), Eastern White Cedar (Thuja occidentalis) and White Ash (Fraxinus americana), all characteristic of wet rich woodland, were specifically excluded. 2. Black Spruce forest. The problem in segregating stands of this forest-type was choosing a minimum percentage crown closure for Black Spruce. Since the mean percentage woody biomass contribution of Black Spruce at the Black Spruce forest sampling points was 53.1% (Sobey and Glen 2002), it was decid- ed to set both a high value and also one that made ecolog- ical sense: the criterion used in the final sort was that for a stand to be assigned to Black Spruce forest, at least 50% of its canopy had to be Black Spruce. From the earlier trial sortings it had also become evi- dent that it was necessary to exclude any stands con- taining American Elm, White Ash or Eastern White Cedar, species virtually absent from this forest-type. 3. Wet rich woodland. In the early trial sortings an attempt was made to select stands of this forest-type on the percentage crown closure of its most impor- tant tree species: Red Maple (Acer rubrum) — a mini- mum crown closure of = 25% was set. However, the problem was that Red Maple was also present at high 5. “Disturbed forest” levels in the still unsorted “disturbed conifer-dominated forest’. It was thus decided to make use of the fact that 80.2% of the sampling points of the TWINSPAN wet rich woodland occurred on imperfectly- and poorly-drained soil series (Sobey and Glen 2002). In the end the criterion used was firstly to select all stands containing the minor tree species diagnostic of this for- est-type (as evident in the TWINSPAN classification): ie., White Ash, American Elm and EasternWhite Cedar, as well as all stands classified as “swampy” by the photo-interpreters. Then, since an overall factor dis- tinguishing the TWINSPAN wet rich woodland sam- pling points from the disturbed conifer-dominated forest was a higher level of total hardwoods (Sobey and Glen 2002), we selected all stands with a minimum total hardwood crown closure of 50%, in combina- tion with a requirement that all such stands be on poorly- and imperfectly-drained soils. Having selected the stands that appear to be derived from pre-European settlement forest-types, the next step was to separate the remaining stands into the two types of successional and disturbed forest: i.e., White Spruce woods and disturbed conifer-dominated forest. 4. White Spruce woods. Here it was a matter of selecting the minimum percentage crown closure of White Spruce for a stand to be assigned to White Spruce woods. Given the fact that the mean percentage contri- bution of White Spruce to the woody biomass of TWINSPAN White Spruce woods sampling points was 62.2 % and in the disturbed conifer-dominated forest it was 16.7% (Sobey and Glen 2002), a middle level (40%) was chosen. 5. Disturbed conifer-dominated forest (or simply “disturbed forest”). The residual stands should by default correspond to the remaining TWINSPAN for- est-type (disturbed conifer-dominated forest), even if 2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 509 TABLE 2. The number of hectares and stands (also as percentages) assigned to each stand-type in the computer-sorting. A percentage breakdown of the stand area by drainage class is also given, as is its status in 1935 as either forest-covered or cleared land as determined from a 1935 aerial photographic survey of the whole island. (In brackets after the forested per- centage is the equivalent parameter for the TWINSPAN forest-type sampling points of the 1991 field survey — those bolded are within + 20% of the stand value.) STAND-TYPES Upland Black Hardwood Spruce Forest Forest HECTARES 55 043 33 106 Percentage of area Dei 13.1 NUMBER OF STANDS 16 170 8 049 Percentage of stands 19.5 9.7 DRAINAGE CLASS (% of area) Organic 0.04 6.8 Poorly-drained 5.8 54.8 Imperfectly-drained 39 16.8 Well-drained (medium) LG) 1.1 Well-drained (coarse) 74.4 9.0 Rapidly-drained 12.9 10.2 Unclassified 0.28 s3 STATUS in 1935 (% of area) Forested 93.8 TP (86.4) (67.5) Cleared land 6.2 22.8 all are not in fact dominated by conifers. In the trial sortings an additional fifth sort was carried out in order to separate out the residual stands that were indeed conifer-dominated. This was done by sorting on a minimum conifer crown closure of = 50% — this selected 42% of the residual stands. The other group was thus termed “disturbed hardwood-dominated for- est’, i.e., comprising residual stands with hardwoods > 50% crown closure. However, in the end, the distinc- tion between hardwood and softwood domination in the residuals was not considered important and these stands have been simply termed “disturbed forest”. Results Table 2 presents the results of the final computer sorting, expressed in terms of the area in hectares and the number of aerial stands assigned to each stand- type, both also expressed as percentages. Table 2 also gives a breakdown of the soil drainage classes for each of the stand-types and their status in 1935 (as “cleared land” or “forested”), as determined from analysis of the earliest aerial photographic survey of the island (see Glen 1997). The tree species make-up of the five stand-types is shown in Table 3 and the computer-printed maps showing their distribution on the island are shown in Figures 3 to 7. In the sorting, the 1990 “high forest” of the island was divided into five stand-types — see Appendix | for summary descriptions of each of these stand-types. These fall naturally into two groups, each of which occupies about half of the island’s forested area: (1) White Wet Rich Spruce Disturbed Woodland Woods Forest TOTALS 42 192 57 982 65 259 253 582 16.6 22.9 Dwi 100 13 863 22 709 22 166 82 957 16.7 27.4 26.7 100 0.78 0.58 0.86 1.4 70.1 11.6 14.5 25.8 24.6 8.1 7.8 10.7 0.74 2.8 5.0 2.4 2D) 48.3 46.2 35.6 13 213 24.1 14.9 0.18 IED eS) 0.93 75.8 37.6 67.4 69.0 (79.5) (24.4) (74.2) (69.5) 24.2 62.4 32.6 31.0 the three stand-types which appear to be recognizable descendants (even if heavily-modified) of pre-European settlement forest-types (upland hardwood forest, Black Spruce forest and wet rich woodland) — these make up 51.4% of the area under high forest; (2) the two other stand-types (old field White Spruce woods and “disturbed forest’), representing more disturbed and/or successional types not likely to have been impor- tant in the pre-European forest — these make up the other 48.6%. It is likely that before the period of European settlement, land surfaces now under these two latter stand-types would have been covered by one of the three “primary” stand-types. Evaluating the validity of the stand-types One way of assessing the validity of the products of the sorting is to compare, as in Table 3, the tree species composition of each stand-type (in the form of per- centage crown closure) with the equivalent parameters of the corresponding TWINSPAN forest-type: the per- centage contribution to woody biomass in the 1200 variable-radius plots of the ground survey, and the con- tribution of each of the tree species to the percentage crown closure in those stand polygons in which the 1200 variable-radius plots were located. Another way to assess the validity of the sorting is to compare the drainage properties and the status in 1935 of the aerial stand-types with those of the TWINSPAN-based forest- types (Tables 2 and 4). The data required for these comparisons were extracted using the Database pro- gram “FoxPro” from the 1991 field survey database 510 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 3. The mean percentage crown closure (+ S.E.) of the principal tree species in each stand-type (based on all the stand-polygons of each stand-type including those in which the species was not recorded). To enable comparison of these values with the equivalent parameters for the TWINSPAN forest-types, placed below in round brackets is the mean percentage contribution (+ S.E.) of each tree species to the total tree biomass in the variable-radius plots of the corresponding TWINSPAN forest-type; and in square brackets the mean percentage crown closure of the tree species (+ S.E.) in those stand- polygons in which the variable-radius plots were located. For crown closure values (unbracketed) > 10%, a bolded percent- age within the brackets indicates that there is “good agreement” between the two values (i.e., the bracketed value lies within + 20% of the unbracketed value), while an italicized bolded percentage indicates a “notable discrepancy” between the two values (i.e., the bracketed value is > + 50% of the unbracketed value); where crown closures (unbracketed) were < 10%, the differences were assessed subjectively on the basis of their relative magnitudes. STAND-TYPES ALL STANDS Upland Black Wet White (All 1200 plots +) Hardwood Spruce Rich Spruce Disturbed [AIl 1200 sampling Forest Forest Woodland Woods Forest point stands] CONIFERS: Picea mariana 0.22+0.020 71.4+0.18 9.9+0.13 1.1 + 0.034 5.3 + 0.079 10.4 + 0.080 (0.72+0.35) (53.1 +4.9) (3.8 + 1.2) (4.3 + 1.5) (10.1 + 1.4) (5.4 + 0.57) [0.79 + 0.28] [39.6 + 3.8] [13.3 + 1.7] [4.1 + 1.1] [12.4 + 1.3] [9.1 + 0.59] Picea rubens 0.59+ 0.032 0.69 + 0.042 2.9 + 0.072 0.47+0.021 7.1 +0.096 2.7 + 0.036 (0.88 + 0.27) (17.3 +3.8) (0.51 + 0.30) (4.6 + 1.2) (Gfe Wien bi) (2.8 + 0.32) [0.98 + 0.28] [3:3 1:2] (5.4 + 1.1] [3.0 + 1.1] [3.6 + 0.66] [2.6 + 0.28] Picea glauca 4.8 + 0.086 1.6 + 0.068 8.2+0.11 68.1+0.13 10.4+0.078 24.2+0.11 (8.6 + 0.92) (32 £723) (18.0 + 2.2) (62.2+3.4) (16.7+ 1.5) (17.0 + 1.0) [10.3+0.94] [25.1 + 3.4] [7.3 + 0.98] ATDE3:01 ) 7a 13) [17.5 + 0.76] Abies balsamea 9.0 + 0.093 4.8 + 0.086 5.6 + 0.072 3.8 + 0.053 9.0 + 0.089 6.6 + 0.037 (18.9 + 1.2) (6.9 + 1.9) (13.5 + 1.5) (8.4 + 1.7) (24.9 + 1.6) (17.6 + 0.75) [10.2 + 0.60] [7-32] [7.8 + 1.0] [5.6+ 0.77] [11.1 + 0.67] [8.9 + 0.33] Tsuga canadensis 0.12 + 0.012 0 0 0 0 0.023 + 0.0023 (0.32 + 0.16) (0) (0) (0.81 40.54) (0.42+0.25) (0.40 + 0.14) [0] [0] [0] [0] [0] [0] Larix laricina 0.082 +0.0097 8520.11 5.4 + 0.075 6.0 + 0.075 12.8 + 0.16 7.1 + 0.052 (0.48 + 0.18) (9.0+2.1) (7.3 + 1.7) (3.7+0.84) (3.84 0.69) (2.6 + 0.31) [1.5 + 0.35] [9.5 + 1.5] [7.8 + 1.2] [8.8 + 1.6] [7.6 + 0.81] [5.3 + 0.37] Thuja occidentalis 0 0) 1.2 + 0.037 0 0 0.20 + 0.0064 (0.42+0.25) (0.12 + 0.13) (TET) (0) (0.64+0.32) (0.93 + 0.23) [0.023 + — ] [0] [0.38 +— ] [0.15+-] [0.39+0.14] [0.17 + 0.041] TOTAL CONIFERS 14.9 87.3 34.9 80.4 46.3 51.4 (30.4) (94.4) (50.0) (84.1) (64.5) (47.3) [23.8] [84.7] [42.0] [69.6] [52.4] [44.4] BROAD-LEAVES: Acer rubrum Sysifas OBL 4.8 + 0.086 25) 210a3 5.4 + 0.061 20.1 + 0.12 18.7 + 0.063 (33.3+41.4) (2.5 +0.77) (24.6 + 2.4) (4.6 + 1.1) (19.0 + 1.5) (Bales 1) [30.4 + 0.73] [5.1 + 1.1] [26.3 + 1.5] [8341.1] [18.9+0.92] [21.7+0.52] Acer saccharum 21.6+0.10 0 0.45 + 0.030 0 0 4.3 + 0.036 (9.7 + 0.89) (0) (1.7+0.56) (0.59+0.46) (0.28 +0.13) (4.9 + 0.54) [16.7 + 0.76] [0.5 +-] [2.1 + 0.55] [20160] [35.053] [7.7 + 0.38] Betula alleghaniensis 7.8 + 0.070 0) 0.22+0.016 0.013 + 0.0026 0.090 + 0.0024 1.6+0.017 (5.6 + 0.62) (0) (3.3 + 0.84) (0.22+0.13) (1.4+0.40) (3.9 + 0.39) [6.7 + 0.41] [0] (0.71+0.22] [1.0+0.35] [0.92 + 0.23] (3.0 + 0.19] Betula papyrifera 137+ 01087, 2:5 = 01059 12.7 + 0.089 6.3+0.059 13.4+0.088 10.3 + 0.039 (9.2+0.74) (0.63 + 0.36) (5.6 + 0.94) (3.6+0.78) (41+0.60) (7.3 + 0.50) {13.1+ 0.53] [2.5 + 0.65] (11.9+0.97] [6.9+0.85] [11.4+0.66] [10.8 + 0.33] Fagus grandifolia 1.5 + 0.036 0 0.017 + 0.0044 0 0 0.30 + 0.0073 (4.1 + 0.58) (0) (0) (0.37+0.24) (0.23+0.11) (2.7+0.37) [1.1 + 0.19] [0] [0.19 +—] (0.074+-—] [0.21+0.10] [0.49 + 0.074] (continued) on next page 2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. S51] TABLE 3. (continued from previous page) STAND-TYPES ALL STANDS Upland Black Wet White (All 1200 plots +) Hardwood Spruce Rich Spruce “Disturbed [AIl 1200 sampling Forest Forest Woodland Woods Forest” point stands] Populus spp. ¢ 4.6 + 0.075 5.0 + 0.090 21.4+0.15 7.6+0.070 18.8+0.14 I23:2.0/057 (4.1 + 0.65) (1.1 + 1.0) (7.8 + 1.6) (4.7 + 1.2) (7.4 + 1.1) (6.6 + 0.62) [6.9 + 0.53] [6.8 + 1.2] {11.2+1.0] [10.7+1.1] [10.9+0.80] [9.6 + 0.39] Fraxinus americana 0 0 0.32 + 0.019 0 0 0.053 + 0.0031 (0.40 + 0.21) (0) (1.6 + 0.75) (0) (O) (0.46 + 0.16 ) [0] [0] [0] [0] [0] [0] Ulmus americana 0 0 0.84 + 0.035 0 0 0.14 + 0.0058 (0.26 + 0.14) (O) G2 saie1) (0) (0) (0.57 + 0.17) [0.47+ — | [0] (0.13 +-] [0] [0.035+-—] [0.042 + 0.019] Alnus incana 0.047 + 0.011 0.26 + 0.027 4.8+0.11 201039" Os 01052 6.4 + 0.071 (-) (-) (-) (-) (-) (-) [0.16 + 0.12] [0] (4.0+1.2] [1.1+0.63] [0.85 + 0.38] [et e10:22] TOTAL BROAD-LEAVES 85.1 12.5 65.1 19.6 3 47.8 (excluding Alnus) (70.0) (5.3) (50.0) (16.5) (35.0) (52.7) [75.6] [14.9] [56.7] [30.3] [46.9] [55.6] Number of Stands 16,170 8,049 13,863 22,709 22,166 82,957 ¥ For “all 1200 plots” the value in round brackets is the percentage contribution to the total above ground biomass (oven- dry tonnes per hectare) in all 1200 variable-radius plots. + Because it is impossible to distinguish the three native poplar species in aerial photographs (Populus tremuloides, P. balsamifera, P. grandidentata) they were grouped as “Populus spp.” — however almost all of this is likely to have been P. tremuloides (Trembling Aspen). FIGURE 3. The distribution of stands of upland hardwood forest in 1990. a2 THE CANADIAN FIELD-NATURALIST Vol. 118 Ficure 4. The distribution of stands of Black Spruce forest in 1990. FicureE 5. The distribution of stands of wet rich woodland in 1990. 2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. FIGuRE 6. The distribution of stands of White Spruce woods in 1990. FIGURE 7. The distribution of stands of “‘disturbed forest” in 1990. 514 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 4. The soil drainage classes of the stand-types (derived from the 1:10 000 soil series maps of MacDougall et al. (1988)) expressed as the percentage of the stand-type’s total area in each drainage class, ignoring the area of unclassified drainage. These are compared with the drainage classes of the corresponding TWINsPAN forest-types: (1) (in round brackets) the soil drainage class at the sampling points of the 1991 field survey as determined from soil pits, and (2) [in square brack- ets] the drainage class of those soil series polygons in which the sampling points were located, derived also from Mac- Dougall et al. (1988), both values being expressed as the percentage of the total area or of the sampling points in each drainage class. * STAND-TYPES Upland Black Hardwood Spruce SOIL DRAINAGE CLASS Forest Forest Poorly- and imperfectly- 9.8 79.4 drained soils (+ organic)(%) (1.9) (66.3) [12.9] [73.8] Well- and rapidly- 90.2 20.6 drained soils (%) (98.1) (33.8) [87.1] [26.3] White ALL Wet Rich Spruce “Disturbed STANDS Woodland Woods Forest” —_ (or points) 95.7 20.6 23.6 39.2 (58.3) (1.5) (15.9) (18.4) [80.3] [14.0] [37.5] [33.6] 4.3 79.4 76.4 60.8 (41.6) (98.5) (84.1) (81.6) [19.9] [85.9] [62.5] [66.4] *For the meaning of the bolding and italicizing see the caption of Table 3. and the 1990 aerial stand database, and then linked to the “MapInfo” G.LS. for the province. (1) Comparing the composition of the tree canopy To facilitate the overall interpretation of the many comparisons being made, indices were selected for assessing the closeness of the two values: if the equiv- alent parameter for the TWINSPAN-based forest-type lay within + 20% of the aerial-stand value, this was taken to indicate “good agreement” between the two values, whereas if the parameter was greater than + 50% of the aerial-stand value this was considered a “notable discrepancy”. On the basis of these criteria, it is evident that in overall terms there is a reasonably good fit between the mean tree canopy composition of the stand-types based on all of the stand polygons (Table 3), and the canopy composition of those stand poly- gons that contained the 1200 variable-radius plots: there are 29 “good agreements” and only two “notable dis- crepancies” for the crown closure of individual species. However, when the mean tree canopy composition for all of the stand polygons is compared with the per- centage contribution to woody biomass in the variable- radius plots, the match is not as good: here there were 15 “good agreements” and 17 “notable discrepancies”. This greater level of discrepancy is not surprising considering that this latter comparison is between plots of different size (i.e., forest stands of variable size, but measured mostly in hectares (the mean stand area was about 3 hectares) and variable-radius plots with a mean size of about 100 m7), as well as different parame- ters (percentage crown closure as determined from an aerial photograph, with the percentage contribution to woody biomass based on the trunk diameter of the trees within the variable-radius plots). Also relevant here is the greater difficulty in identi- fying tree species from an aerial photograph compared with the examination of actual specimens in a field survey. The photo-identification was based on such characters as texture, shape, colour and site (e.g., wheth- er on upland or lowland). The conifers in particular presented problems: Balsam Fir (Abies balsamea) when present as an understorey species may not be apparent from the air; the three spruce species were frequently identified from their site: i.e., if in upland forest, it was listed as Red Spruce (Picea rubens), in lowland, Black Spruce, and on old-field sites, White Spruce, but this was not an infallible approach. Thus the photo- identification was not 100% accurate: the standard re- quired for the 1990 aerial survey, confirmed by ground checks, was that at least 70% of the species-cover identification in a stand be correct for at least 90% of the time. Bearing these points in mind we observe the following: Upland hardwood stands. In almost all of the compar- isons (Table 3) there is generally a good correspondence between the parameters of the aerial stand-type and those of the TWINSPAN-defined forest-type. Black Spruce stands. There are problems with the spruces (perhaps due to the problem in identifying the spruces, discussed above): in Table 3 the most notable discrepancy is the very low Red Spruce contribution to the aerial stand-type (<1%) compared with its level in the variable-radius plots (17%). Also evident is the fact that Black Spruce is at a somewhat higher level in the aerial stand-type (71%) compared with either of the values for the TWINSPAN-defined forest-type. This is presumably due to the high crown closure level (i.e., 50%) that was set as the criterion for inclusion of Black Spruce stands in this group. Wet rich woodland stands. The chief species, Red Maple, shows no difference in either of the comparisons 2004 (Tables 3). However, there are some discrepancies: the use of a sorting criterion of 2 50% hardwoods must be the reason for the higher overall hardwood contribution in the aerial stand-type compared with the TWINSPAN forest-type, as well as the higher individual contribu- tions of Trembling Aspen (Populus tremuloides) and White Birch (Betula papyrifera), though there is no dis- crepancy for the latter species in the comparison of the crown closures. Concomitantly, the conifer contribution is less in the aerial stand-type than in the variable-radius plots, but again there is less discrepancy for the crown closure comparison. White Spruce stands. In overall terms there is a good correspondence between the two classifications, with the main species, White Spruce, giving a reasonably good fit — this time the better fit is for the variable- radius plot comparison (Table 3). “Disturbed forest” stands. This residual group has a smaller total conifer component than the TWINSPAN “disturbed conifer-dominated forest” (this discrepancy is less evident in the comparison of the crown closures — Table 3), and this is reflected especially in the levels of Balsam Fir (for the variable-radius plot comparison only), and to a smaller degree, in White Spruce and Black Spruce. Oddly, Tamarack (Larix laricina) is con- siderably higher in the aerial stand-type than it is in the variable-radius plots. Concomitantly, the hardwoods (notably Trembling Aspen) are higher in the aerial stand-type, while there is no difference for the levels of Red Maple. (2) Comparing the soil drainage classes and the 1935 Status There is a reasonably good agreement between the aerial stand-types and the TWINSPAN forest-types in their drainage class breakdown (Table 4), especially for the drainage classes as determined from the soil series at the 1200 sampling points. There is less agree- ment with the drainage classes as determined from the soil pits dug at the sampling points during the field inventory. The one showing the least correspondence is the wet rich woodland which has higher levels of poor- ly- and imperfectly-drained soils in the aerial stands, due to the direct use of these drainage classes as a sorting criterion. The status of the stands and sam- pling points in 1935 (1.e., whether they were forested or cleared land at that time) is in broad agreement for most of the forest-types (Table 2), with the greatest discrepancy being for the White Spruce woods. Discussion The TWINSPAN analysis of the 1991 field survey data (Sobey and Glen 2002) indicated the presence of five main forest-types on Prince Edward Island, three of these showing a relationship with pre-European settlement forest-types and two appearing to largely owe their presence to human influences. The objective of the current paper was the production of maps show- SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. S15 ing the total distribution of these five forest-types. This objective has been achieved, and for the first time ever we have fine-scale whole-island maps showing the distribution of the main forest-types. Furthermore, these maps have been produced using largely objective cri- teria and methods, and they are based on the actual for- est present at a specific time (in the summer of 1990), with every mapped stand being locatable on the ground and thus potentially verifiable. How valid are the maps as a picture of the distribu- tion of the five forest-types on the island? Firstly, the maps appear to make ecological and geographical sense: the upland hardwood forest (Figure 3) occurs in areas of higher elevation (in island terms), and on soils having good drainage (Tables 2 and 4). In such areas the map- ping indicates that it is present either as small woodlots surrounded by farmland, or as larger connected parcels (at the back end of farms) — especially in the central and south-eastern hill-lands, where steeper slopes and shallow soils placed restrictions on forest clearance in the past. It is also evident that the Black Spruce forest and the wet rich woodland, which occur primarily on soils with poorer drainage (Tables 2 and 4), are segregated geo- graphically from the upland hardwood forest. These predominate in areas of lower elevation, notably in parts of the east and west of the island (Figures 4 and 5). Though both of these forest-types occur in the same geographical areas, the TWINSPAN analysis indicated that they have very different ground and tree vegetation (Sobey and Glen 2002). They also have very different soil chemical properties — the wet rich woodland has soils that are comparatively base-rich and of higher pH with a lower C/N (carbon-nitrogen) ratio, than the soils of Black Spruce forest, more acid and base-poor with a higher C/N ratio. This dissimilarity was also evident from the fact that there was no overlap in their graphical distributions in the DECORANA ordi- nation (Sobey and Glen 2002). Given their similar geo- graphical distribution on the island, the question is whether in the mapping we have been able, using tree and soil criteria alone, to separate the stands of these two wet forest-types from each other. The differences in their tree composition parameters, evident in Table 3, suggest that we have. Stands of the other two forest-types, the White Spruce woods and the disturbed forest, have a wider and more scattered distribution (Figures 6 and 7), occurring less as larger connected parcels, than as individual small stands, which is consistent with the fact that their occur- rence is largely the product of human activity, which will tend to occur at a scale determined by the own- ership of blocks of land. At the same time there is a geographical aspect to their distribution, with the White Spruce woods prevalent in the central hill-lands and in the eastern part of the island, its spatial distribution evidently determined by the geography of field and farm abandonment, while the disturbed forest is con- 516 THE CANADIAN FIELD-NATURALIST Vol. 118 TABLE 5. For each soil drainage class, the total area (in hectares) under the three ‘primary’ forest stand-types in 1990, and the percentage of this area assigned in the computer-sorting to each stand-type is shown. DRAINAGE CLASS PRIMARY FOREST Poorly- _ Imperfectly Well-drained Well-drained STAND-TYPES Organic drained drained (medium textured) (coarse textured) Rapidly-drained Upland Hardwood Forest 0.88 6.2 12.0 68.7 91.2 64.4 Black Spruce Forest 86.5 35.6 30.7 16.7 6.7 30.6 Wet Rich Woodland 12.6 58.2 57:3 14.6 2a 5.0 Total Hectares 2 620 50 912 18 089 Zl Si 44 865 11 073 centrated largely in the far west and in the east of the island. Although a significant and meaningful ecological picture has emerged from the sorting and mapping, at the same time it is important to point out some limi- tations: a basic one is that any sorting of forest stands into forest-types is a classification procedure, and any process of classification is a simplification. Compound- ing this was the fact that the stand sorting was reliant on the products of three other classification procedures: TWINSPAN, stand delineation (strictly-speaking not a classification procedure, but involving demarcation of the forest area into individual stands), and the soil series classification for the province. Each of these three prior procedures will have their own particular limitations. It should also be pointed out that although three cen- turies of forest destruction and exploitation by Euro- peans have undoubtedly generalized and simplified the island’s forest cover, the five maps are still a sim- plification of what was present in 1990. As noted in Sobey and Glen (2002) there are likely to have been other forest-types present but so uncommon as not to have been picked up in the 1991 field survey in suffi- cient quantity to be recognized as separate types (e.g., areas under Hemlock, White Pine, Eastern White Cedar and White Ash, now occurring as only a few small stands, but likely to have been more widespread in the past). Also, each of the five forest-types that has been recognized may have had sub-types that have been overlooked: e.g., the TWINSPAN classification sub- divided the Black Spruce forest plots into wetter and dryer variants (Sobey 1995), but an attempt in the sort- ing to separate stands of these on their tree canopy com- position was not successful. The upland hardwood may also have variants dominated by particular tree species. The only solution to this problem of both rare and more refined forest-types, is firstly to define them, and then to locate them on the ground and map them separately. It should also be noted that the maps produced present a picture of the forest as it was at one particular time: in the summer of 1990 at the time of the aerial photographic survey. But the forests of Prince Edward Island, like forests the world over, are not fixed and static. Apart from natural factors, they are subject to the effects of continual human interference, for example, clear- and partial-cutting, forest clearance for agricul- ture, fire, and farm abandonment, as well as the natural successional processes resulting from these effects. On the other hand, although the three pre-European settlement woodland types may vary over time in the amount and proportion of the island’s area that they occupy, we should not expect the general pattern of their distributions to change, related as this is to innate soil properties. However, the two successional forest- types (White Spruce woods and disturbed forest) are likely to be only transitional on a site, and we may presume that given sufficient time, most of their sites would develop into one of the primary woodland types. Mapping the pre-European settlement forest of the island Despite the above limitations in the methodology, it is of interest to generalize the results even further by considering the potential for extending the mapping to areas without forest cover in 1990, incidentally, an aspect of all earlier attempts at mapping the province’s forests (Stilgenbauer 1929; Halliday 1937; Rowe 1959; Loucks 1962). Such a mapping may have either of two objectives: (1) the reconstruction of the type of forest likely to have occurred before forest clearance, i.e., the “original-natural” forest sensu Peterken (1996); and/or (2) the prediction of the type of forest that would ultimately occur over the whole island if all areas were allowed to revert naturally to climax forest in the future; i.e., the “future-natural” forest sensu Peterken (1996). Either of these objectives involves trying to predict the climax forest-type on the land area now occupied by White Spruce woodland and the “disturbed forest’, as well as the forest cover on land which in 1990 was clear of forest, i.e., the non-forested area evident in Figure 2, comprising some 43% of the island’s land area (Anonymous 1992). The easiest approach to such a mapping is to take advantage of the close relationship existing between natural vegetation and soils, by utilizing data derived from the comprehensive and fine-scale maps produced by the Prince Edward Island Soil Survey (MacDougall et al. 1988). We need first to examine any associations 2004 SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. S197, FIGURE 8. The conjectured distribution of upland hardwood forest prior to European settlement — based on combining the distribution of upland hardwood stands in 1990 with that of well-drained soils as mapped by the Soil Survey of Prince Edward Island (MacDougall et al. 1988). FIGURE 9. The conjectured distribution of wet rich woodland and Black Spruce forest prior to European settlement — based on combining the distribution of stands of these two forest-types in 1990 with the distribution of imperfectly- and poorly-drained soils (excluding stream complexes) as mapped by the Soil Survey of Prince Edward Island (Mac- Dougall et al. 1988). 518 occurring between the three pre-settlkement forest- types and the soils of different drainage properties: Table 5 shows for each drainage class the percentage of its “primary forest’ component contributed by each of the three primary forest-types. From Table 5 it is evident that well-drained coarse- textured soils (and to a less extent well-drained medium- textured soils) have a strong association with upland hardwood forest (some 91% of the well-drained coarse soils and 69% of the medium soils have such a forest cover). By contrast, poorly and imperfectly drained soils are strongly associated with the two other pre- settlement forest types (wet rich woodland and Black Spruce forest) — some 94% of poorly-drained and 88% of imperfectly-drained sites carry such forests. However, although these two “wet” forest-types (as noted in Sobey and Glen (2002) and above), occupy soils of markedly differing chemical properties (in terms of pH, base status and C/N ratios), such chemi- cal properties were not criteria used to characterize the soil series in the 1988 Soil Survey, and thus the soil series maps cannot be used to differentiate these two forest-types. Then there is the problem presented by the remaining soil drainage class, rapidly-drained soils (Table 5): 64% of such soils carry upland hard- wood forest, but a significant proportion (30%) has a cover of Black Spruce forest, more likely the dryer vari- ant of the Black Spruce forest, shown in the TWINSPAN analysis (Sobey 1995) to have some association with rapidly-drained soils. On the basis of the above relationships, Figure 8 is an attempt to show the approximate distribution of the pre-settlement upland hardwood forest, and pre- sumably also the area that is capable of reverting to such forest in future. The map has been constructed by taking the 1990 distribution of upland hardwood forest (Figure 3) and adding to it the area occupied by well- drained soils (both coarse and medium-textured parent materials). It is likely that additional areas under rapid- ly-drained soils, particularly in the south-eastern hill- lands, would also have carried some form of upland hardwood forest, but given the present state of our information, it is not possible at this stage to distinguish the hardwood-bearing areas of such soils from the Black Spruce areas. Figure 9 is an attempt to show the area occupied before European settlement by the two wet forest-types (the wet rich woodland and the Black Spruce forest). It has been constructed by adding to the 1990 distri- bution of these forest-types (i.e., Figures 4 and 5), the combined distribution of poorly- and imperfectly- drained soils. Given the present state of the informa- tion incorporated within the G.I.S. database for the province, this is about as far as we can go for wet wood- land. Distinguishing between these two woodland types would require data on the chemical properties of the soils. And, as noted above, Table 5 indicates that Black Spruce forest (presumably the dryer variant) will have also occurred on land having rapidly-drained soils. It is THE CANADIAN FIELD-NATURALIST Vol. 118 also likely that it is on such soils that the now largely extirpated pine forests of the island would have occurred in the past. The picture that emerges from these maps is of an island that was mostly covered by upland hardwood forest, which formed a large continuous block in the central part of the island, with smaller but significant blocks in the western and eastern parts. By contrast, the wetter forest-types occurred at lower elevations in specific areas in the west and in a more scattered mosaic pattern in the east. It is thus on these lowland sites that the boreal element of the island’s forests, in the form of Black Spruce forest, had a greater occurrence. Such a division of the landscape, based on soil mois- ture levels, between the northern hardwood forest and the boreal forest, appears to be typical in the boreal- broadleaf ecotone (Pastor and Mladenoff 1992; Scott 1995). It should be noted that compared with earlier attempts at mapping the pre-settlement forest (e.g., Stilgenbauer 1929; Loucks 1962) the maps presented here have a much finer scale of detail in their boundary lines. Also, the validity of the maps, as showing the forest distribution before European settlement, can be test- ed by comparing them with early historical descrip- tions of the forest, especially from the data contained in early maps and surveyors’ field notebooks in the provincial archives. Not only would such a compari- son allow us to ascertain the forest-types that actually occurred on many sites before forest clearance, it may also provide data on the fine-scale differences in the forests of such sites, in some cases even as to par- ticular tree species. However this is beyond the scope of the present study. By contrast, the validity of the maps as pictures of the “future-natural” forest will only be testable from what happens if such areas revert to forest in future. Acknowledgments Doug Sobey thanks Professor Gerry McKenna, then Dean of Science at the University of Ulster, for en- abling him to take six months leave from University duties in 1996 so that he could continue his research into the forests of Prince Edward Island, and the P.E.I. Department of Agriculture and Forestry, Forestry Divi- sion, for allowing full access to the 1990-1992 Forest Inventory data. He also thanks Harry Baglole of the Institute of Island Studies at the University of Prince Edward Island for supporting the continuance of the research associateship which enabled the use of library facilities at the University. Both authors thank Delmar Holmstrom of Agriculture Canada, Charlottetown, for providing advice on the grouping of the soil series into drainage classes. Literature Cited Anonymous. 1992. 1990/1992 Prince Edward Island forest inventory: Summary. Forestry Branch, P.E.I. Department of Energy and Forestry, Charlottetown, Prince Edward Island. 2004 Erskine, D. S. 1960. The plants of Prince Edward Island. Publication 1088. Canada Department of Agriculture. Glen, W. M. 1997. Prince Edward Island 1935/1936 forest cover type mapping. Forestry Division, P.E.I. Department of Agriculture and Forestry, Charlottetown, Prince Edward Island. Halliday, W. E. D. 1937. A forest classification for Canada. Canada Department of Mines & Resources, Ottawa. Loucks, O. L. 1962. A forest classification for the Maritime 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, Prince Edward Island. Pastor, J., and D. J. Mladenoff. 1992. The southern boreal- northern hardwood border. Pages 216-240 in A systems analysis of the global boreal forest. Edited by H. H. Shugart, R. Leemans, and G. B. Bonan. Cambridge University Press, Cambridge, England. Peterken, G. F. 1996. Natural woodland — ecology and con- servation in northern temperate regions. Cambridge Uni- versity Press, Cambridge, England. SOBEY AND GLEN: PRESENT AND PAST FOREST-TYPES OF P.E.I. 519 Rowe, J. S. 1959. Forest regions of Canada. Canada Depart- ment of Northern Affairs and Natural Resources, Forestry Branch, Bulletin 123 [Reprinted 1972 with minor revisions as: Publication No. 1300, Canadian Forestry Service, Depart- ment of the Environment, Ottawa]. Scott, G. A. J. 1995. Canada’s vegetation — a world perspec- tive. McGill—Queen’s University, Montreal and Kingston. 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. Prince Edward Island Department of Agriculture, Fisheries and Forestry, Charlottetown, Prince Edward Island. 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: 585-602. Stilgenbauer, F. A. 1929. The geography of Prince Edward Island. Ph.D. dissertation, University of Michigan, Ann Arbor, Michigan. Watts, S. B. 1983. Forestry handbook for British Columbia (Fourth edition). University of British Columbia, Vancou- ver, British Columbia. Received 20 May 2003 Accepted 4 November 2004 APPENDIX 1. Summary descriptions of the five forest stand-types arising from the computer- sorting of 82 957 forest stands. UPLAND HARDWOOD FOREST 16 170 stands (19.5%); 55 043 ha (21.7%) A stand-type of widespread occurrence on well-drained soils (77% of its area is on such soils — plus an additional 13% on rapidly-drained soils), especially away from the coast and further back from roads, in the central and south- eastern hill-lands, in the north-east peninsula, and in small- er specific areas in the east and west of the island. 94% of its area had a forest cover in 1935. The tree canopy is domi- nated by hardwoods (accounting for 85% of crown closure) — particularly Red Maple (36%), with a notable contribution by Sugar Maple (22%), and smaller amounts of Yellow Birch (8%) and American Beech (2%). There is also a significant component of White Birch (14%) and Trembling Aspen (5%). The minority conifer component (15%) consists largely of Balsam Fir (9%), with some White Spruce (5%). The high contribution to crown closure by Red Maple, and the pres- ence of White Birch and Trembling Aspen, are indicative of a high level of past, and we may presume, continuing, human disturbance. This stand-type is the direct descendant of the pre-European settlement upland hardwood forest postulated by Erskine (1960), though subject to varying levels of past and current human disturbance. BLACK SPRUCE FOREST 8 049 stands (9.7%): 33 106 ha (13.1%) A rather localized stand-type occurring primarily on areas with poor and imperfect drainage — 78% of its area is on such soils — especially in specific parts of the west and east of the island. The tree canopy is dominated by conifers (com- prising 87% of crown closure), above all Black Spruce (71%) — with small contributions by Tamarack (9%) and Balsam Fir (5%). The 13% contributed by hardwoods comes largely from Red Maple (5%) and Trembling Aspen (5%). Most of the area (77%) had a forest cover in 1935. This ‘stand-type corresponds to the pre-settlement lowland Black Spruce forest of Erskine (1960). WET RICH WOODLAND 13 863 stands (16.7%); 42 192 ha (16.6%) A stand-type almost completely confined to areas of poorly- and imperfectly-drained soils (95% by ground area) espe- cially in the west of the island, as well as parts of the east. The tree canopy is dominated by hardwoods (making up 65% of crown closure), but there is also a notable conifer contribution (35%). The chief hardwood species are Red Maple (26%), Trembling Aspen (21%) and White Birch (13%). (White Cedar, American Elm and White Ash are present at low levels). The main conifers are Black Spruce (10%), White Spruce (8%), Balsam Fir (6%) and Tamarack (5%). 76% of its area was under forest cover in 1935. This stand-type appears to be a derivative of the pre-settlement lowland Red Maple forest of Erskine (1960) — though with considerable effects due to cutting and other disturbances. 22 709 stands (27.4%): 57 982 ha (22.9%) A widely scattered stand-type occurring as small often field- shaped parcels on well- and rapidly- drained soils (78% by area is on such soils). It is prevalent in the central hill-lands and even more towards the east of the island. A high pro- portion of its area (62%) was cleared land in 1935. The tree canopy is dominated by a single species, White Spruce, at a high level of crown closure (68%). The remaining species contribute only small amounts: Trembling Aspen (8%), White Birch (6%), Tamarack (6%), Red Maple (5%) and Balsam Fir (4%). This stand-type corresponds to the succes- sional “old field” White Spruce woods of Erskine (1960). WHITE SPRUCE Woops 520 “Disturbed Forest’ 22 166 stands (26.7%): 65 259 ha (25.7%) This residual stand-type (i.e., that left over after the stands of the other four types had been selected) is widely scattered, generally as small individual parcels (rather than large blocks), especially in the east and west of the island, notably on well- and rapidly-drained soils (75% by area occurred on such soils). The tree canopy has a slight predominance of hard- woods (54%) — the principal species being Red Maple (20%), THE CANADIAN FIELD-NATURALIST Vol. 118 Trembling Aspen (19%) and White Birch (13%). Its large conifer component (46%) is made up of Tamarack (13%), White Spruce (10%), Balsam Fir (9%), Red Spruce (7%) and Black Spruce (5%). 33% of its area was cleared land in 1935. This stand-type does not equate with any of Erskine’s pre- European forest-types — its species make-up and distribution suggest rather that it is largely a successional community heavily affected by past and continuing human exploitation. Inventaire printanier d’une frayere multispécifique : l’ichtyofaune des rapides de la riviere Gatineau, Québec ANNIE Comto!s!, FRANCOIS CHAPLEAU!, CLAUDE B. RENAUD?, HENRI FOURNIER*, BRENT CAMPBELL’, et RICHARD PARISEAU? 'Département de biologie, Université d’ Ottawa, Ottawa, Ontario KIN 6N5 Canada Division de la recherche, Musée canadien de la nature, Casier Postal 3443, Succursale D, Ottawa, Ontario K1P 6P4 Canada 3Direction de l’aménagement de la faune de I’ Outaouais, Société de la faune et des parcs du Québec, Gatineau, Québec J8Y 3R7 Canada Comtois, Annie, Frangois Chapleau, Claude B. Renaud, Henri Fournier, Brent Campbell, et Richard Pariseau. 2004. Inventaire printanier d’une frayére multispécifique : ichtyofaune des rapides de la riviere Gatineau, Québec. Canadian Field- Naturalist 118(4): 521-529. Un inventaire ichtyologique printanier a été réalisé dans la riviere Gatineau, dans le premier rapide a l’amont de son embou- chure dans la riviere des Outaouais, pour établir la chronoséquence d’utilisation du site par les especes s’y reproduisant, dont trois espéces de chevaliers. L’échantillonnage des rapides a permis de capturer 2388 poissons répartis en 13 familles et 39 espeéces, dont six ayant fait ’ objet d’une évaluation par le COSEPAC. Seize espéces étaient en état de se reproduire lors de |’ échantil- lonnage. Nous avons observé une reproduction plus ou moins simultanée pour le Chevalier blanc et le Chevalier rouge, suivie de celle du Chevalier de riviére. Egalement, pour ces trois espéces, nous avons observé la capture des miles de stade V plus tot que les femelles, et nous les retrouvons en plus grand nombre. Ceci pourrait s’expliquer du fait que les males précedent les femelles sur les frayeéres, que celles-ci sont reconnues pour rester tres peu de temps sur les sites de fraie et qu’un minimum de deux males par femelle est requis pour la reproduction. Cette étude démontre que les rapides de la riviére Gatineau doivent étre protégés puisqu’ils sont habités par trois especes qui ont obtenu un statut spécial du COSEPAC (Menacée: Fouille-roche gris; situation préoccupante : Lamproie du nord et Chevalier de riviere) en plus de constituer une frayére multispécifique importante. Mots-clés: Rapides, Riviere Gatineau, Outaouais, Fraie, Chevalier blanc, Moxostoma anisurum, Chevalier de riviére, Moxostoma carinatum, Chevalier rouge, Moxostoma macrolepidotum, Lamproie du nord, [chthyomyzon fossor, Fouille-roche gris, Percina copelandi. A spring survey of fishes was conducted in the Gatineau River (Quebec), in the first rapids upstream of its confluence with the Ottawa River, to establish the fish spawning chronosequence, especially for the three species of redhorses. The sampling resulted in the capture of 2388 fish belonging to 13 families and 39 species. Six of these species have been assessed by COSEWIC. Sixteen species were in spawning readiness. Spawning of the Silver Redhorse and Shorthead Redhorse occurred first, followed by the River Redhorse. Stage V males of the three redhorse species were captured earlier and in greater numbers than females. These observations are consistent with other studies which suggest that males precede females to the spawning grounds, that females remain there for a short period and that a minimum two males per female is required for reproduction. This study shows that the Gatineau River rapids must be protected because they harbor three species that have obtained a status by COSEWIC (Threatened: Channel Darter; Special concern: Northern Brook Lamprey and River Redhorse) as well as being an important multi-species spawning ground. Key Words: Rapids, Gatineau River, Outaouais, spawning, Silver Redhorse, Moxostoma anisurum, River Redhorse, Moxostoma carinatum, Shorthead Redhorse, Moxostoma macrolepidotum, Northern Brook Lamprey, Ichthyomyzon fossor, Channel Darter, Percina copelandi. Un inventaire réalisé au printemps 1989 a l’aide de filets de dérive a permis de constater que les rapides de la riviére Gatineau en aval du barrage hydroélec- trique de Farmer’s Point (Gatineau, Québec) (Figure 1) sont utilisés pour la reproduction de I’ Esturgeon jaune, des Percidae (Sander spp. et Perca flavescens), et des Catostomidae (Catostomus spp. et Moxostoma spp.) (Houde et Fournier 1992). Une péche expérimentale réalisée en 1998 (données non publiées) a révélé la présence a ce site d’une concentration exceptionnelle de Chevalier de riviére. Des travaux furent donc entre- pris en 1999 pour décrire cette population et son utili- sation du rapide Farmer’s. L étude ichtyologique de ce secteur de la riviere revét un intérét particulier parce que le barrage agit comme barriére au déplacement des poissons vers l’amont. Ceci pourrait occasionner une concentration de poissons dans le secteur des rapides en aval du barrage, en particu- lier au printemps, lors des migrations reproductrices. Ainsi, le secteur en aval du barrage pourrait étre un site privilégié pour la reproduction des poissons provenant des riviéres Gatineau et des Outaouais. Le but de notre étude est de présenter |’inventaire printanier des espéces de poissons de ce secteur de la riviére alors que plusieurs de celles-ci sont normale- ment en période de fraie. De plus, une attention particu- 521 522 45° 29'21"N 75° 45'17°O t po Sz AUX THE CANADIAN FIELD-NATURALIST Vol. 118 45°28'42°N 75°44'13°O outa ONTARIO Figure 1. Site de la riviere Gatineau prés du pont Alonzo-Wright, se trouvant a 5,2 km de la jonction avec la riviére des Outaouais. liere sera portée sur la répartition dans le temps de la reproduction des trois especes de chevalier (Moxos- toma spp.) qui utilisent le secteur des rapides pour frayer. Site d’échantillonnage La section de la riviére Gatineau (45°29'N, 75°44'O) échantillonnée se trouve entre 6,0 et 5,2 km de l’embouchure sur la riviere des Outaouais (Figure 1). La zone d’échantillonnage couvre une longueur de 800 m et est délimitée en amont par le pont Alonzo- Wright et a l’aval par l’extrémité d’une ile localisée prés de la rive ouest de la riviére. La largeur de la riviére dans ce secteur varie de 170 m en amont 4 330 m en aval et couvre une superficie d’environ 22 ha. La pro- fondeur de |’eau varie de 0,5 a 5 m. Le courant est particuliérement rapide sur une distance d’environ 400 m a l’aval du pont et la vitesse devient beaucoup plus lente par la suite alors que la riviére s’élargit et devient plus profonde. Dans les eaux rapides, le substrat est composé de galets, de cailloux et de gros blocs épars dans le centre de la riviére et de gravier en rive ouest. La rive est a été largement modifiée par des remblayages et la construction de quais et murs de soutenement. En aval, ot la riviére est plus large et le courant plus lent, le fond est constitué de gravier, de cailloux et de galets; en rive ouest, le substrat est composé de sable 2004 CoMTOIS ET AL. : L- ICHTYOFAUNE DE LA RIVIERE GATINEAU 523 TABLEAU I. Espéces recensées lors de l’échantillonnage du rapide Farmer’s, riviére Gatineau, au printemps 1999. Famille Nom latin (Espeéce) Nom commun frangais Nombre d’individus Petromyzontidae Ichthyomyzon unicuspis Lamproie argentée 8 Ichthyomyzon fossor Lamproie du nord | Lampetra appendix Lamproie de I’est 3] Acipenseridae Acipenser fulvescens Esturgeon jaune 19 Hiodontidae Hiodon tergisus Laquaiche argentée 276 Anguillidae Anguilla rostrata Anguille d’ Amérique | Cyprinidae Luxilus cornutus Méné a nageoires rouges 4 Notemigonus crysoleucas Méné jaune D) Notropis atherinoides Méné émeraude 309 Notropis hudsonius Queue a tache noire 2 Rhinichthys cataractae Naseux des rapides | Semotilus corporalis Ouitouche 52 Catostomidae Carpiodes cyprinus Couette 68 Catostomus commersoni Meunier noir 33 Moxostoma anisurum Chevalier blanc 223 Moxostoma carinatum Chevalier de riviere 270 Moxostoma macrolepidotum Chevalier rouge 254 Ictaluridae Ameiurus nebulosus Barbotte brune 9 Ictalurus punctatus Barbue de riviére ay Noturus insignis Chat-fou liséré 3 Esocidae Esox lucius Grand brochet l Esox masquinongy Maskinongé 4 Salmonidae Salmo trutta Truite brune | Salvelinus namaycush Touladi l Percopsidae Percopsis omiscomaycus Omisco 5 Centrarchidae Ambloplites rupestris Crapet de roche 116 Lepomis gibbosus Crapet-soleil 9 Lepomis macrochirus Crapet arlequin 4 Micropterus dolomieu Achigan a petite bouche 17 Micropterus salmoides Achigan a grande bouche 1 Pomoxis nigromaculatus Marigane noire 22 Percidae Etheostoma nigrum Raseux-de-terre noir 1 Etheostoma olmstedi Raseux-de-terre gris 12 Perca flavescens Perchaude 79 Percina caprodes Fouille-roche zébré 315 Percina copelandi Fouille-roche gris 76 Sander canadensis Doré noir 16 Sander vitreus Doré jaune 80 Sciaenidae Aplodinotus grunniens Malachigan 5 et de limon alors que la rive est a aussi été largement modifiée par les activités humaines. Méthodes Léchantillonnage s’est effectué a intervalle de 1 a 5 jours, du 29 avril au 23 juin 1999, entre 19:00 h et 01:00 h. Il s’est fait a aide d’un bateau de péche élec- trique de marque Smith Root 16R. Des voltages de 250 a 1000 V et des ampérages de 0,25 a 3,00 A ont été utilisés selon la conductivité de l'eau. La péche débutait a 20 m des piliers du pont Alonzo-Wright. Une dérive contrélée du bateau était faite sur des tran- sects de 400 a 800 m le long des rives et au centre de la riviere. La période de péche active (lorsque le courant était émis dans |’ eau) variait de 320 4 1732 s. Les pois- sons capturés étaient déposés dans un vivier installé sur le pont du bateau. Une fois le vivier rempli, la péche active était arrétée et les poissons étaient examinés et relachés, sauf pour quelques spécimens qui ont été conservés pour des études ultérieures. Les poissons étaient identifiés, dénombrés et le sexe ainsi que le stade de reproduction étaient déterminés lorsque cela était possible. Pour les chevaliers, le sexe était établi a partir de la présence de tubercules nuptiaux (Scott et Crossman 1974). Le stade de reproduction était déterminé par des caractéristiques morphologiques externes et selon l’index de maturité des gonades de Nikolsky (1963). Celui-ci classe les stades en six catégories. Les stades I, II et Ill, ne sont pas identifiables puisqu ils refletent des états de maturation des gonades qui ne montrent pas de signes externes. Le stade IV est caractérisé par l’identification de gonades fermes et gonflées au toucher de la paroi abdominale sans expulsion d’ceufs ou de sperme. Le stade V est identifié lorsqu’il y a expulsion de produits sexuels par simple pression de l’abdomen. 524 20 — on Nombre d'espéces capturées 3 THE CANADIAN FIELD-NATURALIST 0 0,0 Vol. 118 =i Ol ° = oO oO oa fo) 29-avr 3-mai 6-mai 10-mai 13-mai 17-mai 20-mai 25-mai 27-mai 31-mai 3-juin 7-juin 10-juin 14-juin 16-juin 17-juin 21-juin 23-juin Date FicuRE 2. Evolution temporelle du nombre d’espéces capturées et de la température de l’eau (m) au rapide Farmer’s, printemps 1999. Un poisson au stade VI a un abdomen relativement flasque et sans expulsion massive de produits sexuels par pression abdominale. L’échantillonnage ciblant surtout les chevaliers, |’a- bondance des espéces de petite taille est probablement une sous-estimation de leur présence dans le milieu. Résultats Familles et espéces capturées L’échantillonnage des rapides de la riviére Gatineau a permis de capturer 2388 poissons en 12 heures de péche active réparties sur 18 jours entre le 29 avril et le 23 juin 1999. Les captures se répartissent en 13 fa- milles et 39 espéces (Tableau 1). Les Percidae dominent pour le nombre d’espéces capturées (sept espéces). Les Cyprinidae et les Centrarchidae sont représentés par six espéces chacune. Les Catostomidae dominent pour le nombre de spécimens (848), les chevaliers constituant 88% des captures. Un seul spécimen des Anguillidae (Anguilla rostrata) a été capturé (Tableau 1). D’ailleurs, un seul spécimen a été capturé pour huit especes (Tableau 1) alors que plus de 300 spécimens ont été capturés pour le Méné émeraude (309) et le Fouille-roche zébré (315). Six des espéces capturées ont fait l’objet d’une évaluation par le Comité sur la situation des espéces en péril au Canada (COSEPAC 2003) : l’Esturgeon jaune (non en péril; 1986), le Chevalier de riviére (situ- ation préoccupante; 1987), le Chat-fou liséré (menacé; 1989 et données insuffisantes lors de la révision de 2002), la Lamproie du nord (situation préoccupante; 1991), le Fouille-roche gris (menacé; 1993, 2002) et le Raseux-de-terre gris (non en péril; 1993). Deux de ces especes, le Chevalier de riviére et le Fouille-roche gris ont été capturés en grand nombre dans la riviere Gatineau, soit 270 et 76 individus respectivement (Tableau 1). Comme nous pouvons le constater a la Figure 2, le plus petit nombre d’espéces capturées a été de dix lors de la premiere sortie, le 29 avril. Il a augmenté graduellement pour atteindre un maximum de 21 es- peces le 27 mai. Par la suite, le nombre a diminué, vari- ant de 12 a 17 espéces par sortie. Reproduction Des individus de 16 des espéces capturées étaient au stade V du cycle de reproduction et semblent donc y avoir frayé (Tableau 2). Pour plusieurs de ces espéces, nos captures permettent de délimiter une période de reproduction probable et les températures auxquelles elle se serait produite. Pour quelques espéces, des in- dividus en état de fraie (stade V) étaient présents dés la premiere journée d’échantillonnage ou encore pré- sents la derniére journée. Dans ces cas, il est possible que la période de reproduction s’étende au-dela de la période d’échantillonnage. Des individus des deux sexes de tous les stades avancés de développement des gonades (stades IV, V, VI) ont été observés pour cing especes seulement : le Chevalier blanc (Moxostoma anisurum), le Chevalier de riviere (M. carinatum), le Chevalier rouge (M. macrolepidotum), la Couette (Car- piodes cyprinus) et la Laquaiche argentée (Hiodon tergisus). Reproduction des trois espéces de chevalier L’abondance des captures permet d’établir la chro- noséquence des périodes de reproduction des trois especes de chevaliers dans la riviére Gatineau (Figures 3 et 4). Pour toutes les espéces, des spécimens de 2004 COMTOIS ET AL. : L;: ICHTYOFAUNE DE LA RIVIERE GATINEAU Nn i) Nn TABLEAU 2. Période de capture d’individus en fraie (stade V), selon le sexe et ’espece, au rapide Farmer’s, printemps 1999. Espéce Nombre d’individus fe) Q Ambloplites rupestris 2 3 Aplodinotus grunniens | | Carpiodes cyprinus 35 7 Catostomus commersoni 8 5 Hiodon tergisus 198 2 Ichthyomyzon fossor 0 | Ichthyomyzon unicuspis 2 Lampetra appendix 5 3 Moxostoma anisurum 89 6 Moxostoma carinatum 96 16 Moxostoma macrolepidotum 67 7 Percina caprodes 10 0 Percina copelandi d/ 1 Percopsis omiscomaycus 0 2 Pomoxis nigromaculatus 3 3 1 1 Période de capture Température de eau (°C)! 27 mai — 10 juin 13,5 — 18,5 3— 6 mai 9,0 — 10,0 29 avril — 3 juin 7,0 — 18,0 29 avril — 27 mai 7,0 — 13,5 10 mai — 23 juin 12,5 — 19,0 23 juin 19,0 13 mai — 23 juin 12,5 — 19,0 3 — 20 mai 9,0 — 14,0 3 — 31 mai 9,0 — 17,0 10 mai — 23 juin 12,5 — 19,0 29 avril — 21 juin 7,0 — 19,0 6 — 25 mai 10,0 — 14,0 20 mai — 21 juin 14,0 — 19,0 6 mai 10,0 3 — 13 mai 9,0 — 12,5 3 — 20 mai 9,0 — 14,0 Semotilus corporalis ' Température de l’eau au début et a la fin de la période de capture. stade de maturité IV étaient présents au début de la période d’échantillonnage, le 29 avril, sauf pour les femelles du Chevalier rouge et les males du Cheva- lier de riviere. Pour ceux-ci, les individus au stade [V sont apparus le 3 et le 6 mai respectivement (Figure 3). Des males de stade IV ont été capturés jusqu’au 17 mai dans le cas du Chevalier rouge, jusqu’au 27 mai pour le Chevalier blanc et jusqu’au 7 juin pour le Chevalier de riviere. Les dernieres femelles de stade IV de Chevaliers rouge et blanc ont été capturées le 17 mai, alors qu’elles ont été présentes sur le site jusqu’au 17 juin pour le Chevalier de riviere. (8 enna (34) > a 3 af 7) ee 0 i ee ey o bad ae = > (7) = 3 (96) —— ee Deuces BS ® (16) Qe... 8 SSS SSS STC | D Se) 2 s — (24) | © 3 Ca NER SE CREE RUT SEL Se a Le Ee 2) D) see SSS SSS ES | f - + - } — + —— — : oo 26-avr 1-mai 6-mai 11-mai 16-mai 21-mai 26-mai 31-mai 5-juin 10-juin 15-juin 20-juin 25-juin Date FIGURE 3. Période de reproduction des trois espéces de chevalier selon l’index de maturité de Nikolsky (1963) dans la riviére Gatineau, au printemps 1999. Le carré représente le Chevalier blanc (Mi), le losange, le Chevalier de riviére (@) et le triangle, le Chevalier rouge (a). Les symboles pleins désignent les femelles tandis que les vides représentent les males. Le nombre d’individus capturés est indiqué entre parenthéses. Les lignes verticales correspondent au début et a la fin de la période d’échantillonnage. Le pointillé 4 l’extérieur de la période d’échantillonnage signifie que des individus appartenant aux stades indiqués ont pu se retrouver sur la frayére avant ou apres cette période. 526 Des males au stade V sont apparus dans les cap- tures le 3 mai pour le Chevalier blanc et le 10 mai pour le Chevalier de riviére, 4 une température de 9,0 et 12,5 °C respectivement. Les males Chevalier rouge de ce stade de maturité étaient présents des le début de l’échantillonnage. Les femelles de stade V sont appa- rues le 10 mai pour le Chevalier blanc et le Chevalier rouge (12,5 °C) et le 3 juin pour le Chevalier de rivi- ére (18,0 °C). Les captures de males au stade V sont toujours nettement plus abondantes que celles des fe- melles. Aucune femelle de stade V n’a été capturée aprés le 20 mai dans le cas du Chevalier blanc (14,0 °C) et le 7 juin dans le cas du Chevalier rouge (18,5 °C) alors qu’elles étaient toujours présentes a la fin de I’é- chantillonnage, le 23 juin, dans le cas du Chevalier de riviere, 4 une température de 19,0 °C. Des femelles au stade VI apparaissent dans les cap- tures le 17 mai pour le Chevalier blanc et le Chevalier rouge (12,5 °C) alors qu’elles sont présentes pour le Chevalier de riviére 4 compter du 10 juin (18,5 °C). Les dernieres femelles de stade VI ont été capturées le 31 mai pour le Chevalier blanc (17,0 °C) et le 17 juin pour le Chevalier rouge (19,0 °C), alors qu’elles étaient toujours présentes a la derniere date d’échantillonnage pour le Chevalier de riviere (19,0 °C). Discussion Si l’on ajoute aux 16 espéces citées dans le Tableau 2 celles rapportées par Houde et Fournier (1992), il est probable qu’environ 20 espéces se reproduisent dans le rapide Farmer’s. D’autres espéces capturées pourraient potentiellement utiliser le secteur échantillonné de la riviere Gatineau pour frayer (Scott et Crossman 1974; Bernatchez et Giroux 2000) : le Méné émeraude, le Naseux des rapides, et possiblement le Méné a nage- oires rouges et le Raseux-de-terre gris. Toutefois, la période d’échantillonnage ne correspondait pas a la période de fraie pour certaines espéces (e.g. Méné émeraude) alors que trop peu de spécimens ont été cap- turés pour d’autres afin de déterminer |’ état reproduc- tif (e.g. Naseux des rapides). La frayére multispécifique des rapides de la riviére Gatineau compte autant d’espéces que celle de la riviere aux Pins, a Boucherville (Massé et al. 1988). Les deux frayeres ont toutefois des caractéristiques trés différentes; la frayére de la riviére aux Pins constitue un habitat d’eau calme utilisé particuliérement par les espéces phytophyles. D’ailleurs, seulement trois especes sont communes aux deux frayéres : la Oui- touche, le Meunier noir et le Crapet de roche. Dans les eaux vives, Massé et al. (1988) mentionnent cing autres frayéres entre Carillon (riviére des Outaouais) et la riviere des Prairies qui comptent entre 8 et 27 especes reproductrices. La riviére Gatineau se situe donc dans le milieu du peloton quant a son utilisation multispécifique. Des femelles en état de fraie de Chevaliers rouge et blanc sont apparues sur le site en méme temps le 10 THE CANADIAN FIELD-NATURALIST Vol. 118 mai. Elles y ont été capturées pendant une période de 10 jours dans le cas du Chevalier blanc et de 28 jours dans le cas du Chevalier rouge. La fraie des deux es- peces a donc été en partie simultanée (Figures 3 et 4). Contrairement a nos résultats, Curry et Spacie (1984) ont noté que le Chevalier blanc a amorcé la reproduc- tion une semaine avant le Chevalier rouge dans le Deer Creek, Indiana, en 1979. Toutefois, Mongeau et al. (1986, 1992) ont également observé une simultanéité de la reproduction du Chevalier blanc et du Chevalier rouge dans leur étude sur la riviere Richelieu. En effet, ils ont observé que la reproduction de ces deux espéces débutait 4 la mi-mai pour se terminer a la mi-juin. Nos données semblent tendre vers le méme résultat puisque des individus de stade VI ont été capturés du 17 mai au 21 juin pour les deux espéces (Figure 3). Mongeau et al. (1986, 1992) ont noté que le Chevalier rouge et le Chevalier blanc commengaient a déposer des ceufs a une température de 10,7 °C. Meyer (1962) a indiqué que la température a laquelle la reproduction débutait pour le Chevalier rouge et le Chevalier blanc sur la riviere Des Moines, en Iowa, était 11,1°C et 13,3°C respectivement. Par contre, Burr et Morris (1977) ont observé une température de 16,0°C pour la reproduc- tion du Chevalier rouge dans le ruisseau Big Rock, en Illinois et Hackney et al. (1970), une température de 14,4°C pour le Chevalier blanc dans la riviere Flint, en Alabama. La Figure 2 montre que la température de l’eau au 17 mai était a 12,5°C, ce qui correspond relativement bien aux données de Meyer (1962) et de Mongeau et al. (1986, 1992). Burr et Morris (1977) indiquent que plusieurs es- peces de chevaliers pourraient possiblement frayer a des périodes et des températures similaires et sur des substrats comparables. II peut toutefois y avoir de la ségrégation spatiale entre les espéces. En effet, le Che- valier rouge et le Chevalier doré (Moxostoma erythru- rum) ont été observés en train de frayer simultanément au méme site, mais le Chevalier doré avait une dis- tribution plus étendue en amont, ce qui lui fournissait un habitat non utilisé par le Chevalier rouge (Curry et Spacie 1984). Par ailleurs, une étude comparant la fraie chez le Chevalier doré et le Chevalier noir (Moxostoma duquesnei) a démontré que ces deux espéces se repro- duisaient au méme moment et dans la méme section du Stony Creek, en Illinois (Kwak et Skelly 1992). Les auteurs mentionnent toutefois qu’il existe des dif- férences dans le choix des microhabitats de fraie et dans le comportement reproducteur et que celles-ci servi- raient possiblement a réduire la compétition entre ces espeéces. Bien qu’une étude plus approfondie de l’habi- tat et du comportement reproductif soit nécessaire, il est possible qu’une ségrégation spatiale et/ou compor- tementale se produise pour le Chevalier blanc et le Chevalier rouge dans la riviére Gatineau. Des femelles de stade V du Chevalier de rivieére ont été capturées pour la premiere fois le 3 juin et ce, jusqu’a la derniére journée d’échantillonnage. Sa re- 2004. COMTOIS ET AL. : L;- ICHTYOFAUNE DE LA RIVIERE GATINEAU ayy 35 + Chevalier blanc | Femelle: n= 51 30 7 Male: n= 94 25 Individus non sexés: n= 78 — nn a i Femelle [_] Male Ue eee 25 7 Chevalier de riviére Femelle: n = 110 Male: n = 133 w” @ ie = 20 e | Individus non sexés: n = 27 a) j 3 15 2 GT = = 1055; 5 i § £ Sy 6 ; Zz ] 0 20 7 Chevalier rouge Femelle: n= 70 Male: n= 108 15 - Individus non sexés: n= 76 | 1 10 | ae Oo on cs eee peipiccos nace DS rae are ae) | Sea = i i ke juin [Cl |e ee] a Ln lan => OCG G cs eS - SS SS Ss = gee ee eo Ssocas is A Se SAAN: SAN FiGurE 4. Nombre d’individus capturés par jour selon le sexe pour les trois espéces de chevalier, rivi¢re Gatineau, printemps, 1999. 528 production débute donc plus tard, soit autour du 10 juin (Figure 3) a une température de 18,5 °C. Mongeau et al. (1986, 1992) ont obtenu plus ou moins les mémes résultats puisqu’ils indiquent que le Chevalier de riviere amorce la fraie autour de Ja deuxi¢me semaine de juin a une température de 17,7 °C, une conclusion fondée sur la capture de deux spécimens au stade VI. Hackney et al. (1970) mentionnent également que le Chevalier de riviére a une reproduction tardive vis-a-vis de celle du Chevalier blanc dans la riviére Duck, au Tennessee. La différence entre la période de reproduction de cette espece et celles des deux autres chevaliers présents dans les rapides de la riviére Gatineau est probablement due au fait que le Chevalier de riviére nécessite une température de |’eau plus élevée pour débuter sa reproduction. Par ailleurs, pour ces trois espéces, nous observons des males de stade V a une date plus hative que les femelles au méme stade (Figure 3). Ceci s’explique par le fait que les males du Chevalier blanc, du Chevalier rouge et du Chevalier de riviere précédent les femelles sur les frayeres (Meyer 1962; Hackney et al. 1968, 1970; Scott et Crossman 1974). Ce phénoméne est d’ailleurs observé chez d’autres chevaliers et chez plusieurs autres Catostomidae (Meyer 1962; Scott et Crossman 1974; Jenkins et Jenkins 1980; Page et Johnston 1990; Kwak et Skelly 1992; Cooke et Bunt 1999), Dans nos captures, le nombre de males de stade V est plus grand que celui de femelles pour les trois espéces de chevalier. Ceci n’est pas inhabituel. Hackney et al. (1970) ont observé un rapport des sexes de 4 males : | femelle sur une frayére de Chevalier blanc. De plus, Meyer (1962) note que chez les Chevaliers rouge et blanc, les femelles demeurent sur les sites de fraie beaucoup moins longtemps que les males. La fraie chez les chevaliers, et chez la plupart des Catostomidae, se fait généralement en présence de deux males et d’une femelle (Hackney et al. 1968; Burr et Morris 1977; Jenkins et Jenkins 1980; Page et Johnston 1990; Kwak et Skelly 1992; Cooke et Bunt 1999), d’ot la nécessité d’une présence en plus grand nombre de males sur la frayére tel que nous l’avons observé pour chacune des trois espéces présentes au rapide Farmer’s. Nos travaux ont permis de découvrir le site de re- production de l’une des plus importantes populations de Chevalier de riviére. Un échantillonnage sur quel- ques années serait nécessaire pour préciser la chrono- séquence de fraie de cette espéce et des autres che- valiers utilisant le site. Un échantillonnage sur une plus longue période aurait probablement pu permettre de confirmer la fraie au rapide Farmer’s d’un plus grand nombre d’espéces. Cette étude démontre donc que le rapide Farmer’s constitue une frayére multispécifique importante tant par le nombre d’espéces qui s’y reproduisent que par la présence d’espéces évaluées par le COSEPAC comme ayant une situation préoccupante (Lamproie du nord, Chevalier de riviére) ou méme menacée (Fouille-roche gris). Localisés en plein coeur d’une zone urbaine, ces THE CANADIAN FIELD-NATURALIST Vol. 118 rapides ont une grande valeur écologique et doivent étre protégés. Remerciements Nous remercions Anne Phelps, Annie-Chantal Gui- bord et Kevin Moon pour leur aide sur le terrain. Cette recherche a été partiellement subventionnée par une subvention a la découverte du Conseil de recherches en sciences naturelles et en genie du Canada (CRSNG) [Frangois Chapleau]. Littérature Citée Bernatchez, L., et M. Giroux. 2000. Les poissons d’eau douce du Québec et, leur répartition dans l’Est du Canada. Broquet, Boucherville, Québec. 350 pages. Burr, B. M., et M. A. Morris. 1977. Spawning behavior of the Shorthead Redhorse, Moxostoma macrolepidotum, in Big Rock Creek, Illinois. Transactions of the American Fish- eries Society 106: 80-82. Cooke, S. J., et C. M. Bunt. 1999. Spawning and reproduc- tive biology of the Greater Redhorse, Moxostoma valen- ciennesi, in the Grand River, Ontario. Canadian Field- Naturalist 113: 497-502. COSEPAC. 2003. Résultats des évaluations du COSEPAC, novembre 2003. Comité sur la situation des espéces en péril au Canada, Ottawa, Ontario. 54 pages. Curry, K. D., et A. Spacie. 1984. Differential use of stream habitat by spawning catostomids. American Midland Nat- uralist 111: 267-279. Hackney, P.A., W.M. Tatum, et S.L. Spencer. 1968. Life his- tory study of the River Redhorse, Moxostoma carinatum (Cope), in the Cahaba River, Alabama, with notes on the management of the species as a sport fish. Proceedings of the Annual Conference of the Southeastern Association of Game and Fish Commissioners 21: 324-332. Hackney, P. A., G. R. Hooper, et J. F. Webb. 1970. Spawn- ing behavior, age and growth, and sport fishery for the Silver Redhorse, Moxostoma anisurum (Rafinesque), in the Flint River, Alabama. Proceedings of the Twenty-Fourth Annual Conference, Southeastern Association of Game and Fish Commissioners 1970: 569-576. Houde, P. et H. Fournier. 1992. Travaux de recherche de frayéres dans la riviére des Outaouais et ses principaux affluents au cours du printemps 1989. Ministére du Loisir, de la Chasse et de la Péche, Service de l’aménagement et de l’exploitation de la faune, Hull. 28 pages. Jenkins, R. E., et D. J. Jenkins. 1980. Reproductive behavior of the Greater Redhorse, Moxostoma valenciennesi, in the Thousand Islands region. Canadian Field-Naturalist 94: 426-430. Kwak, T. J., et T. M. Skelly. 1992. Spawning habitat, behav- ior, and morphology as isolating mechanisms of the Gol- den Redhorse, Moxostoma erythrurum, and the Black Red- horse, M. duquesnei, two syntopic fishes. Environmental Biology of Fishes 34: 127-137. Massé, G., R. Fortin, P. Dumont, et J. Ferraris. 1988. Etude et aménagement de la frayere multispécifique de la rivi- ére aux Pins et dynamique de la population de Grand bro- chet, Esox lucius L., du fleuve Saint-Laurent, Boucher- ville, Québec. Ministére du Loisir, de la Chasse et de la Péche, Service de l’aménagement et de l’exploitation de la faune, Montréal, Québec, Rapport technique 06-40. 224 pages. 2004 Meyer, W. H. 1962. Life history of three species of redhorse (Moxostoma) in the Des Moines River, Iowa. Transactions of the American Fisheries Society 91: 412-419. Mongeau, J.-R., P. Dumont, et L. Cloutier. 1986. La biolo- gie du Suceur cuivré, Moxostoma hubbsi, une espeéce rare et endémique a la région de Montréal, Québec, Canada. Ministére du Loisir, de la Chasse et de la Péche, Service de l’aménagement et de l’exploitation de la faune, Montréal, Québec, Rapport technique 06-39. 137 pages. Mongeau, J.-R., P. Dumont, et L. Cloutier. 1992. La biolo- gie du Suceur cuivré (Moxostoma hubbsi) comparée a celle de quatre autres espéces de Moxostoma (M. anisurum, M. carinatum, M. macrolepidotum, M. valenciennesi). Revue canadienne de zoologie 70: 1354-1363. COMTOIS ET AL. : L- ICHTYOFAUNE DE LA RIVIERE GATINEAU 529 Nikolsky, G. V. 1963. The ecology of fishes. Traduit par L. Birkett. Academic Press, New York. 352 pages. Page, L. M., et C. E. Johnston. 1990. Spawning in the Creek Chubsucker, Erimyzon oblongus, with a review of spawn- ing behavior in suckers (Catostomidae). Environmental Biology of Fishes 27: 265-272. Scott, W. B., et E. J. Crossman. 1974. Poissons d’eau douce du Canada. Office des recherches sur les pécheries du Canada Bulletin 184. 1026 pages. Received 3 October 2003 Accepted 21 June 2004 Moths and Butterflies (Lepidoptera) of the Boreal Mixedwood Forest near Lac La Biche, Alberta, Including New Provincial Records Grea R. Pout!, Davip W. LANGor!, JEAN-FRANCOIS LANDRY’, and JOHN R. SPENCE? ' Natural Resources Canada, Canadian Forest Service, 5320 - 122 St., Edmonton, Alberta T6H 3S5 Canada > Agriculture and Agrifood Canada, Eastern Cereal and Oilseed Research Centre, 960 Carling Avenue, Ottawa, Ontario K1A 0C6 Canada > Department of Renewable Resources, University of Alberta, Edmonton, Alberta T6G 2H1 Canada Pohl, Greg R., David W. Langor, Jean-Francois Landry, and John R. Spence. 2004. Moths and Butterflies (Lepidoptera) of the boreal mixedwood forest near Lac La Biche, Alberta, including new provincial records. Canadian Field-Naturalist 118(4): 530-549. Lepidoptera were collected, primarily via UV light trap, for three seasons in the boreal mixedwood forest near Lac La Biche, Alberta. A total of 11 111 specimens were collected, representing 41 families and 438 species. A species list with flight times is presented. The total Lepidoptera community was estimated to be 546 +23.34 species. Abundance and species richness peaked in late July. Thirty-five species constitute new records for Alberta, while one species, Acanthopteroctetes bimaculata, is a new record for Canada, and the first record of the family Acanthopteroctetidae in Canada. Key Words: Moths, butterflies, Lepidoptera, Lac La Biche, Alberta, flight times, Acanthopteroctetes bimaculata, new to Canada. Alberta is blessed with an abundance and variety of forested lands. A large portion of the province is covered by the boreal mixedwood (Figure |) domi- nated by Trembling Aspen (Populus tremuloides), Balsam Poplar (P. balsamifera), and White Spruce (Picea glauca) and containing lesser amounts of White Birch (Betula papyrifera), Balsam Fir (Abies bal- samea), and other species (Beckingham and Archibald 1996). Over the last 15 years Trembling Aspen has increased greatly in value as a commercial tree species and there has been a large increase in forestry activity centered on this resource. A large portion of Alberta’s aspen mixedwood forest is now scheduled for har- vesting over the next 30-40 years (Pratt and Urquhart 1994). It is a major concern that our knowledge of these forests is relatively poor and there is little empirical basis for predicting the impacts of harvesting and other development on non-timber values such as biodiversity. In an effort to obtain baseline data for assessing impacts of forestry practices on biodiversity, and to determine whether old aspen stands in mixedwood forests contain unique species, a multi-agency team of scientists studied the structure and composition of biotic assemblages in aspen-dominated forests of various ages in the vicinity of Lac La Biche, Alberta, between 1993 and 1995. This work is among the most comprehensive biotic inventories in aspen forests in the province. Lists of plants, birds, amphibians, and mammals have been published by Stelfox (1995) and those of ground-dwelling beetles and dead wood- inhabiting beetles by Spence et al. (1997) and Ham- mond (1997), respectively. As part of that study, but- terflies and moths were also sampled. This represents the first concerted effort to inventory Lepidoptera, especially moths, in aspen forests in western Canada. Lepidoptera constitute a major component of boreal forest biodiversity (Danks and Foottit 1989), and are important herbivores and pollinators (Scoble 1992). The abundance of new provincial records among micro- moths (defined here as the primitive and monotrysian groups, and the lower ditrysian superfamilies up to and including the Pyraloidea and Thyridoidea sensu Kristensen 1999) indicates how poorly sampled these groups have been. The attached checklist and flight times provide baseline information for comparison to other studies and to aid in future research. Materials and Methods The study area (Figure 1) is located in Lakeland Provincial Park near Touchwood Lake, east of Lac La Biche (54°51'N, 111°27'W) in the Central Mixed- wood subregion of the Boreal Forest Natural Region (Beckingham and Archibald 1996). Lepidoptera were sampled in two stands: a 65-year-old (“mature”) stand of 269 Ha, containing 83% cover of Trembling Aspen, 15% Balsam Poplar, 2% willow (Salix spp.), and 1% White Birch, and a stand over 130 years old (“old”) of 148 Ha, containing 54% cover of Trembling Aspen, 32% White Birch, 11% Balsam Poplar, and 3% willow. An inventory of vascular plants found around these study sites is included in the lists published by Stelfox (1995). Both stands were of fire origin, and were largely undisturbed by humans. The mature stand is consid- ered to be of rotation age and the old stand is much 530 2004. POHL, LANGOR, LANDRY, and SPENCE: ‘ ae ies - : e cat Edmonton waa ALBERTA 100 km boreal mixedwood Ficure |. Extent of the boreal mixed wood ecoregion in Alberta and location of the study area. older than the planned stand rotation age (60-70 years) for Alberta aspen forests. A comparison between these two stands, based on the light trap catches of Lepi- doptera described here, has been done elsewhere (Pohl et al. 2004). Two 30 watt UV traps were run in each stand, from dusk to dawn for one night approximately every two weeks, from 16 June to 16 September in 1993, 3 May TABLE |. Abundance and diversity of three groups of Lepi- doptera collected near Touchwood Lake, Alberta. Butterflies include the superfamilies Hesperioidea and Papilionoidea; macro-moths include the superfamilies Lasiocampoidea, Bombycoidea, Drepanoidea, Geometroidea, and Noctuoidea; micro-moths comprise all other superfamilies. Group Number of Specimens Number of Species (Proportion) (Proportion) micro-moths 3897 (0350) 201 (0.470) macro-moths 7163 (0.643) 229 (0.523) butterflies 51 (0.005) 8 (0.018) total TOE a GL 438 LEPIDOPTERA NEAR LAC LA BICHE to 5 October in 1994, and 28 May to 28 August in 1995. Traps were hung at approximately 1.6 m from the ground, and activated from dusk until dawn. These samples were augmented by periodic hand collecting at portable UV lights, and a small amount of net col- lecting of day-flying species. Specimens were identi- fied using a wide array of taxonomic publications, and by comparing to specimens in the Canadian Forest Service’s Northern Forestry Centre Research Collec- tion (NFRC) in Edmonton, Alberta, and the Canadian National Collection (CNC) in Ottawa, Ontario. Vouch- er specimens have been deposited at the NFRC and CNC. To obtain an estimate of the total size of the Lepi- doptera community in the study area, a Chao-1 estimate of diversity (Chao and Lee 1992) was calculated as described by Colwell and Coddington (1994). Results A total of 11 111 specimens were collected, repre- senting 41 families and 438 species (Table 1). A list of all species collected appears in Table 2. Some species, particularly micro-moths, are identified here merely as morphospecies, reflecting the lack of knowledge of the group, and the difficulty in making species identi- fications. Examples of some of the micro-moth species collected appear in Figure 2; some of the macro-moths are illustrated in Figures 3 and 4. The Chao-1 calcula- tion estimated the total size of the Lepidoptera com- munity in the study area to be 546 + 23.34. The 1994 data is examined in detail here, since it was the most extensively sampled year and there were no trap failures. In 1994, both abundance (Figure 5) and species richness (Figure 6) peaked dramatically in late July. A major flush of Noctuidae occurred in late July, with the highest species richness occurring throughout July and into early August. There were modest peaks of noctuid species that overwinter as adults, in early May and mid-September. The Geo- metridae and other macro-moths peaked slightly earlier than the Noctuidae, exhibiting maximum abundance in early July, and the greatest richness from mid-June to late July. The abundance of micro-moths peaked in late July, although most of this dramatic peak was a single species, Scoparia biplagialis, with 846 speci- mens. The greatest micro-moth richness occurred throughout July. More modest peaks in micro-moth abundance and richness occurred in mid-June and mid-September. Discussion This study likely did not collect all the species pres- ent in the study area; the Chao-1 estimator suggests that approximately 110 species were missed. Many of these are undoubtedly species that are not easily sampled via light traps. For example, the amount of effort spent collecting butterflies and day-flying moths was minimal and many of these taxa were likely missed. THE CANADIAN FIELD-NATURALIST Vol. 118 Da Ajng ayey — oung oye] I satsods piyjsalasay Ajng piu — sung pr I 2Sotoeds viysasdssy oun oye] — oun pri I Satoeds viyjsasKsay Ajng aye] — oun oye] (aouqn}) D]JaaDUISAd DIYISAIKSAY ysnony ae] — Apne pr Suda DIJASVIAO DIYISAAKSAY Ayne ayey — Apne pr (snoevuur’]) DIJAIADPAOS8 DIYISAIKBAY oun aye] — ounr pruu IO|[IZ, Dasnluod vIYISaIKssy oun oe] , Uva] SA1QD DIYISAAKBAY oune Ayrea — Avyy pr (yosng) DIJad[1IDAS. Saplojnauouodkyny oun oye] — oung pru (souqnH) DIJAISAVI VIUDPAIUUDMS oepnnowouod x Ayng prtu — oung pr I Soloads 1aJIAIOUOT]AY Ayn 3e] b (uneig) D]JAYADUL ADJINAOUOT[AY snsny ae] — oun Ape P uneig MAYID] SIA]JOIOYJJOJOA ysnsny Ie] € Soloeds Mau ,.Sdoo1a2019V,, oun oye] Z ({log 2 Aon) DIOINAIJSD SAOIMAIOIIY. 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VIOUAID] J Ajng 33e] | /Satoeds sryjouns.ivdg ysnsny Ajiea (SUSU9]-)) DUVIDINIVas Siyjouvssvdsy ysn3ny Ajiea (JOY[PAMA) saployjupx s1yjouvnssvds oun pru (snovuurT) DUDAJSIUMU DIN qsnsny Aiea — A[ng oye] ig (UOSUIqOY) puvjdwuosd sayjay oune 3je] (YIOMe}) puvu syayIo) Jaquiaydag piu (snto1ige4) DUDSADULA SIAAJIV oung Aiea — Jsnsny ae] (uosuIqoYy) DAUIJOABIU SIAAJIV poled IsIf4 Sd1ION JouNYy soroads snuay Ayre] ‘Apnys IWIK-IdIY} OY} IOAO IpRUL SUOT}DIT[OD UO pase SI POLlod IYSIP “ISIPJOOYS (E86) ‘[e 19 SOSPpOH IY} SMOT[OJ Bxe] JOYSIY UTYIM BdUANbas sa1dadg “(GHG ) UBSUAISTLY JO VY) SMO[[OJ UONROYISSe]O JOYSIP{ “eLIOq[Yy ‘aye poomyYonoL, Iau pojoa[joo saroeds vsaydopidey Jo isi] (panuyuod) *Z ATAV |, Vol. 118 THE CANADIAN FIELD-NATURALIST 536 Ayn ae] (sudua]a) DIJA8VAI3 NA YjYdlss3y Ajng oe] (13 [9Z) DIJAJOIANA DALY AUSY snsny Ayre p (udyourz) SN]JAYIVA] SNQUIDAD snsny oie] — Apne oye] (1jodo9s) snpjajsad snquivsg ysnsny Ajaewo — A[ng prut (aehq) SIDj4aqV viuopny ysnsny avy — Apne prut IOC sypisvidig viapdoos aepriquivig ysndny Ayes — sung aye] (19][97Z) DIJAIUOAPIAYIO VISOINT sung Ajiwa — Avy Ay.eo (douqnH ) DIJA1AVINSSO1s DIpoydoz ysnony Apres — sung aye] dolUNY 2 BNNINIAY Saplopjanauad DIAJICAOIG ysnony Aiwa — Avyy Oe] (jouosey) Dyjaoiajajqns sisdojakpy Ajng aye] G \S[NH DiJajniag ‘qoid ‘ds sispqgoioy Ayng pruu (1OYIPAA) SIpsnjaucy] DIUOYINOG oeprypeskg Avy Ape (UIeYsuUIs|eAA ) DIJajuagSa1d DIpUuOg oeprursodieg Avy Ape (WIBYSUIS|eAA ) void pyyddjquy Ayng pri (AD4[VAA) SnjAJavpouloy VisUul]|ay] aeploydosaid Av] prul Alpue’] 29 Arpuey louuryey Buon] y oepnionyy oun ae] — sung Ayre vd (puenig) Dyjajoundi.sadsp V1IyIOM depIpoly) Ajng ae] (1ouqnH) DuvIp S1jnasoyy oune pr (aekq) DJauaplgIgo SVALO]DI aepynai0yg Ayng pr (YOLIUIoH ) pnbopixayf vps) Ayn ajey — Avyy aie] (josng) puvjndod pips) Avy, oie] — Avy Ape WUBYSUISTeAA pupjoun] DyoydviH, Joquiaydag piw — jsnsny ary] (peut) puppul] pyouldy ysnsny ary — Apne aye] Z (Hoyeay) pupuowou pyouldy Ajng prtu — oun oye] (1OYIPAA) DUDSsusUudA] DYOUIAT Jaquiaydag piu (ojIvay) pubajppi4o pyouldy Jaquiajydag piu — Avy aye] (o19]D) pijasiu pyouldy Ayn avy — Avyy avy] (WIeYSUIS|eA ) puvoydijzaa pyouldy qsnsny Ajrea — Ayn pr (WIeYSsUIS|eM ) Duvaupjspo pyouldy Jaquiaydag piu — Ayn aye] (snavuurq) puplipuvjos pijouldy ysnsny avy — Ang pr (snovuur’y) pyjauosisy vyouldy Joquiaydag piu YOLulay] DUDIIPDA DPJASUH) Avy plu — Avy Ay.tea (WUeYSUIS|eAA ) DUOSALO DAaJUaXapNnas Ayng oe] []oMog pupunjs0fun DAIYADAAZ Jaquiaydag pri Z (Woy.way) puounjiof piaydv.i1az Ayn aie] URLUdd1 2 BAINNIN sisuappuvs v1aydvs1aZ oun ae] — oune piu YOLIULOH, pjoun[pp piuouosdéy A[ng oye] Z (SUdUa[D) pupjoo1ayps puouosdk&y Ajng aye] — oung pri YOU sluoynjysqns puouosdéH pollog 14314 So]ION Joyny saioads snuan Aprurey “Apnys 1edX-99.1Y) OY} QAO ape SUOTDAT[OI UO pase SI PoLiad WYSIPY “ISIP{IOYS (E€8H]) [B19 SOSPOH OY} SMOTI[OJ exe) JOYSIY UIYM IoUANbas satdadg “(GGG ) UISUIISIFY] JO IVY} SMOT[OJ UONLIYISsL]d JOYSIH “VUOG]Y ‘Oye poomyYoNoOL, Aeau payda][oo satoeds viaydopidayq jo si] (panunyuos) *Z ATAVL, (oe) ae) LEPIDOPTERA NEAR LAC LA BICHE POHL, LANGOR, LANDRY, and SPENCE 2004 Avy] Ay.rea (TDYTVM ) DIADSAN DIIKT Ayng prtu — sung pr (snovuur’y) DIADINJAG UOISI Isnsny Apia — oune ey] (aquany) DIADIJAIOd DIUUADOGOJOLd aun Apiea — Avy pr (ADT[NULIIFFIYIS wz stusq) piapnosndasd sidosjoq Ajng prt — Aeyy oye] (souany) DIADAAD] SISdOPIA] Av], prul (194[BMA ) DIXAJAIJU VDAN|DYJaV oung Apiva — Avy pr (TDATCAA) ppusoxa DIUOpyoyliCQ oun aye] (jayouodnq) piuvjsaddi DIUWPASIG Ayng pruu — sung Apiea (aouqnH) DIADUSIS DIADIDIN Ayng prtu — Aeyy pri ({jes1eog) DIDAIIS]N DIADIDW isnsny Apiea — oune aye] (1dYIVAA ) DIVIIDIIG VIADIDW ysnony Ayia — Ayne prw (UBLUSIOAq) DIADIIAO] DIADIDIN Ayng ayey — Apne pr (S1aquny,) DIVIUUNAG VIVID Ayn prtu — sung prut (as[ny) SIVUIGAIA AUDIO! — WUILUOUU — depLNOWIOIH Ajng prut — ounr 3e] (JOyTeAA) DASOA VIALCQ Ajng pri — Aevyy oye] (preyoed) pvauipig vuvdas1q ysnsny Apiva — Aevyy aye] JONTCAA vypno1v Duddaq oune Ayiwa — Avy] prul (aguany) suapnd paypaying Ang prut (a9uany) saplosoydoJDUkd DANDAYIJOPNAS J Ayng prtu — sung pr (asson) DIdIUIS AUKSOAGDH oeprurdaiqg ysnsny Ayiewo — oune aye] (Ainiq) SIUMAYIAD DIYIAD]ISDG snsny av] — oune ae] (qepoy) 1AIg] NU SIDISY aun aye] — 19q0190 Aye (snovuury) pdoyup sijpyduan Avy] prt — Aeyy Ay.ea (spreMpy) SNAKIDS DIUOSA]Og oepryeyduAnyy oune ae] ({BAnpsiog) DINJUAUD SaL1aagq ovpiuaroh'] aunf 31e] sLuey DaIVAsI]O SAI PL Aqng aye] — Avy oe] ueplor 2 P[LYOsyIOY sisuappuvo o1piddg ovprluorideg ounf oe] (sted) uowanppod snjpydados1ajADD oepruiodsay ysnsny Aiwa — A[ng pr (pyeusaj) ajpusisoquinjd Dwus1JSOjJsltoy) qsnony Aiea — oung ye] (AAV) SISA] DAP| Ajng Ajiva — Avyy oye] pieyorg Sypas1og DISNDAKG oune Ayia (9]01D) sypoiu visnvskg Ajng pruu — sung pru (jasvujny) DIDUOLOD DIUAVJIA] Yd oune ae] IO]]9Z, sypynoave vyspdsisad Aqne oe] (jaseujny) DIDpINd SISassaaq ounf prut (susua[D) sypnovu xudodvsvg ysnsny Ayiea — oun pr (TDyTeM ) sypsaqygo vyyouas Ajng aye] — Ayng pr (OJIeay) Dyjajoundissop visvipad polled Ws Joyny sotoads snuan Ayre] ‘Apnys IeaK-daIY} IY) JOAO IPL SUOTIDAT[OI UO pase SI PoLtad IYSI]T4 “ISIP{IOYS (€86]) ‘Te 19 SOSPOH oy} SMOT[OF BXe] JOYSIY UIYIIM Jouanbas satdadg (GGG) UdSUDISITY JO JY] SMOT[OF UONVOYISse[O JOYSIF{ ‘eMOQLY ‘axe poomyonoy Ieau payda]joo sorseds esajdoprdayqT Jo ysvq (panuyjuod) J ATAVL, THE CANADIAN FIELD-NATURALIST Vol. 118 538 Aq ae] = Avy Oe] OMS” DIIDALJad DUBWIOLIPA FT isnsny Ayre qS[0H NB1093 DIAKWA] Ajng piu — ounr Apvo (AO][NULIDJJIYIS wz stusq) pvaappis piadoydyoy ysnony Ajiwo — Ayne pre (as]nH) DUIJAX S1YJ1NGY ysnsny ae] — Ayn pr (JOP) pypupjdxa siyyjng ysnsny Apia — Apne pr (AOTB AA) pivsndoad siyjyjny ysnsny Ayes — oune prt (s[ny) DSOWAOL DULOAJSSKG ysnony Apiva — Avy aye] (aguony) DIVDIJISAIY VUOAISSKG ysnsny oe] — ounr oyv] ({[esae9d) DIVIIYIDM DUOASSKG Ajng piu (jadvujny) DIDIUNA] DULOASSKG. Joquiajdag piu — ysnsny Ayia (snovuury) DIDANI DUOAISSKG — DVUTUDIB'T — aepLauoayH ysnony Apiwo — oung pruu (A9[YOSOP) piuppisidf pjndorsy Ayng ajyey — oung oye] (JOP) piapjount pindoog Ayn piu (as[ny) pypjjaoup vindoos qsnony Ayteo — Ayn pr (YIoMeH) pippunoquyy vjndoasy Ayn aye — Avyy oe] (sguony) piupulnpuad psoydojrKy oune aq] (paeyord) pypuUuadopunjod Dapp] —-—- IWULY.ID}S — 9epLNawUo0ayH ysnsny Apes — Apne aye] (AdJJBYOS-YSLIOH ) DIADISISAL DAWVIOIVUAN qsnony Ayre — Apne prt (SLI) DIADNIDUL DKIIS Jaquiaydag pit — jsnsny aye] (s9uaND) DIAD]JAISY DUIPQuUIVT Ayn prtu — Avyy aye] (a9uany) DIADAOAIMIND Yulsag Ayng ayey — oung pr TOYIRAA vIvsiAlp VjaduUvy oung aye] — Avyy ov] (aguany) DIADJOOID SIPOSY] sung oye] — Avy pri (souony) DIADSOSO]Yyd SIPOSV] ABA OR] (snovuurq) pidpsaapnd siposv]d aun aye] — oune Apvo JOYICAA DIVUNUAAap DUAaUDIaW ysnsny Apiwo — Avy aye] douany DIADUOIDUI DUAUDIaW oune Apwo — Avy pr IOYIRA piapaydig]p piuajas Joquiajdag pri — ysnsny Avo souany DIADUSDU SOWOUUT ysnsny oye] — Apne pr (souany) vypjsad papduvy Ayn Ajrwo — oung prtu (JO4[CM ) DIADISAUOY OLA ysnsny Ayes — Ang pri (Ain.iq) pjadsos adxjoyjuvx ysnsny Apiea — oune avy] (aauany) DIADUIAS 1] DUAD]YIN| Av IR] (our) DIADUISADU DUAD]YINA Ayn piu (touqnH) DIADSNIGO DUAVDJYINT Ayn piu — oung pr sauany DIADJOIIDA DABQDI ysnsny Ajiea — oune pr aouany DIADUAYJAAA DIAQDI 19q0190 Aj.rea — Jaquiaydag prur (SLR H) DIADIPY SIUUDAT poliog ISIy SO]ON Joyny satoads snuan ATTUue,y “Apnys IdA-da.1Y} OY} IOAO pel SUOTIAT[OI UO pase ST polad IYSIPY “ISIPJIIY (€Q6]) ‘Te 39 SASPOH IY) SMOT[OJ BXL] JOYSIY UIIIM BoUONbas sarsadg ‘(GGG |) UASUAISLFY JO JY) SMOT[OF UONVOYTISsL]O JOYSIF{ “eOqLY ‘Oye poomyonoy, teau payoay[oo saroads vsaidopideyq jo asi] (panuyuor) ‘7 ATA], fag) va) POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 2004 Ayn Ayrea — Avyy prut (SLI H) DUDILIAWUD DUISapo]]|AYq oepiduresoise’'] isnsny Apes — oune pr pieyorg DIDAOUWD DIZZ1]]VIJ aeprUuRsip oune av] IOYICA pIdIIBIAIU DAOYdOGOT oune pri — Ary Apiea (1OMTVAA ) DIDANJYOAID DAVPYID ounr Ayiea — Avyy pr (1O¥TVAA ) DIADIUN] DADPVID Avy piu (preyord) DIDPIAIA SISDIV ounr Ayiea — Aevyy prul pieyorg DIVIVISOIOADA DIDAYNAN| oun ae] — sung pr JOYE piavoyup pigayndny ysnsny oye] — une pr (as[nH) DID Jas vioayndny Ayng prtu — ounr Ayiea (as[nH) pasnfsad pigaynldny ysnsny Apiva — Avyy oye] Aepolqnoqg ppjiuissy pigayydny Ajng ayey — Avypy 21] (touqnH) DIDAAIDS DIDAYIAN|A oun ae] — Aeyy oye] (YJIOMP}) pyposnfqns visayjidny Av P| (aehq) pipiquinjoo pivayndny ysnony Aiea — oung aye] (JD4TPAA) poipuau aydpqny 19q0}90Q Ape (as[nH) pypaonag vsajydosad¢Q Jaquiajdag piu (Uasneyyog) pipuumnynd vylssdy Ajng prt — Aryl 312] (s9uaND) DIDIPAOG]D DIZAPOYIIAT, Ayng pri — Avyy pr (aeAq) 1jpsavad pisnua, ysnsny avy — Avy oye] sing DIIAQUIDI DISNUAA ouny 9}e] (1O4TRM) DAiafiqyy DYAaIpAH isnsny ae] — Ang pru OulsseZ pypuidjp sdajydouaz Aqng aye] — Avy pr (aguond)) vypipawaaqur pidydny Ajng oye] — ung pr (JOT [NJA) vpUsayp aoyssdy ysndny Ajiwa — Avyy pru (aguany) DIDAJSNIVD] OYLOYJUDX Ajng piu — Avy oye] (PID) DIDSNALA, POYAOYJUDX isnsny Ayiea — Ang piu (aadsq) DIADAO]OIAP BOYAOYJUDX isnsny Aiea — oung AT Ivo IO[ARL, DIADSSOL JOYAOYIUDX Ayng ae] (gouaNy) DIVNp1 aOYsOYJUDX ysnany Aiea — sung aye] (1dJJVYOS-YSLLIOH ) DIADSDAQD AOYAOYJUDX ARIA AL] (TOTEM) Dpsafyjnul vajayUy oune Ay.rea — Avy pr souony DIVIPISDA DAJIYUY Ajng aye — Aqne Ayaea (JOC ) DIVIDSDG YULOZ1A Aq Ajng prut — sung pru (AD][NULIAJJIYIS 2 studsq) DIpNjan] viuvs.ivds Ajng piu — Avy Apivo (suony) DIDIPLIYNA DINA|OSAW Jaqo1oQ Aira — Jaquiaydag piu (aguony) pypyisapy vsoydisy, ysnsny ae] — Apne aye] (Siaquny,) DIDIAN{ DUAULOLIPAH] Ajng ayey — Avyy oe] (194014) DIpAAGNA DUAUWIOLI PAL] Ajng aye] — Avyy oe] (ADT ) pIpIOUNUAL DUAWOLIPAL pollog SI SO}ON JouNy satoads snuan Ayre] ‘ApNys IdK-9dAY} DY} JOAO IPLUT SUOTIITTOS UO pase st PoLtad IYSIpy “ISIP{IOYS (ERG) ‘Te 19 SOSPpOH oy) SMOT[OF BXe] JOYSsIY UTyIIM doUENbas satdadg (666) UASUIISITY JO IVY) SMOT[OJ UOTVOYISse]O OSI “ViOq[Yy ‘Oye poomyonoy, 1wau pajoa][09 sardeds vsgydopidsy jo jsv] (panuyuod) *7 AAV, Vol. 118 THE CANADIAN FIELD-NATURALIST 540 Ayn oyey — Apne pra aauany sypnbuidosd pinay OVUI[NALY — oepINIoON ysn3ny Apiewo — Apne pro (UIUS “g'f) pauliyovisf sapouadiy seurueutsdans — aepmnjoony Ajng piu — sung oye] (aauqnH) SYDINBUD SIYI]Vq Ajng prut dauany SIPULIpvADI Duldalg Ajng a1] vd (UIs “g'f) NUDYUDY VjOISOUay|PUd ysnsny Aiea — Avy] oe] 01D sypasjad vINOIKYD ysnsny Ayivo — Apne pr (yt “g‘f) pq]vIN] VYyIwUsO]IUDZ Ayn prt — Avy oye] (IDTV) sypsnupskd punydouanjpyd ysn3ny Ajiwo — A[ng prut ¢€ Jouqny pinwav eau saisads Mau VIP] ysnsny Ayia — Ayn pr Jouqny pinuav VIP] ysnsny Apieo — suns oye] (aguany) SIJDIIAAWUD DIP] BUNULULIOF — OepInjoOoN ysnony avy — Avy pr (TDAP) puppisidf DjoajotN — seuldt1yjo.LIeg — aepinjooN Ayn ayy — Apne pra (TDYIPMA ) pypispyd palyIksvg Ajng oyey — Apne pri (ysnouunqgdsyy 2p soureg) SUDBDA DAIYIASDG depILUeULAT oun aye] — oun piu SLLIeH ppjnoput vduma20ydoT Ayng piu — oune pru (SLLIeH) souayjipd DYIAVID] oune a}e] — Avy ae] JORMA SUD]IUNSSD DIGOJDUSDAY qsnsny Ayiea — Ayn piu (O[[IAQUSAJ-ULIOND ) 1ajuoza] vojdpy ysnsny Apivo — Ang prur pieyord DID|G]Y vIsuauia]D ysn3ny Ayres — Ang pr (2}01D) 4OJONIG DUA] oeplyno1y ysnsny Ajiewa — une aye] (01) saplouyda] vanziyos Ajng Ay.ea (yyUIS “A‘f) STUAODIUN DANZIYIS oune ae] (uospny) DISAPOU DINIANA oune ae] — Aeyy ae] (eAnpsiog) pUulApuadojoos VINIANA oun aye] — Avy oye] (touquUr'T) SIDJUAPIIIO VINIAN Avy Ayivo (01D) laujuy pisiydnjpH Avy, RY] — Avyy pre uospny pynopuan visiydn]H ysnsny Ajava — Aeyy ey] IONE stuoluajdas visiydn]y oun Ajrea — Avyy oye] jovin pip) dus DJUOPOJON Aqne piu — Avy prur pieyorg DSOUL DISOAY oune pruu (YJIWIS “A'f) DSOgq1s DIDPPN oung aye] — Ary oie] (IOyTRM) sypo1dv p1ajsoj]D sung aye] — Avy oye] (01D) DSOBIAJS DAAJSO]D Ajng avy — Avy ayy you DUSISOG]D DAAISO]D oepnuUopo}ON oun ae] (Sinquia}0y) mp8 Saja isnsny Aiwa — Avy pr AQAry IAS1499 SNYJULLAWUS oeprsuryds ysnsny Apiewo — Apne pra JouqnH DIAISSIP DULOSOIDIVDIN pollag IWSIpy SION Joyny satoads snuan ATTUre.y ‘ApS IRIA-J9IY} IY) IOAO OPBU SUOTIAT[OS UO paseq st PoLtad IYSIPY “ISTPIIYO (€QG]) “[e 9 SOSPOH BY) SMOTIOY Bxe] JOYSTY UTYTM dUINbes sa1sadg *(6GGH]) UBSUDISITY, JO JY SMOT[OJ UOTBOYTISsKIO JOYSIF “VUOQTY ‘oye poomyonoy, teu payoayfoo saroads viajydopidayq jo isv] (panuiuod) ‘7 ATAV], 54] IRA NEAR LAC LA BICHE NCE: LEPIDOPTI 4 POHL, LANGOR, LANDRY, and SPE 2004 ISNSNY av] (aakads) pupoiiauy paodiyduy Isnsny ayey — Ayn pr (aouqn}) DIIAUSNS SUYINSDADE Jaquiajidag prw — jsnsny ar] (JOYIVAA) VIDIO}! DISCO Jaquiaydag piu — ysnsny aye] (sauany) DIVIIDU DISICQ snsny Ayiea — Ayng ae] (ujTwIS) pID]IBOI Daubdy Ayng aqey — Avyy Ayiea (JOMTeAA) ppowuos vaupdy avuluapeH — oepinjooN jsnsny ae] — Apne ey] (JOMPA\) pxiful Duapoyouoryy snsny ayy — Apne pra (9}019) DSLAJSIPDG DUaPYYOUlo yy OVUIT][NOND — oepInjoOoN ounr piu (TOYTR A) DIDUSISIA] DUUAULISIAAD FY] aune piu — Aryl aye] JORMA DSSatdull DIJUOLIY oun ae] — sung Area JOYE Djajdul vjgU0LIy snsny Aiea — APY] Ae] (aguany) SIS DAL DIDIUOLIY Ajng piu — Aevyy aye] JOYE VASUs DIMUOLIY snsny Ajiea — oun pr aguony DIDIJOUUL DIIIUOLIY aune ae] — dune Aye douony DUldna DIIUOLIY Ayng aqyey — Aryl ore] OID Jawdf viydvy —- aeua1uosdVy — ovpIndd0N Ajng aie] — oun prur (a99uaND) DINpIG]Y VIPOIVYINT dvUIO.NSNY — sepinjooN ysn3ny Apia — une ae] douonh nupujnd pisn}d Ajng pr (AqIry) vinsuvpjoad DyddAsUuKsS, Aqng aye] — Apne pr (Insusjongd) spyp pydvssuasy ysnsny avy — Apne aye] (Q]01D) DUB ISIPIA1A DYADABUKS A[ng ae] (9019) pidiagsoja0 pydvssuasy Ayn ayey — oung aie] (1DYTRAA) pidup pydvisoiny Ajng prt — oune oye] (UOSUTGOY 2 901) pddpu vydpisojny ysnony ary — Ang aye] (suoydays) pypnovuig vydpssoiny oun aye] — oun pru Insud;OnNO ppigqna pydpasojny Ayn ae] (sound) saproskyoay) XKsajdosoydsogq Ayn ae] (Q101D) psowsof pyduicuvsKy) ysnsny Ajiea (anyioqQ) DP]alauUsa DISKAYIA] Og Ajng ae] (9101D) saploasav DISKAYIVDIG. Ayn prt — oun 39e] aouonty S1JUAIN DIOISOAGY avuUlIsN[d — sepInjooN snsny ae] — snsny Ajies 01D DIIYAAMUIS DIVIOIDD Jaquiajdag piu — jsnsny Ayiea spiempy SIASIAG DIDIOIDD Jaqo1oQ Apiea — Apne aye] JOC pentiun pjps0jvyD Jaquiajdag prut —ysnsny Ayieo JOC DJIAL DIDIOIDD ARYA] IE] (WJIOMeH) DINISNISSDAI DULIBANUID IVUT[RIOJVD — OepINIION Avy pruu STeY ynuny Duaday isnsny aye] — Ang pra (Aye ) sypjoipa puaddy Ajng prur quis piupwojyp puadA yy avuruadAY — oepinjooN polled ISt14 loyny soroads snuan Ayre ‘ApNys IdK-IdAY} OY} JQAO IPRUT SUOTIDOT[OD UO pase SI PoLtod IYSIL4 “ISIPYOOYS (€Q6]) [P19 SASPOH IY} SMOTIOJ BxP} JOYSIY UIIIIM ddUaNbas satsadg “(GHG [) USSUIISTNY JO JLY) SMOT[OJ UONVOYISSeID JOYSI “vIIIQTY ‘aYe'] poomyonoy, Iwou po}da]JOo sarseds esoydopidaT Jo jst] (panuuyuod) *7 ATAVI, Vol. 118 THE CANADIAN FIELD-NATURALIST 542 Ayng oye] — Apne prow ysnsny Ajiva — Apne pr ysnsny yey — Ayn pri oun Aj.ro oun Ay.eo — Avy plu oune Ayrea — Avyy Apieo Avy pr oun Ayre — Avy pra oune ae] Ayne ayey — Apne prot ysn3ny Ajiea — sung aye] Ayng pra Ajng ae] — oune aye] ABI, OP] oung pri Ayn oyey — Apne pra ysnsny Ayjieo — oune piu snsny oe] — Isnsny Aj.ea ysnsny aye] — Isnsny Ayia Jaquiaydag piu — jsnsny oye] Jaquiaydag piu snsny ae] Jaquiajdag piu Jaquiajdag prt — ysnsny Aya oung Aye oung Ape oung Ajiva — Jaquiajdag piu 19q0}9Q Aj.iea Jaquiaydag pr — \sndny ae] Avy] plu — Joquiajdag pri oun ayey — Avy oe] Ayn pr — oung prut snsny Ie] Ajng piu — Avyy oye] ysnsny oe] — snsny Ay.ea Jaquiaydag piu — A[ng prur Joquiajdag piu — Ajng oye] isnsny Ajaiva — Ayn pr oune aye] — oun Avo polled 1Yyst[4 SOON (91019) (yMUIS) (Q]01D) (aguany) (21019) (aguany) (YIU ) (UOSTIIOJ]) sguony (Q]01D) (s9uony) (suaydays) (uruuefuag 2 sou) (1O4TBA\ ) (aguand) (e9uan5) (s9uony) (194[2M\) (01D) urueluog 2 souieg (suoydays) Qpeis.i0197) PIOPTNA 2 surejuoje’T (UOSUIGOY 2 9101) 01D (y}IUIS) (UOSLLIO[\) UOSLLIO[ (UOSLLIOJ[) (UOSTLIOJ]) (UBATT[NS 2 dUBevIq) ysnouunqojy (TOTEM ) (soueny)) 01D (9]01D) (1OX[P AA) souony ysnouunqopy louiny DIADSOL DISADIG] DIDIOISIP VISADIG. DAafiqns DISADIG, DINPIAO SAPOYJAOJOL DSO]OP DAIS 1ISIQ1IY DISOYJAQ DIDSALSAS DISOYJAC DIJIAAL DISOYJAC) DJaNSUL DIUDINIT DIDEAxO DULUIYIA PY Da1o] VIOdUIIVT DAIBIUAI DIOGIUIIDT p20d DIKUuoIsVT XIPDA DIGOUDIDT pjouni[py pAyIUD]aW DAALIAQuAL D1]Og DSOQuIIU DILOg DABIUIISIP DIWOJAYIDAG SUAB]D DIMOJAYIDG pippo4sv adXjojAXx sdaoup pyodajv] d SEUSOUILEL O8DID] DIYJUDX pind xiyjpuy panjad aupydoynT pypunuouul aupydoyT pavdou piuojoyny DUIIAIY VAD]SOWOH DUDUAIS DIOWOYINT DINIDUIAINI DUAIAX ppyjvdnyp visydvyq pysp.ijuos vddky plopuod DyOdospuy pynaiapyjpd X1UojAyD psnjaauoajd pydiouidy DIDUN{UI DISADUT 40] 0IJAP VIBADUT psojnoisad psoydosopyd sipunsauag vixajdny soroads snuayy avUINIION — sepInooN Ayre. ‘Apnys 1edX-Ja.1Y} OY} JOAO ape SUOTIDITIOO UO paseq SI poLtad IYSIT4 ISIP{OoYS (C86) ‘Tk 19 S9SPOH 9} SMOTOJ exe} JOYsIY UTyITM ddUANbas sar9adg “(666 ) UASUAISLNY JO LY] SMOT[OJ UONROYTISsL]O JOYSIF{ “eOqTYy ‘Oye'T poomyonoy, wau payda]joo saroads vsajdopiday jo ist] (panuyuod) *Z ATAV L$, 543 AR LAC LA BICHE 4 IRA NE PIDOPTI POHL, LANGOR, LANDRY, and SPENCE: LE 2004 VLOQ[Y IO} P1ODOI MOU ,, sotoods paqiiosapun . urIe}IIOUN UONRUTUIAJOp Sotoeds ~ UONPUILULIAJOp Satdads & DYRUT OF JUSTOYJNSUL ST ISPI|MOUY SIWOUOXL} JO 9}LIS | :S$ojou ———————————————————————————————————————————————————————————————————————————————————————— ———————0—0—08—0—0—8; 0NSssSsSs0—SsaoaSn”suaoOaSS\>OoOwD-Smomaoamoamom—mm,.! Aqng ayey — Ayu prur isnsny Apiva — oune aye] isnsny aye] — Apne aye] snsny Aiea — Avy Aprea ysnsny Ayiea — A[ne pr Aqng ayey — Avy 1e] Avy prul ysnsny Aiea A[ne ae] snsny Aiea Apne are] isnsny aye] — Apne aye] Ayn are] Ajng pr ysnsny Ajiva — Ayng prur isnsny aye] — Apne prw ysnsny ae — Avyy Ape pouled 1yst[4 SION (ounyleg) (21019) (UOSTIIOJ]) (21019) (TDT[NULIDFFIYIS 2 stusq) (sguony) (TONTEAA) (9}01D) (UOSTLIOJ{) (touqnH) (TOTEM) (uaqjaus) (M019) (snovuurT) (UOSLLIOJ]) (snovuul’y) (sniotiqe,y) JoyINYy sisuaippov DIpI0jJdKD puaypsad Sols DJa,any snjgadyna DAduipjojotd snssaid saplojgajdpuy puispad sapiojaaj}dpuy DjIpuod Saplojoajdy wWNADIYYS sisdapvlaW suodfiondo pjiydouao)y DINIINUA] VSSDUOULLIYOPNAS J pylsadul viqouyovd DIXIUL DIGOUYID MYJMUS DIISAX DUDIUDUULAOU DYSIX WINASIU-I DWUASDSAW DIIIAISD SIOANG DIJNIIO SIOANY unsnp pioydiydvsy soroads snuay Ayre eS 0SSOOOaoawamnaoOorrn ao ‘Kpmys rwak-aary} dy} OAO IpeUL SUOI]DIT[OS UO paseg St POLIod IYSI[Y ISIPPIYO (E861) ‘Te 19 SOSPOH 9} SMOTIOJ BXE] JOYSIY UTYIIM douanbas saradg (666) UASUAISLDY JO WU) SMOT[OJ UONROYIssL]O JOYSIH “BogTy ‘aye] PoomYoNOL eau pa}oa][OI soroods viaydopideq jo isi] (papnjouor) *Z AAV, 544 THE CANADIAN FIELD-NATURALIST Vol. 118 Gelechia lynceella FicureE 2. Microlepidoptera collected near Touchwood Lake, Alberta. All are new Alberta records. (ruler = lcm). The study site lies within the ranges of 67 butterfly species (Bird et al. 1995); many of these certainly occur in the mixedwood habitat but were not collected there. Some nocturnal moths, including some species of Gelechiidae and Oecophoridae (Hodges 1974; Miller 2000), are not attracted to UV light so they would be undersampled as well. If other moth species occur in the area but were not collected, they are probably quite rare, or may be extremely localized in particular micro- habitats, which were not adequately sampled in this study. Macro-moths comprised a greater proportion of individuals than of species, whereas the micro-moths were particularly species-rich, comprising a greater proportion of species than of individuals. These pro- portions (Table 1) were similar to those previously reported for the province of Alberta. In his list of Lepidoptera of Alberta, Bowman (1951) reported 40.8% micro-moths (779 species), 50.9% macro-moths (973 species), and 8.3% butterflies (159 species). In the most recent list of Lepidoptera of North America, Poole (1996) listed 49.3% micro-moths (5743 species), 43.9% macro-moths (5114 species), and 6.9% butterflies (801 species). The proportionally higher representation of micro-moth species in the current study compared to Bowman’s 1951 list reflects the recent increase in knowledge of the group, including the description of many new species. A total of 35 positively identified species are new records for the province, and one (Acanthopteroctetes bimaculata) is a new record for Canada (Table 2). Micro-moths (Figure 2) make up 34 of these new records; a further five micro-moth species and one macro-moth species represent undescribed species. Details for some of these new records and new species appear below. Acanthopteroctetes bimaculata Davis (Acanthopteroctetidae): This primitive moth is known previously from California and Oregon (Davis 1978). The current record represents the first report of this moth, and of the family Acanthopte- roctetidae, occurring in Canada. Nothing is known of its biology. Caloptilia anthobaphes (Meyrick) (Gracillariidae): Previously reported from northern Ontario (Forbes 1923). Nothing is known of its biology. Caloptilia betulivora McDunnough (Gracillariidae) (Figure 2): Previously reported first only from Nova Scotia (McDun- nough 1946) and since been reported from Quebec (Hand- field et al. 1997). It likely occurs across the boreal zone in Canada. Larvae feed in the folded leaves of birch. 2004 POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE Chytonix palliatricula FiGurE 3. An assortment of macrolepidoptera collected from mature aspen forest near Touchwood Lake, Alberta (ruler = 1 cm). Caloptilia canadensisella (McDunnough) (Gracillariidae) (Figure 2): Originally described from Nova Scotia (McDunnough 1956). It has since been reported in Quebec (Handfield et al. 1997; Landry and Landry 1992), and is probably widely dis- tributed but uncollected across the boreal forest. The larvae make large blotch mines on the leaves of Bunchberry (Cor- nus canadensis). Caloptilia coroniella (Clemens) (Gracillariidae): Previously known only from the midwestern U.S.A. (Forbes 1923). Larvae feed on birch. Caloptilia stigmatella (Fabricius) (Gracillariidae): Not reported from western Canada, but GRP [Greg R. Pohl] has collected it quite commonly in Alberta and Saska- tchewan. Larvae feed on willow (Forbes 1923). Argyresthia abies Freeman (Yponomeutidae): Reported in eastern Canada only as far west as northern Ontario (Freeman 1972) but is probably widely distributed across western North America in the boreal forest. It is a twig borer on Balsam Fir (Abies balsamea). Coleophora corylifoliella Clemens (Coleophoridae): Previously reported only in eastern North America (Forbes 1923). Larvae there feed on Corylus americana; in the west they probably feed on Beaked Hazelnut (Corylus cornuta). Coleophora duplicis Braun (Coleophoridae): Previously known from eastern Canada and the midwestern United States (Forbes 1923: Handfield et al. 1997). It feeds on the seeds of Aster (Aster spp.) and Goldenrod (Solidago spp.). Limnaecia phragmitella Stainton (Cosmopterigidae) (Fig- ure 2): A holarctic species. The closest it has been reported to Alberta is Wyoming (Hodges 1978). However, collection records (CNC, NFRC) indicate that the species is transamer- ican. Recent collections made by GRP indicate that it is com- mon in western Canada, at sites where its host plant, Com- 546 THE CANADIAN FIELD-NATURALIST Vol. 118 cronicta innotata Ficure 4. An assortment of macrolepidoptera collected from old growth aspen forest near Touchwood Lake, Alberta (ruler —yluemi)® mon Caitail (Typha latifolia), is available. Larvae feed on the flowers and seeds. Coleotechnites blastivora (McLeod) (Gelechiidae): Reported only from the type locality of Gaspe, Quebec (McLeod 1962). It probably occurs across the boreal zone. 2000 Noctuidae HB Geometridae other macro-moths {) micro-moths no. specimens _ oa o o June July Aug. Sept. trapping date FiGure 5. Abundance of major Lepidoptera groups collect- ed in UV traps through the 1994 trapping season. Larvae are needle webbers of White Spruce, occasionally mining within the needles. Gnorimoschema septentrionella Fyles (Gelechiidae): Miller (2000) reports this species only in eastern North America, as far west as Minnesota. It is a stem-gall maker on asters (Aster spp.). 160 ©) Noctuidae GN Geometridae __| other macro-moths {) micro-moths 120 no. species co oO sy Oo June July trapping date Aug. Sept. FiGure 6. Species richness of major Lepidoptera groups collected in UV traps through the 1994 trapping season. | 2004 Aethes promptana (Robinson) (Tortricidae — Tortricinae): The Touchwood Lake record represents the first record of this species from western Canada. It was previously known from eastern North America as far west as Wisconsin (Sabourin et al. 2002). A prevoius record from Washington (Razowski 1997) was based on a misidentification. Nothing is known of it’s feeding habits. Wockia asperipunctella (Bruand) (Urodidae): As noted by Landry (1998), this Touchwood Lake record represents the first report of this species, and the family Uro- didae, in Alberta. This family is a recently recognised addition to the North American fauna (Heppner 1997). Larvae feed on Trembling Aspen; adults tend to fly during the day or early evening. Alucita lalannei Landry & Landry (Alucitidae): Until recently, this species had been combined with two other species in the genus, and referred to collectively as Alu- cita hexadactyla (Linnaeus) (Bowman 1951; Hodges et al. 1983; Poole 1996). However, the true A. hexadactyla 1s restrict- ed to the Old World. In their description and treatment of A. lalannei, Landry and Landry (2004) designate a specimen from the current study as a paratype. This species is known from Ontario, Manitoba, and Alberta. Adults overwinter, and can be collected from May to September. The larval host plant is unknown, but Landry and Landry (2004) report Lonicera spp. and Symphoricarpos spp. as likely candidates. Parapoynx maculalis (Clemens) (Crambidae): Previously known from eastern North America, only as far west as Lake of the Woods, Ontario (Munroe 1972). How- ever, recent collecting by GRP indicates that it is widespread but extremely localised in Alberta and Saskatchewan. Its larvae are aquatic, and are reported by Munroe (1972) to feed on several species of water lilies (Nuphar, Nymphaea, Brasenia spp.). It likely feeds on other plants as well, since it has been collected recently in southern Alberta, outside the distribution of the aforementioned plants (C. D. Bird, personal commu- nication). Idia new species near aemula Hiibner (Noctuidae — Her- miniinae): Usually mixed in with /dia aemula Hibner in collections. It was reported in eastern North America by Rings et al. (1992), and misidentified as Epizeuxis concisa Walker by Forbes (1954). It remains undescribed and has not been reported previously from Alberta, although it is common across the boreal region. It has been reported to feed on the needles of a variety of conifers (Rings et al. 1992). Phalaenostola hanhami (J. B. Smith) (Noctuidae — Her- miniinae): Reported from eastern Saskatchewan as far west as Regina (Hooper 1988). Nothing is known of its feeding habits. The only similar published study of boreal forest Lepidoptera that the authors are aware of is that of Morneau (2002), which sampled primarily macro-moths near Peace River in northwestern Alberta. In that study, 293 species were collected over three seasons, including 278 species of macro-moths. Although most macro-moth species were common to both studies, 43 species were unique to the present study, and 115 species were unique to the Morneau study. The pres- ence of so many unique species suggests some habitat POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 547 specialization within the boreal mixedwood region. Several of the species unique to the Morneau study, most notably some Arctiidae species, represent an incursion of cordilleran species into that area. Several Lepidoptera inventories of provincial parks in the boreal mixedwood forest of northeastern Alberta have been carried out (Schmidt and Pohl 2001*; Macaulay and Pohl 2002*, 2003*). Although these studies collected from 138 to 295 species, they were each based on brief sampling periods, so they missed significant proportions of the fauna. Catches from the current study shared approximately 50 to 60% of the species collected in those studies. In general, the moth fauna of the boreal forest appears to be less diverse than that found in other forests. Summerville and Crist (2002) collected 512 species of moths in deciduous forests in Ohio. Thomas et al. (1998) collected 624 species of macro-moths in Fundy National Park, New Brunswick, which lies in the Aca- dian Forest Region, as described by Rowe (1972). Both of these forest types are more diverse botanical- ly than the boreal forest, and have more moderate cli- matic conditions (Rowe 1972). Inventories of this nature are a necessary prerequi- site to understand impacts of forestry practices and climate change on biodiversity. However, there is a paucity of similar studies, largely due to the lack of required taxonomic expertise. This argues strongly for increased support of systematics research in Canada, as biodiversity issues continue to increase in importance. Acknowledgments Héctor Carcamo, Tom Clark, James Hammond, Rob Lucas, Phillippa Rodriguez, Peter Shipley, the late Tim Spanton, Kevin Sytsma, and Daryl Williams assisted with field collecting. The Alberta Land and Forest Service (Lac La Biche Region) provided field acco- modation, and the Alberta Research Council provided maps and assisted with site selection. Gary Anweiler, Klause Bolte, P.-T. Dang, Ron Hodges, Don Lafontaine, Doug Macaulay, and Jim Troubridge assisted with identifications. This work was funded by the Canada- Alberta Partnership Agreement in Forestry and by the National Science and Engineering Research Council of Canada (operating grant to JRS). Documents Cited (marked * in text) Macaulay, A. D., and G. R. Pohl. 2002. Survey of Lepi- doptera in the Canadian Shield Ecoregion of northeastern Alberta. II. 2001 Survey of La Butte Creek and Fidler/Greywillow Wildland Provincial Parks. Report to the Alberta Natural Resources Service and Alberta Nat- ural Heritage Information Centre, Parks and Protected Areas Division, Alberta Community Development. 61 pages. Macaulay, A. D., and G. R. Pohl. 2003. Survey of Lepi- doptera in the Canadian Shield Ecoregion of northeastern Alberta. III. 2002 Survey of Colin-Cornwall Lakes Wild- land Park. Report to the Alberta Natural Resources Ser- vice and Alberta Natural Heritage Information Centre, 548 Parks and Protected Areas Division, Alberta Community Development. 44 pages. Schmidt, B. C., and G. R. Pohl. 2001. Survey of the butter- flies and moths (Lepidoptera) of the Canadian Shield natural region of Alberta. 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B., Editor 1995. Relationships between stand age, stand structure, and biodiversity in aspen mixed- POHL, LANGOR, LANDRY, and SPENCE: LEPIDOPTERA NEAR LAC LA BICHE 549 wood forests in Alberta. Jointly published by Alberta Environmental Centre (AECV95-R1), Vegreville, Alberta, and Canadian Forest Service (Project Number 0001 A), Edmonton, Alberta. 308 pages. Summerville, K. S., and T. O. Crist. 2002. Effects of tim- ber harvest on forest Lepidoptera: community, guild, and species responses. Ecological Applications 12: 820- 835. Thomas, A. W., J. Edsall, and D. Clay. 1998. Biodiversity survey of moths of Fundy National Park: I. A checklist of the macromoths of Fundy National Park, and its greater ecosystem. Parks Canada Technical Reports in Ecosys- tem Science, Report 017. 33 pages. Received 20 October 2003 Accepted 27 September 2004 Pilose Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an Enigmatic Endemic of Arctic Canada JAMES G. HARRIS Biology Department, Utah Valley State College, 800 West University Parkway, Orem, Utah 84058 USA; e-mail: harrisji@ uvsc.edu Harris, James G. 2004. Pilose Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an enigmatic endemic of Arctic Canada. Canadian Field-Naturalist 118(4): 550-557. Braya pilosa Hooker, Pilose Braya, has been poorly understood among North American botanists due to a paucity of fruiting specimens for study. This has resulted in confusion about the taxonomic position of the taxon within Braya, and has led to speculation about its generic status. An examination of fruiting specimens from the Royal Botanic Garden Herbarium at Kew, England reveals that B. pilosa is correctly placed in the genus Braya, and that it is a distinctive member of the genus deserving recognition at the specific level. I discuss the historical evidence that B. pilosa may not have been collected since 1850 due to its extremely restricted distribution on the Cape Bathurst Peninsula of the Northwest Territories of Canada. I also present evidence suggesting that B. pilosa is diploid and may be a parent species to some of the more widespread members of the genus, all of which are polyploid. Its closest living relative is probably B. thorild-wulffii. Key Words: Braya pilosa, Pilose Braya, discovery, rediscovery, distribution, collections, rare species, Cape Bathurst, North- west Territories, Canada. Braya pilosa Hooker (Figure 1) is an extremely rare endemic of the Northwest Territories of Canada. It is perhaps the most distinctive and, at the same time, least understood North American member of a taxonomi- cally difficult genus. While some botanists (Harris 1985; Argus and Pryer 1990; Rollins 1993; McJannet et al. 1995) have treated B. pilosa as it was envisioned and described by Hooker (1830), most have placed it into synonymy under B. glabella (Polunin 1959; Welsh 1974; Porsild and Cody 1980; Cody 2000; Warwick et al. 2000*) or B. purpurascens (Scoggan 1978). Oth- ers have reduced it to infraspecific status within B. purpurascens (Schultz 1924; Hultén 1970), applied the name broadly to elements of B. glabella (Porsild 1943, Hultén 1968), or ignored it entirely (Cody 1979; Mulligan 2002). To the best of my knowledge, there are no authentic fruiting specimens of B. pilosa in any North American herbarium. It is this lack of available fruiting material for study that is primarily responsible for the taxonomic confusion that has surrounded this taxon over the past 173 years. When he described the species, Hooker (1830) had only a single collection of flowering individuals with immature silicles from which to work and he was ap- parently uncertain whether or not the plant was actu- ally a Braya. He placed a question mark behind the genus name, lamented the lack of mature fruits that would allow unequivocal generic placement, and stat- ed, “...in all probability, it ought to constitute a new genus.” Without access to mature fruits for examination, Hooker’s suggestion of a new genus would indeed seem justified. The flowers are larger than those in any other Braya species (the petals in some tetraploid B. humilis are as long but not as broad); the immature silicles are ovoid-ellipsoid, unlike those in any other Braya with the exception of B. thorild-wulffii (which had not been discovered in 1830); and the styles are exceptionally long for a Braya, measuring over half the length of the ovary. In addition, Sir John Richardson (1830), who collected the type specimen in 1826 during the second Franklin expedition in search of a north- west passage, described the flowers as fragrant, with a smell similar to lilac blossoms. Fragrance has not been associated with any other member of the genus. Murray (1983*) and Scott et al. (2000*) suggested that B. pilosa is a misnomer applied in error by Hook- er (1830) to a Draba, likely D. corymbosa. However, Hooker’s original description and accompanying illus- tration of the taxon indicate that he is describing some- thing quite different from D. corymbosa. Hooker describes the valves of the immature silicles as “valde convexis,” and again later as “remarkably convex,” descriptions that do not accurately describe the dis- tinctly flattened valves of D. corymbosa. Fortunately, any doubt about the identity of Hook- er’s B. pilosa as a legitimate Braya rather than as a misidentified D. corymbosa 1s resolved by an exami- nation of authentic fruiting specimens. There is a herb- arium sheet at the Royal Botanic Gardens Herbarium (K) in Kew, England (Figure 2) that apparently in- cludes specimens from three separate collections (Sea Coast, Arctic, Richardson s.n., 1848; arctic coast, W. of C. Bathurst, Captn. Pullen s.n., Aug. 1850; and what are likely part of Richardson’s type collection of 1826). The herbarium sheet is so congested that it is impos- sible to be certain which individuals are part of the various collections represented, but all of them are defi- nitely members of the same taxon. Mature silicles on 550 2004 HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA Ficure 1: Illustration of Braya pilosa. 55] Bp these specimens, even though pressed, are clearly ovoid-ellipsoid and perfectly match Hooker’s illus- trations of immature fruits of B. pilosa. In addition to possessing fruits that are strikingly different from those of D. corymbosa, the Kew B. pilosa specimens differ significantly from D. corymbosa in several other attrib- utes. The leaves are much longer, narrower, thicker, more obtuse, and less densely pubescent than those in D. corymbosa; some individuals bear a single cauline leaf while D. corymbosa lacks cauline leaves; the styles are much longer than those of D. corymbosa; and most importantly, the epidermal cells of the silicle septum are typical Braya epidermal cells, characteristically thickened and transversely or obliquely elongated (Harris 1985, Figure 2), in stark contrast to the thin- walled, irregularly shaped, and essentially isodiametric septum epidermal cells typical of Draba. Braya pilosa is unequivocally not a misidentified Draba. I have not seen any authentic specimens of B. pilosa that have been collected since 1850. It is possible that it is extinct, as Argus and Pryer (1990) and McJannet et al. (1995) have suggested, but I think it more likely that its distribution is so limited that it has simply been overlooked by botanists. Hooker’s (1830) description of the type locality of B. pilosa, “Mouth of Macken- zie River, lat. 70°” may be somewhat misleading to those who have looked for B. pilosa over the last cen- tury or so. Porsild, for example, made several collec- tions of Braya between the mouth of the Mackenzie River and Nicholson Island in Liverpool Bay, where Hooker’s location description suggests that B. pilosa should be found. Although Porsild initially identified his collections as B. pilosa (Porsild 1943), none of them show the distinctive ovoid-ellipsoid silicles, exception- ally long styles, and large flowers of this taxon and fall instead within the normal range of diversity in B. glabella. There is evidence that Hooker’s description of the type locality may be his interpretation of Richardson’s broad conception of the Mackenzie Delta rather than a precise description of Richardson’s collection location. In his account of the 1826 journey from the Mackenzie River to Cape Bathurst, Richardson (1828) repeatedly commented on the alluvial deposits and sandbars his party encountered in the area, and correctly attributed them to outwash from the Mackenzie. He clearly viewed the entire region of the Tuktoyaktuk Peninsula, Eskimo Lakes, and Liverpool Bay as a product of a historical Mackenzie Delta. It is likely, however, that Richardson was actually very near Cape Bathurst rather than the mouth of the Mackenzie when he collected the type specimens of B. pilosa. In fact, there is some evidence that all three known collections of B. pilosa (Richardson in 1826 and 1848; Pullen in 1850) may have come from the same stretch of seashore near Cape Bathurst, roughly 200 km northeast of the mouth of the Mackenzie River. John Richardson’s original collection of B. pilosa almost certainly occurred on 18 July 1826, as the east- THE CANADIAN FIELD-NATURALIST Vol. 118 ern detachment of Franklin’s second expedition in search of a northwest passage was approaching Cape Bathurst. This eastern detachment, under Richard- son’s command, was assigned the task of exploring the arctic sea coast by boat from the Mackenzie River east to the Coppermine River, then traveling overland on foot from the Coppermine River to Great Bear Lake. On 18 July, Richardson (1828) records stopping for breakfast “about eight miles” from the narrow passage between Cape Bathurst and the Baillie Islands. He writes, “The air was perfumed by numerous tufts of a beautiful phlox, and of a still handsomer and very fra- grant cruciform flower, of a genus hitherto unde- scribed.” There is little doubt that the “cruciform” flower Richardson is describing is B. pilosa. First, 18 July is a date that an arctic Braya would be expected to be in a flowering rather than a fruiting condition. In addition, the dense clusters of large flowers on his type specimens would certainly be “handsome,” and Rich- ardson’s description of the plant’s fragrance and uncer- tain generic status matches perfectly with Hooker’s (1830). No other cruciferous species included in Hook- er’s Flora Boreali-Americana, which treats all of Rich- ardson’s collections from this expedition, fits Richard- son’s 18 July plant description as well as B. pilosa. Richardson collected B. pilosa a second time in 1848, this time when the plant was in fruit. Captain Sir John Franklin, along with his ships, the Erebus and Terror, and their crews disappeared in the Canadian Arctic in 1845. Franklin’s ships carried sufficient pro- visions to last until the summer of 1848, but by 1847, when nothing had been heard from Franklin, the British Admiralty began mounting search expeditions. In 1848, John Rae and John Richardson were placed in com- mand of one of these search parties. Richardson’s orders were to search the arctic coast and islands east of the Mackenzie River to the Coppermine River and then travel overland from the Coppermine to Great Bear Lake, essentially retracing his 1826 journey. Unfortu- nately, Richardson did not mention his 1848 collection of B. pilosa in his published account. However, I think it is likely that this second collection was made at ex- actly the same place as the first. On 10 August, Rich- ardson was approaching Cape Bathurst again, twenty- two years after his first visit there (Richardson 1851). This is exactly the right season to find Braya in fruit. Given Hooker’s uncertainty about the generic status of B. pilosa due to a lack of fruiting material for exam- ination, it is inconceivable that Richardson would not have taken the opportunity to revisit his original col- lection location to collect fruiting plants which would allow a definitive identification. Braya pilosa was collected for a third and apparently last time in August of 1850 from the “arctic coast, W. of C. Bathurst” by Captain William J. S. Pullen. In 1850, Pullen, also in command of an expedition in search of Franklin, attempted to travel from Fort Simp- son on the Mackenzie River to Cape Bathurst and then on to Banks Island. In mid-August he was in the vicin- 2004 HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA 553 KEW NEGATIVE ‘No. 1408 ITT] cance nasi te en APH aac eS | i \ | “4 7) cvselec Cnesl, Pith bo Bet hial a i hip J280 \ k FF Vi VAY aS ) ase) ) NIAIOR aanqnousey,) ap asajsiurpy @ 10g 2staay G Aap va aa de 3 plac Proook » ace Jaspdi Bi FS—F6y 199 §) five Rr ae af iy, Determinavit FiGuRE 2: Photograph of mixed collections of Braya pilosa on a herbarium sheet at the Royal Botanic Gardens Herbarium (K) in Kew, England. Insert is an enlarged view of mature silicles. 554 THE CANADIAN FIELD-NATURALIST Vol. 118 MSW 129°00W 28°45 WBS SC IBMSW «IBOOW «= 1245W O12S0W SC 121BW12P00W Ss 128°45W «126° Baillie Islands Cape Bathurst (ZL 7PR0N 70°30N IE Scale(km) 0 30 70°15N 70° 15N- SW 12°00W « 12BEW 128° wo0G 27°00 1S 45W 3 12°30W FiGuRE 3: Map of Cape Bathurst Peninsula, Northwest Territories, Canada. The dashed box indicates the probable type locality of Braya pilosa. ity of Cape Bathurst, but found dramatically different ice conditions there than Richardson had two years previously. The pack ice was still so thick that he was unable to proceed past Cape Bathurst, and was forced to retreat back to Fort Simpson. On 8 August, Pullen (1979) recorded that his party “landed in a small bay about 7 miles from Cape Bathurst” to eat dinner. They returned to the same bay about midnight on 10 Au- gust, after having been turned back by the pack ice at Cape Bathurst, and remained camped there until 15 August while they waited to see if ice conditions at the Cape might improve. This places Pullen within one nautical mile of Richardson’s (1828) estimate of his position when I believe he made his first collection of B. pilosa, and at the perfect season to find the plant in fruit. He describes the small bay as being “the only place where we could conveniently land,” which sug- gests that it is likely the same place Richardson had chosen to land for breakfast in 1826 when he discov- ered the plant. In addition, Pullen’s steersman was Neil McLeod, a Hudson’s Bay Company employee who had also accompanied Richardson in 1848. It is probable that McLeod would have mentioned Richardson’s interest in the plant to Pullen and perhaps inspired him to make his own collection during the four days they were camped at the site. If Richardson’s and Pullen’s B. pilosa collections did come from the same location on Cape Bathurst Peninsula, the position of that location would most likely be between about 70° 27' and 70° 28' north lat- itude along the western coast of the peninsula (Figure 3). Or perhaps slightly farther north if their estimates of the distance to Cape Bathurst were based on the distance they actually traveled along the coastline rather than the direct straight-line distance. This section of coastline should be searched carefully for extant populations of B. pilosa. While the rediscovery of B. pilosa would of itself be interesting botanically and historically, the avail- ability of living plants for study would also be tremen- dously valuable to our understanding of Braya. The evolution of the genus is intimately tied to hybridiza- tion and polyploidy (Harris 1985), but that phyloge- netic history is obscured by the fact that diploidy has never been reported in Braya. There is some evidence, however, that B. pilosa may be diploid and, therefore, could have played a critical role in the evolutionary history of Braya. Ornduff (1969) listed several attrib- utes in flowering plants that are correlated with xeno- gamy and a diploid chromosome number. Among them are rotate corollas with large petals, exserted styles, scented flowers, abortive fruits, and a narrow distribu- tion. Braya pilosa has the largest petals and longest styles of any Braya, it is the only species in the genus with scented flowers, it appears to have a fairly high frequency of abortive silicles, and it has a very narrow 2004 HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA SBP) FIGURE 4: Photograph of Braya pilosa specimen collected on 24 July 2004 (J. G. Harris and D. L. Taylor 3644). 556 distribution. Large, rotate flowers, allogamy, abortive fruits, and narrow distributions also have been associ- ated with lower ploidy levels in B. humilis (Harris 1985), though these are tetraploids rather than diploids. Pollen measurements also provide evidence that B. pilosa may be diploid. Rollins (1953) and Bécher (1956) noted a correlation of ploidy level and pollen size in B. humilis, with populations of lower ploidy level having somewhat smaller pollen grains than those of higher ploidy level. Pollen sizes in B. pilosa, B. thorild- wulffii, and B. glabella show a similar correlation. Mea- surements (Harris 1985) indicate that B. pilosa has significantly smaller pollen grains than the tetraploid B. thorild-wulffii, with the largest pollen grains being found in the octoploid B. glabella. If B. pilosa is indeed diploid, its extremely limited distribution may be linked to its ploidy level. Favarger (1961) and Johnson and Packer (1967) found a corre- lation between plant ploidy level and habitat stability. Diploids were more likely to be found on stable habi- tats, while polyploids were more frequent on habitats with a history of disturbance. Areas that remained unglaciated during the Pleistocene, for example, were found to have a significantly higher percentage of diploids in their floras than areas that were glaciated (Favarger 1961; Johnson and Packer 1967). The Cape Bathurst Peninsula is known to have been unglaciated during the Pleistocene (Prest 1969) and probably served as a refugium for plants that moved back onto their former ranges as the ice receded. Braya pilosa likely weathered the Pleistocene on the relatively stable habi- tats available near Cape Bathurst, but it has apparently been unable to expand its range in recent times. It may, however, have served as a parent species to some of the more widespread polyploids in the genus. A like- ly candidate is B. thorild-wulffii, a high arctic North American endemic that shares B. pilosa’s ovoid- ellipsoid fruit shape and fenestrate silicle septae, and appears to be most closely related to it. Braya tho- rild-wulffii is a tetraploid (2n=28) (Holmen 1952; Mulligan 1965, 2002; Bocher 1966; Harris 1985), which is the lowest reported ploidy level in the genus (some populations of B. humilis are also tetraploid). However, Boécher (1966) observed that there appears to be only a single large pair of chromosomes in B. thorild-wulffii, so it is probably not simply an autote- traploid from a single diploid parent. DNA sequence data (Harris, unpublished data) sug- gest that B. thorild-wulffii may have served as a parent to other Braya species of higher ploidy level. If DNA of B. pilosa were available, and if the species is indeed a parent to B. thorild-wulffii, it could provide signifi- cant insight into phylogenetic relationships within Braya and between Braya and other genera in the Cruciferae. Acknowledgments I am indebted to C. Stuart Houston for his help with historical questions about the second Franklin expe- THE CANADIAN FIELD-NATURALIST Vol. 118 dition, Ihsan Al-Shehbaz for his helpful discussions about the taxonomic status of Braya pilosa, and Alvan Bregman of the Rare Book Collections Library at Uni- versity of Illinois at Urbana-Champaign for providing me with transcripts of Richardson’s annotations to his personal copy of Flora Boreali-Americana. 1 thank Sam Rushforth, Renée Van Buren, Kimball Harper, and JaNae Brown Haas for insightful comments on previous drafts of this paper and Melinda Woolf Har- ris for her illustrations of Braya pilosa. Finally, | am indebted to two anonymous reviewers who provided valuable comments and suggestions. Addendum On 24 July 2004, Daniel L. Taylor and I briefly visit- ed Richardson’s presumed type locality of Braya pilosa near Cape Bathurst. We discovered several hundred B. pilosa individuals growing along the coastline on sandy, calcareous soils in an area heavily used by Cari- bou (Ranifer taranclus). The plants were limited to small, bare patches of soil disturbed by Caribou hoofs. The plants on Cape Bathurst Peninsula (Figure 4) are perfect matches for Richardson’s and Pullen’s 19" Century B. pilosa collections (Figure 2). Speci- mens (J. G. Harris and D. L. Taylor 3644) are deposit- ed in the Utah Valley State College Herbarium (UVSC), with duplicate material distributed to the following herbaria: Agriculture and Agri-Food Canada (DAO), Canadian Museum of Nature (CAN), Missouri Botan- ical Garden (MO), New York Botanical Garden (NY). DNA sequencing and cytological studies are currently underway (Harris, unpublished data). Documents Cited (marked * in text) Mulligan, G. A. 2000. Key to the Brassicaceae (Cruciferae) of Canada and Alaska. Electronic publication on the inter- net site, http://members.rogers.com/mulligan4520/key/ Murray, D. F. 1983. in sched., Mouth of River Mackenzie, Richardson s.n., National Herbarium of Canada (CAN) 64149, Ottawa, Ontario. Scott, P. J., S. G. Aiken, and M. J. Dallwitz. 2001. Brassi- caceae of the Canadian Arctic Archipelago: descriptions, illustrations, identification, and information retrieval. Ver- sion: 29" June 2001. Electronic publication on the internet site, http://www.mun.ca/biology/delta/arcticf/ Warwick, S. I., A. Francis, and G. A. Mulligan. 2000. Bras- sicaceae of Canada. Agriculture and Agri-Food Canada, Ottawa. Electronic publication on the internet site, http:// sis.agr.gc.ca/brd/brass/ Literature Cited Argus, G. W., and K. M. Pryer. 1990. Rare vascular plants in Canada: our natural heritage. Canadian Museum of Nature, Ottawa. Bocher, T. W. 1956. Further studies in Braya humilis and allied species. Meddelelser om Gronland 124 (7): 1-29. Bocher, T. W. 1966. Experimental and cytological studies in plant species. IX. Some arctic and montane Crucifers. Biologiske Skrifter Kongelige Danske Videnskabernes Selskab 14(7): 1-74. 2004 Cody, W. J. 1979. Vascular plants of restricted range in the continental Northwest Territories, Canada. National Muse- um of Natural Sciences, National Museums of Canada, Ottawa, Ontario. Cody, W. J. 2000. Flora of the Yukon Territory, 2™ edition. National Research Council of Canada, Ottawa. Favager, C. 1961. Sur ?emploi des nombres de chromo- somes en géographie botanique historique. Berichte des Geobotanischen Institutes der Eidgendssischen Technis- chen Hochschule Stiftung Riibel 32: 119-146. Harris, J. G. 1985. A taxonomic revision of the genus Braya (Cruciferae) in North America. Unpublished Ph.D. thesis, University of Alberta, Edmonton, Alberta. Holmen, K. 1952. Cytological studies in the flora of Peary- land, North Greenland. Meddelelser om Gr@nland 128 (5): 1-40. Hooker, J. D. 1830. Flora Boreali-Americana. 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Chromosome numbers determined from Canadian and Alaskan material of native and natu- ralized mustards, Brassicaceae (Cruciferae). Canadian Field-Naturalist 116(4): 611-622. Polunin, N. 1959. Circumpolar arctic flora. Clarendon Press, Oxford. HARRIS: PILOSE BRAYA: AN ENDEMIC OF ARCTIC CANADA a5) Porsild, A. E. 1943. Materials for a flora of the continental Northwest Territories of Canada. Sargentia IV: 44-47. Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Con- tinental Northwest Territories, Canada. National Museums of Canada, Ottawa. Prest, V. K. 1969. Retreat of Wisconsin and recent ice in North America. Geological Survey of Canada, Department of Energy, Mines and Resources, Map 1257-A. Pullen, W. J. S. 1979. Pages 114-126 in H. F. Pullen. The Pullen expedition: In search of Sir John Franklin: The orig- inal diaries, log, and letters of Commander W. J. S. Pullen. The Arctic History Press, Toronto, Ontario. Richardson, J. 1828. 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The flora of Canada. Part 3. National Museums of Canada, Ottawa. Welsh, S. L. 1974. Anderson’s flora of Alaska and adjacent parts of Canada. Brigham Young University Press, Provo, Utah. Received 20 October 2003 Accepted 6 December 2004 New Records of Vascular Plants in the Yukon Territory VI WILLIAM J. Copy!, CATHERINE E. KENNEDY”, BRUCE BENNETT’, and PHIL CASWELL? ‘Biodiversity, National Program on Environmental Health, Agriculture and Agri-Food Canada, Wm. Saunders Building (49), Central Experimental Farm, Ottawa, Ontario KIA 0C6 Canada *Department of Environment, Government of the Yukon, Box 2703, Whitehorse, Yukon Y1A 2C6 Canada 3P. O. Box 91, Roxbury, New York 12474 USA Cody, William J., Catherine E. Kennedy, Bruce Bennett, and Phil Caswell. 2004. New records of vascular plants in the Yukon Territory VI. Canadian Field-Naturalist 118(4): 558-578. Based on field reconnaissance mainly in 2002 in the southern part of the Yukon and particularly in and adjacent to Kluane National Park, information is provided on geographically significant plant occurrences. Six native taxa, Atriplex alaskensis, Claytonia megarrhiza, Corispermum ochotense vat. alaskanum, Oxytropis arctica, Polemonium acutiflorum forma lacteum and Polemo- nium boreale forma albiflorum, and four introduced taxa. Arabis caucasica, Camelina sativa, Senecio eremophilus, and Setaria viridis are reported new to the known flora of the Yukon Territory. Significant range extensions for 158 native and 21 introduced taxa are included. Parrya arctica, Armoracia rusticana, Atriplex patula and Papaver nudicaule ssp. nudicaule are excluded from the Yukon flora. 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 V (Cody et al. 2003), a consider- able number of plant specimens have been submitted to Cody for identification and confirmation. The major submissions include the following locations and col- lectors: (1) Kluane National Park and vicinity by Phil Caswell for the National Park Service as part of on- going botanical inventories; (2) W. J. Cody and Cath- erine Kennedy spent four days collecting specimens in the Fort Selkirk area assessing the flora of that region; (3) W. J. Cody and Margaret Cody studied changed vegetation along the southern highways; (4) Bruce Ben- nett conducted studies adjacent to the North Canol Road, assisting with rare plant surveys for Yukon- Charley Rivers National Park and interesting areas while boating down the Yukon River; (5) Rhonda Rosie visited sites adjacent to Nisling Lake and Morris Lake for the Yukon Territory Government and Parks Branch; (6) Stu Withers visited former mine sites at Atlin Lake and Deadman Creek in connection with reclamation of vegetation and soil stabilization; (7) with his continuing interest in vegetation habitats Greg Brunner collected plant specimens from areas adjacent to the Klondike River and Dawson City. 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 records recently published (Cody et al. 1998, 2000, 2001, 2002, 2003). The floristic information presented earlier and updated here is essential for biological re- search and ongoing work relating to agriculture, fores- try, sustainable resource management and wildlife management. With additions of six native and four introduced species reported here, the flora now includes 1181 species. The new native species are all rare (as defined by Douglas et al. 1981). The taxa addressed in the body of this paper appear in a synoptic list by Yukon status in alphabetical order. The taxa are then discussed in taxonomic order, as pre- sented in the Flora of the Yukon Territory with citation of specimens and other pertinent information. Common names follow Cody (1996) and Douglas et al. (1998- 2001). Synoptic List by Yukon Status Errata in New Records: (5) Carex stylosa Douglasia arctica Poa nemoralis revised to Poa interior Armoracia rusticana revised to Rorippa barbareifolia Papaver nudicaule ssp. nudicaule revised to Papaver croceum Native Taxa New to the Yukon Territory: (6) Atriplex alaskensis Claytonia megarrhiza Corispermum ochotense var. alaskanum Oxytropis arctica Polemonium acutiflorum forma lacteum Polemonium boreale forma albiflorum Introduced Taxa New to the Yukon Territory: (4) Arabis caucasica Camelina sativa Senecio eremophilus Setaria viridis Range Extensions of Native Taxa within the Yukon Territory: (158) Agrostis filiculmis Allium schoenoprasum ssp. sibiricum Alyssum obovatum Amelanchier alnifolia Arabis holboellii var. retrofracta Arabis holboellii var. secunda Aster alpinus ssp. vierhapperi Aster falcatus 558 Atriplex subspicata Bromus ciliatus Callitriche anceps Cardamine bellidifolia Carex albo-nigra Carex athrostachya Carex atrosquama Carex capillaris ssp. chlorostachys Carex crawfordii Carex eleusinoides Carex foenea Carex limosa Carex loliacea Carex obtusata Carex pachystachya Carex pellita Carex petasata Carex praegracilis Carex rossii Carex supina ssp. spaniocarpa Carex sychnocephala Carex viridula Cassiope tetragona ssp. saximontana Cerastium arvense Chamaerhodos erecta ssp. nuttallii Chenopodium capitatum Cicuta maculata var. angustifolia Cryptogramma crispa vat. acrostichoides Deschampsia brevifolia Descurainia incisa vat. incisa Douglasia alaskana Draba aurea Draba corymbosa Draba crassifolia Draba glabella Draba nemorosa Draba ogilviensis Draba palanderiana Draba porsildii Draba stenoloba Dryas hookeriana Eleocharis uniglumis Elymus alaskanus ssp. alaskanus Elymus glaucus Elymus macrourus Elymus spicatus Elymus trachycaulus ssp. andinus Erigeron caespitosus Erigeron hyperboreus Eriophorum brachyantherum Eritrichium aretioides Eritrichium chamissonis Euphrasia subarctica Festuca brachyphylla Festuca richardsonii Festuca saximontana Gentianella tenella Geum aleppicum ssp. strictum Geum rossii Glyceria striata var. stricta Gymnocarpium jessoense ssp. parvulum Harrimanella stellariana Hordeum brachyantherum Tris setosa ssp. interior Isoetes echinospora 2004 Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 559 Isoetes ?maritima Juncus bufonius Juncus filiformis Kobresia simpliciuscula Koeleria macrantha Lepidium densiflorum var. densiflorum Lupinus nootkatensis Luzula arcuata ssp. unalaschkensis Maianthemum canadense ssp. interius Mertensia paniculata var. alaskana Mimulus guttatus Minuartia dawsonensis Minuartia elegans Minuartia macrocarpa Minuartia yukonensis Montia fontana Myriophyllum verticillatum Nuphar polysepalum Orobanche fasciculata Oxytropis campestris ssp. varians Pedicularis oederi Phacelia mollis Phlox hoodii Phyllodoce glanduliflora Pinguicula villosa Pinus contorta ssp. latifolia Plantago canescens Plantago maritima Poa interior Poa leptocoma Poa secunda Podistera macounii Polygonum lapathifolium Potamogeton alpinus ssp. tenuifolius Potamogeton berchtoldii Potamogeton gramineus Potamogeton pectinatus Potamogeton praelongus Potamogeton subsibiricus Potamogeton zosteriformis Potentilla arguta var. convallaria Primula eximia Puccinellia interior Pulsatilla ludoviciana Pyrola minor Ranunculus aquatilis var. subrigidus Ranunculus occidentalis var. brevistylis Rhinanathus minor ssp. borealis Ribes oxyacanthoides ssp. oxyacanthoides Rorippa barbareifolia Rosa woodsii Salix alaxensis ssp. longistylis Salix glauca vat. acutifolia Salix pseudomyrsinites Salix rotundifolia ssp. dodgeana Sambucus racemosa ssp. pubens Saxifraga caespitosa Saxifraga eschscholtzii Saxifraga foliolosa Saxifraga nelsoniana ssp. pacifica Saxifraga rufopilosa Saxifraga tricuspidata Scirpus rollandii Silene involucrata ssp. tenella Silene williamsii 560 Solidago canadensis vat. salebrosa Spiranthes romanzoffiana Stellaria umbellata Stipa comata Stipa nelsonii ssp. dorei Stipa richardsonii Subularia aquatica ssp. americana Tofieldia coccinea Trientalis europaea Triglochin palustre Typha latifolia Urtica dioica ssp. gracilis Utricularia minor Vaccinium membranaceum Vahlodea atropurpurea Valeriana sitchensis Vicia americana Viola renifolia var. brainerdii Woodsia alpina Woodsia ilvensis Range Extensions of Introduced Taxa within the Yukon Territory: (21) Agropyron pectiniforme Agropyron sibiricum Agrostis gigantea Avena sativa Brassica rapa Bromus inermis Capsella bursa-pastoris Elytrigia intermedia Galeopsis tetrahit ssp. bifida Lepidium densiflorum var. macrocarpum Myosotis scorpioides Papaver croceum Poa annua Rheum rhabarbarum Rumex crispus Silene vulgaris Sinapis arvensis Sonchus arvensis ssp. uliginosus Tanacetum vulgare Thlaspi arvense Trifolium pratense Comments on Native Taxa in the Yukon Flora: (3) Carex stylosa Douglasia arctica Isoetes maritima Deletions of Native Taxa from the Yukon Flora: (1) Parrya arctica Deletions of Introduced Taxa from the Yukon Flora: (3) Armoracia rusticana Atriplex patula Papaver nudicaule ssp. nudicaule Annotated Species List ISOETACEAE Isoetes echinospora Dut., Bristle-like Quillwort — YUKON: On stony shore of small pond south of Morris Lake, 60°20°N 131°40°W, R. Rosie 2030, 8 Sept. 1999 (DAO); washed up on lakeshore, northeast shore of Nisling Lake, 62°02’00"N_ 137°58’30"W, R. Rosie THE CANADIAN FIELD-NATURALIST Vol. 118 02-821, 12 Aug. 2002 (DAO) (determined by D. F. Brunton). The first specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 180 kilo- meters southwest of the vicinity of Francis Lake. The second specimen is from a site about 500 kilometers northwest of Frances Lake and about 430 kilometers northwest of the Morris Lake site. Isoetes ?maritima Underw., Maritime Quillwort — YUKON: washed ashore, WNW side of Trout Lake, 68°49°27"N 138°45'8"W, J. M. Line 2000-72, 28 July 2000 (DAO) (determined by D. F. Brunton & D. M. Britton). The fragmented specimens found washed up along the shore of the lake possessed only immature female spores which were mostly densely tuberculate but could not defi- nitely be determined as /. maritima. If this could be verified by more mature specimens from Trout Lake it would be only the third known location in the Territory and a range exten- sion of about 260 kilometers northwest of Vittrewka Lake (Cody et al. 2001). Isoetes maritima Underw., Maritime Quillwort — It has been brought to our attention by Lori Schroeder (Environment Yukon) that specimen data according to Greg Brunner’s field notebook for his collection #39-99 which was published in Cody et al. (2001) should be: Peel River Wetland Study Turner Lake, 66°11'N 134°14'W, 2 July 1999. PTERIDACEAE Cryptogramma crispa (L.) R. Br. var. acrostichoides (R. Br.) C. B. Clarke, Mountain-parsley — YUKON: Kluane National Park, south-facing scree slope, 60°06.019'N 137°08.532'W, P. Caswell 026, 6 June 2002 (DAO). This rare species which was previously known from only six localities in the Territory is now known from two sites in southern Kluane National Park. ASPIDIACEAE Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvu- lum Sarvela, Nahanni Oak Fern — YUKON: dry rock out- crop, lower Beaver River, 60°09'09"N 124°55'16"W, B. Bennett 97-518, 19 Aug. 1997 (B. Bennett Herbar- ium, photo DAO). The specimen cited above of this rare plant in the Territory is an extension of the known range of about 300 kilometers from a site in the vicinity of Francis Lake (Cody 1996). Woodsia alpina (Bolton) S. F. Gray, Northern Woodsia — YUKON: damp organic soil in rock crevice, Cache Lake, Kluane National Park, 61°12.586'N 139°03.671'W, P. Caswell 575, 24 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory of about 45 kilometers southwest of a site at the north end of Kluane Lake and is new to the flora of Kluane National Park. Woodsia ilvensis (L.) R. Br., Rusty Woodsia — YUKON: steep south-facing bluffs along Klondike River, Dawson Area, | mile north of 64°02'N 137°41'30"W, G. Brunner 435, 2 June 2001 (DAO); dry rocky outcrop, 2004 S Fork bluffs, Klondike River, 64°O1'N 138°9'W, G. Brunner 443-01, 14 July 2001 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimens cited above are from sites about 60 kilometers southwest of a site adjacent to the Dempster Highway mapped by Cody (1996). PINACEAE Pinus contorta Doug}. ex Loud. ssp. latifolia Engelm. ex S. Wats., Lodgepole Pine — YUKON: roadside, North Canol Road KM 284, 62°13'02"N 131°46'10"W, B. Bennett 02-723, 22 Sept. 2002 (DAO); roadside across road from large truck dump, North Canol Road Km 380, 62°50'03"N 130°56'24"W, B. Bennett 02-724, 21 Sept. 2002 (DAO). The specimens cited above are an extension of the known range in the Territory of about 175 kilometers northeast of a site adjacent to the South Canol Road. The second specimen was from a site adjacent to an old house and may have been planted. TYPHACEAE Typha latifolia L., Common Cattail — YUKON: border of small lake below swimming pool, Takhini Hot Spring, 60°52'40.7"N 135°21'30.6"W, Cody & Cody 37692, 10 Aug. 2001 (DAO); same locality (fruiting) Cody 35311, 4 Aug. 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory. Cody (1996) knew it only from the vicinity of Mayo. Cody et al. (2000, 2001) added additional sites in the extreme southeast and in the vicinity of Faro. The specimen cited above which was collected just west of Whitehorse is now from the fourth site in the Territory. POTAMOGETONACEAE Potamogeton alpinus Balbis ssp. tenuifolius (Raf.) Hultén — YUKON: plant fragments washed ashore, Trout Lake, 68°50'N 138°45'W, J. M. Line 2000-9, 29 July 2000 (DAO). The specimen cited above is the northernmost yet found in the Territory. It is an extension of the known range (Cody 1996) of about 85 kilometers NE of a site north of latitude 68°N. Potamogeton berchtoldii Fieb. — YUKON: shallow water of small lake, west side of Dempster Highway Km 93, 64°39'01"N 138°23'01"W, Cody & Cody 38060, 27 July 2002 (DAO). The specimen cited above is from a site about 80 kilo- meters northeast of Dawson City. To the north a single site was previously known from adjacent to the Peel River (Cody 1996). Potamogeton gramineus L., Variable-leaved Pondweed — YUKON: slough off Yukon River with old sunken dredge, 61°56'35"N 135°12'17"W, B. Bennett 02-299, 7 Aug. 2002 (DAO). The nearest site to that listed above known to Cody (1996) is about 75 kilometers to the north. Potamogeton pectinatus L., Sago Pondweed — YUKON: shallow backwater, Dezadeash River, Kluane National Park, 60°45.683'N 137°36.010'W, P. Caswell S05, 16 Aug. 2002 (DAO); locally common in mucky alka- Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 56] line pond, Mile 985 Alaska Highway, 60°48'N 136°45'W, Calder & Kukkonen 28277, 14 Aug. 1960 (DAO). The specimens cited above extend the known range in the Territory about 120 kilometers west of the vicinity of White- horse. The Caswell specimen is the first known from Kluane National Park. Potamogeton praelongus Wulf., White-stemmed Pondweed — YUKON: Enoch Lake, Old Crow Flats, 68°05'00"N_ 140°09'50"W, B. Bennett 95-413, 10 Aug. 1995 (B. Bennett Herbarium, photo DAO); plant fragments washed ashore, Trout Lake, 68°50'N 138°45'W, J. M. Line 2000-140, 28 July 2000 (DAO); Nordenskiold River wetland pond, M. Dennington, Nord #23, 19 July 1984 (B. Bennett Herbarium, photo DAO). Douglas et al. (1981) considered this species rare in the Yukon Territory based on specimens from only two locations. Cody (1996) knew it from five locations. Cody et al. (2000) added three locations. The first specimen cited above is a northwestward extension of about 40 kilometers from the vicinity of Old Crow. The second specimen from about 115 kilometers NNE of Old Crow is the northernmost yet found in the Territory. The third specimen is an extension of the known range in the south of about 125 kilometers northwest of a site just north of Whitehorse. Potamogeton subsibiricus Hagstr. - YUKON: growing in about 3 feet of water, lake west of Dempster Rd. near mile 65, R. T: Porsild 317, 25 July 1966 (AKA, photo DAO) (determined by A. Batten). Cody (1996) knew this rare species in the Territory from a single collection from adjacent to the North Canol Road about 450 kilometers to the southeast of the locality cited above. Potamogeton zosteriformis Fern., Flatstemmed Pond- weed — YUKON: in water by outlet of pond, Pumphouse Pond, 3 km S of Alaska Highway, 60°43'31.3"N 135°09'58.3"W, W. J. Cody 38290, 3 Aug. 2002 (DAO). Cody (1996) knew this rare species in the Territory from only two localities, one southeast of Haines Junction and one just north of 64°N. Cody et al. (1998, 2000 and 2001) added two sites in the south and one north of 68°N. SCHEUCHZERIACEAE Triglochin palustre L., Marsh Arrow-grass — YUKON: silty river bank with Carex utriculata, C. aquatilis and Juncus alpinoarticulatus, Selwyn, Yukon River, 62°48'06"N 138°15'29"W, B. Bennett 02-294, 9 Aug. 2002 (DAO). The specimen cited above is between sites adjacent to the Alaska Highway, Klondike Highway and Dawson City. POACEAE (GRAMINEAE) Agropyron pectiniforme R. & S., Crested Wheat Grass — YUKON: lake shore, Earn Lake, 62°.9'08.3"N 134°23'07.6"W, G. Brunner 557-01, 5 Aug. 2001 (DAO); steep stony slope on east side of highway, Haines Highway, 60°6'3.2"N 136°54'51"W, Cody & Cody 37911, 21 July 2002 (DAO). 562 The first specimen of this introduced species cited above is from a site between the northernmost mapped by Cody (1996) in the vicinity of Mayo and a site adjacent to Faro. The second specimen cited above is an extension of the known range in the Territory of about 85 kilometers south- east of a site in the vicinity of Haines Junction. Agropyron sibiricum (Willd.) P. Beauv. — YUKON: top of bank overlooking road into gravel pit south of Alaska Highway, 60°49'58"N 135°45'49"W, Cody & Cody 38210, 1 Aug. 2002 (DAO). Cody (1996) knew this introduced species in the Territory only from two locations, Whitehorse and Carmacks. A third site was reported from the vicinity of Ross River (Cody et al. 2003). The specimen cited above is from a site about 40 kilo- meters west of Whitehorse. Agrostis filiculmis M. E. Jones — YUKON: open area — abandoned placer mine, Mechanic Creek, 62°20.1'N 137°21.5'W, S. Withers SWOI-062, 6 July 2001 (DAO). The specimen cited above is only the second known in the Territory (Cody 1996) and is from a site about 200 kilometers southeast of the first site in the vicinity of Dawson City. Agrostis gigantea Roth., Creeping Bent Grass — YUKON: Spruce Hill Park, border of playground at head of Engelman Drive, Whitehorse, 60°36'40.3"N 134°55'08.8"W, W. J. Cody 37743, 14 July 2002 (DAO). Cody (1996) knew this introduced species in the Territory from only two localities, Dawson City and Carmacks. An additional collection from a mine site east of Dawson City was reported by Cody et al. (2003). The specimen cited above is from only the fourth known locality in the Territory. Avena sativa L., Oats — YUKON: scattered in grass area around Beringia Museum, Whitehorse, 60°42'36"N 135°4'45"W, Cody & Cody 38105, 30 July 2002 (DAO). Cody (1996) knew this introduced species from only wide- ly separated sites in the Territory, vicinity of Mayo, middle of North Canol Road and east of Watson Lake. Bromus ciliatus L. — YUKON: in moist moss on moun- tain slope above treeline on east side of Dempster Highway about 9 kilometers north of Tombstone Park Campsite, 64°34'49"N 138°15'30"W, Cody & Cody 38047, 26 July 2002 (DAO). The specimen cited above is at the northern limit of this species in the Territory (Cody 1996). The nearest sites previ- ously mapped were about 120 kilometers to the west adjacent to the Yukon River and about 150 kilometers to the southeast in the vicinity of Stewart Crossing. This species was con- sidered rare in the Territory by Douglas et al. (1981). Bromus inermis Leyss., Smooth Brome — YUKON: sand beach, Kusawa Lake Campground, 60°5'10"N 136°8'42"W, Cody & Cody 38144, 31 July 2002 (DAO); gravel ditch between road and steep gravel slope, Haines Highway, 60°6'3.3"N 136°54'51"W, Cody & Cody 37912, 21 July 2002 (DAO). The specimens cited above of this introduced species which are from sites about 100 kilometers southeast of Haines Junction and southwest of Whitehorse, are the most south- western yet found in the Territory. THE CANADIAN FIELD-NATURALIST Vol. 118 Deschampsia brevifolia R. Br. — YUKON: flats 7 miles east of Tagish Bridge, 60°19'5"N 134°10'17"W, Cody & Cody 37940, 23 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 90 kilometers southeast of a site west of Whitehorse. Elymus alaskanus (Scribn. & Merr.) A. Léve ssp. alaskanus — YUKON: in sandy soil of river bank, Nis- utlin River Delta National Wildlife Area, 60°11'N 132°35'W, B. Bennett 98-167, 26 July 1998 (DAO). The specimen cited above is an extension of the known range in the Territory of about 180 kilometers south of a site adjacent to the northern South Canol Road. Elymus glaucus Buckl., Western Rye Grass — YUKON: roadstop in disturbed gravelly soil, Atlin Road just north of BC border, 60°00'09"N 133°47'42.1"W, Cody & Cody 37547, 28 July 2001 (DAO) Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above 1s an extension of about 115 kilometers southeast of a site adjacent to Whitehorse reported by Cody et al. (2002). Elymus macrourus (Turcez.) Tzvelev — YUKON: grow- ing in loose sand on riverbar, Wind River, camp #6, 65°40.46'N 135°11.76'W, B. Bennett 00-853, 7 July 2000 (B. Bennett Herbarium, photo DAO); exposed sandy lakeshore, north end of Atlin Lake, 60°01.5'N 133°50.4'W, S. Withers SWO1-045, 30 June 2001 (DAO). The first specimen of this Amphi-Beringian species cited above is the first known from between latitudes 64°N and 66°N east of the Dempster Highway. The second specimen is an extension of the known range in the Territory of about 200 kilometers southwest of a site east of Haines Junction. Elymus spicatus (Pursh) Gould — YUKON: common at base of large talus slope, Yukon River, Brittania Creek, 62°52'28"N 138°42'56"W, B. Bennett 02-641, 10 Aug. 2002 (DAO). Cody (1996) knew this species in the southwest of the Ter- ritory and disjunct to the extreme northwest. The specimen cited above is an extension of the known range in the Territory of about 85 kilometers west of longitude 137°W. Elymus trachycaulus (Link) Gould ex Shinners ssp. andinus (Scribn. & Smith) A. & D. Léve — YUKON: gravel roadside, Top of the World Highway Km 16, 64°6'6"N 139°38'41"W, Cody & Cody 38035, 38037, 26 July 2002 (DAO); gravel roadside, Top of the World Highway Km 102, 64°05'33"N 140°55'22"W, Cody & Cody 38003, 26 July 2002 (DAO). The specimens cited above extend the known range in the Territory to the west about 140 kilometers from a site east of the Dempster Highway. Festuca brachyphylla Schultes & Schultes f., Short- leaf Fescue — YUKON: Site KPL #106, 62°17'38.7"N 137°48'25"W, G. Brunner 578-01, July 2001 (DAO). The specimen cited above is from a site between sites mapped by Cody (1996) west of Carmacks and adjacent to southern Dempster Highway. 2004 Festuca richardsonii Hooker — YUKON: beside test plot, | km N of Montague Roadhouse, Km 132 Klondike Highway, 61°48'35"N_ 136°7'52"W, Cody & Cody 38082, 28 July 2002 (DAO). This species is frequent in the southern part of the Terri- tory west of longitude 137°W and north of latitude 64°N but has not previously been found adjacent to the Klondike Highway. Festuca saximontana Rydb., Rocky Mountain Fescue — YUKON: gravel beside road, Alaska Highway 8 km west of White River, 62°3'15.6"N 140°38'20.5"W, Cody & Cody 37886, 37887, 20 July 2002 (DAO). The specimens cited above are an extension of the known range in the Territory of about 100 kilometers northwest of a site just northwest of Kluane Lake. Glyceria striata (Lam.) Hitche. var. stricta (Scribn.) Fern., Fowl Marina Grass — YUKON: moist muddy organic soil, along road from Dalton Post to Wade Lakes, 60°07.765'N 137°04.105'W, P. Caswell 650, 4 Aug. 2002 (DAO). This plant was considered rare in the Territory by Douglas et al. (1981). Cody (1996) knew it only in the southeast as far west as just west of Watson Lake. The specimen cited above from between the Haines Highway and Kluane Na- tional Park is from a site about 450 kilometers to the west. It is however known from British Columbia. Hordeum brachyantherum Nevski — YUKON: disturbed area in townsite, Big Salmon Village at confluence of Yukon River, 61°53'N 134°55'W, B. Bennett 02-292, 7 Aug. 2002 (DAO). Cody (1996) considered this species to be introduced at Dawson City but native and rare at Carcross. Sites in the vicinity of Whitehorse (Cody et al. 2001) were considered to be introduced. The specimen cited above was probably also introduced. Koeleria macrantha (Ledeb.) Schultes, June Grass — YUKON: south aspect-xeric-sage/graminoid-forbs of low compact growth, Site YPC103 near confluence of Pelly and Yukon rivers, 62°48'16"N 137°19'35.9"W, G. Brunner 536-01, 3 Aug. 2001 (DAO) (determined by S. Darbyshire); gravel pit just north of the Alaska Highway, Silver City Road, 61°01'22"N 138°19'52"W, B. Bennett 01-005, 29 Apr. 2001 (DAO); lower slope south-facing, dry, Five Fingers Coal Mine, Yukon River, 62°12'21"N 136°20'10"W, B. Bennett 02-295, 8 Aug. 2002 (DAO). Douglas et al. (1981) considered this species rare in the Ter- ritory. The first specimen cited above was collected just south- west of the Pelly River Ranch at 62°50'20"N 137°11'40"W by M. Johansen in 1989 (DAO) (Cody 1996); the second specimen is an extension of about 60 kilometers northwest of a site adjacent to Haines Junction; the third specimen is from a site about 40 kilometers southeast of a site near Minto (Cody 1996). Poa annua L., Annual Blue Grass — YUKON: vicinity of campground, Rancheria, at mile 710 Alaska Highway (60°0S'N 130°36'W), S. L. Welsh & G. Moore 7539, 29 June 1968 (AKA, photo DAO) (determined by R. J. Soreng). Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 563 Cody (1996) knew this introduced species from only five sites in the Territory. The specimen cited above is about 100 kilometers west of a site adjacent to the southern Canol Road. Poa interior Rydb. (P. nemoralis sensu Cody 1996) — YUKON: in subalpine meadow, Onion Lake, ca. 46 mi S of Haines Junction, G. W. & G. G. Douglas 7089, 12 Aug. 1973 (AKA, photo DAO) (determined by R. J. Soreng); common on steep, open, west-facing prairie slopes, Conglomerate Mountain on Dawson-White- horse road, 61°38'N 135°53'W, Calder & Gillett 25770, 22 June 1960 (DAO); common on south-facing, prairie slope with scattered aspen, Mile 30 on road to Dawson from Stewart Crossing, 63°33'N 137°25'W, Calder & Gillett 25037, 4 June 1960 (DAO); grown on open southern slope, Pelly Ranch (ca. 62°49'N 136°34'W), J. Y. Tsukamoto s.n., 23 July 1960 (DAO); silt bluff, south-facing, on Porcupine River, 1.5 km upstream from junction between Rat Indian Creek and Porcupine River, 67°34'N 138°21'W L. Cwyner 977, 22 July 1976 (DAO) (determined by M. Bark- worth); dry open exposed gravel ridge overlooking delta, west side of river on ridge campsite, Lower Blow River Delta, 69°53'N 137°10'W, H. L. Dickson & D. L. Allen 5324, 26 July 1982 (DAO); open meadow- abandoned beaver pond, Deadman Creek, 60°20.2'N 133°03.5'W, S. Withers SWO1-126, 16 July 2001 (DAO) (determined by S. Darbyshire). ; Yukon specimens previously mapped as Poa nemoralis by Cody (1996) have been revised to P. interior. In addition, the specimens cited above which were inadvertently missed have now been mapped. (See new map Figure 1). Poa interior FiGurE 1. New distribution map for Poa interior in the Yukon Territory. 564 Poa leptocoma Trin. — YUKON: steep gravelly slope adjacent to Alaska Highway 3 km east of Snag Junc- tion, 62°12'42.3"N 140°41'33.1"W, Cody & Cody 37863, 20 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory of about 175 kilometers northwest of a site in Kluane National Park. Poa secunda Presl, Sandberg Bluegrass — YUKON: commonly found growing on the base of the bluff and occasionally on midslope, Minto Bluff, 62°36'18"N 136°51'06"W, B. Bennett 02-013, 9 June 2002 (DAO). The specimen cited above is an extension of about 175 kilo- meters northwest of a site north of Whitehorse (Cody 1996). To the north the only other site in the Territory is in the vicin- ity of Dawson City. Puccinellia interior Th. Sor. — YUKON: gravel roadside, Top of the World Highway Km 102, 64°05'33"N 140°55'22"W, Cody & Cody 38002, 26 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 75 kilometers west from the vicinity of Dawson City. Setaria viridis (L.) Beauv., Green Bristlegrass (Fig- ure 2) — YUKON: in crack at edge of Extra Foods park- ing lot, downtown Whitehorse, B. Bennett 95-437, 14 Oct. 1988 (B. Bennett Herbarium, photo DAO). The specimen cited above is a new introduction to the Flora of the Yukon Territory (Cody 1996). This species which is introduced from Eurasia is known across Canada from Newfoundland to British Columbia and has been found at Fort Simpson in the Northwest Territories. The genus Setaria can be separated from the genus Panicum as follows: A. Spikelet surrounded by 1|-many distinct or more or less connate bristles, these forming aninvolucre eaters ce ee ee Setaria B. Spikelet not subtended by bristles Stipa comata Trin. & Rupr., Needle-and-thread — YUKON: 45° silty sand south-facing slope with Arte- mesia frigida and Calamagrostis purpurascens, Dutch Bluff, Yukon River, 61°55'N 135°03.99'W, B. Bennett 02-289, 7 Aug. 2002 (DAO). Cody (1996) knew this species in the Territory from only four sites south of 63°N. Cody et al. (2003) added an addi- tional site just east of Haines Junction. The specimen cited above is from a site about 60 kilometers southeast of Car- macks. Stipa nelsonii Scribn. ssp. dorei Barkworth & Maze — YUKON: treeless rocky slope overlooking lake, Snafu Lake Camp Site, 60°08'10"N 133°48'23.6"W, Cody & Cody 37567, 28 July 2001 (DAO); open area adjacent to Populus tremuloides woodland overlook- ing lake, 7 km east of Tagish Bridge, 60°19'5"N 134°10'17"W, Cody & Cody 37938, 23 July 2002 (DAO). Cody (1996) knew this species from only six sites in the Territory north to about latitude 61°N. Cody et al. (2001) added an additional site in the vicinity of the Carcross Dunes. The specimens cited above are from east and southeast of the Carcross Dunes site. THE CANADIAN FIELD-NATURALIST Vol. 118 FIGURE 2. Setaria viridis, Green Bristlegrass (drawn by Lee Mennell). Stipa richardsonii Link (Achnatherum richardsonii (Link) Barkw.), Spreading Needlegrass — YUKON: on steep slope found on lower and mid-slope with Arabis, Arctostaphylos uva-ursi, Festuca saximontana and Poa, hill across Tagish Road from Crag Lake, 60°15'30"N 134°28'53"W, B. Bennett 01-008, 19 May 2001 (DAO); summit of hill behind Carcross Cutoff, south-facing, pine forest Cowley Creek, 60°35.66'N 134°52.96'W, B. Bennett 99-300, 6 Aug. 1999 (DAO). Douglas et al. (1981) knew this rare species in the Territory from only one locality from adjacent to the Alaska Highway just east of the Atlin Road junction (Calder & Gillett 26477, DAO). Cody (1996) mapped three additional sites in the vicin- ity of Whitehorse. The first specimen cited above is the sou- thernmost yet found in the Territory. Vahlodea atropurpurea (Wahlenb.) Fries, Mountain Hairgrass — YUKON: Site YPN 120, 63°06'39.8"N 133°20'15"W, G. Brunner 568-01, 6 Aug. 2001 (DAO). Cody (1996) knew this taxon from only six localities: adjacent to the North Canol Road, adjacent to the South Macmillan River and near Bennett Lake. Additional sites have since been found east of the Canol Road (Cody et al. 1998 and 2000). The specimen cited above is from a site about 50 kilometers northeast of the Macmillan River site. CYPERACEAE Carex albo-nigra Mack., Two-toned Sedge — YUKON: Volcano Mountain, 62°55'N 137°23'W, G. Brunner 521-01, 3 Aug. 2001 (DAO); moist scree with organic component on north-facing slope, Vulcan Mountain, Kluane National Park, 60°54.712'N 138°29.484'W, P. 2004 Caswell 527, 15 July 2002 (DAO); moist organic soil on southwest facing slope on old semi-abandoned road north of Donjek River bridge, 61°42.209'N 139°48.376'W, P. Caswell 196, 25 June 2002 (DAO). Cody (1996) knew this rare species in the Territory from seven localities south of latitude 63°N, three of which were in Kluane National Park. The first specimen cited above is the northernmost yet found in the Territory. Cody et al. (2000) added another site south of Ross River and Cody et al. (2001) another site in the extreme southeast. Carex athrostachya Olney — YUKON: moist soil by lake, Five Mile Lake Campground, 5 km N of Mayo, 63°39'11.1"N 135°53'14.2"W, Cody & Cody 37641, 1 Aug. 2001 (DAO). Douglas et al. (1981) considered this species rare in the Territory on the basis of a specimen collected by R. T. Porsild on the shores of a small bog pond in the vicinity of Mayo, 63°37'N 135°55'W, on 12 May 1967 (Porsild 1975). The specimen cited above: from nearby is only the second yet found in the Territory. Carex atrosquama Mack. — YUKON: steep gravelly slope adjacent to the Alaska Highway 3 km east of Snag Junction, 62°12'42.3"N 140°41'33.1"W, Cody & Cody 37861, 20 July 2002 (DAO); cleared gravel by highway, Top of the World Highway Km 92, 64°6'6"N 140°46'47"W, Cody & Cody 38024, 26 July 2002 (DAO). The first specimen cited above is an extension of the known range in the Territory about 190 kilometers northwest from the south end of Kluane Lake (Cody 1996) and the second spec- imen is an extension of the known range of about 100 kilo- meters west of the Dempster Highway. Carex capillaris L. ssp. chlorostachys (Steven) Love et al. — YUKON: Klondike River near mouth of Little Klondike, 64°02'00"N 137°41'30"W, G. Brunner 440, 3 July 2001 (DAO). The specimen cited above is the first known to Cody (1996) between latitudes 64°N and 66°N east of the Dempster High- way. It is an extension of about 60 kilometers northeast of a site southwest of the south end of the Dempster Highway. Carex crawfordii Fern. — YUKON: sandy lake shore, Kusawa Lake, 60°34'59.2"N 136°08'26.5"W, W. J. Cody 37718, 13 July 2002 (DAO); meadow adjacent to RCMP foundation, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37749A, 16 July 2002 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). The specimen from south of Whitehorse cited above is an extension of the known range in the Terri- tory of about 250 kilometers south of a site mapped by Cody (1996) from adjacent to the Pelly River. The second specimen is from about 85 kilometers west of a site adjacent to the Pelly River. Carex eleusinoides Turcz. — YUKON: moist organic soil with pumice component, down old road west of Alaska Highway north of Donjek River bridge, 61°42.209'N 139°48.376'W, P. Caswell 205, 2 June 2002 (DAO); gravelly silt near water line, Klondike River, 64°O1'N 137°51'W, G. Brunner 546, 8 July Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 565 2001 (DAO); approximately 2 miles downstream from O’Brian Creek on Klondike River, 64°01'46"N 38°01'00"W, G. Brunner 547, 15 July 2001 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). The first specimen cited above is an extension of the range known to Cody (1996) of about 50 kilometers northwest of the north end of Kluane Lake. The second and third specimens are the first known between latitudes 64°N and 66°N east of the Dempster Highway and are an extension of about 45 kilometers northeast of a site south of the south end of the Dempster Highway. Carex foenea Willd. — YUKON: Halfway between Order- ly House and Robert Luke Cabin, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37773, 17 July 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory. Cody (1996) knew it only from three sites, two of which were in the vicinity of Pelly Crossing. The specimen cited above came from the same area. Cody et al. (1998) re- ported an additional site in the vicinity of Beaver Ridge/ Larsen Creek. Carex limosa L. — YUKON: Sphagnum bog, 64°02'N 137°41'W, G. Brunner 441, 3 July 2001 (DAO). The specimen cited above is only the second known between latitudes 64°N and 66°N east of the Dempster High- way (Cody 1996). Carex loliacea L. — Yukon: damp silty ground, forest Picea mariana, Poa glauca, Alnus incana, Salix sp., north side of Klondike River near Parker Creek, 64°OI'N 137°51'W, G. Brunner 592, 9 July 2001 (DAO); spring alongside road, gravelly, Big Gold Creek/60 Mile River Valley, 64°01'N 140°42'W, G. Brunner 116, 23 July 1991 (DAO). Cody (1996) considered this species uncommon in the Territory although mapped north to the Porcupine River. The specimens cited above are about equally distant east and west of Dawson City. Carex obtusata Lilj., Blunt Sedge — YUKON: British Mountains, 69°26'N 140°07'W, A. Martell 625, 12 July 1979 (DAO); steep south-facing bluffs along Klondike River, | mile north of 64°02'00"N 137°41'30"W, G. Brunner 432, 2 June 2001 (DAO). The first specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is an extension of the known range of about 80 kilometers northwest from sites adjacent to the Babbage River. The second specimen cited above is an extension of the known range in the Territory of about 75 kilometers southeast of a site at Sheep Mountain adjacent to the Dempster Highway (Kojima and Brooke 1985). Carex pachystachya Cham. — YUKON: open meadow, abandoned beaver pond, Deadman Creek, S. Withers SWO1-117, 16 July 2002 (B. Bennett Herbarium, photo DAO) (determined by A. A. Reznicek). This species was not included in The Rare Vascular Plants of the Yukon (Douglas et al. 1981) because it was widespread in Canada. Cody (1996) knew it from only two localities in the extreme southern Yukon. The specimen cited above is from a site about 25 kilometers south south- east of Whitehorse. 566 Carex pellita Muhl. ex Willd. — YUKON: silty river bank with Carex utriculata, C. aquatilis, and Juncus alpinoarticulatus, Selwyn, Yukon River, 62°48'06"N 138°15'29"W, B. Bennett 02-293, 9 Aug. 2002 (DAO); silty mud at edge of river at high water line with Juncus alpinoarticulatus, Brittania Creek, Yukon River, 62°52'28"N 138°42'52"W, B. Bennett 02-671, 10 Aug. 2002 (DAO); silty mud at edge of river at high water line with Juncus alpinoarticulatus, Kirkman Creek, Yukon’ River, (62°59'21;N«.139°23'07" W)) B. Bennett 02-675, 10 Aug. 2002 (DAO); river bank in area frequently scoured by river ice, confluence of 12 Mile River, Yukon River, 64°15'11"N 139°43'18"W, B. Bennett 02-806, 23 Aug. 2002 (DAO); deeply rooted, edge of river in silty sand bank, Yukon River, 64°33'28"N 140°38'24"W, B. Bennett 02-807, 24 Aug. 2002 (DAO). Cody et al. (2000) reported this plant as a rare species in the Territory based on a specimen from a site below Rink Rapids adjacent to the Yukon River. An additional site was reported from Ear Lake, Whitehorse (Cody et al. 2002). The five sites reported above extend the known range in the Terri- tory about 275 kilometers northwest of the Rink Rapids. Bruce Bennett has found that Carex pellita is a common species on the Yukon River between Pink Rapids and White River, less common but can be found downstream and even- tually being replaced by C. saxatilis. Carex petasata Dewey — YUKON: east slope adjacent to Alaska Highway ca. 3 km W of Enger Creek, 62°19'12.05"N 140°49'19.9"W, Cody & Cody 37511, 25 July 2001 (DAO); open, abandoned mine road surface, Viceroy Mine — Big Rock Zone, 64°02'N 138°17.9"W, S. Withers SWO1-057, 3 July 2002 (DAO). The first specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 215 kilometers west of a site west of Carmacks and 245 kilometers northwest of a site adjacent to Haines Junction. The second specimen cited above is at the northern limit in the Territory about 50 kilometers east of a site in the vicinity of Dawson City. Carex praegracilis Boott — YUKON: meadow adjacent to RCMP foundation, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37770B, 17 July 2002 (DAO); sandy lake shore, Kusawa Lake, 60°34'59.2"N 135°08'26.5"W, W. J. Cody 37717, 13 July 2002 (DAO). Douglas et al. (1981) knew this rare species in the Terri- tory from only three localities: Mackintosh, Whitehorse and Carcross. Cody (1996) knew it from 11 sites in the Territory, most of which were south of latitude 61°15'N. The second specimen cited above is a slight extension of range south of Whitehorse. Carex rossii R. Br., Ross’ Sedge — YUKON: decom- posed bedrock, South Fork bluffs, Klondike River, 64°O1'N 138°9'W, G. Brunner 446-01, 14 July 2001 (DAO); steep south-facing bluffs along Klondike River 1 mile north of 64°02'N 137°41'30"W, G. Brunner 433, 2 June 2001 (DAO). The specimens cited above are only the third and fourth collections known from north of latitude 64°N (Cody 1996) THE CANADIAN FIELD-NATURALIST Vol. 118 which are from about 80 kilometers east of a site adjacent to Dawson. To the south, the nearest known site is adjacent to the northern South Canol Road about 300 kilometers to the southeast. Carex stylosa C. A. Mey. — Cody et al. (2000) reported this species which is rare in the Yukon Territory from the Arctic Coast. The specimen on which it was based (Hoefs & Smitts 93-37) was misidentified and the range extension should be deleted. There was also an error in the citation where the latitude should have been 69°23'N not 60°29'N Carex supina Wahl. ssp. spaniocarpa (Steud.) Hultén — YUKON: Steep south-facing bluffs along Klondike River | mile north of 64°02'N 137°41'30"W, G. Brun- ner 434, 2 June 2001 (DAO). The specimen cited above is the first known in the Territory north of 64°N east of the Dempster Highway. The nearest known site is about 75 kilometers to the northwest cited by Kojima & Brooke (1985). Carex sychnocephala Carey — YUKON: damp, fine gravel with organic component, at base of highway embankment, Alaska Highway just north of Copper Joe Creek, 61°19.814'N 138°56.805'W, P. Caswell 570, 21 July 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is an extension of the range known to Cody (1996) of about 175 kilometers north- west by west of a site west of Whitehorse. Carex viridula Michx. — YUKON: on moist shoreline of Nares Lake south of Carcross, 60°9'20"N 134°40'1L0"W, W. J. Cody 38318, 1 Aug. 2001 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). The specimen cited above is an extension of the known range to Cody (1996) of about 80 kilometers south of Whitehorse. Eleocharis uniglumis (Link) Schult. — YUKON: open wet area in old road, adjacent to the Alaska Highway, 61°42.209'N 139°48.376'W, P. Caswell 204, 25 June 2002 (DAO); in wet moss by lake shore, Fish Lake, 60°46'3.3"N 135°03'13"W, W. J. Cody 38246, 2 Aug. 2002 (DAO). The species was considered rare in the Territory by Douglas et al. (1981). The first specimen cited above is an extension of the known range to Cody (1996) of about 150 kilometers northwest of a site northwest of Haines Junction and the second specimen is from a site just south of Whitehorse. Eriophorum brachyantherum Trautv., Short-anthered Cotton-grass — YUKON: Vuntut National Park, vicinity of Snowdrift Camp, 68°21.4'N 139°13.1'W, P. Cas- well PPC-2000-Y-070, 21 June 2000 (DAO). The specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is an extension of about 100 kilometers north from sites adjacent to the Porcupine River. Kobresia simpliciuscula (Wahlenb.) Mack., Simple Kobresia — YUKON: along road track through Picea glauca forest, Minto RV Campground, 62°35'N 136°51'W, Cody & Cody 37678, 6 Aug. 2001 (DAO). 2004 The nearest site in the Territory (Cody 1996) to the speci- men cited above is about 130 kilometers to the southeast adjacent to the Klondike Highway. Scirpus rollandii Fern. — YUKON: moist organic soil, side road west of Alaska Highway, north of Donjek River bridge, 61°42.209'N 139°48.376'W, P. Caswell 202, 25 June 2002 (DAO). Cody (1996) knew this species in the southwest of the Territory where he considered it rare. The specimen cited above is an extension of the known range of about 100 kilo- meters northwest of southern Kluane Lake. JUNCACEAE Juncus bufonius L. s.1., Toad Rush — YUKON: mud bar, Dezadeash River below Haines Junction, Kluane National Park, 60°45.545'N 137°34.470'W, P. Cas- well 791, 16 Aug. 2002 (DAO). The specimen cited above is new to Kluane National Park and is a range extension of about 290 kilometers to the south of a site mapped by Cody (1996) and a site east of Johnson’s Crossing reported by Cody et al. (2001). Juncus filiformis L. — YUKON: in muck by small lake adjacent to the Alaska Highway, 60°51'01"N 135°46'0.5"W, Cody & Cody 38185, 1 Aug. 2002 (DAO). The specimen cited above which is from a site about 75 kilometers west of Whitehorse is the westernmost yet found adjacent to the Alaska Highway. Luzula arcuata (Wahlenb.) Sw. ssp. unalaschkensis (Buch) Hultén — gravel beside road and steep slope, Al- aska Highway Km 1918, 62°16'26.1"N 140°45'6.7"W, Cody & Cody 37851, 20 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 215 kilometers northwest of a site northwest of Haines Junction. LILIACEAE Allium schoenoprasum L. ssp. sibiricum (L.) Celak, Wild Onion — YUKON: Herschel Island, 69°35'N 139°05'W, C. E. Kennedy s.n., 1 July 1985 (Environ- ment Yukon Herbarium, photo DAO). The nearest site known to Cody (1996) was adjacent to the Firth River at 69°13'N 139°35'W, about 30 kilometers southwest of the Herschel Island site. Maianthemum canadense Desf. ssp. interius (Fern.) A. & D. Love, Wild Lily-of-the- Valley — YUKON: com- mon in understory in the riparian white spruce poplar zone, Contact Creek, 60°00'00"N 127°43'44"W, B. Bennett 01-025, 25 June 2001 (DAO). Cody (1994, 1996) suggested that this taxon should be looked for in the Territory and Cody et al. (1998, 2000) re- ported it from the extreme southeast. The specimen cited above is an extension of the known range of about 80 kilo- meters southwest from a site adjacent to the Beaver River hotspring. Tofieldia coccinea Richards., Northern False Asphodel — YUKON: south aspect-xeric-graminoid/forb/lichen meadow, Volcano Mountain, 62°55'N 137°23'W, G. Brunner 521a-01, 3 Aug. 2001 (Environment Yukon, photo DAO). Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON YI 567 The specimen cited above is from the first known site between latitudes 62°N and 64°N. The nearest site to the north is about 175 kilometers to a site adjacent to the Dempster Highway and to the southwest about 190 kilometers adjacent to the Alaska Highway. IRIDACEAE Tris setosa Pall. ssp. interior (Anders.) Hultén, Wild Iris — YUKON: at high water level, confluence of 12 Mile River and Yukon River, 64°15'11"N 139°43'18"W, B. Bennett 02-743, 23 Aug. 2002 (DAO). Cody (1996) knew this rare species in the Territory from only three locations, two from near the western end of the Alaska Highway and one from adjacent to the Takhini River west of Whitehorse. The specimen cited above is an extension of the known range of about 225 kilometers to the north of the sites near the Alaska border. Only four plants were observed. ORCHIDACEAE Spiranthes romanzoffiana Cham. & Schlecht., Hooded Ladies’-tresses — YUKON: tussocks near bog, east of Haines Highway, 60°08.570'N 136°58.469'W, P. Cas- well 453, 13 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory of about 120 kilometers southwest of Whitehorse. SALICACEAE Salix alaxensis (Anderss.) Cov. ssp. longistylis (Rydb.) Hultén — YUKON: beside recreation centre in middle of town, Beaver Creek, 62°22'56"N 140°52'41"W, B. Bennett 01-102, 12 June 2001 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 140 kilometers northwest of a site near the north end of Kluane Lake. Salix glauca L. var. acutifolia (Hook.) C. Schneider — YUKON: Mt. Merrill, 60°06'N 124°45'W, G. Brunner, 1994 (DAO). The specimen cited above is about 90 kilometers east of the easternmost site of Salix glauca sl. mapped by Cody (1996). Salix pseudomyrsinites (Anderss.) Ball ex Hultén (S. novae-angliae Anderss.) — YUKON: along Bennett Lake just above high water mark, Carcross Dunes, 60°10'30"N 134°43'26"W, B. Bennett & G. Argus 99- 548, 29 Aug. 1999 (DAO) (determined by G. Argus). The nearest site of this species known to Cody (1996) is in the vicinity of Whitehorse, about 60 kilometers to the north- west. Salix rotundifolia Trauty. ssp. dodgeana (Rydb.) Argus — YUKON: Dryas alaskensis tundra, Mount Casca border Monument 97, 65°21'25"N 141°00'00"W, B. Bennett & M. B. Cook 02-519, 27 Jane 2002 (DAO) (determined by G. Argus). The specimen cited above from adjacent to the Alaska border is from a site about 70 kilometers northwest of the nearest site known to Cody (1996). URTICACEAE Urtica dioica L. ssp. gracilis (Ait.) Selander, Sting- ing Nettle — YUKON: dwelling mound, Forty Mile Historic Site at the confluence of Forty Mile and 568 Yukon rivers, 64°25'N 140°30'W, C. E. Kennedy 44, July 2000 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 50 kilometers northwest of the vicinity of Dawson City. POLYGONACEAE Polygonum lapathifolium L., Willow Weed, Pale Smart- wood — YUKON: wetland Carex fen, uncommon silty mud beside beaver pond, 60°07'00"N 124°15'2"W, B. Bennett 98-609, 16 June 1998 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). The specimen cited above is an extension of the known range in the Territory of about 700 kilometers southeast of sites in the vicinity of Carmacks. Rheum rhabarbarum L. (R. rhaponticum L.) Rhubarb — YUKON: adjacent to garage near machine shop, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Ken- nedy 37799, 17 July 2002 (DAO). Cody et al. (2001) and (2002) reported Rhubarb growing wild at Silver City and Destruction Bay. The specimen cited above is the third known cited where it was growing wild. Rumex crispus L., Curled Dock — YUKON: moist shore of small lake between Ross River and Campbell Highway, 61°58'7"N 132°38'34"W, Cody & Cody 37961, 24 July 2002 (DAO). The specimen cited above is the sixth site in the Territory for this introduced species. Cody (1996) knew it from the vicinity of Dawson City and it has since been reported from just west of Whitehorse, Haines Junction and adjacent to the Alaska Highway near the Alaska border (Cody et al. 2002). CHENOPODIACEAE Atriplex alaskensis S. Watson, (A. patula sensu Cody (1996), A. patula var. alaskensis (S. Wats.) Welsh), Alaskan Orache (Figure 3) — YUKON: alkaline flat, Takhini Salt Flats, 60°51.2'N 135°43.23'W, Bennett & Parker 98-431, 2 Sept. 1998 (B. Bennett Herbarium, photo DAO) (determined by P. W. Ball); alkaline flats, between the salt flat and the highway in the area of the fence, Takhini Salt Flats, 60°51'23"N 135°42'55"W, B. Bennett 01-168, 23 Aug. 2001 (B. Bennett Herb- arium, photo DAO); occasional in wet alkaline margin of a brackish slough, Mile 484.5 Alaska Highway west of Whitehorse, R. L. Taylor 4094, 14 July 1959 (DAO); common around clay margins of dried up alkaline pond, east of Haines Junction at Mile 985 Alaska Highway, 60°48'N 136°45'W, Calder & Kukkonen 25252, 14 Aug. 1960 (DAO); common around clayey margin of alkaline lake, between Minto and Pelly Crossing on Dawson-Whitehorse road, approximately 62°46'N 136°36'W, Calder & Kukkonen 28095, 10 Aug. 1960 (DAO). Cody (1996) considered the Calder & Cody specimens as introductions from Europe (A. patula). Atriplex alaskensis should now be added to the Yukon Flora and the list of rare plants of the Territory (Douglas et al. 1981). The Taylor spe- cimen cited above was the basis for the drawing published in the Yukon Flora as A. patula. THE CANADIAN FIELD-NATURALIST Vol. 118 Atriplex subspicata (Nutt.) Rydberg — YUKON: grassy area in open alkaline salt ponds, Mayo Road, Dilla- bough’s Grazing Lease, 60°55'54"N 135°10'14"W, B. Bennett 02-649, 19 Aug. 2002 (DAO). Cody (1996) knew this species from a single locality adja- cent to a saline pond just west of Whitehorse. The specimen cited above from just north of Whitehorse is only the second known from the Territory. Chenopodium capitatum (L.) Asch., Strawberry-blite — YUKON: disturbed gravelly soil, roadstop, Atlin Road just north of BC border, 60°00'09"N 133°47'42.1"W, Cody & Cody 37546, 28 July 2001 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 60 kilometers to the southwest from a site in the vicinity of Johnson’s Crossing. Corispermum ochotense Ignatov var. alaskanum Mosyakin (C. hyssopifolium sensu Cody 1996) — YUKON: river bar in loose aluvial sand between Salix exigua and river with Tanacetum bipinnatum, Poten- tilla anserina and Aster falcatus, Yukon River at con- fluence of Nester Creek, 64°38'22"N 140°52'52"W, Bennett & Mulder 02-665, 25 Aug. 2002 (DAO); river bar in loose alluvial sand between Salix exigua and river with Tanacetum bipinnatum, Potentilla anserina and Aster falcatus, Yukon River, 64°22'19"N 140°26'49"W, Bennett & Mulder 02-740, 24 Aug. 2002 (DAO). The specimens cited above are the first yet found in the Territory. PORTULACACEAE Claytonia megarrhiza (A. Gray) Parry, Alpine Spring- beauty (Figure 3) — YUKON: alpine tundra, three plants seen at base of a long scree slope near a very small stream, northwest side of Wade Mountain, Kluane National Park, 61°18'N 139°33'W, P. Caswell 236, 28 June 2002 (DAO). This species, which is new to the Flora of the Yukon Terri- tory (Cody 1996), should be added to the list of rare plants of the Territory (Douglas et al. 1981). Porsild and Cody (1980) knew it from three sites in the Mackenzie Mountains in the Continental Northwest Territories and McJannet et al. (1995) knew it from four sites in that area in the Rare Vascular Plants in the Northwest Territories. To the south Douglas et al. (2002) mapped three sites in the extreme southeast of British Columbia where it was considered rare. Montia fontana L., Blinks — YUKON: aquatic, growing in the shallows (10-30 cm) in a backwater slough, Lewes Marsh, 60°34'N 134°35'W, B. Bennett OO- 1091, 25 June 2000 (DAO) (determined by C. Parker and W. J. Cody). Cody (1996) knew this rare plant in the Territory only from a single locality on the Arctic Coast where it was collected by Erling Porsild in 1934 (CAN, photo DAO). In addition a second specimen was collected by Cody on Herschel Island in 1999 (Cody et al. 2001). This is a circumpolar, low-arctic species, which was considered rare in the Territory by Douglas et al. (1981). To the east of the location cited above it is known from the southern Mackenzie Mountains in the 2004 FiGure 3. Claytonia megarrhiza, Alpine Springbeauty (drawn by Lynne Bartosch). former District of Mackenzie, and to the west in essentially coastal regions of Alaska and British Columbia. CARYOPHYLLACEAE Cerastium arvense L., Field Chickweed — YUKON: wet roadside ditch, Top of the World Highway Km 100, 64°05'53"N 140°54'43"W, Cody & Cody 38011, 26 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers west of the vicinity of Dawson City. Minuartia dawsonensis (Britt.) House — YUKON: steep slope by Alaska Highway, 8 km west of White River, 62°3'15.6"N 140°38'20.5"W, Cody & Cody 37880, 20 July 2002. The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 125 kilometers northwest of a site adjacent to Kluane Lake. Minuartia elegans (Cham. & Schlecht.) Schischk. — YUKON: Dryas alaskensis heath, 30° southwest-facing large talus dolomite boulders, summit of mountain on Canadian side of Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett & M. B. Cook 02-543, 27 June 2002 (B. Bennett Herbarium, photo DAO). The specimen cited above from adjacent to the Alaska bor- der is an extension of the known range of about 75 kilometers northwest of the nearest site in the Ogilvie Mountains. Minuartia macrocarpa (Pursh) Ostenf. — YUKON: al- pine tundra, spotty occurrence in raised rocky mounds, mountain between Kusawa and JoJo Lake, 60°35'47"N 136°15'19"W, B. Bennett 97-681, 19 Sept. 1997 (DAO). The specimen cited above is an extension of the known range in the southwest of the Territory of about 80 kilo- meters southeast of a site northwest of Haines Junction. Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 569 Minuartia yukonensis Hultén — YUKON: coarse gravel and small rocks of a stream’s outwash, Upper Joe Creek, Kluane National Park, 61°12.861'N 139°03.863'W, P. Caswell 585, 24 July 2002 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 225 kilometers west of a site north of Whitehorse. Silene involucrata (Cham. & Schlecht.) Bocquet ssp. tenella (Tolm.) Bocquet — YUKON: meadow between Stone House and river near Fort Selkirk sign, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Ken- nedy 37805, 17 July 2002 (DAO). The specimen cited above is from a site between sites in the vicinity of Mayo and south of latitude 62°. Silene williamsii Britton — YUKON: talus lower slope of southwest-facing bluff, Yukon River, Minto Bluff, 62°37'06"N 136°57'41"W, B. Bennett 02-678, 9 Aug. 2002 (B. Bennett Herbarium, photo DAO); common on base of slope across from Brittania Creek growing with Artemisia frigida and Calamagrostis purpuras- cens, Yukon River, 62°52'28"N 138°42'56"W, Bennett 02-443, 10 Aug. 2002 (B. Bennett Herbarium, photo DAO). This species was considered rare in the Territory by Douglas et al. (1981) on the basis of 3 collections from south and southwest of Dawson City. Silene vulgaris (Moench) Garcke, Bladder Campion — YUKON: gravelly soil in old field with grass and sweet clover, west of Mayo airport, 63°37'01.4"N 135°58'20.9"W, Cody & Cody 37665, 4 Aug. 2001 (DAO). The specimen cited above of this introduced species in the Territory (Cody 1996) was previously known only from an area just east of Dawson. Stellaria umbellata Turcz. — YUKON: wet scree cushion in zinc moss, Wade Mountain, Kluane National Park, 61°18.450'N 139°30.921'W, P. Caswell 354, 7 July 2002 (DAO). Douglas et al. (1981) considered this species as rare in the Territory. Cody (1996) knew it from only three widely sepa- rated sites. The specimen cited above is from a site about 60 kilometers northwest of a previously known site in Kluane National Park. NYMPHAEACEAE Nuphar polysepalum Engelm., Yellow Pond-lily — YUKON: shallow water along shore of lake, Tatchun Lake Campground, 62°17'52.8"N 136°08'11.4"W, Cody & Cody 37626, 30 July 2001 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 120 kilometers to the north, south of Mayo. RANUNCULACEAE Pulsatilla ludoviciana (Nutt.) Heller, Prairie-crocus — YUKON: steep grassy slope on ridge northeast of Wood- burn Creek, Tintina Trench, Ddhaw Ghro, 63°08'N 136°05'W, C. E. Kennedy 4, 27 July 2001 (DAO). 570 The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers east northeast of a site north of the junction of the Pelly and Yukon rivers. Ranunculus aquatilis L. var. subrigidus (W. B. Drew) Breitung — YUKON: near South Fork intake, Klondike River on edge of beaver dam in old ditch, 64°0'30"N 138°12'00"W, G. Brunner 549, 22 July 2002 (DAO). The nearest site known to Cody (1996) of this uncommon variety was about 60 kilometers to the north adjacent to the Dempster Highway. Ranunculus occidentalis Nutt. var. brevistylis Greene, Western Buttercup — YUKON: alpine tundra, valley above treeline, Macmillan Pass, 63°15'N 130°02'W, J. Basinger s.n., June 1982 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is the northernmost yet found in the Territory and is an extension of the known range of about 250 kilometers northeast of a site adjacent to the northern South Canol Road. PAPAVERACEAE Papaver croceum Ledeb. — YUKON: by garbage along trail in Picea glauca, Pinus, Populus tremuloides wood- land, Whitehorse, near Hidden Lake, 60°35'00.5"N 135°51'00.2"W, Cody & Cody 37678A, 9 Aug. 2001 (DAO); in gravel beside car parking spot, 11 Chalet Crescent, Whitehorse, 60°55'54.7"N 135°10'26.5"W, W. J. Cody 37711, 12 July 2002 (DAO) (determined by H. Solstad). This species is a garden escape which has not previously been recorded from the Whitehorse area. In addition Heidi Solstad revised three collections previously determined as Papaver nudicaule (Dawson, Calder & Billard 3121 (DAO), Halfway Lakes area 15 miles north of Mayo, Calder & Gil- lett 4176 (DAO) and Tower Hill, Tagish, R. Rosie 798 (DAO)) to P. croceum. P. nudicaule ssp. nudicaule should be replaced by P. croceum in the flora (Cody 1996, 2000). BRASSICACEAE (CRUCIFERAE) Alyssum obovatum (C. A. Meyer) Turez. (A. ameri- canum Greene) — YUKON: 45° talus southwest-facing slope, Yukon River, | km east of Alaska/Yukon border, 64°41'05"N 140°57'30"W, Bennett & Mulder 02-666, 25 Aug. 2002 (B. Bennett Herbarium, photo DAO) (determined by G. A. Mulligan). Douglas et al. (1981) considered this species too wide- spread to include it in The Rare Vascular Plants of the Yukon. Cody (1996) knew it from only eleven sites north of latitude 62°N. The specimen cited above is a slight extension of the known range in the Territory about 50 kilometers to the northwest of another site adjacent to the Yukon River. Arabis caucasica Willd., Wall Rock-Cress (Figure 4) — YUKON: garden escape, Alsek Road, Hayes Property, 10 km NW of Haines Junction, 64°47'N 137°41'W, B. Bennett 01-041, 12 June 2001 (DAO) (determined by G. A. Mulligan). Submitted to B. Bennett by Carolyn Hayes. This garden escape has not previously been observed in the Yukon Territory. THE CANADIAN FIELD-NATURALIST Vol. 118 re a Ze AL Figure 4. Arabis caucasica (drawn by Lee Mennell). Arabis caucasica can be separated from A. codyi as follows: A. Garden escape; cauline and caudex leaves with similar dentate to subdentate margins, flowering stems spreading to ascending; petals!/6:Simim) longi ee ee ae A. caucasica B. Native species; cauline and caudex leaves not similarly dentate to subdentate; flowering stems erect; petals more than 10 mm long............. A. codyi Arabis holboellii Hornem. var. retrofracta (Graham) Rydb. — YUKON: gravel beside road and steep slope, Alaska Highway Km 1918, 62°16'26.1"N 140°45'6.7"W, Cody & Cody 37855, 20 July 2002 (DAO). The nearest site adjacent to the Alaska Highway known to Cody (1996) is about 150 kilometers to the southeast in the vicinity of Kluane Lake. There is, however, a site adjacent to a river about 60 kilometers to the east. Arabis holboellii Hornem. var. secunda (Howell) Jep- son — YUKON: adjacent to narrow road through woods leading down to Nares Lake southeast of Carcross, 60°9'20"N_ 134°40'10"W, Cody & Cody 37934, 23 July 2002 (DAO) (determined G. A. Mulligan). Cody (1996) knew this taxon from only four localities in the Territory, west of longitude 135°. Cody (2001) added new sites in the vicinity of Whitehorse and Wolf Lake to the east. The specimen cited above is the southernmost yet found in the Territory. , { ' 2004 Armoracia rusticana (Lam.) Gaert., Mess. & Scherb. — YUKON: Hultén (1941-50) reported this species “A specimen of this plant was collected at Stewart R. July 1898 by Anders- son (S). It must have been cultivated or have escaped from cultivation". Hultén (1968) mapped the worldwide distribution of this species with a single dot in the Territory and Welsh (1974) sub Rorippa armorocia (L.) A. S. Hitche. stated “Cul- tivated for the root; in southern Alaska and Yukon, persisting: introduced from Europe". Fortunately this species was over- looked by Cody when preparing the Flora of the Yukon Territory (1996); since the Anderson specimen was recently borrowed from the Swedish Museum of Natural History, Stockholm and was revised by Gerald A. Mulligan to Rorippa barbareifolia (DC.) Kitagawa, an amphi-Berigian, nonarctic species which extends across Alaska to central Yukon Territory, the Porcupine River valley, the central Rich- ardson Mountains and the Fort McPherson area south of the Mackenzie Delta. Brassica rapa L., Bird rape — YUKON: roadside gravel, Alaska Highway, near Kluane Wilderness Camp Km 1791, B. Bennett 01-095, 24 July 2001 (DAO). The specimen cited above of this introduced species is an extension of the known range in the Territory of about 175 kilometers northwest of a site adjacent to the Haines High- way. It may have resulted from roadside seeding. Camelina sativa (L.) Cranz, Falseflax (Figure 5) — YUKON: disturbed site, Whitehorse Shipyards, 60°43'34"N 135°03'12"W, B. Bennett 01-150, 28 Aug. RNA tS a Men AP, SS Yee “has ls yy AY .\ Za FiGure 5. Camelina sativa, Falseflax (drawn by Lee Mennell). Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI S74 2001 (B. Bennett Herbarium, photo DAO) (determined by G. A. Mulligan). The specimen cited above is new to the Flora of the Yukon Territory (Cody 1996). It is a Eurasian introduction which is known in Canada from New Brunswick to British Columbia and north into southern District of Mackenzie. The genus Camelina can be separated from the genus Alyssum as follows: A. — Styles usually 2-3 mm long; silicles oval-elliptic to egg-shaped, usually at least 5 mm long; seeds numer- ous; petals often shallowly bilobed; annual Camelina B. Styles not over 1 mm long; silicles oval in outline, less than 5 mm long; petals notbilobed:perennialy ys ee eae ere Alyssum Capsella bursa-pastoris (L.) Medic., Shepherd’s-purse — YUKON: level gravel area, Campbell Highway Km 380, 62°02'34.2"N 132°52'02.2"W, Cody & Cody 37591, 30 July 2001 (DAO); growing beside Artemi- sia tilesii by narrow road through woodland, Pelly Farm Road, 62°51'22.2"N 137°00'34"W, Cody & Kennedy 37821B, 17 July 2002 (DAO); gravel, Pick- handle Lake, 63°02'22.2"N 138°23'15.8"W, Cody & Cody 37834, 20 July 2002 (DAO). The first specimen cited above of this introduced species is from a site between two sites adjacent to the Campbell Highway (Cody 1996) in the vicinity of Carmacks. One of sites is 175 kilometers to the west and the other is about 185 kilometers to the southeast. The second and third specimens are from sites between Keno and Carmacks (Cody 1996). Cardamine bellidifolia L., Alpine Bittercress — YUKON: Herschel Island, 69°35'N 139°O5S'W, C. E. Kennedy s.n., 20 July 1985 (DAO). The nearest sites known to Cody (1996) were from the north coast about 30 kilometers to the west and near the Firth River about 30 kilometers to the southwest. Descurainia incisa (Engelm. ex A. Gray) Britton var. incisa, Tansy Mustard — YUKON: gravel roadside, rare, Dempster Highway Km 98, 64°42'N 138°24'46"W, Cody & Cody 38056, 27 July 2002 (DAO) (deter- mined by G. A. Mulligan). Cody et al. (2001) reported this species as new to the Territory and Cody et al. (2002) added additional sites adja- cent to the Alaska Highway north of latitude 62°N. The specimen cited above is from the first known site adjacent to the Dempster Highway. Draba aurea M. Vahl, Golden Draba — YUKON: Site KP 110, 62°15\09-6°N) 137°41:03) W. (Gs Brunner 591-01, 8 Aug 2001 (Environment Yukon Herbariam, photo DAO) (determined by G. A. Mulligan). The specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is an extension of the known range of about 50 kilometers from a site south of latitude 62°N. Draba corymbosa R. Br. — YUKON: Herschel Island, 69°35'N 139°05'W, C. E. Kennedy s.n., 10 Aug. 1985 (Environment Yukon Herbarium, photo DAO) (deter- mined by G. A. Mulligan). S72 The nearest site known to Cody (1996) was from near the Firth River about 30 kilometers to the southwest. Draba crassifolia Graham — YUKON: sandy moist soil in eroded area, ridge south of Donjek Glacier, 61°08.277'N 139°31.170'W, P. Caswell 403, 8 July 2002 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range in the Territory (Cody 1996) northwest of sites in Kluane National Park. Draba glabella Pursh — YUKON: steep south-facing bluffs along Klondike River, 1 mile north of 64°02'00"N_ 137°41'30"W, G. Brunner 435, 2 June 2001 (DAO) (determined by G. A. Mulligan). The specimen cited above is from a site about 70 kilo- meters east of a site in the vicinity of Dawson City (Cody 1996). Draba nemorosa L., Wood Whitlow-grass — YUKON: disturbed farm area, Black Sheep airline base, Mayo, 63°35.26'N 135°51.83'W, B. Bennett 00-1193, 2 July 2000 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 110 kilometers northeast of a site west of longitude 137°W and north of the Pelly River. Draba ogilviensis Hultén — YUKON: summit of small peak south of Donjek Glacier, Kluane National Park, 61°08.318'N 139°31.132'W, P. Caswell 392, 8 July 2002 (DAO) (determined by G. A. Mulligan). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is the westernmost yet known in Kluane National Park. Draba palanderiana Kjellm. — YUKON: dry alpine Dryas heath with Dryas alaskensis, Silene acaulis, Synthyris borealis, Podistera macounii and lichens, summit of mountain, calcareous dolomite talus, 65°21.37'N 140°59.34'W, B. Bennett 02-523, 27 June 2002 (DAO). The specimen cited above from adjacent to the Alaska border is an extension of the known range in the Territory of about 60 kilometers to the north of a site mapped by Cody (1996). Draba porsildii G. A. Mulligan — YUKON: wet scree, Wade Mountain, Kluane National Park, 61°18'N 139°33'W, P. Caswell 240, 2 June 2002 (DAO) (deter- mined by G. A. Mulligan). Douglas et al. (1981) considered this species rare in the Territory on the basis of three specimens from Kluane Nation- al Park and one from the South Canol Road southwest of Ross River. Cody (2003) cited a specimen from west of the three sites in Kluane National Park. The specimen cited above is from an additional site in the Park about 70 kilometers north of the last site. Draba stenoloba Ledeb. — YUKON: Donjek Valley, 61°25.689'N 139°53.074'W, R. Maraj s.n., 29 June 2002 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 90 kilometers northwest of a site in Kluane National Park. THE CANADIAN FIELD-NATURALIST Vol. 118 Lepidium densiflorum Schrad. var. densiflorum, Com- mon Pepper-grass — YUKON: roadstop in disturbed gravelly soil, Atlin Road just north of BC border, 60°00'09"N_ 133°47'42.1"W, Cody & Cody 37543, 37551, 28 July 2001 (DAO) (determined by G. A. Mulligan). The specimens cited above are an extension of the known range in the Territory (Cody 1996) of about 125 kilometers southeast of a site just west of Whitehorse. Lepidium densiflorum Schrad. var. macrocarpum G. A. Mulligan — YUKON: adjacent to Catholic Church, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37808A, 17 July 2002 (DAO) (determined by G. A. Mulligan). Cody et al. (2001) reported this variety, which is probably introduced, new to the Territory from the vicinities of Daw- son, Ross River and Pelly Crossing. The specimen cited above is an extension of the known range in the Territory of about 50 kilometers to the west of Pelly Crossing. Parrya arctica R. Br. — Two specimens, one from Pauline Cove, Herschel Island (Cody 36030) and the other from the Bonnet Plume River (Loewen 99-25-93) were recently revised to P. nudicaulis by G. A. Mulligan. These two species can be separated as follows: A. Stamens linear, 1.5 to IEPfoyistVool) KON reteieicn acc bane jaa P. nudicaulis B. Stamens oblong, less than 1h Oimmvlon es kane een OT ate ee P. arctica In addition, Cody (1994) reported P. arctica from Herschel Island on the basis of a specimen collected by P. F. Cooper (433), 9 May 1979 (CAN) as new to the Territory and sug- gested that it should be added to the list of rare plants (Doug- las et al. 1981). This specimen has now been revised to Thlas- piarcticum Porsild by G. A. Mulligan and should be deleted from the flora of Herschel Island and the Yukon Territory. Rorippa barbareifolia (DC.) Kitagawa — YUKON: Pelly Crossing, 62°51'09"N 136°56'05.2"W, Cody & Ken- nedy 37831, 17 July 2002 (DAO) (determined by G. A. Mulligan). This species was considered rare in the Territory by Doug- las et al. (1981). The specimen cited above is the southern- most yet found adjacent to the Alaska Highway. Sinapis arvensis L., Charlock — YUKON: in a well-tend- ed flower bed, Haines Junction, P. Caswell 311, 5 July 2002 (DAO) (determined by G. A. Mulligan). The specimen cited above which was the only plant ob- served in Haines Junction of this introduced species was from only the third site known to Cody (1996) in the Territory. Subularia aquatica L. ssp. americana Mulligan & Calder, Awlwort — YUKON: shallows of large pond east of Alaska Highway north of Sulphur Lake, 61°00.435'N 138°12.729'W, P. Caswell 772, 9 Aug. 2002 (DAO) (determined by G. A. Mulligan). This species was considered rare in the Territory by Doug- las et al. (1981). The specimen cited above is an extension of the known range of about 175 kilometers east of a site in the vicinity of Whitehorse (Cody 1996). 2004 Thlaspi arvense L., Penny Cress — in black muck just inside gate, flats 7 miles east of Tagish Bridge, 60°19'S"N 134°10'17"W, Cody & Cody 37958, 23 July 2002 (DAO) (determined by G. A. Mulligan). The specimen cited above is an extension of the known range of this introduced species of about 80 kilometers south- east of the vicinity of Whitehorse. SAXIFRAGACEAE Ribes oxyacanthoides L. ssp. oxyacanthoides — YUKON: meadow adjacent to RCMP foundation, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37757, 16 July 2002 (DAO). Although quite frequent in the Territory south of latitude 62°N, Cody (1996) knew this shrub from only two sites to the north. Saxifraga caespitosa L.— YUKON: 40° slope talus dolo- mite boulders, Mount Casca border Monument 97, 65°21'25"N 141°00'00"W, B. Bennett & M. Cook 02- SISA, 27 June 2002 (DAO). Cody (1996) mapped this species in the Territory from three distinct areas (between latitudes 60° and 62°, between latitudes 64° and 66°, and north of latutude 68°. The specimen cited above from adjacent to the Yukon-Alaska border is about 70 kilometers from the sites east and southeast. Saxifraga eschscholtzii Sternb. — YUKON: dry alpine heath at summit of mountain on Canadian side of Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett & M. Cook 02-512, 27 June 2002 (DAO). Cody (1996) knew this species only from the British Mountains in the extreme northwest of the Territory. The specimen cited above is an extension of the known range in the Territory of about 460 kilometers to the south. Saxifraga foliolosa R. Br. — YUKON: damp sand of small sandy depression in sedge tundra, ridge south of Donjek Glacier, Kluane National Park, 61°07.383'N 139°31.308'W, P. Caswell 416, 8 July 2002 (DAO). Douglas et al. (1981) considered this species which was known to Cody (1996) from north of latitude 67°N and a single site on the west side of Haines Highway southeast of Haines Junction as rare in the Territory. Cody et al. (2000) reported another site in the south from about 80 kilometers northeast of the site adjacent to the Haines Highway. The specimen cited above is an extension of the range in the south of about 135 kilometers northwest of the site near the Haines Highway. Saxifraga nelsoniana D. Don ssp. pacifica (Hultén) — YUKON: wet organic soil with some fine scree at edge of stream, Tabletop Mountain, Kluane National Park, 61°15.231'N 139°11.115'W, P. Caswell 285, 29 June 2002 (DAO). Douglas et al. (1981) considered this subspecies rare in the Territory. The specimen cited above is an extension of the known range in the Park of about 25 kilometers west of a site west of Kluane Lake. Saxifraga rufopilosa (Hultén) Porsild — YUKON: moist places in 30° southwest facing large talus dolomite boulders, summit of mountain on Canadian side of Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett & M. B. Cook 02-528, 27 June 2002 (DAO). Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI SME: The specimen cited above is from a site about 35 kilometers northwest of the northernmost site south of latitude 66°N known to Cody (1996). Saxifraga tricuspidata Rottb., Prickly Saxifrage — YUKON: steep grassy slope on ridge northeast of Wood- burn Creek, Tintina Trench, Ddhaw, 63°08'N 136°05'W, C. E. Kennedy 7, 27 July 2001 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 100 kilometers east northeast of a site at about longitude 137°W, north of the Pelly River. ROSACEAE Amelanchier alnifolia (Nutt.) Nutt., Saskatoon — YUKON: southwest-facing slope across from Brittania Creek, Yukon River, 62°52'28"N 138°42'56"W, B. Bennett 02-653, 10 Aug. 2002 (DAO). Cody (1996) knew only three sites of this species north of the site reported above: Dawson City, Mayo, and adjacent to the Stewart River. Chamaerhodos erecta (L.) Bge. ssp. nuttallii (Picker- ing ex Rydb.) Hultén — YUKON: common at base of southwest-facing slope, across from Brittania Creek, Yukon River, 62°52'28"N 138°42'56"W, B. Bennett 02-640, 10 Aug. 2002 (DAO). Cody (1996) knew this species in the Territory north to near latitude 64°. The specimen cited above is an extension of the known range in the Territory of about 80 kilometers west of a site adjacent to the Pelly River. Dryas hookeriana Juz. — YUKON: Ogilvie Mountains, Tombstone Range, Yakamaw Creek, watershed east of Angelcomb Peak, 64°36'N 138°14'W, Cody 36808, 36845, 20-22 July 1999 (DAO); Dryas alaskensis tundra, Mount Casca border Monument, 65°21'25"N 141°00'00"W, B. Bennett & M. B. Cook 02-518, 27 June 2002 (B. Bennett Herbarium, photo DAO). The specimens cited above are the first yet known to Cody (1996) from west of the Dempster Highway between latitudes 64°N and 66°N. Geum aleppicum Jacq. ssp. strictum (Ait.) Clausen, Yellow Avens — YUKON: Top of the World Golf Course, 64°3'38"N 139°26'13"W, Cody & Cody 37998, 25 July 2002 (DAO). Douglas et al. (1981) considered this taxon rare in the Ter- ritory. The specimen cited above is an extension of the known range in the Territory of about 200 kilometers west of a site west of Keno City (Cody et al. 2001) and north of sites adja- cent to the Alaska Highway near the Alaska border. Geum rossii (R. Br.) Scr. — YUKON: moist places at sum- mit of the Canadian side of Mount Casca, 65°21'37"N 140°59'34"W, B. Bennett 02-524, 27 June 2002 (B. Bennett Herbarium, photo DAO). The specimen cited above is from a site about 100 kilo- meters northwest of the nearest location in the Ogilvie Moun- tains where it has been collected frequently. Potentilla arguta Pursh ssp. convallaria (Rydb.) Keck, Tall Cinquefoil — YUKON: dry rocky area, South Fork intake, Klondike River, 64°0'30"N 138°12'00"W, G. Brunner 551, 22 July 2002 (DAO). 574 The specimen cited above is only the second collected from north of latitude 64°N in the Territory (Cody 1996). It is from a site about 60 kilometers east of the vicinity of Dawson. Rosa woodsii Lindl., Western Rose — YUKON: Artemi- sia frigida slope, east side of lake, Whitehorse, Hid- den Lakes, 60°41'30"N 135°02'45"W, B. Bennett O1- 118, 11 July 2001 (B. Bennett Herbarium, photo DAO); grassy alkaline slope north of highway on slope, Alaska Highway Km 1588, | km east of Cracker Creek, 60°48'20"N 136°48'00"W, B. Bennett 02-735, 11 Sept. 2002 (DAO); common at base of slope, Yukon River, Brittania Creek, 62°52'28"N 138°42'56"W, B. Bennett 02-639, 10 Aug. 2002 (B. Bennett Herbarium, photo DAO). This is a rare species in the Territory (Douglas et al. 1981). The first specimen cited above is only the fourth known in the Territory (Cody 1996). The second is an extension of the known range in the Territory (Cody 1996) of about 50 kilo- meters west of a site adjacent to the Alaska Highway. The third specimen is an extension of about 125 kilometers north- west of a site reported by Cody et al. (2003) from the vic- inity of Carmacks. FABACEAE (LEGUMINOSAE) Lupinus nootkatensis Donn — YUKON: clearing bet- ween road and poplar forest, west side of Haines High- way, north of cut-off to Dalton Post, 60°09.405'N 136°58.572'W, P. Caswell 440, 13 July 2002 (DAO). The specimen cited above is only the third known from southeastern Kluane National Park and the Yukon Territory. Oxytropis arctica R. Br., Arctic Oxytrope — YUKON: at base of talus slope, Lower Kathleen Lake, along south shore ca. 25 km SE of Haines Junction, 60°33'N 137°16'W, G. W. & G. G. Douglas 8391, 25 June 1975 (Kluane National Park, photo DAO); in alpine fell field, Dezadeash River Valley ca. 16 km WSW of Haines Junction, G.W. & G.G. Douglas 8405, 30 June 1975 (Kluane National Park Herbarium, photo DAO). Cody (1996) suggested that this species should be looked for in the mountains of northern Yukon Territory because it is known in the Canadian Arctic and northern Alaska. Douglas et al. (1999) included the Territory in the distribution of O. arctica presumably on the basis of the specimens cited above and also reported the only collection known in British Colum- bia from Mile 416 Alaska Highway which is about 700 kilo- meters east southeast of the Yukon sites. It should be added to the list of rare species in the Territory. Oxytropis campestris (L.) DC. ssp. varians (Rydb.) Cody, Field Locoweed — YUKON: steep grassy slope on ridge northeast of Woodburn Creek, Tintina Trench, Ddhaw Ghro, 63°08'N 136°05'W, C. E. Kennedy 6, 27 July 2001 (DAO). The specimen cited above is an extension of the known range in the Territory (Cody 1996) of about 110 kilometers northeast of a site adjacent to the Yukon River. Trifolium pratense L., Red Clover — YUKON: road- side, junction of Campbell Highway and Frenchman Lake Road, 62°4'30"N 135°30'55"W, Cody & Cody 37985, 24 July 2002 (DAO). THE CANADIAN FIELD-NATURALIST Vol. 118 The nearest site of this introduced species in the Territory (Cody 1996) is from the vicinity of Whitehorse, about 150 kilometers to the south. Vicia americana Muhl., American Vetch — YUKON: bor- der of grassland and Populus, Macoobs Park, Mayo, 63°35'42.5"N 135°56'57.1"W, Cody & Cody 37668, 5 Aug. 2001 (DAO); west end of parking lot by visitor kiosk, Rancheria, 60°05'16"N 130°36'10"W, B. Ben- nett 01-059, 26 July 2001 (DAO). Douglas et al. (1981) considered this species rare in the Territory on the basis of a single collection from the Larsen Creek hotsprings in the extreme southeast (Scotter & Cody 1979). Additional collections were added from this area by Cody et al. (1998) and from the vicinity of Watson Lake (Cody et al. 2000). The first specimen cited above is an extension of the known range in the Territory of about 550 kilometers to the northwest of Watson Lake. The second collection is an extension of the known distribution of about 120 kilometers west of Watson Lake but was probably introduced at that site. CALLITRICHACEAE Callitriche anceps Fern. — YUKON: on mud substrate at edge of water, old beaver pond east of Haines Highway about 10 kilometers south of Kathleen Lake cutoff, 60°30.838'N 137°04.970'W, P. Caswell 787, 11 Aug. 2002 (DAO). This species was considered rare in the Territory (Douglas et al. 1981). The specimen cited above is from a site about 30 kilometers northwest of a site in Kluane National Park. VIOLACEAE Viola renifolia Gray var. brainerdii (Greene) Fern., Kidney-leaved Violet - YUKON: humus covered with old poplar leaves, poplar and white spruce wood flat, Wade Lakes Road, between Kluane National Park and Haines Highway, 60°07.119'N 137°05.369'W, P. Cas- well 020, 6 June 2002 (DAO). Cody (1996) knew this species as far north as the vicinity of Dawson City. The specimen cited above is an extension of the known range in the Territory of about 150 kilometers southwest of the vicinity of Whitehorse. HALORAGACEAE Myriophyllum verticillatum L. — YUKON: in water be- side dock, Nunatuk Camp, 62°9'49"N 135°48'4"W, Cody & Cody 37989, 24 July 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is from a site about half way between the Pelly River and the north end of Lake Le- barge. APIACEAE Cicuta maculata L. var. angustifolia Hook., Spotted Water-hemlock — YUKON: lush herbaceous meadow on riverbank, Kirkman Creek, Yukon River, 62°59'21"N 139°23'07"W, B. Bennett 02-284, 10 Aug. 2002 (DAO). Douglas et al. (1981) considered this taxon rare in the Territory. The specimen cited above is an extension of the known range of about 175 kilometers to the northwest of Carcross (Cody 1996). Podistera macounii (Coult. & Rose) Mathias & Const. — YUKON: Dryas alaskensis tundra, Mount Casca bor- 2004 der Monument 97, 65°21'25"N 141°00'00"W, Bennett & Cook 02-520, 27 June 2002 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). Cody (1996) knew it only from the Richard- _ son Mountains area in the northeast. Cody et al. (1998) ex- tended the known range in the Territory south into the Daw- son range at latitude 62°26'N. The specimen cited above is an extension of the known range in the Territory of about 275 kilometers to the west from the southernmost site in the Richardson Mountains. PYROLACEAE Pyrola minor L. — YUKON: adjacent to the Catholic Church, Fort Selkirk, 62°46'34.4"N_ 137°23'35.9"W, Cody & Kennedy 37815, 17 July 2002 (DAO). Although quite frequent in the Territory east and north of the Klondike Highway, Cody (1996) also knew this species _ from the vicinity of southern Kluane Lake, to the west. _ ERICACEAE Cassiope tetragona (L.) D. Don ssp. saximontana (Small) Porsild — YUKON: dry alpine Dryas heath, summit of Mount Casca, 65°21'37"N 140°59'34"W, Bennett & Cook 02-522, 27 June 2002 (DAO). The specimen cited above is the northernmost yet found in _ the Territory. The nearest site to the above north of Chapman Lake in the Ogilvie Mountains is about 125 kilometers to the southeast (Cody et al. 2001). | Harrimanella stellariana (Pallas) Coville — YUKON: Morley River watershed, Englishman Range, 8 km SW of Morris Lake, northwest facing slope growing in bedrock controlled draw, 60°23'40"N 131°47'50"W, J. Meikle 02-126, 20 June 2002 (DAO). The specimen cited above is a rarity in the Territory (Doug- las et al. 1981). It is a slight range extension north of the eastern site mapped by Cody (1996). Phyllodoce glanduliflora (Hook). Cov., Yellow Moun- tain-heather — YUKON: subalpine slope by creek, Tank Creek, 60°05'30"N 134°41.9'W, S. Withers SWO1-114, 15 July 2001 (DAO). Cody (1996) knew this species in the Territory only from sites east of longitude 130°W and west of longitude 137°W. The specimen cited above is about 125 kilometers east of a site in Kluane National Park. Vaccinium membranaceum Dougl., Tall Blueberry — YUKON: moist subalpine meadow, Kotaneelee Range, 60°14'31L"N 124°7'19"W, B. Bennett 95-149, 20 June 1998 (B. Bennett Herbarium, photo DAO); subalpine forest, Beavercrow Ridge, 60°12.865'N 124°35.79'W, B. Bennett 98-310, (B. Bennett Herbarium, photo DAO). Cody (1996) suggested that this species should be looked for in the southeastern part of the Territory. Cody et al. (1998) reported four collections from Mount Merrill and Gusty Lakes. The two specimens cited above are from the extreme southeast. PRIMULACEAE Douglasia alaskana (Cov. & Standl.) Kelso — YUKON: eroded alpine slope, Table Mountain, Kluane National Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI 575 Park, 61°15'.231'N 139°11.115'W, P. Caswell 282, 29 June 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory where it was only known in Kluane National Park. The specimen cited above extends the known range in the Park about 75 kilometers to the northwest. Douglasia arctica Hook. — YUKON: Cody et al. (2000) reported this species from a mountain between Kusawa and JoJo Lake, which was a considerable southward range extension. The specimen upon which it was reported was unfortu- nately misidentified and the range extension should be deleted. Primula eximia Greene (P. tschuktschorum Kjellm. ssp. cairnesiana A. E. Porsild) — YUKON: Dempster Highway about Km 96, at about altitude 4000 ft., S. Frisch, spring 2001 (DAO). Douglas et al. (1981) considered this species rare in the Territory on the basis of collections from the northern Rich- ardson Mountains and a site adjacent to the Alaska border at about 63°50'N latitude. The specimen cited above is from a site about 150 kilometers northeast of the Alaska border site where there was “a whole field of them". Trientalis europaea L. s.1., Starflower — YUKON: damp mossy Picea glauca, Alnus incana, Dawson area, 64°02'00"N 137°41'30"W, G. Brunner 439, 30 June 2001 (DAO). The specimen cited above from about 70 kilometers east of Dawson City, is only the second collection in the Territory from north of latitude 64°N (Cody 1996). GENTIANACEAE Gentianella tenella (Rottb.) Boerner — YUKON: alpine barrens on long ridge, Atlas Mountain, Kluane Nation- al Park, 61°14.772'N 139°19.254'W, D. Normandeau s.n., P- Caswell 328A, 6 July 2002 (DAO); patch of al- pine tundra, mountain opposite Donjek Glacier, Kluane National Park, 61°11.859'N 139°23.372'W, P. Cas- well 619, 26 July 2002 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). Cody (1996) knew it from only three sites, two about 40 kilometers west of the specimens cited above and one just north of latitude 62°N. POLEMONIACEAE Phlox hoodii Richards., Moss Phlox — YUKON: dry rocky slope, open habitat, Donjek River watershed near Kluane Glacier, M. Hoefs 02-693, 4 July 2002 (DAO). The specimen cited above from west of the Alaska High- way is the fifth and southernmost in that area. This species was not included in the Rare Vascular Plants of the Yukon (Douglas et al. 1981) because of its widespread distribution. Polemonium acutiflorum Willd. forma lacteum Lepage — YUKON: rare in lush subalpine meadow at 4800 ft.. mountain 4 miles west of Upper Hyland Lake, 62°03'N 128°59'W, Calder & Kukkonen 27889, 3 Aug. 1960 (DAO) (determined by B. Boivin 1970). Lepage (1950) described this white-flowered form on the basis of a specimen collected at Anchorage, Alaska. Cody (1996) unfortunately did not mention this white form occurring in the Yukon Flora. 576 Polemonium boreale Adams forma albiflorum Cody, — YUKON: fox den, sandy/grass area, Herschel Island, site 203, 69°30'N 139°15'W, C. E. Kennedy 262, 16 July 1985 (DAO) (PARATYPE); [NORTHWEST TERRI- TORIES: MACKENZIE: on a sandy portion of a gravel bar, Horton River, N.W.T., 69°42'N 126°56'W, G. W. Scotter 101016b, 6 July 1995 (DAO) (HOLOTYPE) (Cody et al. 2003). Cody (1996) stated that the corolla of this species was blue to violet or rarely white. The white form had not been described at that time. HYDROPHYLLACEAE Phacelia mollis Macb., Macbryde’s Phacelia — YUKON: scattered in dry, open rubble on south-facing slopes near base of bluff, Moosehide Hills, Bluff on N side of Yukon R., 25 km downstream from Dawson, 64°15'41"N 139°36'13"W, C. Roland 93-27, 24 May 1993 (AKA, photo DAO). The specimen cited above is an extension of about 50 kilo- meters to the southwest from the nearest site in the Ogilvie Mountains known to Cody (1996). BORAGINACEAE Eritrichium aretioides (Cham. & Schlecht.) DC. — YUKON: dry alpine Dryas heath, Canada/US Boundary Monument 97, 65°21'25"N 141°00'00"W, B. Bennett 02-019, 27 June 2002 (DAO). The specimen cited above is an extension of the known range in the Territory of about 40 kilometers northwest of the nearest site known to Cody (1996) between latitudes 64° and 66°N. Eritrichium chamissonis DC. — YUKON: vicinity of Snowdrift Camp, Vuntut National Park, 68°21.4'N 139°13.1'W, P. Caswell PPC-Y-076, 19 June 2000 (B. Bennett Herbarium, photo DAO). Cody (1996) knew this rare species in the Territory from only two localities in the British Mountains. The specimen cited above is from a site about 50 kilometers south of a site on Mt. Sedgwick. Mertensia paniculata (Britt.) G. Don var. alaskana (Britt.) Williams — YUKON: adjacent to Joe Robert’s Cabin, Fort Selkirk, 62°46'34.4"N 137°23'35.9"W, Cody & Kennedy 37796, 17 July 2002 (DAO). Cody (1996) knew this variety only from south of latitude 61°30'N and north of latitude 67°30'N. Myosotis scorpioides L., Forget-me-not — YUKON: flower bed at Kluane Park Inn, Haines Junction, 60°44'N 137°31'W, P. Caswell PPC-2000-Y-426, 28 Aug. 2000 (B. Bennett Herbarium, photo DAO). This introduced species was previously known in the Ter- ritory only from the vicinity of Whitehorse (Cody 1996). LAMIACEAE (LABIATAE) Galeopsis tetrahit L. ssp. bifida (Boenn.) Fries, Hemp- nettle — YUKON: Forty Mile Historic Site at the conflu- ence of the Forty Mile and Yukon rivers, 64°25'N 140°32'W, C. E. Kennedy 2, 6 July 2000 (DAO); under large Picea glauca in squirrel midden, Judas Creek, 60°23.4'N 134°07.4'W, S. Withers SWO1-178, 30 Aug. 2001 (B. Bennett Herbarium, photo DAO). THE CANADIAN FIELD-NATURALIST Vol. 118 This introduced species was previously known in the Ter- ritory from only three localities: North Canol Road (Cody 1994, 1996), Km 9 Dempster Highway (Cody et al. 1998) and Whitehorse (Cody et al. 2001). The first specimen cited above is the northernmost yet found in the Territory. SCROPHULARIACEAE Euphrasia subarctica Raup — YUKON: stony soil and moss in cleared area below rest stop, Klondike High- way Km 628, 63°50'18.1"N 137°59'20.5"W, Cody & Cody 37660, 4 Aug. 2001 (DAO). The specimen cited above is the northernmost yet found in the Territory (Cody 1996). It is from a site about 110 kilo- meters northwest/west of Mayo. Mimulus guttatus DC., Yellow Monkeyflower — YUKON: small seep near the road, Km 4.5 Atlin Road, 60°17'30"N 133°50'3"W, S. Withers SWO1-131, 24 July 2001 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is from a site about 80 kilometers southeast of Whitehorse. Pedicularis oederi Vahl., Oeder’s Lousewort — YUKON: Herschel Island, 69°35'N 139°05'W, C. E. Kennedy s.n., 30 July 1985 (Yukon Renewable Government Herbarium, photo DAO). This species, which is frequent in the British Mountains to the south is now known to the north on Herschel Island. Rhinanathus minor L. ssp. borealis (Sterneck) A. Love — Yukon: gravel parking area at foot of trail to Rock Glacier, Haines Highway, 60°27'18"N 137°03'37.6"W, Cody & Cody 37894, 21 July 2002 (DAO). This taxon is known in the Territory only as far north as about latitude 61°30'N. The specimen cited above is the first record from the Haines Highway. OROBANCHACEAE Orobanche fasciculata Nutt. — YUKON: 45° silty sand south-facing slope with Artemisia frigida and Cala- magrostis purpurascens, Dutch Bluff, Yukon River, 61°55.72'N 135°03.99'W, B. Bennett 02-290, 7 Aug. 2002 (DAO). This species was considered rare in the Territory by Doug- las et al. (1981). Cody et al. (2003) reported a new site in Kluane National Park adjacent to the Alaska Highway and southern Kluane Lake. The specimen cited above is from a site about 50 kilometers southeast of Carmacks. LENTIBULARIACEAE Pinguicula villosa L. — YUKON: in Sphagnum tus- sock, bog east of Haines Highway, 60°08.421'N 136°58.499'W, P. Caswell 446, 13 July 2002 (DAO). The specimen cited above is new to southwestern Yukon. It is an extension of the known range in the Territory of about 300 kilometers southwest from a site adjacent to the northern South Canol Road. Utricularia minor L., Lesser Bladderwort — YUKON: shallow water of lake between Ross River and Camp- bell Highway, 61°58'16"N 132°38'06"W, Cody & Cody 37967, 24 July 2002 (DAO). This is a rare species in the Territory that Cody (1996) knew from only four sites between latitudes 60°N and 64°30'N. 2004 Cody et al. (1998, 2000 and 2003) reported new collections from the southeast in the vicinity of La Biche River and in the far north between Trout Lake and the Babbage River. The specimen cited above is from about 150 kilometers north of a site near the south end of the South Canol Road. PLANTAGINACEAE Plantago canescens Adams — YUKON: base of cliffs, talus, south-facing slope, Mount Carmacks, 64°20'08"N 140°12'25"W, B. Bennett 02-820, 24 Aug. 2002 (DAO). The specimen cited above is an extension of the known range in the Territory of about 200 kilometers northwest of a site adjacent to the Klondike Highway (Cody 1996). _ Plantago maritima L., Seaside Plaintain — YUKON: dry sandy gravel slope north of gravel pit south of Alaska Highway, 60°49'58"N 135°45'49"W, Cody & Cody — 38204, 1 Aug. 2002 (DAO). This species was considered rare in the Yukon Territory by Douglas et al. (1981). Cody (1996) knew it from only two localities adjacent to Kluane Lake in the extreme southwest. _ The specimen cited above is from a site about 100 kilometers to the east. _ CAPRIFOLIACEAE _ Sambucus racemosa L. ssp. pubens (Michx.) House, _ Red Elderberry — YUKON: roadside and lakeshore, Wade Lakes, 60°05.0'N 137°19.6'W, R. Maraj s.n., 14 Sept. 2002 (DAO). Douglas et al. (1981) (as var. arborescens) knew this taxon from a single locality in southern Kluane National Park. The specimen cited above extends the known range about 80 kilo- _ meters to the east between the park and Haines Highway. VALERIANACEAE Valeriana sitchensis Bong., Sitka Valerian — YUKON: Site YPN 120, 63°06'39.8"N 133°20'15"W, G. Brun- ner 569-01, 6 Aug. 2001 (Environment Yukon, photo DAO). The specimen cited above is from a site between the North Canol Road and Mayo. ASTERACEAE (COMPOSITAE) Aster alpinus L. ssp. vierhapperi Onno, Alpine Aster — YUKON: steep grassy slope on ridge, northeast of Woodburn Creek, Tintina Trench, Ddhaw Ghro, _ 63°08'N 136°05'W, C. E. Kennedy 2, 27 July 2001 _ (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 near the junction of the Pelly and Yukon rivers. Aster falcatus Lindl., Western Heath Aster — YUKON: lush herbaceous meadow riverbank, Kirkman Creek, Yukon River, 62°59'21"N 139°23'07"W, B. Bennett 02-673, 10 Aug. 2002 (B. Bennett Herbarium, photo DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is from a site about 150 kilometers west of a site adjacent to the Klondike Highway. Erigeron caespitosus Nutt. — YUKON: silty sand south- facing slope, Dutch Bluff, Yukon River, 61°55.72'N Copy, KENNEDY, BENNETT, AND CASWELL: PLANTS IN YUKON VI yd 135°03.99'W, B. Bennett 02-300, 7 Aug. 2002 (DAO); top of slope of open bank, 1 mile upstream of Big Salmon, on Yukon River, 61°51.41'N 134°55.01'W, B. Bennett 02-302, 7 Aug. 2002 (DAO); base of cliffs, talus, south-facing slope, Yukon River, Mount Car- macks, 64°20'08"N 140°12'25"W, B. Bennett & R. Mulder 02-816, 24 Aug. 2002 (DAO). The first specimen cited above is from a site about 75 kilo- meters southeast of a site adjacent to Carmacks and the sec- ond specimen is from a site about 120 kilometers southeast of Carmacks. The third specimen is from a site between two sites near the Alaska border, one to the south about 120 kilo- meters and the other to the north about 100 kilometers (Cody 1996). Erigeron hyperboreus Greene — YUKON: middle of dirt road leading to Copper Joe Creek Road, Kluane National Park, 61°18.688'N 138°56.319'W, P. Caswell 170, 24 June 2002 (DAO). This species was considered as rare in the Territory by Douglas et al. (1981). The specimen cited above is an exten- sion of about 425 kilometers south of a site northwest of Dawson City. Senecio eremophilus Richards., Dryland Ragwort (Fig- ure 6) — YUKON: Pine Creek Campground, 4 miles east of Haines Junction, Mile 1012 Alaska Highway, V. L. Harms 6299, 17 August 1968 (DAO). The specimen cited above is the first known record in the Yukon Territory where it was probably introduced. To the FiGurE 6. Senecio eremophilus, Dryland Ragwort (drawn by J. Looman). 578 west in Alaska, Hultén (1968) knew it only from the vicinity of Tok where he considered it also to be introduced. To the east Porsild and Cody (1980) knew it from the Slave River area north of Fort Smith where it was found in damp woodland meadows and adjacent to roadsides and considered native. To the south it is found in eastern British Columbia east to Manitoba and south into the United States. It can be separated from S. sheldonensis as follows: A. Stem leaves deeply incised, not TeducedipwardSey ss yale. coset S. eremophylus B. Stem leaves with merely repand-denticulate margins, reduced upwards ......... S. sheldonensis Solidago canadensis L. var. salebrosa (Piper) Jones — YUKON: beside test plot, 1 km N of Montague Road- house, Km 132 Klondike Highway, 61°49'35"N 136°7'52"W, Cody & Cody 38073, 28 July 2002 (DAO). Douglas et al. (1981) considered this species rare in the Territory. The specimen cited above is from about 90 kilo- meters southwest of a site mapped by Cody (1996). Sonchus arvensis L. ssp. uliginosus (Bieb.) Nyman, Perennial Sow-thistle — YUKON: roadside gravel, Alaska Highway 3 km west of Kusawa Lake turnoff below microwave tower, 60°46'00"N 136°05'36"W, _ B. Bennett 02-749, 11 Sept. 2002 (DAO). The specimen of this introduced species cited above is from a site between Whitehorse and Haines Junction. Tanacetum vulgare L., Common Tansey — YUKON: in distributed ground along Campbell Highway, Tuchitua River, 60°56'N 129°13'00"W, R. Rosie 1959, 16 Aug. 1997 (DAO); single patch west of the boat launch, Yukon River Bridge, 60°34'20"N 134°40'W, B. Ben- nett 01-063, 19 Aug. 2001 (DAO). The specimens cited above are only the second and third records of this introduced species in the Territory. Cody et al. (1998) reported the first collection from just north of Kath- leen Lake Lodge on the Haines Highway. Acknowledgments We thank Donald Britton and Daniel Brunton for their identification of /soetes; Stephen J. Darbyshire for his identification of several Poaceae; Gerald Mulligan for his identification of Brassicaceae specimens; Heidi Solstad for her identification of Papaver croceum; Greg Brunner, Manfred Hoefs, Jennifer Line, R. Maraj, Rhonda Rosie and Stu Withers for their contributions; Margaret Cody and Douglas Cody for assisting the senior author in the summers of 2001 and 2002 in southern Yukon; artist Lee Mennell for his art work; Paul Catling for reviewing an earlier version of this manuscript, and especially Leslie Cody for the many hours inputting this information on her computer. THE CANADIAN FIELD-NATURALIST Vol. 118 Literature Cited Cody, W. J. 1994. The Flora of the Yukon Territory: Addi- tions, Range Extensions and Comments. Canadian Field- Naturalist 108: 428-479. 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. Cana- dian Field-Naturalist 112: 289-328. Cody, W. J., C. E. Kennedy, and B. Bennett. 2000. New Records of Vascular Plants in the Yukon Territory I. Cana- dian Field-Naturalist 114: 417-443. Cody, W. J., C. E. Kennedy, and B. Bennett. 2001. New Records of Vascular Plants in the Yukon Territory HI. Canadian Field-Naturalist 115: 301-322. Cody, W. J., C. E. Kennedy, B. Bennett, and V. Loewen. 2002. New records of Vascular Plants in the Yukon Terri- tory IV. Canadian Field-Naturalist 116: 446-474. Cody, W. J., C. E. Kennedy, B. Bennett, and J. Staniforth. 2003. New records of Vascular Plants in the Yukon Ter- ritory V. Canadian Field-Naturalist 117: 278-301. Cody, W. J., K. L. Reading, and J. L. Line. 2003. Additions and range extensions to the vascular plant flora of the Con- tinental Northwest Territories and Nunavut, Canada II. Canadian Field-Naturalist 117(3): 448-465. 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. Hulten, E. 1941-1950. Flora of Alaska and Yukon, 1-10. Lunds Universitets Arsskrift, N. F. Aud. 2. Volumes 37- 46. 1902 pages. Hulten, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press. Stanford, California. 1008 pages. Kojima, S., and R. C. Brooks. 1985. An annotated vascular flora of areas adjacent to the Dempster Highway, central Yukon Territory. I. Pteridophyta, Gymnospermae and Monocotyledonae. British Columbia Provincial Museum, Contributions of Natural Science 31: 1-16. Lepage, E. 1950. Variations mineures de quelques plantes du nord-est du Canada et de l’ Alaska. Le Naturaliste cana- dien 77 : 228-231. McJannet, C. L., G. W. Argus, and W. J. Cody. 1995. Rare vascular plants in the Northwest Territories. Syllogeus 73: 1-104. Porsild, A. E. 1975. Materials for a flora of central Yukon Territory. National Museum of Natural Sciences. Publica- tions in Botany 4, Ottawa, Ontario. 77 pages. Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Con- tinental Northwest Territories, Canada. National Museum of Natural Sciences, Ottawa, Ontario. 667 pages. Scotter, G. W., and W. J. Cody. 1979. Interesting vascular plants from southeastern Yukon Territory. Canadian Field- Naturalist 93: 163-170. Welsh, S. L. 1974. Anderson’s Flora of Alaska and Adjacent Parts of Canada. Brigham Young University Press, Provo, Utah, U.S.A. 724 pages. Received 11 December 2003 Accepted 3 February 2004 Hybridization Between a Green Turtle, Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and the First Record of a Green Turtle in Atlantic Canada MICHAEL C. JAMES!, KATHLEEN MARTIN2, and PETER H. DuTTON? ' Department of Biology, Dalhousie University, 1355 Oxford St., Halifax, Nova Scotia B3H 4J1 Canada (Corresponding author) > Nova Scotia Leatherback Turtle Working Group, 2070 Oxford St., Halifax, Nova Scotia B3L 2T2 Canada 3 Southwest Fisheries Science Center, National Marine Fisheries Service, 8604 La Jolla Shores Dr., La Jolla, California 92037 USA James, Michael C., Kathleen Martin, and Peter H. Dutton. 2004. Hybridization between a Green Turtle, Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and the first record of a Green Turtle in Atlantic Canada. Canadian Field- Naturalist 118 (4): 579-582. The Green Turtle (Chelonia mydas) principally occupies tropical and subtropical waters, although juveniles are known to occur seasonally in temperate coastal waters. Collaboration with commercial fishers in eastern Canada yielded the most northerly records of this species in the northwest Atlantic. Here we report on the first confirmed record of a Green Turtle in eastern Canada and on the occurrence of a rare Green Turtle—Loggerhead Turtle (Caretta caretta) hybrid. Hybridization between the Carettini and Chelonini is extraordinary given that these groups have been genetically distinct for 50 million years Or more. Key Words: Green turtle, Chelonia mydas, hybrid, Loggerhead Turtle, Caretta caretta, Atlantic Canada. The Green Turtle (Chelonia mydas) has a broad range in the Atlantic, which includes waters off the continental United States. Presence in temperate waters of the northeastern United States is seasonal, with tur- tles retreating to more southerly latitudes when water temperatures decline (Epperly et al. 1995). During summer and fall, this species regularly occurs as far north as New York (Morreale et al. 1992); however, records of Green Turtles at higher latitudes of the United States are rare. While both Leatherback Tur- tles (Dermochelys coriacea) and Loggerhead Turtles (Caretta caretta) are commonly encountered in waters off Atlantic Canada (e.g., Bleakney 1965), with leath- erbacks occupying both nearshore and offshore waters (James et al. 2005) and loggerheads mainly offshore waters, there were no previous confirmed reports of the Green Turtle in this region. Here we report on photo-documented records of a Green Turtle and a Green Turtle-Loggerhead Turtle hybrid in nearshore waters off Nova Scotia, Canada. These exist as the most northerly confirmed records of Chelonia mydas in the northwest Atlantic. Notification about both turtles initially came through calls to a toll-free phone line established for fishing community members in Atlantic Canada to report sea turtle sightings (Martin and James 2005). A small, live cheloniid turtle was reported on 8 August 1999. The turtle was found in Chedabucto Bay, Nova Scotia (at 45°20'37"N, 61°15'36"W, Figure 1) and was pho- tographed and released shortly before the call was 46 - (ag Chedabucto Bay 450 Nova Scotia 2 4 St Margarets Ite Bay 447 ; | eo | a = A FiGurE 1. Capture locations of juvenile Green Turtle (solid circle) and juvenile Green Turtle x Loggerhead Tur- tle hybrid (solid triangle). placed. The turtle was not measured; however, curved carapace length (CCL) was estimated to be between 30 and 40 cm and its mass approximately 4-5 kg. Species identification was confirmed as C. mydas (juve- nile) upon receipt of several excellent photographs that depicted carapacial scute configuration and scale patterning on the head (Figure 2). The second turtle, also live, was reported on 2 Octo- ber 2001. It was found in St. Margarets Bay, Nova Sco- tia (at 44°34'56"N, 64°03'06"W, Figure 1). The animal 579 580 THE CANADIAN FIELD-NATURALIST Vol. 118 FiGure 2. Juvenile Green Turtle (Chelonia mydas) found in Chedabucto Bay, Nova Scotia, on 8 August 1999. was considered unusual by the inshore commercial fishers who encountered it because they had never seen a cheloniid turtle before. Therefore, they collected it and brought it to shore for examination. One of us (MCJ) responded to the report and exam- ined the turtle (CCL 34 cm; mass 4.07 kg), tentatively identifying it as a juvenile C. mydas. The plastron of the turtle was cream coloured, the cutting edge of the lower tomium was mildly serrated, and it had a pair of large prefrontal scutes on its head (Figure 3), all features characteristic of a Green Turtle. However, while the number and arrangement of carapacial scutes was consistent with those of a Green Turtle, there were two claws on the anterior margin of each front flipper, the costal and vertebral scutes overlapped con- siderably, and the marginal scutes were strongly ser- rated (Figure 3), which suggested that the turtle was possibly a hybrid. At the time of examination, the turtle’s movements were sluggish and it was judged to be mildly hypother- mic. Small juvenile cheloniid turtles foraging in coastal areas of the temperate north Atlantic in the fall are particularly vulnerable to developing hypothermia, as water temperatures can rapidly drop below 20°C (Dav- enport 1997). This animal was recovered from an area where the sea surface temperature was 16.2°C. Given the turtle’s compromised physical condition and the declining water temperatures along the coast of Nova Scotia, the turtle was moved to the animal care facili- ties at Dalhousie University in Halifax, where it was warmed and rehydrated for approximately 60 hours in a freshwater bath at 24°C. It was then transported south by air on the evening of 4 October 2001 for release in Bermudian waters. Subsequent genetic analysis at the U.S. National Marine Fisheries Laboratory in La Jolla, California, confirmed that this turtle was a hybrid. This was deter- mined by sequencing a 391 bp fragment of the control region of mitochondrial DNA (mtDNA) extracted from a blood sample (Dutton 1996). The results revealed that this turtle had Loggerhead Turtle mtDNA, while phenotypically it was principally a Green Turtle. Since mtDNA is maternally inherited, we conclude that this animal was the progeny of a female Loggerhead Turtle and a male Green Turtle. Discussion The extent to which natural hybridization in sea tur- tles occurs has not been determined. However, as genet- ic techniques like the ones employed here are increas- ingly applied, hybridization may prove to be more common than previously thought. Molecular genetics have confirmed hybridization between the Loggerhead Turtle and Kemp’s Ridley Turtle (Lepidochelys kempii) (Karl et al. 1995; Barber et al. 2003), Loggerhead Turtle and Hawksbill Turtle (Eretmochelys imbricata) (Karl et al. 1995; Witzell and Schmid 2003), and Green Turtle and Hawksbill Turtle (Wood et al. 1983; Karl 2004 et al. 1995). In addition to the results presented here, there is only one previous report of hybridization between a Loggerhead Turtle and a Green Turtle (Karl et al. 1995). Hybridization between the Carettini and Chelonini is extraordinary given that these tribes have been genetically separated for 50 million years or more (Bowen et al. 1993; Dutton et al. 1996). It has been suggested that these may be the oldest vertebrate line- ages known to hybridize in nature (Karl et al. 1995). The reproductive status of marine turtle hybrids is not known; however, the identification of potential second-generation hybrids (Karl et al. 1995) suggests that at least some hybrids may be fertile. Reproductive viability in hybrids could have important biological consequences for the conservation of marine turtles (Karl 1996). Both turtles reported here were of the size class typ- ical of neritic foraging populations of Green Turtles (i.e., straight carapace length >25 cm) (Musick and Limpus 1997). Therefore, these animals likely arrived in eastern Canadian waters from coastal foraging habi- tat off the northeastern United States, rather than from pelagic areas. Our current understanding of juvenile Green Turtle distribution in the northwest Atlantic suggests that these turtles occupy areas where sea surface temperature is normally higher than that of coastal Atlantic Canada. In the case of the hybrid turtle, the Loggerhead Turtle component of its genotype may have been responsible for directing the animal to higher latitudes for for- aging, as Loggerhead Turtles are commonly encoun- tered in Atlantic Canadian offshore waters. However, the animal’s presence in nearshore waters off Nova Scotia remains puzzling. Loggerhead Turtles are rarely encountered in coastal areas of Atlantic Canada be- cause sea surface temperatures are normally at or below the lower thermal tolerance limits of this species. There was no formal attempt in eastern Canada to specifically promote the reporting of marine turtle sightings until 1998, when a broad public education program and toll-free reporting line were established JAMES, MARTIN, and DUTTON: GREEN TURTLE IN ATLANTIC CANADA The large pair of prefrontal scutes on the head (left) are characteristic of Chelonia mydas. However, overlapping costal and vertebral scutes and strongly serrated marginal scutes (right) are not characteristic of Chelonia mydas. for this purpose (Martin and James 2005). Continued collaboration with commercial fishers will be key to determining if the records reported here represent accidental occurrences of the Green Turtle in this region, or if small numbers of this species regularly forage along the Scotian Shelf in summer and early fall when inshore water temperatures are highest. Acknowledgments We thank K. Horne, A. Harnish and P. Young for reporting the turtles described here to the Nova Sco- tia Leatherback Turtle Working Group. Thanks are also extended to C. Harvey-Clark for treating the hybrid turtle at Dalhousie University’s Animal Care facilities and to J. Conway (Maritime Aquatic Species at Risk Office), K. Eckert (Wider Caribbean Sea Turtle network) and J. Gray (Bermuda Aquarium Museum and Zoo) for facilitating the transport to and subse- quent release of this turtle in Bermudian waters. We thank S. Mockford (Dalhousie University), R. Le Roux and E. La Casella (National Marine Fisheries Service) for assistance with genetic analysis and R. Myers (Dalhousie University) for helpful comments on earli- er drafts of the manuscript. We acknowledge support from World Wildlife Fund Canada, George Cedric Metcalf Charitable Foundation, Canadian Wildlife Federation, U.S. National Marine Fisheries Service, the Habitat Stewardship Program for Species at Risk, and the Natural Sciences and Engineering Research Council of Canada (scholarship to M.C.J.). This work was conducted under Department of Fisheries and Oceans License #2001-425. Literature Cited Barber, R. C., C. T. Fontaine, J. P. Flanagan, and E. E. Louis, Jr. 2003. Natural hybridization between a Kemp’s Ridley (Lepidochelys kempii) and Loggerhead sea turtle (Caretta caretta) confirmed by molecular analysis. Che- lonian Conservation and Biology 4: 701-704. Bleakney, J. S. 1965. Reports of marine turtles from New England and Eastern Canada. Canadian Field-Naturalist 79: 120-128. 582 Bowen, B. W., W. S. Nelson, and J. C. Avise. 1993. A molec- ular phylogeny for marine turtles: trait mapping, rate assessment and conservation relevance. Proceedings of the National Academy of Science (USA) 90: 5574-5577. Davenport, J. 1997. Temperature and the life history strategies of sea turtles. Journal of Thermal Biology 22: 479-488. Dutton, P. H. 1996. Methods for collection and preserva- tion of samples for sea turtle genetic studies. Pages 17-24 in Proceedings of the International Symposium on Sea Turtle Conservation Genetics. Edited by B. W. Bowen and W. N. Witzell. NOAA Technical Memorandum NMEFS- SEFSC-396. Dutton, P. H., S. K. Davis, T. Guerra, and D. Owens. 1996. Molecular phylogeny for marine turtles based on sequences of the ND4-leucine tRNA and control regions of mtDNA. Molecular Phylogenetics and Evolution 5: 511-521. Epperly, S. P., J. Braun, and A. Veishlow. 1995. Sea turtles in North Carolina waters. Conservation Biology 9: 384-394. James, M. C., R. A. Myers, and C. A. Ottensmeyer. 2005. Identification of high-use habitat and threats to leather- back sea turtles in northern waters: new directions for conservation. Ecology Letters 8: 195-201. Karl, S. A. 1996. Hybridization and taxonomy of marine tur- tles: anonymous nuclear DNA sequence analyses. Pages 99-108 in Proceedings of the International Symposium on Sea Turtle Conservation Genetics. Edited by B. W. Bowen and W. N. Witzell. NOAA Technical Memorandum NMFS- SEFSC-396. THE CANADIAN FIELD-NATURALIST Vol. 118 Karl, S. A., B. W. Bowen, and J. C. Avise. 1995. Hybridiza- tion among ancient mariners: characterization of marine turtle hybrids with molecular genetic assays. Journal of Heredity 86: 262-268. Martin, K., and M. C. James. 2005. Conserving sea turtles in Canada: a model for successful collaboration between fishers and scientists. Chelonian Conservation and Biology 4: 899-907. Morreale, S. J., A. B. Meylan, S. S. Sadove, and E. A. Standora. 1992. Annual occurrence and winter mortality of marine turtles in New York waters. Journal of Herpe- tology 26: 301-308. Musick, J. A., and C. J. Limpus. 1997. Habitat utilization and migration in juvenile sea turtles. Pages 137-163 in The Biology of Sea Turtles. Edited by P. L. Lutz and J. A. Musick. CRC Press, Boca Raton, Florida. Witzell, W. N., and J. R. Schmid. 2003. Multiple recaptures of a hybrid Hawksbill-Loggerhead turtle in the Ten Thou- sand Islands, southwest Florida. Herpetological Review 34: 323-325. Wood, J. R., F. E. Wood, and K. Critchley. 1983. Hybridiza- tion of Chelonia mydas and Eretmochelys imbricata. Copeia 1983: 839-842. Received 26 January 2004 Accepted 6 December 2004 A Snow-tracking Protocol Used to Delineate Local Lynx, Lynx canadensis, Distributions JOHN R. SQUIRES, KEVIN S. MCKELVEY, and LEONARD F. RUGGIERO U.S. Forest Service, Rocky Mountain Research Station, Forestry Science Laboratory, P.O. Box 8089, Missoula, Montana 59807 USA. jsquires @fs.fed.us Squires, John R., Kevin S. McKelvey, and Leonard F. Ruggiero. 2004. A snow-tracking protocol used to delineate local lynx, Lynx canadensis, distributions. Canadian Field-Naturalist 118(4): 583-589. Determining Canada Lynx (Lynx canadensis) distribution is an important management need, especially at the southern extent of the species range where it is listed as threatened under the U. S. Endangered Species Act. We describe a systematic snow- track based sampling framework that provides reliable distribution data for Canada Lynx. We used computer simulations to evaluate protocol efficacy. Based on these simulations, the probability of detecting lynx tracks during a single visit (8 km transect) to a survey unit ranged from approximately 0.23 for surveys conducted only one day after snowfall, to 0.78 for surveys conducted 7 days after a snowfall. If the survey effort was increased to three visits, then detection probabilities increased substantially from 0.58 for one day after snowfall to about 0.95 for surveys conducted 7 days after a snowfall. We tested the protocol in the Garnet Range, Montana, where most lynx were radio-collared. We documented a total of 189 lynx tracks during two winters (2001-2003). Lynx distribution based on snow-track surveys was coincident with the area defined through radio telemetry. Additionally, we conducted snow-track surveys in areas of western Wyoming where lynx were believed present but scarce. We detected a total of six lynx tracks during three winters (1999-2002). In Wyoming , where lynx presence was inferred from a few tracks, we verified species identification by securing genetic samples (hairs from daybeds) along track-lines. Key Words: distribution, forest carnivore, Canada Lynx, Lynx canadensis, snow-track, Montana, Wyoming, surveys. Listing Canada Lynx (Lynx canadensis) as “Threat- ened” in the U. S. under the Endangered Species Act compels land managers to consult with the U.S. Fish and Wildlife Service regarding potential impacts of management actions (Federal Register, Volume 63, Number 130). One of the most fundamental informa- tion needs is to delineate the current distribution of lynx in the contiguous United States. Definitive range determination for rare and elusive carnivores, like the lynx, is difficult and may require several well-tested survey methods. The National Lynx Survey uses hair- collection pads (McDaniel et al. 2000) as a means of detection over large areas including much of the north- ern and western portions of the United States (K. S. McKelvey). DNA analysis of collected hair provides positive species identification (Mills et al. 2000), and is the only tested detection method during non-snow periods for lynx. However, unless multiple samples are obtained from a site, the method cannot distinguish local populations from single dispersing individuals, an important issue for land managers. Snow-tracking provides a survey method for detect- ing lynx during the winter (Halfpenny et al. 1995; Zielinski and Kucera 1995). Unlike hair-pad surveys, snow track surveys do not require a behavioral re- sponse, like rubbing, for detection. Lynx also have many attributes that make them particularly good candidates for snow-track surveys: lynx from southern populations have large spatial-use areas (approximately 150 km? for males and 70 km? for females; Aubry et al. 2000); have high daily travel rates (Ward and Krebs 1985; Mowat et al. 2000); and a distinctive snow-track that facilitates identification (Halfpenny and Biesiot 1986; Forrest 1988). Snow-tracking also provides anecdotal infor- mation, such as identification of family groups, useful for distinguishing local populations from dispersing individuals. Snow tracking has been used extensively to survey lynx and other forest carnivores (Thompson et al. 1989; Stephenson and Karczmarczyk 1989; Beier and Cun- ningham 1996; Becker et al. 1999), but, like other detec- tion techniques, snow-track surveys have inherent strengths and weaknesses. Snow track surveys can be conducted across extensive landscapes at reasonable costs, and the method is sensitive to changes in carni- vore occupancy. However, potential problems with snow-track surveys include track misidentification and difficulties in achieving representative surveys (Half- penny et al. 1995; Zielinski and Kucera 1995). For ex- ample, Aubry and Lewis (2003) found that unscreened records of Fishers (Martes pennanti) that presumably included many “false positives” yielded a different, and much more extensive, distribution when compared to verified records. Misidentifications can also have serious management impacts if the detected species is listed as Threatened or Endangered under the U. S. Endangered Species Act. Here, we describe and evaluate a track-based survey method for lynx and its potential limitations. Our goal was to develop a reliable and representative winter sur- vey method to complement existing summer surveys. Specifically, we needed to improve on existing snow- 583 track survey methodologies to ensure they are repre- sentative in terms of their spatial extent and habitat coverage, and to address problems of track misidenti- fication by incorporating genetic sampling (i.e., scats and hairs frozen in tracks and daybeds). To evaluate reliability we tested these methods both through com- puter simulation and through direct comparisons with radio-telemetry data in an area where most lynx were radio-collared. Study Areas We conducted lynx surveys in the Garnet Mountain Range of western Montana, and in the Wyoming/Salt River Mountains approximately 80 km southwest of Jackson, Wyoming. Approximately 80% of the Garnet Range is forested within the Douglas-fir (Pseudotsuga menziesii) and Subalpine Fir (Abies lasiocarpa) series of the Montana Forest Habitat type (Pfister et al. 1977; Burcham et al. 1999). Dominant tree species include Subalpine Fir, Engelmann Spruce (Picea engelmannii), Lodgepole Pine (Pinus contorta), and Douglas-fir. Ele- vations range from 1160 to 2090 m. The primary use of forested public and private lands is timber management, and much of the area was extensively logged within the past 25 years. The Garnet Range is adjacent to private lands that consist mainly of irrigated hay fields and Big Sage Brush (Artemisia tridentata) — wheat grass (Agropyron spp.) cover types. The Wyoming and Salt River ranges, located in west- central Wyoming, support mixed conifer forests that include Subalpine Fir, Engelmann Spruce, Lodgepole Pine, Whitebark Pine (Pinus albicaulis), Douglas-fir, and Quaking Aspen (Populus tremuloides). Mesic sites are dominated by Subalpine fir and Engelmann Spruce forests in seasonally moist or wet areas, or where sub- irrigation maintains a high water table. Spruce-fir forests are typically co-dominant with Lodgepole Pine. Dry sites are dominated by Lodgepole Pine forests, often intermixed with Douglas-fir and Quaking Aspen. Pre- THE CANADIAN FIELD-NATURALIST Vol. 118 cipitation is mostly in the form of snow; elevations range from 1981 — 3353 m. Methods Modeling Track Detections We estimated the probability of detection relative to search effort using computer simulation to model track detections based on the tortuosity of actual lynx tracks. As part of our research, we __ back- tracked 15 radio-collared lynx and mapped their tracks using a Trimble GEO Explorer II with data points taken at 2 s intervals. We randomly selected one 3-km track from each lynx and calculated the average track tortuosity by dividing total length by the linear dis- tance traveled (Turchin 1998). This result was used when generating simulated tracks to ensure realistic track tortuosity. Computer simulations required that we estimate daily travel distances of lynx. Daily movements as meas- ured by radio telemetry are shorter than the animal’s actual movement because of unrecorded tortuosity. In general, reported daily movements of lynx ranged from 1-9 km as measured using various methods, with most estimates from 1-4 km (Table 1). For mod- eling, we assumed that daily movements varied ran- domly from 1.0-4.0 km (z= 2.5 km). We chose a con- servative estimate of daily movements to better ensure that our model results addressed locating lynx at low densities where tracks were uncommon. A uniformly- distributed random number was drawn from this range for each trial. A simulated day’s movement was com- pleted when the linear distance traveled between the day’s start and the current location exceeded the drawn random number. The program continued in this man- ner counting the number of days elapsed until the survey transect intersected a simulated track, and the number of elapsed days was written to an output file. Simulated tracks began at random locations within hypothetical, circular home ranges of 100 km? with TABLE |. Reported daily distances for lynx movement; only winter movements were included where both summer and win- ter movements were reported. Study Distance (km) Parker et al. 1983 8.8, male 7.6, female 6.5, juvenile Ward and Krebs 1985 2.7 (95% CI = 1.8-3.7), hares common Method Cumulative distance based on 2 hour relocations Daily telemetry locations 2.4 (95% CI = 2.0-2.9), hares uncommon Brittell et al. 1989 1.0, (range = 0.02-7.4), males 0.7, (range = 0.02-4.0), females 3.8 (sd = 0.6), male 3.0 (sd = 0.4) female Apps 2000 Mowat et al. 2000 Squires and Laurion 2000 5-9 reported average of multiple studies 2.8 (range = 2.5-3.3), 4 Montana males Daily telemetry locations Daily telemetry locations Snow tracking Daily telemetry locations 3.2 (range = 2.5-3.9), 3 Montana females 2.7 (range = 0.7-9.5), Wyoming male 2.2 (range = 0.3-5.2), Wyoming female 2004 an initial random direction. The length of simulated tracks was evaluated using travel distances that would be expected if we assumed that 1, 3, or 7 days had elapsed since the most recent snowfall. The survey route was modeled as a straight, 8-km transect, randomly located within the 100 km? home range. The entire transect was incorporated into the home range, an assumption based on the study design in which the simulated survey track would be repli- cated within each cell of a contiguous grid. Hence, at least 8 linear km of survey track would be found in any arbitrary 100 km? circular area within the survey. Snow-track Protocol We used GIS to establish an 8 x 8 km grid across each survey area to define sample units. The size of this sample unit was slightly smaller than a typical female home range (Aubry et al. 2000) and was em- ployed to reduce the chance of missing individuals, but was still efficient for searching large landscapes (Zielinski et al. 1995). Female lynx typically have small- er home ranges than males (Koehler 1990; Poole 1994; Slough and Mowat 1996). Establishing a grid of sur- vey units covering the study areas ensured that our search effort was spatially well distributed across avail- able habitats (Zielinski et al. 1995). After establishing the sample grids, we assigned a high or low priority to sampling units based on dominant habitat-type. Sam- ple units in moist, Subalpine Fir forests were a high priority because these stands are more heavily used by lynx than drier forest types (McKelvey et al. 2000). Our protocol specified that “high” priority units be surveyed at least twice per winter (Halfpenny et al. 1995). Open-forest types, such as dry Lodgepole Pine, Quaking Aspen, and Ponderosa Pine (Pinus pon- derosa), were “low” priority and were only searched once per winter. Open habitat types (large park lands, sage brush, tundra, agriculture) and high elevation sites (tundra, rock, and ice) are rarely used by lynx (Koehler 1990; Aubry et al. 2000; McKelvey et al. 2000) and were not surveyed. Observers primarily searched for animal tracks by traveling 15-20 km/hr on roads and trails using snow- mobiles. We saw no evidence that lynx were reluctant to cross snowed-in logging roads and trails (Squires; unpublished data; see also O’Donoghue et al. 1998). Survey units that could not be surveyed by snowmobile were searched on snowshoes to the extent possible. The protocol specified that we search 10 km in each 8 x 8 km sample unit choosing routes that preferen- tially traversed forested habitats with high horizontal cover. This search effort per sample unit was sufficient to traverse key areas within the unit, but still allowed us to efficiently search multiple sample units per day. High horizontal cover is an important component of Snowshoe Hare (Lepus americanus) habitat, the domi- nant prey species of lynx (Hodges 2000). Sample units that contained only dry, open forests were surveyed using routes that best bisected the search area. SQUIRES, MCKELVEY, and RUGGIERO: SNOW-TRACKING PROTOCOL 585 Our goal was a representative survey of sample units that were spatially well distributed across available habitats, while preferentially searching the best habitats in each unit to maximize the chance of detecting lynx. We recorded the locations of all lynx tracks and the tracks of other carnivores using a Trimble GEO Ex- plorer Il GPS. Multiple crossings known to be the same animal and <100 m apart were recorded as a single detection. We measured (stride, straddle, length, width, depth; Halfpenny et al. 1995) and photographed all rare carnivore tracks to provide documentation. Track mis- identification is a potential problem of snow-based surveys (Zielinski and Kucera 1995; Halfpenny et al. 1995), especially in areas where the species of interest is rare. We collected genetic samples (e.g., hairs, scats) from snow-tracks on the Wyoming study area to pro- vide unequivocal documentation. To collect genetic samples, we followed presumed lynx tracks in search of scats or day beds. We carefully removed loose snow from daybeds until we reached the frozen bed layer. We then thoroughly searched the snow surface and used tweezers to secure hairs frozen in the snow. Hairs were stored in desiccant until the Rocky Mountain Research Station’s Wildlife Genetics Laboratory, Missoula, Mon- tana, extracted and analyzed DNA samples. Snow-track surveys require trained personnel to be effective (Zielinski and Kucera 1995). We trained ob- servers at our long-term lynx study area near Seeley Lake, Montana. Observers back-tracked radio-collared lynx to practice identifying tracks under diverse envi- ronmental conditions (e.g., forest type, snow conditions, topography). Following this training, we assumed that all observers could recognize lynx tracks. Survey Verification In the Garnet Range, we used radio-telemetry data to determine if the distribution of lynx within the study area was coincident with survey results. Lynx were trapped using specially designed box traps baited with carrion, beaver castor, and pie plates and wings for visual lures (Kolbe et al. 2003). Trapping was conduct- ed throughout the winters of 2001-02 and 2002-03. Lynx were chemically immobilized using a syringe pole to administer a mixture of ketamine (10 mg/kg Ketaset®; concentration 100 mg/mL) and xylazine (1 mg/kg; concentration 100 mg/mL); this dose pro- duced predictable immobilization periods (30-40 min- utes) and stable vital signs. We fitted adult lynx with VHF radio transmitters (170 g), weighed, measured, and extracted blood from each for future DNA analy- sis. Kittens were fitted with a padded, 100 g collar that allowed for growth. We monitored lynx movements using aerial telemetry (90% of locations) approximately twice per month augmented with ground-based tel- emetry. We combined all relocation points and calculated a 95% kernel home range to delineate the distribution of the local lynx population based on telemetry (Worton 1989; Seaman et al. 1999). We then overlaid a 95% 586 fixed kernel home range on the distribution of lynx, as delineated by snow tracking, to determine if the two methods yielded coincident distributions. Results Simulations of track detections Computer simulations of track detections indicated different asymptotic relationships between the proba- bility of detection and survey effort (number of visits per winter), based on the time between snowfalls suf- ficient to obliterate tracks (Figure 1). The probability of detecting lynx tracks during a single visit (8 km transect) to a survey unit ranged from approximately 0.23 for surveys conducted only one day after snow- fall, to 0.78 for surveys conducted 7 days after a snow- fall. If the survey effort was increased to three visits, then detection probabilities increased substantially from 0.58 for one day after snowfall to about 0.95 for surveys conducted 7 days after a snowfall. Even a modest sur- vey effort of two visits would detect tracks approxi- mately 80% of the time, provided more than 3 days had passed since the last snowfall. Field test — Garnet Range, Montana During winters 2001-2002 and 2002-2003, we sur- veyed for lynx on 242 km and 438 km of roads and trails, respectively. We documented 37 track detections of lynx in 4 of 12 survey units during winter 2001- 2002, and 37 detections in 4 of 16 survey units during winter 2002-2003. In addition, we observed 115 lynx tracks during winter 2001-2002 and 53 in 2002-2003 in survey units, but not during the formal survey. These detections were made during trapping and related trav- el. We encountered a higher number of track-cross- ings during trapping because we only trapped in areas of high lynx use. The average elevation where we located lynx tracks was 1839 m (SD 97, range 1443 — 2012 m). We trapped five lynx during winter 2001-2002, and two additional individuals during winter 2002-2003. We believed this sample of radio-collared individuals was a local census of lynx in the Garnet Range based on trapping results, recaptures, and field observations, but we could not formally test this perception. Field test — Wyoming and Salt River Ranges, Wyoming We conducted track-surveys for lynx in Wyoming during the winters of 1999-2000, 2000-2001, 2001- 2002. During winter 1999-2000, we surveyed 1055 km in 32 sample units, and we detected one lynx track in the Salt River Range and a second track in the north- eastern portion of the Wyoming Range. In addition, tracks from a family group were observed in the north- eastern portion of the Wyoming Range by Wyoming Game and Fish Department biologists (B. Oakleaf, personal communication 2000). During winter 2000- 2001, we surveyed 1103 km in 37 sample units, and 4 lynx tracks were detected in the northeastern portion of the Wyoming Range, and one set in the southwestern THE CANADIAN FIELD-NATURALIST Vol. 118 © fo) Probability of detecting lynx oO (op) Number of visits FIGURE |. Computer-modeled relationship between the proba- bility of detecting lynx and the number of visits to an 8 km survey transect pixel relative to the number of days since last snow. Salt River Range. We surveyed 1080 km in 34 sample units during winter 2001-2002 and detected no lynx. Two tracks located during the 1999-2000 survey looked like lynx, but were poorly defined with each impression being a pedestal above the snow surface as a result of wind scouring. We followed two tracks for approximately 200 m until we located daybeds and collected five hairs. Two of the hairs contained suffi- cient DNA to amplify and the species identification was verified as lynx. Survey Verification The distribution of lynx in the Garnet Range, as de- fined by a 95% fixed kernel home range of all marked animals (n= 96 locations from 7 individuals, = 14 loca- tions per individual, Figure 2B) combined, subsumed 97% of all tracks detected during snow-track surveys. The general distribution of lynx in the Garnet Range as defined by radio telemetry and snow-track surveys was coincident (Figure 2A). We documented too few lynx detections in the Wy- oming and Salt River ranges to delineate a distribution based on snow-track surveys. However, biologists with the Wyoming Game and Fish Department trapped two lynx in the northeastern portion of the Wyoming Range during the winter of 1996-1997 (B. Oakleaf personal communication 2000). The winter home ranges of these two individuals were restricted to the northeastern por- tion of the Wyoming Range in the same approximate area as 5 of the 7 track detections from our survey. Discussion Snow-tracking has been used to document the pres- ence/absence of carnivores (Zielinski and Kucera 1995), and continues to be an important technique, especially when the organisms of interest are rare and difficult to observe. One primary appeal of using track-based surveys for presence/absence sampling is the method’s 2004 SQUIRES, MCKELVEY, and RUGGIERO: SNOW-TRACKING PROTOCOL 587 - # di a . Ke f j eS er! rae SS re \ i) a as / ; | oe | me, | XX —, ) - a 1 &Pevil Mn } 4% : q ; d ¥ fs f y © Lynx Telemetry Locations s Drummond es 95% Fixed Kernal Home Range << acs velude Grid 4 a Lynx Detections RASS Snow-track Surveys f ia = Lynx Distribution —_ Drummond Survey Grid i | | 95% Fixed Kernal Home Range (Telemetry - See Above) Sa a FicurE 2. The local distribution of lynx in the Garnet Range delineated from: (A) radio telemetry of all individuals com- bined (95% fixed kernal, n = 96 locations from 7 individuals, x = 13 locations per individual) compared to, (B) the local distribution based on sample units with track detections. 588 sensitivity to changes in use patterns, especially for species like lynx that are highly mobile and have large home ranges (Ward and Krebs 1985; Poole 1994; Slough and Mowat 1996; Aubry et al. 2000). Changes in lynx occupancy can be quickly detected across extensive areas using snow-track surveys. For example, general occurrence data from 1842-1998 suggested that lynx had a long-term history of occupancy in the Salt River and Wyoming Ranges (see McKelvey et al. 2000: Fig- ure 8.17), and the perception that lynx were currently distributed throughout the Wyoming and Salt River Ranges was shared by local managers. Snow-track sur- veys conducted in a representative manner throughout the Wyoming and Salt River Mountain ranges provided evidence that lynx were present in 1999, and this result was confirmed using genetic analyses in 2000. How- ever, the low overall detection rate coupled with the lack of tracks in 2001 indicated that lynx were rare and only located in a small area in the northern part of the Wyoming range. In areas with many track detections, snow-track sur- veys can yield more than simple presence/absence data. Results from the Garnet Range demonstrated that snow- track surveys conducted in a representative manner can also be used to delineate the local distribution of lynx. The high overlap in distributions determined through telemetry and survey data suggested that snow-track surveys can at least coarsely define the local distribu- tion of lynx (Figure 2). We acknowledge that survey data are more limited than telemetry data given that only winter movements are documented and the resulting distribution is coarse-scale relative to the sample unit. However, even a coarse-scale understanding of local distributions is useful to habitat managers. Thus, based on our data in the Garnet Range, we believe that snow- track surveys that are spatially well distributed across all available habitats can be used to estimate the local distribution of lynx during the winter. If lynx are abundant in an area, a 5% misidentification rate would have little impact on management decisions. However, in areas where lynx are very rare, a track misidentification can have serious management rami- fications. One way to address the chronic problem of track misidentification is to view the track as a “‘col- lection device” for obtaining genetic samples rather than a primary method of species identification. Genet- ic verification provides definitive species documentation, regardless of the observer’s ability to identify rare car- nivores based on track characteristics. We therefore rec- ommend that genetic identification be included when- ever proper identification of an individual set of tracks is critical. Snow-track surveys are labor intensive, and we have only used this method at a mid or “meso” spatial scale, where we searched all sample units. Such an intensive approach would be impractical across large landscapes, for example a large portion of a state. Although we have not conducted a snow-track based survey at a THE CANADIAN FIELD-NATURALIST Vol. 118 “state” scale, adaptive cluster sampling may provide a promising method of achieving a representative sam- ple across very large landscapes (Thompson 1992). Adaptive cluster sampling is effective for delineating clumped distributions, which we believe is the case for lynx at the southern extent of the species’ range (Aubry et al. 2000). Snow-track surveys are often conducted when track characteristics are most distinctive, usually within a day or two following fresh snowfall. This constraint limits the utility of snow tracking because the proba- bility of detecting lynx and other rare carnivores is a function of the time since last snow. The more time that lynx have to travel and establish long tracks, the more likely it is that their tracks will be detected on a survey route. For example, our simulations suggest almost a 100% probability of detecting lynx in two entries 7 days after a snowfall compared to a 43% probability of detection if the area was searched 1 day from snowfall (Figure 1). In the areas we surveyed, snow accumulations over 5 cm were recorded on 22 and 24 (17-18%) of the days for the Garnet and Wyoming ranges, respectively, and the snow-free periods between storms varied from 4-6 days (U.S.D.A., Nat- ural Resources Conservation Service, SNOTEL data). The ability to correctly identify “old” tracks by incor- porating genetic samples allows biologists to survey for rare carnivores when the probability of detection is high, even under poor tracking conditions. The winter surveys that we used to detect lynx in Wyoming and Montana had the following key ele- ments: (1) sample units were representative in terms of spatial coverage, but still focused on high-quality habitats that maximize lynx detections; (2) search effort per survey grid was defined and consistent across the survey area; (3) search intensity was consistent with simulated results indicating a high probability of detection, (4) technicians were well trained in track identification and data recording; (5) track identifica- tions were rigorously documented and spatially ref- erenced; and (6) lynx tracks were searched for genetic samples to confirm identification. We believe that sur- veys having these characteristics can be used to rigor- ously define the range of local lynx populations, and could be extended to the regional scale using adap- tive cluster sampling. Literature Cited Apps, C. D. 2000. 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Received 4 February 2004 Accepted 17 August 2004 Hummock Vegetation at the Arctic Tree-line near Churchill, Manitoba JORG TEWS Institute of Geography, University of Erlangen-Niirnberg, Kochstr. 4/4, 91054 Erlangen, Germany Current address: Geomatics and Landscape Ecology Research Laboratory (GLEL), Department of Biology, Carleton University, 1125 Colonel By Drive, Ottawa, Ontario K1S 5B6 Canada Tews, Jérg. 2004. Hummock vegetation at the arctic tree-line near Churchill, Manitoba, Canadian Field-Naturalist 118(4): 590-594. Hummocks, small earth or peat mounds, are widely distributed in the arctic and develop as a consequence of biomass accumulation and cryoturbation in the active layer. There is general agreement that the type of vegetation covering peat hummocks may alter the accumulation rate of organic material and thus hummock growth and local carbon sink dynamics. Studies on hummock plant community compositions from the arctic are very scarce. Here, I present results of a case study from the arctic tree-line near Churchill, Manitoba (Canada). Vegetation composition, hummock height and soil moisture content were recorded in 40 peat hummocks located along a tree-line gradient from open forest to tundra. Based on a cluster analysis I found three moss-dominated types of hummock vegetation, according to (1) a Tomenthypnum nitens (golden fuzzy fen moss) type on low hummocks, (2) a Hylocomium splendens (stair-step moss) type on medium-sized hummocks, and (3) a Pleurozium schreberi (red-stemmed feathermoss) type on hummocks higher than 60-70 cm. I found hummock height to increase towards the forest interior with decreasing water content of the upper organic layer on the hummock top. This is indicated by a significant change in vegetation composition towards drought resistant moss species on higher hummocks. Furthermore, species richness decreased with increase in hummock height. Based on evidence from historical tree-line invasion the overall results suggest that hammock height increases due to peat accumulation over the course of time resulting in a typical change in plant community composition. Key Words: peat hummocks, Tomenthypnum nitens, Golden Fuzzy Fen Moss, Hylocomium splendens, Stair-step Moss, Pleu- rozium schreberi, Red-stemmed Feathermoss, subarctic, forest-tundra ecotone, Hudson Bay Lowlands, Manitoba. Hummocks are small, up to 1 m high soil mounds widely distributed in the northern boreal, sub-arctic and arctic permafrost regions (e.g., Lundquist 1969; Mackay 1980). Generally, two types of hummocks can be classified: earth hummocks and peat hummocks. Earth hummocks may develop as a result of frost heave and cryoturbation processes where the organic layer overlies fine-grained frost-susceptible soils (Quinton et al. 2000). In contrast, peat hummocks grow as the result of the accumulation of organic material and where the surface of the uppermost mineral layer inside the hummock is positioned below the surrounding ground level (see Dredge 1992). Both hummock types may be perennially frozen or, as a result of warmer summer climate and lower latitude, completely thawed (Zoltai and Pettapiece 1974; Tarnocai and Zoltai 1978). As a transitional form they may be partially frozen with ice lenses remaining in the hummock core during the summer period. Even though hummocks are a common landscape feature and hummocky terrain covers a large proportion of Arctic Canada (Tarnocai and Zoltai 1978), scientific studies focusing on the vegetation structure of peat hummocks are very scarce. This lack of empirical data and also the potential role of peat hummocks for the carbon dynamics in the northern hemisphere empha- size the need to investigate the spatial distribution of major hummock plant community types. Besides the prevailing local climate, carbon storage in the arctic is strongly influenced by vegetation composition and succession (see Camill et al. 2001). Thus, in order to appreciate carbon dynamics in the northern hemisphere acknowledge of potential carbon sinks (such as peat hummocks) and their major plant community types are necessary. This enables evaluation of ecological factors which may alter accumulation or decomposition rates of organic material and related carbon dynamics. For example, mosses which often dominate hummock veg- etation have the potential to play a key role in modifying decomposition rates and the thermal and hydrological regime of arctic soils (Beringer et al. 2001). Here, I present results of a case study on vegetation composi- tion, succession and physiognomy of 40 peat hum- mocks located along an arctic tree-line transect from tundra to open forest in the Hudson Bay Lowlands near Churchill, Manitoba (Canada). Methods Study area The study site is situated at the open forest tree-line near Twin Lakes, a flat-topped glacial kame deposit approximately 25 km southeast of the town of Churchill (Figure 1). The open forest vegetation is com- posed of a mix of Tamarack (Larix laricina [Du Roi] K. Kock) and White Spruce (Picea glauca [Moench] Voss) with interspersed Black Spruce (Picea mariana [Mill.] Britt Sterns & Pogg). The present tree line north of Twin Lakes is extended into a wet sedge fen 590 2004 roy udson|Bayss S = es SiS apa ov Twin Lakes FiGuRE |: Location of the study site in the Hudson Bay Lowlands near Churchill in northeastern Manitoba (Canada). predominated by Carex aquatilis (Water Sedge) and Carex limosa (Mud Sedge). The current position of the tree-line has moved up to 150 m towards the open _ sedge fen within the last 70 years (see Scott et al. 1987). _ Here, the current tree-line is composed of L. laricina which established during the latest forest invasion. _ Whereas young L. laricina tree-line stands are char- acterized by small hummocks, the open forest interior towards Twin Lakes is dominated by mature P. glauca trees on typically hummocky terrain with large hum- _ mocks and inter-hummock, water-filled troughs. The nomenclature for plants follows after Porsild and Cody (1980). _ Sample design During July 1999 I established a 250 m transect from the open sedge fen towards the forest interior. I sam- pled each hummock (total 40) that occurred within a 5 m wide corridor along the transect line. For each hummock I measured its height from the top to the base. Vegetation composition was studied by using a _ frame placed on top of each hummock. The frame size 0.5 * 0.5 m was small enough to cover the tops of the smallest hummocks. For larger hummocks I chose homogeneous parts of the vegetation on the hammock top. Vascular plant cover within the sampling frame was then estimated for each plant species separately _ using the decimal Londo-scale (Londo 1984). Within each frame soil samples were taken from the upper organic layer (5 cm — 20 cm depth) during one sam- pling day and then water content gravimetrically deter- mined (samples were 24h oven-dried at 95°). Statistical analysis To group species datasets Mulva’s minimal variance clustering technique using van der Maarel’s coefficient was used (Wildi and Orléci 1996). Simple linear regres- sion was used to examine the relationship between hummock height as an independent variable and water content, species richness, and similarity of species com- TEWS: HUMMOCK VEGETATION AT THE ARCTIC TREE-LINE 59] position (species turnover rate) as the dependent vari- ables. For the species turnover rate between sample i and j hummocks were ranked by height and / defined according to: B= where / is the number of species that disappeared between sample i and j and g the number of new species. l+g i+] eqn |, Results Hummocks are a dominant micro-topographical fea- ture at the open forest tree-line. For the transect area I found a mean density of 320 hummocks ha’!. However, density was significantly higher in mature P. glauca stands towards the open forest. The majority of hum- mocks were predominately covered by either one of the moss species Tomenthypnum nitens (Golden Fuzzy Fen Moss), Hylocomium splendens (Stair-step Moss) or Pleurozium schreber (Red-stemmed Feathermoss) whereas herbaceous plants where less frequent. This was confirmed by a resemblance matrix of a cluster analysis for 40 sample plots (Figure 2). The Tomen- thypnum nitens-group was mainly composed of Poly- gonum viviparum (Alpine Bistort), Eguisetum varie- gatum (Variegated Scouring-rush), Andromeda polifolia (Dwarf Bog-rosemary), and Carex aquatilis (Water Sedge), indicating somewhat wet conditions (see Table 1). In contrast, hammocks with Hylocomium splendens had a more or less Dwarf Shrub dominated cover with species such as Vaccinium vitis-idaea (Lingonberry), Ledum groenlandicum (Common Labrador Tea), and the lichen Cladina rangiferina (Grey Reindeer Lichen). The third major group was dominated by Pleurozium schreberi associated with Betula glandulosa (Dwarf Birch) and the grass Calamagrostis canadensis (Blue Joint), indicating somewhat dry conditions. The species groups that were revealed by the clus- ter analysis showed significant affiliation in terms of transect position and respective hummock height. The Tomenthypnum nitens-group was mainly found on low hummocks (see Table 1) located near the tree-line (Fig- ure 3a). Here, the organic layer on hummock tops was mostly saturated (Figure 3b), typical for the hydro- logical situation near the sedge fen. Hummocks with Hylocomium splendens had medium-sized heights and an intermediate transect position, whereas Pleu- rozium schreberi-hummocks with heights above 60 cm where predominately found in the open forest interior. Overall, there was an increase in hummock height towards the open forest (Figure 3a, R? = 0.57). Increase in height had a significant negative effect on organic layer water content (Figure 3b, R*= 0.46). In terms of patterns of plant species richness, species number de- creased with increase in hummock height (Figure 3c, R? = 0.42; see also Table 1). Total number of plant species found, including mosses and lichens, was 45, and 8.1 species per sample plot on average. More- over, intermediate hummock heights were indicated 592 -4.66E-02 4.66E-01 9.79E-01 31 26 22 34 28 oa 27 Tomenthypnum 20 nitens - Gruppe 21 Pleurozium schreberi - Gruppe Hylocomium splendens - Gruppe a ee | ee eee] 9.79E-01 4.66E-01 -4.66E-02 FiGuRE 2: Cluster analysis of 40 hummock vegetation sam- ples. Three moss-dominated plant community types are distinctive according to the occurrence of Tomenthyp- num nitens, Hylocomium splendens and Pleurozium schreberi. Numbers for each sample indicate respec- tive hummock height given in cm. by a relatively high species turnover rate when plots were ranked by height (Figure 3d), i.e., species com- position showed a higher variation than vegetation of either low or high hummocks. Discussion The results of this field study from the arctic tree- line near Churchill indicate a significant relationship between hummock height and the position along the tree-line gradient on one hand, and hummock height and water content, vegetation type, species richness, THE CANADIAN FIELD-NATURALIST Vol. 118 and species turnover on the other hand. Increasing hummock height towards the forest interior seemed to reduce moisture availability for mosses on the hum- mock tops and facilitate the establishment of species- poorer communities with drought-resistant vascular plants. Interestingly, Zomenthypnum nitens, Hylocomi- um splendens and Pleurozium schreberi are ubiqui- tous moss species with a wide ecological distribution (Nicholson and Gignac 1995). However in this study they showed distinct distribution patterns in relation to hummock height and soil water availability within a relatively small area. The plant community composition of peat hammocks described here are the first inventory of hummock vegetation in the Hudson Bay Lowlands, the largest contiguous wetland in North America (Boudreau and Rouse 1995). Other published studies are concerned with the Mackenzie delta region where earth hummock physiology and plant species composition is com- pletely different. Thus, they are difficult to compare (see e.g., Zoltai and Pettapiece 1974). It is unclear whether peat hummocks accumulate organic material and increase in size regardless of the local environ- ment or whether this is driven by the micro-topogra- phy such as the establishment of trees. However, the local tree-line extension near Twin Lakes suggests that once young trees establish on formerly open tundra, peat hummocks may develop where shading and increased moisture from trapped snow coincide with feather moss establishment (Scott and Hansell 2002). Moreover, the occurrence of rotten tree stumps in the subsoil of the former centre of large, degraded hummocks (J. Tews personal observation) may support the latter hypothesis and is additional evidence that these hummock are formed by organic matter accumulation, not by cryo- genic processes as is the case with earth hummocks. Based on this study I hypothesize that hummock plant community composition at the tree-line near Twin Lakes is changing in the course of hummock growth. This is important as accumulation rates of peat (and thus carbon dynamics) are strongly influenced by local vegetation succession (Camill et al. 2001). Bello and D’Souza (2000) found that with increase in hum- mock height, accumulation rates of organic material decrease. For an average height of 20 cm they estimated an accumulation rate of 20 g m? year’, for 60 cm hum- mock height only 5 g m? year". These results largely confirm growth rates found for Tomenthypnum nitens and Hylocomium splendens (see Busby et al. 1978). In general, decomposition rates increase with decreas- ing water saturation during the short summer period. However, due to this rather small array of data, current available data may not support proper estimates of community-specific accumulation rates necessarty to model large-scale spatial and temporal carbon dynam- ics in the arctic. Thus, based on the knowledge of the predominant types of hummock plant communities, more studies on community-specific accumulation rates are needed. | H 2004 TEWS: HUMMOCK VEGETATION AT THE ARCTIC TREE-LINE 593 | Hummock height vs. gradient position Water content a) @ 120 b) 100,-——_____ == Ss ~ R“=0.462, p<0O. 5 80 a areal oO 3) Oo 80 2 60 ie SH Lges gs 4 40 E ro oe i) & 20 2 60 © e = 1 {0) x 50 0 ee Ale 0 20 40 60 80 100 120 Gradient [m] *f a Tuda ————— > Open forest ummock eight [cm] Species richness 8 — turnover rate oii @) 4s a R2= 0.4209, p < 0.€ rian R* = 0.2563, p < 0.C 8 oe FS 8 10 206 oa = = B04 ra | che ca 0.2 5 Z, 0 0 0 20 40 60 80 100 120 0 20 40 60 80 100 120 Hummock height [cm] Hummock height [cm] | Ficure 3: Simple regression analysis for: (3a) hummock height vs. gradient position along the tree-line from open tundra to forest; (3b) water content of upper humus layer on hummock top vs. hummock height; (3c) total number of plant species vs. hammock height; (3d) species turnover rate vs. hummock height. Another current environmental issue makes the mat- agreement that global climate change will alter decom- | ter even more complex: global climate change. On __ position rates and carbon storage (e.g., Gorham 1997; | average, climate change in the arctic may yield more _ Earle et al. 2003). Dormann and Woodin (2002) point- | precipitation and warmer summers (see Sonesson ed out that the driver of future change in arctic vege- | 2002). Based on these assumptions, there is general _ tation is likely to be increased nutrient availability, aris- Table 1: Typical species composition of hummocks at the arctic tree-line near Churchill, Manitoba. The table shows 27 characteristic hammock samples out of a total of 40 samples (13 samples with less significant community affiliation are not | shown). Three moss-dominated plant community types are evident according to Tomenthypnum nitens, Hylocomium splendens ) and Pleurozium schreberi. Dp ies BER Sy 6s ONO AP Aa AS 14y SANG 17; 16H 19 4 (20): 21) 22) 23) 242526 27 ‘Hummock height 21 19 26 28 34 31 25 20 25 37 22 22 27 98 47 98 73 77 61 10595 76 11277 84 67 80 # of species OMe Onno SSN Oron iO CON niOnn GR Gus GuSie ae fai nlem OMS) a9 hater Ouro _Tomenthypnum nitens Sei GUORMOeTOLOnaSrwtOe GiB bid OO Li29 Polygonum viviparum Moana Ae Onn Aw Aan Ql a2 | Equisetum variegatum Ana 2s AL 4 OE TAuL eit 4 °.2 4 | _Scirpus caespitosus ne 3 4 4 Carex aquatilis 1 1 _ Andromeda polifolia 2 1 Platanthera obtusata _Oxyecocus microcarpus Rubus acaulis _ Pedicularis lapponica Salix reticulata _ Aulacomnium palustre Hylocomium splendens i iS _Vaccinium vitis-idaea CEC, Bey Abi 40 Wig ln eg! Ledum groenlandicum 4.4 2: _Ledum decumbens 2 ES TY 1 _Cladina rangiferina. c4iuat i PY year sar Pleurozium schreberi (4 StS OS _ Betula glandulosa Calamagrostis canadensis io aA 594 ing for example from temperature-induced increases in mineralization. In particular, the response of plant growth to rising CO, levels appears to depend on nutri- ent availability (Heijmans et al. 2002). In the arctic vegetation nitrogen is tightly controlled by the moss layer (e.g., Li and Vitt 1997; Sommerkorn et al. 1999). Moreover, mosses such as Pleurozium schreberi have been reported to be able to fix nitrogen in symbiosis with a cyanobacterium (i.e. Nostoc sp.) (Deluca et al. 2002). Thus, increase in average summer temperatures due to climate change and the resulting increase in min- eralization rates may potentially decrease local carbon storage with severe consequences for the global carbon cycle. However, scientific research is far from estab- lishing realistic future scenarios. For example, in the context of our study, it was recently shown that indi- vidual bryophyte species displayed contrasting respons- es to changes in the nutrient supply and that they should not be grouped as a single functional type (Gordon et al. 2001). From this it is evident that further detailed hummock vegetation studies are needed, particularly in the arctic where the impact of climate change is likely to be most effective. Acknowledgments I thank the Churchill Northern Studies Centre and the Northern Research Fund (NRF) for logistical and financial support. Special thanks go to C. Oswald and P. Brown for their great help during the field work. I appreciate the constructive comments on this manu- script by two anonymous reviewers. Literature Cited Bello, R., and A. D’Souza. 2000. Half-century carbon accu- mulation rates in a Churchill, Manitoba Sedge Fen. In: Circumpolar Ecosystems IV, Churchill Northern Studies Centre, page 16. Beringer, J., A. H. Lynch, F. S. Chapin, M. Mack, and G. B. Bonan. 2001. The representation of arctic soils in the land surface model: The importance of mosses. Journal of Climate 14: 3324-3335. Boudreau, L. D., and W. R. Rouse.1995: The role of individ- ual terrain units in the water balance of wetland tundra. Climate Research 5: 31-47. Busby, J. R., L. C. Bliss, and C. D. Hamilton.1978. Micro- climate control of growth rates and habitats of the boreal forest mosses Thomenthypnum nitens and Hylocomium plendens. Ecological Monographs 48: 95-110. Camill, P., J. A. Lynch, J. S. Clark, J. B. Adams, and B. Jordan. 2001. Changes in biomass, aboveground net pri- mary production, and peat accumulation following per- mafrost thaw in the boreal peatlands of Manitoba, Canada. Ecosystems 4: 461-478. Deluca, T. H., O. Zackrisson, M. C. Nilsson, and A. Sellstedt. 2002. Quantifying nitrogen-fixation in feather moss car- pets of boreal forests. Nature 419: 917-920. Dorman, C. F., and S. L. Woodin. 2002. Climate change in the Arctic: using plant functional types in a meta-analy- sis of field experiments. Functional Ecology 16: 4-17. Dredge, L. A. 1992. Field guide to the Churchill Region, Manitoba. Geological Survey of Canada, Miscellaneous Report 53. THE CANADIAN FIELD-NATURALIST Vol. 118 Earle, L. R., B. G. Warner, and R. Aravena. 2003. Rapid development of an unusual peat-accumulating ecosystem in the Chilean Altiplano. Quaternary Research 59: 2-11. Gordon, C., J. M. Wynn, and S. J. Woodin. 2001. Impacts of increased nitrogen supply on high Arctic heath: the importance of bryophytes and phosphorus availability. New Phytologist 149: 461-471. Gorham, E. 1997. Northern peatlands: role in the carbon cycle and probable responses to climatic warming. Ecological Applications 1: 182-195. Heijmans, M. M. P. D., H. Klees, W. de Visser, and F. Berendse. 2002. Response of a Sphagnum bog plant community to elevated CO2 and N supply. Plant Ecology 162: 123-134. Li, Y. H., and D. H. Vitt. 1997. Patterns of retention and uti- lization of aerially deposited nitrogen in boreal peatlands. Ecoscience 4: 106-116. Londo, G. 1984. The decimal scale for releves of perma- nent quadrats. Pages 45-49 in Sampling methods and taxon analysis in vegetation science — Handbook of Vegetation Science 4. Edited by R. Knapp. Junk, The Hague, Neth- erlands. Lundquist, J. 1969. Earth and ice mounds: a terminological discussion. Pages 203-215 in The Periglacial Environment. Edited by T. L. Pewe. Montreal. Quebec. Mackay, J. R. 1980. The origin of hummocks, Western Arctic Coast, Canada. Canadian Journal of Earth Science 17: 996- 1006. Nicholson, B. J., and L. D. Gignac. 1995. Ecotope dimensions of peatland bryophyte indicator species along gradients in the Mackenzie River Basin, Canada. Bryologist 98: 437-451. Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Con- tinental Northwest Territories, Canada. National Museum of Natural Sciences, Ottawa. Quinton, W. L., D. M. Gray, and P. Marsh. 2000. Subsurface drainage from hummock-covered hillslopes in the Arctic tundra. Journal of Hydrology 237: 113-125. Scott, P. A., R. I. C. Hansell, and D. C. F. Fayle.1987. Establishment of white spruce population and responses to climatic change at the tree-line, Churchill, Manitoba, Canada. Arctic and Alpine Research 19: 45-51. Scott, P. A., and R. I. C. Hansell. 2002. Development of white spruce islands in the shrub zone of the forest-tun- dra. Arctic 55: 238-246. Sommerkorn, M., M. Bolter, and L. Kappen.1999. Car- bon dioxide fluxes of soils and mosses in wet tundra of Taimyr Peninsula, Siberia: controlling factors and contri- bution to net system fluxes. Polar Research 18: 253-260. Sonesson, M., B. A. Carlsson, T. V. Callaghan, S. Halling, L. O. Bjorn, M. Bertgren, and U. Johanson. 2002. Growth of two peat-forming mosses in subarctic mires: species interactions and effects of simulated climate change. Oikos 99: 151-160. Tarnocai, C., and S. C. Zoltai.1978. Earth hummocks of the Canadian arctic and subarctic. Arctic and Alpine Research 10: 581-594. Wildi, O., and L. Orléci.1996. Numerical Exploration of Community Patterns. SPB Academic Publishing, The Hague, 171 pages Zoltai, S. C., and W. W. Pettapiece. 1974. Tree distribution on perennially frozen earth hummocks. Arctic and Alpine Research 6: 403-411. Received 17 February 2004 Accepted 24 December 2004 Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus banksiana, Growth in Manitoba _ Brock Epp and JACQUES C. TARDIF'! | Centre for Forest Interdisciplinary Research (C-FIR) and Department of Biology, University of Winnipeg, 515 Avenue Portage, Winnipeg, Manitoba R3B 2E9 Canada ‘Corresponding author | Epp, Brock, and Jacques C. Tardif. 2004. Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus banksiana, growth in Manitoba. Canadian Field-Naturalist 118(4): 595-601. | The Lodgepole Pine Dwarf Mistletoe (Arceuthobium americanum Nutt. ex Engelm.) is an important pathogen of Jack Pine | (Pinus banksiana Lamb.). Dwarf Mistletoe alters tree form, suppresses growth, and reduces volume and overall wood quality of its host. Stem analysis and a 3-parameter logistic regression model were used to compare the growth of heavily and lightly to non infected Jack Pine trees. At the time of sampling, no significant reduction in diameter at breast height and basal area were observed in heavily infected trees. However, a significant reduction in height and volume and an increase in taper were observed in heavily infected trees. Growth models predicted a 21.1% lower basal area, 23.4% lower height and 42.1% lower volume by age 60 for the high infection group. Key Words: Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, Jack Pine, Pinus banksiana, stem analysis, logistic regression, basal area, height, volume, tree growth, productivity, Manitoba. Le Faux—gui du pin (Arceuthobium americanum Nutt. ex Engelm.) est un important agent pathogeéne du Pin gris (Pinus banksiana Lamb.). Le faux—gui modifie le défilement des arbres hotes, réduit leur croissance et leur volume marchand ainsi que la qualité du bois. L’analyse de tige et la régression logistique a trois-paramétres ont été utilisés afin de comparer |’accroissement entre les Pins Gris sévérement atteints et faiblement atteints. Au moment de |’ échantillonnage, aucune réduction significative du diamétre a hauteur de poitrine ou de la surface terriére ne fut observée chez le groupe sévérement atteint. Toutefois, les arbres atteints ont enregistré une baisse significative de la hauteur et du volume ainsi que du coefficient de défilement. Les modeéles de régression ont prédit une perte de 21,1% en surface terriére, de 23,4% en hauteur et de 42,1% en volume pour les pins gris fortement atteints par le faux—gui a un age de 60 ans. Mots clés: Le Faux—gui du pin, Arceuthobium americanum, Pin gris, Pinus banksiana Lamb., analyse de tige, régression logistique, surface terriére, hauteur, volume, croisssance, productivité, Manitoba. Jack Pine (Pinus banksiana Lamb.) is an important commercial tree species because of its pole-like growth form, and as a pioneer species, it is relatively easy to regenerate following harvest (Rudolph and Laidly 1990; Sims et al. 1990). An important pathogen of Jack Pine is Lodgepole Pine Dwarf Mistletoe (Arceuthobium americanum Nutt. ex Engelm.). There are five species of Dwarf Mistletoe known in Canada, of which A. americanum is the most widely distributed, extend- ing from British Columbia to southeastern Manitoba (Hawksworth and Wiens 1996). Individual species of Dwarf Mistletoe are generally host-specific, able to infect only a few tree species. In the case of A. amer- icanum, other susceptible hosts include Lodgepole Pine (Pinus contorta Doug}. ex Loud.) and Ponderosa Pine (Pinus ponderosa Dougl. ex P. & C. Laws.). As of 1996, there were 670 000 ha of forest with severe A. americanum infections in Alberta, Saskat- chewan and Manitoba (Brandt et al. 1998). In Mani- toba, approximately 8.7% of mature Jack Pine stands in important growing regions were infected by Dwarf Mistletoe (Baker et al. 1992). This has resulted in a loss of up to 7.9% of the total pine volume in these regions (525 224 m? out of 6 648 405 m°) and a loss of up to 70.3% within the infected stands. Aside from the sub- stantial volume losses, there is also a reduction in wood quality, making wood less merchantable. Wood of Lodgepole Pine infected by A. americanum exhibited greater longitudinal shrinkage and was weaker in strength when compared to uninfected trees because of the production of tree-rings having a lower percentage of latewood, shorter tracheid length, and higher resin content (Piirto et al. 1974). Generally, Dwarf Mistletoe deprives its host of water and nutrients, thus reducing height and diameter growth as well as seed production, and weakening the tree (Franc and Baker 2000). Dwarf Mistletoe alters tree form by disrupting apical domi- nance (Tinnin and Knutson 1980) through formation of witches’ brooms. Because Dwarf Mistletoe derives all of its nutrients from host tissue and fixes little or no carbon dioxide for its own use, witches’ brooms act as sinks for metabolites produced in other parts of the host (Hull and Leonard 1964a, 1964b). Douglas-fir and Western Larch infected by A. douglasii and A. laricis, respectively, exhibited a significant increase in leaf to sapwood ratios in heavily infected trees, altering resource 395 596 allocation processes and reducing the overall water- use efficiencies (Sala et al. 2001). The long incubation period characterized by the life cycle of A. americanum (Hawksworth and Wiens 1996) poses a problem in the early detection of initial infections. In forest management situations, the current most effective methods of treatment are removal of heavily infected trees and isolation of infected stands by planting buffer zones of incompatible host species (Franc and Baker 2000). Even-aged silvicultural systems are often effective in controlling Dwarf Mistletoe, as the entire overstory may be removed. No one has quantified volume losses on individual Jack Pine trees due to infection by Dwarf Mistletoe. One study examined the effect of A. americanum on the growth of Lodgepole Pine (Baranyay and Safranyik 1970). Within Manitoba, Baker et al. (1992) studied the impact of Dwarf Mistletoe on Jack Pine trees at the stand scale, focusing on loss of wood volume in the forest stands. The objective of this study was to com- pare the growth of heavily infected Jack Pine trees to that of non-infected or lightly infected Jack Pine trees based on differences in diameter at breast height (dbh), basal area, height, volume, and stem form. We hypoth- esized that there was a significant reduction in total cumulative dbh, basal area, height, and volume growth rate in trees infected with Dwarf Mistletoe. Methods The sampling area is located about 97 km north- east of Winnipeg, Manitoba (Figure |) within Belair Provincial Forest (50°38'N, 96°29'W, 250 meters above sea level). The underlying bedrock consists of Archean granites and gneisses (Manitoba Geological Survey 2002*). The closest meteorological station is located at Pine Falls about 21 km from the sampling area. The climate is continental (Burton et al. 1998). The mean annual temperature for Pine Falls is 2.1°C, and the total annual precipitation is 538.5 mm (Envi- ronment Canada 2002*). In winters, the average min- imum temperature reaches —23.5°C in January, and in summers, the average maximum temperature reaches 25.3°C in July. The sampling site consists of a small, open, fire- originated Jack Pine stand, and is bound by Provincial Highway 11 to the south and a recent cutover to the north. The sample site is located near the south-eastern limit of the lodgepole pine dwarf mistletoe geograph- ical distribution in Manitoba (Baker et al. 1992). Physically, the site is characterized by level, homoge- neous terrain. The site is also homogeneous with regards to stand origin, slope variation and understory vege- tation. Due to the small size of the infected stand, and the localized nature of the mistletoe infections, the number of trees for sampling was limited. There are epicentres of Dwarf Mistletoe infections scattered throughout the area. Within the sampling site, there is a relatively even scattering of heavily infected trees THE CANADIAN FIELD-NATURALIST Vol. 118 intermixed with lightly and non-infected trees (Fig- ure 1). Three transects were established approximately 50 m apart, oriented in a north-south direction, extending north from Provincial Hwy. 11. Ten sampling points were randomly established along the three transects, and the surrounding area was divided into four quad- rants. One of the four quadrants was randomly select- ed in which the two nearest trees to the point were select- ed; one heavily infected tree and the nearest lightly or non-infected tree between 40 and 60 years of age. Trees with noticeable fire scars, fungal infections, or injuries were avoided to minimize the effects of other disturbances on tree growth. Trees that exhibited exces- sive branching or forking on the main stem were also avoided to simplify stem analysis. Trees were selected up to 50 m from plot centre, as long as they remained within the stand boundaries. If no trees matching our criteria were found within the selected quadrant, sam- pling was done in the next quadrant in numerical order. This method helped to eliminate bias by selecting the closest trees to the sampling points. Once candidate trees were selected, they were numbered, marked with a north-orientation line, marked at 0.5 m and 1.3 m, and locations were recorded with a GPS unit (Figure 1). Sampling took place in late July — early August 2002. Prior to felling, the infestation index of the trees was estimated using the Hawksworth 6-class system (Hawksworth 1977). Infection was most easily deter- mined from the ground by the presence of witches’ brooms. Trees with an infestation index of 0 to 3 were classified as lightly infected, whereas trees with an index of 4 to 6 were classified as moderately to heavily infected. In order to take into account the competition of other trees, the dbh and distance of the nearest tree in each cardinal direction were measured. Following felling of the trees, tree height was record- ed and stems were sectioned at 0 m, 0.5 m, 1.3 m, and then 1-m intervals until stem diameter was less than 1 cm. Sections at points of the stem where branches emerged were avoided, and the adjusted height was recorded. For each cross-section, diameter inside-bark and outside-bark were measured along two diameters perpendicular to each other. At the laboratory, age of each cross-section was measured, and the pointer-year method of cross-dating was used to validate ages (Yam- aguchi 1991). Following dating, stem analysis was per- formed on each tree to determine annual increments. Ring width measurements were done using the Win- DENDRO™ y. 2002a program (Régent Instruments Inc. 2002*), and annual increments were calculated us- ing XLSTEM™ 1.3a (Régent Instruments Inc. 1999*). Each cross-section was scanned with a high-resolution scanner and saved as a digital image. An image reso- lution of 800 dpi was used, except for cross-sections exhibiting high levels of suppression, where a resolu- tion of 1600 dpi was used. For one tree, suppression was too great for ring detection by the WinDENDRO™ Sast atchewan @ Winnipeg 100° 96° g2° Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 597] 0 Low-infection O High-infection 50 Meters FicurE 1: Location of the study area and distribution of the sampled trees. The lower right panel illustrates the relative posi- tion of the sampled trees and their infection level in the jack pine stand. Empty circles correspond to heavily infected trees, and empty squares correspond to lightly or non-infected trees. program, so measurements were performed manually at a magnification of 50x using a Velmex measuring stage to a precision of 0.001 mm. Ring widths were measured, starting at the pith, along four radial paths in the north, east, south and west directions. Data obtained were examined for differences between the two infection groups with regards to age, competi- tion index, dbh, basal area, height, volume and stem form. The competition index was modified from Hegyi’s distance weighted size ratio index (Avery and Burk- hart 2002): i Sy DIST, jal where C/I; is the competition index for the subject tree, D; is the dbh of the jth of four competitor trees (N, E, S, and W directions), D, is the dbh of the subject tree, and DIST, is the distance between the subject tree and the jth competitor tree. Basal area at dbh was calculated for each tree by using the formula for the area of a circle. Tree volume without bark was calcu- lated by applying Smalian’s formula for volume (Avery and Burkhart 2002) to each segment. The volume of each segment was then added to obtain tree volume. Stem form expressions were determined using the Girard form class (Avery and Burkhart 2002), with the quadratic mean diameter taken at the section nearest 5 metres. Because each group had a relatively low sample size (N=10), the non-parametric Mann-Whitney U-test was performed on each variable to determine significant differences between samples from each infection group (a = 0.05). It should be noted that the Mann-Whitney U-test and the Wilcoxon two-samples test yield the same statistic and give the same results (Sokal and Rohlf 1997). To limit the effect of environ- mental variability, lightly or non-infected and heavily- infected trees were also carefully matched in pairs based on the distance separating them (Figure 1). Nine pairs were formed with a mean distance of 9.49 m (standard deviation = 4.83 m) between lightly or non-infected and heavily-infected trees. A Wilcoxon signed-rank test (Sokal and Rohlf 1997) was used to test for sig- nificant differences between infection groups. Paired- sample tests are more powerful than independent- sample tests. The cumulative dbh, basal area, height and volume average growth curves produced for each tree were compared between infection groups. First, predicted growth curves were modelled for each growth variable and tree using non-linear regression. Best fit to actual cumulative growth curves for each variable and tree were obtained using a three-parameter logistic model of the form: a SD 1+ ext ] where y is the value of the subject growth variable, x is the year of growth, a is the asymptotic level of 598 growth, b is the growth constant, and xp is the inflection point. Because each of these three parameters defines the shape of the growth curves, significant differences between parameters for the two infection groups were determined using the Mann-Whitney U-test (a = 0.05). This was repeated for each growth variable. Second, the best three-parameter logistic model was determined for each variable and infection group using the 10 trees as replicates. The resulting predicted growth curves were plotted to age 60. Because trees were not all of the same age and the number of observations decreased with time, only data covering the first 45 years of growth were used in all the regression analyses. Results The two infection groups showed no significant dif- ference in age, competition index, dbh, and basal area (Table 1). There was, however, a significant difference in height and form class (Table 1). Specifically, trees in the high infection group were 20.2% shorter and THE CANADIAN FIELD-NATURALIST Vol. 118 had more stem taper. Both independent- and paired- sample tests yielded divergent results regarding total volume, and when partially controlling for environmen- tal variability no significant difference in total volume was observed as indicated by the Wilcoxon signed-rank test. The analysis of the coefficients from the logistic regressions derived from each tree in each infection group revealed that the high infection group reached a significantly lower maximum basal area, height, and volume, as predicted by the asymptotic values of the regression model (Table 2). Maximum dbh did not dif- fer significantly between groups. The logistic regres- sion model, projected to age 60, predicts a 21.1% lower basal area (Figure 2B), a 23.4% lower height (Figure 3A), and a 42.1% lower volume (Figure 3B) for the high infection group. Results showed that there was a significantly higher growth rate for dbh and basal area in the high infection group (Table 2). This was also observed for volume but TABLE |: General characteristics of heavily infected and non-infected or lightly infected trees (n = 10 for each group) at time of sampling (Max = Maximum value, Min = Minimum value, SEM = Standard Error of the Mean). The first column of probabilities (P!) is based on the Mann-Whitney U-test (n=10 for each group). The second column of probabilities (P?) is based on the Wilcoxon signed-rank test (n=9 pairs). Low infection High infection Max Min Mean SEM Max Min Mean SEM P! Pp? Infestation index 2.0 0.0 12 0.29 6.0 4.0 2 0.20 0.000 0.007 Tree age 48.0 42.0 46.3 0.62 82.0 44.0 50.0 3.59 O28 022: Competition index 3.34 0.77 1.85 0.30 1.89 0.66 1.29 0.15 0257) .O!515: Diameter at breast height (cm) 19205) W310 od 0.38 L960 SES ON eliS 45 0.83 0.544 0.515 Basal area (m7?) 0:022> O!012. ~0:017;- 0:00 0.024 0.007 0.016 0.00 0.544 0.594 Height (m) 14.10 10.90 12.65 0.39 L330 830 ea lORO 4s 3054: 0.004 0.008 Volume (m?) 0.16 0.09 0.12 0.01 0.17 0.04 0.08 0.01 0.019 0.110 Form class 0.83 0.64 0.73 0.02 0:76" 10335 0.56 0.04 0.004 0.008 TABLE 2. Values of the three parameters of the predicted logistic growth curves used to model diameter at breast height, basal area, height, and volume for heavily infected and non-infected or lightly infected Jack Pine trees (n = 10 for each group). The three parameters are the theoretic maximum (a), growth (b), and the inflection point (x9) (Max = Maximum value, Min = Minimum value, SEM = Standard Error of the Mean). The first column of probabilities (P') is based on the Mann-Whit- ney U-test (n=10 for each group). The second column of probabilities (P?) is based on the Wilcoxon signed-rank test (n=9 pairs). Low infection group High infection group Max Min Mean SEM Max Min Mean SEM P! pz Diameter at breast height (cm) a 22: OS na WLANs lif2,9 0.85 QO0:97 18287 2 18.97 PAIL 0.059 0.260 b — 84) '-3230) 1 <2:46%) 10M3 -2.46 -4.29 -3.26 0.19 0.005 0.015 Xo 36:220 23133) 28:46 1.50 Qe oie MOD3ie 524 1.19 0.023 0.028 Basal area (m7) a 0.071 0.019 0.038 0.0061 0.032 0.007 0.016 0.0026 0.002 0.038 b -2.32 -3.51 =O Ams ORS -3.07 -5.27 -3.84 0.20 0.001 0.008 Xo OB Man) S464) 5538/8 Soe 39.18 24.84 31.59 1.72 0.001 0.011 Height (m) a 24.37 14.99 19.59 1.01 20.63 9.28 13.90 1.2] 0.008 0.015 b -1.71 -2.64 -1.98 0.09 -1.44 -2.65 -2.07 0.12 0.326 0.374 Xo AD OI 24938 = 32°66 1.74 49.34 20.56 27.58 DATA 0.034 0.038 Volume (m?) a 2237 2 OM2OF 1Or604s O97, 0.214 0.042 0.097 0.021 0.001 0.015 b -2.98 -4.36 -3.60 0.13 -3.12 -6.07 -4.09 0.25 0.082 0.028 Xo 111.79 40.48 60.48 6.51 48.74 30.68 36.59 1.97 0.001 0.011 2004 P<0.0001 16 y = 16.6392/[1+(x/27.2775) 242" adj. ° = 0.96 aaa, P<0.0001 eso 5 e = a4 + : . y = 13,8979/{1+(x/23.3196)?°%) iw adj. r = 0.88 3 P<0.0001 5 6 5 4 (S) 2 0 0.020 y = 0.0274/[1+(x/45.0484)22743) B) adj. r = j. F = 0.92 P <0.0001 Wa ~ 0.015 Vy , nN | i os © Pe ® | | © 0.010 gf g q © # jaa) ¥ y = 0.0170/[1+(x/33.1868) 74767] 0.005 # SAE ey 0 10 20 30 40 50 60 Tree age FIGURE 2: Average cumulative dbh growth (A) and average cumulative basal area growth (B) for each of the two infec- tion groups. Dashed lines represent predicted growth up to year 60 for the infection groups, based on the 3-parameter logistic regression models depicted. Vertical bars represent standard errors of the mean. The black dashed line and empty circles indicates the low infection group, and the dark grey dashed line and empty triangles indicates the high infection group. only after the Wilcoxon signed-rank test. It should be noted that in a logistic regression, there is an inverse relationship between the growth constant and the actual growth rate. The significantly lower growth constant observed in the high infection group indicates a faster rate of growth. No significant difference in the growth rate for height was observed. However, all four cumu- lative growth variables of the high infection group reached their inflection point significantly sooner than the low infection group (Table 2), suggesting that the high infection group reached its maximum growth rate sooner. Dbh (Figure 2A) and basal area (Figure 2B) of the two infection groups were not predicted to diverge until after 45 years of age, while height growth curves began to diverge at about 20 years of age (Fig- ure 3A), and volume growth curves began to diverge at 35 to 40 years of age (Figure 3B). Results indicate a higher level of variability in the basal area and volume of the high-infection group when compared to the low-infection group, as indicated by the lower value of the adjusted 1 for each logistic regres- sion model. This may indicate variation in environmen- tal conditions at early stages of growth for the trees, or Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 599 16 y = 18.4039/(1+(x/30.4334) 1%") 14 A) a0 f 087 iS , 12 E en: ‘er Be antl 5 : agp a 6 fi Fy = 13.2629/(1+(x/25.9869)29"% oH r=0.92 il Bsc 4 os 24 a 0 es geet 0.16 ] B) y = 0.2048/[1+(x/46.9484)? 7] 0.14 | adh O82 s 0.12 i “E 0.10 x 2 0.08 4 § ee S 0.06 0.04 ft 0.02 sf Pi y = 0.101 1/[1+(x/38.4295)?7""4 0.00 soon? PE OOOR | 0 10 20 30 40 me 2 Tree age FIGURE 3: Average cumulative height growth (A) and average cumulative volume growth (B) for each of the two infection groups. Dashed lines represent predicted growth up to year 60 for the infection groups, based on the 3-parameter logis- tic regression models depicted. Vertical bars represent stan- dard errors of the mean. The black dashed line and empty circles indicates the low infection group, and the dark grey dashed line and empty triangles indicates the high infec- tion group. variation in genetic resistance to mistletoe infection. Examination of the stem profiles also revealed that some trees in the high-infection group exhibited little or no evidence of growth reduction despite a high infection index (not presented). These trees could have been recently infected by Dwarf Mistletoe. Discussion No significant differences in tree age or competition levels between the two infections groups were observed indicating that Dwarf Misteltoe was was the key factor affecting Jack Pine growth. The lack of a significant difference in dbh between the heavily infected and lightly infected Jack Pine trees at the time of sampling likely reflects the young age of the subject trees and the duration of the infection period. Tinnin et al. (1999) found that there was a significant reduction in the diameter growth of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) heavily infected by A. douglasii (infection index = 5 and 6) when compared to non-infected trees (infection index = 0) for trees hav- ing mean ages between 78 and 84 years. In Lodgepole Pine, diameter growth was not significantly affected 600 at stump height, but significant decreases in diameter growth were apparent at heights further up the stem because of greater stem taper (Baranyay and Safranyik 1970). Our study also showed that stem taper was sig- nificantly more pronounced in the high infection group. At the time of sampling, basal area in the high infec- tion group was not significantly different than that of the low infection group. Comparison of the logistic regression coefficients provided similar results to dbh, except that the high infection group was predicted to reach a significantly lower basal area compared to the low infection group as trees continue to grow after age 45. Assuming that basal area increment after the age of 45 years will continue to follow that predicted by the logistic regression model, the projected maximum basal area of the high infection group was approxi- mately 57.9% lower than that of the low infection group. Pierce (1960) also found a 68.5% reduction in basal area for heavily infected Douglas-fir, and a 41.0% reduction in moderately infected Douglas-fir. Of the significantly affected growth variables, height increment showed a reduction first, i.e., between the ages of 20 and 25 years. Pierce (1960) also found that differences in height were statistically significant be- tween all infection classes, in contrast to diameter. This corresponds with the highly significant 20.2% lower tree height in heavily infected trees in this study. The greater sensitivity of this variable is likely due to the initial response of trees to Dwarf Mistletoe infections. Dwarf Mistletoe infected trees accumulate a high level of biomass in their brooms, which detracts the allo- cation of resources from biomass production elsewhere in the tree (Tinnin and Knutson 1980). In contrast to the other growth variables, height of the high-infection group had similar variability to the low-infection group, as indicated by similar adjusted r values in the logistic regression curves. This suggests that height growth is less sensitive to temporal changes in factors such as stand density and competition (Avery and Burkhart 2002). Height growth diverged between the two infec- tion groups at a much earlier age than volume, diame- ter, or basal area, further suggesting that height growth begins to show significant decreases shortly after the development of brooms and loss of apical dominance (Tinnin and Knutson 1980). At the time of sampling, there was a 33.3% lower average total volume in the high infection group, which was mostly due to a significant reduction in tree height. However, despite early reduction in height growth, volume growth of the heavily infected trees was not significantly affected until 30 to 40 years of age. Our values very closely correspond with those reported by Baranyay and Safranyik (1970), where a group of infected Lodgepole Pine trees with an average age of 37 showed a 35.5% reduction in volume. From the logistic growth model, it was found that the predicted maximum average volume of the high infection group was 83.9% lower than the low infection group. This THE CANADIAN FIELD-NATURALIST Vol. 118 projected value exceeded the estimated volume reduc- tion in infected stands made by Baker et al. (1992), which was between 53.4% and 70.3% depending on the level of potential crown closure. However, it should be noted that our findings reflected the volume loss in individual trees, and not the volume loss as averaged through an entire stand. In conclusion, this study confirmed the significance of the impact of Dwarf Mistletoe on the growth of com- mercial Jack Pine forests, and the potential economic loss due to severe infection. Despite early reduction in height growth, volume growth of the heavily infected trees was not significantly affected until 30 to 40 years of age. Further study on the effects of Dwarf Mistletoe over a wider range of age classes and site indexes would be beneficial in modelling the impact of the parasite. Acknowledgements This project was part of an undergraduate honours thesis course at the University of Winnipeg, conducted by the first author. We thank the following people for their contributions: F. Conciatori, G. Sayer, R. Moodie, R. Westwood, D. Ko Heinrichs, the University of Win- nipeg Department of Biology, and the Centre for For- est Interdisciplinary Research. Special thanks go to K. Knowles from Manitoba Conservation and R. Staniforth for their comments on a previous draft of this paper. Documents Cited (marked * in text) Environment Canada. 2002. Canadian climate normals 1971-2000. 3 March 2003. Manitoba Geological Survey. 2002. Manitoba geology. Mani- toba Industry, Trade and Mines. 27 March 2003. Régent Instruments Inc. 1999. XLSTEM™ 1.3a. Québec, Quebec. Régent Instruments Inc. 2002. WinDENDRO™ vy. 2002a. Québec, Quebec. Literature Cited Avery, T. E., and H. E. Burkhart. 2002. Forest measure- ments. Fifth edition. Mcgraw-Hill. New York. 456 pages. Baker, F. A., M. Slivitsky, and K. Knowles. 1992. Impact of dwarf mistletoe on jack pine forests in Manitoba. Plant Disease 76: 1256-1259. Baranyay, J. A., and L. Safranyik. 1970. Effect of dwarf mistletoe on growth and mortality of lodgepole pine in Alberta. Canadian Forestry Service. Department of Fish- eries and Forestry. Publication Number 1285. Brandt, J. P., R. D. Brett, K. R. Knowles, and A. Sproule. 1998. Distribution of severe dwarf mistletoe damage in west-central Canada. Natural Resources Canada, Canadian Forestry Service, Northern Forestry Centre, Edmonton, Alberta. Special Report 13. Burton, V. B., D. R. Zak, S. R. Denton, and S. H. Spurr. 1998. Forest ecology, Fourth edition. John Wiley & Sons Inc. New York. 774 pages. Franc, G. D., and FE. A. Baker. 2000. Dwarf mistletoes affect- ing pines and firs of Wyoming. University of Wyoming and Utah State University. Cooperative Extension Service. B- 1083. 2004 Hawksworth, F. G. 1977. The 6-class dwarf mistletoe rating system. USDA Forest Service, General Technical Report RM-48. Hawksworth, F. G., and D. Wiens. 1996. Dwarf mistletoes: biology, pathology, and systematics. Agricultural Handbook Number 709. USDA Forest Service. Hull, R. J., and O. A. Leonard. 1964a. Physiological aspects of parasitism in mistletoes (Arceuthobium and Phoraden- dron). 1. The carbohydrate nutrition of mistletoe. Plant Physiology 39: 996-1007. Hull, R. J., and O. A. Leonard. 1964b. Physiological aspects of parasitism in mistletoes (Arceuthobium and Phoraden- dron). Ul. The photosynthetic capacity of mistletoe. Plant Physiology 39: 1008-1017. Pierce, W. R. 1960. Dwarf mistletoe and its effect upon the larch and Douglas-fir of western Montana. Montana State University. School of Forestry. Missoula, Montana. Bul- letin Number 10. 37 pages. Piirto, D. D., D. L. Crews, and H. E. Troxell. 1974. The effects of dwarf mistletoe on the wood properties of lodge- pole pine. Wood and Fiber 6: 26-35. Rudolph, T. D., and P. R. Laidly. 1990. Pinus banksiana Jack Pine. Jn Silvics of North America: |. Conifers; 2. Hard- woods. Agriculture Handbook 654. U.S. Department of Agriculture, Forest Service, Washington, DC. Volume 2. Technical coordinators, Burns, R. M., and B. H. Honkala. 877 pages. Epp AND TARDIF: EFFECS OF LODGEPOLE PINE DWARF MISTLETOE 60) Sala, A., E. V. Carey, and R. M. Callaway. 2001. Dwarf mistletoe affects whole-tree water relations of Douglas fir and western larch primarily through changes in leaf to sapwood ratios. Oecologia 126: 42-52. Sims, R. A., H. M. Kershaw, and G. M. Wickware. 1990. The autecology of major tree species in the north central region of Ontario. Technical Report 48, Northwestern Ontario Forest Technology Development Unit, Ontario Ministry of Natural Resources, Thunder Bay, Ontario. 126 pages. Sokal, R. R., and F. J. Rohlf. 1997. Biometry. W.H. Free- man and Company, New York. Tinnin, R. O., and D. M. Knutson. 1980. Growth charac- teristics of the brooms on Douglas-fir caused by Arceutho- bium douglasii. Forest Science 26: 149-158. Tinnin, R. O., C. G. Parks, and D. M. Knutson. 1999. Effects of Douglas-fir dwarf mistletoe on trees in thinned stands in the pacific northwest. Forest Science 45(3): 359-365. Yamaguchi, D. K. 1991. A simple method for cross-dating increment cores from living trees. Canadian Journal of Forest Research 21: 414-416. Received 10 February 2004 Accepted 24 December 2004 Notes Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus Evan S. RICHARDSON! and RYAN K. BROOK? ‘Canadian Wildlife Service, 5320-122 St., Edmonton, Alberta T6H 3S5 Canada Faculty of Environment, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada Richardson, Evan S., and Ryan K. Brook. 2004. Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus. Canadian Field-Naturalist 118(4): 602-603. We observed a Polar Bear (Ursus maritimus) excavating an Arctic Fox (Alopex lagopus) den on 24 June 1998, 3 km inland from the Hudson Bay coast (58°40'N, 93°12'W), near Cape Churchill in Wapusk National Park, Manitoba. To our knowledge this is the first observed excavation of an Arctic Fox den by a Polar Bear. Key Words: Arctic Fox, Alopex lagopus, Polar Bear, Ursus maritimus, den, behaviour, Hudson Bay, Wapusk National Park, Manitoba. The western Hudson Bay Polar Bear (Ursus mar- itimus) population remains on shore from approximate- ly mid-July through early November due to complete annual melting of the sea ice (Stirling et al. 1977). During this period, bears mainly remain relatively inactive (Knudsen 1978; Latour 1981; Lunn and Stir- ling 1985) and subsist primarily on stored fat reserves (Nelson et al. 1983; Derocher et al. 1990; Ramsay et al. 1991). However, during the summer months, polar bears are opportunistic and occasionally feed on a variety of terrestrial food sources including grasses, sedges and berries (Knudsen 1978; Lunn and Stirling 1985, Derocher et al. 1993), Thick-billed Murres (Uria lomvia) (Donaldson et al. 1995), Canada Geese (Bran- ta canadensis) (Russell 1975) and their eggs (Smith and Hill 1996), Snow Goose eggs (Anser caerulescens) (Abraham et al. 1977), Willow Ptarmigan (Lagopus lagopus) (Miller and Woolridge 1983), seabirds (Stemp- niewicz 1993), microtine rodents (Russell 1975; Miller and Woolridge 1983), and Caribou (Rangifer tarandus) remains (Brook and Richardson 2002). We observed an adult male Polar Bear excavate an Arctic Fox den near Nestor | field camp (58°40'N, 93°12'W), near Cape Churchill in Wapusk National Park, Manitoba on 24 June 1998 at 11:00 CST. The bear was initially sighted on a large beach ridge inves- tigating one of the camp buildings and then moved to an Arctic Fox den located approximately 350 m south- east of camp. The bear moved around the site, inves- tigated several of the den entrances and then stopped at one entrance and began to excavate the den. The bear stopped digging every 10 —15 seconds to put its head into the enlarged entrance, continued this behav- iour for approximately 3 minutes, then stopped dig- ging and spent approximately 2 minutes investigating several other den entrances, but did not dig them out. The bear then moved away from the den and down the eastern side of the beach ridge, where it could no longer be observed. The bear was not observed eating anything, although while its head was in the entrance, it could not be observed. Several fox pups were occu- pying the den at the time and we assumed the pups were in the den during the observation although the location of the vixen at the time was unknown. We visited the following day and found a large pit dug by the bear. The excavation was 55 cm at its deepest and was approximately 70 cm wide and 80 cm in length. Arctic Fox pups could still be heard inside the den confirming that the den had not been abandoned and there was no evidence any fox pups were killed. The top of the den was littered with Canada and Snow Goose remains, consisting mostly of feet (98) and other waterfowl remains such as bones and feathers. Cached prey items inside the den were visible from several den entrances and consisted primarily of goose remains. Food remains found at fox dens in the region included both adult and juvenile geese, ducks, Caribou remains and in one instance, a Muskrat (Ondatra zibethicus) (Richardson and Brook, personal observations, 1999). Polar Bears are known to excavate subnivean lairs in search of young Ringed Seal (Phoca hispida) pups (Stirling and Archibald 1977). They also excavate earth and snow dens for reproduction, as well as open pits for resting (Clark 1996; Clark et al. 1997). Although adult male bears are known to excavate open pits on coastal beach ridges (Clark 1996), we suggest that the bear was most probably trying to gain access to a poten- tial food source. The bear may have been attracted to the fox den for several reasons. In areas where prey is abundant, Arctic Foxes cache large quantities of food at den sites for later consumption (Sklepkovych and 602 2004 Montevecchi 1996; Garrott et al. 1984). As a result, other predators may be attracted to den sites by their smell (Prestrud 1992). At active den sites, character- | istic barks of arctic fox pups can be heard from with- _ in the den when it is disturbed (Eberhardt et al. 1983; Richardson and Brook, personal observations, 1999). Bears are adept at locating food sources by smell (Lunn and Stirling 1985) and it seems unlikely that barking attracted the bear to the site, but the sound may have stimulated further investigation. Prey remains at den _ sites may provide a direct energy source for bears, how- | ever the energetic cost of excavating a den would not likely be repaid in the capture of a small Arctic Fox pup. Although, Macpherson (1969) notes that Arctic Fox dens may be occasionally excavated by Grizzly Bears | (Ursus arctos) and Wolves (Canis lupus), we are not aware of any other published reports of Polar Bears | excavating Arctic Fox dens. | Acknowledgments These observations were made while conducting research supported by Wapusk National Park, the Manitoba Department of Conservation, the Churchill _ Northern Studies Centre, the Northern Studies Training _ Program of the Department of Indian and Northern Affairs Canada, the Canadian Department of Fisheries and Oceans, the Western Canada Service Centre of _ Parks Canada, Wat’chee Lodge, the Canadian Wildlife Federation, and the Manitoba Chapter of the Wildlife Society. We would like to thank I. Stirling and N. J. Lunn for their valuable comments on the manuscript. Literature Cited Abraham, K. F., P. Mineau, and F. Cooke. 1977. Unusual predators of snow goose eggs. Canadian Field-Naturalist 91: 317-318. Brook, R. K., and E. S. Richardson. 2002. Observations of polar bear, Ursus maritimus, predatory behavior toward caribou, Rangifer tarandus, in Wapusk National Park, Manitoba. Arctic 55: 193-196. Clark, D. A. 1996. Terrestrial habitat selection by Polar Bears (Ursus maritimus Phipps) in the Western Hudson Bay Lowlands. M.Sc. thesis, University of Alberta, Edmonton, Alberta. Clark, D. A., I. Stirling, and W. Calvert. 1997. Distribution, characteristics and use of earth dens and related excavations by polar bears on the western Hudson Bay lowlands. Arctic 50: 158-166. Derocher, A. E., R. A. Nelson, I. Stirling, and M. A. Ram- say. 1990. Effects of fasting and feeding on serum urea and serum creatine levels in polar bears. Marine Mammal Science 6: 196-203. Derocher, A. E., D. Andriashek, and I. Stirling. 1993. Ter- restrial foraging by polar bears during the ice-free period in western Hudson Bay. Arctic 46: 251-254. Donaldson, G. M., G. Chapdelaine, and J. D. Andrews. 1995S. Predation of thick-billed murres, Uria lomvia, at two NOTES 603 breeding colonies by polar bears, Ursus maritimus, and walruses, Odobenus rosmarus. Canadian Field-Naturalist 109: 112-114. Eberhardt, L. E., R. A. Garrott, and W. C. Hanson. 1983. Den use by arctic foxes in northern Alaska. Journal of Mammalogy 64: 97-102. Garrott, R. A., L. E. Ebehardt, and W. C. Hanson. 1984. Arctic fox denning behaviour in northern Alaska. Cana- dian Journal of Zoology 62: 1636-1640. Knudsen, B. 1978. Time budgets of polar bears, Ursus mar- itimus, on North Twin Island, James Bay, during summer. Canadian Journal of Zoology 56: 1627-1628. Latour, P. B. 1981. Spatial relationships and behaviour of polar bears (Ursus maritimus Phipps) concentrated on land during the ice-free season of Hudson Bay. Canadian Jour- nal of Zoology 59: 1763-1774. Lunn, N. J., and I. Stirling. 1985. The significance of sup- plemental food to polar bears during the ice-free period of Hudson Bay. Canadian Journal of Zoology 63: 2291- 2297. Macpherson, A. H. 1969. The dynamics of Canadian arctic fox populations. Canadian Wildlife Service Report Series 8. 52 pages. Miller, G. D., and D. R. Woolridge. 1983. Small game hunt- ing behaviour of polar bears, Ursus maritumus. Canadian Field-Naturalist 97: 93-94. Nelson, R. A., G. E. Folk Jr., E. W. Pfeiffer, J. J. Craig- head, C. J. Jonkel, and D. L. Steiger. 1983. Behavior, biochemistry, and hibernation in black, grizzly, and polar bears. International Conference on Bear Research and Management 5: 284-290. Prestrud, P. 1992. Denning and home-range characteristics of breeding arctic foxes in Svalbard. Canadian Journal of Zoology 70: 1276-1283. Ramsay, M. A., R. A. Nelson, and I. Stirling. 1991. Season- al changes in the ration of serum urea to creatinine in feed- ing and fasting polar bears. Canadian Journal of Zoology 69: 298-302. Russell, R. H. 1975. The food habits of polar bears of James Bay and southwest Hudson Bay in summer and autumn. Arctic 28: 117-129. Sklepkovych, B. O., and W. A. Montevecchi. 1996. Food availability and food hoarding behaviour by red and arc- tic foxes. Arctic 49: 228-234. Smith, A. E., and M. R. J. Hill. 1996. Polar bear, Ursus mar- itimus, depredation of Canada goose, Branta canadensis, nests. Canadian Field-Naturalist 110: 339-340. Stempniewicz, L. 1993. The Polar bear, Ursus maritimus, feeding in a seabird colony in Frans Josef Land. Polar Research 12: 33-36. Stirling, I., and W. R. Archibald. 1977. Aspects of predation of seals by polar bears. Journal of the Fisheries Research Board of Canada 34: 1126-1129. Stirling, I., C. Jonkel, P. Smith, R. Robertson, and D. Cross. 1977. The ecology of the polar bear (Ursus mar- itimus) along the western coast of Hudson Bay. Canadian Wildlife Service Occasional Paper 33: 64 pages. Received 24 June 2003 Accepted 2 November 2004 604 THE CANADIAN FIELD-NATURALIST Vol. 118 Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed Duck, Clangula hyemalis MARTHA DowsLey! and ANDREW CIRTWILL? ‘Department of Geography, McGill University, 805 Sherbrooke Street West, Montreal, Quebec H3A 2K6 Canada °515 Roosevelt Drive, Kingston, Ontario K7M 5Y2 Canada Dowsley, Martha, and Andrew Cirtwill. 2004. Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed Duck, Clangula hyemalis. Canadian Field-Naturalist 118(4): 604-605. At Presqu’ile Point, Presqu’ile Provincial Park, Ontario on 23 March 2003, while interpreting the waterfowl migration for park visitors, we witnessed two adult Great Black-backed Gulls attack and kill a male Long-tailed Duck. Key Words: Great Black-backed Gull, Larus marinus, Long-tailed Duck, Clangula hyemalis, predation Great Black-backed Gulls, Larus marinus, are known to be scavengers and to take eggs, nestlings, and fledg- lings of other birds. However, their efficacy as preda- tors is less well documented. On 23 March 2003 we witnessed two adult Great Black-backed Gulls attack and kill a Long-tailed Duck, Clangula hyemalis. We were using a spotting scope (power 45x) to observe diving ducks located 200 to 250 m from shore off Presqu’ile Point, Ontario (44°00'N, 77°41'W) as they fed and rested at the front edge of ice in Presqu’ile Bay. The ducks routinely lifted off when a predatory bird passed overhead. After one such take-off, we ob- served a pair of adult Great Black-backed Gulls flying above the ducks. One male Long-tailed Duck failed to lift off and the gulls flew down and began to harass it. We did not see the initial separation of the duck from the flock, so we were unable to observe whether it was healthy and merely caught under water when the rest of the flock flew, or whether it was already injured when attacked. The attack proceeded in what appeared to be a coor- dinated effort by the gulls. One Great Black-backed Gull flew in circles overhead when the duck dived and swam under water. When the duck surfaced, the gull alighted on the water and tried to bite him. The sec- ond gull then took off from the water and flew over- head, watching the duck as he dived again to avoid the attack of the first gull. When the duck resurfaced, the second gull attacked him while the first gull again took off and watched as the duck tried to make another escape under water. The Great Black-backed Gulls continued to take turns flying and attacking the duck for 20-25 minutes. At the end of this time, one gull succeeded in grabbing the wing of the Long-tailed Duck and both gulls then pecked at the body of the duck. Less than 5 minutes later the duck was dead, and its limp body could be seen floating on the water. The adult Great Black-backed Gulls were then joined by an immature Great Black- backed Gull and all three gulls began to eat the dead duck. We could not continue observations as the wind and currents had pushed the duck’s carcass out into Lake Ontario, beyond the useful range of our spotting scope. Great Black-backed Gulls have a varied diet, but their main food sources are fish and seabirds (both often consumed as carrion) (Buckley 1990). Their predation of birds is focused on the eggs, nestlings, newly fledged young and sick or injured individuals (Beaman 1978; Mawhinney and Diamond 1999). Healthy adult birds are rarely killed, with the notable exception of preda- tion in some seabird colonies, such as those of the Atlantic Puffin (Fratercula arctica) in Northern Europe (Beaman 1978), and Newfoundland (Nettleship 1972). Great Black-backed Gulls are considered to be pred- ators of waterfowl (Good 1998); however, observations of Great Black-backed Gulls attacking and killing adult ducks are quite rare. One notable record reported by Cleghorn (1942) was of an adult Great Black-backed Gull killing a female Goldeneye (Bucephala clangula) and an immature Great Black-backed Gull killing anoth- er unidentified duck the same day. Cobb (1957) observed a single adult Great Black- backed Gull kill an injured Ruddy Duck, Oxyura Jamaicensis, in Rhode Island. In that case the method of attack was similar to what we witnessed at Presqu ile. The gull hovered above the duck as it dived and then attempted to grab it when it resurfaced (Cobb 1957). Our report corroborates the evidence that Great Black- backed Gulls will fly above diving ducks, presumably to watch them under water, and attack them when they resurface. Addy (1945) witnessed a group of Great Black- backed Gulls attack and kill an adult American Black Duck, Anas rubripes. The gulls chased and swooped at groups of American Black Ducks until one adult gull injured a duck. The gulls then worried the duck by lifting it from the water by its neck and by grasping its back. After about half an hour the duck died, and was consumed, mainly by the gull which had first attacked it (Addy 1945). This observation does not appear to have been a coordinated attack on the duck as in the case we witnessed, because one gull of a group carried out most of the attack and was the pri- mary consumer of the carcass. In our observation, the two Great Black-backed Gulls showed a level of coordination in their attack that is On 2004 unrecorded in the literature. The method they used of taking turns flying overhead and attacking the duck seems to be an efficient means to harass their prey to | the point of exhaustion and death. Literature Cited Addy, C. E. 1945. Great Black-backed Gull kills adult Black Duck. Auk 62: 142-143. Beaman, M. A. S. 1978. The feeding and population ecology of the Great Black-backed Gull in northern Scotland. Ibis 120: 126-127. Buckley, N. J. 1990. Diet and feeding ecology of Great Black-backed Gulls (Larus marinus) at a southern Irish breeding colony. Journal of Zoology: Proceedings of the Zoological Society of London 222: 363-373. GLYNNIS A. Hoop and TIM NEUFELD NOTES 605 Cleghorn, J. D. 1942. Great Black-backed Gull killing Amer- ican Goldeneye. Auk 59: 584-585. Cobb, S. 1957. Great Black-backed Gull (Larus marinus). Auk 74: 498. Good, T. 1998. Great Black-backed Gull. The birds of North America (330). Edited by A. Poole and F. Gill in The Birds of North America, Inc., Philadelphia Pennsylvania. Mawhinney, K., and A. W. Diamond. 1999. Using radio- transmitters to improve estimates of gull predation on Com- mon Eider ducklings. Condor 101: 824-831. Nettleship, D. N. 1972. Breeding success of the Common Puffin (Fratercula arctica) on different habitats at Great Island, Newfoundland. Ecological Monographs 42: 239- 268. Received 7 May 2004 Accepted 4 October 2004 | First Record of Mountain Lions, Puma concolor, in Elk Island National | Park, Alberta | Elk Island National Park, R.R. #1, Site 4, Fort Saskatchewan, Alberta T8L 2N7 Canada, e-mail: [Hood] glynnis.hood@pc.ge.ca | Hood, Glynnis A., and Tim Neufeld. 2004. First record of Mountain Lions, Puma concolor, in Elk Island National Park, Alberta. Canadian Field Naturalist 118(4): 605-607. Several sightings of Mountain Lions (Puma concolor) and wildlife mortalities consistent with predation by Mountain Lion have occurred in and adjacent to Elk Island National Park from February 2003 to present. These are the first recorded Mountain | Lions (locally called Cougar) sightings since the area encompassing the park was protected in 1906. Key Words: Mountain Lion, Cougar, Puma, Panther, Catamount, Puma concolor, Elk Island National Park, Alberta, range, first record. Along with its current range, Mountain Lion (Puma concolor) historically occupied central and eastern Canada (Banfield 1974). In present day Aiberta how- ever, Mountain Lion (locally called Cougar) popula- tions are mainly concentrated in the Rocky Mountains and the foothills (Soper 1964; Banfield 1974; Smith 1993; Pattie and Fisher 1999). By 1961, Bird described many large predators, including the Mountain Lion, as being extirpated from the aspen parkland of Canada’s prairie provinces. Soper (1964) referenced individual _ reports of Mountain Lions north of Edmonton, Alberta _ at Boiler and Grand Rapids on the Athabasca River and in the Peace River Region in northwestern Alberta. _ He also noted occasional sightings in southern Alberta along the Milk, Bow, and South Saskatchewan River systems, as well as in the Cypress Hills. Smith (1993) added additional sightings near the towns of White- court and Athabasca, Alberta. Sightings of Mountain Lions are increasing in areas where they were previously unreported. Occasional sightings have been documented to the north in Wood Buffalo National Park (Gau et al. 2001), and to the south and west of Elk Island National Park (EINP) near Cooking Lake, Pigeon Lake, and Sherwood Park (Smith 1979). However, until 2003, there had never been a documented Mountain Lion sighting in EINP (Soper 1940*; Soper 1951; Burns and Cool 1984*; Parks Canada Warden Service unpublished notes). Elk Island lies between longitude W112°57'00" and W112°46'45" and latitude N53°42'57" and N53°30'37" (Figure 1) and covers an area of 196 km*. The park is located in the Aspen Parkland Natural Subregion (Achuff 1994) in the Beaver Hills of east-central Alberta and is home to several species of large ungulates. Given the known distances that Mountain Lions would have to travel to access the park, Burns and Cool (1984) predicted “occurrences of cougar in Elk Island National Park [EINP] in future remain a slim possibility”. The closest source populations to EINP are in the Rocky Mountains foothills and the Swan Hills region of north-central Alberta (Pattie and Fisher 1999). These populations are approximately 200 km away from EINP, which is well within the 500 km dispersal distance observed for Mountain Lions (Logan and Sweanor 1999). On 8 February 2003 at 11:00 h one of us observed an adult Mountain Lion approximately 10 km south of the boundary of EINP at Islet Lake in the southern reaches of the Cooking Lake — Blackfoot Grazing, Wildlife and Provincial Recreation Area (Figure 2). 606 ,EBERTA Elk Island National Park Edmonton @ , =» Calgary == Mountain lion N distribution A in Alberta 6 i 10 = w Es FIGURE |. Location of Elk Island National Park in east-central Alberta. The shaded area indicates the distribution of Mountain Lion in Alberta (based on Pattie and Fisher 1999). In late March, a conservation officer found tracks of one large Mountain Lion and a much smaller Moun- tain Lion together near the northern boundary of the Provincial Recreation Area — an area that bounds the southern extent of EINP. On 26 July 2003 at 14:00 h a group of Young Canada Works students working in the park reported seeing an adult Mountain Lion just south of Highway 16 in the southern half of EINP. An adult Mountain Lion was also seen by park vis- itors on two different occasions in the northcentral section of the park on 29 July 2003. The first visitor reported seeing a Mountain Lion at 07:30 h beside the main road through the park and a second visitor saw a Mountain Lion while on a nearby trail at 21:40 h. In the latter case the visitor reported seeing the Moun- tain Lion watching a herd of Bison (Bison bison) approximately 5 km from the trailhead. Equestrians saw two Mountain Lions together in the north central part of the Provincial Recreation Area around the same time. Further sightings were reported in the fall of 2003 when a park visitor observed an adult Mountain Lion while hiking on 17 September. Another park visitor reported seeing a Mountain Lion cross the main park road on 2 October. In mid-October, a hiker reported seeing a Mountain Lion at the southern part of the Provincial Recreation Area between Push Lake and Islet Lake. On 20 November 2003 a park warden report- THE CANADIAN FIELD-NATURALIST Vol. 118 Elk Island N nal.Park ‘{ » atio iN (i Sighting Ungulate Mortality CookingLake - Blackfoot Grazing, Wildlife and Provincial Recreation Area FiGure 2. Mountain Lion sightings and wildlife mortalities consistent with predation by Mountain Lion in Elk Island National Park and the Cooking Lake — Black- foot Grazing, Wildlife and Provincial Recreation Area since February 2003. ed seeing a Mountain Lion crossing a secondary road in the east-central part of EINP. In 2004, a cyclist observed a Mountain Lion sitting beside the main road in the northern part of the park (13 July). The most recent confirmed sighting in the park was on 24 October 2004 when a park resident observed a Mountain Lion in the main part of the park (Figure 2). On two occasions, acreage owners on lands east and north of the park heard estrous calls of a Moun- tain Lion (October 2003 and August 2004). The land- owner who reported the August 2004 calls observed a small Mountain Lion on his property a few days later. In addition to sightings, four ungulate mortalities with wounds consistent with Mountain Lion predation (Ross et al. 1997) have been documented in the park since the winter of 2002-2003 (Figure 2). Prey species were: 2 American Elk (Cervus canadensis), 1 Moose (Alces alces), and 1 White-tailed Deer (Odocoileus virginianus). In all cases the carcasses had signs of wounds and fatal injuries to the neck and, in some cases, large puncture marks in the hide. Apart from Coyotes (Canis latrans), EINP lacks any large resident predators. In addition, the park has very high ungulate densities (approximately 13 ungulates per km7; Parks Canada unpublished data) and is home to American Elk, Moose, Bison (Bison bison), White-tailed Deer, and Mule Deer (Odocoileus hemionus). The park also has a large Beaver (Castor canadensis) population and many resident medium and small mammal species | including porcupine (Erethizon dorsatum), Muskrat | (Ondatra zibethicus) and Red Squirrel (Jamiasciurus | hudsonicus). Hare (Lepus spp.) are at low densities in the park (Super 1951). The park provides adequate undeveloped habitat and food resources to support an adult Mountain Lion (Spalding and Lesowski 1971; Ross and Jalkotzy 1992; Ross et al. 1997; Robinson et al. 2002). Currently, park staff translocate Bison and American Elk away from the park to reduce the impact of intense herbivory on vegetation. The presence of large predators, such as Mountain Lions, may help to reduce the ungulate den- | sities in the park, and therefore moderate the amount of | active management of ungulate populations by park | staff. The park is completely fenced and the abundance of prey species might offer enough resources to support a | future Mountain Lion population. Given that success- | ful reproduction in Mountain Lions is related to the | availability of food (Pierce et al. 2000), the likely pres- 'ence of a mated pair within EINP suggests such a scenario 1s not unrealistic. Acknowledgments We thank Conservation Officer Ed Whitelock and Park Warden Olaf Jensen for providing us with addi- ( tional information on Mountain Lion sightings. We also thank three anonymous reviewers for their edito- rial comments. Documents Cited | Burns, G. R., and N. L. Cool. 1984. A biophysical inventory of the non-ungulate mammals of Elk Island National Park. Report prepared for Parks Canada by the Canadian Wild- life Service, Edmonton, Alberta, Canada. 195 pages. Soper, J. D. 1940. Preliminary final report on Elk Island Na- tional Park, Alberta, Canada. unpublished report. Depart- ment of Mines and Resources Lands, Parks and Forest Branch, National Parks Bureau. Edmonton, Alberta, Cana- da. 46 pages. Literature Cited Achuff, P. L. 1994. Natural regions, subregions and natural history themes of Alberta — a classification for protected areas management, revised and updated December 1994. NOTES 607 Alberta Environmental Protection, Edmonton, Alberta, Canada. 72 pages. Banfield, A. W. F. 1974. The mammals of Canada. Univer- sity of Toronto Press, Toronto, Ontario, Canada. 438 pages. Bird, R. D. 1961. Ecology of the aspen parkland of western Canada in relation to land use. Canada Department of Agri- culture, Ottawa, Ontario, Canada. 155 pages. Gau, R. J., R. Mulders, T. Lamb, and L. Gunn. 2001. Cougars (Puma concolor) in Northwest Territories and Wood Buffalo National Park. Arctic 54: 185-187. Logan, K. A., and L. Sweanor. 1999. Puma. Pages 347-77 in Ecology and management of large mammals in North America. Edited by S. Demarais and P. R. Krausman Prentice-Hall, Englewood Cliffs, New Jersey. Pattie, D., and C. Fisher. 1999. Mammals of Alberta. Lone Pine Publishing. Edmonton, Alberta, Canada. 240 pages. Pierce, B. M., V. C. Bleich, and R. T. Bowyer. 2000. Social organization of Mountain Lions: does a land-tenure sys- tem regulate population size? Ecology 81: 1533-1543. Robinson, H. S., R. B. Wielgus, and J. C. Gwilliam. 2002. Cougar predation and population growth of sympatric mule deer and white-tailed deer. Canadian Journal of Zoology 80: 556-568. Ross, P. I., and M. G. Jalkotzy. 1992. Characteristics of a hunted population of cougars in southwestern Alberta. Journal of Wildlife Management 56: 417-426. Ross, P. I., M. G. Jalkotzy, and B. M. Festa. 1997. Cougar predation on bighorn sheep in southwestern Alberta dur- ing winter. Canadian Journal of Zoology 75: 771-775. Smith, H. C. 1979. Mammals of the Edmonton area. Provin- cial Museum of Alberta Natural History Occasional Paper Number 2. Queens Printer, Edmonton, Alberta, Canada. 34 pages. Smith, H. C. 1993. Alberta mammals, an atlas and guide. The Provincial Museum of Alberta. Edmonton, Alberta, Cana- da. 238 pages. Soper, J. D. 1951. The mammals of Elk Island National Park, Alberta Canada. Wildlife Management Bulletin Series 1, Number 3. Canadian Wildlife Service, Ottawa, Ontario, Canada. 27 pages. Soper, J. D. 1964. The mammals of Alberta. The Hamley Press, Ltd. Edmonton, Alberta, Canada. 402 pages. Spalding, D. J., and J. Lesowski. 1971. Winter food of the cougar in south-central British Columbia. Journal of Wild- life Management 35: 378-381. Received 19 December 2003 Accepted 23 December 2004 608 THE CANADIAN FIELD-NATURALIST Vol. 118 First Confirmed Occurrence of a Wolf, Canis lupus, South of the St. Lawrence River in Over 100 Years Mario VILLEMURE! and HELENE JOLICOEUR? 'Université de Sherbrooke, Département de Biologie, Sherbrooke, Québec JIK 2R1 Canada Current address: 11A, chemin St-Francois, St-Mathieu-du-Parc, Québec GOX INO Canada *Société de la faune et des parcs du Québec, Direction du développement de la faune, 675, boulevard René-Lévesque Est. . 11" floor, box 92, Québec, Québec GIR 5V7 Canada Villemure, Mario, and Helene Jolicoeur. 2004. First confirmed occurrence of a Wolf, Canis lupus, south of the St. Lawrence River in over 100 years. Canadian Field-Naturalist 118(4): 608-610. A large canid was snared near Sainte-Marguerite-de-Lingwick, Québec, in January 2002. DNA analysis confirmed the ani- mal to be a Wolf (Canis lupus). Wolves were extirpated from this region around 1850-1900 and this is the first confirmed observation since then. Key Words: Wolf, Canis lupus, range, dispersal, St. Lawrence River, Québec. Historically, Wolves (Canis lupus) were distributed across most of North America. They were extirpated in the southern portion of their range by the turn of the century (Nowak 1983). In Québec, Wolves disap- peared from the south shore of the St. Lawrence River around 1850-1900 (Peterson 1966). The extermination of the Wolf and the development of agriculture in the mid-1900 facilitated the extension of Coyote (Canis latrans) range. Wolves are now mostly limited to the northern and less populated regions of North America (Wayne et al. 1992). On 19 January 2002, a male Wolf was trapped near the village of Sainte-Marguerite-de-Lingwick (45°36'15"N, 71°17'15"W) in the Eastern Townships of southern Québec. The Wolf weighed 29.1 kg, sim- ilar to the weight of an adult from Papineau-Labelle (Potvin 1986*) or a yearling from the Laurentides region (Jolicoeur 1998%*). A tissue sample was collected from the temporal muscle for genetic identification of the species. DNA analyses were performed by the Natural Resources DNA Profiling and Forensic Center (Trent University, Peterborough, Ontario K9J 7B8 Can- ada) following the method described in Wilson et al. (2000). The Eastern Townships sample was profiled at the mitochondrial DNA (mtDNA) control region and 8 microsatellite loci. The genetic profile was compared to samples of canids representing the Eastern Wolf (C. lupus lycaon) from Algonquin Provincial Park; West- ern Coyotes (C. latrans) from Texas, North Carolina and Ohio; and Wolves (C. lupus) from Pukaskwa Na- tional Park, northeastern Ontario and the Laurentide Wildlife Reserve region north of Québec City. The above populations were used to compare the Eastern Townships sample against C. lupus lycaon, C. latrans and C. lupus to assess the species-of-origin or hybrid genotype. The sample had a mtDNA consistent with C. 1. lycaon/ C. latrans and the microsatellite genotype suggested 95.0% shared ancestry with Eastern Wolves from Algonquin Provincial Park. The sample had a DNA profile consistent with an Eastern Wolf. The Eastern Wolf is generally described as being smaller than other Gray Wolf subspecies (Nowak 1995, 2002). Although it has been proposed as a distinct species (C. lycaon) by Wilson et al. (2000), its status as a subspecies is still generally accepted. Some authors suggest it may result from hybridization between C. rufus and C. Tupus (Nowak 2002). In Québec, the Eastern Wolf is found mostly in the southern deciduous and mixed forests (Jolicoeur and Henault 2002*) Coyote genes found in the mitochondrial DNA of the specimen have been reported before in Wolves in eastern Canada (Lehman et al. 1991; Wilson et al. 2000). Wolves and Coyotes are most likely to interbreed when Wolf den- sity is low relative to Coyotes and when the species are similar in size. In southern Québec, male Coyotes weigh 14.6—18.7 kg (Fortin and Huot 1995*; Dumond and Villard 2000; Villemure 2003*). Wolf dispersal has been monitored in southern Qué- bec (Messier 1985; Potvin 1987: Jolicoeur 1998*; Villemure 2003*, but Wolves have not been docu- mented crossing the St. Lawrence River. Harrison and Chapin (1998) identified two potential corridors link- ing Wolf populations north of the St. Lawrence River to potential habitat in Maine and New Hampshire. However, movements of Wolves south of their current range are thought to be unlikely because of potential barriers such as the St. Lawrence Seaway and regions with high human population, high road density and intensive agriculture (Wydeven et al. 1998). Potential core habitat for Wolves has been identified in New England as well as the Eastern Townships and Beauce regions of Québec (Harrison and Chapin 1998; Mlade- noff and Sickley 1998; Carroll 2003*; Jolicoeur and Etcheverry, in preparation”). Wolves are highly mobile and frequently move over long distances (Van Camp and Glukie 1979; Fritts 1983: Mech 1987). Some of the longest dispersal move- ments documented (460-555 km) occurred across a mixture of forest, farmland, and 4-lane highways in the upper Midwest (Mech et al. 1995; Wydeven et al. 1995). A Wolf in Alberta crossed rivers 0.5 - 2.0 km ' wide during summer (Van Camp and Glukie 1979), | and Wolves have crossed 24 km on a frozen lake (Mech 1966). Wolves have increased in Minnesota (Fuller et ) al. 1992), and since the mid-1970s have naturally re- colonized portions of northern Wisconsin and, more recently, Upper Michigan (Mech and Nowak 1981; Fuller et al. 1992). Wolves from Canada have recolo- nized Montana for at least the past decade (Boyd et al. 1995). In a review of Wolf dispersal and recoloniza- | tion, Wydeven et al. (1998) reported that 31 to 63% of dispersing Wolves successfully settled and formed pairs in new territories. It is unclear whether Wolves have begun re-estab- _ lishing in the Eastern Townships, or whether this Wolf was simply a dispersing individual. While the Eastern Townships may be a sink habitat for Wolves, this Wolf was caught only 30-50 km from potential Wolf habi- tat in Maine (Carroll 2003*). Further investigation is necessary to confirm other possible Wolf observa- tions in these regions. Reporting of Wolf sightings or accidental Wolf captures south of the St Lawrence river should be encouraged. | Acknowledgments We are grateful to Laurent Cloutier, trapper and for- mer Eastern Townships Wildlife Protection Officer, for providing the specimen and details of the capture location. DNA analysis were funded by the Société de la Faune et des Parcs du Québec. Thanks to C. | Carroll, M. Festa-Bianchet, C. Reining and P. Struh- | sacker for comments on an earlier draft of this manu- | script. Documents Cited (marked * in text) Carrol, C. 2003. Carnivore restoration in the Northeastern U.S. and Southeastern Canada: a regional-scale analysis of habitat and population viability for wolf, lynx, and marten. Progress report 1: Wolf viability analysis. Prepared for The Wildlands Project. Orleans, California: Klamath Cen- ter for Conservation Research. 38 pages. Fortin, C., and J. Huot. 1995. Ecologie comparée du coyote, du lynx et du renard roux au pare national Forillon. Report prepared for Parks Canada. Département de Biologie, Universite Laval. Ste-Foy, Québec. 291 pages. _ Jolicoeur, H. 1998. Le loup du massif du lac Jacques-Cartier. Québec, Ministére de l’Environnement et de la Faune, Direction de la Faune et des Habitats, Direction de la conservation et du patrimoine écologique. 132 pages. Jolicoeur, H., and M. Hénault. 2002. Repartition géogra- phique du loup et du coyote au sud du 52° paralléle et estimation de la population de loups du Québec. Québec, Societe de la faune et des pares du Québec, Direction du développement de la faune et Direction de l’aménagement de la faune des Laurentides. 51 pages. | Jolicoeur, H., and P. Etcheverry (in preparation). Habitat potentiel pour le loup sur la rive sud du St-Laurent. Québec, Sociéte de la faune et des parcs du Québec, Direction du développement de la faune. Potvin, F. 1986. Ecologie du loup dans la reserve de Papi- neau-Labelle. Québec, Ministére du Loisir, de la Chasse et de la Péche, Direction générale de la Faune. 103 pages. NOTES 609 Villemure, M. 2003. Ecologie et conservation du Joup dans la region du parc national de la Mauricie. Mémoire de Maitrise. Département de Biologie, Universite de Sherbrooke. Sher- brooke, Québec, Canada. 89 pages. Literature Cited Boyd, D. K., P. C. Paquet, S. Donelon, R. R. Ream, D. H. Pletcher, and C. C. White. 1995. Transboundary move- ments of a recolonizing wolf population in the Rocky Mountains. Pages 141-146 in: Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts and D. R. Seip. Canadian Circumpolar Insti- tute, Occasional Publication Number 35, 642 pages. Dumond, M., and M. A. Villard. 2000. Demography and body condition of coyotes (Canis latrans) in eastern New Brunswick. Canadian Journal of Zoology 78: 399-406. Fritts, S. H. 1983. Record dispersal by a wolf from Min- nesota. Journal of Mammalogy 64: 166-167. Fuller, T. K, W. E. Berg, G. L. Radde, M. S. Lenarz, and G. Blair Joselyn. 1992. A history and current estimate of wolf distribution and numbers in Minnesota. Wildlife Society Bulletin 20: 42-55. Harrison, D. J., and T. G. Chapin. 1998. Extent and con- nectivity of habitat for wolves in eastern North America. Wildlife Society Bulletin 26: 767-775. 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 sympat- ric North American gray wolf populations. Evolution 45: 104-119. ‘ Mech, L. D. 1966. The wolves of Isle Royale. U.S. National Park Service, Fauna Series 7. Mech, L. D. 1987. Age, season, distance, direction, and social aspects of wolf dispersal from a Minnesota pack. Pages 55-74 in Mammalian dispersal patterns: The effects of social structure on population genetics. Edited by B. D. Chepko-Sade and Z. Tang Halpin. University of Chicago Press. Chicago. 342 pages. Mech, L. D., S. H. Fritts, and D. Wagner. 1995. Minnesota Wolf dispersal to Wisconsin and Michigan. American Mid- land Naturalist 133: 368-370. Mech, L. D., and R. M. Nowak. 1981. Return of the gray wolf to Wisconsin. American Midland Naturalist 105: 408-409. Messier, F. 1985. Solitary living and extraterritorial move- ments of wolves in relation to social status and prey abun- dance. Canadian Journal of Zoology 63: 239-245. Mladenoff, D. J., and T. A. Sickley. 1998. Assessing poten- tial gray wolf restoration in the northeastern United States —a spatial prediction of favorable habitat and potential population levels. Journal of Wildlife Management 62: 1-10. Nowak, R. M. 1983. A perspective on the taxonomy of wolves in North America. Pages 10-19 in: Wolves in Canada and Alaska: their status, biology, and management. Edited by L. N. Carbyn. Canadian Wildlife Service. 145 pages. Nowak, R. M. 1995. Another look at wolf taxonomy. Pages 375-397 in: Ecology and conservation of wolves in a chang- ing world. Edited by L. N. Carbyn, S. H. Fritts and D. R. Seip. Canadian Circumpolar Institute, Occasional Publi- cation Number 35, 642 pages. Nowak, R. M. 2002. The original status of wolves in eastern north america. Southeastern Naturalist 1: 95-130. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press. Toronto. 465 pages. 610 Potvin, F. 1987. Wolf movement and population dynamics in Papineau-Labelle reserve, Québec. Canadian Journal of Zoology 66: 1266-1273. Van Camp, J., and R. Glukie. 1979. A record long distance move by a wolf (Canis lupus). Journal of Mammalogy 60: 236-237. Wayne, R K., N. Lehman, M. W. Allard, and R. L. Honey- cutt. 1992. Mitochondrial DNA variability of the gray wolf: Genetic consequences of population decline and habitat fragmentation. Conservation Biology 6: 559-569. Wilson, P. J., S. Grewal, I. D. Lawford, J. N. M. Heal, A. G. Granacki, D. Pennock, J. B. Theberge, M. T. The- berge, D. R. 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 evolutionary history THE CANADIAN FIELD-NATURALIST Vol. 118 independent of the gray wolf. Canadian Journal of Zoology 78: 2156-2166. Wydeven, A. P., T. K. Fuller, W. Weber, and K. Mac Donald. 1998. The potential for wolf recovery in the Northeastern United States via dispersal from Southeastern Canada. Wildlife Society Bulletin 26: 776-784. Wydeven, A. P., R. N. Schultz, and R. P. Thiel. 1995. Moni- toring of a recovering gray wolf population in Wisconsin. Pages 147-156 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, Occa- sional Publication Number 35. 642 pages. Received 11 August 2003 Accepted 24 December 2004 Is Cost of Locomotion the Reason for Prolonged Nesting Forays of Snapping Turtles, Chelydra serpentina? SHANE R. DE SOLLA and KIM J. FERNIE Canadian Wildlife Service, Environment Canada, Canada Centre for Inland Waters, 867 Lakeshore Road, P.O. Box 5050, Burlington, Ontario L7R 4A6 Canada de Solla, Shane R., and Kim J. Fernie. 2004. Is cost of locomotion the reason for prolonged nesting forays of Snapping Turtles, Chelydra serpentina? Canadian Field-Naturalist. 118(4): 610-612. Prolonged nesting forays were observed in five gravid Snapping Turtles (Chelydra serpentina), in 1999 and 2001. For all observations, the females began exploratory nest excavations but failed to oviposit. Subsequently, all five females sought refuge either by burying themselves in substrate, or by seeking shade under vegetation, presumably to wait until the following day to resume nesting activities. By contrast, most observations of failed nesting resulted in the females abandoning the nesting site and returning immediately to water. Although prolonged nesting attempts in other turtle species (i.e. kinosternids) likely are associated with rainfall or predation risk, we speculate that these prolonged nesting attempts in Snapping Turtles reduced the cost of terrestrial travel. Key Words: Snapping Turtles, Chelydra serpentina, oviposition, nesting behaviour, cost of locomotion, Ontario Nesting behaviour of Snapping Turtles (Chelydra serpentina) has been described in detail elsewhere (see Ernst et al. 1994), and is generally characterized by behaviour that minimizes exposure on terrestrial habitats. Briefly, nesting behaviour starts with emer- gence from water, followed by nest site selection. Females typically dig with their front limbs, and often poke their noses into the substrate, presumably to deter- mine if the soil type and/or hydration are adequate. Subsequently, the rear limbs are used first to excavate the nest cavity, then position the eggs during oviposi- tion, and finally to bury the eggs. Subsequently, the female leaves the nesting area, and returns to the water. Although Snapping Turtles have been known to travel more than 0.5 km overland to nesting sites (Obbard and Brooks 1980), normally they nest at a site much closer to the water. Regardless if the female was suc- cessful in nesting or not, generally she returns to water immediately afterwards. Nesting normally occurs in the early morning or late evening, although females may nest in the afternoon following rain. By mini- mizing the distance or time traveled on land, female Snapping Turtles may avoid energetic costs, or the risks of dehydration or predation. Baudinette et al. (2000) found that terrestrial locomotion was 2.6 times more energetically costly than aquatic locomotion for the Murray Short-necked Turtle (Emydura macquarii). Snapping Turtles are particularly vulnerable to water loss compared to other turtles (Ernst 1968) due to their exposed skin. Although depredation of adults is rare, adult females are occasionally taken by large predators, such as bears or coyotes (Ernst et al. 1994). Here we document five instances of prolonged nesting forays in Snapping Turtles, which have not previously been documented, and discuss possible reasons for this behaviour. Study Site Observations were made from two locations in Ontario, both known for many years to be nesting sites for Snapping Turtles. The first nesting site, the north- western shoreline of Coote’s Paradise, Hamilton (43°16'N, 79°56'W), consists primarily of a communi- ty vegetable garden and a wood chip pile on Ontario Power Generation property. The nesting site is sepa- rated from the open water by approximately 150 m, first by a steep hill with thick shrub cover, and sec- -ondly by dense cattails. The second nesting site, at ‘! Wheatley Provincial Park (42°5'N, 82°26'W), consists of a series of sand, dirt, and wood chip piles in a main- tenance area. Although normally open water is relative- ly close to the nesting site (~ 50 m), in 2001 when the observations were made, water levels were extremely low, and most water in the park had drained into Lake Erie, leaving bare mudflats. We observed many turtle | tracks in the exposed mud leaving the park into Lake Erie (de Solla, personal observation). Thus, at both sites, the nesting females did not have nearby access to water. | Nesting Observations De Solla et al. (2001) reported the deaths of two gravid Snapping Turtles that buried themselves in a | composting wood chip pile at Cootes Paradise, Hamil- ton, on 7 and 9 June 1999. Both gravid turtles remained in the wood chip pile until their deaths. Results of post- mortem examinations were consistent with death due to hyperthermia (de Solla et al. 2001); however, no reasons for their behaviour were suggested. The behav- iour of both females was consistent for females search- | ing for suitable nesting sites, except after failing to | successfully nest, both buried themselves in the wood chip pile. A third female Snapping Turtle was also | observed 8 June 1999 on the wood chip pile. Follow- | ing apparently unsuccessful attempts to nest, the female | was observed to be motionless for a few hours under | the shade of burdock (Arctium spp.) and other vege- | tation, which provided the only shade in the area. De Solla did not touch her until later in the morning, but | the turtle was found to be gravid. She remained in the shade after being disturbed. On 9 June 2001 at 1045, a female turtle was found on a storage area for sand, dirt, and wood chips at Wheatley Provincial Park. There were numerous tracks on both the wood chips and dirt piles, possibly by more than one female. There was abundant shade at this site. She was sitting in shade under vegetation on a small dirt pile, with no apparent movement, and no evidence of nesting. She was left alone until 1200, at which time the female was about 2 m from her previ- ous location, further in shade under vegetation. She did not move by 1300. At this time she was palpated, found to be still gravid, and a blood sample was taken. She was released at the same spot she was found, where she remained for at least one hour, but she was not subsequently observed. No temperature readings were taken; however, the weather was sunny and hot. On 12 June 2002, a female was seen in the vegetable garden at Coote’s Paradise at approximately 2130. Nearby were at least 10 digging attempts in freshly cultivated soil, and she was observed digging with her front limbs and poking her head into the soil. She was observed for approximately an hour, but did not show any signs of nesting. At approximately 2230, she NOTES 611 was observed to have buried herself, with only the top of her carapace showing. There was relatively little shade by vegetation available. At approximately 0745 hrs, 13 June 2002, a female was seen walking very near where the female from the previous night was observed. This female was heavily covered with dirt, suggesting that she was the same female that had buried herself. No nesting attempts were made by 1000 hrs; subsequently we left and she was not observed again. Discussion The five observations of nesting attempts by Snap- ping Turtles had two common features; the delay was prior to oviposition, and the turtles sought terrestrial refuge by burying themselves in soil or wood chips, or by staying in shade. We have no observations of females remaining at the nesting ground after suc- cessful oviposition. We are unaware of any published reports of such behaviour in Snapping Turtles. Various kinosternid species prolong their nesting forays, typically by burying themselves before and/or after oviposition (Iverson 1990; Burke et al. 1994; Wil- son et al. 1999) adjacent to the nest. The duration of nest attendance in species other than Chelydra appears to be linked to the timing or duration of rainfall (Burke et al. 1994: Ernst et al. 1994). Intensive disturbance may also delay successful oviposition, and competitive nesting by Green Turtles (Chelonia mydas) at high densities may delay oviposition by a few days (Jessop et al. 1999). Typically, after a failed nesting attempt Snapping Turtles return to the wetland immediately following cessation of nesting behaviour (de Solla, persional observation). We speculate that the prolonged nesting attempts we observed in Snapping Turtles was to avoid the cost of making a second trip between the water and nest- ing site. All instances of prolonged nesting attempts that we observed occurred a considerable distance from water. Abandoning a nesting attempt would therefore result in a relatively long journey to the water, and back again to the nest site for a second oviposition attempt. To avoid this cost, once a female aborts a nesting attempt, she may burrow under substrate or hide under vegetation until the following day. Females nesting closer to wetlands are more likely to leave the nest site and come back at a later time. At the Wildlife Research Station (Algonquin Park), Snapping Turtle nesting has been monitored on a dam at the southern end of Lake Sasajewun for > 25 years, and all observed nesting attempts that failed were followed by the imme- diate withdrawal to the water (~ 5 m) by the female (R. J. Brooks, University of Guelph, personal com- munication). Similarly, at eight other nesting sites in Ontario that the authors visited, all of which were close to wetland habitat, females withdrew to water following a failed nesting attempt. We did not count the number of these observations, but de Solla has frequently observed this behaviour. 612 In two of the five observations of prolonged nesting attempts, the turtles died of hyperthermia after bury- ing themselves in composting wood chips (de Solla et al. (2001). We speculate that in most cases where females bury themselves, they use non-composting material, and thus are not at risk of hyperthermia. Acknowledgments We thank Wheatley and Algonquin Provincial Parks, and Ontario Power Generation for permission to enter their properties to look for turtle nests. Ron Brooks and anonymous reviewers gave valuable comments on earlier drafts. Literature Cited Baudinette R. V., A. M. Miller, and M. P. Sarre. 2000. Aquatic and terrestrial locomotory energetics in a toad and a turtle: a search for generalisations among ectotherms. Physiological and Biochemical Zoology 73: 672-682. Burke V. J., J. W. Gibbons, and J. L. Greene. 1994. Pro- longed nesting forays by common mud turtles (Kinosternon subrubrum). American Midland Naturalist 131: 190-195 THE CANADIAN FIELD-NATURALIST Vol. 118 de Solla S. R., D. Campbell, and C. A. Bishop. 2001. Hyperthermia induced mortality of gravid snapping tur- tles, Chelydra serpentina, and eggs in a wood chip pile. Canadian Field-Naturalist 115: 510-512. Ernst, C. H. 1968. Evaporative water-loss relationships of turtles. Journal of Herpetology. 2: 159-161. Ernst, C. H., J. E. Lovich, and R. W. Barbour. 1994. Turtles of the United States and Canada. Smithsonian Institution Press. Washington. Iverson, J. B. 1990. Nesting and parental care in the mud turtle, Kinosternon flavescens. Canadian Journal of Zoology 68: 230-233. Jessop, T. S., C. J. Limpus, and J. M. Whittier. 1999. Plasma steroid interactions during high-density green turtle nest- ing and associated disturbance. General and Comparative Endocrinology 115: 90-100. Obbard M. E., and R. J. Brooks. 1980. Nesting migrations of the snapping turtle (Chelydra serpentina). Herpetologica 36: 158-162. Wilson D. S., H. R. Mushinsky, and E. D. McCoy. 1999. Nesting behavior of the striped mud turtle, Kinosternon baurii (Testudines: Kinosternidae). Copeia. 1999: 958-968. Received 11 October 2003 Accepted 23 November 2004 Significant Vascular Plant Records from the Hamilton Area, Ontario CARL ROTHFELS Royal Botanical Gardens, 680 Plains Road W., Burlington, Ontario L7T 4H4 Canada Rothfels, Carl. 2004. Significant vascular plant records from the Hamilton area, Ontario. Canadian Field-Naturalist 118(4): 612-615. Four additions to the known vascular flora of Ontario (Aesculus pavia, Ambrosia x helenae, Anthriscus caucalis, Verbena bonariensis) and ten other provincially significant records are discussed. Of the 14 taxa listed, two (Actaea x ludovici and Ambrosia X helenae) are native. Key Words: adventive species, flora, rare species, distribution, Hamilton, Halton, Ontario. Field and herbarium work centred around Royal Botanical Gardens on the border between Hamilton and Burlington, Ontario (43°29.00'N, 79°88.00'W), has resulted in the following 16 provincially significant vascular plant records in 2002. For locally significant records, see Rothfels (2003). All records are supported by specimens in the Royal Botanical Gardens herbar- ium (HAM). Taxa preceded by an asterisk (*) are not native to the Hamilton Region. Taxon names are followed by their subnational rank (Srank) as of 1998, where appli- cable (see Newmaster et al. 1998). An Srank of S1 indicates a very rare native taxon, with generally five or fewer occurrences in Ontario; a rank of SE1 is similar, but refers to a non-native taxon; and a rank of SEH (historical) indicates a non-native species that has typically not been recorded in Ontario in the past 20 years (Newmaster et al. 1998). The names used follow Kartesz and Meacham (1999*). Actaea x ludovici B.Boivin. Hybrid Baneberry S1 RANUNCULACEAE One clump of this hybrid was collected from the Devil’s Punchbowl Environmentally Significant Area, in Hamilton (43°11.80'N, 79°38.50'W). Both parental species (Actaea rubra (Aiton) Willd. and Actaea pachypoda Ell.) were in close proximity. Despite the overlapping ranges of the parental taxa, the hybrid is rare (Voss 1985), perhaps due to differing phenology (Pringle, personal communication). In Canada, it is recorded from Ontario and Quebec (Kartesz and Meacham 1999*). (C. J. Rothfels 305). * Aesculus pavia L. Red Buckeye HIPPOCASTANACEAE This species is an addition to the flora of Ontario (Newmaster et al. 1998). Over ten young trees were found naturalized in the Red Hill Creek escarpment valley, City of Hamilton, by Megan Ogilvie and Albert Garofalo. These plants are likely spreading from trees 2004 | planted in 1927 (Bruce Duncan, personal communica- ) tion). A species of the southern United States, it | reaches its northern limit in Kentucky, West Virginia, and southern Illinois (Kartesz and Meacham 1999*),. | It differs from A. glabra Willd. and A. hippocastanum | L., the other two species reported from Ontario, by its / smooth fruit (Edmondson 1997), among other features. | It is, due to its reddish flowers, perhaps most likely to | be confused with the frequently-planted Aesculus _ xcarnea Hayne, which differs in its size (A. pavia is a | small tree or large shrub, A. xcarnea is a large tree), _ leaf shape (A. pavia has lanceolate leaves unlike the | obovate leaves of A. xcarnea) and petal shape (A. _ pavia has dissimilar petals with two broad lateral _ petals and two long-stalked spoon-shaped petals; all _ petals of A. xcarnea are large and shaggy) (Krussman _ 1984). (M. Ogilvie, A .Garofalo et al. s.n.; HAM 16221). _ * Aethusa cynapium L. Fool’s-parsley SE1 APIACEAE This species was found new for both the City of Hamilton (Goodban 1995) and Lambton County (Tiedje and Tiedje 2002) in 2002. In Hamilton, it is fairly common along wooded paths along the base of the Niagara Escarpment in the Devil’s Punchbowl Envi- ronmentally Significant Area (43°12.20'N, 79°40.70'W). _ The Hamilton specimen was determined only to the _ specific level. The Lambton County specimen was _ determined as A. cynapium ssp. cynapioides (M. Bieb.) _ Nyman, the larger woodland subspecies (using Tutin 1968) and it was common on the Ausable River flood- plain in the Rock Glen Conservation Area (43°05.10'N, 81°06.80'W). Aethusa cynapium has been found in the Great Lakes States, and in Quebec, New Brunswick, and Nova Scotia (Kartesz and Meacham 1999*). The early Canadian reports describe this species as a garden weed “introduced with garden seeds from Europe” (Macoun 1883), and list it from Hastings and North- umberland Counties in Ontario, and as “occasional” in New Brunswick (Macoun 1883). (J. Rothfels & J. Shearer 243; C. J. Rothfels 274). * Amaranthus blitum L. Purplish Amaranth SEH AMARANTHACEAE This species is new for Halton Region (Varga et al. 2000). A purple-leaved amaranth, it is scattered irreg- ularly in the lawns at Royal Botanical Gardens Centre (43°17.40'N, 79°59.70'W) where it appears to be dis- persed by Canada Geese. Scoggan (1978) reports this species as introduced in waste ground in Elgin County, Huron County, and Waterloo Regional Municipality in Ontario, and from Masson, Montreal and Quebec City in Quebec. (C. J. Rothfels & D. Gugler 303). Ambrosia x helenae Rouleau Hybrid Ragweed ASTERACEAE This taxon is an addition to the flora of Ontario (Newmaster et al. 1998). It is a hybrid between Com- mon Ragweed (Ambrosia artemisiifolia L.) and Giant Ragweed (Ambrosia trifida L.). Although the ranges NOTES 613 of these two species overlap frequently, the hybrid is very rare (Wagner 1958). It was collected along the north shore trails of the Cootes Paradise Nature Sanc- tuary in Hamilton (43°16.90'N, 79°54.10'W). The type locality is in Quebec (Wagner 1958), and it has also been found in Michigan, New York State, and Ohio (Kartesz and Meacham 1999*). (D. Gugler s.n.: HAM 15959). * Anthriscus caucalis M. Bieb. Bur-chervil APIACEAE This species is an addition to the flora of Ontario (Newmaster et al. 1998). It was abundant on disturbed ground near the Royal Botanical Gardens’ Laking Garden (43°17.50'N, 79°53.30'W). Shortly after the discovery, the site was mowed by the owner. This species is known in Canada only from Nova Scotia, but is scattered widely across the United States (Kartesz and Meacham 1999*), and is easily distinguished from its more common congener (A. sylvestris (L.) Hoffm.) by the hooked hairs on the fruit (e.g., Gleason and Cronquist 1991). (C. J. Rothfels & J. L. Reader 115). * Anthyllis vulneraria L. Lady’s-fingers SE1 FABACEAE This species is new for the City of Hamilton (Good- ban 1995). It was fairly common, but local, in disturbed, weedy, poorly drained ground (a pipeline right of way) in the Beverly Swamp Environmentally Significant Area (43°21.10'N, 80°06.80'W). It has been reported from British Columbia, Ontario, Quebec, Newfound- land, and New Brunswick (Kartesz and Meacham 1999*). Scoggan (1978) notes that it is “locally intro- duced into clover fields and waste places,” a description that concords closely with this report, and lists records from Oxford, Waterloo and Wellington Counties in Ontario, from “slaty banks of the Restigouche River near Matapedia” in Quebec, and from Newcastle in New Brunswick. (C. J. Rothfels 143). * Cardamine impatiens L. Narrow-leaved Bittercress SE1 BRASSICACEAE This species is new for the City of Hamilton (Good- ban 1995). Four plants were found along the north shore of the Cootes Paradise Nature Sanctuary, near a small boat-storage facility, along a partially disturbed forest edge (43°16.90'N, 79°54.00'W). This is an up- right Cardamine, with distinctly auriculate pinnately- divided leaves (e.g. Voss 1985). Mulligan (2002) only saw a single Canadian specimen, collected by J. M. Weber in 1980 for Port Credit, Ontario, and notes that this species is “sporadic and uncommon” in the United States. This species was not listed by Scoggan (1978). (C. J. Rothfels 88). * Coronopus didymus (L.) J.E.Sm. Lesser Wart-cress SEI BRASSICACEAE This strange little mustard is new for the City of Hamilton (Goodban 1995). It is a common weed in some of the Royal Botanical Gardens’ Rock Garden 614 flower beds (43°17.20'N, 79°53.60'W). This is the second confirmed record for Ontario (Oldham personal communication). It first established a foothold in the east — Scoggan (1978) lists five counties in Nova Scotia and two in New Brunswick containing this species — but it also occurs across the country (a single record each in British Columbia, Alberta, Quebec, and Newfoundland (Scoggan 1978). The first Canadian records are from Gaspé Basin, Quebec, and North Sidney, Nova Scotia, in 1862 and 1883, respectively (Mulligan 2002). Coronopus didymus can be differen- tiated from Coronopus squamatus (Forssk.) Aschers. (which is also weedy) by its wrinkled cordate fruits that are notched at the summit. The fruits of C. sguamatus are conspicuously apiculate at the summit (e.g. Holmgren 1998). (C. J. Rothfels 71). * Euonymus fortunei (Turcz.) Hand.-Mazz. Wall- creeper SE] CELASTRACEAE This species has been known from Royal Botani- cal Gardens’ sanctuary lands for several years (Smith 2003), but has not been collected until this year. It is widespread but local in the Cootes Paradise Nature Sanctuary, where it occasionally forms small dense patches on the oak-hickory slopes (43°1620'N, 79°55.10'W). It seems to be increasing in southern Ontario, and may become a problem in some natural areas (Oldham, personal communication 2003). (C. J. Rothfels 235). * Macleaya cordata (Willd.) R. Br. Plume-poppy SEH PAPAVERACEAE This species is new for the City of Hamilton (Good- ban 1995). This large plant is occasionally planted as an ornamental. Its occurrence on the east shore of the Cootes Paradise Nature Sanctuary (43°16.80'N, 79°53.50'W) is mysterious; it could be a persistent population from some unknown source or it could be “spontaneous”. Voss (1985) describes it as “seldom escaping” in Michigan, and Scoggan (1978) lists it as a “garden escape” that is “scarcely established” in Nor- folk County in Ontario and Missisquoi County in Quebec. (C. J. Rothfels 318). * Myrrhis odorata (L.) Scop. Scented Myrrhis SE1 APIACEAE This large umbellifer is new for Halton Region (Varga et al. 2002). It is escaping from the Royal Bot- anical Gardens’ Scented Garden into the neighboring ravines (43°17.50'N, 79°52.60'W). Currently, its spread is limited, and will be tracked in future years. It is reported from two other Ontario locations: St. Thomas, Elgin County, and Manitoulin Island, Manitoulin District (Pringle 1994); as well as from British Colum- bia, Nova Scotia, Michigan, Pennsylvania, and Oregon (Kartesz and Meacham 1999*). The smooth fruit and large size help differentiate this species from the native Osmorhiza species (Pringle 1994). (C. J. Rothfels 190). THE CANADIAN FIELD-NATURALIST Vol. 118 * Sorghum bicolor (L.) Moench ssp. bicolor. Sorghum SE] POACEAE This species is new for the City of Hamilton (Good- ban 1995). It was common with other weeds on freshly disturbed ground at the Royal Botanical Gardens’ outdoor compost facility (43°17.10'N, 79°53.70'W). It has been found in Quebec and most U.S. states, and is a noxious weed in three states (Kartesz and Meacham 1999). (C. J. Rothfels 317). * Verbena bonariensis L. Purpletop Vervain VERBE- NACEAE This species is an addition to the flora of Ontario (Newmaster et al. 1998). One plant was found on the Osprey Marsh Christmas tree carp barrier (a row of old Christmas trees erected in the mud to prevent carp from travelling from Grindstone Creek into Osprey Marsh), in the Hendrie Valley Nature Sanctuary, Halton Region (43°17.40'N, 79°52.90'W). It might have ar- rived in the treads of the machinery used to place the trees in the winter. This species is not in the Royal Botanical Gardens’ cultivated plants database, and thus has theoretically never been planted on RBG property. One individual of this species was seen at the Childrens’ Garden in Westdale (not operated by RBG) in 2002, approximately 3.5 km from the Osprey Marsh population. This species is introduced to the southern States, and has also been reported from Ore- gon, New York State, New Jersey, and New Hampshire (Kartesz and Meacham 1999*). It has not been re- ported from Canada. (C. J. Rothfels & I. Vaithilingam 435). A previously-overlooked specimen was also uncovered at HAM. It is an undated specimen from a “river bank” at “Lake Erie,’ and was identified by J. S. Pringle in 1964 (K. Stanley s.n.). In light of the 2002 discovery, it seems reasonable that this record could also have been “spontaneous”. Discussion The eight taxa found new for Hamilton add to the current published list of 1304 species (Goodban 1995), and the four new taxa for Halton bring that region’s tally to 1305 (Varga et al 2000). Additionally, four of the taxa are new for Ontario. Twelve of the fourteen records discussed are non-native occurrences, several of which, especially Aethusa cynapium, Euonymus fortunei, and Sorghum bicolor, should be watched for invasive tendencies. Acknowledgments Thanks to Jim Pringle for assistance in the deter- mination of Aethusa cynapium, Amaranthus blitum, Anthriscus caucalis, Cardamine impatiens, Coronopus didymus, and Macleaya cordata; thanks to Jim Pringle and Michael J. Oldham for helpful earlier reviews of this manuscript, and to Megan Ogilvie, Albert Garofalo and Dean Gugler for submitting their records. Con- tribution from Royal Botanical Gardens Number 1 14. 2004 NOTES 615 Documents Cited (marked * in text) Pringle, J. S. 1994. Notes on some adventive and naturalized Kartesz, J. T., and C. A. Meacham. 1999. Synthesis of the species in the flora of Ontario. Field Botanists of Ontario North American Flora, Version 1.0. Chapel Hill: North Newsletter. 7(4): 10-14. Carolina Botanical Garden. Compact disc. Rothfels, C. J. 2003. Significant 2002 Hamilton Study Area Literature Cited: Edmondson, J. R. 1997. Aesculus. Pages 155-157 in The / European garden flora Volume V. Edited by J. Cullen, J. C. M. Alexander, C. D. Brickell, J. R. Edmondson, P. S. Green, V. H. Heywood, P. —M. Jorgensen, S. L. Jury, S. G. Knees, V. A. Matthews, H. S. Maxwell, D. M. Miller, E. C. Nelson, N. K. B. Robson, S. M. Walters, and P. F. Yeo. Cambridge University Press, United Kingdom. Gleason, H. A., and A. Cronquist. 1991. Manual of vascular plants of Northeastern United States and adjacent Canada. New York Botanical Garden. Bronx, New York. 910 pages. _ Goodban, A. G. 1995. The vascular plant flora of the Regional Municipality of Hamilton-Wentworth, Ontario. First edition: September 1995. Hamilton Region Conservation Authority. 86 pages. Holmgren, N. H. 1998. Illustrated Companion to Gleason and Cronquist’s Manual. New York Botanical Garden. Bronx, New York. 937 pages. Krussmann, G. 1984. Manual of cultivated broad-leaved trees & shrubs. B. T. Batsford Ltd., London. 448 pages. Macoun, J. 1883. Catalogue of Canadian plants: Part 1. Geological Survey of Canada. Ottawa. Mulligan, G. A. 2002. Weedy introduced mustards (Bras- sicaceae) of Canada. Canadian Field-Naturalist. 116: 623- 631. Newmaster, S. G., A. Lehela, P. W. C. Uhlig, S. McMurray, and M. J. Oldham. 1998. Ontario Plant List. Ontario Ministry of Natural Resources, Ontario Forest Research Institute, Sault Ste. Marie, Ontario, Forest Research Information Paper Number 123, 550 pages + appendices. Plant Records from the Royal Botanical Gardens Herbar- ium (HAM). Wood Duck 56: 155-161. Scoggan, H. J. 1978. The flora of Canada. National Museums of Canada. Ottawa. Four Parts; 1711 pages. Smith, T. W. 2003. Checklist of the Vascular Flora of Royal Botanical Gardens. Contribution of Royal Botanical Gar- dens Number 113. Royal Botanical Gardens, Burlington, Ontario. 126 pages. Tiedje, J., and D. Tiedje. 2002. Vascular plants of Lambton County, Ontario. 11' Edition: November 2002. Published by the authors. 66 pages. Tutin, T. G. 1968. Aethusa. Pages 339-340 in Flora Europaea Volume 2. Edited by T. G. Tutin, V. H. Heywood, N. A. Burges, D. M. Moore, D. H. Valentine, S. M. Walters, and D. A. Webb. Cambridge University Press. 455 pages. Varga, S., D. Leadbeater, J. Webber, J. Kaiser, B. Crins, J. Kamstra, D. Banville, E. Ashley, G. Miller, C. Kingsley, C. Jacobsen, K. Mewa, L. Tebby, E. Mosley, and E. Zaje. 2000. Distribution and status of the vascular plants of the greater Toronto Area. Ontario Ministry of Natural Resources, Aurora District. August 2000. 102 pages. Voss, E. G. 1985. Michigan Flora: Part II Dicots (Saururaceae to Cornaceae). Cranbrook Institute of Science Bulletin 59. University of Michigan Herbarium. 724 pages. Wagner, W. H., Jr. 1958. The hybrid ragweed, Ambrosia artemisiifolia x trifida. Rhodora 60: 309-316. Received 10 November 2003 Accepted 21 December 2004 Do Rhinoceros Auklet, Cerorhinca monocerata, Fledglings Fly to the Sea from Their Natal Burrows? JAMES L. HAYWARD and JERE K. CLAYBURN Biology Department, Andrews University, Berrien Springs, Michigan 49104-0410 USA Hayward, James L., and Jere K. Clayburn. 2004. Do Rhinoceros Auklet, Cerorhinca moncerata, fledglings fly to the sea from their natal burrows? Canadian Field-Naturalist 118(4): 615-617. The mode of departure of Rhinoceros Auklet fledglings from their nest burrows has remained uncertain. Both walk-down and fly-down hypotheses have been proposed. Here we use the unique geography of Protection Island, Washington, to eval- uate the fly-down hypothesis. Some fledglings raised on Protection Island do appear to walk to the water, but our results suggest that many of the island’s fledgling Rhinoceros Auklets fly to the sea. Key Words: Rhinoceros Auklet, Cerorhinca monocerata, fledgling departure, Washington. Rhinoceros Auklets (Cerorhinca monocerata) are the most abundant puffins in the waters of the Pacific Northwest. Unlike Tufted and Horned puffins (Frater- cula cirrhata and F: corniculata, respectively), which also breed along western North America, members of most populations of Rhinoceros Auklets leave their nesting colonies before dawn to feed and return after sunset (Wilson and Manuwal 1986; but see Thoresen 1980). Consequently, some basic information on the behavior of these birds remains obscure, despite grow- ing knowledge of their breeding biology (Richardson 1961; Scott et al. 1974; Leschner 1976; Summers and Drent 1979; Vermeer 1978, 1979, 1980; Thoresen 1980, 1983; Wilson and Manuwal 1986; Wilson 1993; Gaston and Duchesne 1996). One elusive piece of information involves the mode of departure from nest sites by newly-fledged Rhinoc- eros Auklets. Young Atlantic Puffins (F. arctica) walk, 616 flutter, or fly to the water from their burrows (Lockley 1934: Richardson 1961; Gaston and Jones 1998), where- as fledgling Cassin’s Auklets (Ptychoramphus aleuti- cus) fly to the sea after making short “practice flights” (Manuwal 1974). Leschner (1976) found Rhinoceros Auklet fledglings that were too small to fly “crouched under rocks or crevices in the morning” on Destruction Island, Washington; this suggested to her “that fledg- lings walk or flutter to the water.” Similarly, Wilson (1977) observed fledgling Rhinoceros Auklets on Pro- tection Island, Washington, “walking down the grass covered slopes” to the sea. But Richardson, (1961) who also worked on Protection Island, wrote that “flight [capability] ... appears to develop precociously,’ and that like the “half-grown young” of murres, fledgling Rhinoceros Auklets “fly down to the water.” On Protection Island, hundreds of Rhinoceros Auk- lets nest along a sandy cliff that rises above Violet Point, a gravel spit that contains a large Glaucous-winged Gull (Larus glaucescens) colony. Each summer, scores of auklet fledglings are found dead within a relatively defined area of the gull colony, across an artificial channel and marina from the cliff. The distribution of dead fledglings and the unique semi-natural geography of Protection Island allowed us to evaluate Richard- son’s fly-down hypothesis. Methods Protection Island, Jefferson County, Washington (48°08'N, 122°55'W), consists of a 35—76-m high pla- teau surrounded by steep, grass-covered, sandy cliffs ideally suited for Rhinoceros Auklet burrowing. The largest Rhinoceros Auklet colony in Washington is located here, with approximately 27 549 burrows (Wil- son and Manuwal 1986). Violet Point, a gravel spit that extends 800 m east from the upper island, contained around 5 100 of Glaucous-winged Gull nests during this study (J. G. Galusha, personal communication). In July and August 1992, 1993, and 1997 the location of each auklet fledgling found dead on Violet Point was plotted by measuring its north or south distance from a transect that measured its eastward distance from the base of the island’s east-facing cliff. To estimate age at death, culmen and tarsal lengths, if still present, were measured in 1992 and 1993. The height of the highest (60 m) nesting burrows on the cliff in relation to the distribution of dead fledglings on Violet Point was used to approximate a range of presumed fledgling flight path angles trigonometrically. Results One hundred and twenty-one dead fledglings (43 in 1992, 11 in 1993, and 67 in 1997) were counted between 106 m east of the cliff base, at the western margin of the gull colony, and 398 m east of the cliff base, toward the center of the gull colony (Figure 1). The fledglings apparently died as a result of hitting the spit or being attacked by resident gulls. No dead fledglings were THE CANADIAN FIELD-NATURALIST Vol. 118 A. Aerial view Strait of Juan de Fuca Protection Island 60-m cliff with auklet burrows Violet Point (Exeter Of quill COL Cy) _—_—_—_——_——_——[$ (478 m - longest estimated fledgling flight) See! (111 m - shortest estimated fledgling flight) bt (292 m - range Of dead fledglings) B. Side view Ficure |. A. Aerial view of Violet Point, Protection Island, Washington. Dots indicate locations of dead fledgling Rhinoceros Auklets found during July and August, 1992, 1993, and 1997. Lines below indicate approximate extent of the spit occupied by the gull colony, extent of fledgling auklet corpses, and estimated extents of fledgling auklet flight distances. B. Side view of Violet Point showing the estimated range of Rhinoceros Auklet fledgling flight paths, sloping from 7°—18° from the highest (60 m) natal burrows above the gull colony; fledgling flights from lower burrows would exhibit progressively shallower angles. found beyond 398 m. The pooled mean (+ SD here and below) distance of fledgling corpses from the base of the nest cliff was 303 + 58.0 m (n = 54, range = 106-398 m). Mean culmen length was 24.4 + 3.0 mm (n = 35), and mean tarsus length was 37.9 + 1.6 mm (n = 48); Rhinoceros Auklets of this size are about 45 days old (Wilson and Manuwal 1986). During late July and early August, we occasionally found live Rhinoceros Auklet fledglings hiding in grass between the nesting cliff and the channel/marina, and we reg- ularly observed fledglings swimming in the water of the marina/channel. Discussion Large numbers of dead fledglings found in gull ter- ritories east of the marina/channel water and across from the nesting cliff support the hypothesis that these fledglings flew down and over the water from their natal burrows. Alternate means of reaching the east side of the marina/channel were highly unlikely: (1) walk/flutter down slope, enter the marina/channel water from the west shore, leave the water on the east shore, enter the gull colony, be killed; (2) walk/flutter 2004 down slope, enter the colony north of the marina/ channel, walk/flutter through a gauntlet of gull terri- tories to the east side of the water, be killed; (3) walk/ flutter down slope, enter the colony north of the marina/channel, be killed, then be transported to dis- tant points east of the water by gulls. With respect to (3), it should be noted that fledging Rhinoceros Auk- lets exhibit an average mass of 360 g (Wilson and Manuwal 1986), 34% of the average mass of 1051 g (Vermeer 1963) for adult Glaucous-winged Gulls in this region; thus it appears unlikely that auklet corpses could be moved very far from the place where they were killed. Our data, with those of Richardson (1961), Lesch- ner (1976), Wilson (1977), and Wilson and Manuwal (1986), support the following scenario for the depar- ture of fledgling Rhinoceros Auklets from their nest burrows on Protection Island: In late July and early August, some fledgling auklets walk and flutter, but also many fly directly away from their natal burrows. Fledglings cannot sustain prolonged flight, however, and from the highest nests lose altitude at a rate of 13-32 m/100-m distance, with a slope of 7°-18° (Figure 1B); fledgling flights from lower burrows would exhibit progressively shallower angles. Fledg- lings from preferred nest sites located directly above the ocean (Richardson 1961) reach the water success- fully. Fledglings that fly from nest sites along the cliff above Violet Point hit the gull colony where they are killed, unless they veer to the ocean or are fortunate enough to land in the marina/channel. Whether this scenario applies to fledgling departures at other Rhi- noceros Auklet colonies remains unknown. Acknowledgments We thank Robert Edens, Jean Takekawa and Kevin Ryan for permission to work on Protection Island National Wildlife Refuge, Ulrich Wilson for valuable discussions about Rhinoceros Auklets, Shandelle Hen- son for comments on the manuscript, and Walla Walla College Marine Station for logistical support during this study. Funding was provided by faculty grants to JLH from Andrews University. Literature Cited Gaston, A. J., and S. B. C. Dechesne. 1996. Rhincoceros Auklet: Cerorhinca monocerata. The Birds of North Amer- NOTES 617 ica (212). American Ornithologists’ Union, Washington, D. C., and Academy of Natural Sciences, Philadelphia, Pennsylvania. 20 pages. Gaston, A. J., and I. L. Jones. 1998. The Auks: Alcidae. Oxford University Press, Oxford. 349 pages. Leschner, L. L. 1976. The breeding biology of the Rhinoceros Auklet on Destruction Island. M.S. thesis, University of Washington, Seattle. 77 pages. Lockley, R. M. 1934. On the breeding habits of the puffin: With special reference to its incubation and fledgling peri- ods. British Birds 27: 214-223. Manuwal, D. A. 1974. The natural history of Cassin’s Auk- let (Ptychoramphus aleuticus). Condor 76: 421-431. Richardson, F. 1961. Breeding biology of the Rhinoceros Auklet on Protection Island, Washington. Condor 63: 456— 473. Scott, J. M., W. Hoffman, D. Ainley, and C. F. Zeillemaker. 1974. Range expansion and activity patterns in Rhinoceros Auklets. Western Birds 5: 13-20. Summers, K. R., and R. H. Drent. 1979. Breeding biology and twinning experiments of Rhinoceros Auklets on Cle- land Island, British Columbia. Murrelet 60: 16-22. Thoresen, A. C. 1980. Diurnal land visitations by Rhinoceros Auklets. Western Birds 11: 154. Thoresen, A. C. 1983. Diurnal activity and social displays of Rhinoceros Auklets on Teuri Island, Japan. Condor 85: 373— SU Vermeer, K. 1963. The breeding ecology of the Glaucous- winged Gull (Larus glaucescens) on Mandarte Island, B. C. Occasional Papers of the British Columbia Provin- cial Museum Number 13. 104 pages. Vermeer, K. 1978. Extensive reproductive failure of Rhi- noceros Auklets and Tufted Puffins. Ibis 120: 112. Vermeer, K. 1979. Nesting requirements, food and breeding distribution of Rhinoceros Auklets, Cerorhinca monocer- ata, and Tufted Puffins, Lunda cirrhata. Ardea 67: 101— 110. Vermeer, K. 1980. The importance of timing and type of prey to reproductive success of Rhinoceros Auklets Cerorhinca monocerata. Ibis 122: 342-350. Wilson, U. W. 1977. A study of the biology of the Rhinoceros Auklet on Protection Island, Washington. M.S. thesis, Uni- versity of Washington, Seattle. 98 pages. Wilson, U. W. 1993. Rhinoceros Auklet burrow use, breeding success, and chick growth: Gull-free vs. gull-occupied habi- tat. Journal of Field Ornithology 64: 256-261. Wilson, U. W., and D. A. Manuwal. 1986. Breeding biology of the Rhinoceros Auklet in Washington. Condor 88: 143— 156. Received 6 February 2004 Accepted 25 October 2004 A Tribute to Thomas Henry Manning 1911-1998 BRENDA CARTER R.R. #4. Merrickville, Ontario KOG 1NO Canada Carter, Brenda. 2004. A tribute to Thomas Henry Manning 1911-1998. Canadian Field-Naturalist 118(4): 618-625. I first met Tom Manning on his Merrickville farm in the spring of 1965, during lambing time. Andrew and Betty Macpherson who had both travelled with him in the Arctic, brought me out to see Arctic Foxes. These captive animals were part of a research program on the influence of light on moult timing. On the way to. Merrickville, I listened to their stories about Tom’s legendary explorations and scientific expeditions. I was therefore prepared for the prospect of meeting an Arc- tic hero. Instead, I found a quiet, stocky man in his fifties, dressed in faded and ragged overalls tending his sheep. The stub of a hand-rolled cigarette hung from his lower lip — perpetually unlit. He did not seem pleased at extra company and bluntly asked why I had been brought out. Later, over glasses of his lethal home- made cider, I realized that here was a man who was much more than he seemed. By the end of that day we had become firm friends — a friendship that was to last until his death in November of 1998. Few knew Tom Manning well; and few indeed were given a chance to comprehend, as a whole, the com- plexity and sheer bulk of his phenomenal life’s work. Except for a very few close friends, Tom was deeply private, and even withdrawn. He was a relentless and dogged scientist who spared no effort to find either the original source of any information, or to redo any questionable measurements — even if they were his own. Applying his logical and inquiring mind to all kinds of life-forms, natural phenomena and expedi- tion challenges, Tom also had the skill, intuition and knowledge to assess and react appropriately in critical situations. He did so with a rapidity I found astonishing. His mental brilliance, awesome physical endurance, good looks and charm, combined with great modesty and fondness for red sportscars, contributed to a re- markable personality. On the 22 of December, 1911, Thomas Henry Man- ning was born into a gentleman’s life. Even as an infant, he defied prediction and showed his amazing capacity for endurance. Except for a surgeon who “did not want to commit murder on Christmas Day” and reluctantly operated, Tom might soon have died, a victim of pyloric stenosis. Tom was the only child of Dorothy (nee Ran- dall) whose brother was the last gentleman jockey in England, and Thomas Edward Manning, (TEM) who managed the family brewery and was cricket Captain for Northamptonshire. Their privileged son was neither destined nor inclined to follow in his father’s foot- steps. Tom once told me that his first childhood memory was of being put out on a blanket in the sun before he could walk. He remembered his excitement and free- dom at being outside. Tom never lost his passion for open air which propelled him into a life of exploration and adventure. His scientific career began and ended with eggs. As a boy, he had a passion for collecting eggs, and this hobby directed him into a career as a biologist, rather than a brewery owner as his father expected. A friendly gardener sparked his interest in nests, and Tom undoubtedly learned to keep notes and data for his tro- phies. This mixture of intense interest and attention to detail stood him in good stead as he completed, at the age of seventy, his last pioneering scientific work on weight lost by eggs during incubation for the Canadian Field-Naturalist. Tom was educated at Harrow, and like his father, attended shooting parties and rode with the famous Pytchley Hunt. There were, however, some interesting dissimilarities. He cleaned the skulls of the mammals and birds shot, drying them in his mother’s linen cup- board. Eventually, he escaped from the hunt meets be- cause he disliked the formalities and confining clothes. While at Harrow, Tom took a cycling trip with a school friend and discovered an unsuspected physical trait. As the pair wound their way through country lanes toward the southern coast of England, he found himself farther and farther ahead. With characteristic independence and detachment from social niceties, he left his friend far behind and continued on, solely interested in how far he could travel in one day. At the end of that journey, he was surprised by his endurance. Although he would never admit it, this ability was more than unusual. After his death, I discovered a buried manuscript, transcribed from a 1931 diary. It gives an account of his first expedition to Iceland and the Faroe Islands with Harrow school friend, “Twitch” Mitchell. Tom’s descriptions of unidentified gulls and seabirds revealed how scrupulous his note-taking was at the young age of twenty. Equally remarkable, for a twenty-year old, was the care he took in logistical planning; from deciding on exact food supplies and arranging trans- 618 CARTER: THOMAS HENRY MANNING Thomas Henry Manning with Massey Medal presented by the Royal Canadian Geographical Society in 1977 with portrait of former Government-General Vincent Massey in background. portation, to sorting out necessary visas. This thor- pithy comments about the unfortunate “Twitch”, dis- oughness remained as long as I knew him. Having _ gruntled bureaucrats and miserable weather. travelled with him, I could also recognize in that early After Harrow, Tom read Natural Sciences at Jesus diary his trademark dry and ironic humour, hidden in College, Cambridge. This academic subject captivat- 620 ed him and provided a continuum with his childhood interest in natural history. Owing to several circum- stances, the most prominent being boredom, he never took his degree. Instead, in the summer of 1932, at the age of twenty-one, he embarked on a serious journey, walking the length of the Atlantic coast of Norway on his own. Joined later by Reynold Bray, his close school friend from Harrow, the pair proceeded east- ward into Finland, making a winter trip by reindeer sledge through Lapland. In 1932, near the end of their trip, they trespassed into Russia. Tom told me they were arrested by villagers armed with pitchforks, who sus- pected them of spying. They subsequently spent three weeks in a Leningrad jail, being fed on increasingly weakened soup, until the British Consul secured their release. Exploration, travel and adventure were now Tom’s calling. From 1933-1935, he made a solitary expedi- tion to Southampton Island, northern Hudson Bay, to study a Snow Goose colony. He mapped the island under the auspices of the Royal Geographical Soci- ety and collected birds and mammals for the British Museum (Natural History). His longest journeys were made by dogsled in winter. From the Inuit of Coral Har- bor (on the island’s southern coast) he learned how to build an igloo, drive a dog team and hunt seal and caribou. When I once asked why he had an aversion to dumplings, he replied that during that first summer he had been given flour and baking powder by the Hud- son’s Bay Company Manager. Given no instructions and being too proud to ask what he should make, he existed on a tiresome diet of dumplings until rescued by the Inuit who showed him how to make bannock. Tom had no sooner returned to England in 1935 than he started to organize a larger expedition to South- ampton Island, and Foxe Basin. The five members of the British-Canadian Arctic Expedition, consisting of Tom, Graham Rowley, Reynold Bray, Pat Baird, Richard (Dick) Keeling and Peter Bennett (who later replaced Dick); arrived in Churchill in May of 1936. A smaller team consisting of Tom, Graham and Pat sailing in the Polecat, a 30-foot open whaleboat, established their main base at Coral Harbor. Each pursued his own investigation in biology, archaeology and geology. The expedition was later marred by the tragic death of Reynold Bray, who drowned after being blown out to sea in a collapsible boat. By the summer of 1938, Tom was alone in the field, at a camp in the Cape Dorset area of southern Baffin Island. He sent a message by Inuit dog team and radio to Montreal to Miss Ella Wallace Jackson (Jackie), a nurse from Nova Scotia whom he had met on the ship that took him back to England in 1935. He invited her to join him in the north. Jackie accepted, and the day after she arrived they were married, using a ring made from a brass fitting by the ship’s engineer. A few days later they headed north in a whaleboat. For the next eighteen months they conducted mapping and zoolog- THE CANADIAN FIELD-NATURALIST Vol. 118 Tom Manning photograph by Borg Mesch, Kiruna, north- ern Sweden, 1932. ical studies on the west coast of Baffin Island. Jackie later published two books, /gloo for the Night, The University of Toronto Press, 1946 and A Summer on Hudson Bay, Hodder and Stoughton Ltd., 1949 about their travels together. Although Jackie and Tom spent many years together, the couple separated in the 1960s. Tom often visited her in Ottawa where she still resides. Late in 1939, Tom had a dream that caused him to head south; he and Jackie arrived in Cape Dorset on 2 January 1940 to learn that World War II had been rag- ing for four months. Anxious to join the war effort, they discovered that the annual resupply ship — due in August — could not get them to Montreal before October. Tom decided they would make their own way back, continuing around the Foxe Basin coast of Baffin Island, hoping to reach Churchill by August. It was 20 January 1941, however, before they reached Churchill, after an epic journey of nearly 2500 miles by boat and dog team, mapping as they went. Tom received a commission in the Royal Canadian Navy Voluntary Reserve, but was soon seconded to the U.S. Army Engineers to provide advice on the sit- CARTER: THOMAS HENRY MANNING 62) Tom Manning and Reynold Bray (1911-1936) on expedition 21 October 1932. Photograph T. Dahill of Kiruna, northern Sweden. | ing, construction and operation of an airfield and | weather stations on Southampton Island. He spent the | rest of the war on defence-related geodetic surveys in | the north, as well as the design and development of | Arctic clothing and equipment for the armed services. Tom served in the navy from 1941 to 1945, leaving | with the rank of Lieutenant Commander. Following the war, Tom’s services were sought by the Defence Research Board, the Geodetic Survey, the _ Geographical Branch, and other agencies of the Cana- _ dian Government. Assignments included the exploration _ of islands discovered in Foxe Basin by air photographs in 1948, a survey of Banks Island, an oceanographic survey in the Beaufort Sea, and many coastal surveys. Throughout his travels, Tom continued to collect spec- _imens of birds, mammals, plants and occasionally _ insects. The National Museum of Canada was one of the fortunate recipients of these valuable specimens. _ In 1949 Tom introduced the seventeen year old Andrew Macpherson to the Arctic. They became fast friends and made several expeditions together, collaborating on many reports of their findings. These journeys, with their combined achievements in the fields of sci- ence and exploration, are well known. Accounts of Tom’s work have appeared over the last few years in newspapers and journals around the world. In these articles writers have paid extensive tribute to his work. Many of his colleagues and acquaintances noticed that Tom was as much at ease at a formal dinner as on the snow-scoured rock of the Arctic Barrens. However, he preferred the familiar landscape of the Barrens and was able to make himself comfortable in surroundings and circumstances that many civilized people could not endure. After leaving the North, in 1941, Tom lived briefly in Montreal. Later, he and his wife bought a house in Ottawa on Linden Terrace where they entertained many of their northern friends. Tom started the Arctic Circle Club, 14 November 1947, when the guests no longer fitted around the huge oak dining-table that he had made himself. He was an expert craftsman, making cabinets for his house and building book shelves for his huge library. He and Jackie also spent hours patiently binding many of the collected volumes. When I first met him in 1965, Tom had given up his Arctic exploration and much of his scientific work. In the tradition of his grandfather T. A. Manning (TAM), he had become a farmer, an occupation which had 622 always interested him. However, Tom’s grandfather in Northampton, had forsaken farming and instead, found- ed a brewery continued by his son TEM. The prospect of this confining tradition was one of the forces that drove Tom to some of the most remote regions on the planet. However, he made some reconciliation with his heritage about 1960, by purchasing and working over 500 acres of hayfield, pasture and woodlot in the Merrickville countryside, some fifty km south of Otta- wa. He, and his Arctic friends, built a cabin on the largest portion of property which he also used for pasture. Tom went on to renovate a stone house and rework the fer- tile land into a self-sustaining farm with a huge and immaculate vegetable garden, cattle, sheep, chickens and geese. Tom and I talked a great deal about his northern trips. I finally convinced him that he should go back to his scientific work, and that I would make an excellent assis- tant! He cut back on the livestock and farming opera- tions, making them manageable for summer and occa- sional fieldwork. Our first trip was to western Canada collecting birds, mammals and plants. We travelled in a truck, which Tom had skillfully rebuilt as a home on wheels, complete with woodstove. He said it remind- ed him of caravan holidays as a child, and he always took immense pleasure in its rugged simplicity — often preferring to sleep in it, even when it was parked at home. My suggestion led to four decades of collaboration. I usually acted as assistant, and collected and prepared the birds with the hunting and skin preparation skills that Tom had taught me. There was ample opportunity for me to paint landscapes and wildlife studies on all the expeditions we made; and Tom was always encour- aging and supportive of my work. Occasionally I could provide help with illustrations or maps for his scientific papers. My first trip North with him was to Cape Henrietta-Maria, tagging polar bears for the Canadian Wildlife Service, under the direction of Dr. Charles Jonkel. Although I saw no bears, I fell in love with the land and, from then on, understood and shared Tom’s passion for this fabulous country. I was soon to see many bears, and we worked together from 1968 to 1973 on the Circumpolar Polar Bear Project. During these travels, I had a firsthand chance to see Tom’s unique skills. As always, he collected as many types of specimens as he could, and kept extensive field notes and descriptions of everything we encoun- tered. Unlike many Arctic explorers of his time and previously, Tom had a broader range of interests, and documented everything he considered important. At the time, I did not fully comprehend either Tom’s skills or his decision-making process. Later, through field experience, I began to understand the brilliance of his mind and abilities. On one occasion he piloted an open boat in dense fog through seven miles (about 11 km) of the unforgiving waters of James Bay, using dead reckoning and expe- THE CANADIAN FIELD-NATURALIST Vol. 118 = Tom with beached hull of former 1949 expedition ship Nauya behind him, Igloolik, 1983, Photograph by Brenda Carter. rience to arrive at South Twin Island, exactly on target. On another occasion, attempting an open water cross- ing on the Belcher Islands, we ran into a dangerous pack-ice jam. This hazard was further compounded by an early snowstorm and a five-foot (1.5 m) tide rip. To stay meant being stranded. To leave meant facing rough water in an overloaded canoe. Tom’s choice lit- erally meant life or death. As usual, he made the right decision — elegant in its simple, straightforward truth. Despite the blinding snow he found a path through the heavy waves and got us safely ashore. He always kept part of his mind tuned to the essential core of any sit- uation, a trait which was the quiet binding of his life. From 1975 to 1986, Tom continued his collecting trips by truck in the summer, spending winters writing papers (a task he disliked) interspersed with his favour- ite seasonal activities of gardening, cutting fence-posts and filling the woodshed from his hardwood bush. Renewing his interest in his boyhood pursuit of eggs, he began to locate and weigh the eggs of different species. He could often be seen in nearby swamps with his odd, long-legged wooden table, a set of scales and an umbrella to shade the eggs. We both enjoyed the nest hunting and Tom became very interested in the results of his weight loss experiments, but the statistics proved daunting. I often wonder if a modern comput- 2004 er would have helped him with his puzzles. However, his resulting papers were successful, and the most re- | quested from scientists around the world. In 1983 Tom and I made our last trip north together. As we left Iqaluit for Igloolik, we had a poignant ex- change. Looking down at the shorelines and islands that | Tom had explored and mapped as a young man, | said that I thought I had come into his life fifteen years too late, and asked him what he felt as he looked down. He | said that he thought he also had arrived in the North fifteen years too late! At the age of seventy-five, he | continued to dream of an unmapped place and an era when the Arctic was truly beyond the reach of ordi- nary travel. On that trip, I noticed that Tom was walking badly; _ and for the first time since I met him, I realized that I | was able to outwalk him. This was startling and worry- | ing and we soon learned that he had Parkinson’s dis- | ease. However, he continued to work with great energy | on the farm. When Parkinson’s disease forced him to | give up his Arctic work, Tom donated his extensive col- lection of rare Arctic books along with his field journals and papers to the newly formed library in Iqaluit. He | also made a very generous gift of one million dollars | to the University of Cambridge towards their new lib- | rary at the Scott Polar Research Institute. He is remem- | bered there in a special room, the “Manning Archives”. Then, following a broken hip, Tom had to have an | operation. This trauma caused the Parkinson’s disease | to accelerate. After 10 years of his usual bravery and | independence, Tom became extremely ill. However, he | exhibited the same stoic patience that he had while | waiting out a blizzard. One of his last projects was to | slowly dig two new flower beds. His dry humour never | vanished, but the struggle became very difficult. In | November of 1998, at the age of 86 he died. He was in | hospital in Smiths Falls near his home, and remarkably | at peace. He left many legacies behind, including a | gift of $100 000 to the Ottawa Field Naturalist Club. | The council decided that 80% of interest from this bequest would go to assist authors with publication charges for the Canadian Field-Naturalist, primarily _ for northern papers. Tom was a renowned Arctic explorer and zoologist | whose life was characterized by a love of adventure, _ self-reliance, and an ever-inquiring mind. In fact as | Graham Rowley describes in his wonderful book, Cold Comfort: My Love Affair with the Arctic, (McGill-— Queen’s University Press, 1996) what Tom accom- _ plished will never be equalled. He was also a wonder- ful companion and brilliant teacher, showing by exam- ple how to use intuition and logic to unravel a crisis and how to see both facts and beauty in the natural world. | He was quiet to the point of being taciturn, and slow to | join in the mindless customs of society. But, his loy- alty to his friends and to his family in England was complete. Those who were privileged to know him well miss him in many, many ways. CARTER: THOMAS HENRY MANNING 623 Other tributes to Tom have appeared in several news- paper articles as well as an obituary in Arctic 52 (1): 104-105 [March 1999] by Andrew H. Macpherson. Tributes to his disciple Andrew Hall Macpherson (1932- 2002) and Manning’s role as his mentor have appeared in The Daily Telegraph (London, England) 22 June 2002 and in Arctic 55(4): 403-406 (December 2002) by Frank L. Miller. Awards 1944 — Bruce Medal of the Scottish Geographical Society and the Philosophical Society of Edinburgh. 1948 — Patron’s Medal of the Royal Geographical Society 1959 — Guggenheim Fellowship held for Arctic research 1974 — Officer of the Order of Canada 1977 — Massey Medal of the Royal Canadian Geographical Society 1977 — Queen’s Jubilee Award 1979 — Honorary LLD from McMaster University 1992 —Doris Huestis Speirs Award in recognition of his pioneering work on birds in the Canadian North Positions in Scientific and Professional Societies Fellow: Royal Geographical Society; Royal Canadian Geo- graphical Society (also a term as Director) Arctic Institute of North America (also Executive Director 1955-1956). Member: American Ornithological Union; Wilson Ornitho- logical Club; Cooper Ornithological Club; Wildlife Society; American Society of Mammalogists; Society for American Archaeology. Founder: Arctic Circle Club; 1947 (also Secretary 1948-1950) Research Associate: Canadian Museum of Nature (1970-1990) Bibliography Publications Manning, T. H. 1936. Some notes on Southampton Island. Geographical Journal 88: 232-242. Manning, T. H. 1942. Remarks on the physiography, Eskimo and mammals of Southampton Island. Canadian Geograph- ical Journal 25: 17-35. Manning, T. H. 1942. Notes on some fish of the eastern Canadian Arctic. Canadian Field-Naturalist 56: 128-129. Manning, T. H. 1942. Blue and Lesser Snow geese on South- ampton and Baffin Islands. Auk 59: 158-175. Manning, T. H. 1943. Notes on the coastal district of the east- ern barren grounds and Melville Peninsula from Igloolik to Caper Fullerton. Canadian Geographical Journal 26: 84-105. Manning, T. H. 1943. The Foxe Basin coasts of Baffin Island. Geographical Journal 101(5, 6): 225-251. Manning, T. H. 1943. Notes on the mammals of south and central west Baffin Island. Journal of Mammalogy 24: 47-59. Manning, T. H. 1943. Notes on the birds of Southampton Island, Baffin Island and Melville Peninsula. Auk 60: 504- 536. [prepared partially from the notes of R. J. O. Bray, who died on the expedition] Manning, T. H. 1944. Hunting implements and methods of the present day Eskimos of north-west Hudson Bay, Mel- ville Peninsula, and south-west Baffin Island. Geographi- cal Journal 103: 137-152. Manning, T. H., and E. W. Manning. 1944. The preparation of skins and clothing in the eastern Canadian Arctic. Polar Record 28: 156-169. 624 Manning, T. H. 1944. Lesser Snow and Blue Geese on South- ampton Island. Auk 61: 146-147. Manning, T. H. 1946. Ruins of Eskimo stone houses on the east side of Hudson Bay. American Antiquity 11(3) 201- 202. Manning, T. H. 1946. Bird and mammal notes from the east side of Hudson Bay. Canadian Field-Naturalist 60: 71-85. Manning, T. H. 1947. Explorations on the east coast of Hud- son Bay. Geographical Journal 109: 58-75. Manning, T. H. 1948. Eskimo stone house ruins on the east side of Hudson Bay: A correction. American Antiquity 13 (3). Manning, T. H. 1948. Pipestems of the Caribou Eskimos. American Anthropologist 50: 162-163. Manning, T. H. 1948. Notes on the country, birds and mam- mals west of Hudson Bay between Reindeer and Baker lakes. Canadian Field-Naturalist 62: 1-28. Manning, T. H. 1949. The birds of northwestern Ungava. Pages 155-224 in A Summer on Hudson Bay by Mrs. T. H. Manning. Hodden and Stoughton Ltd. Manning, T. H. 1949. The varying lemming in captivity in Ottawa. Arctic Circular 3: 20-21. Manning, T. H. 1950. Tidal observations in Arctic waters: Notes on the tides along the south Hudson Bay and west James Bay coasts. Arctic 3: 95-100. Manning, T H. 1950. Eskimo stone houses in Foxe Basin. Arctic 3: 108-112. Manning, T. H. 1950. A mixed Cape Dorset-Thule site on Smith Island, east Hudson Bay. Annual Report of the National Museum of Canada 1949-50, Bulletin 123: 64-71. Manning, T. H., et al. 1950. The voyage of C. G. M. V. Nauja to Foxe Basin in 1949. 158 pages, plus maps, photographs. Manning, T. H. 1951. Remarks on the tides and driftwood strand lines along the east coast of James Bay. Arctic 4: 122-130. Manning, T. H. 1952. Birds of the west James Bay and south- erm Hudson Bay coasts. National Museum of Canada Bul- letin 125: 1-114. Manning, T. H., and D. F. Coats. 1952. Notes on the birds of some James Bay islands. Annual Report of the National Museum Canada 1950-51, Bulletin 126: 195-207. Manning, T. H., and A. H. Macpherson. 1952. Birds of the east James Bay coast between Long Point and Cape Jones. Canadian Field-Naturalist 66: 1-35. Manning, T. H. 1953. Narrative of an unsuccessful attempt to circumnavigate Banks Island by canoe in 1952. Arctic 6: 170-197. Manning, T. H. 1953. Notes on the fish of Banks Island. Arctic 6: 276-277. Manning, T. H. 1954. Remarks on the reproduction, sex ratio, and life expectancy of the varying lemming, Dicrostonyx groenlandicus, in nature and captivity. Arctic 7: 36-48. Manning, T. H. 1956. The northern red-backed mouse, Cleth- rionomys rutilus (Pallas), in Canada. National Museum of Canada Bulletin 144: 1-67. Manning, T. H. 1956. Narrative of a second Defence Research Board expedition to Banks Island, with notes on the coun- try and its history. Arctic 9(1, 2): 1-77. Manning, T. H., E. Q. Hohn, and A. H. Macpherson. 1956. The birds of Banks Island. National Museum of Canada Bulletin 143: 1-144. Manning, T. H., and A. H. Macpherson. 1958. The mammals of Banks Island. Arctic Institute of North America Tech- nical Paper 2: | -74. THE CANADIAN FIELD-NATURALIST Vol. 118 Macpherson, A. H., and T. H. Manning. 1959. The birds and mammals of Adelaide Peninsula, N.W.T. National Museum of Canada Bulletin 161: 1-63. Manning, T. H. 1960. The relationship of the Peary and bar- ren ground caribou. Arctic Institute of North America Tech- nical Paper 4: 1-52. Manning, T. H. 1961. Comments on “Carnivorous Walrus and Some Arctic Zoonoses”. Arctic 14(1): 76-77. Manning, T. H., and A. H. Macpherson. 1961. A biological investigation of Prince of Wales Island, N.W.T. Transactions of the Royal Canadian Institute 33: 116-239. Macpherson, A. H., and T. H. Manning 1961. Pack dogs in the Canadian Arctic. Polar Record 10: 509. Manning, T. H. 1964. Age determination in the polar bear, Ursus maritimus Phipps. Canadian Wildlife Service Occa- sional Paper 5: 1-12. Manning, T. H. 1964 Geographical and Sexual Variation in the Long-tailed Jaeger, Stercorarius longicaudus Vieillot. Biological Papers of the University of Alaska. (7): 1-16. Manning, T. H. 1969 Wreckage on Banks Island. Beaver Autumn: 21. Manning, T. H. 1971. Geographical variation in the polar bear Ursus maritimus Phipps. Canadian Wildlife Service Report Series 13: 1-26. Manning ,T. H. 1974. Variations in the skull of the bearded seal. Biological Papers of the University of Alaska (16). Manning, T. H. 1976. Birds and mammals of the Belcher, Sleeper, Ottawa and King George islands, Northwest Ter- ritories. Canadian Wildlife Service Occasional Papers 28: 1-42, maps, tables. Morrison, R. I. G., T. H. Manning, and J. A. Hager. 1976. Breeding of the Marbled Godwit, Limosa Fedoa, in James Bay. Canadian Field-Naturalist 90: 487-490. Morrison, R. I. G., and T. H. Manning 1976. First breed- ing records of Wilson’s Phalarope for James Bay, Ontario. Auk 93(3): 656-657. Manning, T. H., and B. Carter. 1977. Incidence of runt eggs in the Canada Goose and Semipalmated Sandpiper. Wilson Bulletin 89: 469. Manning, T. H. 1978. Measurements and weights of eggs of the Canada Goose, Branta canadensis, analyzed and compared with those of other species. Canadian Journal of Zoology 56: 676-687. Manning, T. H. 1979. Density and volume corrections of eggs of seven passerine birds. Auk 96: 207-211. Manning, T. H. 1981. Analysis of weight lost by eggs of ele- ven species of birds during incubation. Canadian Field- Naturalist 95: 63-68. Manning, T. H. 1981 Birds of the Twin Islands, James Bay, N.W.T., Canada. National Museum of Natural Sciences, Syllogeus 30: 1-50. Manning, T. H. 1982. Daily Measurements of Variation in weight loss of eggs of seven Passerine Species before and during natural incubation. National Research Council of Canada 60: 3143-3149. Barker, A. J., J. L. Eger, R. L. Peterson, and T. H. Manning. 1983. Geographic variations and taxonomy of Arctic Hares. Acta Zoologica Fennica 174: 45-48. Unpublished Reports Manning, T. H. 1944. Descriptive report to the Geodetic Service of Canada on a survey of the east Hudson Bay coast. 64 pages plus maps, photographs. 2004 ' Manning, T. H. 1945. Report on Exercise Lemming to the Geodetic Service of Canada. 21 pages plus 4 pages, plus photographs. |Manning, T. H. 1945. Descriptive report to the Geodetic Service of Canada on a survey of northern Manitoba and southeast Keewatin. 54 pages, plus photographs. Manning, T. H. 1945. Technical report to the Geodetic Ser- vice of Canada on a survey of northern Manitoba and southeast Keewatin. 179 pages, plus maps, photographs. Manning, T. H. 1946. Report to the Geodetic Service of Canada on Exercise Muskox and on the Cambridge Bay ___ observation station. 25 pages, plus photographs. | Manning, T. H. 1946. Descriptive report to the Geodetic Service of Canada on a survey of Ungava, east Hudson Bay, and the south coast of Hudson Strait. 133 plus 49 pages, photographs and maps. | Manning, T. H. 1946. Technical report to the Geodetic Ser- vice of Canada on a survey of Ungava, east Hudson Bay, | and the south coast of Hudson Strait. 90 pages, maps. | Manning, T. H. 1947. Preliminary report on a background | study of the caribou, Rangifer caribou caribou (Gmelin) and Rangifer arcticus caboti Allen of the Labrador Peninsula and the Province of Quebec north of the St. Lawrence. (Mimeo). | Manning, T. H. 1947. Descriptive report to the Geodetic Ser- vice of Canada on a survey of the west James Bay and southern Hudson Bay coasts. 166 pages, plus maps, pho- tographs. | Manning, T. H. 1947. Technical report to the Geodetic Ser- vice of Canada on a survey of the west James Bay and south- ern Hudson Bay coasts. 64 pages, plus maps. | Manning, T. H. 1948-49. Reports to Defence Research Board. 1. South coast of Hudson Bay, 147 pages, plus photographs, maps, charts, etc. II. West side of James Bay, 185 pages, plus photographs, etc. FV. East coast of James Bay, 295 pages, plus photographs, etc. Professional Experience | 1931 (summer) — Traveled in Iceland and Faro Islands. 1932-33 (winter) — Traveled through Norway, Sweden, Fin- land, and Russia (Bergen to Murmansk) on foot and with reindeer. | 1933-35 — Mapped Southampton Island and also studied and made collections of birds and mammals. 1936-41 — Leader, surveyor and ornithologist on the British Canadian — Arctic Expedition, working in Southamp- ton and Baffin islands and Repulse Bay areas: includ- ed two years in Foxe Basin which was circumnavi- | gated by whale boat. 1941-45 — Royal Canadian Navy. Seconded as an advisor to the U.S. Army in connection with the sighting and con- struction of the air field and establishment by tractor train of outlying weather stations on Southampton | Island; and later to the Geodetic Service (Canada) to | fix ground controls for air photographic survey of northern Canada (until 1947). | 1944 — Geodetic work in Ungava by aircraft and Peterhead | boat. j 1945 — Geodetic work in west Hudson Bay by aircraft. | 1946 — Geodetic work in Ungava, east Hudson Bay and west Hudson Strait by aircraft and Peterhead boat. _ 1947 — Geodetic work by canoe along the west James Bay and southern Hudson Bay coasts. CARTER: THOMAS HENRY MANNING 625 1948 — Consultant to Defence Research Board. 1949 — Leader, Geographical Bureau’s expedition in C. G. M. V. Nauja to “new” islands in Foxe Basin: includ- ed circumnavigation of Foxe Basin. Soundings and tidal observations taken. 1950 — Canoe journey to collect birds and mammals along east James Bay. 1951 — Leader, Defence Research Board expedition to the Beaufort Sea and Master C.G.M.V Cancolim (includ- ed “circumnavigation” of Alaska). About 4000 miles of soundings taken. 1952 — Leader, Defence Research Board expedition to Banks Island: included attempted circumnavigation of the Island by canoe. 1953 — Completion of circumnavigation of Banks Island. 1954 — Worked up results of Banks Island Expedition under contract with Defence Research Board, and others. 1960 (June to July) — West Coast of North America with Diana Rowley 1967 — Introduction of caribou from Coats Island to South- ampton Island for Canadian Wildlife Service. 1968 (June 14 — August) — Western Canada collecting by truck assisted by Brenda and Charlotte Carter; (Sep- tember) — Cape Henrietta-Maria with Brenda Carter, Dick Russell and Chuck Jonkel for Canadian Wildlife Service. 1969 — Alaska collecting by truck assisted by Z. E. Ellshoff 1970 (July 21 — August) — South Twin Islands, polar bear trapping with Brenda Carter for Canadian Wildlife Service. : 1971 (May 2 — September) — Belcher Islands, Sleeper Islands and Ottawa Islands with Brenda Carter for Canadian Wildlife Service. 1972 (May — June) — UK and Norway measuring bear skulls with Charlotte Carter; (September 10 — Octo- ber 18) -North Twin Island, polar bear trapping with Brenda Carter for Canadian Wildlife Service. 1973 (March 22 — April 17) — Fort George and Belcher Islands, helicopter polar bear tagging with Brenda Carter for Canadian Wildlife Service; (May 2 — Au- gust 1) — North Twin Island, polar bear work with Brenda Carter for Canadian Wildlife Service; (August 13 — September 12) — Foxe Basin helicopter work with Brenda Carter for Canadian Wildlife Service. 1975 (May 15 — July 16) — Collecting at North Point, James Bay. 1977 — Yukon collecting by truck and making a floral and faunal inventory of the Dempster Highway assisted by Jan Rowell and Wendy Earl (who departed Daw- son City). Robert Porsild and his wife were visited. 1980 — New Brunswick collecting by truck. 1982 — Manitoulin Island collecting by truck assisted by Doug Perkins 1983 — Igloolic Wildlife description. Advisor to adult Inuit for Igloolic Education Council. 1983 — USA (Mississippi, Louisiana, Nevada, Utah and Texas) collecting by truck assited by S. C. Tuutz. 1986 — Yukon collecting by truck (age 77) assisted by Brid- get Ryan. Most of the specimens collected on these trips were sold either to the National Museums of Canada or Carleton University. A Tribute to Victor Kent Prest 1913-2003 A. S. DyKE!. D. A. Hopcson!, E. L. BouSFIELD?, and R. E. BEDFORD? : Terrain Sciences Division, Geological Survey of Canada, 601 Booth Street, Ottawa, Ontario K1A OE8 Canada ~ 301-540 Dallas Road, Victoria, British Columbia V8V 4X9 Canada ; 524 Penhill Avenue, Ottawa, Ontario K1G 0V6 Canada Dyke, A. S., D. A. Hodgson, E. L. Bousfield, and R. E. Bedford. 2004. A Tribute to Victor Kent Prest 1913-2003. Canadian Field-Naturalist 118(4): 626-629. With the death of Dr. Victor Kent Prest on 2003 Sep- tember 26, Canada lost one of its most distinguished, best-known, and most influential Quaternary geolo- gists. Vic was active as a leader of Canadian Quater- nary science for a remarkably long time and he posi- tively influenced the careers of almost everyone else in the field in this country as well as his many friends abroad. His work became basic to discoveries by other scientists in related fields such as plant and animal biogeography, both terrestrial and aquatic*>°. He came to personify the Quaternary Geology of Canada both domestically and internationally. This identity arose from his unstinting and deep interest in the subject, his long view of the evolution of ideas, his intimate knowledge of the ever-growing knowledge base, and especially the very genuine interest he took in the work of all others. Tentative, junior researchers and graduate students always met with as much respect and encour- agement as did his senior colleagues and Vic often took the time to send notes of encouragement and compli- ment. Victor K. Prest was born in Edmonton Alberta, 2 April 1913, the only child of John (Jack) Prest and Elizabeth Buckley (Prest). His early education was obtained at the MacKay Avenue School in Edmonton. In 1925 the family moved to Toronto for a brief period, and relocated in London, Ontario, where Vic completed his early schooling at Tecumseh Public School and London South Collegiate High School. In 1930 the Prests moved to Winnipeg where Victor completed his secondary education at Kelvin High School. Unreal- ized by Vic, his future wife Patricia Horder was also attending Kelvin, in the same grade, but in a different classroom. Her father, a Canadian Pacific Railway offi- cial, had been transferred from Montreal to Winnipeg in 1926. Vic received his B. Sc. (Honours) from the University of Manitoba in 1935 and his M. Sc. a year later. Growing stronger during these university days was his friendship with Patricia Horder who was active in music, drama, and art, and in the Canadian Girl Guide Association. Vic worked as a summer assistant for the Manito- ba Mines Branch in 1934 and the Geological survey of Canada in 1935 and 1936. While registered for Ph.D. studies at the University of Toronto, Vic was Party Chief, Ontario Department of Mines, from 1937- 1940, mainly mapping bedrock outcrops in Northern Ontario. In 1941 Vic received his Ph.D. degree at “Tor- onto” where he also served as an assistant in both the Geology Department and the Royal Ontario Museum and later as Lecturer. The newly minted Dr. Prest joined the International Nickel Company as a geolo- gist for a year (1941-1942). Vic and Pat were married in Toronto in 1942 and moved immediately to Harbour Grace, Newfoundland, where Vic served with the RCNVR from 1942-1945. Their first child, Sherron (Sherry) Gail was born there in 1944. Following the surrender of Nazi Germany in early May, 1945, Lieutenant Prest was a member of the naval party accepting the surrender of submarine U190 in St. John’s Harbour. During the next five years Vic was a permanent staff member of the Ontario Department of Mines, based in Toronto, where his son Wayne Horder Prest was born in 1946. In 1950, he transferred to the Geological Sur- vey of Canada in Ottawa. Vic served as Chief of the Pleistocene Engineering and Groundwater Section, first in a third-floor office of the Victoria Memorial Museum building and, following its completion in 1960, in the newly constructed Geological Survey building on Booth Street. Vic formally retired from government service in 1979, but maintained a con- sultantship office at the Booth St. building until very recently. Dr. Victor K. Prest is probably best known because of the Glacial Map of Canada (Prest et al. 1968), a work of such veracity that it has yet to be superceded for the country as a whole. This map followed an ear- lier version initiated by J. Tuzo Wilson, to which Vic contributed as co-compiler (Wilson et al. 1959). The 1968 Glacial Map was followed by the beautiful blue- shaded “pancake map” depicting the Retreat of Wis- consin and Recent Ice in North America (Prest 1969), and his masterful Quaternary Geology chapter in the Geology of Canada volume (Prest 1970). That trilogy was the masterpiece of a generation, which adorned many an academic hallway and classroom. Its impact places it among the most outstanding contributions of the Geological Survey of Canada (GSC) and fixes it as its premier Quaternary contribution. It was the most 626 2004 DYKE, HODGSON, BOUSFIELD, and BEDFORD: VICTOR KENT PREST 627 VICTOR KENT PREST cogent, persuasive, and attractively illustrated synthesis available and it has served as the baseline for almost everything else that has followed. As a great builder of Quaternary science in Canada, Vic had wide knowledge of geological and other sci- _ ences. He resumed mapping the Precambrian geology of northern Ontario for ODM (1945-1950) where he authored more than 30 maps and annual reports. He was one of the first to combine Quaternary and bedrock mapping with the release of his map of Red Lake- Lansdowne House. His first Quaternary publication was the Pleistocene geology of the Vermillion River system with special reference to placer gold (Prest 1949). Vic’s early Pleistocene colleagues at GSC includ- ed Jack Armstrong, Wes Blake, Jr., Bruce Craig, Bob Fulton, John Fyles, Nelson Gadd, Eric Henderson, Owen Hughes, Bert Lee, and Archie Stalker, all of whom were to become major regional authorities, and Jaan Terasmae and Bob Mott, leaders in the develop- ment of Quaternary paleoecology in Canada. Addi- tion of his academic and provincial geological survey friends, Bill Matthews (University of British Colum- bia), Earl Christiansen (Saskatchewan), Alexis Drei- manis (University of Western Ontario), Con Gravenor (University of Windsor), Paul Karrow (Waterloo Uni- versity), Pierre LaSalle (Quebec), and John Elson (McGill University) largely completes the Prest cohort of Canadian Quaternary geologists. Although he departed senior management for full- time research in 1964, even those who joined the Sur- 628 vey later, and many in outside agencies, identified the Quaternary group with Vic Prest more than with any- one else. Vic became a sort of mentor-at-large, with great moral authority and prestige, and engendered pride in the work of others. By building Quaternary science at the Survey and by mentoring and infor- mally supervising graduate students, Vic profoundly contributed to the Canadian Quaternary community. Vic was an enthusiastic participant in numerous field excursions in southern Canada, the northern USA, Alaska, and internationally. His GSC fieldwork took him on a High Arctic voyage that reached Ellesmere Island (Prest 1958), mapping projects in Ontario (Prest 1963), Quebec (Prest 1966; Prest and Hode-Keyser 1977) and the Maritimes. He had a special passion for the Maritimes, where his map of Prince Edward Island (Prest 1973) remains seminal, and where, with his pro- tégé Douglas Grant, he formulated the concept of an Appalachian Ice Complex (Prest and Grant 1969; Prest et al. 1972; Prest 1977) to encapsulate the style of regional glaciation, as distinct from Laurentide glacia- tion. This model is still favoured by regional geolo- gists. The Magdalen Islands in the Gulf of St. Law- rence long held his interest, because of the perplexing lack of direct evidence of Late Wisconsin glaciation and its record of older events (Prest et al. 1976). This evidence led him to portray on his ice recession map what Doug Grant termed “Prest’s Problematical Pleis- tocene Prong”, a reference to the glaciological improb- ability of leaving the Magdalen Islands ice free while allowing ice to extend to the edge of the Continental Shelf. The portrayal revealed Vic’s preference for field evidence over theory. Vic formally retired from GSC in 1978, and formed Veekay Consultants (1978-1994) to receive Ontario Geological Survey (OGS) surficial mapping contracts in northwestern Ontario and a Department of Indian Affairs and Northern Development contract to work in the Bebensee Lake area (Prest 1985). He continued fieldwork into his seventies, including long traverses into dense bush around the famous Red Lake gold mining camp. This work yielded a series of detailed maps (e.g., Prest 1980) that formed the basis for recent drift prospecting programs. His major syntheses includ- ed Canada’s Heritage of Glacial Features (Prest 1983), the Late Wisconsin Glacier Complex paper and map (Prest 1984), a synthesis of glacial lake history in the Lake Ontario basin with Ernie Muller (Muller and Prest 1985), and maps of North American paleogeog- raphy (Dyke and Prest 1987). His two final publications dealt with the history of interpretation of the Laurentide Ice Sheet (Prest 1990) and a more specific passion, the long-distance dispersal of the distinctive erratics that he termed “omarsi” from the central part of the ice sheet (Prest et al. 2000). Vic was an accomplished athlete and maintained a life-long interest in several sports. During the half cen- tury of his residence in Ottawa, Vic’s love of tennis THE CANADIAN FIELD-NATURALIST Vol. 118 became super-seded by a passionate devotion to curl- ing. He was a charter member of the City View Curl- ing Club, founded in 1957, and President the follow- ing year. Over the years, he won many curling bonspiels both in men’s and mixed rink events, and was ever a formidable shot-maker and curling strategist. He was Honorary Governor General’s Curling Club President (1972), Canadian Branch (Royal Caledonia Curling Club), Council of Management (1978-1980) and Life Member (1986). With support from the Ottawa Val- ley Curling Association, Vic introduced Junior Curl- ing in Eastern Ontario in 1975, and co-ordinated play with the Canadian Branch in Quebec. He also intro- duced “Little Rock” curling in 1983, and organized Junior and Little Rock bonspiels for the OVCA over a period of ten years. After more than 50 continuous years of winter curling, he “pushed” his last rock at age 89. Vic’s professional career accomplishments received due recognition in the form of the Johnston Medal from the Canadian Quaternary Association (the first awarded), the Gold Medal from the Royal Canadian Geographical Society, the Kirk Bryan and the Distin- guished Career Awards from the Quaternary Geology and Geomorphology Division of the Geological Society of America (1993), election as a Fellow to join the distinguished scientists of the Royal Society of Cana- da, and election as Honorary Member of the Interna- tional Union for Quaternary Research. Vic and Pat enjoyed more than sixty years of mutual accomplishment. He loved the out-of-doors and experi- enced great pleasure whenever he had a chance to stand on a good piece of Precambrian bedrock. Vic was also very generous in his support of charitable organiza- tions, and agencies promoting wildlife conservation and the natural sciences. He will be greatly missed by his family, and by his many friends and scientific col- leagues. References Dyke, A. S., and V. K. Prest. 1987. The Late Wisconsin and Holocene history of the Laurentide Ice Sheet. Géogra- phie physique et Quaternaire 41: 247-264. Muller, E. H., and V. K. Prest. 1985. Glacial lakes in the Ontario basin. Edited by P. F. Karrow, and P. E. Calkin in Quaternary Evolution of the Great Lakes. Geological Association of Canada, Special Paper 30: 213-229. Prest, V. K. 1939. Geology of the Keezhik-Miminiska Lake area. Ontario Department of Mines, Annual Report 48, Part 6, 8 pages. Prest, V. K. 1941. The Precambrian of the Miminiska-Port Hope area. Unpublished Ph.D. thesis, University of Toronto, 93 pages. Prest, V. K. 1949. The Pleistocene geology of the Vermillion River system near Capreol, District of Sudbury, Ontario (with special reference to placer gold), Ontario Department of Mines Open File. Prest, V. K. 1952. Notes on the geology of parts of Ellesmere and Devon islands. Geological Survey of Canada, Paper 52-32, 15 pages. 2004 | Prest, V. K. 1963. The Red Lake-Lansdowne House area, northwestern Ontario — Surficial geology. Geological Sur- vey of Canada, Paper 63-6, 23 pages. Prest, V. K. 1966. Glacial studies, northeastern Ontario and northwestern Quebec. In Report of Activities May to Octo- ber 1965. Geological Survey of Canada, Paper 66-1: 202- 203. Prest, V. K. 1969. Retreat of Wisconsin and Recent ice in North America. Geological Survey of Canada, Map 1257A, scale 1:5 000 000. Prest, V. K. 1970. Quaternary geology of Canada. Edited by R. J. W. Douglas in Geology and Economic Minerals of Canada. Economic Geology Series (1): 676-764. Prest, V. K. 1973. Surficial deposits of Prince Edward Island. Geological Survey of Canada, Map 1366A, scale 1:253 444. Prest, V. K. 1977. General stratigraphic framework of the Qua- ternary in eastern Canada. Géographie physique et Qua- ternaire 31: 7-14. | Prest, V. K. 1980. Quaternary geology of the Red Lake area, District of Kenora. Ontario Geological Survey, Preliminary Map 2484, scale 1:50 000. Prest, V. K. 1983. Canada’s heritage of glacial features. Geo- logical Survey of Canada, Miscellaneous Report 28: 119 pages. Prest, V. K. 1984. Late Wisconsinan Glacier Complex. Edited by R. J. Fulton, in Quaternary Stratigraphy of Canada — A Canadian contribution to IGCP Project 24. Geological Survey of Canada, Paper 84-10: 21-38. Prest, V. K. 1984. Late Wisconsinan Glacier Complex. Geo- | logical Survey of Canada, Map 1584A, 1:7 500 000. Prest, V. K. 1985. Glacial geology of the Bebensee Lake map- area (NTS 86M), Great Bear Lake region, Northwest Ter- ritories. Edited by J. A. Brophy in Contributions to the Geology of the Northwest Territories, 63-70. Prest, V. K. 1990. Laurentide ice-flow patterns: a historical review, and implications of the dispersal of Belcher Islands erratics. Géographie physique et Quaternaire 44: 113-136. Prest, V. K., and J. A. Donaldson. 1963. Bedrock-drift dis- tribution, Red Lake-Lansdowne House area, Ontario. Geo- logical Survey of Canada, Map 4-1963. Prest, V. K., J. A. Donaldson, and H. D. Moores. 2000. The Omar story: The role of Omars in assessing glacial his- DYKE, HODGSON, BOUSFIELD, and BEDFORD: VICTOR KENT PRES1 tory of west-central North America. Géographie physique et Quaternaire 54: 257-270. Prest, V. K., and D. R. Grant. 1969. Retreat of the last ice sheet from the Maritime Provinces — Gulf of St. Lawrence region. Geological Survey of Canada, Paper 69-33, 15 pages. Prest, V. K., D. R. Grant, H. W. Borns, I. A. Brookes, R. H. MacNeill, J. G. Ogden, J. F. Jones, T. W. Hennigar, and M. L. Parsons. 1972. Quaternary geology, geo-morphology and hydrogeology of the Atlantic Provinces. XXIV Inter- national Geological Congress, Excursion Guidebook A61- C61, 79 pages. Prest, V. K., D. R. Grant, and V. Rampton. 1968. Glacial Map of Canada. Geological Survey of Canada, Map 1253A, scale 1:5 000 000. Prest, V. K., and J. Hode-Keyser. 1977. Geology and engi- neering characteristics of surficial deposits, Montreal Island and vicinity, Quebec. Geological Survey of Canada, Paper 75-29, 29 pages. Prest, V. K., J. Terasmae, J. V. Matthews, and S. Lichti- Federovich. 1976. Late-Quaternary history of Magdalen Islands, Quebec. Maritime Sediments, 12(2): 39-59. Wilson, J. T., G. Falconer, W. H. Mathews, and V. K. Prest. 1959. Glacial Map of Canada. Geological Associ- ation of Canada, scale 1:3 801 600. Examples of research publications in periph- eral fields of science for which the work of V. K. Prest was basic: Bousfield, E. L., and M. L. H. Thomas 1975. Postglacial changes in distribution of littoral marine invertebrates in the Canadian Atlantic Region. Proceedings of the Nova Sco- tia Institute of Science 27, Supplement 3: 37-42, 8 figures. Dadswell, M. J. 1974. Distribution, ecology, and postglacial dispersal of certain crustaceans and fishes in eastern North America. National Museum of Canada Publications in Zoology 11: 110 pages, 20 maps. Pielou, E. C. 1991. After the Ice age. University Chicago Press. Chicago. 366 pages. Acknowledgments For providing biographical material, photographs, and per- sonal recollections, the authors are especially grateful to Patricia Prest (wife) and Sherry Armstrong (daughter). Book Reviews ZOOLOGY Birds of Australia (Seventh edition) By Ken Simpson and Nicolas Day. 2004. Princeton Univer- sity Press, 41 William Street, Princeton, New Jersey, USA, 08540-5237. iv + 92 pages, U.S.$39.50. When Simpson and Day’s first edition appeared (as Birds of Australia, later changed to The Princeton Field Guide to the Birds of Australia) it meant Australia had a bird guide that was equivalent to North America’s National Geographic guide (Dunn 2002). It provided complete coverage of Australian birds in color, with accompanying text and range maps. Curiously the print- ing was, and still is, poorer than that of the National Geographic. As with the original this book covers all six states (including Tasmania), three territories and 10 island territories. The new edition has a double page map of the continent, but does not show the island territories. This revised edition has undergone significant changes from the original. Although much of the con- tent is the same it has been reorganized. More impor- tant 19 of the plates have been repainted (actually I counted 24 plates with changes). The illustrations now include more plumage variations than before, such as female, juvenile, winter and different races. In addition to over 2000 colour images there are almost 1000 sup- plementary black-and-white drawings. One noticeable difference is that the illustration of the dead White- throated Needletail has been replaced with a represen- The Bird Almanac By David M. Bird. 2004 Key Porter Books Ltd. 70 The Espla- nade, Toronto, Ontario, MSE 1R2 Canada. 460 pages, Can $24.95 Paper. Within this book is a massive amount of bird-related information! This review could actually end right here. However, I will elaborate somewhat. Information in this book is presented in three ways: line drawings (restricted to the anatomy chapter), tables/ lists as well as glossary-style entries. These are all ap- propriate and lead the reader to finding information rather quickly. There is both birding information and ornithological information (and a massive amount of overlap which is shared by both). A glossary of over 20 pages contains over 1000 terms — surely the word you're looking for must be there! There are simply too many categories of information to give more than a smattering of examples here. Both traditional and genetically-based classifications of bird families are given, followed by the massive list of all tation of a live, flying one. Some extra space was gen- erated on the main plates by moving the vagrants to a separate section. This is a better method of dealing with these rare birds. The introductory material has been reorganized to give a more logical flow. Two small sections — on DNA and prehistoric birds — have been replaced by more pertinent material. All the range maps have been up- dated. The result is the book is slightly smaller than the original, so the book remains portable in the field. Within the field information I noted only a few points I would question. The authors tend to lump species that more recently have been separated. These include the Royal and Macaroni Penguins, the Lesser and Greater Snow Petrels and the Yellow and Crimson Rosellas. The Paradise Parrot is still included, despite not having been seen since 1927. This revised edition, with the original, simpler title of Birds of Australia and the expansion of information, makes this field guide a more useful book than the original. Reference 'Dunn, Jon L. 2002. National Geographic Field Guide to the Birds of North America: 4'" edition Revised and Updated. National Geo- graphic Society, Washington, D.C. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada known bird species. Significant people are listed in sev- eral tables, ornithological award recipients, world-class listers, Taverner Cup winners, bird artists and more. Bird watching clubs, ornithological societies, magazines and journals are listed from sources around the planet. To compare this book with Leahy’s The Birdwatch- er’s Companion, a recent tome of similar intent, would be to have The Bird Almanac come on top. The former is essentially all in dictionary format, and therefore lacks the comparative ease (or the ease of comparisons) of the thematically-organized, tabular format in Alma- nac. There are more in-depth definitions and descrip- tions in Companion (it is also a much bigger book), and the bibliography is better- organized, but I believe birders will much more enjoy flipping through Almanac. RANDY LAUFF Department of Biology, St. Francis Xavier University, Anti- gonish, Nova Scotia B2G 2W5 Canada 630 2004 Book REVIEWS 63] Handbook of Birds of the World: Volume 9 Cotingas to Pipits and Wagtails Edited by Josep del Hoyo, Andrew Elhott and David Christie. 2004. Lynx Edicions, Barcelona, Spain. 850 pages, illus- trated. Cloth $195 US. This volume of the Handbook of Birds of the World covers 819 species, including Cotingas, Manakins, Tyrant-flycatchers (over 50 % of the book), New Zea- land Wrens, Scrub-birds, Lyrebirds, Larks, Swallows and Martins and Pipits and Wagtails. With the publi- cation of this volume Handbook has now covered 60% of the world’s 197 families of birds. Under the current schedule Volume 10 will be published in 2005 and the final volume, 16, is due in 2011. It has become very difficult to review these books. Most reviewers, including myself, have already praised the content, the artwork, the scholarship, the format and so on. The quality of all aspects has been remarkably consistent. This volume is no different and it is equiv- alent to the other volumes. Volume 9 opens with an essay on ornithological nomenclature by Richard Banks. This is a description of the history, the rules of nomenclature and some of the pitfalls. The author has tabulated some of the most frequently used species names (cinereus and its vari- ants is the most used). He also notes that after Linnaeus, the originator of the nomenclature system, Philip Lutley Sclater, a British bio-geographer and taxonomist, has officially named the most birds. This is a timely essay for Volume 9 because it pre- cedes an unusual situation. Research has shown that a proposed sub-division of the Tyrannidae had not been formally recognized. Therefore the editors published a description of the proposed tribe as required by inter- national protocol. While HBOW is a little different than the usual scientific journal, there is no reason why it should not publish this text. The breadth of coverage starts with the wildly col- oured and sometimes oddball Cotingas, not just the dazzling Cock-of-the-Rock but the bellbirds and um- brellabirds too. The manakins are smaller but also bril- liant. [remember stopping dead at the sight of a Red- capped Manakin and being so transfixed I almost missed a much rarer flycatcher; the Sulphur-rumped. But in this edition the flashy birds are overshadowed by the “little brown jobs - LBJs.” Just under half of the 429 Tyrannids are olive-grey-brown. Most of the larks and pipits are LBJs too. In fact, about 75% of the species in this volume fall into this category. This includes the dozen or so notorious Empidomax flycatchers and the two dozen tropical Elaenias. To separate these birds you need a good view plus either their song or geographical location to stand a chance at identification. Earlier this year, a colleague and I spent 15 minutes studying a close and very still (it was drizzling) Mountain Elaenia before we were satisfied with its identity. Even when the Tyranid is brightly coloured with a yellow underside and a black-and-white striped head, then you have 14 look-alikes to contend with. Again geography helps but separating Social and Lesser Kiskadee flycatchers can be tricky. One of the qualities I have learnt to appreciate more and more about HBOW is the perspective I get by hav- ing the entire world taxonomy and distribution at my fingertips. For example, when I encountered a new species, one of the eight African longlclaws (in Mac- ronyx) I was able to read about their relationship with other pipits. This despite my tendency to regard them as little meadowlarks (of the purely America Sturnella), from their appearance and habits. HBOW brings a focus to the biogeography and family ties to these widespread families. This access to worldwide taxonomy also raises ques- tions. How can the Temminck’s (Horned) Lark of north- ern North Africa be a different species from the Horned Lark of Europe and North America? The variation in plumage of the latter easily overlaps the paleness of the former (especially Eromophila alpestris bicornis, E.p. brandti and E.p. teleschowi). Indeed, some years ago, I saw a Horned Lark in Manitoba that was remark- ably pale and very close to Temmink’s Lark. I have never doubted it was nothing more than a pale version of our common Horned Lark. So I wonder why is Tem- minck’s separate from the Horned Lark, which in turn is split into 42 subspecies? I look forward to Volume 10 to see how the authors handle the stonechats, in particular the very distinctive Madagascan Stonechat. The editors are continuing to maintain their uni- form, high standard and have settled in to a one- per-year production. In this volume they have made another useful change. Previously the plates identi- fied the species by a number only. You needed to turn to the text to match the number with a name. The names of all the species depicted on a plate are now given along with the species number in a footnote opposite the plate. Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada British Columbia: A Natural History. Revised and Updated Richard Cannings, and Sydney Cannings. 2004. Greystone Books, Douglas and McIntyre, Vancouver, British Colum- bia, Canada. 341 pages, 209 figures. $39.95. This paperback reprint is a gem. As the title indicates, it is a revised and updated version of a hardcover book first published in 1996, which won the Bill Duthie Bookseller’s Choice Award for the “best work pub- lished in British Columbia” that year, “The Lieutenant- Governor’s Award for Historical Writing in British Columbia,” and the Science in Society Award of the 632 Canadian Science Writers’ Association for the best gen- eral work. It is an informative, attractive, extremely well-organized book. The Cannings brothers (identical twins), their broth- er, Robert, and their late father, Steve, have each con- tributed masterful photographs. All four are well-known naturalists who have done much to encourage and fur- ther the study of natural history through print, radio, and television in western Canada. The Cannings’ book succeeds at a number of levels. Visually it is a delight, with superb, eye-catching maps and photographs, enhanced by informative captions. On initial browse, one is stimulated by the pithy, in- formative, boxed vignettes, such as “when is a seagull not a seagull?” and “Western Sandpipers.” These vig- THE CANADIAN FIELD-NATURALIST Vol. 118 nettes will please even the most jaded of naturalists. Finally, the main text provides a wealth of well-organ- ized and thoughtfully developed information. Topics such as geology, the ice age, forests, mountains, grass- lands, and water are clearly and incisively presented, as the Cannings’ knowledge and enthusiasm shine through. British Columbia is most fortunate to have such mas- terful and able presenters; nature lovers everywhere will benefit from this book. Only superlatives apply. I cannot think of a single word of criticism. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada Field Guide to Bird Nests and Eggs of Alaska’s Coastal Tundra By T. D. Bowman, 2004. U.S. Fish and Wildlife Service Anchorage, Alaska. Published by Alaska Sea Grant Col- lege Program. 81 pages, U.S. $25.00 Paper This little publication is an interesting add-on for the field practitioner in Alaska and the arctic. This region includes some of the most productive bird nesting areas in North America, and probably in the world. It shows in detail nest and egg features of 70 bird species (loons, waterfowl and crane, jaegers, gulls and terns, owls, ptarmigan, alcids, shorebirds and passerines). Tavern- er’s Canada Goose and Cackling Canada Goose receive separate chapters. The species included in this guide were selected based on their occurrence in Alaska and by their “relative abundance”. Some rare species like Sanderling, Surfbird and Yellow-rumped Warbler are unfortunately not included, which will not really help to bring us any closer to their nest discoveries. This is a convenient book designed to be used in the field. Bird species in this book are ordered by egg size, which makes an interesting presentation. The book design and the photo arrangements are done nicely, but sometimes I find the egg photos a little overdone and repetitive; sometimes wing photos are hard to differ- entiate. For each species, the photos show the bird (often male and female), bird on the nest, the actual nest, eggs and (down) feathers; but this format is not always strictly followed. Personally, I find it somewhat a shortcoming that chicks are rarely presented in this Guide to Hawk Watching in North America By Donald S. Heintzelman. 2004. Globe Pequot Press, Guild- ford, Connecticut. 425 pages, U.S. $16.95 Paper. The Guide to Hawk Watching in North America, written by ornithologist Donald Heintzelman, is an up- dated version of his A Guide to Hawk Watching in North America, published in 1979. The new edition includes updated information about “hawk” (including vultures, eagles, harriers and falcons) biology, identi- fication and seasonal migration movements, as well guide. Of interest is the concept of a “Sizing Chart’, which “provides a quick reference to help reduce the number of prospective species to which an egg could belong”. Further, the book offers for geese “Parting Shots...” in order to identify escaping birds. A quick reference guide is provided for “Dark Goose Nests.” Six short introduction text pages provide basic back- ground about the species and the book concept. I ad- mire the challenges and amount of field work involved in compiling the specific nest and egg colour photos (over 450). A great photo collection is presented which includes not only, those done by the author but also contributions from over 70 other contributors. As I tend to be creative while in the field, I miss some free pages to write on during field work. The competent author has over 15 years of experi- ence working in the arctic, and is inspired by the “...dedication to improving the reliability and accu- racy of scientific data...”. Therefore, it can be hoped that these photos eventually might occur on the internet /www for a free use by the general and interested public. Overall, I think this interesting book serves it pur- pose well as a quick field reference and fills a vacant niche. I am sure it will also prove useful for Arctic regions of Canada as well as Russia. FALK HUETTMANN Institute of Arctic Biology, Institute of Wildlife Biology, University of Alaska-Fairbanks, Fairbanks Alaska 99775 USA as numerous additional sites to watch raptors. The new edition adds a separate section on where to view Bald Eagles. The book is divided into two main parts. The first concentrates on general information, including species accounts for a wide variety of raptors other than owls, tips on how to identify and study hawks, a description of the migration seasons, an overview of different types of hawk watching, an introduction to field equipment for the activity, and an explanation of the mechanics of 2004 hawk flight. The second part of the book focuses on hawk watching sites, including U.S. hawk migration watch sites, Canadian hawk migration watch sites, Bald Eagle viewing areas, and places to view raptors out- side the migration season. The species accounts provide excellent identification information on new world vultures, ospreys, kites, hawks, eagles, harriers, caracaras and falcons. Each species 1s described in terms of wingspread, total length, field recognition features, flight style, voice, nest, eggs, maximum recorded longevity, food, habitat, and North American range. It makes for an excellent species sum- mary, even if it lacks references to the bird’s conser- vation status. I found the chapter on mechanics of hawk flights particularly interesting. Although my general under- standing of hawk migration was fairly good when I started reading this book, I had never looked into it in great detail. So I was fascinated to read about the influ- ence of general weather conditions, and the use migrat- ing raptors make of deflective updrafts, lee waves, ther- mals, thermal streets, squall lines, and leading lines. I was also interested to read about hawk migration watch sites throughout Canada, including Alberta’s Canmore Collegiate High School, Nova Scotia’s Brier Island near Digby, Ontario’s Holiday Beach Migration observatory near Windsor, and Quebec’s Morgan Book REVIEWS 633 Arboretum at the west end of Montreal — a sampling of Canadian migration watch sites with high ratings. Canadian entries in the Bald Eagle watch sites section include British Columbia’s Active Pass in the Gulf Is- lands, Pacific Rim National Park on Vancouver Island, the Fraser Valley Bald Eagle Festival near Mission, the Squamish Valley, the Greater Vancouver Area, and Prince Rupert. The section on other raptor viewing areas also offers sites in Canada, including Amherst Island and Wolfe Island in Eastern Ontario, both featuring Rough-Legged Hawks and various owls during the winter season. The other Canadian sites for concentrations of raptors out- side the migration season are the Yukon’s North Klon- dike Highway and South Klondike Highway. Both areas offer year-round opportunities to see a wide vari- ety of eagles, hawks, falcons and owls. The Guide to Hawk Watching in North America is an extensive and practical information source for any- one interested in raptor watching on this continent. My only objection about the book is its title, which I feel should be changed to Guide to Hawk Watching in the United States and Canada, since it lacks any ref- erence to sites outside those two countries. R. SANDER-REGIER RR5 Shawville, Quebec JOX 2YO Canada Locust: The Devastating Rise and Mysterious Disappearance of the Insect that Shaped the American Frontier Jeffrey A. Lockwood. Basic Books, New York. Hardcover. 294 pages, Can. $39.00 Paper. Locust is much more than an account of a single species of insect. In fact, it is everything the dust cover promises: “A fascinating detective story” that delves into “history, culture, religion, and especially ecology, interwoven by the life story of a common insect. ...with vivid prose, epic thoroughness and scientific precision.” I would add geography and science to that list. Lock- wood writes engagingly. He shares his years of detec- tive work, providing details that only a professional entomologist could. His historical delvings put most historians to shame, and his writing skills exceed those of almost any living science writer. The book opens in Dodge County, Nebraska, in July 1875, a drought year, with clouds of locusts obscuring the sky, their wings crackling like a horrific blaze. Limbs of willows bent to the ground under the weight of the insects, as adjacent cornstalks were stripped bare. When a sheet of insects six inches thick passed over a perpendicular ledge of rock, they caused a roar- ing noise similar to a cataract of water. In Utah, locust eggs were counted at 743 million eggs per acre. The voracious insects literally ate the clothing off human limbs, then entered homes to eat objects such as win- dow blinds. When domestic chickens gorged on the locusts, their eggs and flesh became inedible. Farmers and their families lost their gardens and crops, and were on the brink of starvation. The U.S. Army under General Ord saved many farm family lives by far exceeding normal army routine. Ord issued thousands of infantry coats, shoes and mil- itary blankets, as well as large amounts of army rations. Lawmakers apportioned money to distribute wheat seed for planting the following spring. Without Ord, thou- sands would have died. North America was blessed with hundreds of species of grasshoppers, but only a single species of locust, the Rocky Mountain locust, Melanoplus spretus, named by Benjamin Dann Walsh back in 1866, caused such vast destruction. Ingenious but rather ineffective ma- chines such as suction machines and flame throwers were invented to combat the locust; two men and a team of horses could incinerate ten acres of locust-infested fields in a day. One Minnesota community alone had a thousand coal-tar hopperdozers that could harvest 150 000 locusts per hour. On the scientific front, three entomologists did their best to help combat the locust. Charles Valentine Riley was the state entomologist for Missouri. Cyrus Thomas and Alphaeus Spring Packard, Jr., held similar posts in Illinois and Massachusetts. Riley was able to show that the locust had the potential to increase its popu- lation 100-fold from one generation to the next; he argued that for every bushel of locust eggs destroyed, 100 acres of crop could be saved. He founded the Na- 634 tional Insect Collection and through his efforts a branch unit in economic ornithology arose. This office became the Bureau of Biological Survey and later metamor- phosed into today’s U.S. Fish and Wildlife Service. Locust has important Canadian content. Entomolo- gist Norman Criddle of Manitoba devised a widely- used mixture of copper acetoarsenite, molasses and horse manure to combat later outbreaks of somewhat less harmful grasshoppers. Criddle also collected the world’s last two specimens of Melanoplus spretus on 19 July 1902. Three Saskatchewan men, Paul Riegert, Bill Chapco, and Bob Randell, also helped solve prob- lems discussed in later chapters. Why did the Rocky Mountain grasshopper become extinct? Five different theories in turn held sway: the spread of alfalfa; the demise of the Bison; changes in weather; overgrazing of grasslands; widespread prairie fires — but each was discredited, though two were later revisited as contributing factors.. To allow DNA studies and do radiocarbon dating, Lockwood determined to find locust specimens en- tombed in glacial ice for a century or more. His first try, in 1987, yielded a quarter pound of dried grass- hopper parts; when results were submitted to a leading entomological journal, the editor rejected the paper and informed Lockwood that “you have mistaken natural history for science.” In 1988, Lockwood and colleagues collected 134 specimens of twenty species of grasshopper, but no locust, on the glacier. In 1989, they obtained 4 mg of grasshopper parts, including mandibles that appeared to match those of extant pre- served specimens of Melanoplus spretus. Finally, in Mammals of the World: A Checklist By A. Duff and A. Lawson. 2004. Yale University Press, P.O. Box 209040 New Haven, Connecticut 06520-9040 USA. 312 pages, U.S.$45.00. Perhaps I should have bought this book some time ago, or at least its predecessor; Mammal Species of the World (Edited by D. E. Wilson, and D. M. Reeder, 1993, Smithsonian Institution Press), which contains the names of the recognized species of mammals doc- umented at that date. So many times I have struggled to understand which species are present in a specific area. With birds this is fairly easy to resolve. English names are reasonably standard and scientific names are the ultimate guide. With birds I have rarely had to probe recent literature to catch up on the latest splits. With mammals this is much more difficult. It took some time to clarify the species and distribution of fur seals. The confusion of English names for wilde- beest left me very perplexed. My latest book on African mammals was particularly mystifying. It is a transla- tion from German to English and uses different Eng- lish names for the plates and the text. The scientific names are not current. Starting with this book it took a lot of work to create a list of lemurs that showed the THE CANADIAN FIELD-NATURALIST Vol. 118 1990, at the melting edge of glaciers, they collected 250 locust bodies, including 14 males with well-preserved abdomens and genitalia, allowing unequivocal identi- fication and DNA analysis. These insect bodies had taken about 150 years to travel 300 m as the ice moved from the crevasses where they had been entombed. Why, indeed, had Melanoplus spretus become extinct? For once, the unplanned effects of human activ- ity had an effect for the better. Lockwood explains that the “base locality” of the locust was, between out- breaks, restricted to a few relatively small areas in val- leys within the Rocky mountains, where eggs could be deposited in sand and gravel. Following European settlement, floods occurred more often due to overgraz- ing on the slopes, irrigation periodically flooded the valleys, and the locust eggs lost their vitality. Plough- ing and harrowing destroyed the eggs. The new alfal- fa crops were inimical to development of the locust nymphs. The enigma has been solved. Lockwood is a consummate writer. His eminently readable book is a detective story, keeping the reader in suspense to the final chapter. Admittedly this review has given away the ending and thus has spoiled some of that suspense, but the pleasure of this book lies more in its details and insights than it does in the suspense. I strongly recommend this book to everyone with an interest in a good story well told. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada currently known species and sub-species. With Mam- mals of the World this exercise is simple. This current list contains 5069 species. From the Wilson and Reader list of 4629 the authors have sub- tracted 41 species that were extinct prior to 1800. These include such creatures as Steller’s Sea Cow, hunted to extinction by 1768. More recently extinct species are left in, presumably as, in theory, there is a chance of still finding one alive. The Thylacine comes to mind. Duff and Lawson have added 522 “new” species. A large portion of these come from two sources. Juliette Clutton-Brock’s work on domestic animals has elevat- ed numerous creatures to species status (the Domestic Pig goes from Sus scrofa to Sus domesticus for exam- ple). The second source of change is the assignment of species status to numerous sub-species (Colin Groves of the Australian National University has been active in this exercise). Finally, 41 species have been demoted in synonymy. Sadly this loses us the Queen of Sheba’s Gazelle (now a subspecies of the Arabian Gazelle), a romantic loss at least. The changes in all the above categories are fully explained in the appendices. Also included are the nomenclatural changes in scientific names. 2004 The authors say they have followed Wilson and Coles (Common Names of Mammals of the World. 2000. Smithsonian Press) version of the English names with corrections and conversion to “well-established names.” Generally they do not list alternative names so an animal like the Cougar, or Mountain Lion, or Catamount only gets listed as Puma. When I used the index, being uncertain of the author’s choice, I looked up Lion (Panthera leo) knowing that Cougar would be nearby. The most odd name I found was Sewellel, the Chinook Indian name for the Mountain Beaver. This was the first time I had seen this name although Audubon used it on his painting of Aplodontia rufa. Naturally I compared the list of lemurs that I had created to that of Duff and Lawson. I was not surprised to find many that I had as sub-species had been elevat- ed to full species. Otherwise the lists were the same except for Grey-brown Mouse Lemur (Microcebus griseorufus). R. M. Rasoloarison, S. M. Goodman, and J. U. Ganzhorn first described this cute, hamster-like lemur in 2000. I have since been fortunate to see this little beast in South-western Madagascar. This omis- BooK REVIEWS 635 sion is surprising as the author’s references go up to 2002, but to be fair this is a family undergoing con- stant change. I was also surprised that Canada was not mentioned in the range of the Red Wolf (Canis rufus). John The- berge’s work has shown that Algonquin wolves are closer to Red Wolves than the more common Timber Wolf (Canis lupus). This information is far more wide- ly known. For those of us that like to travel and see mammals in their native habitat this book is a good and useful guide. However, it is only a list and, although it will not resolve all taxonomic questions, it will bring clarity and order to your research and therefore it is a valuable reference book. As it is based on Wilson and Reeder’s out-of-date book it is more up-to-date, but I understand a revised edition of Wilson and Reeder will be published soon — should we wait? Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada Parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey Louis A. Somma, 2003. Krieger Publishing Company, Mal- abar, Florida. x + 184 pages, U.S.$33.50 Cloth. This book is exactly what it purports to be. It is a summary and exhaustive bibliography of literature on parental behaviour in lizards, snakes, amphisbaenians, tuatara and turtles. Passing reference is also made to literature on other vertebrates including dinosaurs. This book focuses exclusively on behaviour as opposed to physiological adaptations and so, while parental behaviour in some species of snakes and lizards is well known, many readers will be astonished to find turtles included. Yet Somma cites references that provide some evidence of parental behaviour for seven species of turtles. Fifteen categories of parental behaviour are discussed including defence, thermo- and hydro-regulation, assis- tance during hatching and facilitated feeding. For those unfamiliar with research in this fascinating field the range and diversity of parental behaviours in these reptiles will intrigue and astonish you. Unfortunately, description and discussion of these behaviours is lim- ited to the first 11 pages of the book. This is followed by 46 pages of tables summarizing the results and guiding the reader to literature on various topics and taxa. The remainder of the book consists of 100 pages of refer- ences and an index to subjects and taxa. Readers should be aware that the index is inconsistent as to whether it includes references to taxa within the tables (for example, it does for Trachemys stejnegeri malonei but not for Eumeces fasciatus). The primary purpose of the book seems to be to raise the profile of parental behaviour in non-avian reptiles and encourage further research on this understudied phenomenon. It is unfortunate that having read such an enormous amount of information on this topic, Somma does not provide more insights into its evolution and ecological ramifications but perhaps the field is too young for such a synthesis. Certainly for anyone inter- ested in embarking on research into this topic it is an invaluable and relatively inexpensive reference. For those who are most keen on this topic, Somma has also published an addendum to the book (Somma, 2003). Reference Somma, L. A. 2003. Parental Behaviour in Lepidosaurs and Turtles: Source Addendum. Bulletin of the Chicago Herpetological Soci- ety 38(4): 65-76. CAROLYN SEBURN 7210 Clarenda Street, Ottawa, Ontario K2B 7S5 Canada Prairie Ghost: Pronghorn and Human Interaction in Early America By Richard E. McCabe, Bart W. O’Gara, and Henry M. Reeves. 2004. 176 pages, U.S.$29.95 Cloth. The Pronghorn, Antilocapra americana, formerly called “antelope,” is the “most American” of the con- tinent’s terrestrial wildlife, since it is found nowhere else. It is the world’s second fastest land animal, and perhaps the most inquisitive. Protuberant eyes allow it a nearly 360-degree field of vision. Large lungs, heart and trachea permit it to achieve great speed. This historical look at the Pronghorn is thoroughly researched, with informative tables, extensive refer- ences, and well-chosen, sumptuous illustrations. It 636 delves deeply into anthropology. The Pronghorn rep- resented fleetness, alertness, and pertinacity, and was a catalyst in the social structure and welfare of most plains tribes. It is no wonder that many place names today derive from this one species, but who would have expected 108 such instances in Arizona and 79 in Wy- oming? Two detailed appendices list dates of eyewit- ness accounts between 1540 and 1896, and the names of the Pronghorn in each native language. Historical accounts go back to Sahagun in 1569 and Hernandez in 1651, both in Mexico. Lewis and Clark popularized it during their exploratory journey of 1804-1806, and George Ord gave the Pronghorn its binomial Latin name in 1815. Before Europeans reached the Americas, aborigines hunted the swift-footed Pronghorn with three instru- ments: sling, atlatl, and bow-and-arrow. A well-condi- tioned, determined native could sometimes, in spite of his much slower pace, outlast and eventually tire and kill a Pronghorn. More often, stalking, pursuing, sur- rounding, luring, calling, impaling, ambushing, netting, driving with v-shaped fences, setting prairie fires, and making pitfalls, were the methods used to kill them for food. Pronghorn hunting required more prepara- tion and more co-operative effort than did hunting of the larger Bison. The Pronghorn was then one of the natives’ most important food sources, especially where Bison were scarce near the edge of their range. An antelope skin was thinner and lighter than that of a Bison, and thus more suitable for clothing. A native would obtain about 45000 calories from a 43-pound Pronghorn carcass. Pronghorn bones were used as toys, rattles, awls, pipes, fishhooks and decorations. Sinew served as strings for bows, and for sewing. A skin stretched tight- ly over a section of hollowed tree formed a drum. Marrow was rubbed on sunburned or chapped lips and THE CANADIAN FIELD-NATURALIST Vol. 118 skin. A Pronghorn fawn was used as bait on branches above a pit where an Indian patiently waited to catch a Golden Eagle for its feathers. Images of the Prong- horn were used to decorate native pottery, on the walls of caves at ten known sites, and as effigies. Hunting became much easier after guns spread north from Mexico and guns and ammunition were obtained in trade, but this soon led to squandering of the once- precious resource by a veritable army of 5000 white hunters. From 1874 through 1877, more than 100 000 Pronghorn hides were shipped from the plains annually. As the Bison almost vanished, the Pronghorn became even more important as a food source, and their num- bers dwindled as well, from about forty million to fewer than 15000 in 1910. Bereft of their two largest natural food sources, native peoples were overwhelmed, sub- jugated, displaced, and pauperized of their culture, iden- tity and social options, losing much of their spirit and vitality. Although three of the four last sections tell a grim story, dealing with the near demise, in turn, of the Bison, the people, and the Pronghorn, the book ends on a positive note with the birth of the conservation ethic. In 1887, Theodore Roosevelt convened a dinner meet- ing in Manhattan to launch the Boone and Crockett Club; he and the other founders had a strong affection for the Pronghorn, which eventually benefitted from the conservation ethic and philosophy of this fledgling movement. This attractive, scholarly, modestly-priced book belongs in major libraries. It would make a perfect gift for anyone interested in history, geography, anthro- pology, or big game hunting on the plains. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Canada A Guide to the Birds of St. Helena and Ascension Island By N. McCulloch. 2004. The Royal Society for the Protection of Birds, The Lodge, Sandy, Bedfordshire, SG19 2DL Great Britain. iv + 92 pages, 11 GBP. The booklet is intended for the visitor and has three parts. The first is an overview of island history from time immemorial to present. The second is a site guide giving where to find birds on these islands. The last part is the bird accounts. The bird accounts cover 31 species for St. Helena and 46 for Ascension that a visitor is likely to see in a day trip around each island. Of these 11 are seabirds and 10 are shorebirds on St. Helena and 13 are seabirds and 16 shorebirds on Ascension So only about 30 % of the island species are land birds, with most of them being introduced. There have been 41 other attempts at introduction on St. Helena and 9 on Ascension The account of the history is fascinating and so typical of remote islands, especially after its “discov- ery” by humans. This is not good bedtime reading though. The destruction of the islands ecosystem is the usual unpleasant tale of wanton destruction misguided mistake and foolish carelessness. There have been 41 failed attempts to introduce everything from a white-eye to an ostrich to St. Helena and 9 attempts on Ascension. This includes the introduction of House Sparrows in 1986 (to be company for a ship-assisted vagrant who arrived in 1985). Will we never learn? On a happier note there are programs underway to restore habitat for around 40 species of endemic plant and 9 species of creepy crawlies (one — the Golden Sail Spider — is illustrated) as well as birds. There are also programs to reduce cats, goats and the like. The bird finding section is typical of current field guides, portraying the bird on the left and giving infor- mation on the right. It covers 28 species. Half of these are seabirds. The remainder are land birds, almost all introduced. The English names are fairly consistent with other texts, but there are no references to alter- 2004 | native names. For example Maderan vs. Band-rumped Petrel or Parasitic Jaeger vs. Arctic Skua. The text here along with the rest of the book is good and clear. The author has added a table of accidental species for the two islands (31 on St. Helena and 46 on Ascen- sion. This did not quite match my own list, but I think the difference is due to the political boundary that in- cludes Tristan de Cunha and Gough Islands, thus add- ing several Antarctic species) The artwork consists of watercolours that are refined sketches of birds during their normal activities. This gives a better idea of their jizz than the more formal- ized field guide style of the recent books, and is more _ akin to the style of postwar books. Indeed, my favour- ite is a perky field sketch of a Java Sparrow that fronts | the section of land birds on which you can almost | count the small number of brush strokes. The reader | can compare this with the more “finished” plate in the accounts section. I found it ironic that the endemic Madagascar Fody | was introduced, not from Madagascar but from Mau- | ritius (where it is an introduced threat to the endemic | BOTANY BooK REVIEWS 637 Mauritius Fody) and became so numerous it was part of the St. Helena cage bird trade. How convoluted we make the world. So why go to these remote places? First there is the attraction of wild oceanic islands. While they are no longer the lush paradise first seen by the Portuguese they still are dramatic. The rich brown cliffs surround- ed by blue sea set off the white, guano-capped islands. Second, however diminished, there are good seabird colonies; always exciting places. And last, for the hard core, they are the only places to see Ascension Island Frigate and Wirebird. While remote, these islands are not inaccessible. There are some tours that include them on a cruise, although these are very expensive. Routine commercial sailings leave from Cardiff, Wales and Cape Town, South Africa a few times a year. They dock in St. Helena and Ascension for a day or two — enough time with this guide to see most of the islands birds Roy JOHN 2193 Emard Crescent, Beacon Hill North, Ottawa, Ontario K1J 6K5 Canada | Flora and Climatic Conditions of the North Pacific: A Collection of Scientific Papers | Edited by A. N. Berkutenko, H. G. Lumsden, and D. Lumsden. 2001. Institute of Biological Problems of the North, North- East Scientific Center, Far East Branch, Russian Academy of Sciences. Magadan. 189 pages, No price available. The subtitle — A collection of papers — better indi- | cates the contents of this book than does its lead title. | What we have here is a nicely produced small book | consisting of papers on flora and vegetation, for the | most part, but also one paper each on fungi and eth- nobotany, two on seed biology, and one on the influ- | ence of air masses from the Sea of Okhotsk on sum- mer temperatures in Japan. An odd mix, but perhaps | the inevitable result of the need to collect sufficient | papers, achieve critical mass, and gain publication. | To one interested in floristics and taxonomy, several _of these papers are informative and useful. Since the | book is entirely in English, it opens to a wider audi- ence than usual the results of botanical studies in the Russian Far East. The prime mover for this collection was A. N. Berkutenko who wrote entirely or con- _ tributed to six of the collected 15 papers. Since it is difficult to generalize the disparate contributions, I will give a précis of each. Yakuboy et al. provide a brief sketch of the physical _ setting, a history of botanizing, and an annotated check- | list of 235 species for the flora of Avachinsky volcano. | The next paper by Mochalova describes the very small flora of very small islands of the Commander archipelago and the effects of bird colonies on the veg- etation there. Two papers by Khoreva and by Berku- tenko et al. discuss the vegetation and flora of two islands in the Sea of Okhotsk and provide checklists for the Yams Islands and Nedorasumenia Island, the latter with the unexpected occurrence of the Asiatic shrub Caragana jubata. Sinelnikova gives a synopsis of the plant cover and a checklist for 454 species of vascular plants found at the Orotuk field station in the upper Kolyma River region. The station lies in larch taiga near the Koly- ma floodplain. The Chosenia arbutifolia and Populus suaveolens of the gallery forest and the Pinus pumila communities farther upslope are among the memorable botanical images I took away from my trip to the region. A short paper by Berkutenko and Khoreva, in a structure now familiar, provides a sketch of the veg- etation and a checklist to the 98 species found at the Mount Kamenny Venets nature monument, which in- cludes the endemic willow Salix magadanensis. Thus ends the first 117 pages. From here on, the papers have less to do with each other or to what has gone before. A list of 161 macromycetes in the Magadan Preserve by Sazanova is followed by a report by Berkutenko and Yukawa of the first record for the orchid Liparis kumokiri for mainland Russian Far East.Next Misako proposes that morphological variation in the Japanese Sanguisorba tenuifolia has originated from hybrids between S. parviflora and S. officinalis occurring in coastal Russian Far East. In a brief, idiosyncratic but interesting essay, Berkutenko contrasts with ethnobotan- ical anecdotes the changes that have taken place in people’s live since Krasheninnikov reported on 18" century habits of the local people on Kamchatka. She includes some differences between practices in the Russian Far East and Alaska as well. This paper is fol- 638 lowed by comments on the germination of 78 species by Andrianova and Berkutenko: which taxa require pretreatments and which kinds of treatments are effi- cacious. A paper by Kryukov reports briefly on the relative germination success of 27 species following different periods of storage. Haese describes a phytoso- ciological analysis of coastal tundra using the methods of Braun-Blanquet, and provides tables typical of this analysis. Hanno and Oka in a short paper that reads more like a proposal noted that the Sea of Okhotsk affects the climate of eastern Japan. Pachomov and ENVIRONMENT The Russian Far East By Josh Newell. Second Edition, 2004. Daniel and Daniel, Publishers, McKinleyville, California, USA. 486 pages, U.S.$59.95. Paper. This is a massive and thorough compendium. The subtitle reads, “A reference guide for conservation and development.” The format is a series of chapters each concerning one of the major administrative divisions of the huge region, abbreviated as “RFE.” These divi- sions are: Primorsky Krai, Khabarovsk Krai, Jewish Autonomous Oblast, Amur Oblast, Republic of Sakha, Magadan Oblast, Chukotsky Autonomous Okrug (Chut- kotka), Koryak Autonomous Okrug (Koryakia), Kam- chatka Oblast, Sakhalin Oblast. There are over 50 maps, and a host of tables, figures and photographs, as well as an index. Vegetation is organized into the classic Tundra and Taiga formations. This book divides Tundra into two parts: “Arctic Tundra” and “Tundra” (referred to in much Russian ecological literature as “High Arctic” and “Low Arctic.”) Taiga is defined correctly as “the large mass of the boreal forest that forms the heart of the RFE.” The southern taiga, which in much of Russ- ian ecological literature is known as Ussuri Taiga, is also known as “Dark Taiga” because of the high per- centage of spruce and pine. This division serves to differentiate it from the northern part of the Taiga which is widely known in the Reindeer literature as “Light Taiga’” because of the very large percentage of the forest cover of deciduous larch. The Reindeer literature, and much ecological literature, also designates the southern-most tundra, combined with the northern-most taiga as ‘‘Forest- Tundra.” This recognition of it as a separate entity is undoubtedly because the Forest-Tundra is particularly important as winter pasture for the semi-domesticated Reindeer. The Ussuri taiga is relatively familiar to English- speaking biologists because of translations of works by such authors as Sdobnikov and Arseniev, as well as Kurosawa’s famous 1975 film about Arseniev and Dersu. The number and distinctiveness of the species of plants and animals of the RFE is legendary, espe- cially the Ussuri taiga. THE CANADIAN FIELD-NATURALIST Vol. 118 Sinelnikova present six years of observations on the effect of artificial warming on the growth of four tun- dra plants at or near the field station described by Sinelnikova (see above). I fear these studies in support of the International Tundra Experiment (ITEX) proj- ect will be lost to the [TEX community in these pages; it belongs with others of its ilk. DAvID F. MURRAY University of Alaska Museum of the North, Fairbanks, Alaska Each of these 10 administrative regions receives a section of the book. Each begins with an overview, ranging from a paragraph or so to a number of pages, followed by extensive entries on: location, size, climate, geography and ecology (including carbon stocks), major ecosystems, protected areas and their problems, bio- diversity hotspots, political status, natural resources, main industries, infrastructure, foreign trade, economic importance in the Russian Federation, general outlook. This latter section is particularly interesting for such items as a full-page table of major environmental issues and problem areas of each of the regions: fishing, energy, timber, mining, agriculture. There are pertinent discussions of the weaknesses of various Russian gov- ernmental regulations concerning oil, gas and mining operations compared to the (already-weak) United States and Canadian regulations. There is discussion of various schemes with poten- tially-severe effects as well as prospects for a sustain- able economy, ENGOs, the UN Global Environment Facility, foreign government aid agencies, other prom- ising sectors such as ecotourism and NTFP (Non Tim- ber Forest Products). There is rather detailed consid- eration of the degradation of zapovedniks (“protected areas”) by tourism, poaching (salmon, caviar), bear killing (for paws and gall bladders). The forests of Kamchatka are particularly vulnerable and critical for mitigating floods and protection of salmon spawning grounds. (Remember British Colum- bia?). In the RFE the largest emitter of atmospheric CO, is fossil fuels combustion, but second place is de- forestation leading to loss of carbon-rich boreal forest and replacement by pioneer types of forests and shrub- by vegetation. More than 90 percent of the logging in the RFE is by clearcut. Even plantation-forests do not recover pre-logging stocks of CO,; managed planta- tion-forests usually contain only 4 to 4 the carbon that undisturbed forests do. Activities to “Save Tropical Forests” are having adverse effects on the taiga of the RFE... “Plywood manufacturers are promoting Russian larch (tamarack) as a green alternative to tropical luan timber and have been steadily increasing levels over the past decade... |More than 98 percent of all Russian larch grows on /some form of permafrost (continuous, discontinuous, | sporadic) making large-scale logging of the species an | unwise proposition” (page 31). “The sheer size of the Siberian and RFE forests and | the diversity of their plant and animal life and habitats | make these forests a tremendously important factor in |}Russia and the world. Loss of habitat, mostly from | forest exploitation, fire, disease and inappropriate man- | agement is the most serious threat to the unique bio- diversity of climax forests. The survival of endangered | species, such as the Siberian Tiger ... depends on the /maintenance of large, undisturbed forest areas. Frag- ‘ile permafrost areas must be recognized as environ- ‘mentally critical and the forests’ large contributions to carbon sequestration must not be jeopardized. Such environmental considerations are currently inadequate- ly incorporated in the planning process or not properly ) addressed in forest management and harvesting activ- |ities. As a general requirement for attending to these environmental concerns large forested areas must re- ‘main undisturbed and forests outside protected areas /need appropriate and complementary management.” (page 32) The Russian Far East has the endangered Siberian Tiger (Panthera tigris altaica) while Canada has the }endangered Woodland Caribou (Rangifer tarandus caribou) (Pruitt and Baskin 2004; Schaefer 2003). Pages 35 — 39 also bring problems in the RFE directly home to Canada: “A major problem is the lack and loss of | field scientists and field workers because of dramatic cuts in government funding...” We can compare the situation in the RFE with the massive cuts in scientif- ic staff of Canadian Wildlife Service, National Parks | Canada, Canadian Meteorologial Service and the Cana- | dian Museum of Nature, beginning with the Mulroney Conservatives and not restored by later governments. In Canada, the problem is compounded by the growing infatuation of university biology, botany and zoology | departments with “computer models” or “keyboard ecology” instead of learning about real animals and | plants (Ehrenfeld 1993; Futuyma 1998; Noss 1998). __ The book is almost overwhelming in its information: history of European invasions, lists (with comments) of major species of plants and animals, lists of endan- gered species (usually with comments regarding threats to survival), lists of protected areas by type, size, date of establishment, biodiversity hotspots, human econ- -omy and environmental impact. The sections on Indigenous Peoples and their decline _in numbers, relations and their problems with “modern” BooK REVIEWS 639 extractive economy (mining, logging, etc.) is particu- larly instructive because of the great resemblance to the history of similar Aboriginal groups and their ex- ploitation or rejection by Canadian governments and industries. We read much in the western press about the wide- spread pollution and environmental destruction dur- ing the Soviet era, but “While the focus has been on nuclear, air, and water pollution and, somewhat less, on the wanton waste of resources caused by inefficient production, scholars have largely ignored what may be the most significant environmental legacy from the Soviet era, and what is Russia’s greatest legacy to the planet: wilderness.” (page 29). Although the Russian Far East is well over the shoul- der of the world from Canada, the unity of the taiga (so-called “boreal forest’) is evident by the similarity of its animals and plants to those in Canada. We can also see in this compendium disturbing similarities to the problems affecting the Canadian taiga (Pruitt and Baskin 2004; Schaefer 2003). Comparison of amelio- rating efforts will be valuable. There are many places in Canada which should have copies of this valuable compendium: Federal and Provincial departments of Conservation, Natural Resources, Forestry, Wildlife, Aquatic Resources, Fisheries. All University libraries, the libraries of all ENGOs concerned with conservation and sustainable use of renewable resources, anthropol- ogy and aboriginal activists, everyone teaching Field Biology, Ecology, Boreal Ecology, Resource Manage- ment, Forestry, Wildlife Management, Fisheries Man- agement or Sustainable Development course or seminar should have access to a copy. Literature Cited: Ehrenfeld, D. 1993. Beginning again (pages 65-72). Oxford, Oxford University Press. Pages xiv + 216. Futuyma, D. 1998. Wherefore and whither the naturalist? American Naturalist 151(1): 1-6. Noss, R. F. 1998. Does conservation biology need natural history? Wild Earth 8: 10-14. Pruitt, W. O., Jr., and L. M. Baskin 2004. Boreal Forest of Canada and Russia/Taiga of Canada and Russia. Bilingual, English and Russian, parallel texts. Pensoft Publishers, Sofia, Bulgaria, illustrated: 163 pages ISBN 954-642-199-5. Schaefer, J. 2003. Long-term range recession and the persistence of Caribou in the taiga. Conservation Biology 17(5): 1435-1439. WILLIAM O. PRUITT, JR. Department of Zoology and Taiga Biological Station, Univer- sity of Manitoba, Winnipeg, Manitoba R3T 2N2 Canada 640 THE CANADIAN FIELD-NATURALIST Vol. 118 Resource and Environmental Management in Canada: Addressing Conflict and Uncertainty Edited by Bruce Mitchell. 2004. Third Edition. Oxford Uni- versity Press, Don Mills, Ontario, 608 pages. Can $46.50 Paper. Canada is a resource-rich country. The nation’s his- tory and folklore are a story of resource exploitation and the people who trapped, fished, farmed, mined and lumbered. In many ways, much has not changed as evidenced by the national wealth accrued from natural resources, the dependence of many towns and regions on primary resource use, and the growing social con- flicts centred on the type and intensity of resource use and conservation. It is generally acknowledged that all natural ecosys- tems are more or less human-dominated. Recent, well- cited studies have estimated that humans appropriate over 30% of the total terrestrial net primary produc- tion. This is a remarkable level of consumption for humans that represent roughly 0.5% of the total het- erotroph biomass on Earth. Spatially-explicit studies have identified appropriation levels that range from > 6% for South America to > 80% for south-central Asia. North America falls in between with a human appropriation level of 23%. Given these levels of human domination of the earth’s resources, it is understandable that the notions of conflict and uncertainty would be chosen as thematic foci for a book on resource and environmental man- agement; conflict among differing values and ideolo- gies, and uncertainties over adequate scientific and social knowledge. A long-standing textbook, now in its third edition, Resource and Environmental Management is intended primarily for undergraduate university students. The book’s 20 chapters are broken down into three parts: (1) emerging concerns, (2) enduring concerns, and (3) contemporary responses to these concerns. Emerging concerns include ecosystem health and ecological integrity, globalization and neo-conservatism, First Nations and resources, feminist perspectives, climate change, and water security. Enduring concerns focus on traditional resource issues such as fisheries, agricul- ture, forestry, wildlife management, mining, and parks and protected areas. The third part addresses some of the approaches used to develop resource and environ- mental visions. Among others, these include search conferences, ecosystem approaches, adaptive manage- ment, environment impact assessment, participatory approaches, governance issues, and environmental jus- tice. The majority of contributing authors are geogra- phy and environmental studies academics. In any collection of contributed essays, consistency in style and quality is always a challenge. Conflict and! uncertainty act as foundational themes and tie together all chapters. Most, if not all authors, attend to this theme and regularly make reference to other chapters in the book. Many of the chapters have been rewritten and updated for this edition and seven new chapters; have been added. The editor has prefaced each major; section with a good synthesis and summary of each) chapter thus helping the reader to situate the individ- ual contributions. Authors provide a sound scientific and technical analysis of each resource issue, generally up-to-date resource statistics, and policy implications. The strength of the edited volume is the authors’ attention to the social, cultural and political dimensions of resource and environmental management. A former forest ecolo- gy professor of mine was wont to proclaim: “Forestry is more about people than it is about trees.” The same can be said for all other resource issues. Because of the essential human dimension, ecosystem-based natural resource management is not rocket science — it’s much | more complicated than that. The amalgam of biophys- ical, cultural and social complexities fuel conflict and promote uncertainty. This inherent complexity of re- source and environmental management is highlighted throughout the text. Well discussed, as well, are a number of significant mechanisms that attempt to deal with the often intractable and inherently stochastic dimensions of institutional human behavior. I found some of the essays unnecessarily long- winded and somewhat pedantic. Twenty chapters weigh- ing in at over 600 pages do not make for easy or nec- essarily enjoyable reading, especially given the diversity of chapter topics. The chapter on forest management is noticeably remiss in addressing contemporary sus- tainable forest management paradigms. No mention is given, for example, of the current developments in the emulation of natural forest landscape disturbances, a key concept in sustainable forest management. As an edited book it does not possess the thrill or the fast-paced narrative of a single author. As a textbook, it has all the information, but you may find yourself simply dipping into chapters of specific interest. JOHN MCCARTHY Holy Rosary Parish, NIE 1V6 Canada 175 Emma Street, Guelph, Ontario d 2004 | MISCELLANEOUS By Jonathan A. Edlow. Second Edition, 2004. Yale Universi- ty Press, New Haven and London. 304 pages. U.S.$29.95 Paper. There have been few newly-emerging infectious dis- ;eases that have generated as much controversy as to their etiology and treatment as Lyme disease. From the “conventional” viewpoint, the disease is easily and accurately diagnosed, and short-term antibiotic treat- ment is effective; in the “alternative” point of view, ‘the diagnosis of late-presenting cases is often missed _ due to inaccurate diagnostic tests, and long-term antibi- _ Otic treatment of such cases is necessary for a cure. This readable book gives a fair and balanced account of the differing points of view of these two camps. It covers | the history of the discovery of Lyme disease, the elu- cidation of its causative agent, and the development of diagnostic tests and of treatments for the disease. | The disease was first recognized in North America when, around 1975, doctors at the Yale University School of Medicine noticed a new array of symptoms in patients from the area of Lyme, Connecticut. These symptoms consisted of a growing, ring-shaped rash (the bull’s-eye) at the site of a tick bite, and swollen joints. Sometimes the rash and the arthritis had ap- peared without the patient’s being aware of a tick bite. The author, Jonathan Edlow, M.D., is Vice-Chairman of the Department of Emergency Medicine, Beth Israel Deaconess Medical Center, and Assistant Professor of Medicine, Harvard Medical School. He himself describes the book as “a medical detective story” that _| flowed from a combination of his interest in infectious diseases, and the fact that relatives of his who had | moved to the Lyme area in the 1980s were stricken with symptoms of a bizarre arthritis, and/or rashes following _| tick bites. The author starts with a detailed and interesting j; account of how, in the 1970s, residents of the Lyme area were experiencing a baffling array of physical symp- toms, including rashes, unexplained neurological symp- toms and swollen joints, that eluded diagnosis by their "| family doctors. Then two women, who together with ; their families and neighbours, had been ill for several years, were separately urged by their doctors to visit the Rheumatology Clinic at the Yale School of Medi- cine in New Haven. After hearing their stories, the doc- tors at the clinic realized that about thirty-five cases ) : : ae clustered in the Lyme area were presenting similar ! | Edited by Peter Kareiva and Simon A. Levin. Princeton University Press, 41 William Street, Princeton, New Jer- sey 08540. 427 pages. U.S.$53.00 Paper. The ecological impact of the economic system we subscribe to could be viewed as the largest of all pos- BOOK REVIEWS 64] Bull’s Eye: Unraveling the Medical Mystery of Lyme Disease symptoms, and that an investigation into “what was happening at Lyme” was required. At the same time, U.S. Navy doctors at the nearby base in Groton, Con- necticut, observed several patients with ring-shaped rashes which spread over a large area. Their case report in The Journal of the American Medical Association (JAMA), was read by a family doctor at the eastern end of Long Island who had patients with tick bites surrounded by a rash, who thought that his patients might have the same disease. He realized it was similar to a disease that had been recognized in Europe since 1910 as erythema migrans (EM). In the European reports, the disease was associated with tick bites and thought to have a spirochaete as the causative agent. Having set the scene, the author describes the iso- lation of this causative agent by Willy Burgdorfer at the Rocky Mountain Laboratories, U.S. Public Health Service, who demonstrated that a spirochaete found in the deer tick, Ixodes scapularis, caused Lyme disease. The organism was later named Borrelia burgdorferi in his honour. But this tick can carry other pathogens (e.g., Babesia spp.) causing a co-infection in some patients, complicating both diagnosis and treatment. He further describes how, over time, a definite schism arose between the two groups of doctors maintaining the differing viewpoints mentioned. Patient advocacy groups arose who mainly supported the “alternative” position and, in 1993, a U.S. Senate Committee held hearings on the subject. The author sums up the cur- rent position by saying that debate remains about the best way to diagnose the disease, the utility of long- term antibiotic treatment, and the safety of a vaccine. A drawback in the author’s style arises when he uses analogies to try to explain scientific concepts. One example is that of a five-page description (complete with 5 Tables!) of the separation of peaches, apples, oranges and nectarines by machine, in an attempt to explain the meaning of the terms sensitivity and speci- ficity in assessing the accuracy of laboratory tests. The result is confusion. Appendix A lists the symptoms of Lyme disease, Appendix B lists tick-borne diseases in humans and animals. The book does not have a bibliography but. following the appendices, gives a list of sources for each chapter. PEARL PETERKIN #801 — 240 Brittany Drive, Ottawa, Ontario K1K OR7 Canada The Importance of Species: Perspective on Expendability and Triage sible species removal experiments. Unfortunately for us, this experiment is unreplicatable—we only get one shot at it. From the mid-point things don’t look good. When the consequences of our personal actions and public policies are compared to the cataclysmic impact of an asteroid (we are in the midst of the sixth and greatest wave of extinctions on the planet after all) you know we're in trouble. Still, the overwhelming number of species facing imminent elimination has lead some to ask if we need to protect every one? Surely of the millions of species we “share” Earth with there must be a few that we could afford to lose? Especially con- sidering the expense and inconvenience of conserva- tion programs. “The importance of species” sets out to answer that question. Or at least to summarize the collective experiences of the ecological community in examin- ing the idea of species expendability. The initial impe- tus for this volume came from a symposium to hon- our the accomplishments of Robert Paine. Dr. Paine’s pioneering work on the importance of species began nearly 40 years ago. His investigations were based on a simple idea: if you want to understand the role of a particular species in a community, remove it and see what happens. He works primarily in rocky inter-tidal zones. It was there he discovered that some species are absolutely critical to that ecosystem: remove one of these “keystones” and the community collapses. This conclusion has since been reached by many other ecologists working in different systems, and the book provides a nice introduction to their work. However, while keystone species have been identified in a variety of settings, their redundant counterparts have been harder to locate. The strength of this book lies in the diversity of voices that converge on a few common conclusions. The most critical outcome of a great deal of research is the recognition that identifying truly redundant spe- cies is an extremely difficult and perhaps impossible task. Each paper presents new obstacles. Louda and Rand present a particularly nasty problem in the first chapter. Native thistles of the Great Plains may ini- tially appear to be expendable as a direct result of their being essential! They argue that the native thistles serve to inoculate the community against the invasion of their exotic relatives. The presence of the native thistles sup- ports populations of herbivorous insects which readily switch to feeding on exotic thistle species that have been introduced to the region. In areas where this has been most successful the exotic thistles have been al- most completely excluded from the community. Under these conditions the exotic thistles do not appear to pose a serious threat to the community, so the value of the native thistles is not readily apparent. More generally, the environmental context may determine the relative importance of a species in a community. A keystone predator at one location may have negligible influence at another (Chapter 2). Sim- ilarly, the influence of a species may vary greatly over time (Chapter 3). Think of a herbivorous insect that spends nine years in small, widely scattered populations of minor importance, only to erupt in the tenth year to devastate thousands of hectares of forest (and feed THE CANADIAN FIELD-NATURALIST Vol. 118 millions of birds...). At a broader temporal scale, Stephen Palumbi (Chapter 15) notes that important ecological associations can evolve rapidly. A species that is redundant now may become an important com- ponent of its ecosystem in a few decades. The blink of an eye in evolutionary terms, but an eon measured in funding cycles. Taking a different approach Daniel Simberloff (Chapter 11) examines the impacts of the extinctions or near-extinctions of species one could reasonably assume to be important community mem- bers (i.e., American Chestnut, American Elm, Bison etc.). He reveals two related challenges: we have very little data to objectively assess the impacts of these losses, and there is no agreement on what constitutes a significant “ecosystem consequence”. As you would expect from a book with 28 authors the treatment is a little uneven. Given the emphasis on detailed natural history knowledge that many authors present as critical to the question, I was not convinced of the value of theoretical models in predicting which species are expendable. Shahid Naeem’s (Chapter 6) discussion of ecosystem reliability models starts from the assumption that species are interchangeable, effec- tively sidestepping the entire issue. The most troubling essay is perhaps the most impor- tant one in evaluating the concept of expendability. Mary Ruckelshaus, Paul McElhany, and Michael Ford collaborate in reviewing their efforts to conserve Pacific salmon species as part of the American National Marine Fisheries Service. Given that five of six of these pre- sumably economically important species are threatened or endangered, you might expect they would approach their task with the objective of maximizing the viability of the various stocks. On the contrary, they outline the process they followed in determining how many more populations they can afford to lose. This is a chilling glimpse of the possible future of conservation biology. The emphasis is not on how best to protect and enhance our biological heritage. Rather, conservation goals are set in a manner that minimizes their impact on eco- nomic objectives. Egbert Leigh’s contribution, “Social conflict, bio- logical ignorance, and trying to agree which species are expendable” (chapter 12) provides an insightful review of the entire debate. He identifies a critical need for basic natural history research if we are to have any hope of understanding the relative importance of indi- vidual species. More importantly, he reminds us “that social conflict, whether latent or blatant, is the biggest single obstacle to sound conservation. Suppose for a moment that we were Laplacean demons, able to assess the ecological consequences of each and every extinc- tion. Would this circumstance really cause society to close ranks behind conservation? As the world seems organized to destroy any sense of community among human beings, a greater knowledge of biology is not likely to make it easier for society to agree on conser- vation policy.” Leigh reinforces the value of natural 2004 | history in dealing with biological issues, while prop- | erly identifying the question of species expendability as part of a larger social problem. This is an important book, and deserves the critical attention of conservation biologists. In an age when we _ are pressured to apply a business model to all aspects of our society, it is important to examine the conse- | quences in a rigorous way. The essays in this book NEW TITLES +Available * Assigned Zoology *Common Birds of Ontario. By D. Sept. 2004. Calypso | Publishing, Box 1141, Sechelt, British Columbia VON 3A0. | 93 pages, Can. $12.95. ) +Birding in Venezuela (revised edition). By M. Goodwin. | 2004 Lynx Edicions Barcelona Spain. 332 pages, U.S.$25.60 | Paper. | Birds of the Middle East (Reprint of 1996 edition). By R. | Porter, S. Christensen and P. Schiermacker-Hansen. 2004. Princeton University Press, 41 William Street, Princeton, | New Jersey, USA, 08540-5237. 480 pages, U.S.$35. Paper, | U.S.$65 Cloth. | +Bumblebee Economics (2™ Edition). By B. Heinreich. | 2004. Harvard University Press, 79 Garden Street, Cambridge, Massachusetts 02138 USA. 245 pages, U.S.$19.95 Paper. _ *Damselflies of Alberta — Flying Neon Toothpicks in the | Grass. By John Acorn. 2004. University of Alberta Press, ! Ring House 2, Edmonton, Alberta T6G 2E1 Can. $29.95. _ Experimental Approaches to Conservation Biology. Edit- _ ed by M. S. Gordon and S. M. Bartol 2004. University of California Press, 2120 Berkeley Way, Berkeley, California 94704-1012. 358 pages, U.S.$75. *Frogs of Australia. By J. Turner. 2004. Pensoft Publishers. Acad. G. Bonchevy Str., Bl. 6, 1113 Sofia, Bulgaria. 276 pages, Eur 34.50 Cloth. *How the Earthquake Bird got its name and Other Tales of an Unbalanced Nature. By H. Shugart. 2005. Yale Uni- versity Press, P.O. Box 209040, New Haven, Connecticut 06520-9040. 227 pages, U.S. $30. Cloth. Hummingbirds. By L. Aziz. 2004. Firefly Books Ltd., 3680 Victoria Park Avenue, Toronto. 64 pages, Can. $9.95. Paper. *Mammals of Australia. By J. Turner. 2004. Pensoft Pub- lishers. Acad. G. Bonchev Str., Bl. 6, 1113 Sofia, Bulgaria. 216 pages, Eur 38.50 Cloth. *The Peregrine Falcon Survey in Canada. Edited by U Banasch and G. Holroyd. 2004. Canadian Wildlife Service 4999 — 98 Ave., Edmonton, Alberta T6X 2X3 Canada. No price available. Seabirds: a natural history. By A. Gaston. 2004. Yale Uni- versity Press, Box 209040, New Haven, Connecticut 06520. 224 pages, U.S.$45.00. BOOK REVIEWS 643 allow us to do that. The motivation behind identifying redundant species is that it would allow us to maximize the bang per conservation buck. However, cost-benefit analysis is of very limited value when you cannot quan- tify either costs or benefits. TYLER SMITH 155 Avenue Vanguard, Pointe Claire, Quebec H9R 3T4 Canada *Spiders of Australia. By T. Hawkeswood. 2004. Pensoft Publishers. Acad. G. Bonchev Str., Bl. 6, 1113 Sofia, Bulgaria. 264 pages, Eur 34.95 Cloth. *Venomous Reptiles of the Western Hemisphere. By J. Campbell and W. Lamar. 2004. Cornell University Press, 512 East State Street, Ithaca, New York 14850. 1032 pages (2 volumes), U.S.$149.95. *Whales and Dolphins of the World. By M. Simmonds. 2004. The MIT Press, Five Cambridge Centre, Cambridge, Massachusetts 02142. 160 pages, U.S.$29.95 Cloth. Botany Atlas of Plants of Nunavik Villages. By M. Blondeau and C. Roy. 2004. Editions Multimondes, 930 rue Pouliot, Sainte- Foy, Quebec G1V 3N9. Can. $34.95. *Boreal Forest of Canada and Russia. By W. Pruitt and L. Baskin. 2004. Russian Academy of Science, Leniasky pr., 33, Moscow V-71, Russia. 163 pages, Price not known. +Flower Chronicles (reprinted). By E. B. Hollingsworth. 2004 (original 1958). University of Chicago Press, 1427 East 60" Street, Chicago, Illinois 60637. 300 pages, U.S.$16. *Gathering Moss — A Natural and Cultural History of Mossess. By R. Kimmerer. 2004. Oregon State University Press, 102 Adams Hall, Corvallis, Oregan 97331. 176 pages, U.S.$17.95 Paper. Environment *Experimental Approches to Conservation Biology. Edit- ed by M. Gordon and S. Bariol 2004 University of Califor- nia Press. 343 pages, not illus. U.S.$ ? +Rendevous with the Wild — The Boreal Forest. Edited by J. Raffan (ed). 2004. BostonMills Press — Firefly Books Ltd.. 3680 Victoria Park Avenue, Toronto, Ontario M2H 3K1 188 pages, Can. $39.95 Miscellaneous The New Amateur Naturalist. By Nick Baker .2004. Harper Collins HarperCollins Publishers Ltd., Bloor Street East, 20" Floor, Toronto, Ontario M4W 1A8. 288 pages, £14.99 — approx. U.S.$38 Cloth. Wildlife Spectacles. By Russell A. Mittermeier et al. 2004. Conservation International 1919 M Street, NW Suite 600, Washington, DC 20036. 324 pages, £39.50 — approx. U.S.$74. News and Comment RENEW. 2004. Recovery of Nationally Endangered Wildlife in Canada. Annual Report (14) Contents (36 pages): Highlights of 2003-2004 [Of 160 Endangered, 108 Threatened, and 21 Extirpated species on the November 2003 COSEWIC list: 206 species are wholly or partially covered by recovery plans (completed or in develop- ment); 117 species are included in ecosystem or multispecies recovery, to the extent that their range overlaps with the geo- graphic area; 53 show stable or increasing population trend. $30.7 million was expended on recovery (salaries + expenses): Employment was equivalent to about 142 salaried and 23 vol- unteer people working full-time; 258 organizations made finan- cial contributions; 348 organizations participated on recovery teams] — Report from the Co-Chairs — Figures [4] — Species Recovery — Responsible Jurisdictions — Financial Contributors — Funding per Target — Canadian Wildlife Directors Commit- tee [contacts for species at risk]. The report was produced by the Canadian Wildlife Service of Environment Canada in cooperation with the provinces, territories, Fisheries and Oceans Canada, Parks Canada Agency, Wildlife Management Boards and many individuals and non-government groups. Readers are cautioned (inside front cover): “This informa- tion is carefully reviewed but the Canadian Wildlife Service disclaims responsibility for the accuracy of the information contributed”. Calendar: Endangered Reptiles of Canada 2004 and 2005 For 2004, 12 (one for each month) striking black-and- white drawings by various artists portray five turtles, 1 lizard and seven snakes whose surviving Canadian populations are designated as Endangered (Blue Racer), Threatened (Black Ratsnake, Eastern Foxsnake, Stinkpot, Eastern Ribbonsnake [Nova Scotia population], Blanding’s Turtle [Nova Scotia population], Eastern Massasauga), Special Concern (Wood Turtle, Five-lined Skink, Eastern Ribbonsnake [Ontario pop- ulations], Eastern Milksnake, Spotted Turtle), or Under Review (Prairie Rattlesnake, Blanding’s Turtle [Quebec, Ontario] populations). For each species there are brief remarks on ap- pearance, occurrence, habitat and risks to its survival. A 13% species, the Pacific Pond Turtle is featured on the cover as designated “extirpated” despite the lack of credible records that native populations ever existed in Canada with the spec- ulative remark “There is no biogeographic basis for the Pacific pond turtle’s range not to have extended in Canada, so cur- rent efforts to rebuild populations in Washington State may result in the return of the Pacific pond turtle to southwestern British Columbia”. The fine artwork was contributed by Joe Crowly (3), Kathryn Peiman (3), Mandi Eldridge (2), Kevin Kerr (2), Sonya Amin, Marisa Bonofiglio, and Sarah Ing- werson. In the 2005 version the black-and-white art is replaced by colour photography. The cover head of Western Rattlesnake (Crotalus oreganus) from British Colombia [sic] is followed by 7 snakes, 6 freshwater turtles, and one lizard each opposite a monthly calendar page covering December 2004 to Decem- ber 2005. Each has additional cameo shots of the same species and a paragraph that highlights its status and characteristics. Canadian Species at Risk November 2004 The latest version of the booklet Canadian Species at Risk prepared after the November 2004 COSEWIC assessment meeting is available on-line in both PDF and HTML formats Species represented are Eastern Ratsnake (Elaphe obsoleta) [= Western Ratsnake Elaphe spiloides of one recent study], Eastern Foxsnake (Elaphe gloydi) [= Elaphe vulpina gloydi\, Wood Turtle (Glyptemys insculpta), Spiny Softshell Apalone spinifera), Stinkpot (Sternotherous odoratus), Massasauga (Sisturus catenatus), Eastern Hog-nosed Snake (Heterodon platirhinos), Five-lined Skink (Eumeces fasciatus), Blanding’s Turtle (Emydoidea blandingi), Spotted Turtle (Clemmys gut- tata), Racers (Coluber constrictor: 3 subspecies), Snapping Turtle (Chelydra serpentina), Great Basin Gopher Snake (Pituophis catenifer deserticola). The back of the calender features 3 species supposedly extirpated from Canada: Timber Rattlesnake (Crotalus horridus), Eastern Box Turtle (Terra- pene carpolina carolina) and Pacific Pond Turtle (Actinemys marmorata). Not mentioned 1s that the latter two were likely introductions to Canada. Two other forms are not illustrated but listed: Pacific Gophersnake (Pituophis catenifer catenifer) and Pigmy Short-horned Lizard (Phrynosoma douglasii). These calendars have been produced by “the Brooks lab” at the Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1 Canada; (519) 824-4120 extension 58360; e-mail brookslab@hotmail.com. The selections for both cal- endars are based on the latest status designations by the reptile subcommittee (for which Brooks in chair) of the Committee on the Status of Endangered Wildlife (COSEWIC). See website: www.cosewic.ga.ca. All proceeds from sale of calendars goes to support the Kawartha Turtle Trama Centre in Peterborough, Ontario, established in 2002 to treat injured native turtles and release them back in the wild (website: www.kawarthaturtle. org) and reptile conservation research. on the web site http://www.cosewic.qc.ca/eng/sct5/indexe.cfm Paper format of this publication will now be available only once a year, following the spring assessment meeting. 644 2004 Number 65 (Focus on Australia), October 2004. Contents: | DAPTFE Seed Grants 2005 — The Impact of Habitat Modifica- | tion on the Striped Marsh Frog Limnodynastes peronii (Amy Gye) — Conservation Status of Australian Frogs (Jean-Marc Hero, Care Morrison, Graeme Gilespie, Dale Roberts. Paul Horner, David Newell, Ed Meyer, Keith McDonald, Frank Lemckert, Michhhhael Mahony, Michaelll Tyler, Will Osborne, Harry Hines, Steve Richards, Conrad Hoskin, Naomi Doak and Luke Shooo) — New Funds available from the DAPTE (Tim Halliday) — Alaska Conference Summary (Deborah Rudis — Froglog Shorts. | Number 66, December 2004. Contents: New Funds avail- _ able from the DAPTF (Tim Halliday & Don Church) — U.S. | Cricket Frog Symposium (Michael Lannoo and Ralph Grundel) _ — Press Release: Sunburt Frogs a Myth: Pond Scum offers | Natural Sunscreen (Larry Licht) — Global Amphibian Assess- / ment (Simon Stuart, Janice Chanson, Neil Cox and Bruce | Young) — Atelopus mucubajiensis still survives in Andes of _ Venezuela. Prelimnary Report (Cesar L. Barrio-Ammmmoros, | December 2004 Editor’s Note — Thank You from the Meeting Organizer — _ Overview of the Annual Meeting [Edmonton, Alberta, 24-27 _ September 2004] — 2004 Silver Salamander Award Winners | [Laura Frits, Steve Brechtel] — 2004 Best Student Presenta- _ tion [Connie Browne] — Meeting Field Trip: Tyrrell Museum and Reptile World — Meeting Field Trip: Garter Snake Dens | Marine Turtle Newsletter (106) October 2004. 28 pages: EDITORIAL Conceptual Problems _ with the ICUN Red Listing Assessment for the Green Turtle: _ Move Over Raine Island — GUEST EDITORIAL: Sea Turtles, | Red Listing, and the Need for Regional Assessments — _ ARTICLES: The Trade of Marine Turtles in the Toliara region, _ Souty West Madagasgar — NOTES: Rat Eradication as Part of a Green Turtle (Chelonia mydas) Conservation Prograamme in Indonesia — Fibropapillomatosis and Multiple Fibromas in a Green Turtle from the South Coast of Pernambucoo State, Brazil — First Record of a Leatherback Turtle in Syria — MEETING REPORTS — IUCNMTSG UPDATE — ANNOUNCEMENTS — NEWS & NEWS AND COMMENT 645 | Froglog: Newsletter of the Declining Amphibian Populations Task Force (65, 66) Fundacion AndigennA — The Afelopus Initiative: Conserving Endangered Tropical Andeannn Amphibians (Ariadne Angulo, Paul Salaman and Jose Vicente Rodriguez — Book Reviews — Froglog Shorts. Froglog is the bi-monthly newsletter of the Declining Amphibian Populations Task Force of The World Conserva- tion Union (IUCN)/Species Survival Commission (SSC) and is supported by The Open University, The World Congress of Herpetology, and Arizona State University. These issues were edited by John WIlkinson and Jeanne McKay (65) and Jeanne McKay (66), 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. Pub- lication is also supported by Peace Frogs www.peacefrogs.com and by RANA and the US National Science Foundation grants DEB-0130273. _ The Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile Conservation Network/Reseau Canadien de Conservation des Amphibiens et des Reptiles 9(1) — List of Presentations titles at 2004 Annual Meeting — Attendees of the 2004 Annual Meeting [55]. - Membership in CARCNET/RECCAR (contact Bruce Pauli, Canadian Wildlife Service, National Wildlife Research Centre, Carleton University, Raven Road, Ottawa, Ontario K1A 0H3. Web site: http://www.carcnet.ca/). LEGAL BRIEFS — RECENT PUBLICATIONS. The Marine Turtle Newsletter is edited by Brendan J. Godley and Annette C. Broderick, Marine Turtle Research Group, Centre for Ecology and Conservation, University of Exeter in Cornwall, Tremough Campus, Penryn TR10 9EZ United Kingdom; e-mail MTN @seaturtle.org; Fax +44 1392 263700. Subscriptions and donations towards the production of the MTN can be made online at or postal mail to Michael Coyne (online Editor) Marine Turtle Newsletter, 1 Southampton Place, Durham, North Carolina 27705 USA (e-mail: mcoyne @seaturtle.org). WARS 1983-2002 Special Annual Report Wildlife Accident Reporting and Mitigation The British Columbia Ministry of Transportation has published as a CD this report on the Wildlife Accident Reporting System (WARS) which describes its work and achievements of the last 20 years as one of the pioneers and leaders in wildlife accident reporting and mitigation. The WARS system in used in British Columbia to: 1) quantify the magnitude of wildlife-related vehicle colli- sions by species 2) identify accident-prone locations and accident trends 3) direct accident mitigation efforts (such as wildlife exclu- sion fencing and wildlife crossing structures) to problem locations 4) evaluate the effectiveness of accident risk profiles for highway corridors 5) develop species-specific accident risk profiles for high- way corridors 6) advance mitigation efforts to protect rare and endangered species 7) establish policies and strategies for accident issues and mitigation iniatives Copies of the CD can be obtained from Leonard Sielecki, R. P. Bio, MCIP, Engineering Branch, Environmental Manage- ment, British Columbia Ministry of Transportation, P.O. Box 9850, Station Provincial Government, Victoria British Columbia V8W 9T5 [location address: 4B — 940 Blanshard Street, Victoria] e-mail: leonard.sielecki@ gems9.gov.be.ca: telephone (250) 356-2255. The Ottawa Field-Naturalists’ Club Awards for 2003 IRWIN M. BRopo, CHRISTINE HANRAHAN, BEVERLY MCBRIDE, and SHEILA THOMSON At the Club’s Annual Soirée, held on 24 April 2004, at St. Basil’s Church in Ottawa, awards were once again given to members, and one non-member, who distinguished them- selves by accomplishments in the field of natural history and conservation, or by extraordinary activity within the Club. Daniel Strickland — Honorary Member The Ottawa Field-Naturalists’ Club takes great pleasure in honouring Daniel Strickland, not only for his renowned stud- ies on Gray Jays, but also for the major role that he has played in fostering an appreciation and enjoyment of natural history by the general public. Dan earned his Bachelor of Science degree from the Uni- versity of Toronto, followed by his Master of Science from the University of Montreal, choosing as his thesis the life history of the Gray Jay, which he studied in La Verendrye Park in Quebec. For more than thirty years, Dan has been studying and docu- menting Gray Jay behaviour in Algonquin Park, including the dynamics of the Gray Jay family unit, its nesting behaviour, winter survival and food gathering strategies. He is recognized world-wide as an authority on Gray Jays, and has co-authored the account on Gray Jays in the prestigious series, The Birds of North America. In 1970, Dan became Chief Naturalist for Algonquin Park, a position he held until his retirement in 2001. Under his leadership, Algonquin Park’s nature interpretation program has become renowned continent-wide. Tens of thousands of park visitors each year enjoy the Park’s nature walks, public Colin Bowen — George McGee Service Award 2003 When Colin Bowen first joined the Birds Committee 13 years ago, committee members recognized a man who would do an excellent job at whatever he took on. Since then, he has put in countless hours of solid, thorough work to keep the OFNC’s many bird-related materials coherent and accessible. Colin is extremely well-organized. With his involvement, information management projects get done effectively. In the mid-90s the Birds Committee urgently needed to consolidate and properly archive over thirty years’ worth of bird records and publications that were scattered in homes around the region. Colin formed the backbone of a team that worked for almost three years to bring it all together. He continues to maintain the archives so that, now, anyone can easily find what they need. As secretary of the Bird Records Sub-committee, Colin has a major task on his hands. Sub-committee membership has changed many times since it first reconstituted itself in the mid-90s, but Colin has stayed. He receives all rare bird reports and prepares and redistributes them to committee members for review. He tracks all debates and decisions at meetings and prepares the sub-committee’s minutes and reports. On top of all that, he set up and maintains the extensive cross-referenced bird records database and photo collection. A visit to the OFNC’s website will reveal fine examples of Colin’s efforts to bring Ottawa bird resources to the Internet As is frequently the case, there isn’t a winner for every poten- tial award and, this year, the Anne Hanes Natural History Award was not given. The following citations for those who did receive an award, however, were read to the members and guests assembled for the event. wolf howls, and natural history slide presentations. In addition, | seventeen self-guided nature trails, dozens of natural history publications on subjects as diverse as butterflies, mushrooms, mammals and lichens, as well as a world-class interpretive centre, all had their genesis under Dan Strickland’s tenure as Chief Naturalist. In Algonquin Park’s lively and informative publication, The Raven, Dan shared his broad knowledge and his fascination with nature in more than 300 articles. The inter- pretive programs he developed have contributed immeasurably to the goal of fostering public understanding and appreciation of nature. Not the least of Dan Strickland’s legacies is his impact on a whole generation of young park naturalists, trained under his guidance, who have moved on to new parks or to new fields of endeavour, profoundly influenced by Dan’s insight and his enthusiasm for natural history. In recognition of his contributions to our knowledge of the natural history of Gray Jay, as well as his outstanding career in the field of nature interpretation and public education, it is a pleasure to confer Honorary Membership in the Ottawa Field-Naturalists’ Club on Dan Strickland. and therefore to a broader community. Visitors can see the on- line. tabular seasonal Ottawa bird checklist, the beautifully tabulated Christmas Bird Count results or the often-consulted Ottawa birding site guide, complete with maps. Colin has volunteered for the Peregrine Falcon Watch and the Ontario Breeding Bird Atlas. He has helped coordinate many Christmas and Fall Bird Counts. When out of town bird- ers contact the club looking for birding information Colin is one of the generous volunteers who offer hours of their time to take them to our fine birding spots. Colin does so much work for the Birds and Bird Records committees that we wonder how he finds time to do anything else... but he does. He is a devoted and highly-valued volun- teer at the Canadian Museum of Nature and, together with his wife, Pat, were the featured volunteers in the museum’s recent newsletter. He is a grandfather, an amateur genealogist, a trav- eller, and is taking up woodworking. He can also fix or build just about anything. The OFNC thanks Colin for his tremendous contribution, and for retiring from his job with Bell Canada so he would have more time to come and work with us. He has truly never stopped, and we are proud to present him with this year’s George McGee Service Award. 646 2004 CLUB AWARDS 647 Daniel Brunton — Conservation Award for Members, 2003 The recipient of the 2003 Conservation Award for Mem- bers, Daniel Brunton, is an individual well known to mem- bers of the Ottawa Field-Naturalists’ Club through his active role in Club affairs for over 20 years, including a term as President. Indeed, his name is familiar to many residents of the Ottawa region because he is frequently consulted and quoted as an expert naturalist and conservationist by the Ottawa media. Dan has been an ecological consultant since 1979. The award he is receiving today, however, has to do with only one of his many activities on behalf of wildlife and wild spaces in the region, namely, his role in the establish- ment of the RiverKEEPER project for the Ottawa River. Dan has done almost 200 studies along the river, both as a profes- sional and amateur naturalist, making him eminently qualified to take this initiative. The problems associated with keeping track of the posi- tive and negative activities that impact the well-being of the Ottawa River demanded a new approach at coordination. With federal, two provincial, and dozens of municipal juris- dictions coming into play, and the high level of pollution af- fecting this crucial waterway, an initiative with vision was needed. Dan Brunton, together with some like-minded con- servationists, particularly George Brown, a Law professor at the University of Ottawa, and John Almstedt, got together to tackle the problem and recommended that the United States- based RiverKEEPER Alliance would provide the best frame- work within which to proceed. In the spring of 2001, a new Canadian chapter of RiverKEEPER was established with Dan as President. Its mission was to oversee the activities and legislation affecting the health of the river, and to report to the public. This would be done by hiring a full time KEEPER. Dan worked tirelessly to secure funding for the project and, after much work, succeeded in getting a three-year Trillium grant. In 2002, Laura Van Loon was appointed River KEEPER, launching the active phase of the project, and, in 2003, Dan was able to step down as President and allow others to carry on. As aresult of Dan Brunton’s efforts and vision, the public now has a full-time ombudsman to look after the river, advo- cating compliance with environmental laws, responding to citizen complaints, identifying problems that affect the river, and suggesting remedies. We are all beneficiaries of these efforts and have Dan Brunton to thank for helping to get the project started. The OFNC is therefore very pleased to pres- ent him with this year’s Conservation Award for Members. Friends of the Jock River — Conservation Award for Non-Member, 2003 The OFNC Conservation Award, non-member, is given to a group or individual who has made an outstanding contribu- tion in the cause of natural history conservation in the Ottawa Valley, with particular emphasis on activities within the Ottawa District. The Jock River originates in the Goodwood Marsh approx- imately 50 kilometers west of Ottawa and empties into the Rideau River after winding through fields, villages, and the provincially significant Richmond Fen. Many species of fish, birds, mammals and other wildlife depend on this waterway for food, breeding sites and shelter. Over the years large areas of riparian vegetation have been destroyed causing severe silting of the river, which in turn has had a serious impact on flora and fauna of the river and its shoreline. The Friends of the Jock River, a non-profit, volunteer environmental organ- ization, was formed to look at ways of protecting, restoring, and enhancing the Jock River and its watershed. Working with local landowners, government agencies, recreational users and businesses, the Friends have engaged in numerous projects aimed at restoring the health of the river’s ecosystem. The number and types of work they have under- taken is truly impressive. The Friends have been planting native trees and shrubs at various sites along the river and this is helping restore sections of the river’s shoreline to a more natural forested state. One example of their work is the Twin Elm restoration project. This project combines restoration planting with fencing of Pearl Peterkin -OFNC Member of the Year, 2003 This award is given to a member judged to have made sig- nificant contributions to the Club during the previous year. Looking back over the previous recipients of this award, it is illuminating to see the many and varied services performed for the Club. From rejuvenating committees, leading an ex- ceptional number of walks, editing Trail & Landscape, web site design and maintenance, the list is endless. But one thing is clear: joining the ranks of these hard-working, dedicated souls is no small thing. livestock from the riverbank and installation of a nosepump to meet their needs. Another important project undertaken by the group is The Rideau Valley Conservation Authority (RVCA) Stream Corridor Riparian Vegetation Survey. With training provided by the Conservation Authority, they col- lected data according to OMNR stream survey protocols to determine the ecological health of the river and its shoreline habitat. These data will become part of the RVCA database, which, in turn, will be made available to the Friends on CD to help them plan further riparian restoration projects. The Friends of the Jock River are involved with issues affecting the entire Jock River Watershed, thus they have been monitoring the ecology of the Richmond Conservation Area and advocating environmentally sound sewage treatment in Munster. The group is working on detailed guidelines for all future projects, including erosion control, buffering run-off, and en- hancing biodiversity in the watershed. They produce a quar- terly newsletter, hold regular meetings to which the public is invited, and maintain an informative website. For their dedicated, inspired, hard-working and thought- ful approach to protecting the health of the Jock River and its watershed, the OFNC is very pleased to give the Friends of the Jock River this Conservation Award. [Brian Finch, the present President; Eric Snyder, the Secretary; and Neil Barrington, the Past President, accepted the award on behalf of the Friends of the Jock River.] About ten years ago, it was decided to comb through back issues of Trail & Landscape and select those issues contain- ing articles of significant interest and value (for example, special issues on Butterflies, Reptiles and Amphibians, etc.) and retain a stockpile of these issues to sell. Since that orig- inal exercise, however, many more articles of interest have been published and Pearl Peterkin decided it was time to go through another selection process. She brought her sugges- tion to the Education and Publicity Committee who heartily 648 endorsed it. A suggestion was made to bundle together all issues on a particular topic and sell them as sets. This meant dealing with the entire 35 year run of Trail & Landscape, find- ing all the relevant issues, combining them, creating multiple sets, pricing them, finding adequate storage space at the Fletch- er Wildlife Garden centre and, of course, transporting the fin- ished products to the monthly meetings. During the past year, Pearl was often found ensconced in the back room of the Fletcher Centre, surrounded by stacks of Trail & Landscape, creating order out of chaos. She has also spent a lot of time there organizing the Education and Publicity Committee’s items so that they are more easily and quickly found. But it wasn’t only during 2003 that Pearl was active in Club affairs. Over the 15 years that Pearl has been an OFNC mem- ber, she has been involved with several club committees. In the 1990s she volunteered for three years with the Fletcher Wildlife Garden, becoming one of the stalwarts of this ener- getic group. In 2000 she took over as Chair of the Education and Publicity Committee at a time when the committee was at a low ebb with no leader. During her three years as Chair she successfully rejuvenated the committee, rounding up new members, finding judges for the Science Fair (which members of this Committee have been involved with for some time), finding volunteers to staff displays at various events, and do- ing all the other myriad jobs that keep a committee running THE CANADIAN FIELD-NATURALIST Vol. 118 smoothly. Once the committee was up and running smoothly, Pearl decided to step down from that role and concentrate on other aspects of Education and Publicity committee work. Many people probably recognize Pearl, even if they don’t know her name, because she is the friendly face behind the OFNC sales table at the monthly meetings. It requires a seri- ous commitment to attend every meeting, and, when people rely on you to be there, the responsibility is that much greater. Each month Pearl has to gather together the various sale items and, with the help of some other volunteers, transport them to and from the meeting. She also has to keep on top of new items (such as the new Bird Checklist), and make sure the stock of regular items is in good supply. This dedication has paid off, however, because during the last year the sales table has become a focus of much interest and activity. In her spare time (what spare time, you may ask), Pearl is an enthusiastic and long time member of the Rideau Trail Association. She also volunteers for Bird Studies Canada’s Loon Survey. The OFNC is fortunate to have such enthusiastic and com- mitted volunteers as Pearl Peterkin. For hard work on the Trail & Landscape project as well as all her involvement with the Education and Publicity Committee and her dedication to the club, we are pleased to give Pearl Peterkin the Member of the Year award for 2003. . Errata for The Canadian Field Naturalist 118(3) | Inside front cover bottom: and marine algae, and, still lower, the algae take over. This Cover. Intertidal zone on rugged rock headland of the Queen — caption and photo (the latter in colour) have previously Charlotte Islands, British Columbia. Each dominant species —_ appeared as Figure 64, page 77, in Lichens of North America creates a distinct zone. The white lichen is mainly Coccotrema _ by I. M. Brodo, S. D. Sharnoff, and S. Sharnoff. 2001. Yale maritimum and the black one is almost entirely Verrucaria University Press, New Haven Connecticut. See article by maura. Below the Verrucaria is a zone bare of both lichens — Brodo and Sloan on lichen zonation, pages 405-424. On page 408, Figure 2, the islands were omitted. 132°W 131°W 53°N 53°N 18 Limestone Island Moresby Island Ep Gwaii Haanas National Park Reserve and Haida Heritage Site and proposed National Marine Conservation Area 52°N 17 Treat Bay 52°N 50 Kilometres 132°W 131°W FiGuRE 2. Map of the Gwaii Haanas area showing Limestone Island (location 18) to the north and Treat Bay (location 17) to the south of the insets. 649 Index to Volume 118 Compiled by Leslie Cody Abies balsamea, 46,95,124,251,268,274,361,388,401,510,530 lasiocarpa, 191,355,584 Abrostola urentis, 541 Acanthopteroctetes bimaculata, 532 Acanthopteroctetidae, 532 Acasis viridata, 539 Acer, 52 negundo, 309 rubrum, 41,96,136,136,251,277,361,508 saccharinum, 41,124,251 saccharum, 361,507 spicatum, 362 Achigan a grande bouche, 523 a petite bouche, 523 Achillea millefolium, 166 Achnatherum richardsonii, 564 Acipenser fulvescens, 523 Acipenseridae, 523 Acleris albicomana, 534 britannia, 534 celiana, 534 emargana, 535 forbesana, 534 logiana, 534 nigrolinea, 535 obligatoria, 534 schalleriana, 534 variana, 534 Acorn, J.H., Review by, 468 Acossus centerensis, 534 populi, 534 Acrobasis betulella, 536 Acrocercops sp., 532 astericola, 532 Acronicta fragilis, 541 grisea, 541 impleta, 541 impressa, 541 innotata, 541 vulpina, 541 Acronictinae, 541 Actaea X ludovici, 612 Actitis macularia, 444 Adalia bipunctata, 312 Adela purpurea, 532 Adelidae, 532 Adelolecia koalensis, 406 Aechmophorus spp., 210 Aepyceros melampus, 185 Aesculus pavia, 612 Aethalura intertexta, 537 Aethes promptana, 535 promptana, 547 Aethusa cynapium, 613 Agabus sp., 431 arcticus, 431 confines, 431 thomsoni, 431 Agelaius phoeniceus, 201 Agelaius phoeniceus, Staging in Eastern South Dakota, Spring Dispersal Patterns of Red-winged Blackbirds, 201 Aglais milberti, 537 Agonopterix gelidella, 533 Agriphila ruricolella, 536 vulgivagella, 536 Agropyron spp., 400,584 cristatum, 300 pectiniforme, 560 repens, 300 sibiricum, 560 Agrostis sp., 169 filiculmis, 558 gigantea, 560 Aira praecox, 177 Aix sponsa, 131,230 Alaska, Changes in Loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) Populations in the Lower Mata- nuska-Susitna Valley, 210 Alaska, Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Pre- serve, 247 Alaska, Seed Dispersal by Brown Bears, Ursus arctos, in Southeastern, 499 Alaska, The Heather Vole, Genus Phenacomys, in, 438 Alberta: Distribution and Status, The Barred Owl, Strix varia in, 215 Alberta, First Record of Mountain Lions, Puma concolor, in — Elk Island National Park, 605 Alberta, Including New Provincial Records, Moths and But- terflies (Lepidoptera) of the Boreal Mixedwood For- est near Lac La Biche, 530 Alberta, Winter Habitat Use by Wolves, Canis lupus, in Rela- — | tion to Forest Harvesting in West-central, 368 | Alca torda, 258 Alces alces, 40,57,61,96,134,185,248,368,.606 Alder, 46,247,268,401 Speckled, 253 Alewife, 319 Alfalfa, 361 Algae, 320 Allium schoenoprasum ssp. sibiricum, 558 Alnus sp., 46,247,401 crispa, 268 i incana, 253 incana ssp. rugosa, 136 incanatable, 511 rugosa, 268 Alopecurus carolinianus, 170 Alopex lagopus, 602 Alopex lagopus, den by a Polar Bear, Ursus maritimus, Exca- vation of an Arctic Fox, 602 Alosa pseudoharengus, 319 | | | 650 | 4 | 1 — 2004 sapidissima, 77 _ Alucita lalannei, 536 | Alucitidae, 536 Alyssum americanum, 570 | obovatum, 558 | Amandinea coniops, 411 Amaranth, Purplish, 613 Amaranthus blitum, 613 Ambloplites rupestris, 523 _ Amblyptilia pica, 536 | Ambrosia x helenae, 613 Ameiurus nebulosus, 523 Amelanchier sp., 196 alnifolia, 558 | Amirault, D.L., 448 i Amirault, D.L., J. Kierstead, P. MacDonald and L. MacDon- nell. Sequential Polyandry in Piping Plover, Chara- drius melodus, Nesting in Eastern Canada, 444 | Ammodytes sp., 262 | Ammophila breviligulata, 444 _ Amphipacifica: Journal of Aquatic Systematic Biology 3(4), 294 | Amphipod, 77 _ Amphipoea americana, 541 | Ampithoe vallida, 79 | Anabeana sp., 320 | Anacampsis conclusella, 534 | Anaplectoides prasina, 543 | pressus, 543 | Anas crecca, 131 | platyrhynchos, 130 | rubripes, 129,268,604 _ Anas rubripes, Broods, Extraordinary Size and Survival of | American Black Duck, 129 _ Anathix puta, 542 _ Anatis labiculata, 312 _ Ancylis comptana, 535 i diminutana, 535 fuscociliana, 535 laciniana, 535 subaequana, 535 Andres, B.A., 210 Andromeda polifolia, 591 _ Andropogon gerardii, 342 Andropolia contacta, 542 Anguilla rostrata, 523 Anguille d’ Amérique, 523 Anguillidae, 523 Annotated List of the Arctic Marine Fishes of Canada, 156 Anser caerulescens, 602 Antelope, Roan, 185 | Anthoxanthum odoratum, 177 Anthriscus caucalis, 613 Anthyllis vulneraria, 613 | Anticlea multiferata, 539 vasiliata, 539 Apamea cogitata, 541 commoda, 541 Aplectoides condita, 543 Aplodinotus grunniens, 523 Apotomis capreana, 535 deceptana, 535 infida, 535 removana, 535 Apple, 299,361 INDEX TO VOLUME 118 65] Apple, Crab, 125 Apps, C.D., 191 Arabis caucasica, 558 holboellii var. retrofracta, 558 holboellii var. secunda, 558 Aralia nudicaulis, 46,365 Arbutus menziesii, 174 Arbutus, 174 Arceuthobium americanum, 595 douglasti, 595 laricis, 595 Arceuthobium americanum, on Jack Pine, Pinus banksiana, Growth in Manitoba, Effects of Lodgepole Pine Dwarf Mistletoe, 595 Archilochus colubris, 85 Archilochus colubris, Entanglements in Burdock, Arctium spp., at Delta Marsh, Manitoba, Ruby-throated Hum- mingbird, 85 Archips argyrospila, 535 Arctiidae, 540 Arctium spp., 85,253,611 lappa, 85 minus, 85 tomentosum, 85 Arctium spp., at Delta Marsh, Manitoba, Ruby-throated Hum- mingbird, Archilochus colubris, Entanglements in Burdock, 85 Ardea herodias, 231 Argyresthia sp., 532 abies, 532 conjugella, 532 goedartella, 532 oreasella, 532 pygmaeella, 532 Aristida basiramea, 276 Aristida basiramea Engelm. ex Vasey: A New Addition to the Flora of Quebec, Forked Three-awned Grass, 276 Armoracia rusticana, 558 Arrow-grass, Marsh, 561 Arrowhead, 318 Artemisia tridentata, 300,584 Arthonia phaeobaea, 410 Arthurs, W.R., Review by, 475 Asaphocrita sp., 533 Ascaphus truei, 447 Ascaphus truei, American Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, 446 Ash, 253 Black, 310 Green, 309 Mountain, 362 White, 361,508 Aspen, Quaking, 211,584 Trembling, 46,62,96,218,251,268,274,277,316,342,355 361,370,388,5 15,530 Asphodel, Northern False, 567 Aspicilia caesiocinerea, 411 supertegens, 424 Aspidotis densa, 166 Aster spp., 251 alpinus ssp. vierhapperi, 558 falcatus, 558 Aster, 251 Alpine, 577 Gulf of Saint Lawrence, 105 652 Western Heath, 577 Aster, Symphyotrichum laurentianum, (Fernald) Nesom, Esti- mation of Seed Bank and Seed Viability of the Gulf of Saint Lawrence, 105 Atkinson, J., Review by, 283 Atriplex alaskensis, 558 patula, 560 patula sensu, 568 patula var. alaskensis, 568 subspicata, 559 Auklet, Cassin’s, 616 Rhinoceros, 615 Auklet, Cerorhinca monocerata, Fledglings Fly to the Sea from Their Natal Burrows?, Do Rhinoceros, 615 Austrocedrus chilensis, 132 Autographa ampla, 541 bimaculata, 541 mappa, 541 rubida, 541 Avena sativa, 361,560 Avens, Yellow, 573 Awlwort, 572 Aythya collards, 268 marila, 268 Bacidia alaskensis, 412 Bacidina inundata, 424 Baeus sp., 122 Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Araneae, Lyco- sidae) in Southeastern Idaho, Occurrence of Parasi- toid Wasps, 122 Baillie, S.M., C.D. Wilkerson and T.L. Newbury. “Ashkui” Vernal Ice-cover Phenomena and Their Ecological Role in Southern Labrador, 267 Balanus, 406 Ball, P.W., 266 Baneberry, Hybrid, 612 Barbotte brune, 523 Barbue de riviére, 523 Barker, W., Review by, 474 Barley, 361 Barnacle, 406 Bartier, P.M., 77 Basilarchia arthemis, 537 Bass, Largemouth, 113 Basswood, 310 American, 253 Batrachedra praeangusta, 533 Batrachedridae, 533 Bear, 134,443 Barren-ground Grizzly, 239 Black, 41,96,370 Brown, 247,499 Grizzly, 117,239,247,370,603 Polar, 395,602 Bear, Ursus maritimus, Excavation of an Arctic Fox, Alopex lagopus, den by a Polar, 602 Bears, H. Parasite Prevalence in Dark-eyed Juncos, Junco hyemalis, Breeding at Different Elevations, 235 Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Preserve, Alaska, Interactions of Brown, 247 Bears, Ursus arctos, in Southeastern Alaska, Seed Dispersal by Brown, 499 THE CANADIAN FIELD-NATURALIST Vol. 118 Bears, Ursus arctos, in the Central Canadian Arctic, Move- ments of Subadult Male Grizzly, 239 Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, and Northern Bering Seas, Observations of Habitat Use by Polar, 395 Beaver, 40,129,248,267,606 Eurasian, 434 Beaver, Castor fiber, Collapsing Burrow Causes Death of a Eurasian, 434 Bedford, R.E., 626 Beech, American, 41,361,507 Beetle, 388 Carabid Intertidal, 79 Colorado Potato, 226 Convergent Lady, 309 Lady, 309 Ladybird, 225 Parenthesis Lady, 309 Seven-spotted Lady, 309 Thirteen-spotted Lady, 309 Transverse, 309 Water, 425 Beetles (Coccinellidae) in Overwintering Sites in Manitoba*, Density and Survival of Lady, 309 Beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Col- lected Along the Horton and Thelon Rivers in the Arc- tic Central Barrens of Canada, Predaceous Water, 425 Beetles (Coleoptera: Coccinellidae) in Prince Edward Island, Use of Eelgrass, Zostera marina, Wrack by Three Species of Ladybird, 225 Bennett, B., 558 Bentgrass, 169 Berberis spp., 132 Beringer, J., 185 Berube, D.K.A., 386 Besma quercivoraria, 538 Betula, 52 alleghaniensis, 41,361,508 glandulosa, 591 papyrifera, 41,46,95,136,211,251,268,274,355,388,510, 530 populifolia, 277 Bibarrambla allenella, 533 Bidens sp., 180 Bigler, L.L., 65 Birch, Dwarf, 591 Gray, 277 Paper, 211,268,355 White, 41,46,95,251,274,388,515,530 Yellow, 41,361,507 Bird rape, 571 Bison bison, 115,606 Bison, 115,606 Bison, Bison bison, Toward Elk, Cervus elaphus, and Wolves, Canis lupus, Unusual Behavior by, 115 Bison bison, Toward Elk, Cervus elaphus, and Wolves, Canis lupus, Unusual Behavior by Bison, 115 Biston betularia, 537 Bistort, Alpine, 591 Bittercress, Alpine, 571 Narrow-leaved, 613 Blackbird, Red-winged, 201 Yellow-headed, 207 Blackbirds, Agelaius phoeniceus, Staging in Eastern South Dakota, Spring Dispersal Patterns of Red-winged, 201 i — 2004 Bladderwort, Lesser, 576 | Blarina brevicauda, 402 Blastodacna curvilineella, 533 _ Bleptina caradrinalis, 540 Blueberry, 251 Lowbush, 195 Tall, 575 Velvet Leaf, 195 | Blueberry, Vaccinium angustifolium, and Velvet Leaf Blue- berry, Vaccinium myrtilloides, Production in Eastern Ontario, Effect of Fire Intensity and Depth of Burn on Lowbush, 195 Blueberry, Vaccinium myrtilloides, Production in Eastern Ontario, Effect of Fire Intensity and Depth of Burn on Lowbush Blueberry, Vaccinium angustifolium, and Velvet Leaf, 195 _ Bluegill, 318 | Bluegrass, 300 Kentucky, 176 Sandberg, 564 Bluestem, Big, 342 Bobcat, 40 Bog-rosemary, Dwarf, 591 | Bombycilla cedrorum, 232 Bonasa umbellus, 41,232 _ Bondia crescentella, 536 _ Boreal Dip Net/L’Epuisette Boreal: Newsletter of the Canadian | Amphibian and Reptile Conservation Network — Re- seau Canadien de Conservation des Amphbiens et des | Reptiles, The, 155,484,645 - Botrychium sp., 365 | Bousfield, E.L, 626 | Bowen, K.D. Frequency of Tail Breakage of the Northern Watersnake, Nerodia sipedon sipedon, 435 | Bowerbankia gracilis, 79 Brachylomia algens, 542 discinigra, 542 , Brachythecium spp., 253 | Bracken, 253 Branta canadensis, 268,602 Brassica rapa, 560 Braya glabella, 550 humilis, 550 pilosa, 550 purpurascens, 550 thorild-wulffii, 550 Braya, Pilose, 550 Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an Enigmatic Endemic of Arctic Canada, Pilose, 550 Braya pilosa Hooker (Cruciferae; Brassicaceae), an Enigmat- ic Endemic of Arctic Canada, Pilose Braya, 550 Brisson, J. Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: A New Addition to the Flora of Quebec, 276 Bristlegrass, Green, 564 British Columbia, Bird Communities of the Garry Oak Habitat in Southwestern, 376 British Columbia, Introduced Marine Species in the Haida Gwaii (Queen Charlotte Islands) Region, 77 British Columbia, Lichen Zonation on Coastal Rocks in Gwati Haanas National Park Reserve, Haida Gwaii (Queen Charlotte Islands), 405 British Columbia, 1972-2002, Sea Otter, Enhydra lutris, Sight- ings off Haida Gwaii/Queen Charlotte Islands, 270 INDEX TO VOLUME 118 653 British Columbia, Observations of Above-Surface Littoral For- aging in Two Sea Ducks, Barrow’s Goldeneye, Buceph- ala islandica, and Surf Scoter, Melanitta perspicillata, in Coastal Southwestern, 264 British Columbia waters: a reassessment, Status of marine turtles in, 72 Brodo, I.M. and N.A. Sloan. Lichen Zonation on Coastal Rocks in Gwaii Haanas National Park Reserve, Haida Gwaii (Queen Charlotte Islands), British Columbia, 405 Brome, Barren, 176 California, 174 Downy, 300 Rip-gut, 174 Smooth, 300,342,562 Soft, 174 Bromus spp., 175 carinatus, 174 ciliatus, 559 hordeaceus, 174 inermis, 300,342,560 rigidus, 174 sterilis, 176 tectorum, 174,300 Brook, R.K., 602 Broom, Scotch, 177 Brunton, D.F. Origins and History of The Ottawa Field- Naturalists’ Club, 1 Brush, Big Sage, 584 Bryotropha sp., 534 Bryozoan, 79 Bubo virginianus, 222,273 Bucculatricidae, 532 Bucculatrix sp., 532 canadensisella, 532 Bucephala clangula, 267,604 islandica, 264 Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in Coastal Southwestern British Columbia, Observa- tions of Above-Surface Littoral Foraging in Two Sea Ducks, Barrow’s Goldeneye, 264 Buckeye, Red, 612 Budworm, Spruce, 361 Bulrush, 318 Bunchberry, 46 Bunting, Indigo, 232 Bur-chervil, 613 Burdock, 85,253,611 Common, 85 Burdock, Arctium spp., at Delta Marsh, Manitoba, Ruby- throated Hummingbird, Archilochus colubris, Entan- glements in, 85 Burnett, D., Review by, 151 Burrows, F.G.M., 95 Buteo lagopus, 268 Buttercup, Western, 570 Butterflies (Lepidoptera) of the Boreal Mixedwood Forest near Lac La Biche, Alberta, Including New Provin- cial Records, Moths and, 530 Butterfly, 388 Buttons, Brass, 169 Cabera erythemaria, 538 variolaria, 538 Caenurgina crassiuscula, 541 654 Calafate, 132 Calamagrostis canadensis, 591 Calendar: Endangered Reptiles of Canada 2004 and 2005, 644 Callitriche anceps, 559 Callizzia amorata, 539 Caloplaca, 420 chlorina, 424 citrina, 411 exsecuta, 420 flavogranulosa, 411 inconspecta, 406 litoricola, 406 marina, 420 rosei, 411 verruculifera, 411 Caloptilia alnivorella, 532 anthobaphes, 532 betulivora, 532 canadensisella, 532 coroniella, 532 stigmatella, 532 Caloreas occidentella, 536 Camas, Common, 175 Great, 175 Camassia leichtlinii, 175 quamash, 175 Camelina sativa, 558 Campaea perlata, 538 Campbell, B., 521 Campion, Bladder, 569 Canachites canadensis, 41 Canada, Conservation Evaluation of Howell’s Triteleia, Tri- teleia howellii, an Endangered Lily in, 174 Canada, Conservation Evaluation of Lemmon’s Holly Fern, Polystichum lemmonii, a Threatened Fern in, 164 Canada, Conservation Evaluation of Small-flowered Lipo- carpha, Lipocarpha micrantha (Cyperaceae), in, 179 Canada, Conservation Evaluation of the Pacific Population of Tall Woolly-heads, Psilocarphus elatior, an Endan- gered Herb in, 169 Canada, Hybridization between a Green Turtle, Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and the First Record of a Green Turtle in Atlantic, 579 Canada, Pilose Braya, Braya pilosa Hooker (Cruciferae; Bras- sicaceae), an Enigmatic Endemic of Arctic, 550 Canada, Predaceous Water Beetles (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Collected Along the Horton and Thelon Rivers in the Arctic Central Barrens of, 425 Canada, Sequential Polyandry in Piping Plover, Charadrius melodus, Nesting in Eastern, 444 Canada, Stranding of a Pygmy Sperm Whale, Kogia brevi- ceps, in the Northern Gulf of St. Lawrence, 495 Canada 2004 and 2005, Calendar: Endangered Reptiles of, 644 Canadian Arctic, Movements of Subadult Male Grizzly Bears, Ursus arctos, in the Central, 239 Canadian Arctic Islands Using Morphology and Chloroplast DNA Data, A Systematic Analysis of the Alpine Saxi- frage Complex (Saxifragaceae) in the, 326 Canadian Field-Naturalist, Advice for Contributors to The, 158,294,494 THE CANADIAN FIELD-NATURALIST Vol. 118 Canadian Rocky Mountains, Predation on Two Mule Deer, Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the Southern, 191 Canadian Species at Risk, 484,644 Candelariella sp., 411 Canis domesticus, 247 latrans, 41,96,116,243,278,306,341,368,370,606,608 lupus, 42,57,61,95,115,127,134,194,247,278,350,361, 368,603,608 lupus lycaon, 608 rufus, 608 Canis latrans, Evidence for the Use of Vocalization to Coor- dinate the Killing of a White-Tailed Deer, Odocoileus virginianus, by Coyotes, 278 Canis latrans, in a Heterogeneous Environment, The Influ- ence of Prey Availability and Vegetation Character- istics on Scent Station Visitation Rates of Coyotes, 341 Canis latrans, Severe Chronic Neck Injury Caused by a Snare in a Coyote, 243 Canis lupus, at Katmai National Park and Preserve, Alaska, Interactions of Brown Bears, Ursus arctos, and Gray Wolves, 247 Canis lupus, in and near Pukaskwa National Park, Ontario, Demographic patterns and limitations of Grey Wolves, 95 Canis lupus, in Relation to Forest Harvesting in West-central Alberta, Winter Habitat Use by Wolves, 368 Canis lupus, Pack Density, Record High Wolf, 127 Canis lupus, South of the St. Lawrence River in Over 100 Years, First Confirmed Occurrence of a Wolf, 608 Canis lupus, Unusual Behavior by Bison, Bison bison, Toward Elk, Cervus elaphus, and Wolves, 115 Capelin, 262 Capreolus capreolus, 134 Caprimulgus vociferus, 230 Capsella bursa-pastoris, 560 Carcinus maenas, 82 Cardamine bellidifolia, 559 impatiens, 613 Carduelis pinus, 378 tristis, 378 Caretta caretta, 579 Caretta caretta, and the First Record of a Green Turtle in At- lantic Canada, Hybridization between a Green Turtle, Chelonia mydas, and Loggerhead Turtle, 579 Carex spp., 247,318,400,501 albo-nigra, 559 aquatilis, 591 athrostachya, 169,559 atrosquama, 559 capillaris ssp. chlorostachys, 559 crawfordii, 559 eleusinoides, 559 feta, 170 foenea, 266,559 glareosa, 266 hoodii, 266 houghtonii, 197 lapponica, 266 limosa, 559,591 loliacea, 559 magellanica ssp. irrigua, 266 marina, 266 obtusata, 559 pachystachya, 266,559 pellita, 559 petasata, 559 praegracilis, 559 rossii, 559 stylosa, 558 supina ssp. spaniocarpa, 559 sychnocephala, 559 i} unilateralis, 169 viridula, 559 Caribou, 39,192,241,362,602 Woodland, 57,61,96,119,368 ‘Caribou, Rangifer tarandus caribou, in Saskatchewan, Mor- phology of Female Woodland, 119 : Caripeta divisata, 538 | Carpiodes cyprinus, 523 _ Carpodacus purpureus, 378 Carposinidae, 536 Carter, B. A Tribute to Thomas Henry Manning (1911-1998), 618 Carterocephalus palaemon, 537 Case, R., 239 Cassiope tetragona ssp. saximontana, 559 | Castor canadensis, 40,129,136,248,267,606 fiber, 434 | Castor fiber, Collapsing Burrow Causes Death of a Eurasian Beaver, 434 Castors, 136 Caswell, P., 558 -Catharus fuscescens, 232 | Catillaria chalybeia, 408 | Catocala briseis, 541 relicta, 541 semirelicta, 541 unijuga, 541 Catocalinae, 541 Catostomidae, 521 Catostomus spp., 521 commersoni, 319,523 | Cattail, 253,318 Common, 545,561 | Cedar, Eastern White, 95,253,361,508 Yellow, 159 Centaurea diffusa, 300 Centipede, 389 Centrarchidae, 523 Cephalophus monticola, 185 | Cepphus grylle, 258 Cerastium arvense, 559 | Cerorhinca monocerata, 615 | Cerorhinca monocerata, Fledglings Fly to the Sea from Their Natal Burrows?, Do Rhinoceros Auklet, 615 | Certhia americana, 384 Cervus canadensis, 606 elaphus, 115,132,185,368 elaphus nelsoni, 132 nippon, 185 Cervus elaphus, and Wolves, Canis lupus, Unusual Behavior by Bison, Bison bison, Toward Elk, 115 Cervus elaphus, in Patagonia, Observations of Interactions between Puma, Puma concolor, and Introduced Euro- pean Red Deer, 132 Chamaecyparis nootkatensis, 159 Chamaerhodos erecta ssp. nuttallii, 559 | Chapdelaine, G., 256 INDEX TO VOLUME 118 655 Chapleau, F., 521 Charadrius alexandrinus, 445 melodus, 444,448 vociferus, 445 Charadrius melodus, egg viability after seawater immersion, Piping Plover, 448 Charadrius melodus, Nesting in Eastern Canada, Sequential Polyandry in Piping Plover, 444 Charlock, 572 Chat-fou liséré, 523 Chelonia mydas, 72,579,611 Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and the First Record of a Green Turtle in Atlantic Canada, Hybridization between a Green Turtle, 579 Chelydra serpentina, 610 Chelydra serpentina?, Is Cost of Locomotion the Reason for Prolonged Nesting Forays of Snapping Turtles, 610 Chenopodium capitatum, 559 Cherry, Choke, 253,362 Sand, 196 Chevalier blanc, 523 de riviere, 521 doré 526 rouge, 523 Chickadee, Black-capped, 232 Boreal, 48 Chestnut-backed, 378 Chickweed, Field, 569 Chilocorus stigma, 312 Chionodes continuella, 534 lugubrella, 534 mediofuscella, 534 occlusus, 534 psilopterus, 534 sattleri, 534 terminimaculella, 534 Chipmunk, Yellow-pine, 299 Chipmunks, Tamias amoenus, in Old Field and Orchard Habi- tats, Population Dynamics of Deer Mice, Peromyscus maniculatus, and Yellow-pine, 299 Chondrus crispus, 226 Chorda filum, 226 Chordaria flagelliformis, 226 Choreutidae, 536 Choreutis diana, 536 Choristoneura albaniana, 535 conflictana, 535 fumiferana, 361,535 rosaceana, 535 Choristostigma plumbosignale, 537 Chrysanympha formosa, 541 Chytolita petrealis, 540 Chytonix palliatricula, 542 Cicuta maculata vat. angustifolia, 559 Cinclus mexicanus, 443,446 Cinclus mexicanus, Preys Upon Larval Tailed Frogs, Asca- phus truei, American Dipper, 446 Cinquefoil, Tall, 573 Ciona intestinalis, 79 Ciprés, 132 Cirsium muticum, 253 Cirtwill, A., 604 Cladara atroliturata, limitaria, 539 Cladina rangiferina, 539 591 656 Clam, Butter, 77 Manila, 78 Native Littleneck, 77 Soft-shell, 78 Clangula hyemalis, 604 Clangula hyemalis, Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed Duck, 604 Clayburn, J.K., 615 Claytonia megarrhiza, 558 Cleavers, 175 Clemensia albata, 540 Clepsis clemensiana, 535 melaleucana, 535 persicana, 535 virescana, 535 Clintonia borealis, 362 Clintonia, Yellow, 362 Cliostomum griffithii, 424 Clostera albosigma, 540 apicalis, 540 strigosa, 540 Clover, 253,362 Red, 574 Cluff, H.D., 239 Coad, B.W., Reviews by, 281,282,469 Cobb, L.M. and V.A. Cobb. Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Ara- neae, Lycosidae) in Southeastern Idaho, 122 Cobb, V.A., 122 Coccinella septempunctata, 225,309 transversoguttata, 310 transversoguttata richardsonii, 309 Coccotrema cucurbitula, 420 maritimum, 406 Cochylis nana, 535 Cody, W.J., C.E. Kennedy, B. Bennett, and P. Caswell. New Records of Vascular Plants in the Yukon Territory VI, 558 Cody, W.J., Reviews by, 147,289,476 Coenophila opacifrons, 543 Coir6n Amargo, 132 Dulce, 132 Colaptes auratus, 358 Coleophora sp., 533 alnifoliae, 533 comptoniella, 533 corylifoliella, 533 dextrella, 533 duplicis, 533 glaucicolella, 533 mayrella, 533 mcedunnoughiella, 533 persimplexella, 533 pruniella, 533 rosaevorella, 533 Coleophoridae, 533 Coleotechnites atrupictella, 534 blastivora, 534 florae, 534 piceaella, 534 Collema fecundum, 411 novozelandicum, 420 Colletia spinosissima, 132 Comptonia peregrina, 196 THE CANADIAN FIELD-NATURALIST Vol. 118 Comtois, A., F. Chapleau, C.B. Renaud, H. Fournier, B. Camp-) bell, et R. Pariseau. Inventaire printanier d’une frayére multispécifique: l’ichtyofaune des rapides de la riv- iére Gatineau, Québec, 521 Contopus cooperi, 378 virens, 232 Cook, E.R., Reviews by, 138,141,467 Cook, J.A., 438 Copepod, 77 Corispermum hyssopifolium sensu, 568 ochotense var. alaskanum, 558 Cormorant, Double-crested, 257 Great, 257 Corn, 253 Cornus canadensis, 46 racemosa, 342 stolonifera, 253 Coronopus didymus, 613 Corophium acherusicum, 77 Corvus brachyrhynchos, 232 caurinus, 378 corax, 273 Corylus cornuta, 197,251 COSEWIC Assessment Results November 2003, 155 Cosmopterigidae, 534 Cossidae, 534 Cotton-grass, Short-anthered, 566 Cottontail, Eastern, 342 Cottonwood, Black, 211,355 Plains, 309 Cottus bairdi, 323 cognatus, 323 Cotula coronopifolia, 169 Couette, 523 Cougar, 159,370 Cougars, Puma concolor vancouverensis, Morphology and Population Characteristics of Vancouver Island, 159 Couleuvre brune, 136 mince, 135 rayée de l’est, 135 Couleuvre mince, Thamnophis sauritus septentrionalis, au Québec, Premiéres mentions de la, 135 Cowbird, Brown-headed, 378 Coyote, 41,96,116,243,278,306,341,368,606,608 Western, 608 Coyote, Canis latrans, Severe Chronic Neck Injury Caused by a Snare in a, 243 Coyotes, Canis latrans, Evidence for the Use of Vocalization to Coordinate the Killing of a White-Tailed Deer, Odocoileus virginianus, by, 278 Coyotes, Canis latrans, in a Heterogeneous Environment, The Influence of Prey Availability and Vegetation Characteristics on Scent Station Visitation Rates of, 341 Crab, European Green, 82 Crambidae, 536 Crambus leachellus, 536 perlellus, 536 Crapet arlequin, 523 de roche, 523 soleil, 523 Crassostrea gigas, 77 Crataegus sp., 365 Creeper, Brown, 378 Cress, Penny, 573 ii) | 2004 | Créte, M., 360 | Crow, American, 232 Northwestern, 378 | Cryptocala acadiensis, 543 | Cryptococcus spp., 41 Cryptogramma crispa vat. acrostichoides, 559 Cryptosula pallasiana, 79 Cuculliinae, 541 Cudweed, Lowland, 169 / Cumming, E.E. Habitat Segregation Among Songbirds in Old-Growth Boreal Mixedwood Forest, 45 Cyanocitta cristata, 231 Cyclophora pendulinaria, 538 Cydia flexiloqua, 536 populana, 536 | Cynosurus echinatus, 176 Cyperus sp., 180 squarrosus, 179 Cyperus, Awned, 179 | Cyprinidae, 523 Cytisus scoparius, 177 Dactylis glomerata, 174,300 ‘Daigle, C., M. Créte, L. Lesage, J.-P. Ouellet, and J. Huot. Summer Diet of Two White-tailed Deer, Odocoileus virginianus, Populations Living at Low and High Den- sity in Southern Québec, 360 | Danthonia californica, 169 spicata, 277 _| Daoust, P.-Y. and P.H. Nicholson. Severe Chronic Neck Injury Caused by a Snare in a Coyote, Canis latrans, 243 | Darbyshire, S.J. The Northern True Katydid, Pterophylla camellifolia (Orthoptera: Pseudophyllidae), at Ottawa, Ontario, 124 ) Darter, Iowa, 319 Johnny, 319 Rainbow, 319 | Dasychira plagiata, 540 vagans, 540 Daucus carota, 342 Deer, 368 Columbia Black-tailed, 162 European Red, 132 Mule, 132,162,185,191,370,606 Red, 185 Roe, 134 Sika, 185 White-tailed, 40,96, 127,185,278,360,370,606 | Deer, Cervus elaphus, in Patagonia, Observations of Inter- actions between Puma, Puma concolor, and Introduced European Red, 132 Odocoileus hemionus, by a Canada Lynx, Lynx cana- densis, in the Southern Canadian Rocky Mountains, | Predation on Two Mule, 191 Deer, Odocoileus virginianus, by Coyotes, Canis latrans, | Deer, Evidence for the Use of Vocalization to Coordinate the Killing of a White-Tailed, 278 Odocoileus virginianus, Populations Living at Low and High Density in Southern Québec, Summer Diet of Two White-tailed, 360 | Deer, | Deer, Odocoileus virginianus, with Medetomidine and Ket- amine, and Antagonism with Atipamezole, Immobi- lization of Clover-trapped White-tailed, 185 | Dendroica castanea, 48 cerulea, 229 INDEX TO VOLUME 118 657 coronata, 48,86,232,378 fusca, 48 magnolia, 48 pensylvanica, 232 petechia, 86,232 virens, 48,232 Dendroica cerulea Management on Eastern Ontario Bird Species, Predicting the Effects of Cerulean Warbler, 229 Depressariodes ciniflonella, 533 Dermacentor variabilis, 42 Dermochelys coriacea, 72,579 Deschampsia brevifolia, 559 Descurainia incisa vat. incisa, 559 Desmognathus ochrophaeus, 136 de Solla, S.R. and K.J. Fernie. Is Cost of Locomotion the Rea- son for Prolonged Nesting Forays of Snapping Turtles, Chelydra serpentina?, 610 Desroches, J.-F. et I. Picard. Extension de I’ aire de distribution connue de la Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du Québec, 441 Desroches, J.-F. et R. Lapar. Premiéres mentions de la Cou- leuvre mince, Thamnophis sauritus septentrionalis, au Québec, 135 Diachrysia aereoides, 541 Diarsia dislocata, 542 rosaria, 542 rubifera, 542 Dibb, A., 191 Dicamptodon copei, 448 ensatus, 448 Dichomeris levisella, 534 Dicrostonyx hudsonius, 403 richardsoni, 403 Digrammia rippertaria, 537 Dioryctria reniculelloides, 536 Dipper, American, 443,446 Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, Ascaphus truei, American, 446 Dock, Curled, 568 Dodecatheon hendersonii, 175 hendersonii ssp. hendersonii, 175 Dogtail, Hedgehog, 176 Dogwood, 253 Gray, 342 Dolichomia thymetusalis, 536 Doré jaune, 523 noir, 523 Dorosoma cepedianum, 318 Douglas, G.W., 179 Douglas, G.W. Conservation Evaluation of Lemmon’s Holly Fern, Polystichum lemmonii, a Threatened Fern in Canada, 164 Douglas, G.W. and J.L. Penny. Conservation Evaluation of Howell’s Triteleia, Triteleia howellii, an Endangered Lily in Canada, 174 Douglas, G.W. and J.M. Illingworth. Conservation Evalua- tion of the Pacific Population of Tall Woolly-heads, Psilocarphus elatior, an Endangered Herb in Canada, 169 Douglasia alaskana, 559 arctica, 558 Dowsley, M. and A. Cirtwill. Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed Duck, Clangu- la hyemalis, 604 658 Draba aurea, 559 corymbosa, 550,559 crassifolia, 559 glabella, 559 nemorosa, 559 ogilviensis, 559 palanderiana, 559 porsildii, 559 stenoloba, 559 Draba, Golden, 571 Drepana arcuata, 537 bilineata, 537 Drepanidae, 537 Dryas hookeriana, 559 Duchesne, L.C. and S. Wetzel. Effect of Fire Intensity and Depth of Burn on Lowbush Blueberry, Vaccinium angustifolium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, Production in Eastern Ontario, 195 Duck, American Black, 129,604 Black, 268 Longtailed, 604 Ring-necked, 268 Ruddy, 604 Wood, 131,230 Duck, Anas rubripes, Broods, Extraordinary Size and Survival of American Black, 129 Duck, Clangula hyemalis, Two Great Black-backed Gulls, Larus marinus, Kill Male Longtailed, 604 Ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in Coastal South- western British Columbia, Observations of Above- Surface Littoral Foraging in Two Sea, 264 Duiker, Blue, 185 Dulse, 226 Dutton, P.H., 579 Dyke, A.S., D.A. Hodgson, E.L. Bousfield, and R.E. Bedford. A Tribute to Victor Kent Prest (1913-2003), 626 Dysstroma citrata, 538 formosa, 538 hersiliata, 538 truncata, 538 walkerata, 538 Eagle, Bald, 267 Earthworm, 388 Ecliptopera silaceata, 538 Ectropis crepuscularia, 537 Editor’s Report for Volume 117 (2003), 296 Eelgrass, 225,318 Eelgrass, Zostera marina, Wrack by Three Species of Lady- bird Beetles (Coleoptera: Coccinellidae) in Prince Edward Island, Use of, 225 Egira dolosa, 542 Eider, Common, 258 Eilema bicolor, 540 Elachista sp., 533 adempta, 533 albicapitella, 533 Elachistidae, 533 Elaphria alapallida, 542 Elderberry, Red, 577 Eleocharis elliptica, 266 uniglumis, 559 Elk, 115,368 American, 132,606 THE CANADIAN FIELD-NATURALIST Vol. 118 Elk, Cervus elaphus, and Wolves, Canis lupus, Unusual Behavior by Bison, Bison bison, Toward, 115 Elm, American, 310,508 Elodea canadensis, 318 Elymus alaskanus ssp. alaskanus, 559 glaucus, 559 macrourus, 559 repens, 253 spicatus, 559 trachycaulus ssp. andinus, 559 Elytrigia intermedia, 560 Empidonax difficilis, 378 minimus, 86,232 Emydura macquarii, 610 Enargia decolor, 542 infumata, 542 Engeman, R.M., 201 Enhydra lutris, 270 Enhydra lutris, Sightings off Haida Gwaii/Queen Charlotte Islands, British Columbia, 1972-2002, Sea Otter, 270 © Ennominae, 537 Ennomos magnaria, 538 Eosphoropteryx thyatyroides, 541 Ephebe lanata, 424 Epilobium angustifolium, 253,365 Epinotia castaneana, 536 criddleana, 536 lindana, 536 momonana, 536 nisella, 536 rectiplicana, 536 solandriana, 536 transmissana, 536 trigonella, 536 Epirrhoe alternata, 539 Epirrita autumnata, 539 Epp, B. and J.C. Tardif. Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus banksiana, Growth in Manitoba, 595 Equisetum spp., 46 variegatum, 591 Equus cabalus, 370 Erannis tiliaria, 538 Erethizon dorsatum, 607 Eretmochelys imbricata, 580 Erickson, W.R. Bird Communities of the Garry Oak Habitat in Southwestern British Columbia, 376 Erigeron caespitosus, 559 hyperboreus, 559 Eriophorum brachyantherum, 559 Eritrichium aretioides, 559 chamissonis, 559 Ermine, 243 Ernst, J., 56,61 Errata for The Canadian Field-Naturalist 118(3), 649 Erratum Canadian Field-Naturalist 118(2), 485 Esocidae, 523 Esox lucius, 523 masquinongy, 523 Espino Negro, 132 Esturgeon jaune, 523 Etheostoma caeruleum, 319 exile, 319 nigrum, 319,523 olmstedi, 523 2004 | Eubaphe mendica, 539 Euchlaena marginaria, 538 obtusaria, 538 | tigrinaria, 538 _Eudonia albertalis, 536 | Eueretagrotis perattenta, 543 _Euhyponomeutoides gracilariella, 532 Eulia ministrana, 535 | Eulithis explanata, 538 propulsata, 538 | xylina, 538 Eulogia ochrifrontella, 536 _Euonymus fortunei, 614 | Euphrasia subarctica, 559 | Euphyia intermediata, 539 | Eupithecia anticaria, 539 i| assimilata, 539 columbiata, 539 i| perfusca, 539 ravocostaliata, 539 satyrata, 539 stellata, 539 | subfuscata, 539 | Euplexia benesimilis, 542 | Eurois astricta, 543 i occulta, 543 Eustrotiinae, 541 Euthyatira pudens, 537 | Everes amyntula, 537 Everett, K.T. and K.A. Otter. Presence of Cavities in Snags Retained in Forest Cutblocks: Do Management Poli- i cies Promote Species Retention?, 354 | | Evergestis pallidata, 537 Fagus, 52 i grandifolia, 41,361,507 Falseflax, 571 | Feathermoss, Red-stemmed, 591 | Felis concolor, 370 | Ferguson, C., 225 Ferguson, S.H. and D.K.A. Berube. Invertebrate Diversity under Artificial Cover in Relation to Boreal Forest Habitat Characteristics, 386 “Fern, 159 | Bracken, 196,251 Kruckeberg’s Holly, 164 | Lemmon’s Holly, 164 Mountain Holly, 164 | Nahanni Oak, 560 | Ostrich, 46,253 | Sensitive, 253 Sweet, 196 | Fern, Polystichum lemmonii, a Threatened Fern in Canada, Conservation Evaluation of Lemmon’s Holly, 164 | Fernie, K.J., 610 | Fescue, Rocky Mountain, 563 i Short-leaf, 562 . Festuca brachyphylla, 559 pallescens, 132 | richardsonii, 559 saximontana, 559 Filatima abactella, 534 _ Finch, Purple, 378 Fir, Balsam, 46,95,251,268,274,361,388,401,514,530 Douglas, 159,355,376,584,595 | i | INDEX TO VOLUME 118 659 Sub-alpine, 191,355,584 Fireweed, 253 Fisher, 56,96,583 Flicker, Common, 379 Northern, 358,378 Flounder, Starry, 248 Flueck, W.T. Observations of Interactions between Puma, Puma concolor, and Introduced European Red Deer, Cervus elaphus, in Patagonia, 132 Flycatcher, Least, 232 Olive-sided, 378 Pacific-coast, 378 Pacific-slope, 378 Least, 86 Fool’s-parsley, 613 Forget-me-not, 576 Forshner, S.A., P.C. Paquet, F.G.M. Burrows, G.K. Neale, K.D. Wade and W.M. Samuel. Demographic patterns and limitations of Grey Wolves, Canis lupus, in and near Pukaskwa National Park, Ontario, 95 Fouille-roche gris, 523 zébré, 523 Fournier, H., 521 Fox, 58,273 Arctic, 602 Red, 40,69,96,244,249 368 Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus, Excavation of an Arctic, 602 Fragaria sp., 365 Fraser, S., 225 Fratercula arctica, 258,604,615 cirrhata, 615 corniculata, 615 Fraxinus sp., 136,253 americana, 361,508 nigra, 310,365 pennsylvanica, 309 Frog, Red-legged, 448 Tailed, 447 Froglog: Newsletter of the Declining Amphibian Populations Task Force, 155,294,484,645 Frogs, Ascaphus truei, American Dipper, Cinclus mexicanus, Preys Upon Larval Tailed, 446 Fucus gardneri, 406 serratus, 226 Fungus, Nectria, 41 Furcellaria lumbricalis, 226 Furcula modesta, 540 occidentalis, 540 scolopendrina, 540 Fuscidea cyathoides, 420 intercincta, 421 mollis, 424 thomsonii, 406 Fuscopannaria leucostictoides, 411 maritima, 411 thiersii, 421 Galeopsis tetrahit ssp. bifida, 560 Galium aparine, 175 Garbary, D.J., S. Fraser, C. Ferguson, and R.F. Lauff. Use of Eelgrass, Zostera marina, Wrack by Three Species of Ladybird Beetles (Coleoptera: Coccinellidae) in Prince Edward Island, 225 660 Gau, R.J., B.D. McLoughlin, R. Case, H.D. Cluff, R. Mulders, and F. Messier. Movements of Subadult Male Grizzly Bears, Ursus arctos, in the Central Canadian Arctic, 239 Gaultheria procumbens, 196 shallon, 159 Gavia spp., 210 immer, 210 pacifica, 210 stellata, 258 Gavia spp.) and Red-necked Grebe (Podiceps grisegena) Populations in the Lower Matanuska-Susitna Valley, Alaska, Changes in Loon, 210 Gawn, M., Review by, 473 Gelechia dyarella, 534 lynceella, 534 Gelechiidae, 534 Gelis sp., 122 Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in South- eastern Idaho, Occurrence of Parasitoid Wasps, Baeus sp. and, 122 Gemsbok, 185 Gende, S.M., 499 Gentianella tenella, 559 Geometridae, 537 Geothlypis trichas, 86,232 Geum aleppicum ssp. strictum, 559 rossti, 559 Giberson, D.J., 425 Gillespie, L.J., 326 Ginns, J., Review, by, 146 Gladders, A., 159 Glaux maritimus, 226 Glen, W.M., 504 Gluphisia avimacula, 540 lintneri, 540 septentrionis, 540 Glyceria striata var. stricta, 559 Glycine max, 253 Gnaphalium palustre, 169 Gnatcatcher, Blue-grey, 232 Gnorimoschema septentrionella, 534 vastificum, 534 Goat, Mountain, 370 Goby, Round, 318 Tubenose, 318 Goby, Neogobius melanostomus Pallas (Gobiidae), and Eco- tone Utilisation in St. Clair Lowland Waters, Ontario, Description of Age-O0 Round, 318 Goh, K.M.L., 159 Goldeneye, 604 Barrow’s, 264 Common, 267 Goldeneye, Bucephala islandica, and Surf Scoter, Melanitta perspicillata, in Coastal Southwestern British Colum- bia, Observations of Above-Surface Littoral Foraging in Two Sea Ducks, Barrow’s, 264 Goldenrod, 253 Tall, 342 Goldfinch, American, 378 Goodwin, C.E., Review by, 138 Goose, Canada, 268,602 Snow, 602 Gracillariidae, 532 THE CANADIAN FIELD-NATURALIST Vol. 118 Grand brochet, 523 Graphiphora augur, 543 Grapholita lunatana, 536 Grass, 400 Annual Blue, 563 Cheat, 174 Creeping Bent, 562 Crested Wheat, 561 Forked Three-awned, 276 Fowl Marina, 563 Indian, 342 June, 563 Orchard, 174,300 Quack, 300 Western Rye, 562 Wheat, 584 Grass, Aristida basiramea Engelm. ex Vasey: A New Addition to the Flora of Quebec, Forked Three-awned, 276 Grayling, Arctic, 111 Grayling, Thymallus arcticus, Consumption of Shrews, Sorex spp., by Arctic, 111 Grebe, Red-necked, 210 Grebe (Podiceps grisegena) Populations in the Lower Mata- nuska-Susitna Valley, Alaska, Changes in Loon (Gavia spp.) and Red-necked, 210 Greya politella, 532 Griselda radicana, 536 Grosbeak, Rose-breasted, 232 Grouse, 41 Ruffed, 232 Guillemot, Black, 257 Gull, 249,442 Bonaparte’s, 443 Glaucous-winged, 443,616 Great Black-backed, 257,604 Herring, 257,443,449 Mew, 443 Ring-billed, 258 Gulls, Larus marinus, Kill Male Longtailed Duck, Clangula hyemalis, Two Great Black-backed, 604 Gulls, Larus spp., Foraging at Pink Salmon, Oncorhynchus gorbuscha, Spawning Runs, 442 Gulo gulo luscus, 56,61 Gulo gulo luscus, in the boreal forest, Effects of mid-winter snow depth on stand selection by Wolverines, 56 Gulo gulo luscus, Resting Sites and Caching Behavior in the Boreal Forest, Wolverine, 61 Gyalecta jenensis, 406 Gymnocarpium jessoense ssp. parvulum, 559 Gypsonoma adjuncta, 536 fasciolana, 535 salicicolana, 536 substitutionis, 536 Gyrinus opacus, 431 Habrosyne scripta, 537 Hadeninae, 541 Haemoproteus, 236 Hahn, A., 159 Hairgrass, Early, 177 Mountain, 564 Haliaeetus leucocephalus, 267 Hamr, J., 251 2004 Hansen, D.J. Observations of Habitat Use by Polar Bears, Ursus maritimus, in the Alaskan Beaufort, Chukchi, and Northern Bering Seas, 395 Hansen, L.P., 185 Haploa lecontei, 540 Hare, 607 Snowshoe, 41,58,191,278,585 Hare, J.F., G. Todd, and W.A. Untereiner. Multiple Mating Results in Multiple Paternity in Richardson’s Ground Squirrels, Spermophilus richardsonii, 90 | Harrimanella stellariana, 559 Harris, A.G., 179 Harris, J.G. Pilose Braya, Braya pilosa Hooker (Cruciferae; Brassicaceae), an Enigmatic Endemic of Arctic Cana- da, 550 Harris, S.A. and P.W. Ball. New Records of Cyperaceae and Juncaceae from the Yukon Territory, 266 Harrisimemna trisignata, 541 Hawk, Rough-legged, 268 _ Hay, 361 | Hayward, J.L. and J.K. Clayburn. Do Rhinoceros Auklet, Cerorhinca monocerata, Fledglings Fly to the Sea from Their Natal Burrows?, 615 Hazel, Beaked, 251 _ Healy, C. and L.J. Gillespie. A Systematic Analysis of the Alpine Saxifrage Complex (Saxifragaceae) in the Can- adian Arctic Islands Using Morphology and Chloro- plast DNA Data, 326 Hedeoma hispida, 277 Helcystogramma fernaldella, 534 Hellinsia homodactylus, 536 _ Hemicarpha, 179 | Hemlock, 41,361,499 Eastern, 251,508 Mountain, 159 Western, 159 Hemp-nettle, 576 Hepialidae, 532 Heracleum, 501 Herminiinae, 540 Heron, Great Blue, 230 Herteliana alaskensis, 408 Hesperiidae, 537 Heterocodon rariflorum, 170 Heterocodon, 170 Hillia iris, 542 Hinam, H.L., S.G. Sealy, and T.J. Underwood. Ruby-throated Hummingbird, Archilochus colubris, Entanglements in Burdock, Arctium spp., at Delta Marsh, Manitoba, 85 | Hiodon tergisus, 523 Hiodontidae, 523 Hippodamia convergens, 309 parenthesis, 309 tredecimpunctata, 225,309 Hippotragus equinus, 185 Hirundo rustica, 230 Hodgson, D.A., 626 Hogue, E.J., 299 Holodiscus discolor, 176 Homan, H.J., G.M. Linz, R.M. Engeman, and L.B. Penry. Spring Dispersal Patterns of Red-winged Blackbirds, Agelaius phoeniceus, Staging in Eastern South Dako- ta, 201 Homarus americanus, 82 INDEX TO VOLUME 118 66] Homoglaea hircina, 542 Homohadena badistriga, 5: infixa, 541 Homosetia fasciella, 532 Honeysuckle, Hairy, 176 Hood, G.A. and T. Neufeld. First Record of Mountain Lions, Puma concolor, in Elk Island National Park, Alberta, 605 Hop-clover, Small, 170 Hordeum brachyantherum, 559 vulgare, 361 Horse, Wild, 370 Houston, C.S., Reviews by, 142,290,465,466,470,47 1,481,631, 633,635 Huckleberry, Blue, 198 Huettmann, F., Reviews by, 478,632 Hummingbird, Ruby-throated, 85 Rufous, 378 Hummingbird, Archilochus colubris, Entanglements in Bur- dock, Arctium spp., at Delta Marsh, Manitoba, Ruby- throated, 85 Huot, J., 360 Hydrelia albifera, 539 Hydriomena furcata, 539 perfracta, 538 renunciata, 539 ruberata, 539 Hydroporus appalachius, 431 geniculatus, 431 hudsonicus, 432 morio, 431 Hydrozoan, 78 Hygrotus novemlineatus, 430 HAyles gallii, 540 Hylocichla mustelina, 232 Hylocomium splendens, 591 Hypatopa titanella, 533 Hypena atomaria, 541 edictalis, 541 humuli, 541 Hypeninae, 541 Hypenodes fractilinea, 540 Hyperaspis benedetti, 312 Hyppa contrasta, 542 Ichthyomyzon fossor, 523 unicuspis, 523 Ictaluridae, 523 Ictalurus punctatus, 523 Icterus galbula, 232 Idaea rotundopennata, 538 Idaho, Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in South- eastern, 122 Idia aemula, 540 americalis, 540 Illex illecebrosus, 497 Illingworth, J.M., 169 Ilybius erichsoni, 431 Impala, 185 Indian’s Dream, 166 Ipimorpha pleonectusa, 542 Tridopsis larvaria, 537 Tris setosa ssp. interior, 559 662 Iris, Wild, 567 Isoetes ?maritima, 559 echinospora, 559 Isopod, 389 James, M.C., K. Martin, and P.H. Dutton. Hybridization be- tween a Green Turtle, Chelonia mydas, and Logger- head Turtle, Caretta caretta, and the First Record of a Green Turtle in Atlantic Canada, 579 Jay, Blue, 231 Gray, 273 John, R., Reviews by, 140,143,147,150,282,288,291,630,631, 634,636 Joint, Blue, 591 Jolicoeur, H., 608 Jones, J., W.J. McLeish, and R.J. Robertson. Predicting the Effects of Cerulean Warbler, Dendroica cerulea Man- agement on Eastern Ontario Bird Species, 229 Junco hyemalis, 235,381 hyemalis cismontanus, 235 hyemalis hyemalis, 235 hyemalis oregonus, 235 Junco, Dark-eyed, 235,381 Junco hyemalis, Breeding at Different Elevations, Parasite Prevalence in Dark-eyed Juncos, 235 Juncos, Junco hyemalis, Breeding at Different Elevations, Parasite Prevalence in Dark-eyed, 235 Juncus spp., 400 bufonius, 170,559 filiformis, 559 Stygius SSp. americanus, 267 Juniper, Common, 166 Juniperus communis, 166 Kalmia, 501 angustifolia, 196 Katydid, Northern True, 124 Katydid, Prerophylla camellifolia (Orthoptera: PseudophyI- lidae), at Ottawa, Ontario, The Northern True, 124 Kemp, J.P. and C.R. Lacroix. Estimation of Seed Bank and Seed Viability of the Gulf of Saint Lawrence Aster, Symphyotrichum laurentianum, (Fernald) Nesom, 105 Kenagy, G.J., 111 Kennedy, C.E., 558 Kent, D., 264 Kierstead, J., 444 Killdeer, 445 Kinglet, Golden-crowned, 48,86 Ruby-crowned, 48,86 Kittiwake, Black-legged, 257 Knapweed, Diffuse, 300 Kneteman, J., 368 Kobresia simpliciuscula, 559 Kobresia, Simple, 566 Koeleria macrantha, 559 Kogia breviceps, 495 simus, 495 Kogia breviceps, in the Northern Gulf of St. Lawrence, Cana- da, Stranding of a Pygmy Sperm Whale, 495 Kohlmeyera complicatula, 420,424 Korscheltellus gracilis, 532 Kuzyk, G.W., J. Kneteman, and F.K.A. Schmiegelow. Winter Habitat Use by Wolves, Canis lupus, in Relation to Forest Harvesting in West-central Alberta, 368 THE CANADIAN FIELD-NATURALIST l’Esturgeon jaune, 521 Labidesthes sicculus, 319 Labrador, “Ashkui” Vernal Ice-cover Phenomena and Their | Ecological Role in Southern, 267 Labrador, First Record of a Barred Owl, Strix varia, in, 273 Lacanobia radix, 542 Lace, Queen Anne’s, 342 Lacinipolia lorea, 542 renigera, 542 Lacroix, C.R., 105 Lacroix, D.L., K.G. Wright, and D. Kent. Observations of ' Above-Surface Littoral Foraging in Two Sea Ducks, . Barrow’s Goldeneye, Bucephala islandica, and Surf | Scoter, Melanitta perspicillata, in Coastal Southwest- ern British Columbia, 264 Lactuca serriola, 300 Ladies’-tresses, Hooded, 567 Lady’s-fingers, 613 Lagopus lagopus, 602 Lambdina fiscellaria, 538 Lamna nasus, 245 Lampetra appendix, 523 Lamproie argentée, 523 de lest, 523 du nord, 523 Lamy, R., 270 Landry, J.-F., 530 Langor, D.W., 530 Lapar, R., 135 Laquaiche argentée, 523 Larch, Western, 595 Larentiinae, 538 Lariviere, S., Review by, 139 Larix laricina, 63,136,370,510,590 Larus spp., 249,442 argentatus, 258,443,449 canus, 443 delawarensis, 258 glaucescens, 443,616 marinus, 258,604 philadelphia, 443 Larus marinus, Kill Male Longtailed Duck, Clangula hye- malis, Two Great Black-backed Gulls, 604 Larus spp., Foraging at Pink Salmon, Oncorhynchus gor- buscha, Spawning Runs, Gulls, 442 Lasiocampidae, 539 Lasionycta poca, 542 antigReke225 Lauff, R.F., Reviews by, 144,145,288,630 Laura, 132 Laurel, Sheep, 196 Lecanora sp., 411 actophila, 420 contractula, 424 dispersa, 421 helicopsis, 420 muralis, 411 poliophaea, 424 | poliophaea ssp. poliophaea, 421 straminea, 411 tenera, 424 . : Lecidea conferenda, 424 Lecidella scabra, 424 stigmatea, 411 stigmatea f. stigmatea, 424 Mi — 2004 Ledum groenlandicum, 591 Lemming, Brown, 440 Varying, 403 Lemmus trimucronatus, 440 Lenga, 132 | Lepidium densiflorum var. densiflorum, 559 densiflorum var. macrocarpum, 560 Lepidochelys kempii, 580 Lepomis gibbosus, 523 macrochirus, 318,523 Leptinotarsa decemlineata, 226 Lepus spp., 607 americanus, 41,58,191,278,585 Lesage, L., 360 Leslie, J.K. and C.A. Timmins. Description of Age-O Round Goby, Neogobius melanostomus Pallas (Gobiidae), and Ecotone Utilisation in St. Clair Lowland Waters, Ontario, 318 | Lettuce, Prickly, 300 Leucania insueta, 542 Leucocytozoon, 236 Lichen, 408 Grey Reindeer, 591 Lichina pygmaea, 419 Lily-of-the-Valley, Wild, 362,567 Lim, B.K., Reviews by, 286,472 Limnaecia phragmitella, 534 _ Limnodrilus monothecus, 78 Limonium nashii, 226 Lingonberry, 591 Linz, G.M., 201 Lion, Mountain, 159,605 Lions, Puma concolor, in Elk Island National Park, Alberta, First Record of Mountain, 605 Lipocarpha micrantha, 179 Lipocarpha, Small-flowered, 179 Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada, Conservation Evaluation of Small-flowered, 179 Lipocarpha micrantha (Cyperaceae), in Canada, Conservation Evaluation of Small-flowered Lipocarpha, 179 Lithacodia albidula, 541 Litholomia napaea, 542 Lithomoia germana, 542 Lithophane innominata, 542 petulca, 542 Lobophora nivigerata, 539 Lobster, Atlantic, 82 Locoweed, Field, 574 _ Lolium perenne, 170,174 Lomatia hirsuta, 132 Longcore, J.R. and D.G. McAuley. Extraordinary Size and Survival of American Black Duck, Anas rubripes, Broods, 129 Lonicera hispidula, 176 Lontra canadensis, 96 Loon, 210 Common, 210 Pacific, 210 Red-throated, 256 Loon (Gavia spp.) and Red-necked Grebe (Podiceps grise- gena) Populations in the Lower Matanuska-Susitna Valley, Alaska, Changes in, 210 Lophocampa maculata, 540 Lophodytes cucullatus, 131 Lotus corniculatus, 365 INDEX TO VOLUME 118 663 Lousewort, Oeder’s, 576 Lupinus nootkatensis, 559 Lutra canadensis, 267 Luxilus cornutus, 523 Luzula arcuata ssp. unalaschkensis, 559 Lycaenidae, 537 Lycia ursaria, 537 Lycopodium complanatum, 196 Lymantriidae, 540 Lynx, 57,96,134 Canada, 191,583 Lynx canadensis, 57,96,191,583 lynx, 134 rufus, 40 Lynx canadensis, Distributions, A Snow-tracking Protocol Used to Delineate Local Lynx, 583 Lynx canadensis, in the Southern Canadian Rocky Mountains, Predation on Two Mule Deer, Odocoileus hemionus, by a Canada Lynx, 191 Lynx, Lynx canadensis, Distributions, A Snow-tracking Pro- tocol Used to Delineate Local, 583 Lynx, Lynx canadensis, in the Southern Canadian Rocky Mountains, Predation on Two Mule Deer, Odocoileus hemionus, by a Canada, 191 Lyonetia prunifoliella, 533 Lyonetiidae, 533 Macaria bitactata, 537 brunneata, 537 loricaria, 537 signaria, 537 ulsterata, 537 MacDonald, P., 444 MacDonald, S.O., A.M. Runck, and J.A. Cook. The Heather Vole, Genus Phenacomys, in Alaska, 438 MacDonnell, L., 444 Macleaya cordata, 614 Madden, E.M., 400 Mahonia aquifolium, 176 Maianthemum, 500 canadense, 196,362 canadense ssp. interius, 559 Maier, T.J., 400 Maitén, 132 Malachigan, 523 Malacosoma disstria, 540 Mallard, 130 Mallotus villosus, 262 Malus sp., 125 domestica, 299 pumila, 361 Manitoba*, Density and Survival of Lady Beetles (Coccinel- lidae) in Overwintering Sites in, 309 Manitoba, Effects of Lodgepole Pine Dwarf Mistletoe, Arceu- thobium americanum, on Jack Pine, Pinus banksiana, Growth in, 595 Manitoba, Hummock Vegetation at the Arctic Tree-line near Churchill, 590 Manitoba, Ruby-throated Hummingbird, Archilochus colubris, Entanglements in Burdock, Arctium spp., at Delta Marsh, 85 Manning (1911-1998), A Tribute to Thomas Henry, 618 Maple, Manitoba, 309 Mountain, 362 Red, 41,96,251,277,361,508 664 Silver, 125 Sugar, 41,124,251,361,507 Marigane noire, 523 Marine Turtle Newsletter, 156,294,484,645 Marmot, Hoary, 440 Vancouver, 403 Marmota caligata, 438 vancouverensis, 403 Marten, 58,63 American, 96,438 Pine, 243 Martes americana, 58,63,96,243,438 pennanti, 56,96,583 Martin, K., 579 Maskinongé, 523 Matteuccia struthiopteris, 46,253 Maytenus boaria, 132 McAlpine, D.F., S.A. Orchard, K.A. Sendall, and R. Palm. Status of marine turtles in British Columbia waters: a reassessment, 72 McAuley, D.G., 129 McCarthy, J, Reviews by, 150,152,640 McIntyre, R.T., 115 McKelvey, K.S., 583 McKnight, J., L. Thomas, and D.L. Amirault. Piping Plover, Charadrius melodus, egg viability after seawater im- mersion, 448 McLeish, W.J., 229 McLoughlin, P.D., 239 McNicholl, M.K., Review by, 285 Meadow-foxtail, Carolina, 170 Measures, L., B. Roberge, and R. Sears. Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the Northern Gulf of St. Lawrence, Canada, 495 Mech, L.D. and S. Tracy. Record High Wolf, Canis lupus, Pack Density, 127 Mech, L.D., R.T. McIntyre and D.W. Smith. Unusual Beha- vior by Bison, Bison bison, Toward Elk, Cervus ela- phus, and Wolves, Canis lupus, 115 Medicago sativa, 361 Megasema c-nigrum, 543 Melanchra adjuncta, 542 Melanitta perspicillata, 264 Melanitta perspicillata, in Coastal Southwestern British Columbia, Observations of Above-Surface Littoral Foraging in Two Sea Ducks, Barrow’s Goldeneye, Bucephala islandica, and Surf Scoter, 264 Meleagris gallopavo silvestris, 251 Meleagris gallopavo silvestris, Behavior in Central Ontario During Winter, Wild Turkey, 251 Melica subulata, 176 Méné a nageoires rouges, 523 émeraude, 523 jaune, 523 Meningeal Worm, 40 Mephitis mephitis, 40,69 Merganser, Hooded, 131 Mertensia paniculata var. alaskana, 559 Mesoleuca ruficillata, 539 Messier, F., 239 Metalepsis salicarum, 543 Metanema determinata, 538 inatomaria, 538 Meunier noir, 523 Meyer, T.M., 185 THE CANADIAN FIELD-NATURALIST Vol. 118 Mice, Peromyscus maniculatus, and Yellow-pine Chipmunks, Tamias amoenus, in Old Field and Orchard Habitats, | Population Dynamics of Deer, 299 Micranthes kumlienti, 326 nivalis, 326 tenuis, 326 Micropterus dolomieu, 523 salmoides, 113,523 Microtus, 345 longicaudus, 300 montanus, 300 ochrogaster, 342 pennsylvanicus, 342,400 Milfoil, Eurasian, 318 Millipede, 388 Mills, T.K. and B.A. Andres. Changes in Loon (Gavia spp.) and Red-necked Grebe (Podiceps grisegena) Popula- tions in the Lower Matanuska-Susitna Valley, Alaska, 210 Millspaugh, J.J., B.E. Washburn, T.M. Meyer, J. Beringer, and L.P. Hansen. Immobilization of Clover-trapped White-tailed Deer, Odocoileus virginianus, with Med- etomidine and Ketamine, and Antagonism with Ati- pamezole, 185 Mimulus guttatus, 559 Mink, 96 European, 185 Minnow, Bluntnose, 318 Minuartia dawsonensis, 559 elegans, 559 macrocarpa, 559 obtusiloba, 166 yukonensis, 559 Minutes of the 125" Annual Business Meeting of the Ottawa Field-Naturalists’ Club 13 January 2004, 486 Mistletoe, Lodgepole Pine Dwarf, 595 Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus banksiana, Growth in Manitoba, Effects of Lodgepole Pine Dwarf, 595 Mizuhopecten yessoensis, 78 Mniotilta varia, 230 Molothrus ater, 382 Mompha sp., 533 albapalpella, 533 terminella, 533 Monarda punctata, 277 Monkeyflower, Yellow, 576 Monopis laevigella, 532 spilotella, 532 Montia fontana, 559 Moore, J.W. and G.J. Kenagy. Consumption of Shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus, 111 Moose, 40,57,61,96,134,185,248,368,606 Moose Sickness, 40 Morrissey, C.A. and R.J. Olenick. American Dipper, Cinclus mexicanus, Preys Upon Larval Tailed Frogs, Ascaphus truei, 446 Moss, 253 Golden Fuzzy Fen, 591 Irish, 226 Stair-step, 591 Moth, 388 Moths and Butterflies (Lepidoptera) of the Boreal Mixedwood Forest near Lac La Biche, Alberta, Including New Provincial Records, 530 2004 | Mountain-heather, Yellow, 575 Mountain-parsley, 560 | Mouse, 342 Deer, 299,342,402 Great Basin Pocket, 300 Meadow Jumping, 403 Western Harvest, 300 White-footed, 299,342 Moxostoma spp., 521 anisurum, 523 carinatum, 523 erythrurum, 526 macrolepidotum, 523 Mulders, R., 239 | Mulinum spinosum, 132 ! Mullein, Great, 300 Muntz, E.M. and B.R. Patterson. Evidence for the Use of Vocalization to Coordinate the Killing of a White- Tailed Deer, Odocoileus virginianus, by Coyotes, Canis latrans, 278 Murray, D.F., Review by, 637 ' Murre, Common, 257 Thick-billed, 602 Musaraigne cendrée, 441 fuligineuse, 441 Musaraigne fuligineuse, Sorex fumeus, dans le nord-est du Québec, Extension de |’ aire de distribution connue de la, 441 _ Muskrat, 267,602,607 | Mussel, Bay, 264 - Mustard, Tansy, 571 Mustela spp., 96,306 erminea, 243 lutreola, 185 nivalis, 403 vison, 96 | Mya arenaria, 78 Myelopsis subtetricella, 536 _ Myosotis scorpioides, 560 | Myriophyllum spicatum, 318 verticillatum, 559 Myrrhis odorata, 614 Myrrhis, Scented, 614 Mythimna oxygala, 542 _ Mytilicola orientalis, 77 | Mytilus edulis, 264 trossulus, 264 Nadata gibbosa, 540 | Naemia serriata, 227 Naseux des rapides, 523 Navarretia squarrosa, 169 Neale, G.K., 95 Nectria coecinea vat. faginata, 41 Needle-and-thread, 564 Needlegrass, Spreading, 564 Nemapogon acapnopennella, 532 roburella, 532 Nematocampa resistaria, 538 Nemophila parviflora, 176 Nemophila, Small-flowered, 176 Neneo, 132 Neogobius melanostomus, 318 INDEX TO VOLUME 118 665 Neogobius melanostomus Pallas (Gobiidae), and Ecotone Utilisation in St. Clair Lowland Waters, Ontario, Des- cription of Age-O Round Goby, 318 Neotelphusa praefixa, 534 Nepticulidae, 532 Nerodia sipedon sipedon, 435 Nerodia sipedon sipedon, Frequency of Tail Breakage of the Northern Watersnake, 435 Nettle, Stinging, 567 Neufeld, T., 605 Newbury, T.L., 267 Nguyen, L.P., J. Hamr, and G.H. Parker. Wild Turkey, Melea- gris gallopavo silvestris, Behavior in Central Ontario During Winter, 251 Nicholson, P.H., 243 Niditinea orleansella, 532 Nites betulella, 533 Noctuidae, 540 Nostoc sp., 594 Notemigonus crysoleucas, 523 Nothofagus antarctica, 132 pumilio, 132 Notocelia culminana, 535 Notodonta simplaria, 540 Notodontidae, 540 Notropis atherinoides, 319,523 hudsonius, 318,523 Noturus insignis, 523 Nova Scotia, Continuing Environmental Change: An Example from, 39 Nuphar polysepalum, 559 Nuthatch, Red-breasted, 384 Nycteola frigidana, 540 Nymphalidae, 537 Nymphalis antiopa, 537 Nire, 132 Oak, 342 Bur, 310 California Black, 383 English, 125 Gambel, 383 Garry, 174,376 Red, 124,251 Oak Habitat in Southwestern British Columbia, Bird Com- munities of the Garry, 376 Oat, 361,562 Oatgrass, California, 169 Oceanodroma leucorhoa, 258 Oceanspray, 176 Ochrolechia subplicans, 411 subplicans ssp. hultenii, 420 subplicans ssp. subplicans, 420 Odocoileus spp., 368 hemionus, 132,162,185,191,370,606 hemionus columbianus, 162 virginianus, 40,96, 127,185,278,360,370.606 Odocoileus hemionus, by a Canada Lynx, Lynx canadensis, in the Southern Canadian Rocky Mountains, Preda- tion on Two Mule Deer, 191 Odocoileus virginianus, by Coyotes, Canis latrans, Evidence for the Use of Vocalization to Coordinate the Killing of a White-Tailed Deer, 278 666 Odocoileus virginianus, Populations Living at Low and High Density in Southern Québec, Summer Diet of Two White-tailed Deer, 360 Odocoileus virginianus, with Medetomidine and Ketamine, and Antagonism with Atipamezole, Immobilization of Clover-trapped White-tailed Deer, 185 Oecophoridae, 533 Oenothera biennis, 253 Olenick, R.J., 446 Olethreutes glaciana, 535 metallicana, 535 Olethreutinae, 535 Oligia illocata, 541 mactata, 541 Oligochaete, 78 Omisco, 523 Oncorhynchus gorbuscha, 442 keta, 248 nerka, 111,249,443 Oncorhynchus gorbuscha, Spawning Runs, Gulls, Larus spp.. Foraging at Pink Salmon, 442 Ondatra zibethicus, 267,602,607 Onion, Wild, 567 Oniongrass, Alaska, 176 Onoclea sensibilis, 253 Ontario Bird Species, Predicting the Effects of Cerulean War- bler, Dendroica cerulea Management on Eastern, 229 Ontario, Demographic patterns and limitations of Grey Wolves, Canis lupus, in and near Pukaskwa National Park, 95 Ontario, Description of Age-O Round Goby, Neogobius mela- nostomus Pallas (Gobiidae), and Ecotone Utilisation in St. Clair Lowland Waters, 318 Ontario During Winter, Wild Turkey, Meleagris gallopavo silvestris, Behavior in Central, 251 Ontario, Effect of Fire Intensity and Depth of Burn on Low- bush Blueberry, Vaccinium angustifolium, and Velvet Leaf Blueberry, Vaccinium myrtilloides, Production in Eastern, 195 Ontario Natural Heritage Information Centre Science and Information Newsletter 9(1) Winter 2004, 156 Ontario: rabies management implications, Seasonal home ranges of raccoons using a common feeding site in rural eastern, 65 Ontario, Significant Vascular Plant Records from the Hamil- ton Area, 612 Ontario, The Northern True Katydid, Prerophylla camellifolia (Orthoptera: Pseudophyllidae), at Ottawa, 124 Opegrapha sp., 411 gyrocarpa, 406 Operophtera bruceata, 539 Oplopanax, 500 horridum, 500 Orache, Alaskan, 568 Orchard, S.A., 72 Oreamnos americanus, 370 Oregon-grape, Tall, 176 Oreodytes laevis, 427 sanmarkii, 427 Oreta rosea, 537 Oriole, Baltimore, 232 Orobanche fasciculata, 559 Orthofidonia exornata, 537 Orthosia hibisci, 542 revicta, 542 segregata, 542 THE CANADIAN FIELD-NATURALIST Vol. 118 Oryx gazella, 185 Osprey, 267 Otisorex, 400 Ottawa Field-Naturalists’ Club Awards for 2003, The, 646 Otter, K.A., 354 Otter, River, 96,267 Sea, 270 Otter, Enhydra lutris, Sightings off Haida Gwaii/Queen Char- - | lotte Islands, British Columbia, 1972-2002, Sea, 270 } Ouellet, J.-P., 360 Ouitouche, 523 Ovenbird, 232 Ovis canadensis, 370 Owl, Barred, 215,273 Great Horned, 222,273 Owl, Strix varia in Alberta: Distribution and Status, The — | Barred, 215 | Owl, Strix varia, in Labrador, First Record of a Barred, 273 Oxytrope, Arctic, 574 Oxytropis arctica, 558 campestris ssp. varians, 559 Oxyura jamaicensis, 604 Oyster, Pacific, 77 Pachnobia imperita, 543 mixta, 543 Palm, R., 72 Palmaria palmata, 226 Palthis angulalis, 540. Pandemis canadana, 535 Pandion haliaetus, 267 Panic-grass, Yellow, 197 Panicum xanthophysum, 197 Panthera tigris, 185 Papaver croceum, 558 nudicaule ssp. nudicaule, 558 Papilio canadensis, 537 Papilionidae, 537 Paquet, P.C., 95 Parapoynx maculalis, 537 Parastichtis suspecta, 541 Pardosa lapidicina, 123 moesta, 122 sternalis, 122 Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) 1n Southeastern Idaho, Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders, 122 Pardosa sternalis (Araneae, Lycosidae) in Southeastern Idaho, Occurrence of Parasitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and, 122 Parectopa pennsylvaniella, 532 Parelaphostrongylus tenuis, 40 Pariseau, R., 521 Parker, G.H., 251 Parmelia saxatilis, 406 Parornix conspicuella, 532 | Parrya arctica, 560 | Parsnip, Cow 196 Partridge, S.T., 247 Passerina cyanea, 232 Patagonia, Observations of Interactions between Puma, Puma _ concolor, and Introduced European Red Deer, Cervus elaphus, in, 132 — ——S—= 2004 Patinopecten sp., 82 | caurinus, 78 Patterson, B.R., 278 Pediasia dorsipunctella, 537 | Pedicularis oederi, 559 _Pelonaia corrugata, 79 Penny, J.L., 174 #))), Penry, L.B., 201 | Pepper-grass, Common, 572 | Perca flavescens, 521 , Perchaude, 523 | Percidae, 521 | Percina caprodes, 523 | copelandi, 523 | Percopsidae, 523 « Percopsis omiscomaycus, 523 | Perisoreus canadensis, 273 | Perispasta caeculalis, 537 _Perizoma basaliata, 539 | Pero honestaria, 538 | Perognathus parvus, 300 Peromyscus spp., 342 leucopus, 299,342 | maniculatus, 299,342,402 | Peromyscus maniculatus, and Yellow-pine Chipmunks, Tamias | amoenus, in Old Field and Orchard Habitats, Popu- | lation Dynamics of Deer Mice, 299 | Perry, N.D., D.T. Stewart, EM. Madden, and T.J. Maier. New | Records for the Arctic Shrew, Sorex arcticus and the i] Newly Recognized Maritime Shrew, Sorex mariti- mensis, 400 | Peterkin, P., Review by, 641 | Petrel, Leach’s Storm, 258 | Petromyzontidae, 523 | Phacelia, Macbryde’s, 576 | Phacelia mollis, 559 Phalacrocorax auritus, 258 carbo, 258 j Phalaenophana pyramusalis, 540 Phalaenostola hanhami, 540 | Phaneta awemeana, 535 convergana, 535 parmatana, 535 Phenacomys intermedius, 438 intermedius intermedius, 438 ungava, 438 i Phenacomys, in Alaska, The Heather Vole, Genus, 438 Pheosia rimosa, 540 Pheucticus ludovicianus, 232 Phillips, F., 273 Phleum pratense, 253 Phlogophora periculosa, 542 Phlox hoodii, 559 Phlox, Moss, 575 Phlyctaenia coronata, 537 Phoca hispida, 395,602 Phragmatobia assimilans, 540 Phyllodesma americana, 539 Phyllodoce glanduliflora, 559 Phyllonorycter sp., 532 martiella, 532 Physcia caesia, 411 Physeter macrocephalus, 495 Picard, [., 441 Picea spp., 57,61 INDEX TO VOLUME 118 667 engelmannii, 191,584 glauca, 46,95,211,218,249,251,274,361,370,388,401, 504,530,590 glauca X engelmannii, 355 mariana, 62,95,268,274,370,388,504,590 rubens, 41,52,510 sitchensis, 499 Picoides pubescens, 358 Pieridae, 537 Pieris oleracea, 537 Pimephales notatus, 318 Pine, Eastern White, 251 Ground, 196 Jack, 95,196,253,388,595 Lodgepole, 58,62,191,355,370,499,561,584,595 Ponderosa, 198,300,383,595 Red, 196,251 White, 41,196,277 Whitebark, 584 Pine, Pinus banksiana, Growth in Manitoba, Effects of Lodge- pole Pine Dwarf Mistletoe, Arceuthobium america- num, on Jack, 595 Pinguicula villosa, 559 Pinus spp., 60 albicaulis, 584 banksiana, 95,196,253,388,595 contorta, 58,62,191,370,499,584,595 contorta ssp. latifolia, 559 contorta var. latifolia, 355 ponderosa, 198,300,383,595 resinosa, 196,251 strobus, 41,196,251,277 Pinus banksiana, Growth in Manitoba, Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, 595 Pipilo maculatus, 378 Piranga ludoviciana, 378 Piranga olivacea, 232 Pisaster ochraceus, 265 Pissodes strobi, 41 Pitcher, K., 270 Placopsis gelida, 424 lambii, 406 Plagiobothrys scouleri, 170 Plagodis alcoolaria, 538 phlogosaria, 538 pulveraria, 538 Plaintain, Seaside, 577 Plantago spp., 247 canescens, 559 maritima, 559 Plasmodium spp., 237 Platarctia parthenos, 540 Platichthys stellatus, 248 Platynota idaeusalis, 535 Platyphyllum concavum, 125 Platypolia anceps, 542 Plemyria georgii, 538 Pleurozium schreberi, 591 Plover, Piping, 444.448 Snowy, 445 Plover, Charadrius melodus, egg viability after seawater im- mersion, Piping, 448 Plover, Charadrius melodus, Nesting Sequential Polyandry in Piping, in Eastern Canada, 444 668 Plume-poppy, 614 Plusia putnami, 541 Plusiinae, 541 Plutella xylostella, 533 Plutellidae, 533 Poa spp., 300 annua, 560 compressa, 277 interior, 558 leptocoma, 559 nemoralis, 558 pratensis, 176 secunda, 559 Podiceps grisegena, 210 Podiceps grisegena) Populations in the Lower Matanuska- Susitna Valley, Alaska, Changes in Loon (Gavia spp.) and Red-necked Grebe, 210 Podistera macounii, 559 Poecile atricapillus, 232 hudsonicus, 48 rufescens, 378 Pohl, G.R., D.W. Langor, J.-F. Landry, and J.R. Spence. Moths and Butterflies (Lepidoptera) of the Boreal Mixed- wood Forest near Lac La Biche, Alberta, Including New Provincial Records, 530 Point Pelee Natural History News 3(4), 155 Polemonium acutiflorum forma lacteum, 558 boreale forma albiflorum, 558 Polia imbrifera, 542 nimbosa, 542 Polioptila caerulea, 232 Polix coloradella, 533 Polychidium muscicola, 424 Polychrysia esmerelda, 541 Polygonatum pubescens, 196 Polygonia satyrus, 537 Polygonum lapathifolium, 559 viviparum, 591 Polystichum kruckebergii, 164 lemmonii, 164 mohrioides, 164 scopulinum, 164 Polystichum lemmonii, a Threatened Fern in Canada, Con- servation Evaluation of Lemmon’s Holly Fern, 164 Polytrichadelphus lyellii, 421 Pomoxis nigromaculatus, 523 Pond-lily, Yellow, 569 Pondweed, 318 Curly, 318 Flatstemmed, 561 Sago, 561 Variable-leaved, 561 White-stemmed, 561 Popcornflower, Scouler’s, 170 Poplar, Balsam, 62,218,268,530 Populus spp., 52,57,61,511 balsamifera, 62,136,218,268,530 balsamifera ssp. trichocarpa, 355 deltoides, 309 grandidentata, 136 tremuloides, 46,62,96,136,211,218,251,268,274,277, 316,342,355,361,370,388,5 15,530,584 trichocarpa, 211 Porbeagle, 245 Porcupine, 607 THE CANADIAN FIELD-NATURALIST Vol. 118 Porina chlorotica, 421 pacifica, 406 Porpidea speirea, 415 carlottiana, 411 contraponenda, 406 speirea, 406 thomsonii, 406 Poszig, D., C.D. Apps, and A. Dibb. Predation on Two Mule » Deer, Odocoileus hemionus, by a Canada Lynx, Lynx : canadensis, in the Southern Canadian Rocky Moun- tains, 191 Potamogeton spp., 318,365 alpinus ssp. tenuifolius, 559 berchtoldii, 559 crispus, 318 gramineus, 559 pectinatus, 559 praelongus, 559 subsibiricus, 559 zosteriformis, 559 Potentilla arguta ssp. convallaria, 573 arguta var. convallaria, 559 Prairie-crocus, 569 Prest (1913-2003), A Tribute to Victor Kent, 626 Primrose, Evening, 253 Primula eximia, 559 tschuktschorum ssp. cairnesiana, 575 Prince Edward Island, A Mapping of the Present and Past Forest-types of, 504 Prince Edward Island, Use of Eelgrass, Zostera marina, Wrack by Three Species of Ladybird Beetles (Coleoptera: Coccinellidae) in, 225 Prionoxystus robiniae, 534 Procyon lotor, 65 Prodoxidae, 532 Propylea quatuordecimpunctata, 225 Proterorhinus marmoratus, 318 Protitame virginalis, 537 Protoboarmia porcelaria, 537 Protolampra rufipectus, 543 Protolithocolletis lathyri, 532 Protorthodes oviduca, 542 Protothaca staminea, 77 Pruitt, W.O., Jr., Review by, 638 Prunus sp., 136 pennsylvanica, 365 pumila, 196 serotina, 365 virginiana, 253,362 Pseudacris maculata, 136 Pseudexentera oregona, 536 Pseudohermonassa tenuicula, 543 Pseudosciaphila duplex, 535 Pseudothyatira cymatophoroides, 537 Pseudotsuga menziesii, 159,376,584 menziesii var. glauca, 355 menziesii var. menziesii, 599 Psilocarphus elatior, 169 tenellus var. tenellus, 169 Psilocarphus elatior, an Endangered Herb in Canada, Con- servation Evaluation of the Pacific Population of Tall Woolly-heads, 169 Psyllobora virginimaculata, 312 Ptarmigan, Willow, 602 Pteridium aquilinum, 196,251 ieee 2004 ' Pterophoridae, 536 Pterophylla, 125 camellifolia, 124 Pterophylla camellifolia (Orthoptera: Pseudophyllidae), at Ottawa, Ontario, The Northern True Katydid, 124 Ptychoramphus aleuticus, 616 | Puccinellia interior, 559 | Puffin, Atlantic, 257,604,615 Horned, 615 | Tufted, 615 | Pulsatilla ludoviciana, 559 Puma concolor, 132,605 concolor vancouverensis, 159 |Puma, 132 | Puma concolor, and Introduced European Red Deer, Cervus elaphus, in Patagonia, Observations of Interactions between Puma, 132 ‘Puma concolor, in Elk Island National Park, Alberta, First Record of Mountain Lions, 605 Puma concolor vancouverensis, Morphology and Population | Characteristics of Vancouver Island Cougars, 159 Puma, Puma concolor, and Introduced European Red Deer, Cervus elaphus, in Patagonia, Observations of Inter- | actions between, 132 | Punctelia stictica, 424 | Pyralidae, 536 ! Pyrausta borealis, 537 nicalis, 537 | Pyrenocollema halodytes, 410 | sublitorale, 424 Pyrola minor, 559 Quackgrass, 253 : Quail, California, 378 Québec, Extension de I’aire de distribution connue de la Mus- araigne fuligineuse, Sorex fumeus, dans le nord-est du, 44] Quebec, Forked Three-awned Grass, Aristida basiramea Engelm. ex Vasey: A New Addition to the Flora of, 276 Québec, Inventaire printanier d’une frayére multispécifique: Vichtyofaune des rapides de la riviére Gatineau, 521 Québec, Premiéres mentions de la Couleuvre mince, Tham- nophis sauritus septentrionalis, au, 135 Québec, Summer Diet of Two White-tailed Deer, Odocoileus virginianus, Populations Living at Low and High Den- sity in Southern, 360 —Quenouilles, 136 Quercus spp., 342,365 alba, 342 candicans, 383 gambellii, 383 garryana, 174,376 kelloggii, 383 laurina, 383 macrocarpa, 310,342 i obtusata, 383 robur, 125 rubra, 124,251,342 rugosa, 383 ' Queue 4 tache noire, 523 - Quillwort, Bristle-like, 560 Maritime, 560 INDEX TO VOLUME 118 669 Raccoon, 65 Raccoons using a common feeding site in rural eastern On- tario: rabies management implications, Seasonal home ranges of, 65 Radal, 132 Ragweed, Hybrid, 613 Ragwort, Dryland, 577 Rail, J.-F. and G. Chapdelaine. Fifteenth Census of Seabird Populations in the Sanctuaries of the North Shore of the Gulf of St. Lawrence, 1998-1999, 256 Rainette faux-grillon boréale, 136 Raisin, Northern Wild, 253 Ramalina siliquosa, 420 Rana aurora, 448 Randa, L.A. and J.A. Yunger. The Influence of Prey Avail- ability and Vegetation Characteristics on Scent Sta- tion Visitation Rates of Coyotes, Canis latrans, in a Heterogeneous Environment, 341 Rangifer tarandus, 39,119,192,241,362,602 tarandus caribou, 57,61,119,368 tarandus tarandus, 96,185 Rangifer tarandus caribou, in Saskatchewan, Morphology of Female Woodland Caribou, 119 Ranunculus aquatilis var. subrigidus, 559 occidentalis var. brevistylis, 559 Raphia frater, 541 Raptor, 306 Raseux-de-terre gris, 523 noir, 523 Raspberry, 251 Raum-Suryan, K., K. Pitcher, and R. Lamy. Sea Otter, Enhy- dra lutris, Sightings off Haida Gwaii/Queen Charlotte Islands, British Columbia, 1972-2002, 270 Raven, 273 Razorbill, 257 Redcedar, Western, 159 Redstart, American, 232 Regulus calendula, 48,86 satrapa, 48,86 Reindeer, 185 Reithrodontomys megalotis, 300 Renaud, C.B., 521 RENEW. 2004. Recovery of Nationally Endangered Wildlife in Canada. Annual Report (14), 644 Retinia burkeana, 535 Rettie, W.J. Morphoiogy of Female Woodland Caribou, Rangifer tarandus caribou, in Saskatchewan, 119 Rheum rhabarbarum, 560 Rhigognostis interrupta, 533 Rhinanathus minor ssp. borealis, 559 cataractae, 523 Rhizocarpon, 411 cinereovirens, 421 geminatum, 411 haidensis, 411 hensseniae, 408 hochstetteri, 411 maritimum, 421 Rhubarb, 568 Ribes, 500 oxyacanthoides ssp. oxyacanthoides, 559 Richardson, E.S. and R.K. Brook. Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus, 602 Rinodina gennari, 408 670 Rissa tridactyla, 258 Rivula propinqualis, 540 Rivulinae, 540 Roberge, B., 495 Robertson, R.J., 229 Robin, American, 381 Rock-Cress, Wall, 570 Rockweed, 406 Rorippa barbareifolia, 558 Rosa blanda, 253 woodsii, 559 Rosatte, R.C., 65 Rose, Smooth Wild, 253 Western, 574 Rosell, F., 434 Rothfels, C. Significant Vascular Plant Records from the Ham- ilton Area, Ontario, 612 Rubus spp., 251,365,500 ideaus, 197 spectabilis, 500 Ruggiero, L.F., 583 Rumex crispus, 560 Runck, A.M., 438 Rush, 400 Toad, 170,567 Russell, 1904-1998, A Tribute to Loris Shano, 451 Ryegrass, Perennial, 170,174 Sabia conica, 78 Sagebrush, 300 Sagittaria sp., 318 Salal, 159 Salamander, Cope’s, 448 Pacific Giant, 448 Salamandre sombre des montagnes, 136 Salix spp., 57,136,180,211,247,253,268,356,365,370,530 alaxensis ssp. longistylis, 559 amagdaloides, 309 glauca var. acutifolia, 559 novae-angliae, 567 pseudomyrsinites, 559 rotundifolia ssp. dodgeana, 559 Salmo clarki, 448 salar, 77 trutta, 113,523 Salmon, Atlantic, 77 Chum, 248 Pink, 442 Red, 249 Sockeye, 111,443 Salmon, Oncorhynchus gorbuscha, Spawning Runs, Gulls, Larus spp., Foraging at Pink, 442 Salmonidae, 523 Salsify, Yellow, 300 Salvelinus namaycush, 523 Sambucus, 501 racemosa, 500 racemosa ssp. pubens, 559 Samuel, W.M., 95 Sander spp., 521 canadensis, 523 vitreus, 523 Sander-Regier, R., Reviews by, 153,474,477,478,632 Sandlance, 262 Sandpiper, Spotted, 444 THE CANADIAN FIELD-NATURALIST Vol. 118: Sandwort, Alpine, 166 Sanicle, Pacific, 174 Sanicula crassicaulis var. crassicaulis, 174 Sargassum muticum, 77 Sarrothripinae, 540 Sarsasparilla, Wild, 46 Saskatchewan, Morphology of Female Woodland Caribou, , Rangifer tarandus caribou, in, 119 Saskatoon, 573 Saxidomus giganteus, 77 Saxifraga, 326 caespitosa, 559 eschscholtzii, 559 foliolosa, 327,559 gaspensis, 327 hieracifolia, 336 nelsoniana ssp. pacifica, 559 nivalis, 326 nivalis var. gaspensis, 327 nivalis var. labradorica, 327 nivalis var. rufopilosa, 327 nivalis var. tenuis, 326 rufopilosa, 327,559 tenuis, 326 tricuspidata, 559 Saxifrage, Prickly, 573 Scale, Woolly Beech, 41 Scallop, Weathervane, 82 Scaup, Greater, 268 Schinus patagonicus, 132 Schizoporella unicornis, 79 Schizura leptinoides, 540 unicornis, 540 Schmelzer, [. and F. Phillips. First Record of a Barred Owl, Strix varia, in Labrador, 273 Schmiegelow, F.K.A., 368 Schoen, J.W., 247 Sciaenidae, 523 Scirpus spp., 318 americanus, 226 cyperinus, 402 maritimus, 226 rollandii, 559 Scoparia biplagialis, 531 Scopula ancellata, 538 frigidaria, 538 junctaria, 538 limboundata, 538 Scoter, Surf, 264 Scoter, Melanitta perspicillata, in Coastal Southwestern British Columbia, Observations of Above-Surface Littoral Foraging in Two Sea Ducks, Barrow’s Gold- eneye, Bucephala islandica, and Surf, 264 Scouring-rush, Variegated, 591 Scymnus (Pullus) brullei, 312 Seal, Ringed, 395,602 Sealy, S.G., 85 Sears, R., 495 Sea Star, Ochre, 265 Seaweed, 77 Sargassum, 78 Seburn, C., Reviews by, 149,635 Seburn, D., Reviews by, 140,143,468,471 Sedge, 318,400 Blunt, 565 === Green-sheathed, 170 Houton’s, 197 Mud, 591 One-sided, 169 Ross’, 566 Slender-beaked, 169 Two-toned, 564 i Water, 591 Seiurus aurocapilla, 232 noveboracensis, 232 | Selasphorus rufus, 378 | Selenia alciphearia, 538 | Semibalanus spp., 406 | Semioscopis inornata, 533 Semotilus corporalis, 523 | Sendall, K.A., 72 Senecio eremophilus, 558 | Sergeant, D.E., Review by, 144 | Serviceberry, 196 | Setaria viridis, 558 | Setophaga ruticilla, 232 | Seymour, K.L. A Tribute to Loris Shano Russell, 1904-1998, | 451 | Shackleton, D.M., 159 | Shad, American, 77 | Gizzard, 318 | Sharpe. F., 434 | Shaverdo, H.V. and D.J. Giberson. Predaceous Water Beetles lected Along the Horton and Thelon Rivers in the (Coleoptera: Adephaga: Dytiscidae, Gyrinidae) Col- | Arctic Central Barrens of Canada, 425 | Sheep, Bighorn, 370 Shepherd, German, 247 | Shepherd’s-purse, 571 | Shiner, Emerald, 319 | Spottail, 318 | Shooting-star, Broad-leaved, 175 | Shrew, 111 | Arctic, 400 | Gaspé, 403 | Maritime, 400 ] \ Masked, 400 Northern Short-tailed, 402 Pygmy, 402 Smoky, 400 Tundra, 400 | Shrew, Sorex arcticus and the Newly Recognized Maritime Shrew, Sorex maritimensis, New Records for the Arctic, 400 || Shrew, Sorex maritimensis, New Records for the Arctic Shrew, Til Sorex arcticus and the Newly Recognized Maritime, 400 Shrews, Sorex spp., by Arctic Grayling, Thymallus arcticus, Consumption of, 111 Sicya macularia, 538 Silene involucrata ssp. tenella, 559 vulgaris, 560 | williamsii, 559 Silverside, Brook, 319 | Sinapis arvensis, 560 | Sinoe sp., 534 Siskin, Pine, 378 | Sisymbrium altissimum, 300 | Sitta canadensis, 384 Skunk, Striped, 40,69 INDEX TO VOLUME 118 671 Skunkweed, 169 Sloan, N.A., 405 Sloan, N.A. and P.M. Bartier. Introduced Marine Species in the Haida Gwaii (Queen Charlotte Islands) Region, British Columbia, 77 Slug, 388 Smartwood, Pale, 568 Smerinthus cerisyi, 540 Smith, C.M., Reviews by, 284,287 Smith, D.W., 115 Smith, T., Reviews by, 146,148,480,641 Smith, T.S., S.T. Partridge, and J.W. Schoen. Interactions of Brown Bears, Ursus arctos, and Gray Wolves, Canis lupus, at Katmai National Park and Preserve, Alas- ka, 247 Smith, T.W., G.W. Douglas and A.G. Harris. Conservation Evaluation of Small-flowered Lipocarpha, Lipocarpha micrantha (Cyperaceae), in Canada, 179 Snail, 78,388 Snake, Butler’s Garter, 435 Eastern Garter, 435 Garter, 448 Northern Ribbon, 435 Snowberry, Common, 175 Sobey, D.G. and W.M. Glen. A Mapping of the Present and Past Forest-types of Prince Edward Island, 504 Solidago spp., 253,365 altissima, 342 canadensis var. salebrosa, 560 Solomon’s Seal, Hairy, 196 Somateria mollissima, 258 Sonchus sp., 365 arvensis ssp. uliginosus, 560 Sorbus americana, 362 Sorex spp., 111,400 araneus, 113 arcticus, 400 arcticus arcticus, 402 arcticus laricorum, 402 arcticus maritimensis, 400 cinereus, 111,400,441 fumeus, 400,441 gaspensis, 403 hoyi, 112,402 maritimensis, 400 monticolus, 111,438 tundrensis, 112,400 yukonicus, 112 Sorex arcticus and the Newly Recognized Maritime Shrew, Sorex maritimensis, New Records for the Arctic Shrew, 400 Sorex fumeus, dans le nord-est du Québec, Extension de l’aire de distribution connue de la Musaraigne fuligineuse. 44] Sorex maritimensis, New Records for the Arctic Shrew, Sorex arcticus and the Newly Recognized Maritime Shrew, 400 Sorex spp., by Arctic Grayling, Thymallus arcticus, Con- sumption of Shrews, 111 Sorgastrum nutans, 342 South Dakota, Spring Dispersal Patterns of Red-winged Blackbirds, Agelaius phoeniceus, Staging in East- ern, 201 Sow-thistle, Perennial, 578 Soybean, 253 672 Spargania luctuata, 539 Sparganothis sp., 535 reticulatana, 535 xanthoides, 535 Sparrow, Chipping, 378 Song, 378 White-crowned, 378 Spartina spp., 400 alterniflora, 226 patens, 226 Spence, J.R., 530 Spermophilus spp., 90 beecheyi, 90 beldingi, 90 brunneus, 91 columbianus, 90 parryii, 91 richardsonii, 90 tridecemlineatus, 90 Spermophilus richardsonii, Multiple Mating Results in Mul- tiple Paternity in Richardson’s Ground Squirrels, 90 Sphagnum, 499 Sphingidae, 540 Spider, Wolf, 122 Spiders Pardosa moesta and Pardosa sternalis (Araneae, Lycosidae) in Southeastern Idaho, Occurrence of Par- asitoid Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf, 122 Spilonema revertens, 410 Spiraea latifolia, 253 Spiranthes romanzoffiana, 560 Spizella passerine, 378 Springbeauty, Alpine, 568 Springtail, 389 Spruce, Black, 57,61,95,268,274,370,388,504,590 Engelmann, 191,584 Norway, 124 Red, 41,52,514 Sitka, 499 White, 46,57,61,95,211,218,249,251,274,355,361,370, 388,401,504,530,590 Squid, 497 Squires, J.R., K.S. McKelvey, and L.F. Ruggiero. A Snow- tracking Protocol Used to Delineate Local Lynx, Lynx canadensis, Distributions, 583 Squirrel, Arctic Ground, 91 Belding’s Ground, 90 California Ground, 90 Columbian Ground, 90 Idaho Ground, 91 Red, 607 Richardson’s Ground, 90 Thirteen-lined Ground, 90 Squirrels, Spermophilus richardsonii, Multiple Mating Results in Multiple Paternity in Richardson’s Ground, 90 Starflower, 575 Starling, European, 378 Stellaria umbellata, 560 Stenolechia sp., 534 Sterna caspia, 258 hirundo, 258 paradisaea, 258 Sternotherus odoratus, 136 Sterrhinae, 538 Stethorus punctum, 312 THE CANADIAN FIELD-NATURALIST Stewart, D.T., 400 Stictotarsus griseostriatus, 430 Stigmella sp., 532 Stipa comata, 560 nelsonii ssp. dorei, 560 richardsonii, 560 speciosa var. major, 132 Stoffel, M.J., Review by, 284 Storeria dekayi, 136 Storm-Petrel, Leach’s, 259 Strawberry-blite, 568 Strepsimaninae, 540 Streptopus, 500 amplexifolius, 500 Strix varia, 215,273 Strix varia in Alberta: Distribution and Status, The Barred Owl, 215 | Strix varia, in Labrador, First Record of a Barred Owl, 273 Sturnus vulgaris, 378 Subularia aquatica ssp. americana, 560 Sucker, White, 319 Sullivan, D.S., 299 i Sullivan, T.P., D.S. Sullivan, and E.J. Hogue. Population Dy- namics of Deer Mice, Peromyscus maniculatus, an Yellow-pine Chipmunks, Jamias amoenus, in Old dl Field and Orchard Habitats, 299 | Swallow, Barn 230 i! Violet-green, 379 Swammerdamia caesiella, 532 Sweet, Meadow, 253 Sylvilagus, 345 floridanus, 342 Symphoricarpos albus, 175 Symphyotrichum laurentianum, 105 Symphyotrichum laurentianum, (Fernald) Nesom, Estimation } of Seed Bank and Seed Viability of the Gulf of Saint | Lawrence Aster, 105 Synaptomys borealis, 438 Synclita obliteralis, 537 Syncopacma sp., 534 Syndemis afflictana, 535 Syngrapha alias, 541 octoscripta, 541 rectangula, 541 viridisigma, 541 Takats Priestley, L. The Barred Owl, Strix varia in Alberta: Distribution and Status, 215 Tamarack, 63,370,515,590 Tamias amoenus, 299 Tamias amoenus, in Old Field and Orchard Habitats, Popu- lation Dynamics of Deer Mice, Peromyscus manicu- latus, and Yellow-pine Chipmunks, 299 Tamiasciurus hudsonicus, 607 Tanacetum vulgare, 560 Tanager, Scarlet, 232 Western, 378 Taniva albolineana, 535 Tansey, Common, 578 Taraxacum officinale, 365 Tardif, J.C., 595 Taxus canadensis, 41,365 Tea, Common Labrador, 591 Teal, Green-winged, 131 Teleiodes proximella, 534 IF 1 ek ITE 2004 SS Telfer, E.S. Continuing Environmental Change: An Example from Nova Scotia, 39 | Tephromela atra, 424 Tern, Arctic, 257 Caspian, 258 Common, 257 Tews, J. Hummock Vegetation at the Arctic Tree-line near i} Churchill, Manitoba, 590 Thamnophis spp., 448 | butleri, 435 sauritus, 135,435 | sauritus septentrionalis, 135 sirtalis, 435 sirtalis sirtalis, 135 Thamnophis sauritus septentrionalis, aa Québec, Premieres mentions de la Couleuvre mince, 135 | Thistle, Swamp, 253 Thlaspi arvense, 560 Thomas, L., 448 Thomsen, L.R., F. Sharpe, and F. Rosell. Collapsing Burrow | Causes Death of a Eurasian Beaver, Castor fiber, 434 Thrush, Swainson’s, 382 ! Wood, 232 | Thryomanes bewickii, 378 ‘f | Thuja occidentalis, 96,136,253,361,508 | plicata, 159 _Thymallus arcticus, 111 thymallus, 113 Thymallus arcticus, Consumption of Shrews, Sorex spp., by | Arctic Grayling, 111 Tick, American Dog, 42 Tiger, 185 Tilia americana, 253,310 _| Timmins, C.A., 318 | Timothy, 253 | Tineidae, 532 | Tinline, R.R., 65 | Todd, G., 90 Tofieldia coccinea, 560 | Tomenthypnum nitens, 591 Tortricidae, 534 | Tortricinae, 534 | Tortue musquée, 136 Totton, S.C., R.C. Rosatte, R.R. Tinline, and L.L. Bigler. Sea- sonal home ranges of raccoons using a common feed- ing site in rural eastern Ontario: rabies management implications, 65 | Touladi, 523 _Towhee, Spotted, 378 | Tracheoniscus rathkei, 388 i Tracy, S., 127 Tragopogon dubius, 300 | Trechus obtusus, 79 | Trichodezia albovittata, 539 | Trientalis europaea, 560 Trifolium spp., 253,362 dubium, 170 | pratense, 560 Triglochin spp., 247 maritima, 226 palustre, 560 Trillium sp., 365 Triphosa haesitata, 539 |. Triteleia grandiflora, 174 grandiflora var. howellii, 174 INDEX TO VOLUME 118 673 howellti, 174 Triteleia, Howell’s, 174 Triteleia howellii, an Endangered Lily in Canada, Conserva- tion Evaluation of Howell’s Triteleia, 174 Triteleia, Triteleia howellii, an Endangered Lily in Canada, Conservation Evaluation of Howell’s, 174 Troglodytes aedon, 378 Trout, Brown, 113 Cutthroat, 448 Truite brune, 523 Trypanosoma, 236 avium, 237 Tsuga spp., 499 canadensis, 41,251,361,508 heterophylla, 159 mertensiana, 159 Tubularia crocea, 78 Tumble-mustard, Tall, 300 Tunicate, 79 Turdus migratorius, 381 Turkey, Eastern Wild, 251 Turkey, Meleagris gallopavo silvestris, Behavior in Central Ontario During Winter, Wild, 251 Turnock, W.J. and I.L. Wise. Density and Survival of Lady Beetles (Coccinellidae) in Overwintering Sites in Manitoba*, 309 Turtle, Green, 72,579,611 Hawksbill, 580 Kemp’s Ridley, 580 Leatherback, 72,579 Loggerhead, 579 Murray Short-necked, 610 Snapping, 610 Turtle, Caretta caretta, and the First Record of a Green Tur- tle in Atlantic Canada, Hybridization between a Green Turtle, Chelonia mydas, and Loggerhead, 579 Turtle, Chelonia mydas, and Loggerhead Turtle, Caretta caret- ta, and the First Record of a Green Turtle in Atlantic Canada, Hybridization between a Green, 579 Turtle in Atlantic Canada, Hybridization between a Green Tur- tle, Chelonia mydas, and Loggerhead Turtle, Caretta caretta, and the First Record of a Green, 579 Turtles, Chelydra serpentina?, Is Cost of Locomotion the Reason for Prolonged Nesting Forays of Snapping. 610 Turtles in British Columbia waters: a reassessment, Status of marine, 72 Tylothallia biformigera, 424 Typha spp., 136,318 latifolia, 253,546,560 Udea itysalis, 537 Ulmus americana, 310,508 Underwood, T.J., 85 Untereiner, W.A., 90 Update on Quebec Amphibian and Reptile Atlas Project: Rana- Saura 6(2), 294 Uraniidae, 539 Uria aalge, 258 lomvia, 602 Urodidae, 536 Ursus spp., 443 americanus, 41.96,370 arctos, 117,134,239,247,370,499,603 maritimus, 395,602 674 Ursus arctos, and Gray Wolves, Canis lupus, at Katmai Nation- al Park and Preserve, Alaska, Interactions of Brown Bears, 247 Ursus arctos, in Southeastern Alaska, Seed Dispersal by Brown Bears, 499 Ursus arctos, in the Central Canadian Arctic, Movements of Subadult Male Grizzly Bears, 239 Ursus maritimus, Excavation of an Arctic Fox, Alopex lago- pus, den by a Polar Bear, 602 Ursus maritimus, in the Alaskan Beaufort, Chukchi, and Northern Bering Seas, Observations of Habitat Use by Polar Bears, 395 Urtica dioica ssp. gracilis, 560 Utricularia minor, 560 Vaccinium sp., 159,251,500 alaskense, 500 angustifolium, 195 globulare, 198 membranaceum, 560 myrtilloides, 195 ovalifolium, 500 vitis-idaea, 591 Vaccinium angustifolium, and Velvet Leaf Blueberry, Vac- cinium myrtilloides, Production in Eastern Ontario, Effect of Fire Intensity and Depth of Burn on Low- bush Blueberry, 195 Vaccinium myrtilloides, Production in Eastern Ontario, Effect of Fire Intensity and Depth of Burn on Lowbush Blue- berry, Vaccinium angustifolium, and Velvet Leaf Blue- berry, 195 Vahlodea atropurpurea, 560 Valerian, Sitka, 577 Valeriana sitchensis, 560 Vallisneria americana, 318 Veery, 232 Venerupis philippinarum, 78 Venusia cambrica, 539 pearsalli, 539 Verbascum thapsus, 300 Verbena bonariensis, 614 Vermivora celata, 381 chrysoptera, 232 peregrina, 48 Vernalgrass, Sweet, 177 Verrucaria sp., 406 amphibia, 411 degelii, 411 durietzii, 420 epimaura, 406 erichsenii, 410 halizoa, 424 maura, 408 mucosa, 410 sandstedei, 406 schofieldii, 406 silicicola, 424 striatula, 406 Vervain, Purpletop, 614 Vetch, American, 300,574 Common, 174 Viburnum, 500 cassinoides, 253 Vicia americana, 300,560 sativa, 174 THE CANADIAN FIELD-NATURALIST Vol. 118 Villemure, M. and H. Jolicoeur. First Confirmed Occurrence of a Wolf, Canis lupus, South of the St. Lawrence River in Over 100 Years, 608 Viola praemorsa ssp. praemorsa, 175 renifolia var. brainerdii, 560 Violet, Kidney-leaved, 574 Yellow Montane, 175 Vireo cassinii, 378 flavifrons, 232 gilvus, 232 olivaceus, 231 solitarius, 48 Vireo, Blue-headed, 48 Cassin’s, 378 Red-eyed, 231 Warbling, 232 Yellow-throated, 232 Vole, 342 Heather, 438 Long-tailed, 300 Meadow, 342,400 Montane, 300 Prairie, 342 Vole, Genus Phenacomys, in Alaska, The Heather, 438 Vulpes vulpes, 40,58,69,96,244,249 273,368 Wade, K.D., 95 Wall-creeper, 614 Warbler, Bay-breasted, 48 Black-and-white, 230 Black-throated Green, 48,232 Blackburnian, 48 Cerulean, 229 Chestnut-sided, 232 Golden-winged, 232 Magnolia, 48 Myrtle, 48 Orange-crowned, 381 Tennessee, 48 Townsend’s, 382 * Yellow, 86,232 Yellow-rumped, 48,86,232,378 Warbler, Dendroica cerulea Management on Eastern Ontario Bird Species, Predicting the Effects of Cerulean, 229 WARS 1983-2002 Special Annual Report Wildlife Accident Reporting and Mitigation, 645 Wart-cress, Lesser, 613 Washburn, B.E., 185 Wasp, Parasitoid, 122 Wasps, Baeus sp. and Gelis sp., in the Egg Sacs of the Wolf Spiders Pardosa moesta and Pardosa sternalis (Ara- neae, Lycosidae) in Southeastern Idaho, Occurrence of Parasitoid, 122 Water-hemlock, Spotted, 574 Watersnake, Northern, 435 Watersnake, Nerodia sipedon sipedon, Frequency of Tail Breakage of the Northern, 435 Waterthrush, Northern, 232 Waterweed, Canada, 318 Waxwing, Cedar, 232,378 Weasel, 96,306 Least, 403 Weathervane X Japanese Scallop Hybrid, 78 Weed, Willow, 568 2004 Weevil, White Pine, 41 Wetzel, S., 195 Whale, Dwarf Sperm, 495 Pygmy Sperm, 495 Sperm, 495 Whale, Kogia breviceps, in the Northern Gulf of St. Law- rence, Canada, Stranding of a Pygmy Sperm, 495 Wheatgrass, Crested, 300 Whip-poor-will, 230 Whitlow-grass, Wood, 572 Wilkerson, C.D., 267 Williston, P., Review by, 290 Willow, 57,211,247,253,268,356,370,530 Peachleaf, 309 Willson, M.F. and S.M. Gende. Seed Dispersal by Brown Bears, Ursus arctos, in Southeastern Alaska, 499 Willson, M.F. Gulls, Larus spp., Foraging at Pink Salmon, Oncorhynchus gorbuscha, Spawning Runs, 442 Wilson, S.F., A. Hahn, A. Gladders, K.M.L. Goh, and D.M. Shackleton. Morphology and Population Char- acteristics of Vancouver Island Cougars, Puma con- color vancouverensis, 159 Wintergreen, 196 Wise, I.L., 309 Wockia asperipunctella, 536 Wolf, 115,127,134,278,368,603,608 Eastern, 608 Gray [Grey], 42,57,61,95,194,247,350,361,608 Wolf, Canis lupus, Pack Density, Record High, 127 Wolf, Canis lupus, South of the St. Lawrence River in Over 100 Years, First Confirmed Occurrence of a, 608 Wolverine, 56,61 Wolverine, Gulo gulo luscus, Resting Sites and Caching Behavior in the Boreal Forest, 61 Wolverines, Gulo gulo luscus, in the boreal forest, Effects of mid-winter snow depth on stand selection by, 56 Wolves, Canis lupus, at Katmai National Park and Preserve, Alaska, Interactions of Brown Bears, Ursus arctos, and Gray, 247 Wolves, Canis lupus, in and near Pukaskwa National Park, Ontario, Demographic patterns and limitations of Grey, 95 Wolves, Canis lupus, in Relation to Forest Harvesting in West-central Alberta, Winter Habitat Use by, 368 Wolves, Canis lupus, Unusual Behavior by Bison, Bison bison, Toward Elk, Cervus elaphus, and, 115 Wood-Peewee, Eastern, 232 Woodlouse, 388 Woodpecker, Downy, 358 Woodsia alpina, 560 ilvensis, 560 Woodsia, Northern, 560 Rusty, 560 Woolly-heads, Slender, 169 Tall, 169 INDEX TO VOLUME 118 675 Woolly-heads, Psilocarphus elatior, an Endangered Herb in Canada, Conservation Evaluation of the Pacific Pop- ulation of Tall, 169 Wren, Bewick’s, 378 House, 378 Wright, J.D. and J. Ernst. Effects of mid-winter snow depth on stand selection by Wolverines, Gulo gulo luscus, in the boreal forest, 56 Wright, J.D. and J. Ernst. Wolverine, Gulo gulo luscus, Rest- ing Sites and Caching Behavior in the Boreal Forest, 61 Wright, K.G., 264 Xanthia tatago, 542 Xanthocephalus xanthocephalus, 207 Xanthorhoe abrasaria, 539 decoloraria, 539 ferrugata, 539 fossaria, 539 iduata, 539 lacustrata, 539 Xanthoria, 420 candelaria, 411 parietina, 420 Xanthotype sospeta, 538 Xenolechia aethiops, 534 Xestia normaniana, 543 smithii, 543 Xylena curvimacula, 542 Xylotype arcadia, 542 Yarrow, 166 Yellowthroat, Common, 86,232 Yew, Canada, 41 Yponomeutidae, 532 Ypsolopha canariella, 533 dentiferella, 533 Yukon Territory, New Records of Cyperaceae and Juncaceae from the, 266 Yukon Territory VI, New Records of Vascular Plants in the, 558 Yunger, J.A., 341 Zanclognatha lutalba, 540 Zapus hudsonius, 403 Zea mays, 253,365 Zeiraphera canadensis, 536 fortunana, 536 unfortunana, 536 Zenophleps alpinata, 539 Zonotrichia leucophrys, 378 Zophodia grossulariella, 536 Zostera marina, 225 Zostera marina, Wrack by Three Species of Ladybird Beetles (Coleoptera: Coccinellidae) in Prince Edward Island, Use of Eelgrass, 225 676 Index to Book Reviews Botany Berkutenko, A.N., H.G. Lumsden, and D. Lumsden. Flora and Climatic Conditions of the North Pacific: A Col- lection of Scientific Papers, 637 Beresford-Kroeger, D. Arboretum America, A Philosophy of the Forest, 474 Boulet, B. Les champignons des arbres de l’est de |’ Amerique du Nord, 146 Clark, L.J. Lewis Clarks’s Field Guide to Wild Flowers of the Sea Coast in the Pacific Northwest, 476 Clark, L.J. Wild Flowers of Field & Slope in the Pacific Northwest, 289,476 Clark, L.J. Wild Flowers of Forest & Woodland in the Pacific Northwest, 289,476 Clark, L.J. Wild Flowers of the Mountains in the Pacific Northwest, 289,476 DiGregorio, M.J. and J. Wallner. Cape Cod Wildflowers: A Vanishing Heritage, 475 Gleason, H.A. and A. Cronquist. Manual of Vascular Plants of Northeastern United States and Adjacent Canada: Second Edition, 147 Martin, N.D. and N.M. Martin. Biotic Forest Communities of Ontario, 146 Martin, P. The Wild Orchids of North America, North of Mex- ico, 147 Nicholls G. and R. Lupp. Alpine Plants of North America: An Encyclopedia of Mountain Flowers from the Rockies to Alaska, 474 Trelawny, J.G. Wild Flowers of the Yukon, Alaska & North- western Canada, 2nd Edition, 289 Waldron, G. Trees of the Carolinian Forest: A Guide to Species, Their Ecology and Uses, 148 Environment Boelens, B. and M. Watkins. Whose Bird? 150 Bryson, M.A. Visions of the Land: Science, Literature, and the American Environment from the Era of Exploration to the Age of Ecology, 478 Conner, J.K. and D.L. Hartl. A Primer of Ecological Genetics, 480 Czajkowski, C. Snowshoes & Spotted Dick; Letters from a Wilderness Dweller, 151 Dietrich, W. Natural Grace: The Charm, Wonder, & Lessons of Pacific Northwest Animals & Plants, 152 Frelich, L.E. Forest Dynamics and Disturbance Regimes: Stud- ies from Temperate Evergreen-Deciduous Forests, 150 Josephson Weddell, B. Conserving Living Resources in the Context of a Changing World, 477 Mitchell, B. Resource and Environmental Management in Canada: Addressing Conflict and Uncertainty, 640 Newell, J. The Russian Far East, 638 Suzuki, D. and Dressel, H. Good News for a Change: How Everyday People are Helping the Planet, 149 Wolfe, D.W. Tales from the Underground: A Natural History of Subterranean Life, 153 Young, A.G. and G.M. Clarke. Genetics, Demography and via- bility of Fragmented Populations, 478 Miscellaneous Burnett, J.A. A Passion for Wildlife: The History of the Cana- dian Wildlife Service, 290 THE CANADIAN FIELD-NATURALIST Vol. 118 Edlow, J.A. Bull’s Eye: Unraveling the Medical Mystery of Lyme Disease, 641 Jennings, J. The Canoe: A Living Tradition, 290 Kareiva, P. and S.A. Levin. The Importance of Species: Per- spective on Expendability and Triage, 641 Snetsinger, P. Birding on Borrowed Time, 291 Yochelson, E.L. Smithsonian Institution Secretary Charles Doolittle Walcott, 481 Zoology Attenborough, D. The Life of Mammals, 286 Bird, D.M. The Bird Almanac, 630 Bowman, T.D. Field Guide to Bird Nests and Eggs of Alaska’s Coastal Tundra, 632 Bumstead, P.E. Canadian Skin and Scales: A Complete Ency- clopedia of Canadian Amphibians and Reptiles, 467 Cannings, R. and S. Cannings. British Columbia: A Natural History. Revised and Updated, 631 Carroll, D. Self-Portrait With Turtles: A Memoir, 471 Cox, B. Conversations with an Eagle, 283 Crump, M. In Search of the Golden Frog, 282 del Hoyo, J., A. Elliott and D. Christie. Handbook of Birds of the World: Volume 9 Cotingas to Pipits and Wagtails, 631 Ducey, J.E. Birds of the Untamed West: The History of Bird- life in Nebraska, 1750 to 1975, 466 Duff, A. and A. Lawson. Mammals of the World: A Checklist, 634 Dunne, P. Pete Dune on Bird Watching: The How-to, Where- to and When-to of Birding, 473 Early, C. Sparrows and Finches of the Great Lakes and Eastern North America, 288 Early, C. Warblers of the Great Lakes and Eastern North Amer- ica, 288 Eisner, T. For Love of Insects, 468 Feldhamer, G.A., B.C. Thompson and J.A. Chapman. Wild Mammals of North America: Biology, Management, and Conservation (Second edition), 472 Ferrari, A. & A. Sharks, 282 Ferri, V. Turtles and Tortoises, 143 Heide-Jérgensen, M.P., O. Wiig, and D.G. Pike. Belugas in the North Atlantic and the Russian Arctic, 144 Heintzelman, D. All-Weather Hawk Watcher’s Field Journal, 471 Heintzelman, D.S. Guide to Hawk Watching in North Amer- ica, 632 Johnsgard, P.A. North American Owls, Biology and Natural History, Second Edition, 284 Johnston, D.W. The History of Ornithology in Virginia, 470 Jones, H.L. Birds of Belize, 140 Joynt, A. and M.G. Sullivan. Fish of Alberta, 281 Limbert, M. The Uses and Curation of Birds’ Egg Collec- tions: An Examination and Biography, 465 Linder, G., S.K. Krest, D.W. Sparling. Amphibian Decline: An Integrated Analysis of Multiple Stressor Effects, 138 Lockwood, J.A. Locust: The Devastating Rise and Mysterious Disappearance of the Insect that Shaped the American Frontier, 633 McCabe, R.E., B.W. O’ Gara, and H.M. Reeves. Prairie Ghost: Pronghorn and Human Interaction in Early America, 635 2004 McCulloch, N. A Guide to the Birds of St. Helena and Ascen- sion Island, 636 Melvin, E.F. and J.K. Parrish. Seabird Bycatch: Trends, Road- blocks, and Solutions, 285 Morton, M.L. The Mountain White-Crowned Sparrow: Migra- tion and Reproduction at High Altitude, 284 O'Toole, C. The Firefly Encyclopedia of Insects and Spiders, 145 Perrins, C. The Firefly Encyclopedia of Birds, 139 Pianka, E.R., L.J. Vitt. Lizards: Windows to the Evolution of Diversity, 141 Pough, F.H., R.M. Andrews, J.E. Cadle, M.L. Crump, A.H. Savitsky and D.D. Wells. Herpetology: Third Edition, 140 Richards, J.M., Y.T. Tymstra and A.W. White. Birds of Nuna- vut: A Checklist, 465 Rising, J.D. Geographic Variation in Size and Shape of Svan- nah Sparrows (Passerculus sanwichensis), 287 Sharpe, R.S., W.R. Silcock and J.G. Jorgensen. Birds of Neb- raska: Their Distribution and Temporal Occurrence, 466 INDEX TO VOLUME 118 677 Simpson, K. and N. Day. Birds of Australia (Seventh edition), 630 Sinclair, P.H., W.A. Nixon, C.D. Eckert and N.L. Hughes. Birds of the Yukon Territory, 288 Somma, L.A. Parental Behavior in Lepidosaurian and Tes- tudinian Reptiles: A Literature Survey, 635 Stewart, K.W. and D.A. Watkinson. The Freshwater Fishes of Manitoba, 469 Stuart, C. & T. Birds of Africa: From Seabirds to Seed-eaters, 138 Swarth, C.W., W.M. Roosebrug and E. Kiviat. Conservation and Ecology of Turtles of the Mid-Atlantic Region: A Symposium, 468 Waldbauer, G. What Good are Bugs? Insects in the Web of Life, 144 Wheeler, B. Raptors of Eastern North America, 143 Wheeler, B.K. Raptors of Western North America, 142 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 read- ers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. The journal also publishes natural history news and comment items if judged by the Editor to be of interest to readers and sub- scribers, 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 IRO Canada 678 | TABLE OF CONTENTS (concluded) Volume 118 Number 4 | Tributes | A tribute to Thomas Henry Manning 1911-1998 BRENDA CARTER | A tribute to Victor Kent Prest 1913-2003 A. S. Dyke, D. A. Hopcson, E. L. BOUSFIELD, R. E. BEDFORD ‘Book Reviews | ZOOLOGY: Birds of Australia (Seventh edition) — The Bird Almanac — Handbook of Birds of the World: Volume 9 Cotingas to Pipits and Wagtails — British Columbia: A Natural History Revised and Updated —A Field Guide to Bird Nests and Eggs of Alaska’s Coastal Tundra — Guide to Hawk Watching in North America — Locust: the Devastating Rise and Mysterious Disappearance of the Insect that shaped the American Frontier — Mammals of the World: A Checklist — Parental Behavior in Lepidosaurian and Testudinian Reptiles: A Literature Survey — Prairie Ghost: Pronghorn and Human Interaction in Early America — A Guide to the Birds of St. Helena and Ascension Island | BoTaNy: Flora and Climatic Conditions of the North Pacific: A Collection of Scientific Papers | ENVIRONMENT: The Russian Far East — Resource and Evironmental Management in Canada: Addressing Conflict and Uncertainty | MISCELLANEOUS: Bull’s Eye: Unraveling the Medical Mystery of Lyme Disease; The Importance of Species: | Perspective on Expendibility and Triage q | NEw TITLES News and Comment | RENEW 2004 Recovery of Nationally Endangered Wildlife in Canada Annual Report (14) — Calendar: | Endangered Reptiles of Canada 2004 and 2005 — Canadian Species at Risk November 2004 — Froglog: Newsletter of the Declining Amphibian Populations Task Force (65, 66) — The Boreal Dip Net/L’Epuisette Boreale: Newsletter of the Canadian Amphibian and Reptile Conservation Network/ Reseau Canadien de Conservation des Amphibiens et des Reptiles 9(1) December 2004 — Marine Turtle Newsletter (106) — WARS 1983-2002 Special Annual Report Wildlife Accident Reporting and Mitigation The Ottawa Field-Naturalists’ Club Awards for 2003 Errata The Canadian Field-Naturalist 118(3) | Index to Volume 118 COMPILED BY LESLIE CODY | Advice to Contributors | Mailing date of the previous issue 118(3): 21 October 2005 2004 618 626 630 637 638 641 643 644 646 649 650 678 THE CANADIAN FIELD-NATURALIST Volume 118 Number 4 2004) Articles Stranding of a Pygmy Sperm Whale, Kogia breviceps, in the northern Gulf of St. Lawrence, Canada LENA MEASURES, BENOIT ROBERGE, and RICHARD SEARS Seed dispersal by Brown Bears, Ursus arctos, in southeastern Alaska Mary F. WILLSON and Scott M. GENDE A mapping of the present and past forest-types of Prince Edward Island D. G. SoBey and W. M. GLEN Inventaire printanier d’une frayere multispécifique: ’ichtyofaune des rapides de la riviere Gatineau, Québec ANNIE COMTOIS, FRANCOIS CHAPLEAU, CLAUDE B. RENAUD, HENRI FOURNIER, BRENT CAMPBELL, and RICHARD PARISEAU Moths and butterflies (Lepidoptera) of the boreal mixedwood forest near Lac La Biche, Alberta, including new provincial records GREG R. POHL, DAvID W. LANGOR, JEAN-FRANCOIS LANDRY, and JOHN R. SPENCE Pilose Braya, Braya pilosa Hooker (Cruciferae, Brassicaceae), an enigmatic endemic of arctic Canada JAMES G. HARRIS New records of vascular plants in the Yukon Territory VI WILLIAM J. Copy, CATHERINE E. KENNEDY, BRUCE BENNETT, and PHIL CASWELL Hybridization between a Green Turtle, Chelonia mydas, and a Loggerhead Turtle, Caretta caretta, and the first record of a Green Turtle in Atlantic Canada MICHAEL C. JAMES, KATHLEEN MARTIN, and PETER H. DUTTON A snow-tracking protocol used to delineate local Lynx, Lynx canadensis, distributions JOHN R. SQUIRES, KEVIN S. MCKELVEY, and LEONARD F. RUGGIERO Hummock vegetation at the arctic tree-line near Churchill, Manitoba JORG TEWS Effects of Lodgepole Pine Dwarf Mistletoe, Arceuthobium americanum, on Jack Pine, Pinus banksiana, growth in Manitoba Brock Epp and JACQUES C. TARDIF Notes Excavation of an Arctic Fox, Alopex lagopus, den by a Polar Bear, Ursus maritimus EVAN RICHARDSON and RYAN K. BROOK Two Great Black-backed Gulls, Larus marinus, kill male Longtailed Duck, Clangula hyemalis MARTHA DowsLey and ANDREW CIRTWILL First record of Mountain Lions, Puma concolor, in Elk Island National Park, Alberta GLYNNIS A. Hoop and TIM NEUFELD First confirmed occurrence of a Wolf, Canis lupus, south of the St. Lawrence River in over 100 years Mario VILLEMURE and HELENE JOLICOEUR Is cost of locomotion the reason for prolonged nesting forays of Snapping Turtles, Chelydra serpentina? SHANE R. DE SOLLA and KIM J. FERNIE Significant vascular plant records from the Hamilton area, Ontario CARL ROTHFELS Do Rhinoceros Auklet, Cerorhinca monocerata, fledglings fly to the sea from their natal burrows JAMES L. HAYWARD and JERE K. CLAYBURN 49° 49¢ 50- 521 530 550 558 579 583 590) 595 602 604 605 608 61k 612 615 (continued on inside back cover) ISSN 0008-3550 ~< By “4 b = om) a5 w ES + =z tt he oO ne) wo Tf | I i Printed IUAIEIICITT I Ni sqeyan stat i < 7 nak . ates ‘ : ater ar te ae z fons gary AE cpa iasace® Madre foment pretend rere Pirie awacota OT renee SMES parernnerrrencs oe feveee at samen ne te é Feta Dacre apart ied coasts eSomanagtas Petey Lenqup yee aks ke SON ‘ > seas : ‘ f 3 sient 210 biepetverine : : : : : HAST ores ret i : : waaay santerseneertencsagn emesene! ef : C wt Saag pe ed altsTn Gh : v ne eg nny ae sas pei . 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