PPh eal fie a) erie 4 brane Mies! ry Hey aera Pere nT Ee a ee oa) aired aha Hat eN ANG Gn? fae BLEUE ND faye Mey ie en Wid aed TRIN epee Meet | PA ne Prony aen ts ne nad Ted peg siti ag GALEN POU aU Mats Pa eae a Sila Meg iain taer eit eka Cite ay tes Pee ge hk rtreenr Trane nesta rene athens PU dy ed ha Y an int Danneel it Abaya Rte tS meh r ike 2 i etait i Mas HARVARD UNIVERSITY e Library of the Museum of Comparative Zoology rn te ero he | The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Special Issue: A Life with Birds: Percy A. Taverner, Canadian Ornithologist, 1875-1947 JOHN L. CRANMER-BYNG Volume 110, Number 1 January-March 1996 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patron His Excellency The Right Honourable Roméo LeBlanc, P.C., C.C., C.M.M., C.D., Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Clarence Frankton Don E. McAllister Hugh M. Raup Irwin M. Brodo Claude E. Garton Stewart D. MacDonald Loris S. Russell William J. Cody W. Earl Godfrey Verna Ross McGiffin Douglas B.O. Savile Ellaine Dickson C. Stuart Houston Hue N. MacKenzie Pauline Snure William G. Dore George F. Ledingham Eugene G. Munroe Mary E. Stuart R. Yorke Edwards Thomas H. Manning Robert W. Nero Sheila Thomson Anthony J. Erskine 1996 Council President: David W. Moore Ronald E. Bedford Jeff Harrison p : : : Fenja Brodo Cendrine Huemer Vice-Presidents: Michael Murphy i Caine Naa INGER onne | William J. Cody Patricia Narraway If Recording Secretary: David Smythe Francis R. Cook Frank Pope 1 Corresponding Secretary: Eileen Evans Ellaine Dickson Tom Reeve i Treasurer: Gillian Marston oe Ces este Hoppe I 3 Alan German Chris Traynor Christine Hanrahan Ken Young Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box |) P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. For information on Club activities telephone (613) 722-3050. it 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. Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211 Copy Editor: Wanda J. Cook Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1JO Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9 Associate Editors: Robert R. Anderson Robert R. Campbell W. Earl Godfrey Warren B. Ballard Brian W. Coad William O. Pruitt, Jr. Charles D. Bird Anthony J. Erskine Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor with the exception of book reviews which should go directly to Book Review Editor. : Subscription rates for individuals are $23 per calendar year. Libraries and other institutions may subscribe at the rate of | $38 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $23 includes a subscription to The Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $5.00 to cover postage. | 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. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Date of this issue: January-March 1996 (April 1996). | Subscriptions and Membership | | Back Numbers and Index Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, | and The Ottawa Naturalist, 1887-1919, and Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager. Cover. John Macoun (left) and Percy Taverner (right) on a field outing, probably upriver of Chaudiére Falls, Ottawa, in | June 1911. The photographer may have been C. H. Young. (Reproduced courtesy of the Canadian Museum of | Nature, number J-5535). | THE CANADIAN FIELD-NATURALIST Volume 110 1996 THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA a eine os mere co i iy . Ve itty The Canadian Field-Naturalist Volume 110, Number 1 January—March 1996 A Life with Birds: | Percy A. Taverner, Canadian Ornithologist, 1875-1947 JOHN L. CRANMER-BYNG Professor Emeritus, Department of History, University of Toronto, Toronto, Ontario MSS 1A1 Correspondence address: 92 Greenbelt Crescent, Richmond Hill, Ontario L4C 5R8 Cranmer-Byng, John L. 1996. A life with birds: Percy A. Taverner, Canadian ornithologist, 1875-1947. Canadian Field- Naturalist 110(1): 1-254. From 1911 to 1942 Percy Taverner was Ornithologist at The National Museum of Natural Sciences in Ottawa, now called The Canadian Museum of Nature. He helped lay the foundations of scientific ornithology in Canada by building up the necessary collections of birds at the museum, and studying their distribution, working through a network of people who collected specimens and gathered ornithological information from across the country. He was a leading advocate of the need for conservation and wild bird protection, and played a major role, through his research and recommendations, in the creation of a national park at Point Pelee, and bird sanctuaries along the North Shore of the Gulf of St. Lawrence and at Bonaventure Island off the southern coast of the Gaspé Peninsula. Perhaps Taverner’s most far-reaching contribution was as an educator of public thought. His wide knowledge was passed to the public in his books, Birds of Eastern Canada (1919), Birds of Western Canada (1926) and Birds of Canada (1934). He presented scientific information about his birds in their many plumages and habitats, and at the same time conveyed to the reader his own sense of appreciation of the birds he was describing. In this way he helped to make the study of birds and their habitats popular recreation. He was an active council member of the Ottawa Field-Naturalists’ Club and an influential Associate Editor and frequent contributor to The Canadian Field-Naturalist. Throughout his life and career, his consistent devotion to the study of birds and their behaviour, his achievements and difficulties at the National Museum, his bird expeditions to different parts of Canada, and his family life, show a warm-hearted person with a sense of humour who was a tireless writer of letters to his many friends and col- leagues. Key Words: Percy Algernon Taverner, 1875-1947, ornithologist, National Museum of Canada, 1911-1942, Birds of Canada. Preface and Acknowledgements Sometimes, by chance, something happens quite unexpectedly to cause you to take a turn in a new the species of birds likely to be seen in Canada — P. A. Taverner Birds of Canada (1934). direction. This is what happened to me in May 1981 at the Federation of Ontario Naturalists’ Annual Meeting at the University of Western Ontario, London. When registering we were each given a book called W. E. Saunders — Naturalist. This con- tained several photographs, one of which was of W.E. Saunders talking to his old friends J. H. Fleming and P. A. Taverner, both well known ornithologists of their time. Fleming donated his large collections of birds and ornithological books to the Royal Ontario Museum in Toronto. Taverner held the post of ornithologist at the National Museum of Canada from 1911 until his retirement in 1942. Also, he wrote the first book about most of ©The Ottawa Field-Naturalists’ Club and John L. Cranmer-Byng. In the photograph Taverner appeared as a tall, rather distinguished looking man with a neatly trimmed beard and moustache. Something about him, and his cheerful looking friends, made me want to know more about him. The photograph showed that it was taken at the meeting of the American Ornithologists’ Union at Charleston, South Carolina in 1937 (see page 183). When I returned to Toronto I looked for a biog- raphy of Taverner, but I was disappointed, there was none. All I could find was a substantial “obit- uary” notice in The Auk of 1948, but this gave me enough information to confirm my resolve to find out more. From this point onwards I was commit- ted to Taverner the ornithologist, and Taverner the person. Early in the 1980s my own future was of immi- nent retirement, and ideas already in my mind began to take shape. Here was this ornithologist without a 2 THE CANADIAN FIELD-NATURALIST biography, and here was I, a historian at Toronto University soon to have time and energy to spend on a project of my choice. My professional training as a historian, and a long standing fascination with birds, persuaded me to join my two life interests and attempt to write a biography of P. A. Taverner. A preliminary search showed that there was indeed enough material for a biography, but it was not until I had made a survey of the mass of Taverner material in the National Museum, Ottawa, and the Taverner papers in the Royal Ontario Museum, Toronto, that I realized the wealth of detail they both contained. As time went by it became clear that my task was to write about more than the life of one Canadian ornithologist. It must include a look at events, personalities and achievements that were part of the development of ornithology in Canada in the first half of the twentieth century. What started as a pastime during the early years of retirement devel- oped into a challenge which has taken up the best part of ten years. Taverner wrote well in the mass of letters, articles and other items that he wrote. But he had one weak- ness — erratic spelling. The same was true of his friend J. H. Fleming. Rather than retain Taverner’s spelling mistakes and typing errors these have been corrected for the sake of clarity. It would have been beyond my abilities to write adequately about the ornithology in Taverner’s life had I not been fortunate to receive help and guidance from many ornithologists, professional and amateur, without whose encouragement I could not have com- pleted this work. I wish to thank a number of indi- viduals and institutions for invaluable help. W. Earl Godfrey, Henri Ouellet, Michel Gosselin and Stewart D. MacDonald at the National Museum of Natural Sciences, Ottawa (now the Canadian Museum of Nature) gave me much help and encour- agement, and made my visits to the ornithology department a pleasure. I am particularly grateful to Earl Godfrey for his insights into the career chal- lenges faced by his predecessor and invaluable advice in clarifying my attempts to realistically por- tray Taverner’s professional life. The staff of the National Archives of Canada, private manuscript and government record groups, gave me much help. Ross James, of the Royal Ontario Museum, Toronto Introduction The year 1926 was an interesting one for some ornithologists in North America. In that year the American Ornithologists’ Union (AOU) held its 44th annual meeting for the first time in Canada. The chairman of the committee in charge of local arrangements in Ottawa was Percy Taverner, Ornithologist at the National Museum of Canada. Vol. 110 (ROM), read my entire manuscript twice, and not only reviewed bird names but also offered helpful editorial comments. To him I am extremely indebt- ed. At the ROM I was given every assistance to work in the museum library and archive. I take this opportunity to sincerely thank Julia Matthews and staff of the library. Heather McCallum and staff of the Metropolitan Toronto Library, Theatre Department, Taverner Collection, were also helpful to my work. At the time when I began my research I came into contact and exchanged help with a number of others working on biographies of Canadian naturalists. In particular I wish to thank Marianne G. Ainley who was beginning a life of William Rowan, for the stim- ulating discussions we had on our research subjects; also W. A. Waiser who was writing a life of John Macoun, and Richard Mackie who was working on a biography of Hamilton Mack Laing. People who helped me to find information about Taverner’s home life were his stepson, Karel Wiest, with whom I had two meetings just before he died, Corwin Ferguson, a relative of Karel Wiest, and Marthe Kent (nee Wiest); family friends who gave me useful information were Jean York, Anne Whitmore, Barbara Reynolds, Stuart Jenness. Especially I wish to acknowledge the considerable information I have received about Taverner family life from Karin Lumsden (nee Porsild), and Elizabeth Lloyd whose help and encouragement I gladly acknowledge. People to whom I am grateful for the opportunity of discussing Taverner’s career: Daniel F. Brunton, Harry Lumsden, Martin K. McNicholl, and Robert B. Stewart. Other people whose assistance was of importance to me are: Lise Anglin, A. W. F. Banfield, Fred Bodsworth, H. Deichmann, C. Stuart Houston, Trevor Levere, Farley Mowat, and Angus H. Shortt. Special mention should be made of Terry Shortt and Louise de Kiriline Lawrence who gener- ously shared their memories of Taverner with me before they died. Lastly, to my wife Margaret, who has prepared my manuscript for publication, from start to finish, and who shares with me a love of the natural world. To her this book is dedicated with affection and admiration. For three years previously Taverner and his bird- artist friend Allan Brooks who lived in the Okanagan Valley, British Columbia, had been working on an illustrated book titled Birds of Western Canada. When the AOU accepted an invi- tation to hold its annual meeting for 1926 in Canada Taverner and Brooks planned that the print- 1996 ing of their book should coincide with the AOU meeting in Ottawa. The meeting began early in October with busi- ness sessions in the banquet room of the Chateau Laurier hotel. The presentation of scientific papers occupied three days and was held in the National Museum. In addition to papers there were several exhibitions of paintings, photographs and publica- tions. The catalogue of paintings included 440 items by about 50 artists. On registering, each member was presented with a copy of Taverner’s handsome- ly illustrated Birds of Western Canada, given cour- tesy of the Canadian Government as a souvenir of the occasion. The dinner, held at the Chateau Laurier Hotel, was attended by about 180 guests and was an occa- sion for fun, much of it thanks to the sharp mind of Percy Taverner. After the formal proceedings about 60 people went on an excursion by motor bus to the Gatineau Hills. On arrival the majority of the mem- bers walked to the top of King’s Mountain to enjoy views of the fall colours, while the remainder visited the grounds of the summer home of the Right Honourable W. L. Mackenzie King, Prime Minister of Canada. The AOU meeting was a prestigious event for Canada’s capital and the National Museum. For Taverner it presented a challenge which he faced with enthusiasm, humour and hard work. It gave him the pleasure of renewing his circle of ornithological contacts, and of knowing that Birds of Western Canada, his second book, would immediately reach an informed audience. It was a high point in his career. Taverner’s chief work at the museum was to build up a collection of specimens, birds mounted for dis- play in galleries, and skins arranged in drawers for research purposes. He started with about 5000 speci- mens which had been collected by John Macoun of the Geological Survey. Taverner made a number of field expeditions to various parts of Canada, includ- ing the St. Lawrence estuary, the prairie provinces and British Columbia. He also employed younger men such as Dewey Soper and Mack Laing to under- take collecting expeditions for the museum, one of which resulted in the discovery of the nesting grounds of the Blue Goose on Baffin Island by Soper in 1929. When Taverner retired in 1942 the collec- tion numbered 35 000 specimens including many series showing specific variations and geographic ranges of Canadian birds. It was this collection which served as the basis for Taverner’s books and many articles, and the range maps which he devised where his skill as a draftsman was seen to advantage. A major contribution to Canadian ornithology was his method for showing the distribution of birds across Canada. Large topographical maps in a binder were kept on a table where bird records were plotted. CRANMER-BYNG: A LIFE WITH BIRDS 3 The information from each was recorded on cards arranged by species, and this is still consulted by ornithologists today. Taverner also made an important contribution to the conservation of wildlife and its habitat through his involvement in the stuggle to protect migratory birds in North America. In 1915 he presented recom- mendations to the federal Commission of Conservation for the creation of national parks at Point Pelee, and at Percé Rock and Bonaventure Island off the Gaspé Peninsula. In support of the rec- ommendations he made a study of the stomach con- tents of the Double-crested Cormorants nesting on Percé Rock. He also made a study of the gannets nesting on Bonaventure, published in 1918, which contained a strong plea that the island should be pre- served permanently as a bird sanctuary. Percé Rock and Bonaventure cliffs were declared federal migra- tory bird sanctuaries a year later. In addition Taverner was one of several dedicated ornithologists who encouraged amateur collectors of birds’ eggs and skins to change their hobby and instead become students of the life histories and behaviour of birds. This led to the growth of the recreation of bird watching. Another far reaching contribution made by Taverner was in stimulating the general public towards an interest in birds and an understanding of how they live in the wild. He did this by a series of handbooks beginning with Birds of Eastern Canada 1919. Here he was ahead of his time in using “field marks” as an aid to telling one species from another in the field. He had the ability to combine scientific information with an appreciation of birds in their many plumages and habitats. Birds of Eastern Canada was so successful that Taverner was asked to produce a companion volume Birds of Western Canada 1926. Later he amalgamated the material in the two books and produced Birds of Canada 1934. The name P. A. Taverner was well known in Canada during most of his career because of his three books, and even after his death his name continued to be known for another 20 years or more until a com- pletely new book, The Birds of Canada by Earl Godfrey, was published by the museum in 1966. Since he was well known among naturalists across Canada for over 50 years, why has no biogra- phy been written about Percy Taverner until now? Near the end of his life an ornithological colleague at the Royal Ontario Museum, Ontario, suggested that Taverner should use his retirement to write his auto- biography. Taverner replied, “I do not feel that there has been enough of general interest in my life for an autobiography”. His judgment that the story of his life was not sufficiently interesting for a book may have been correct in the mid 1940s when the Second World War was coming to a climax. But now, 50 years later, when bird watching has become increas- 4 THE CANADIAN FIELD-NATURALIST ingly popular and when preservation of habitat and biodiversity have become issues of global concern, his biography should be of interest to a wide public. Taverner was one of the ornithologists of the 1920s and 1930s who played a major part in making Part I — Early Years CHAPTER 1. Percy Fowler A boy was born in the small town of Guelph in southern Ontario on 10 June 1875. His father, Edwin Fowler, was principal of a private school in which his mother, Emily Buckley, taught.! The child was christened Percy Algernon at St. George’s Anglican Church, Guelph.” Information on the background of Percy’s mother and father is meagre. Few of the details usually available to a biographer about the subject of his study are available in this case. For information on his first years there are only two sources based on family reminiscences. One is a typed, four-page “Biographical Outline” compiled by Taverner himself, in which he said nothing about his parents except that he was “of English born parentage on both sides”.° The other source is a nine- teen page “In Memoriam” notice written by W. L. McAtee, a longtime friend, printed in The Auk soon after Taverner’s death. According to this account, Percy’s mother was a survivor of the great Chicago fire of 1871.4 Some information can be gleaned from a few newspaper articles written about him and from a few snippets of information embedded in Taverner’s extensive ornithological correspondence which has been preserved. But almost no “family” letters appear to have survived. A feature article about Taverner that appeared in a Toronto newspaper in 1944 gave the information that his mother “had been a child in school in Chicago at the time of the fire, lost her home and entire family in the disaster but had stayed on at her school and even- tually taught there.”? Another scrap of information comes from a series of articles on Percy Taverner, his mother and his sister written for an Ottawa newspa- per in 1958, eleven years after his death. According to this source his mother was born in Manchester, England, and emigrated to Canada when very young.° As a result of this clue I was able to obtain a copy of her birth certificate from England, which gives her year of birth as 1854. Her father, George Buckley, was shown as a “book keeper” (clerk).’ Using this information I attempted to trace Emily Buckley’s family in Chicago but without any success. Chicago’s records of this period are incomplete and difficult to trace. Some were destroyed in the fire of 1871; thus there are no documents to corroborate the statement that Percy’s mother was orphaned during the Chicago fire, and no evidence to show that her family lived in Chicago. Vol. 110 the study and enjoyment of birds and their habits a popular recreation. In this he was in the first rank, and as a museum builder he was way ahead of the facilities available to him. Information on Percy’s father is equally vague. If Emily Buckley was at school in Chicago in 1871 but was in Guelph in 1875, how and why did she make the journey from Chicago to Guelph? One possible explanation is that Edwin Fowler was living in Chicago during this period, that he met Emily Buckley, and that together they went to Guelph. A search of the Chicago records seemed promising. According to the U.S. Census returns for Chicago in 1870 a man by the name of Edwin Fowler was liv- ing in Ward 12. This man was born in England, was twenty-eight years old, and had a wife, Mary Fowler, who was twenty-one years old and was also born in England. In the Chicago Street Directory for 1871 he was shown as a teacher at Dyhrenfurth College, a business school. In the 1873 Directory he was listed as superintendent of the college, and as boarding at 160 North Carpenter. The last mention of Edwin Fowler in the Chicago records was in the 1874 Directory, where he was shown as residing with. the ‘president: of: the ‘college, (Julius Dyhrenfurth, at 160 North Carpenter.® According to an account of some of the private schools in Guelph in the nineteenth century, a boarding school known as the Guelph Academy was sold to Edwin Fowler, B.A. This account states he had attended Oxford University.” My enquiry about Edwin Fowler addressed to the registrar of Oxford University received a negative reply. “I have searched our records for the nineteenth century but regret that I am unable to find that he was ever a member of, or obtained any degree from Oxford University.”'° The Guelph assessment rolls show that in 1874 Edwin Fowler took a three year lease of the Guelph Academy. The only Guelph city directo- ry that mentioned Fowler was for the year 1875-76. The entry reads: “Fowler, E. Principal Guelph Academy, 175 Woolwich St., h. same Fowler, Mrs. E. directress ladies department, Guelph Academy.” |! There is no record in either Chicago or in Guelph to prove that Edwin Fowler and Emily Buckley were married. But this means little. They might have been married in Chicago without any record of the marriage surviving. They could also have been married in Guelph without any record being pre- served. !” { 1996 An advertisement for Fowler’s academy, printed in the Wellington County Atlas of 1877, gives as principal Edwin Fowler B.A., and as directress Mrs. Edwin Fowler; it lists three members of staff by name. The school had facilities “for both sexes in Classics, Mathematics, Languages, English, Drawing, Painting etc. Boarders received and stu- dents prepared for College.”'!? Apparently Fowler’s Academy was well regarded.'* Hugh Douglas, writ- ing in 1960, stated that “Elderly ladies have told me that Mrs. Fowler deserted her husband, went to New York, and became a well known actress on the Broadway stage. Her stage name was Ida Van Cortland.”!> This was not mere gossip, though the reference to Broadway is an exaggeration. This account implies that Emily Fowler was to blame for deserting her husband and “going on the stage”. It is true that in 1877 Emily Fowler joined Mrs. Morrison’s Stock Company in Toronto where she first took a part as a member of the ballet.'° Not until near the end of Percy’s life was there an indication that anyone who had known his parents was still alive. In a letter to an old friend, Hamilton Mack Laing, Percy mentioned that he had received a letter from Mrs. Sharman. “Of course I never met her personally but she seems to be the only one I know of who knows anything of my CHAPTER 2. Percy Taverner When Emily Fowler came from Guelph to Toronto she found employment as a member of Charlotte Morrison’s stock company playing at the Grand Opera House.! At that time Toronto was a city of about 250 000, with several theatres which pro- vided a certain amount of cultural entertainment. Whether Emily had any friends in Toronto is not known but she was young, had a good figure and attractive looks, as well as considerable vivacity and intelligence. Under the stage name of Ida Van Cortland she soon began to play small parts. In a stock company novices learned how to act while on the job by watching rehearsals, and by studying the technique of visiting “stars” who played the leading roles. Her first chance to speak a few lines came in September 1877 in a production of As You Like It. By the end of the season Ida had played as many as forty different small parts.’ When Mrs. Morrison’s company disbanded in the summer of 1878 owing to financial problems Ida went, with a letter of introduction, to seek work in New York. She became a member of William Nannary’s company on a tour to Atlantic Canada in January 1879, and began to receive favourable press notices.? In September a young actor called Albert Tavernier came from New York to join the com- pany. He was the same age as Ida and had started his CRANMER-BYNG: A LIFE WITH BIRDS 5) early days and forbears. She must be a remarkable per- sonage, at her age to retain such interests and clear fac- ulties. Her writing and letters are those of a young woman. Strange too how different life lines cross. Not only did she know my mother as a young woman but also is closely connected with other people who fortu- itously had a great influence on my life.”!7 Soon after Percy Taverner died, Mrs. Sharman wrote to Mack Laing mentioning his earliest years. In doing so she let slip the only information about his parents that now exists: “... the recollections of myself a twelve year old girl pupil — flashes of memory — of charming people who were unsuited to one another — seem better left with Mrs. Taverner, who could if she thought best, add what she pleased.”!® But Taverner’s widow left no information on his earliest years. It seemed to have been a closed sub- ject to his mother, his half sister and his wife. Percy Taverner had an unpromising start in life — parents who parted when he was two years old, a father whom he never saw again, and a mother who, at the age of twenty-three, started a career on the stage in order to support herself and her infant son. But Ida Van Cortland, as she now called herself, had intelli- gence and character, qualities that were to sustain her in the ups and downs of life which she and her son had to face. This is the subject of the next chapter. own career in the theatre at the same time as her, but in Cleveland, Ohio. His father, Joseph William Taverner, was a well known teacher of elocution and dramatics who coached several famous public speak- ers. His son, therefore, had a good start for the the- atrical profession.* Albert was handsome and per- sonable. Ida and Albert fell in love: At the end of the year the Nannary Company disbanded and they both returned to New York to find employment. From August 1880 to April 1881 Ida toured in the northern and southern States while Albert acted in various New York theatres. In June 1881 Albert Tavernier and Ida Van Cortland were married.° Following their marriage they toured for a season with W. J. Florence and his wife in The Mighty Dollar, and a few other plays. Florence’s perfor- mance as an ignorant and self-seeking politician in The Mighty Dollar was an excellent character study which Albert adopted later as a model for his own interpretation of the part.° In 1883 Albert and Ida decided that the organizational side of the theatre was changing so rapidly that touring companies had now supplanted resident companies, and that the future lay with touring companies presenting a few well chosen and well presented plays. They made the bold decision to set up their own company, with Ida as the leading lady and Albert as actor-manager.’ Meanwhile, little Percy had an unsettled child- hood, travelling from place to place with his mother and learning his first lessons from her. When she married again Percy was informally adopted by his stepfather and exchanged the name Fowler for that of Tavernier.® His earliest connected memories were of a private kindergarten school at Highland Falls, near West Point, New York where he boarded with a few other children for one fall, winter and spring. These memories concerned natural objects such as the first spring flowers, watching orioles at their nest in a tree in the front yard, and monthly numbers of a maga- zine in which there was a story about a naturalist and a Scarlet Tanager with a picture of the bird.” It is likely that Percy attended this school from the fall of 1881, when his mother and stepfather left New York to tour with the Florences’ company, until their return in the spring of 1882. This would be during Percy’s sixth year since his seventh birthday occurred in June 1882. The earliest photograph of Percy to have survived shows the head and shoulders of a small boy of about six years old, his hair neatly brushed, wearing a smart polka-dot shirt and a matching tie. This photograph might have been taken for his mother to have with her while on tour.'° The Taverniers’ big venture began in August 1883 with their own company, the New York Comedy Company.!'! Each season the Company would start in the fall on an itinerary which would last until some time in the following summer. One season they toured the Atlantic provinces and states; another season they would tour mainly in southwest- ern Ontario and the Great Lakes.'* The visit of a touring company, especially to towns in rural areas, was a local event. Members of the Company brought with them a touch of a wider world, a hint of ele- gance and culture, and on the stage the drama of pas- sion, of anger, of sorrow. Colourful costumes, music and well spoken words roused members of the audi- ence from the routine of their narrow lives. The local newspapers reported the performances of visiting companies as did the Dramatic Mirror. On the whole the Taverniers’ Company received favourable notices. Albert and Ida were seriously involved in their profession; they wished to provide quality entertainment, well acted, well spoken and produc- tions that were well dressed. They consciously aided this image by attending church on Sundays, giving benefit performances for local causes, attending civic functions and even producing plays written by local amateurs. In this way they aimed to establish a “home audience” in the towns they visited so that their annual appearance was an event for their audi- ence to look forward to.'? Percy’s earliest memories were of travelling on circuit with the company — of temporary residences with a theatrical family in Halifax, with a family in Brooklyn, and with a min- ister’s family by the Erie Canal.!4 THE CANADIAN FIELD-NATURALIST Vol. 110 By the summer of 1885 Percy had turned ten years old and his mother sent him to a private school in Port Huron, Michigan. Here he boarded with a Mr. and Mrs. C. F. Smith and their children who treated him as one of their own.!> But however kind- ly he was treated it would have been impossible to make up for not having a father and the lack of a permanent home with brothers and sisters and cousins of his own. However, this was soon to be remedied. Albert’s father, Joseph William Taverner, the pro- fessor of elocution, died intestate in Jamaica in 1885. Among his assets was an island in Lake Muskoka opposite the holiday resort of Beaumaris.!° Since he left no will, possession of the island would have to be divided equally between his two sons, Henry W. of San Francisco, and Joseph E. of Mount Vernon, N. Y., and his five married daughters. Rather than do this his heirs worked out a family agreement by which each one relinquished their share in favour of Ida Tavernier in return for five hundred dollars Canadian money to be paid by her. No family papers have survived to explain why the island was sold to Ida only, rather than to Albert and Ida jointly. All that exists is the original deed of sale.'’ Albert and Ida with Percy first visited the cottage on the island for one night in the summer of 1886.!° At least Percy could now look forward to summer holidays at a cot- tage on a lake, and this should have given him a feel- ing of belonging somewhere. Another change occurred in Percy’s life when his mother withdrew from the Company’s tour early in 1887 due to “an approaching domestic event” as the Dramatic Mirror delicately phrased it. In June 1887 she gave birth to a daughter in the town of Markham north of Toronto.'!? Percy now had a half-sister, Ida, 12 years younger than himself, with whom to share family life, especially in the summer holidays. Presumably Percy’s mother went to Markham for the birth of her child because her sister-in-law, Florence Knill, with her husband Edwin G. Knill who was a physician, lived there. They had a young family of their own, and some years later bought a one acre lot on Gibraltar Island.”° Ida grew up with first cousins who came to stay near to her family in the summer. During the late 1880s the Tavernier Company was becoming well known and drawing reasonably good audiences and receiving good notices. The the- atre business was growing at this period owing to an expanding population and the growth of cities. Communications were improving with the network of new railway lines that was being laid out. All this made for a greater demand for theatrical perfor- mances, which in turn caused an increase of compe- tition. The Taverniers managed to keep abreast of this by playing the lucrative Great Lakes circuit, especially in the cities where they were always well CRANMER-BYNG: A LIFE WITH BIRDS Taverner family photo taken at Ann Arbor, Michigan, 1889. Percy age 13, Ida Clare age 2 (Courtesy of the Metropolitan Toronto Reference Library, original in the collection of Murray Edwards 8 THE CANADIAN FIELD-NATURALIST received such as Hamilton and Toronto.”! Although in the late 1880s they were doing well in the public’s estimation artistically, the account books show that financially they were not all that prosperous. Their finances were not improved when Albert Tavernier indulged his ambition to be an actor-manager by tak- ing a five-year lease on the Hibbard Opera House in Jackson, Michigan. Apparently this was an unprof- itable venture and he had to give up the lease after only one year.” In the summer of 1889 Percy turned fourteen and it was time for him to go to high school. His mother and stepfather must have given some thought to his future education and a possible career. One thing was certain — with his stammer he would never want to become an actor.” It was decided that he should prepare for entry into Ann Arbor High School, probably because it was recognized at that time as the leading high school in the state of Michigan. It was established in 1856 largely to pre- pare students in Ann Arbor for the University of Michigan. The relation between Ann Arbor High School and the university was close, and a number of university teachers also taught at the school.” Many students from outside the city enrolled in order to take advantage of its connection with the university. Board and lodging with families in the town was available and Percy lived with the family of Wallace Bliss. A portrait in sepia of the Tavernier family taken by a photographer of Ann Arbor in 1889 exists showing his mother on one side of a chesterfield, his stepfather on the other side and little Ida Clare, aged two, in the middle. Percy is standing behind, his hair cut short, looking straight at the camera with his eyes fully open and his mouth a fraction open. It is a non-committal look, as though he may be about to break into a smile, or that he may not trust the photographer.”> Percy attended grade eight in Ann Arbor for one year before entering the High School in the fall of 1890. Ann Arbor High was an imposing imitation Gothic building of undressed stone and brick with turreted slate roofs which blended well with the “academic” Gothic architecture of Michigan University in the 1890s.7° We know nothing about his time at high school either from Percy or from his family, and can only adduce a little information from the Ann Arbor High School “Catalogues”.?’ These show that Percy A. Tavernier was enrolled in the First Year program in 1890-91, and that the student enrolment for that year was 200. He attended the Second Year program in 1891-92 and the Third Year in 1892-93. The cata- logue for that year showed that he was in the English course. Students in this program would be required to take five science courses, two of which must be in botany and physics. The other three courses could be chosen from: astronomy, chemistry, physiology or another full course in physics. What was not offered Vol. 110 was biology, a subject that would have been very useful to Percy in the future had he been able to study it. In this program two years of Latin, French or German could be taken instead of Physiology, Chemistry, Astronomy, and English History. French would have been useful to him in later life if he had chosen to take it at Ann Arbor High. Other subjects included were: Arithmetic, Algebra, Geometry, English Literature, Old English, Grammar, History. Programs required four years of study in order to obtain a diploma that would admit students into the University of Michigan. One major problem Percy would have faced in high school was that much of the class work was heard orally in daily recitation periods. In this he would have been at a disadvan- tage. Moreover, in his third year he would have been required to take “rhetoric”. If this meant “skill in the effective use of speech” and required practice in “verbal communication” then Percy would have been at a double disadvantage. The school encour- aged participation in various extra-curricular activi- ties such as those of the literary society in “speaking, discussion, and parliamentary practice”, as well as a society for debating and dramatics. Ironically none of these activities would have been ones in which Percy could have participated satisfactorily, except perhaps as a stage hand. There is no evidence that Percy took any part in football or athletics though there is plenty of evi- dence from the annual year book of that period, The Omega, that students took part in sports of various kinds. In fact no records appear to have survived concerning Percy’s school days at Ann Arbor High. It was almost as if there was nothing he wished to record or remember from that period. We can only guess that because of his stammer he may have felt “out of things”. One of the rules laid down by the school board stated that “Hazing, rushing, pumping, concerted riots, and disorderly conduct on the street, or on the school or public grounds shall subject pupils to suspension; and certificates of graduation may be withheld from any pupil ... engaging in such practices.”?® A boy such as Percy was likely to have been subject to some form of ridicule simply because his speech sounded comic. This may have . had the effect of turning him into a “loner” at that time.”? Luckily young Taverner took up a recreation that he could enjoy in a quiet, uncompetitive way. What exact year he became interested in the study of birds is not clear but it was certainly before May 1893 when he began to keep regular bird notes. He proba- bly started by collecting birds’ eggs as most boys did in that period. One day, just by chance, he met a man on the street who happened to be A. B. Covert the taxidermist of the University of Michigan Museum. When Covert realized that the boy was seriously interested in birds he invited him to come and see 1996 what went on behind the scenes in the zoology sec- tion of the Museum. Here he met two other young men with the same interest — George Prey and Robert Walcott, who were constantly in the field together collecting birds for study. This was proba- bly in 1892 when the Museum was busy making a Michigan natural history exhibit for the World’s Columbian Exhibition to be held at Chicago in 1893. The scene inside the Museum was “entrancing” and Percy gradually became interested in the more seri- ous side of bird study.*° Covert taught him the rudi- ments of taxidermy, and Percy became a constant visitor at the Museum and an unofficial, unpaid assistant. In this way he began to develop a system- atic knowledge of birds and an appreciation of their qualities. On weekends and holidays he was out- doors birding with two or three others with the result that his spare time was happily filled and he got enough exercise by tramping the countryside around Ann Arbor. By the time he began keeping a note- book he had already gained experience through col- lecting birds and he knew that his ambition was to be a student of ornithology. Although he never studied ornithology at the University he did study it unoffi- cially behind the scenes in the university museum. Percy Taverner approached his chosen recreation in the spring of 1893 with seriousness as can be seen from the “Preface” he wrote for his first book of bird notes. This, he explained, was to be a daily record of his collecting trips with anything he felt worth recording such as the weather, early spring arrivals, early eggs, and rare birds identified. In a second book he proposed to list the birds he saw by species showing year and place together with notes taken from his first book on what he had observed of their behaviour. In a third book he intended to keep statis- tics such as arrival and departure dates on migration, length of incubation of eggs, and information on what food a bird had been eating after he had dis- sected it. As a result he hoped to classify all his notes in a way that would make it easy to find any- thing that he might want. His notes would be useful for his own education. He began book | with some cautions that he intended to carry out. These were safety precautions about handling a gun in the field “that I have learned by experience.”*! But first he cautioned himself about his purpose. “Never forget you are a student, not a mere ‘collector’.” Then fol- lowed twelve points about hunting birds with a gun such as “never fool with a gun whether it is loaded or not; never point your gun at any living creature unless you want to kill it; learn to handle a gun care- fully from habit.” Someone at the Museum had trained him well. But in addition he was well aware that birds are living organisms not just museum specimens that have been killed for identification purposes. “Remember ornithology is not a dry study it is full of life and poetry and beauty”, and he CRANMER-BYNG: A LIFE WITH BIRDS 9 advised that one should study a newly killed bird as it lay in one’s hand before it is skinned. One should try to imagine it in its native tree tops. A quarter of an hour of this would give one a truer idea of the bird’s build than a week of measure- ments. This was revealing of Percy’s attitude towards birds and their study one month before his eighteenth birthday.” The first entry in his bird notes was made on 7 May 1893 walking down the Ann Arbor river when he recorded: White-throated Sparrows; Maryland [Common] Yellowthroat; Chewink [Rufous-sided Towhee]; Yellow Warbler; [Eastern] Kingbird; Rose-breasted Grosbeak; Carolina Rail [Sora]; Long-billed [Louisiana] Waterthrush; stake driver [American Bittern]; Florida Gallinule [Common Mcorhen].** Two days later he recorded a Scarlet Tanager, a Red-headed Woodpecker, a Blue-gray Gnatcatcher and some Bobolinks. From this time until 1910 Percy would spend much of his spare time walking the fields and marshes, forests and shores observing, collecting and recording. A week later he was out with Covert and Walcott when an uncom- mon bird was collected which was identified as a Yellow-throated Warbler. Although he was able to examine it closely in the Museum he missed the great satisfaction of identifying an uncommon bird alive in the wild.** At Four-mile Lake near Dexter in late June he recorded six nests of Least Bittern, one of which had a set of six eggs. Meanwhile his moth- er and stepfather were on Gibraltar Island for a week in May 1893 preparing for a three-month summer season at the Silurian Casino in Waukesha County, Wisconsin, the site of a health spa.*> Percy’s bird records show that he probably visited them at the spa in July 1893 when he recorded seeing a Pied-billed Grebe at Grass Lake in Waukesha County. From early August until mid-November he was staying on Gibraltar Island, and recorded shooting partridge [Ruffed Grouse] from time to time. By now Percy had completed three years at Ann Arbor High and had turned eighteen in June 1893. But his movements during the school year 1893-94 are not clear. He remained at Beaumaris until mid- November which may indicate that he returned to Ann Arbor late for some reason.*° According to his bird notes he was at Ann Arbor in the first half of 1894. He described the nuptial display flight of a male Marsh Hawk [Northern Harrier] observed on 25 April which shows that his powers of observation were sharp and that already he had developed a clear style in writing English with a pleasant rhythm to it. “He would fly along about twenty feet from the ground and then suddenly drop down, turning over and round, sometimes on his back and sometimes on his side, until within five feet of the ground when he would rise again and do it over again. All this while uttering a low gurgling note. After a while he sailed 10 THE CANADIAN FIELD-NATURALIST off and was joined by his mate.”*’ In mid June he was birding with Covert and Walcott southward along the railway track in Ann Arbor and found a nest of young Bobolinks; a male Henslow’s Sparrow, adult and young Chipping Sparrows and Yellow-breasted Chats. According to the log-book at Camp Coo-ee Percy was there from July into November, while Albert Tavernier’s sisters Clara from California and Daisy from Detroit occupied the house. The Knill and Power families also visited during the summer. Altogether there were four aunts, two uncles, and three cousins of Ida Clare’s at the island during the summer. After the last relative had left Percy stayed at the Prowse’s who owned the Beaumaris Hotel. In early November he went to live in Guelph where his stepfather had taken a two-year lease of the newly opened Royal Opera House as manager.*® Percy was office boy, since he recorded in his notebook: “At five p.m. I closed the box office and taking a lunch for supper went up the Grand River until dark.” He made several trips down the local rivers and was out with his gun collecting birds on most days in May. From Guelph he went to Beaumaris where he spent the summer of 1895 with his mother, stepfather and stepsister Ida, now aged eight, and members of the Company. One of them had his mandolin with him and another had his guitar. On 28 June the Company played The Mighty Dollar at Bracebridge for the benefit of the orchestra while on II June they played Forget-me-not at Rosseau. They then went to Parry Sound, twenty-four miles by stage coach, to play there for a week.*° In July Mrs. Smith, with whom Percy had boarded at Port Huron, arrived to stay and they celebrated the “glorious fourth” with a picnic at Bala. “Had big time & got towed back by the supply boat and ended up the day by two immense bonfires. The Stars & Stripes very much in evidence.” Another time Ida Clare and Mrs. Smith went with neighbours, the Fearmans, in their steam yacht “Tris” to Judge Mahaffy’s. Someone held a “bee” and got his verandah up while the people at Solid Comfort Camp gave their “at home and annual jollification” which they all attended. In August there was a fancy dress “hop” at the Hotel and everyone at the camp went over in costume. Albert went as Bardwell Slote, a character in The Mighty Dollar.*! The summer of 1895 should have been a memo- rable one for Percy. He had a gun and a rowboat for recreation, plenty of social life, and no further anxi- eties about school. Whether or not he had any ambi- tion to study at Michigan University is not known.” All we know is that Percy was a young man with no career in view. However, a temporary job was easy to find. When the Tavernier Company took the road for the 1895-96 season Percy accompanied it as assistant property man at $5.00 per week. Little Ida Tavernier was listed as one of the “artistes” and Vol. 110 received $1.00 per performance. This season Ida Van Cortland was probably managing her own com- pany which would have put a heavy load on her shoulders.*° The Company started the tour at Barrie in mid- September, was in Guelph for a week, then went to Ottawa followed by Sault Ste Marie, followed by Ashland (Wisconsin) and Brainerd (Minnesota), and then Winnipeg which was their furthest stop west. This was quite a challenge for Percy, not because of his job, but because of the continually changing opportunities for observing birds. His ornithological notes expanded considerably and while the others rested or rehearsed Percy went for a walk or “took a tramp”. At Ashland he recognized an Arctic Three- toed Woodpecker [Black-backed Woodpecker], but did not recognize a sparrow with a suggestion of a crest. He made a passable watercolour sketch of it and considered it a Lincoln’s Sparrow. At Brainerd he noted that the Ruffed Grouse on sale in the mar- ket were all of the grey variety. In Muskoka there were both red and grey colours. He shot two “sand- pipers” which he could not identify. All he had to identify them with was Afsgar’s pocket key and could only guess that they might be Robin Snipe [Red Knot]. At Winnipeg the first thing he did on arrival was to go to the shop of A. Hine and Sons, taxidermist, and get the local bird news. One thing he learned was that Whooping Cranes were reported to be common but so wary that they were hard to shoot. He saw three specimens for sale in Hine’s shop. When he went on a hunt with one of Hine’s boys he found that the Prairie Chicken [Greater Prairie-chicken] was very common “but so wild as to be impossible to approach.” An entry in his notes from South Dakota began: “There is a barber in town who is also a taxidermist. I have spent every vacant minute over at his shop ... I mounted two owls and a ruffed grouse for him and gave him some points on bird work.’ The barber-taxidermist gave Percy skins of a Night Heron [Black-crowned Night Heron], Prairie Chicken [Greater Prairie-chicken] and Lark Bunting. Percy’s notes for 1896 start with a short self- examination. He wrote: “This begins another year with the birds. Have I learned anything in the last year? I think so. But such a little out of a great deal that might of [have] been learned.” He decided that he should give special attention to drawings of parts of birds — head, bill, wings, tail, feet — with the object of getting information for the classification of birds by species, and also the relation of plumage to flight. Under an entry added to the Preface to his Ornithological Notes he noted that he had carried out the intention he wrote down in May 1893 as regards Volumes one and two and that he would call his illustrations “Volume three”. He aimed to col- lect a picture of each of “our birds” signed, dated, 1996 named and numbered with the American Ornithologists’ Union Check-list number, and a sketch showing the distance between the point of the bill and the eye.*° Although he did not have a guide book of any kind with him in the field, and certainly no bird illustrations, he did mention using a field glass, though this presumably would have been a single lens with low powered magnification and not good light. To be certain of identifying a bird correctly it was necessary to shoot a specimen and key it out. This is where measurements and sketches of the salient parts of a bird were useful. Percy took his specimens and drawings seriously, and went about it in a systematic way. To a great extent it was a question of learning from his own collection of skins, though he could continue to return to the Museum at Michigan University for specific information, referring to books, and learn- ing from the experience of others. At the beginning of February Percy left the Company in Iowa and visited Ann Arbor. While there he took the opportunity to look at Audubon’s plates in The Birds of America, a copy of which was in the University Library. Percy was not over- awed by the great bird artist; in fact he wrote what he thought, or possibly what he had heard from oth- ers. “It is a big thing for a beginner to criticize Audubon but I think he is an over estimated man. The work shows an immense undertaking but a great deal of fault can be found with it. There is only one plate that really pleased me - a nest of Barn Swallows. The rest all show too little artistic value and idealization.’ From here he went to join Albert at the Royal Opera House, Guelph, where he looked for birds regularly in the spring. This included a trip to a local heronry. Percy and a friend wanted to shoot “fish cranes” [Great Blue Heron] as they were called and after much “soft soaping” the farmer on whose land the trees stood gave them permission to shoot three only. Apparently all the farmers owning land includ- ed within the limits of the heronry signed an agree- ment to allow no one on it because twenty-five or thirty birds had often been slaughtered “without thyme or reason”. As it turned out Percy only got two birds.** Meanwhile Ida Van Cortland and the Company had traversed several mid-west states in winter and arrived in Detroit in May. They spent a month in Hamilton and arrived in St. Thomas late in July, where they had a good reception. “Business has steadily increased. W. J. Butler and Albert Tavernier are both great favorites, and the singing of Wilfred Lucas is always a much welcome number of the pro- gramme.” The Dramatic Mirror announced that the Tavernier Dramatic Company would close its season at St. Thomas on | August 1896. It also said that Ida CRANMER-BYNG: A LIFE WITH BIRDS 11 Van Cortland would spend the summer holiday at Beaumaris, Lake Muskoka, and that Miss Van Cortland would not star next season, but would like- ly accept an engagement.*” The log book of Camp Coo-ee, written in Percy’s handwriting, shows that Albert came from Hamilton but stayed only two days. This was the last time that his name appeared in the log book. It was a sad but perhaps inevitable ending to the marriage, and part- nership, between Albert and Ida. But Ida Van Cortland was now forty-two years old and her daughter, Ida Clare, was nine in mid 1896. Ida had to think seriously about Ida Clare’s education, and about her own career. Albert Tavernier, although a congenial man and a good actor, had artistic tastes rather than business ability and his two attempts to become a successful actor-manager had failed. To succeed as a manager in the theatre in the 1890s one needed to be a businessman with capital. Albert appears to have worked hard and had no obvious vices. But the fact remains that the Tavernier Company made only a limited amount of money and that Ida’s share would not be enough to support her once she ceased to act.*? There seems to have been no sudden quarrel because they remained acting together until the end of the season of 1896. But they both had to think of the future. Albert Tavernier had to find another engagement. Ida took an engagement with Wait’s Comedy Company which toured in New England during the season 1897-’98. Young Ida went with her mother.*! The only member of the family who had no work and no income for the future was Percy. He did a lit- tle taxidermy work locally until September when he got a commission to mount about thirty skins of New Zealand birds at Grimsby, Ontario.°? This brought him close to Hamilton where lived Thomas MclIlwraith, the senior ornithologist in Ontario at this time and a founder of the American Ornithologists’ Union. Young Taverner called on old Mcllwraith and they talked about birds.°? Back in Beaumaris Percy had plenty of time to wonder about the drifting apart of his mother and stepfather, and the implica- tions for him. The island, and the cottage on the point opposite Beaumaris, was still his mother’s, a place where he and his mother and half-sister could always expect to spend a summer holiday. Otherwise the outlook was uncertain, as bleak, perhaps, as the weather of November in Muskoka country. Percy remained there as long as he could but when it grew too cold to stand, and the crossing from the island to the village by rowboat became too rough, and when he was down to his last few dollars he knew he had to go. The last entry in his bird notes was dated 12 November. He took the train to Toronto to face a new world without his small family and the extended family of the Tavernier Company to support him. 12 THE CANADIAN FIELD-NATURALIST CHAPTER 3. Getting a Start in Life Percy Taverner first met J. H. Fleming in December 1896 when he came down to Toronto from Muskoka “broke and aimless”. He was intro- duced to Fleming by James Grand of the firm of Grand and Toy, whom he had known for some years in Muskoka.! Percy hoped that Fleming would give him a job in the shop of Oliver Spanner and Co., taxidermists, on Yonge Street. Fleming, whose fami- ly seed business was on Yonge and Elm, just north of Spanner’s shop, had a business agreement with Spanner. This was not as a financial investment but as a way of providing him with facilities for prepar- ing his specimens and aiding his ornithological inter- ests. Although involved in his family business his absorbing ambition was to form a collection of mounted birds and study skins on a world-wide scale.” In those days a taxidermist’s shop was the meeting place for the shooting fraternity who brought in the trophies they wanted mounting, and the rarer specimens that they wanted identifying. Spanner’s shop was a natural centre for the exchange of ornithological information in the Toronto region and Fleming was at the centre of it. Although there was no job there for Taverner, for some reason Fleming took an interest in him and gave him the use of rooms above the shop. Looking back on this meeting Taverner wrote “I was a rather raw youth with more enthusiasm than knowledge and only vague glimpses of the great field ahead, and I have often wondered why he took an interest in me.” The collection of water-colour drawings of birds Taverner had made in an amateur way may have roused Fleming’s interest in him, “...perhaps he recognized some glimmerings of promise in my uatrained ornithological enthusiasm.”? Fleming was only three years older than Taverner but had been elected an Associate of the American Ornithologists’ Union in 1893. The two young men developed a close friendship and collaboration which was to last until Fleming’s death nearly 50 years later. Taverner was also befriended in Toronto by W.G. A. Lambe, founder of a company of grocery brokers, an original member of the Argonaut Rowing Club, and a collector of birds’ skins. He bought Percy’s colour-wash drawings, and nearly every Sunday Percy had dinner either at the Flemings’ or the Lambes’. He later recalled “From Lambe I was introduced to an old-world-like courtesy that I had not seen before”. Both Lambe and Grand gave him “very material aid” for which he was always grate- ful.4 Percy was a regular visitor to 267 Rusholme Road, the Fleming family home since 1892. Here, in front of the gold-fish pool in the greenhouse, cigars in hand, Fleming and Taverner sat and talked many times after Sunday dinner. From Fleming Percy gained his first broad view beyond parochial fields and was introduced to a wider outlook. Here they Vol. 110 discussed ornithological standards and ideals. “From Fleming I gradually absorbed the foundations of sci- entific ornithology as it was developing at that time ... It was as good, perhaps better than a university, at least it was more broadening.”° Here, also, Percy could examine Fleming’s nucleus of a collection, which already contained some interesting specimens, including birds from outside North America. By the time of his death in 1940, Fleming had built up one of the largest and finest private collections of birds in the world.® By taking odd taxidermy jobs, and with the help of friends, Taverner managed to exist. All his spare time he spent at the shop in the company of Oliver Spanner, his assistant Fred Dippie, Fleming and any- one else who happened to be visiting. “Fleming usu- ally occupied a chair in the corner keeping an ear open for the bell announcing a customer (or the game warden) downstairs, when he descended to attend to business or to pump a visitor for informa- tion...”’ In this atmosphere, blue with tobacco smoke, they discussed many things, but mainly ornithological matters. Percy had fallen on his feet. Suddenly he was a member of a circle of kind and congenial men with whom he had something in com- mon, and even his stammer ceased to matter much. Taverner was no outsider to the bird talk and anec- dotes that flowed around him. He, too, had experi- ences of his own that he could relate, such as the September day at Beaumaris in 1894 when he shot a bird he did not recognize. He sent the skin to his mentor, A. B. Covert at Ann Arbor, for identifica- tion. Covert sent it to Robert Ridgway of the Smithsonian Institution who identified it as an immature male Greenland [Northern] Wheatear. Covert claimed that he had taken this specimen. The Smithsonian kept it and credited it to Covert.® Another bird record which showed that Taverner was a serious and knowledgable bird man was his letter published in the Gravenhurst Banner about a Sharp-tailed Grouse killed near Beaumaris in October 1896, the first reported capture of this species east of Sault Ste. Marie.’ Not all the activity and conversation in Spanner’s shop was centred on birds. Spanner designed, built and raced his own boats almost weekly throughout the season. At times the conversations centred on yachts and yachting, on technicalities such as load waterlines, beam, sail areas and length overall. Taverner often crewed for Spanner in his 19 foot class boat.!° It was good fun and many years later he still hankered after “the feel of the stick and the pull of the sheet”.!' During the years 1897 and 1898 Taverner was often in Spanner’s shop as birds were brought in to be “made up” for collectors. He had the opportunity to see birds in various plumages — spring breeding, fall molt and winter plumage, as 1996 CRANMER-BYNG: A LIFE WITH BIRDS 13 4 The taxidermy shop of Oliver Spanner and Company at 358 Yonge Street, Toronto, which Taverner frequented to obtain the latest bird sightings. Galbraith Photo, Toronto, Ontario, undated, but wall calendar reads 1902. (Reproduced courtesy of the Royal Ontario Museum Library and Archives, number 0001264.) well as the chance to study their anatomy closely,!* such as the male Cory’s Least Bittern Spanner obtained. In his notebook Taverner recorded some bird outings in the spring in Toronto with Fleming, or with Lambe, or others. Once Spanner and Taverner cycled out to the western suburb of Etobicoke in early April where they got two Ruby- crowned Kinglets and an Eastern Phoebe. In May 1897 he was at Muskoka recording the spring migra- tion. That summer he was in his boat near Crown Island when he saw two large hawks with pointed wings fly round him screeching loudly. He guessed they were Duck Hawks [Peregrine Falcons] and that the white on the cliffs marked the site of a nest. He decided that if he could screw his courage up to the point he would have himself lowered to where he could look at the cliff face. Later he heard that two local boys had climbed down and taken the eggs. However, in 1898 he found another Peregrine’s nest off Crown Island that it was possible to look into. When a clutch of four eggs was in the nest he took a photograph of them. In 1897 his mother and sister were not listed in the guest book as being at the cot- tage, but a family of ten from Pittsburgh rented it from early July until mid-September. Percy then moved into the cottage and left on 23 December. It must have been a cold stay all alone on Gibraltar Island, one that Percy remembered all his life, and the story of how he managed to fend for himself was one that was handed down orally, and probably gained in the telling.'> Back in Toronto in time for Christmas he caught up on the bird news from Fleming and Spanner. The following spring (1898) Percy went to the cottage again though according to the log-book his mother and sister were not present at any time. Much of the summer he spent doing photographic work among the tourists. Percy kept a notebook in which he recorded his negatives numbered from one to 148, with a short title for each subject. At the back of the book was a list of photos giving date, size, type of paper and mounts, and a number as though prints had been sold. There were some pictures of the Muskoka area though more were of Ann Arbor.'* He must have been busy with photography and taxi- dermy jobs because his bird notes were very scanty 14 THE CANADIAN FIELD-NATURALIST and then petered out altogether. Early in December he was sent a [Northern] Hawk Owl for mounting by a friend in Beaumaris which had been shot by two of his boys. They had seen it perched on a tree as they walked to school. When they returned it was still there so they went home, fetched a gun, went back and shot it. Taverner commented with his tongue in his cheek: “A truly unsophisticated bird.” Percy returned to Toronto a few days before Christmas in time to spend Christmas with his mother and sister Ida in Port Huron. The year 1899 marks a major change in Taverner’s life. This was the year in which he kept no bird notes, not even a spring migration list, but settled down to learn a profession on which he could depend for a living. He boarded with the family of C. F. Smith at 513 St. Clair Street, Port Huron, where he had previ- ously lived while at Junior school. It was like coming home. Percy had already shown an ability to draw quite well and now he was to put this promise to the test. He took a correspondence course in architecture while working in the office of George L. Harvey, architect, for the sum of $3.00 a week.'® In the spring, summer and fall he went out early in the morning bird watching, spent working hours in his employer’s office tracing plans and drafting buildings, and at night supplemented his tiny salary by rowing passen- gers across the St. Clair River to Sarnia after the reg- ular ferry had stopped. The fare was 25 cents, and drunks who had missed the ferry boat were regular customers. Percy kept a cudgel in his boat for self defence if necessary. Potentially more dangerous were the numerous freighters between which Percy had to steer a course.'’ He also took on odd jobs of commercial and private taxidermy. In his photo- graphic notebook, dated 1899, he copied out four pages of notes from Rowley’s “Art of Taxidermy” which were mainly of chemical formulae. In spite of starting to study for a career Percy did not give up practising taxidermy, as witness a letter he wrote to Ernest Seton-Thompson early in 1900 about the pos- sibility of taxidermy being raised to the position of a high art. At this time Thompson-Seton (the form of his name which he used from 1902 onwards) was known as a wildlife artist, though his book, Wild Animals I Have Known, published in 1898, was to make him better known as a naturalist writer. Seton answered that “the taxidermist must learn form and expression, just as the sculptor or painter does. As a matter of fact he must be a sculptor... The best art is always suggestive not realistic ... Its strongest quality then, is to appeal to the imagination.” He ended by stating his opinion that taxidermy could never rank among the higher arts.!® Percy continued taxidermy work, not as an art but as a means of barter. The fol- lowing advertisement appeared in the magazine American Ornithology at this time. “Old Magazines Wanted — I will be prepared this spring [1901] to Vol. 110 exchange mounted birds and scientific skins from this locality for back numbers of magazines. Address: P. A. Taverner, Port Huron, Mich.”!? Another step which Taverner took at this time was to write to John Macoun, Naturalist, The Geological Survey of Canada, with several records of birds taken in the Muskoka area. These were intended for incorporation in the Catalogue of Canadian Birds which Macoun was in the process of compiling.”° Taverner wrote to him in December 1899 with infor- mation on the Sharp-tailed Grouse movement and mentioned a Wilson’s Petrel [Wilson’s Storm Petrel]. Macoun replied asking Taverner to write a note on the petrel and send any notes that he had on geographical distribution that would add to the infor- mation in Thomas MclIlwraith’s Birds of Ontario, as well as notes on nesting habits of birds and the con- struction of their nests.*! This was an important step forward for Taverner’s recognition in ornithological circles. John Macoun, although primarily a botanist, was the senior government naturalist in Canada at this time, and as such was responsible for compiling the first comprehensive annotated list of birds recorded in Canada. Taverner was an unknown enthusiast who had already collected a few relatively rare species and was keeping detailed records in the Muskoka area. He realized the value of being in touch with a man of John Macoun’s prestige and having his records accepted in such an important compilation. One further development in Taverner’s efforts to be accepted as a reliable amateur ornitholo- gist was his election to associate membership in the American Ornithologists’ Union.” As regards his family life in Port Huron not much is known. The 1900 United States Population Census shows that there were six people living at 513 St. Clair Street: Charles Smith, his wife Maria, his son John, his grandson Stanley and granddaughter Gale, and Percy Tavernier. Percy was shown as being born in Canada, his parents as both born in England. The year of his immigration into the States was shown as 1885, his profession as architect, and his marital status as single.”> John Smith is shown as “widdower”, his pro- fession as travelling salesman. Charles Smith was born in New York; Maria Smith was born in Canada, her father was born in England and her mother in Ireland. Mr. Smith’s occupation was given as “Superintendent of street sprinklers”. Percy’s sister described his con- tinuing interest in natural history at this time. “His room in Port Huron was a veritable magnet for con- genial spirits... The trays of eggs and artificial birds’ eyes were fascinating to me, and all his explanations of birds and animals, in fact everything. He always seemed an encyclopedia. In fact he was a great reader of that very work and often had three or four volumes ... under his bed.”7+ Percy’s mother had left the stage at the end of the 1897 season as far as we know. She herself said that 1996 when she found a demand for other work that she could do she “quietly left the field to those who had the power to amuse and attract by newer methods”. Percy described her new vocation as “organizational work for a fraternal insurance company”.?° From several sheets of headed notepaper that Percy used in 1904 we know that Ida Van Cortland was employed by the Independent Order of Foresters.”’ This is not quite such a dramatic change of occupation as may seem on the surface. Ida’s experience organizing the Tavernier Company and travelling on circuit through Michigan and Illinois would have given her the abili- ty to organize salesmen selling insurance for the Foresters. We know that by 1902 Ida Van Cortland was in Chicago in this capacity. Whether she was already working for the Foresters while living in Port Hurton is not known.”® Information gleaned from the cottage log-book is of interest for this period. In 1899 it shows Ida Van Cortland and Ida Clare spending the summer at the cottage on Point Coo-ee with eleven visitors alto- gether, including a Mrs. Ella T. Nash who came to Muskoka as a representative of Campbells Illustrated Journal. She hoped to have an article on the Muskoka Lakes in its October 1899 issue com- missioned by the Grand Trunk Rail Road and Navigation Co. She planned to use the material gath- ered during the summer for descriptive articles for various magazines in the spring of 1900, together with illustrations supplied by the G.T.R. One evening in August 1899 she gave a talk to a large audience at the Prowses’ hotel at Beaumaris on “palmistry”, and for several days subsequently she was kept busy reading hands. Percy was not listed as visiting but Stanley Smith, aged ten, was there. For the years 1901-1903 the cottage at Point Coo-ee was rented out, while the Taverner family used the shan- ty at Kamp Kozy Korner. In 1901 the family occu- pied the same place during August and Percy expended a lot of energy trying to destroy poison ivy which had taken over the Point where they hoped to build a new cottage. At some time between 1901 and early 1902 Percy’s mother and sister moved to Chicago where Ida worked in the central office of the Independent Order of Foresters. Percy was still in Port Huron in August 1901”, but during the winter of 1901-1902 he took a night course in Chicago*®, and during the spring of 1902 he was keeping an active bird migra- tion list there.*! All Percy himself said about his stay in Chicago was that he was a draftsman in various architectural offices, but spent much spare time bird watching with opera glasses in the city parks, as well as some minor collecting, mainly about Roby Lake on the outskirts of the city.** However, his sister was more forthcoming and wrote: “In Chicago Percy had a very severe pneumonia, nearly didn’t make it. It was then he grew a beard; the Doctor CRANMER-BYNG: A LIFE WITH BIRDS iI) thought he had better not remove it during the winter and he wore it always thereafter”.** Most likely this was during the winter of 1902- 1903 because he wrote at the beginning of May 1903 of “my late sickness” as if it had occurred then.*4 Certainly by early March he was out watching birds with a companion and explaining in his ornithologi- cal notes how and where they went as though he was seeing some of these streets for the first time. The disjointed and simple notes in which he had recorded what he had seen in the period 1893-1898 now, after a lapse of four years, suddenly blossomed again. But the style had been transformed in the intervening years; the brash young naturalist had matured. He was now more than a collector of bird skins, he was also a collector of precious moments and impressions inspired by the songs and movements of birds in their natural surroundings. He compared the fresh looking plumage of what he considered to be newly arrived Song Sparrows on migration with the plumage of Horned Larks, grimy from the city’s smoke by which he judged that they had spent the winter in Chicago.*° Walking in Jackson Park one April morning he saw a male White-throated Sparrow, and heard a single whistle come through the fog from across the lagoon. “His ‘hard times Canada, Canada, Canada’ went right to my heart. It is the first I have heard since leaving Muskoka and his clear whistle brought back memo- ries of gorgeous sunsets through black pines upon placid lake and for a moment all the glamour of the northern wilds was about me.”*° Percy no longer wrote for utilitarian reasons only; he wrote to com- municate his feelings and his appreciation of nature to others. His notes now can be classed as a journal. An exciting few days occurred at the beginning of May 1903 when his friend Fleming came to Chicago to join an AOU excursion to California. Fleming stayed at the Taverners’ so that the two men had plenty of opportunity to talk about ornithology. The first night they: sat up and talked birds and other things until 2 a.m. The next morning they went to see Professor Whitman, well known as a pigeon fancier, who had species from all over the world in his aviaries — Band-tailed from Western North America; Ringed-neck Turtle Doves [Ring- necked Doves]; ground doves from Australia, New Zealand and South America; Malacca Pigeons; Blue Rocks [Rock Doves] from Europe and many others whose names Taverner promptly forgot. But to Taverner the gems of the Professor’s collection were twelve Passenger Pigeons. “This day I have seen real live Passenger Pigeons. Think of it. I am almost humbled with my good fortune. And grand birds they are. These seemed as strong and lively as though they were not about the last of their race but as they were when their numbers filled the heavens and their shadow blotted out the landscape from sight.”*/ In the afternoon Fleming and Taverner went to the Field Museum and were full of admiration for the 16 THE CANADIAN FIELD-NATURALIST taxidermy work of the Virginia deer and Quagga groups. When the museum closed at 4 p.m. they walked home through the shrubbery on one side of Wooded Island and noted Palm and Myrtle [Yellow- rumped] Warblers, White-throated Sparrows and Blue-gray Gnatcatchers as common, and one Scarlet Tanager. In the lagoon were [Common] Loon, Horned Grebe, and Blue-bill [scaup] and Red-breast- ed Mergansers.*® On the way Percy and Fleming had a talk about Ernest Thompson Seton. They both agreed that from what they read it seemed that Seton had passed his zenith and that his current serial in the Ladies Home Journal was a failure. In the evening Percy went with Fleming to the home of Ruthven Deane where they met some of the bright lights of the bird world — T.S. Palmer, Frank Chapman, J. A. Allen and C. H. Merriam, to name only a few.” Percy wrote that he was so bewildered by the pres- ence of the great ones that he could hardly remember who was there and who was not. This was the first time that he had seen any famous ornithologists, and he was somewhat overawed, but at the same time reassured to see that they walked and talked like ordinary mortals. He went with them to the depot to see them board their private excursion train. When the train arrived the gates were opened and they all filed through except the odd man out — Percy Taverner. It was a poignant moment which he described with feeling in his journal. As he watched them climb into the Pullman coaches he knew that they were bound on the most delightful excursion that America could offer to a man of his interests. It tasted a little bitter to realize that the money spent during his “late sickness” would have carried him with them and back again and satisfied one of the strongest desires of his life.” He felt a little “sad for himself” by the tone of his entry. Although by now they would have forgotten him and his almost unpro- nounceable name, yet the memory of having met them would urge him on to work, and one day, if he lived, he would join an AOU trip himself. Taverner was not “down in the dumps” for long. In June he went to Roby Lake and recorded that King Rails were “common”. He found the nest of one with a young bird just coming out of the shell, a little black ball with shiny black eyes. The parent bird came off the nest at Percy’s feet with an amaz- ing splashing and cackling. Later Taverner went back to the nest and had a good look at the bird and identified it with certainty. He counted the remains of twelve eggs. With a companion he found several small rails’ nests but all had hatched and the young gone. Later they found a nest with one Least Bittern’s egg, as well as flushing five or six of this species. His comment in his notes was enthusiastic: “This has been a day indeed — a day to remem- ber.”4! During the heat of late summer he did little collecting in the field. Instead he made use of the Field Columbian Museum for learning more about Vol. 110 ornithology. One day he visited Professor Dearborn who described the method developed by the taxider- mist Akerley for making models in clay and papier mache, and coated with shellac, which were both strong and relatively light. This information was to be useful to Taverner several years later.*” His ornithological education was also broadened by his correspondence with Fleming, giving perspec- tives on a wider world. The group of 40 members of the AOU under their leader, C. Hart Merriam, whom he saw off at the depot in May in two special Pullman cars visited Santa Fe, the Grand Canyon, the Mojave Desert, San Bernardino and Los Angeles before attending the AOU meeting at San Francisco. Fleming wrote to Percy with his person- al impressions of some of the participants. “Dr. Merriam is simply a wonderful man full of infor- mation, and the success of the trip is mostly due to him. Chapman was not visible in the smoking room after. He has a wife big enough to squash him a huge woman, he also has his mother with him. [ liked Dr. Allen he was very quiet and nice. He smokes when permitted. Mrs. Allen was very nice but she kept a close eye on him. I got to like her. The real revelation of the trip was Fuertes the artist as modest unassuming a fellow as one would want to meet he is a genius. I don’t like all his work but it is based on a thorough knowledge. He has a big future before him.”*+ Such a letter would serve to make Percy feel as though he might be able to get to know these distin- guished men one day, that they were quite human. It may also have made him determined to earn their approval so that eventually when he attended an AOU meeting they would know who he was. Judging from a few remarks in his journal entries he was suffering from a sense of inferiority perhaps caused by his stammer and, in addition, by a lack of self confidence resulting from an insecurity of iden- tity. He had not yet made his mark. In October 1903 he attended a lecture at the Field Museum on bird migration which was illustrated with slides, and with maps showing breeding, tran- sient and winter quarters of various birds. One that impressed Taverner was the Black-bellied Plover which, in the American continent, was capable of migrating from its arctic breeding grounds south as far as Patagonia. This excited Percy’s imagination and set his mind working. In his journal he recorded that after coming home he started to ponder the problem of migration, and asked himself the ques- tion “why do these birds travel this awful distance?” He argued that it cannot be lack of food in their southern grounds because they support themselves all winter there and surely must be able to do the same during the summer. He then sketched out a the- ory to explain this in his journal, using notes he had made from books and articles on natural history problems such as the migration of birds.** He impul- sively sent his first thoughts on the cause and origin 1996 of migration to Fleming for criticism.*° Fleming showed them to Dr. Brodie who said that Taverner was rather mixed up over the extent of the Ice Age, and Fleming advised him to read some modern European writings on the subject before offering them as a paper. He encouraged Taverner to keep thinking. “Go ahead and think all you can. You are able to transfer your thoughts to paper, a blessing few possess.” Taverner went ahead, but instead wrote to J. A. Allen who advised him to send his paper to the AOU Secretary before 13 November of that year. +” It was accepted as a short paper for the Twenty First AOU Meeting in Philadelphia. Since Percy did not attend the meeting his contribution was given to Dr. T. S. Palmer, Assistant in Charge of Game Preservation, the US Biological Survey, who read it aloud on his behalf. This was a typical action by Taverner as a younger man. He acted impetuous- ly rather than cool-headedly. However, the AOU members listening to it received it well and, accord- ing to Fleming’s report they applauded. There was only one comment, namely that A. R. Wallace, the independent discoverer of the concept of evolution, had referred to a somewhat similar theory in his Geographical Distribution of Animals.** But this was only the beginning. Taverner now wanted to have it published in The Auk but to do that Allen, the editor, said that a number of basic changes would have to be made. T. S. Palmer wrote suggesting that he should read a chapter by Professor W. K. Brooks on “Migration in its Bearing on Lamarckism”.*? In reply to a further query from Taverner about the pos- sibility of publishing it in The Auk, Allen jumped on him saying that it was not a new hypothesis; that he himself had put forward the same ideas as hypothe- ses ten years ago; that Taverner’s statement that no glacial traces had as yet been discovered in the southern hemisphere were “grossly in error”? In a second letter he advised Taverner to revise his paper and to discriminate clearly between what he consid- ered to be new ideas and what was common proper- ty.°! Finally he wrote saying that the revisions were satisfactory and it was now all right for publication in The Auk.* It was published as an eleven page arti- cle in The Auk with the title “A Discussion of the Origin of Migration”. In it he discussed several dif- ferent hypotheses, among which was that of Alfred Russell Wallace who suggested that migrants in the spring were searching for soft-bodied insects suitable for nestlings; that as the season advances in the trop- ics it becomes dryer and dryer and such insects soon disappear. Taverner speculated that it was the need for large quantities of food that drove them north. As an article summarizing the various views of that time it was found to be useful. It was a useful exercise in hypotheses. Towards the end of 1903 Taverner’s creative energy suddenly began to blossom and his physical CRANMER-BYNG: A LIFE WITH BIRDS IL energy to match the workings of his mind. In his “Journal of Bird Observations” for 28 November 1903 he noted that he had written to Dr. Palmer thanking him for reading his paper and for his criti- cism. He took the opportunity to mention his own scheme for studying the migrations of individual birds by tagging (banding) them, and asked Palmer if he thought that a general movement for tagging migrants by an organized scheme could be started. This was fortunate timing. Palmer replied that Dr. Paul Bartsch of the Smithsonian had made some experiments in tagging 100 night herons in two colonies near Washington, and had a paper in the press discussing the results. Palmer also showed Taverner’s letter to Professor Wells Cooke of the Game Preservation Department, who was in charge of bird migration and distribution data. He suggested that Taverner should write to both men outlining his scheme. Taverner proposed that the Migration Committee of the Biological Survey should issue small rings made of aluminum wire or some light non-corrosive metal. The rings should be stamped with a number and a record kept of those to whom they were sent.>* In a letter to Fleming he said that the whole body of ornithologists would have to participate in order to get any results. He also mentioned that he had designed a brick trap for capturing ground-feeding birds. In his Journal he drew a draftsman’s plan of perspective views of the trap with measurements.°° In the same letter he said he was undergoing an extensive course of reading in order to get his ideas together and to satisfy himself. He mentioned that he was thinking about the property of matter, the unity of nature, the tendency to bring in electricity to explain anything complicated. He was sending Fleming a separate paper on evolution.°° Fleming seemed to be acting as his “supervisor of studies” to whom he submitted essays for criticism — and encouragement. Taverner had a lot of productive energy at this time because he had a good deal of spare time. In fact he mentioned being laid off work during the past winter.°’ So he started a number of projects, one of which was to make sketches for an extension to Fleming’s house for a bird room which he sent him for comments and return.** He also found time to write a short paper on feathers and sent it to the acknowledged expert on the subject, Dr. Jonathan Dwight. This was typical of Taverner at this period in his life. He would get enthusiastic about a problem, read it up as best he could, study it in the field if possible or in a museum, and then pro- duce something on paper which could hardly fail to be superficial. But nevertheless he sent it off to an expert to read. Dwight treated him kindly in his reply, told him where he was wrong but gave him encouragement when he praised him for being “so close an observer”.°? 18 THE CANADIAN FIELD-NATURALIST By the beginning of 1904 he knew that his mother was to be transferred to Detroit in the spring by the Independent Order of Foresters. He told Fleming the news and commented that they hoped to buy a place in the suburbs and settle there for good since his mother “had been given complete charge of the state’. Percy welcomed the prospect of moving to Detroit and wrote: “Detroit will make a good centre for bird work it is easy to reach Port Huron and the transitional fauna and also Ann Arbor and the Carolinian ... I believe the Mich[igan] Ornith.[ological] Club is also progressive and has its headquarters in Detroit. Altogether I am looking forward to the change with a great deal of plea- sure. Another great improvement will be in being nearer - to Muskoka.” ©° There was exciting news from the University of Michigan museum at Ann Arbor. When Percy knew that he would be living in Detroit he wrote to his friend Norman Wood the taxidermist there. Wood was glad to hear the news and sent Percy a Museum Report and a copy of the Michigan Ornithological Part II — Apprenticeship Years CHAPTER 4. Ornithologist-in-the-Making The move from Chicago to Detroit was yet anoth- er upheaval for Percy, his mother and sister; they had already done enough moving for a normal life-time. But there is little indication that they regarded Chicago as a permanent home, and if they were merely exchanging one large American city for another fast-growing modern one they were, at least, returning to an area that they were more familiar with — southeastern Michigan. Detroit was no dis- tance by interurban rail from Ann Arbor and Port Huron where they had good friends, places where Percy had lived and would feel at home. For Percy Detroit had the advantage of being nearer to Toronto and the opportunity of seeing his good friend Fleming from time to time. They arrived in Detroit at a good time of the year — early spring, and Percy soon began to explore the neighbourhood for birding spots. In his journal for 6 April 1904 he listed sixteen species seen, among which were [Eastern] Bluebird, [Rufous-sided] Towhee, [Eastern] Meadowlark, Vesper Sparrow, Fox Sparrow, and Purple Finch.! He was off to a good start, exploring a new location in the spring of the year, doing what he enjoyed most and transform- ing his observations into notes for his journal. It did not take him long to renew old friendships and make new ones through the Michigan Ornithological Club (MOC), of which he had previously been a member. He attended the annual meeting of the club at Ann Arbor early in April when he heard two papers on Vol. 110 Club Bulletin. The main bird news was that in June 1903 Wood had spent nearly three weeks in the south of Michigan and had found three colonies of breeding Kirtland’s Warblers. He had the only pho- tos of this rare warbler to be taken from life in vari- ous postures. He promised Percy a copy of his account of this discovery and asked for a copy of Percy’s article on the origins of migration.°! The Taverner family had lived in Chicago for only three years, not sufficient time to put down roots and make a circle of friends. Their memories of those years were likely to have been clouded by Percy’s nearly fatal illness. Chicago, in the first decade of the twentieth century, was a booming city of industries, railways, dockyards and immigrants packed into suburbs. It was also a financial centre of expanding wealth and population, new architectural styles, and labour unrest. It was a city in which the air was regularly polluted; it was not the best place for someone whose lungs had recently been weak- ened by pneumonia.° the Kirtland’s Warbler.2, Wood asked him to come and look for Kirtland’s Warblers but he reluctantly had to refuse. As he explained to Fleming: “What with my lay off last winter and the expences and enforced idleness consequent on our moving I’m afraid my bank ac’t [account] will not permit it. Too bad to lose such a chance too, isn’t it.”* This was not the first time, and it wasn’t to be the last, when Percy lost an excellent birding opportuni- ty owing to lack of money. However, the rare Kirtland’s Warbler gave him an interesting topic to investigate and a problem on which to write a short article. His paper “The Origin of Migration” was published in The Auk in 1904? and this slight success in the field of speculation probably encouraged him to focus on the origin of the Kirtland’s Warbler. Based on information in the two talks he had heard, he speculated on how the Kirtland’s could have orig- inated. He came to the conclusion that from present knowledge it seemed likely that “Kirtland’s Warbler is what remains of a once far more widely distributed species wintering along the Gulf States and spread from thence to the Bahama Islands. They migrated up the Mississipi Valley and perhaps also up the Eastern States, which would account for our records at Washington and elsewhere along that line. The breeding grounds then likely covered the greater part of the Alleghanian Transition Zone. Changed conditions ren- dered this continental portion of the southern habitat untenable for a permanent winter residence, and the species became extinct there, leaving the Bahaman indi- viduals the only surviving representatives of the race.” 1996 Taverner offered no suggestion as to what the “changed conditions” were.° By publishing papers and taking an active part in an ornithological club Taverner became known as a serious amateur ornithologist studying the birds of the Michigan-Ontario border. In this way he was able to hold up his head and not let his stammer, lack of higher education, and money weigh on him too heav- ily. By nature he was resilient and quick to enjoy life in the company of congenial friends, especially if this meant field trips or just short tramps in the country observing and recording the natural world. Through the MOC he quickly made a friend with whom he was to spend a great deal of time in the next few years. Bradshaw H. Swales was born in Detroit in June 1875, the same month and year as Taverner. He graduated from the University of Michigan with a degree in law, and by the time they met in April 1904 had already published some forty bird notes in vari- ous journals, based on his knowledge of Michigan birds. He was also making a collection of skins. Both men had a strong attachment to “scientific” ornitholo- gy as opposed to egg collecting and casual list mak- ing and they soon teamed up. Through Swales, Percy met Walter Barrows, Professor of Zoology at Michigan Agricultural College, East Lansing, a rec- ognized ornithologist, with whom he had recently corresponded about the [Northern] Wheatear record claimed by Covert. They first met at Swales’ house where they talked birds until after midnight, mainly about Barrows’ projected List of Birds of Michigan. Barrows wanted the notes that Swales and Taverner were collecting on the birds of St. Clair County. This was a fruitful meeting for Percy since Barrows became another professional ornithologist friend with whom he corresponded regularly, and from whom he could learn much. He was also a useful “patron” for Percy’s drawings. From the beginning of May until mid-July 1904 Swales and Taverner birded whenever they were free, and Taverner’s journal contains regular descrip- tions, some quite extensive, written with verve and sharp images. Taverner was enjoying the spring weather and the new countryside, and his pleasure is reflected in his writing. They were birding on 4 May and Taverner noted that the big migration day nearly always occurred around that time. To him 4 May was the greatest day of the whole year. He described seeing his first hummingbird of the year and con- cluded “Well this great day has come and gone.”’ There was a new zest and feeling of satisfaction about what he saw and heard. He could hardly wait to get away from work as his entry for 7 May shows: “Saturday is my afternoon off — off to the woods.” With two new acquaintances he went to Gross Point where he noted: “Birds, birds everywhere and all singing”. By 21 May he noted that the spring migra- tion seemed almost over. But Swales and Taverner CRANMER-BYNG: A LIFE WITH BIRDS 19 continued to go out regularly, as breeding birds were active. On 30 May they found two nests of Louisiana Waterthrushes and put bands on seven young birds about to leave the nest.* Early in June they made the first of many visits to the St. Clair mud flats where they began a new project of record keeping. They travelled on the Port Huron Rapid Railway System to Pearl Beach and stayed the night at the Wellington. They spent considerable time on the St. Clair flats in the next few years obtaining valuable information for Barrows’ Michigan list. Apart from Swales, Percy’s first impressions of some of the other members of the MOC were not encouraging. In mid-April they went birding with J. Clair [Clare] Wood and two others along the Ann Arbor-Ypsilanti Electric Railroad. The others had some hawk nests marked and soon had taken three sets of Red-shouldered Hawks’ eggs. Meanwhile Taverner and Swales sat and watched a Turkey Vulture sailing about through their glasses, the first he had seen since his early days at Ann Arbor. Taverner gave his impressions of the others in his journal: “First impressions are hardly to be trusted but I am afraid that the ornithologists of Detroit are mostly “col- lectors” and egg collectors at that. Clair Wood seems to have had the most experience of any of them but he makes but small pretentions towards science, and I believe that he is apt to jump to conclusions. He keeps no notes and has faith in his memory — a great fault with a naturalist. Swales seems to be the only one here with any appreciation of the meaning of scientific work and he bewailed the inaccuracies that have unavoidably crept into the records from the above causes. He cited two cases of Yellow Chat | Yellow-breasted Chat] breed- ing records here in Wayne [County] but in neither case were the parent birds seen. Identification was solely from nest and eggs.” Taverner reveals as much about himself as the others. His ambition was to study birds in all their aspects, their relationship to each other and to their habitat, and to study this scientifically, taking noth- ing for granted. It was a justified avocation and in trying to live up to it he got a great deal of quiet sat- isfaction. The others behaved like boys, and his jour- nal shows them doing it. “As the rest of the party rushed off from one hawk tree to another and intent only apparently of [on] gathering a lot of eggs Swales & I found the pace too fast for any observations so we dropped behind and proceeded to “naturalize” according to our own ideals which was [sic] in this case to sit on a rail fence in the warm sun and dis- cuss ornithology in general and listened [sic] to the Winter Wren as he tried to recall his old breeding song or watched the grand majestic flight of our only Vulture — the Turkey Buzzard [Turkey Vulture]. We gained notes on two species. Wood, Wisner & Wood got sever- al egg shells. Who gained the most [?].”!° Taverner was ready for his annual holiday in Muskoka in late August, but compared with some 20 THE CANADIAN FIELD-NATURALIST other years when he was there in spring and summer there were few birds to be seen. Leaving Beaumaris early in September he joined Fleming in Toronto, and the following day they went to visit Klugh at his cottage.'' The next day Taverner continued to London and spent the afternoon with Saunders look- ing at his bird skins, and taking notes from his List of Birds of Western Ontario.!” There are some indications at this time that Percy was bored with being a draftsman. On the other hand when Fleming wanted plans for an extension to his house in Toronto Taverner set to with enthusiasm. Again, when Fleming needed maps made with data on them about irruptions of Brunnich’s Murre [Thick-billed Murre], for making slides which he could take with him when he attended the Fourth International Ornithological Congress in England in June 1905, Taverner was willing enough to make them. “Do not hesitate to send your map data to me. It is really a comparatively short job to make the maps. I will trace them off and mark them up in pret- ty short order. Of course I am pretty busy but I never found my time so well occupied but that I could not get in a little more work.”'? This might be the motto for the whole of the rest of Taverner’s life; he was well on the way to becoming a compulsive work horse. Family life seems to have settled down well for all three Taverners in Detroit. His half-sister, now a woman of twenty, took piano lessons from a Miss Martha Hohly of the Michigan Conservatory of Music. Martha was born in Toledo in 1880 and came to Detroit in 1900 where she taught piano and also acted as accompanist at concerts. In December 1906 she was invited to accompany the French violinist Henri Ern who was sharing a concert at which the French composer Camille Saint-Saens was the piano soloist, and the program was mainly of his composi- tions. Thus Martha played the accompaniment to Saint-Saens’ well known Introduction and Rondo Capriccioso with the maestro nearby.!* Ida would have known about the concert and very likely would have attended it with her family, especially since in 1907 she was offering piano lessons herself.!> Cultural life in Detroit during the first decade of the twentieth century was flourishing. There were plays, operas, concerts of various kinds, recitals, vaudeville entertainment as well as amateur performances tak- ing place regularly. A look at the entertainment pages of the Detroit News Tribune for a single month in 1908 will give an idea of the diversity of entertainment available to the people of Detroit. As an acting family the Taverners would have been interested to see some of the plays and operas, and some of the “stars” who performed in them. Letters between Fleming and Taverner were not entirely about birds, and Fleming reported from Toronto: “Saw Ben Greet’s Company in Julius Caesar yesterday, most interesting; the plays are produced as they were in Vol. 110 Shakespeare’s time. I saw him at the University a year or so ago playing in the open air. One gets a much better idea of Shakespeare when the scenic effects are not pre- sent to distract the attention.””!° At the end of this letter Fleming’s strong sense of humour shines through. He wanted to let his friend know that he was forgetful about signing his letters. “P.S. Please sign your letters. You may put W.P. (with- out prejudice) if you like as the lawyers do, and feel yourself safe.” Though life was by no means perfect it was rea- sonably good. They had friends, such as the Smith family in Port Huron, with whom they spent Christmas Day 1905. Other friends, such as W. E. Saunders and family, encouraged the Taverners’ fondness for music. The Saunders stayed with the Taverners when they went to Detroit to hear the Boston Symphony Orchestra, and Ida and the Saunders’ daughter got acquainted.!7 A major event for the Taverners occurred in 1907 when Martha Hohly married Jacob Merton Wiest, the Sunday editor of the Detroit News Tribune. His photograph shows him to be a handsome young man. He had a good voice and was a soloist of some repute locally. The Wiests and the Taverners became friends and used to take part in amateur musical/operatic concerts. They formed a strong team. Ida Van Cortland had plenty of experience in acting and could help in the production of amateur performances. Martha could play anything required on the piano, Jacob Wiest had natural acting ability as well as a good voice, Percy was a useful handy- man in stage design and lighting and Ida C. could act and help wherever needed.!® But for Percy ornithology took first place and absorbed most of his surplus time and energy. The journal which he kept regularly during these years at Detroit contained a number of quite long entries. Above all he seems to have derived so much intense pleasure from his birding rambles that he wanted to preserve them permanently as happy memories. Sometimes Percy designated a certain date as a red- letter day. For instance the entry for 2 October 1904: “This has been a day. A sparrow day and a day above all other days. This should all be written in red ink.”!? Then for the next seven pages he related what he had seen as though he was compelled to tell it in full even if no one was likely to read it. Another of these special days occurred on 5 August 1905. “A Red Letter day to an ornithologist is one in which he discovers something new — at least to him — within the way of habit of some before known species or a new species altogether. Today I did the latter.”2° Such days were recorded red-hot as the words tumbled into his mind and the excitement of what he had seen moved him. Sometimes, when he had been for a “tramp of exploration” he included a small sketch map neatly drawn. In the spring he usually included a list of the first wild flowers seen during his April walks, before being overwhelmed by the 1996 full warbler migration in May. Another entry might convey something of the tension that built up inside him when he had spent all day in the office bent over his drawing board drafting plans while outside the birds were away and flying. He was like a coiled spring until, suddenly, it was time to go and he made a dash for it. “This evening took a hurry-up jaunt to the country and back by way of Palmer Park woods. Left the office at 5:30 and took the car right out there. It is not dark until about seven o’clock now and I had an hour or so of day- light before me.”?! Apart from his constant companion — his journal — there was a regular flow of letters between Fleming and himself which continued until Fleming’s death in 1940. After he reached Detroit his letters to Fleming began to mature, perhaps because he was stimulated by the new areas he was discovering, the rarer birds seen and new people he met. Many of these letters had an urgency about them, a sense of excitement that complemented Fleming’s cautious approach. It is true that at the same time he was writing regularly to others such as Professor Barrows and Norman Wood, to mention only a few, but these were run-of-the-mill ornitho- logical letters, while letters to Fleming, and from Fleming, were special. They were always about bird matters in some form or other, but often contained bits of information on their families, friends, gar- dens, health and personal opinions. Taverner, like Fleming, had a light touch, a gleam of humour that was part of his style. He first started typing letters to Fleming in late 1905 when he said at the end of a let- ter: “My handwriting is slowly improving but this machine cannot spell correctly.” Taverner’s erratic spelling was something no typewriter could cure. However, his practical mind was pleased with the information that the signs for sex, a useful thing for ornithologists, could be put on a new machine free of charge. He also began to use a typewriter to copy out his rough bird notes which would have the double advantage of being easier for him to keep and easy for others to read. It also meant that he had a carbon copy for other use. Apart from the information Taverner gained through his correspondence with Fleming and other amateur ornithologists, and through letters and con- versation with professional museum men, he had also begun to educate himself in the principles and practice of ornithology through reading. Although he could not afford to buy a collection of books and journals, nevertheless he borrowed from Fleming and scanned trade catalogues for bargains he could afford. His first major step in self education was to read Elliot Coues’ treatise Key to North American Birds, designed as a manual of North American ornithology which would enable the reader to identi- fy and label the specimens he obtained. The first sec- CRANMER-BYNG: A LIFE WITH BIRDS Dl tion was devoted to “field ornithology”, that is: how to collect and preserve bird specimens for study. Coues explained the rationale of field ornithology. “The true ornithologist goes out to study birds alive and destroys some simply because that is the only way of learning their structure and technical characters”. The reader is admonished to “shoot an unknown bird on sight...” and to collect skins as duplicates since they can be used as a medium of exchange. The best way to determine the exact geographical distribution of a species is “to have specimens from as many different and widely separated localities as possible”, as even the commonest bird has a special value if collected outside its normal range. “But let all your justifiable destruction of birds be tem- pered with mercy; your humanity ... should never per- mit you to take life wantonly.”* Another book which he relied on in this period and which influenced him was Robert Ridgway’s first volumes of The Birds of North and Middle America. In this period he was continually reading articles and notes as they appeared in The Auk, the most widely read of the ornithological journals at that time. As a member of the AOU he would have received a quarterly copy since early 1903. At the same time he bought “back numbers” whenever he could find them. Another advance that Percy made while in Detroit was in the technical side of his “calling”, namely the equipment he used and his skill in using it. As a boy, and even later, he had to make do with opera glasses instead of field glasses because of the cost. But once settled in a steady job in Detroit he felt the need of adequate equipment. On the red-let- ter “sparrow day” in October 1904 he noted in his journal that opera glasses failed to bring out the colour of birds well on cloudy days. As a result he wrote to a friend at the cottage-colony at Lake Muskoka, John Brashear, enquiring about the cost of field glasses. Since his friend was a partner in the firm of John A. Brashear Co. Ltd. of Allegheny, Pa., makers of astronomical and physical instruments, he did not lack for good advice. The best quality would cost about $48 a pair, but for about $10 it was possi- ble to buy an adequate pair with good lighting.*4 The next advance in equipment came in June 1909 when Fleming asked him, in a letter, what field glasses he was using. Although gaps in the correspondence exist we know that Fleming had recently invested in a pair of binoculars and wanted Percy also to benefit from having a pair. Flemming wrote: “Ever since I got those prism binoculars, I have seen the advisability of your having a pair, and if you will put that $35.00 in the savings bank I think I can fix you out with a pair, that is if you are not too particular as to make or above a gift from an old friend. I have recently got a pair of less power as I felt it was unwise for me to use glasses of too high a power as my eyes are not too strong. In fact I should not use glasses of any kind.”?> 22 THE CANADIAN FIELD-NATURALIST But aspiring to be a scientific ornithologist with only a pair of glasses and no gun was impractical and Percy was never without a gun if he could help it. At this point in his career he owned a .410 — a small shotgun with short barrel, light and handy for carrying, and ideal for collecting small birds. For larger birds he invested in a double-barrelled twelve gauge hammerless shotgun.*° Since nearly all Taverner’s ornithological friends and acquaintances collected birds he was able to get practical advice when he wanted it. W. E. Clyde Todd, of the Carnegie Museum, Pittsburgh supplied the name of a reliable maker of auxiliary tubes, and much detailed technical advice on powder and shot and the tech- nique for loading one’s own shells.”’ Being a handy man Taverner could make up his own shells to suit his own purpose and to get the best result. He was always ready to try out an “improved” product in order to see its effect in practice. “I used Dupont Smokeless powder in my shells today for the first time and found it very satisfactory — no smoke and very little noise and increased penetration and range.”* The perfect puff for a new product. In addition to finding birds, identifying them with field glasses, or shooting them first and then identi- fying them later with Coues’ Key to North American Birds and other books, there was also the laborious work of dealing with the dead specimens. In the field it was necessary to stop bleeding from nostrils, beak and pellet holes by inserting a wad of cotton or paper, and sawdust to dry up the blood. The feathers had to be cleaned if possible, and smoothed down before placing the specimea carefully into a collect- ing bag — though Percy always used a fisherman’s creel instead. On reaching camp or home it was nec- essary to skin and “make up” one’s specimens. There was no refrigeration available in the field, and decay and insects would quickly damage them. We have already seen that in the period c. 1894-1900 Taverner did a fair amount of taxidermy work, and that he was reasonably experienced at it. Now, in Detroit, with new opportunities he collected regular- ly — one might say insatiably — not to earn money, but to enlarge his own study collection. He wanted not only to build up a collection of various species and subspecies but also series of particular speci- mens, say of breeding females, in order to have a range of examples of plumages with which to identi- fy the birds he was seeing in the field and recording in his field notes. His journals contain many com- ments on his success, or lack of it. Sometimes he fired too soon — he should have crept farther for- ward; sometimes he held his fire too long, the bird was frightened by something and flew just as he was about to fire. By late 1905 he was able to report to Fleming: “Have been rather surprised to find I have collected and made nearly five hundred skins this year past. Not bad is it? I have the nucleus of a fair Vol. 110 little local collection.” By early 1911 this collection had increased to over one thousand skins.”? Having skinned the bird the skin was treated with arsenic powder to prevent moths and beetles damag- ing it later. Professional taxidermists usually suf- fered from a skin rash caused by this powder, and Percy was no exception. By late 1905 he had a rash all over his face and neck which at first he thought was the result of contact with poison ivy, but later he realized it was caused by the constant use of arsenic poison when preparing his bird skins.*° Before mak- ing up a skin Taverner, as a serious ornithologist, would usually measure the wing span of the bird, and the total length from the tip of the bill to the tip of the tail. He would also dissect the specimen in order to determine its sex, since the plumage of the juvenile male often resembles the female of the species. Also in spring it was sometimes helpful to determine the readiness or not of a specimen for breeding. He might also check the contents of the stomach to discover, if possible, what the bird had recently been eating, and in this way assess its eco- nomic value. Having done all this he would make out a label containing the date; his own serial collec- tion number; scientific name of the bird; place of capture; name or initials of collector; any remarks. Only then would he start the process of skinning.*! In the midst of all this intense activity Taverner became involved in a new enterprise which gave his own ornithological studies a push forward, and brought a new sense of purpose to ornithology in the Detroit — London region. In the fall of 1904 Fleming was preparing the paper which he was to present at the Fourth International Ornithological Congress to be held in England in June 1905. The subject was on irruptions of Brunnich’s Murre [Thick-billed Murre] in Eastern North America.*? At the same time Taverner was working on an article in which he aimed to present an acceptable hypothesis to explain why numbers of western forms of species that migrated regularly along the Mississippi Valley could also be found as stragglers in southeastern Ontario while in Michigan these forms were not reported.*? Both men were frustrated by the lack of sufficient data to round out their studies satisfactori- ly. While Fleming was able to supply Taverner with a few interesting records from the Toronto area for: American Avocet; Snowy Plover; Swainson’s Hawk; Le Conte’s Sparrow; Nelson’s Sharp-tailed Sparrow, Taverner was unable to add anything to the few existing records for the Brunnich’s Murre [Thick- billed Murre] in the Detroit area. In their correspon- dence at this time both men agreed that a survey of the birds of the Great Lakes would be of great importance, that this would have to be carried out by a number of competent observers over a number of years, and edited by one man who could judge the accuracy of their identifications.**4 1996 Taverner seized this opportunity to put forward a plan that he and Swales had been discussing. Replying to a letter from Fleming he wrote: “You speak of the lack of material for a survey of the Great Lakes. Now it is just to gather such material that we want to start a new club.” But how could its members keep in touch with each other, and main- tain their interest when they would be living a long way apart? Regular meetings would be impossible. Taverner proposed that it might be a correspondence type of club confined to members living in provinces and states bordering the Great Lakes, and that the club policy should be guided on strictly scientific lines. He went into considerable detail for a pro- posed organization, explained that the Michigan Ornithological Club would not be a suitable medium through which to publish material (“the club is a club of boys’), but suggested that if The Auk would not publish this material, then The Wilson Bulletin, which had a good reputation, might do so. He asked Fleming to come to London for a weekend meeting with Saunders and others to discuss the project.* During January and February there was a regular exchange of views about forming a club and eventu- ally Taverner, Swales and A.B. Klugh spent the last weekend of February 1905 at Saunders’ home. Fleming was unable to attend but was regarded as a founding member because of his keen interest in the project. At this first meeting it was only decided to set up an organization and call it the Great Lakes Ornithological Club (GLOC). A draft constitution was soon drawn up and sent to the five founding members by means of a Bulletin edited by Saunders and circulated from member to member with their comments. No regular meetings were proposed but a field trip to Point Pelee in May 1905 was planned.*° The location and climate of Point Pelee makes it a unique natural area. It is a narrow peninsula in the shape of a triangle, situated in the extreme southern Ontario, and jutting some nine miles (14 km) into Lake Erie from its base along the Ontario shore. This is the most southern point of the Canadian mainland and is blessed with a relatively warm climate which favours the growth of a variety of Carolinian forest trees and shrubs. In Taverner’s time there were open areas along the west side of the Point where squat- ters had built their homes and existed by marginal farming combined with fishing and wildfowling. As a result a variety of habitats could be found all with- in the small space (15 sq km) of the Point — wet and dry hardwood forests, cedar thickets, brushy tangles, dry fields, extensive cattail marshes of various degrees of wetness and containing ponds. In addition there were sandy beaches, with sand dunes on the east side. Among the Carolinian trees and plants were: black walnut, chestnut oak, hackberry, hicko- ry, juniper, red mulberry, western prickly pear, sum- mer grape and buttonwood; all in all a place ideally CRANMER-BYNG: A LIFE WITH BIRDS 23} suited to support a vast flock of birds during the spring and fall migrations.°7 W.E. Saunders was responsible for identifying Point Pelee as the most favourable locality for field work by the newly formed club. He had first visited the Point in 1882 when he was excited by its flora and fauna — cactus, sassafras, Carolina Wren, [Northern] Cardinal and small flocks of Passenger Pigeons, one of which he collected.** He had visited the Point several times since then collecting and making observations, and he realized the uniqueness to bird migration of this slender wedge of land point- ing into Lake Erie. Taverner and Swales now heard about these attractions at first hand, and talked them over in anticipation. When May arrived there were two young ornithologists eager for new ground to explore, both fired up with anticipation for the trip to Point Pelee. This is how Taverner described the excitement of that first experience: “May 13 1905. At last this trip has come off. It is a diffi- cult place to reach from here. We had to get up, catch two street cars and the ferry, pass customs and catch a 6.15 a.m. train that put us into Leamington at about 7.30. There we waited for Saunders’ train at 8.49. He came all right and on time and we took a rig that was to drive us to the east base of the Point ... As we left Leamington we passed through a very slightly hilly country the most noticeable features being the stumps of peach orchards killed the winter before last. Stray red cedars showed here and there and the fences were full of White-crowned Sparrows. A little stream or ditch that we crossed was covered over the surface with black evil-odored oil. It was fed from artesian wells the driver told us. We came down a road that struck the point somewhere in the middle and turned towards the east shore. Pretty well across the point as we drove along listening to the various sounds a loud whistling ‘chip chip chip r-r-r-r-r’ reached us and call[ed] our attention. The horse was stopped and we listened for a repetition of the sound, when we identified [it] as a Yellow-breasted Chat, and a minute after the bird appeared in the thicket just across the fence. The guns were put together and we went after the bird. We worked about for about twenty minutes with the result — | Chat taken by Saunders — 3 or 4 more seen & heard, | Bob- white seen and a Whiporwill taken by me. The Indigos were common and we watched a couple of males singing on the wing most prettily. On leaving here we drove on to the shore line & dis- missed the buggy.”*? They now walked slowly down the wide sandy beach of the east shore where the dunes rose up almost devoid of trees except for a few scattered cot- tonwoods. This was wader and shore-bird ground and here they came across about twenty-five Piping Plover, and many nesting hollows scraped in the sand but none with eggs. Taverner recorded Dunlin, Least Sandpiper, and a full-plumaged male Hudsonian Godwit which he was able to examine in his hands. Terns were recorded; Common, a few Caspian and one Black Tern; also Bonaparte’s Gulls and single Herring Gulls flying past. 24 THE CANADIAN FIELD-NATURALIST “Throughout the trip I think Orchard Orioles were more numerous than Baltimores [Northern Orioles] and both were very common. Near six o’clock we passed along the end of the lake [pond] and crossed to the East [=West] shore and made camp. We had supper and mea- sured the birds and made a few of them into skins, then wrapped ourselves in our oil-cloths and, with our feet to the good fire, slept the best part of the night. A Whipporwill called near us and we timed him. He whip- porwilled 15 times in 20 seconds, calling as regular as a pendulum beat.””” The place where they bivouacked was near to the present Visitor Centre in the cedars beside the west- shore road — just across from Post Wood. When they woke warblers were singing all round them, Bay-breasted and Black-throated Green being com- mon. After breakfast they loaded their provisions and equipment on their backs and started along the west shore. Taverner continued his journal: “This side of the Point is lined along the shore with beautiful thickets of red cedar & juniper between. A road runs along here parallel with the lake & separated from it with a screen of this evergreen. ... On the inland side of the road are broad fields — sandy but supporting a thrifty-looking farming community. There are farm houses at intervals all along here until the lake and the marsh behind it approaches too close to the lake shore to allow of cultivation. This is in striking contrast to the other side where there was but one building to be seen the whole way.”*! He noted a suggestive feature of each farm house was the array of pound nets hanging in the yard showing that the farmers combined farming with fishing. As they walked Taverner was taken by sur- prise. “I never saw so many warblers in my life.” In one big walnut tree he noted: 2 Bay-breasted, | Parula, a Myrtle, a Magnolia, a Redstart and 2 Blackburnian Warblers. “The fauna is peculiar, Carolinian. Not long after starting I found a lot of cactus growing in the sandy meadows.” In the mar- gin of his journal he drew a sketch of one. [Northern] Cardinals were not uncommon and a Connecticut Warbler showed itself. Later Swales and Taverner watched a Bald Eagle with “a superb white head and tail” which a Kingbird was harassing, div- ing at it persistently. Through his glasses he saw two eaglets in the nest. By now it was time to be thinking of returning home — on foot, not by vehicle. “We started in towards Leamington about dark. It was a six mile tramp and tired as we were with the unusual load on our backs it seemed like sixteen to us. However, about 9.30 we arrived there & put up at a hotel. Had a good cold bath & rub down and were soon asleep in a good bed and far more snugly ensconsed than we ever expected to have been in such a small town.”*? It had been a memorable trip for the three of them even though during the day Saunders lost his purse, and Taverner lost his pipe — “a major calamity in times of stress”.*4 The Great Lakes Ornithological Club was now launched, its second Bulletin was circulating to its six founding members during May, and another trip Vol. 110 to Pelee was planned for the fall of 1905. This was a more ambitious venture, planned for a fortnight, in order to give ample time for observing the fall migration, and a tent was rented from a London (Ontario) firm.** Saunders and Barrows hoped to come but Fleming was doubtful. Taverner devoted thirty pages in his Journal to recording the events. It was headed, rather grandly: “Sept. 4. 1905. At Camp Coues, Point Pelee, Essex Co. Ont.” “Made camp here today. Klugh is the only one of the crowd that finally showed up. Barrows and Saunders have, I fear, finally backed out & Swales is detained by business for a few days. I hardly expect that Fleming will get here. Klugh is greatly interested in the botany of the locality and is discovering a treasure at every step. Never saw as many raptors as are to be seen about here. One or more hawks are continually to be seen — more Sharp-shinned, I think, than any other though the Sparrow [American Kestrel] & Red-tailed and Marsh Hawks [Northern Harrier] are in evidence. This seems just as it should be expected. Down the road on the west side is coming a continual stream of warblers.”*» They had camped in the same place as in May, among the Red Cedar close to the west shore of the lake. This was valuable learning experience for Taverner because there were so many birds intent on their feeding so close at hand and so little alarmed by human beings, even when they were carrying guns; also because he had plenty of time to collect speci- mens, take them back to the nearby tent, study their plumage, skin and dissect them. The perfect holiday for a young ornithologist. For instance, he collected a Chipping Sparrow and thought from the streaked appearance of the crown that it was an immature bird only to find on skinning it by the complete ossifica- tion of the skull that it was an adult. From a look at Jonathan Dwight’s Passerine Plumages he gathered that this was the normal fall plumage of the species. There were no professional and highly experienced ornithologists visiting Point Pelee at that time. W. E. Saunders was the best person for identification, but when he was not there Taverner and Swales had to do their best with their own resources. At least they could bring their skins home and hope that Saunders or a professional at Ann Arbor would be able to identify them with certainty. In the middle of this intense birding Taverner was alert enough to spot a plant in flower that he did not recognize. Klugh iden- tified it as Apios tuberosa — ground nut. A major migration wave must have come in during the night for in his Journal for 5 September he reported: “Today the first day [of] field work at Camp Coues. Got up about 6.30 and searched along the road running in front of the camp. Never saw birds so thick in my life over such an expanse of territory. Usually in the fall the warblers cruise about in companies but here on the point the whole place seems to be occupied by one large company.” 1996 While Taverner found Blackpoll the commonest, with Magnolia, Blackburnian and Black-and-white Warblers plentiful, Klugh wandered off into the shrubbery bordering the cleared land nearby and col- lected a rarer warbler — a Connecticut. The long account Taverner wrote of their fort- night there is full of the exploring they did along the east and west shores, in the dry fields, the marshes and lakes within them, the wooded areas and the tip of the Point. It is also a vivid and detailed account of the birds they saw, collected, or failed to collect, with sometimes a reference to plants or insects. To read it is rather like spending time in September at Point Pelee oneself, among that special habitat of sand dunes, dense bush with climbing vines, wet thickets and marshy woodlands. Here among many other species they found Carolina Wrens, a juvenile Prairie Warbler, a male [Northern] Cardinal in moult and a King Rail.*° Another day they came across juvenile Carolina Wrens which constituted the first record of breeding in Ontario.*” Swales joined them on 7 September. Although the warbler wave had moved through the Point, never- theless during the next few days, between them, they found: Ruby-crowned Kinglet; Brown Thrasher; House Wren; Olive-backed Thrush [Swainson’s Thrush] and a few Gray-cheeked Thrushes among other birds. They now explored the extreme end of the Point and found the habitat very different. “The large deciduous trees end about half a mile or so from our camp & the whole end of the point is covered with the same red cedar, scrub & juniper that clothes the rest of the west shore. Here we found warblers common again. It is interesting to see how the numbers have var- ied. The day after we came we found the woods alive with warblers. The next day there were as many. The day after they were slightly fewer and today, in the vicinity of the camp, they were all gone but still quite a number [are] left at the extreme end of the point as though they were but leftovers that had reached the jumping off place.”** On the east coast, on 15 September, they collected a bird that they could not recognize. Back in camp, with the aid of Coues’ Key, and much difficulty, they tentatively called it a [Red] Knot, but knew that it would take further study to make certain of their identification. This is a good example of the difficul- ty facing field ornithologists of that period in identi- fying birds with reasonable accuracy. They lacked specially designed pocket field guides with well coloured illustrations that they could carry in the field with them. With glasses still limited in field of vision and clarity of light the serious student of ornithology had to collect specimens in order to bring them home for study with the Key, and to com- pare them with skins in museums and private collec- tions. It is worth saying again that descriptions and illustrations in the high-grade field guides that enthu- siasts use today were developed from the skins, in various ages and plumages, of specimens that were CRANMER-BYNG: A LIFE WITH BIRDS 25 collected at the time Taverner and his friends were exploring and collecting at Point Pelee and else- where. They could not have been written and illus- trated from sight records. A few miles southwest of Point Pelee lies Pelee Island, with a somewhat similar land formation and vegetation. Here Lynds Jones spent a week in the fall - of 1905 making observations on the islands, four days being spent on Pelee Island. He described the migrational conditions as follows: “I found the birds migrating practically everywhere along the line of the islands, but the largest and best defined stream was across Pelee Island, with a well marked convergence to its southern point, thence across to Middle [Island], and beyond to Kelley’s Island, thence across to Marblehead. ... No birds were seen crossing the lake except in a line with the islands.”*? Lynds Jones was invited to join the GLOC and in his letter of thanks for “the privilege of joining with you in this very effective way of keeping in touch with other bird lovers”, said how glad he was to be cooperating with them in their survey of the area. “My studies of the birds and plants of the islands makes it clear that the lake is a great climatic levelling factor upon the portions of land bordering heey What Saunders and Jones already suspected was borne out by the experience of Taverner, Swales and Klugh during the fall migration of 1905, namely that Point Pelee served as a migration funnel in the fall for the much wider hinterland north of its base, while in the spring it again acted as a funnel for a concen- trated mass of birds which then spread throughout the much wider area to the north of its base. To Taverner and Swales this was an exciting discovery, and presented them with a challenge that would require much close study over the next few years. The first task they set themselves was to compile an annotated list of the birds of Point Pelee. Taverner wrote to Saunders proposing that a list of the birds of Point Pelee should be undertaken, and asking for his help and criticism in compiling it.°! He realized this would be a major undertaking, but one that badly needed doing. It was agreed that members of the club should receive recognition for their observa- tions while the actual preparation of the list should be done by Taverner. In a letter to Fleming Taverner wrote that he wanted to make it more than just an annotated check-list by expanding on the biological importance of Point Pelee to naturalists as a northern pocket of Carolinian influence and its importance as a migration route. Taverner presumed that he would be sole editor and that Swales would let him use the observations Swales had made at Pelee during their past visits. But Swales wanted it to be under their joint authorship so that he would receive an equal share of the credit. Taverner wanted to draw a num- ber of generalizations from the check-list which would be his alone. As he frankly told Fleming, from a purely selfish standpoint he wanted to write the list 26 THE CANADIAN FIELD-NATURALIST himself so as to show what he could do. This led to a certain coolness between the two men but there was little chance that their conflicting aims would seri- ously endanger their friendship. They each needed the companionship of the other. Both appealed to Fleming for advice, who suggested the basis for a compromise which they accepted, and their tiff in a teacup soon subsided. Taverner recovered from his disappointment at having to publish the Pelee list under joint authorship, and in The Wilson Bulletin rather than in the better known Auk. Among the various topics discussed in the early issues of the Bulletin was one initiated by Taverner on “The Tagging of Birds”. Before moving to Detroit in March 1904 Taverner had already become interested in the possibilities of learning more about migration and distribution by “tagging” birds with bands on their legs. Although he did not originate the idea himself his practical mind turned to a workable method for applying it as widely and effectively as possible in North America. Stimulated by the new opportunities open to him in Michigan he wrote to the editor of the Bulletin of the Michigan Ornithological Club pointing out that valu- able information might result if birds were systemati- cally tagged. He suggested that if those undertaking field work were to tag each young bird they came across, while older birds were trapped and tagged in the shrubbery of one’s immediate neighbourhood, results could be obtained with the minimum of time and labour. He explained the kind of bands he pro- posed to use for small birds, how they would be marked, and the results recorded. It was certainly a bold scheme and showed considerable optimism on Taverner’s part. In a letter to The Auk he wrote: “Mr. P. A. Taverner, of 95 North Grand Boulevard, W., Detroit, Michigan, proposes to attach small aluminum bands to the tarsus of young birds, in the hope that some of the birds thus tagged may afterward fall into the hands of ornithologists and be reported. The tag, for the sake of brevity of address, will be inscribed ‘Notify the Auk, N.Y.,’ to which any such discoveries should be reported for publication.’>? In this way the American Ornithologists’ Union, through its journal The Auk, was associated with bird banding in America from its earliest stage. Also, in a hand written note dated 30 March 1905 he gave an account of his own involvement in bird tagging to members of the GLOC in its Bulletin, and a summa- ry of attempts to carry out banding in North America to that date. With this he included sketch plans for a brick trap for catching ground haunting birds. Fleming was responsible for using the first num- bered and recorded band in Canada. His Journal for 24 September 1905 reads “In the garden in the after- noon I caught a Robin [American Robin] which I will release with a ring Number 1.”>? Taverner supplied thirty tags to Fleming, fifty to Klugh and twenty-eight to Saunders in May 1905. Vol. 110 But the mere handing out of bands was of no signifi- cance, and it was with a note of pride that he made an announcement in the GLOC Bulletin of January 1906 that a banded flicker [Northern Flicker] had been shot and the tag returned by the finder. This warranted a notification under “General Notes” in The Auk which read: “Tagging Migrants. — In accordance with the scheme outlined in “The Auk,” XXI, p. 410, I have been placing aluminum tags upon the tarsus of nestling birds, and have induced others to follow my example in the work. This past spring several field workers have been using tags supplied by me and this winter the first result has been attained. May 29 Mr. Chas. Kirkpatrick of Keota, Keocuck Co., Iowa, tagged a nest of half grown Flickers (Colaptes auratus) near his home. Dec. 25 Mr. J. E. Ross took No. 123 of this series at Many, Sabine Co., Louisiana. The bird was not saved but I secured the tag from the collec- tor and have identified it as one of my issue ... ” ~4 Between May 1905 and June 1908 Taverner appears to have issued about four hundred bands.°° Though the return address was printed by means of a die the number of each band had to be done by hand. Taverner found that hand numbering was too labori- ous to be practical. When during the winter of 1907- 1908 a bird club in New Haven (Conn.) was orga- nized for the purpose of banding birds, Taverner vol- unteered to hand over his records and bands to the club and to act as an advisory member of its commit- tee on banding.°° At the meeting of the American Ornithologists” Union in November 1908, Dr. Leon Cole called the attention of members to the value of tagging as a means to obtaining information on the movements of individual birds. In this paper he reviewed the work done by Taverner, himself and others to that date.*’ Although Taverner ceased active banding in 1908 another individual effort began near Leamington, Ontario, the following year. The event was recorded in The Auk (1910) by W. E. Saunders under the title “Another tagged bird heard from’. He reported that Mr. J. T. Miner of Kingsville, Ontario, had a number of wild ducks in semi-domestication along with geese and pheasants. In the fall of 1909 his Black Ducks attracted a wild one of that species on 5 August. It soon became very tame and was banded. It was free to come and go, but did not migrate until 15 December. It was shot in South Carolina on 15 January 1910 and the sportsman who found the band returned it to Mr. Miner.*® During his years in Detroit Percy was continually reading and thinking about ornithology. He had read and re-read Darwin on the evolution of the species, so when, in a letter, Fleming claimed that ornitholo- gy, as it then existed, was not a science he forced Taverner to set down his thoughts on the subject clearly. This he did in a letter of November 1905. The work that ornithologists do rarely has great sig- 1996 nificance, he argued. Yet even the collection of data is of value. At this point Taverner launched into an eloquent exposition of science as the key to the plan of creation. “Strictly speaking there is but one science — the science of the plan of the creation. Some study it in the stars and are astronomers, others in the earthly strata and are geol- ogists or in the different forms of life and are zoologists. Only by piecing together the results of all branches will we ever hope to reach the ultimate goal — that of a per- fect understanding of the creation. The advanced studies of one age are the primary ones of the next and it is only by working out the minor details of the broad base of the pyramid that we can ever hope to achieve and locate the capstone above. Our little data help solve questions of distribution and hence dispersal of life. This, when thoroughly under- stood will undoubtedly throw some light upon the origin of life. The great problem then will be the origin of mat- ter. We are attacking it from all points — astronomy, chemistry, physics, entomology, bacteriology, geology, botany and perhaps last of all ornithology. No matter how little the light is that it throws upon the great ques- tion — if it throws any or if it even promises to do so it is a science. I hope I have made myself clear. Perhaps you will see why I am not interested in the so called Life histories of birds and why I am interested in the specula- tive side of things. To my mind the mere collection of data and records with no other end in view is on a par with a collection of postage stamps or tin tags. As it is I look upon every little fact that I record as a possible aid in the solution of a problem that may in an infinitesimal way help to answer what are we, where did we come from, where are we going, and why are we coming or going so? It has been said The only study of man is man. I say the study worthy of man is this creation and if I am only aiding that study in the most microscopic way I feel that I am doing scientific work and ornithology is a sci- ence. Q:E.D. There my mind is relieved. I have lectured all evening in a most pedantic manner but I feel easier.”>” Although these ideas are in no way original at least they show that he was reading about science and creation, and thinking about the subject. Another problem which was much on Taverner’s mind at this time was concerned with nomenclature and the system by which the AOU Check List of North American birds was kept up to date. A particu- larly contentious problem concerned the growing number of subspecies distinguished by taxonomists. Taverner discussed with his friends the harm which he felt was being caused to ornithology by creating too many new races within a species, each with its Own separate name, based on only slight variations. This was a highly technical question and one that a novice should have had the sense to avoid but Taverner did the opposite. He wrote to Dr. J. A. Allen of the Department of Mammalogy and Ornithology at the American Museum of Natural History in New York. Allen was a founder of the American Ornithologists’ Union and its president for the first seven years of its existence. CRANMER-BYNG: A LIFE WITH BIRDS Di He was also the editor of The Auk and was much involved in the problems of nomenclature, and the efforts to stabilize scientific names and the style of the AOU Check List. At this time Allen was fifty- eight years old and was looked up to as perhaps the most influential ornithologist of that time. Taverner certainly went to the top authority in presenting his dissatisfaction with the system of designating sub- species. He must have foreseen the consequences and was either too sincere, or too foolhardy to flinch from his purpose. His letter began with a claim to be presenting the views of ornithologists of the “Middle West” concerning some modifications he wished to see made to the format of the next edition of the Check-list (Third Edition published 1910). He pro- ceeded to make a number of criticisms.*! This was a challenge to the authority and wisdom of the AOU and the working of its committee on the nomenclature of North American birds. Taverner compounded this by implying that the committee had accepted a large number of subspecies which it had subsequently found necessary to discard. Allen replied with restraint while refuting what Taverner had written. This should have quietened Taverner but it did not. He was foolish enough to raise the issue again. This time Allen spoke sharply and put him in his place. “T cannot discuss so large a subject as the law of priority and a statute of limitation in a letter. The best I can do is to send you a copy of the A.O.U. Code and call your attention to the discussion there given of both these top- ics — pp. 32-39. You may have seen this already and if you have it would be doubtless a waste of time for me to say more. Now let me be plain-spoken. I take it that your actual experience in working out questions of nomenclature has necessarily been quite limited. You are disgusted with changes of names and are looking for some easy way to get rid of such annoyances. Do you reflect that hundreds of others, for the last fifty years, have had the same dis- gust and annoyance, and that in the light of a hundred-fold more experience and in every way a much broader equip- ment for the consideration of such questions than you have had, have almost unanimously reached the conclu- sion that the law of priority is the only basis of hope for future stability in nomenclature? No statute of limitation can be formulated that will receive general approval. The matter has been up before and thoroughly considered, with the result that such a proposition has been over- whelmingly disapproved as wholly impracticable.”° Eventually Taverner accepted the law of priority but he never gave up challenging any subspecies that, in his opinion were determined on very flimsy grounds. In this he showed another side of his char- acter — an extreme stubbornness in the face of authority, or as some people felt, a strong streak of perverseness. The Taverners lived in rented accommodation when they moved to Detroit in March 1904. A year later they moved to another address, and six months later to yet another address. Percy did not complain 28 THE CANADIAN FIELD-NATURALIST about accommodation in his letters to Fleming, but he must have felt at a disadvantage when he com- pared his rooms in other people’s homes with his friends who had homes of their own. Both Percy and his mother wanted a garden of their own which they could tend year by year and which would repay them simply by flourishing. But how to achieve it? There was only one way that would quickly solve their problem, and that was to sell the cottage and land on Gibraltar Island in Lake Muskoka. It was a sad decision to make — the three had been going there fairly regularly since 1887 — but it was an obvious one. They now appeared to be permanently settled in Detroit, and since they spent almost the whole of each year living there it made sense to raise the capital for a home of their own by selling the cot- tage and land. This task fell to Percy as a man who had some acquaintance with buying and selling property through his work in an architect’s office. He wrote to Edward Prowse, proprietor of the Beaumaris Hotel, enclosing a plan of Gibraltar Island, and asking if he could dispose of the lots which he had numbered. In reply Prowse said that he might have found a buyer, someone who offered $1300 for the two lots, Point Coo-ee and Kamp Komfort, or $2000 cash for all the island except the lots already sold. Percy made fur- ther enquiries, writing to Fleming and to John Brashear, his friend and neighbour in the Beaumaris cottage community, about his plan to sell. Brashear replied that an advertisement put in any one of the Pittsburg newspapers might bring a purchaser. Eventually the cottage and land were sold for $2500. Percy went to Beaumaris in June 1906 to clear out their possessions, and by taking an early train from Windsor was able to spend three hours with Fleming in Toronto at Spanner’s shop where they “talked ornithology”. Thence to Muskoka wharf birdwatching from the train as he went (he saw a Red-headed Woodpecker at Guilford) and then by steamer to Beaumaris. The next day he went for a ramble in the old familiar area round Camp Comfort. He saw a male Purple Finch in bright plumage which was probably nesting nearby, found the nest of a Wood Thrush with eggs and saw [Dark-eyed] Juncos feeding Cowbirds. One Whip-poor-will was calling in the evening.“ So Percy said goodbye to the only permanent home that he had known up to this point in his life. What his feelings were we can only guess since nothing in his journal nor in his surviving let- ters gives a clue. But he had recently turned thirty- one years old and needed a home where he could plan for a more comfortable and permanent way of life for his mother, sister and himself. With the money from the Muskoka property Percy and his mother were able to buy a house and garden in the village of Highland Park just outside the boundary of Detroit city. The address was 55 Vol. 110 Elmhurst Avenue, Highland Park, Michigan, and the purchase price was $2900. Although closing day was fixed for | July 1906 the Taverners did not get pos- session until early September. This was frustrating since it meant they had to remain in rented accom- modation in Detroit through the hottest part of the summer just when they thought they were moving to the countryside. Percy was doubly annoyed because, as he wrote to Fleming, the move would occur right in the middle of the fall migration when he wanted to be at Pelee. Even after they got possession their problems were not over since they discovered that the contractor had not fixed the locks of the house properly, there was a leak in the roof and the toilet had not yet been installed. From the initial agree- ment it was clear that the property formed part of a new subdivision which was still in the process of completion. The vendor agreed to build walkways round the house and a fence on each side, and to put down sod from the rear line to the street line. Also he agreed to supply a water closet to be placed in the basement.® At last Percy was able to tell Fleming of his change of address which drew a sly witticism from Fleming at Percy’s expense in a letter which began “T trust the statute of limitation will apply to your address: none other to be published for 10 years.’ This was what Taverner also hoped but just as he and his mother and sister were settling into their own home outside the city he received a letter from Fleming, mainly about birds, but ending with this cryptic sentence: “I have a matter I want to talk over with you and hoped to have been able to see you, as it may be of importance but I will have to put off referral to it tonight.”°’ This aroused Taverner’s curiosity and in a letter containing questions about how to start a garden at the new house, he asked “What is it you want to talk over. You say that it may be of importance.”°* Fleming’s answer, when it arrived, was totally unexpected and fell like a bomb in the Taverner household. For the first time in his life Taverner heard that there might be a chance for him to exchange the career of architectural drafts- man for that of professional ornithologist. On a recent visit to Ottawa Fleming had met with John Macoun, and it was at this point that the future career of Percy Taverner came into the orbit of forces shaping the newly emerging Victoria Memorial Museum. Until then Taverner’s only con- tact with Macoun had been by letter containing bird notes that the “Professor” had invited him to send. This had been a discouraging experience for Percy who was upset by Macoun’s apparent laxness in acknowledging his contributions to the Catalogue of Canadian Birds. [See Chapter 3]. Taverner, howev- er, was unaware of the difficulties that the natural history branch of the Geological Survey faced through lack of staff and inadequate funds.°? 1996 In the fall of 1906 prospects for the museum were not very encouraging but at least there was some sign of forward movement. In a letter to Taverner Fleming said that A. P. Low, newly appointed Acting Head of the Geological Survey,”° intended to get rid of its taxidermist, Samuel Herring. Fleming had heard this from Macoun who also told him that: “They want some one to take hold of the museum, not necessarily a taxidermist, some one of sufficient artistic taste and the needful knowledge to arrange the new museum and take the burden off the hands of the present staff. In fact they want a curator but there was nothing said about salary only that the appointment lay between Low and himself and they wanted the right man ...” In the final sentence he said: “Write soon and best tear up the letter light your pipe with it.”7! In his reply Taverner said he realized it was only a pipe dream as yet and might never materialize, and that he and his mother and sister had only just settled in Highland Park. “Though I am a single man I am far from free to do just as my fancy dictates. I am as necessary to my mother and sister as they are to me.” He then stated his own position. “T have always wanted to do museum work and think that I have some of the qualifications for it. If I ever achieve anything it will be along such lines. Architecture will never offer me anything but a living as I do not take enough interest in it to develop any latent talent | might have in that direction. However I am no longer a taxider- mist. I do not care to do a taxidermist’s work and beside were it otherwise the effect of the arsenic upon me would render it impossible for me to do so regularly ... As for remuneration, I know that there is no great for- tune to be made from such positions but of course I should expect anyway as much as J can make draught- ing. Twelve hundred a year would be a minimum but I think that such a sum coupled with the idea of doing congenial work would be a large inducement. Were I alone in the world I am afraid that a smaller sum with a permancy and the thoughts of the pleasure of the work would be more than I could resist.” A few days later it was no more than a pipe dream, when Fleming received a letter from Low stating that what the Geological Survey wanted was a practical taxidermist; that in those circumstances Taverner would not be suitable for the position.’* In a letter which shows Percy’s trait of modesty (some might call it lack of confidence) he wrote a detached self-appraisal. “Now in the first place you overrate my ability as an architectural designer. I am not a good designer. I have done some good little things but they were more the inspiration of the moment than a good criterion of my general ability. No one knows my failings in this line bet- ter than myself. In the steady grind of office work I am not there with the goods. I am a good draughtsman and do not think that I need be out of a position for any length of time. Neither am I good for the outside work — I am the worst handler of men possible nor can I hustle for business. So I can therefore see no prospect of success in any business of my own or in partnership in any. As CRANMER-BYNG: A LIFE WITH BIRDS 29 things are I do not want it either. | am able to make a comfortable living as it is and can expect a slow perhaps but steady advance in salary without the worry and fret of business responsibility. My present employer is a slave to his business — I don’t want any of that. I intend to get as much out of this life as I can and do not think that I can do so in a race after money. All I want is enough to live on and that I think I can get. Were I a married man I should of course look on things differently.” He agreed with Fleming in the matter of clothes, that he should pay more attention to his appearance. He admitted to absolute indifference to what other people thought of him, and confessed that he was bored by people from whom he could not get a sin- gle intellectual spark.” Life in the Taverner household resumed its nor- mal course. Percy could turn his mind “back to the drawing board” (literally) while finding a very con- genial outlet for his surplus energies in the satisfac- tion he obtained through ornithological projects and, from now onwards, in gardening. Mrs. Taverner could look forward to indefinite time for herself and her family in the new home, as well as in creating a garden of their own. Since Percy had no previous experience of gardening he relied initially on advice from friends, especially from Fleming whose practi- cal experience was useful.” As for Fleming’s sound advice on getting a better job in architecture this may have had some influence on Percy in spite of his strong views on not wanting to get into a “race after money” and not being “a slave to his business”. From what scanty information exists on his professional work it appears that he changed firms in 1906 and, that by the time of writ- ing his letter of self-appraisal on his ability as “an architectural designer’, he was working for William E. N. Hunter who had an office in Detroit from 1906 onwards. Taverner is listed in the Detroit City Directory of 1906 (dated August) and in the 1907 Directory as working for W. E. N. Hunter. This was the man he referred to as “a slave to his business’’.’° For Percy Taverner his first two-and-a-half years in Detroit marked a considerable change in his way of life. These years were ones of much wider scope and deeper satisfaction in his chosen recreation of amateur ornithologist. His life now was full of inter- ests and opportunities thanks to his passion for bird study. By the end of 1906 he had made some progress in his bird banding scheme, had carried out collecting and record keeping far more extensively and systematically than previously, and he was working on a major study of bird migration and dis- tribution in one unique geographical area of the Great Lakes. Also by late 1906 his home life was more comfortable and satisfying than at any time previously. As well, he had caught a glimpse of what he might possibly achieve in the future, with good luck and hard work: a professional position in a nat- ural history museum. The challenge he now faced 30 THE CANADIAN FIELD-NATURALIST was to produce a solid and worthwhile bird study which would be published in a good ornithological journal, and to extend very considerably his own knowledge and experience in the world of ornitholo- ‘ gy. The next four years at Detroit were critical for Taverner and his future career; they were what might be called his “ornithologist-in-the-making years”. The Great Lakes Ornithological Club gave Taverner, Swales and a few others a reason to visit Point Pelee regularly, to keep systematic records of their sightings and collecting, as well as to “write up” material in the Bulletin.’’ One obvious by-prod- uct was recording species not previously known at Pelee and rare in Ontario. For instance Fleming, on one of his infrequent visits to Pelee, collected a Chuck-will’ s-widow, the first record for Ontario.’® Taverner was responsible for the first Canadian record of a Blue-winged Warbler, taken there in September 1906”? ,while the first [Northern] Mockingbird recorded at Point Pelee was also col- lected in 1906.°° The movement northwards of the [Northern] Cardinal into Canada from the American side of the border had hardly started at the beginning of the century, but the records kept by members of the Club allowed the authors of “The Birds of Point Pelee” to claim, in 1907, “Point Pelee and its vicinity boasts of being the only locality in the Dominion of Canada where the Cardinal is regular and com- mon.”®! The first record of the Bewick’s Wren at Point Pelee dates from 1909.°*? Although Taverner was not able to visit Pelee more than a few times each year he obtained useful information and a succession of specimens from Bert Gardner, a resident wildfowler on whose land mem- bers of the GLOC camped and where, in 1908, they built their shack.** One spring Taverner brought back from Pelee an injured male Wood Duck which Gardner had saved. Writing to Fleming he said: “It is doing well and is the gentlest and prettiest little pet you ever saw.” At the end of the letter he wrote: “Have just given my Wood Duck a bath in the tub and you should see how he enjoys it.””*4 Percy had kept some pets while living in Port Huron but it was only now, when they had a home and garden of their own, that he could keep birds as pets. In another letter to Fleming he wrote: “I have a tame Sparrow Hawk [American Kestrel] now. He is very tame and will stand all the handling we want to give him. His favorite perch is on my head but it is not quite safe to let him stay there. He has never been house-broken.®° Although Taverner shot birds as specimens for his study collection, or for exchange for other speci- mens, there is no evidence that he enjoyed shooting them as a sportsman. Nor was he free to shoot birds whenever he liked, and at Point Pelee he had to have a permit ready to show the game warden. To obtain a permit, he needed a form signed by two people certi- fying that he was a proper person to collect for scien- Vol. 110 tific purposes. Writing to ask Saunders to endorse his application he reported: “For the first time in my life I had to show my permit to the game warden. Someone in Leamington had com- plained on us and he had to question us. He did not notice, however, it had expired — lucky for us.”°° Swales, also, had been in the habit of getting Fleming to endorse his application, but the warden refused to issue him one. “If Taverner can have one over here I don’t see why I can’t especially as I have considerable property in Canada.”8” His feeling of injustice was all the more because Taverner had been issued a permit to collect for sci- entific purposes in the State of Michigan by the game warden stationed at Sault Ste Marie.** But the matter did not end here. The game warden tackled them at the Point in the fall of 1907 and warned them against shooting at a mark even, on Sundays. Taverner explained the reason to Fleming. “I think some of the people in Leamington complained on us. They cannot understand any one preferring to shoot small birds instead of ducks. I imagine that they think that we used the permit as a cloak to other shooting.”®? From the correspondence and records that have survived among the Taverner, Fleming and GLOC papers in the Royal Ontario Museum one can get an idea of Taverner’s continuous ornithological activi- ties. From the fall of 1906 there was the immediate vicinity of his new home to explore, as well as regu- lar expeditions with Swales to such places as St. Clair Flats, the Ecorse mud flats and the Huron River at Rockwood. Weekend excursions were made to Point Pelee with Swales several times a year, as well as the annual camps there. Although telephone com- munication at this time was not sufficiently devel- oped to make possible the equivalent of the birding “hot-line” of today it was easy and quick to send a postcard with a useful tip-off of recent arrivals. For instance Saunders sent Taverner a card from London in November 1906 which said: “Look out for W. W. Crossbills [White-winged Crossbills] on Hemlocks. I got 9 out of 50 yesterday.” It cost one cent to send and reached Taverner at Highland Park, Michigan, the next day.?? Not only was Taverner very busy collecting birds, dissecting their organs for information, and making notes of what he observed — what he referred to as “field work” — he also spent time regularly on writing up his Journal from notes, and keeping up his species ledger. His journal and species ledgers are still of value for information they supply on distribution of particular species. For instance, a species that is now a very rare permanent resident in southwestern Ontario — the Bobwhite — was even then subject to fluctuations in number. After birding in March 1907 with Swales he noted in his Journal that the most interesting species seen was Bobwhite. They saw a covey of twelve. These were the first quail that he had seen in the county of Wayne since he arrived in 1996 CRANMER-BYNG: A LIFE WITH BIRDS 31 a Taverner preparing a skin in the Great Lakes Ornithological Club “shack” at Point Pelee, 30 May 1909. Photograph by Detroit News Tribune (now Detroit News) 27 June 1909. (Reproduced from print courtesy of the Royal Ontario Museum, Toronto.) 32 THE CANADIAN FIELD-NATURALIST the spring of 1904. The winter of 1903-04 pretty well exterminated them in southern Michigan.?! We can also learn a little about conditions of life on Point Pelee from his Journal. There was a lifesaving station on the west side of the Point. In November 1906 a ship was driven ashore in a storm and the local Life Saving Corps took eighteen people off the hulk.’” The tower near the tip of the point was used not only as a lookout but also as a very convenient place from which to view the fall migration. Fleming was very concerned about the effects of alien species on native species in North America. He was especially worried about the spread of [European] Starlings and wrote: “I see Bird Lore is discussing the Starling question. The harm has gone too far to stop and I foresee the passing of the [Eastern] Bluebird. Urban ornithology of the future will consist of English Sparrows [House Sparrows], Starlings and possible Jackdaws. The Swallows and Martins will go.””? An overly pessimistic forecast. From what both Taverner and Swales wrote they were both careful not to accept records of birds seen without supporting evidence. Taverner was suspi- cious of the reliability of John Clare Wood of the Michigan Ornithological Club,* while Swales was very critical of Lynds Jones’, “big days” when he and one other would claim to have seen as many as 130 different species without any proof to show for it.°> Taverner’s point of view was clearly stated in his Journal when he wrote that he had shot a Carolina Wren, the first authentic specimen for the county, and one of the first for the State of Michigan. “In this respect I class nothing as absolutely authentic unless the specimen has been taken and viewed by some one of undoubted authority.”° He regarded taking a bird’s life as a necessity when a specimen was required to authenticate a record. One other use of his Journal was for him to record ideas and problems which he considered needed fur- ther study. For instance while at Point Pelee in early 1907 he collected two very pale-coloured Great Horned Owls. He was already aware of the wide dif- ference in the colour of plumage of the owls, and now, looking at them closely, he wrote to Fleming: “T think that the distribution of the Horned Owls will have to be studied with breeding birds. The[y] migrate largely and in the winter the forms must be pretty well mixed up. There seem to be but few breeding birds in collections.” In other words it is not possible to make a reli- able study of the breeding distribution of a species or particular form (e.g., western or northern Great Horned Owl) in a region by using records from only winter months or during migration. They need to be based on records made of breeding birds of that particular region.”” During this period ornithology had become, for Taverner, a serious undertaking, a semi-professional Vol. 110 occupation more than a recreation or a hobby. Taverner’s ambition was to write a major paper that would make use of the material he had been collect- ing since coming to live in Detroit. He wanted to prove that he could write about birds in an extended work, and he wanted to have that work published in a journal where it would be read by the majority of serious ornithologists. The subject he had in mind was an annotated list of the birds of Point Pelee; the journal he aimed for was The Auk. Before examining “The Birds of Point Pelee” it may be helpful to say something about Taverner’s ability to write well at this time, and also to discuss his collaboration with Swales as joint authors. Two extracts from his Journal will show his powers of description. Here is a female Wilson’s Phalarope observed through glasses in May 1906. “Tt floated very high in the water and glided about with- out any signs of effort striking right and left with its graceful straight bill like an expert fencer handling his rapier. It seemed to be continually picking up food from the water and its head was constantly going as it struck with a thrust-like action of its bill on either hand.” He then described its plumage with the diagnostic marks. “T saw the dark of the back of the neck shading into red on the sides and with the pearl grey of the median stripe on the head — I watched it for nearly half an hour ....”?° By contrast here is a flock of Semipalmated Sandpipers in flight. “As a clock they flew as with one mind wheeling abso- lutely together and wheeling at the same angle on the turn. One instant there would be a mass of scurrying brown dots looking like a bunch of mosquitos in the dis- tance then the next moment they would wheel and the brown dots would simultaneously flash into snowy whiteness in the sunshine. In such times the glistening underparts would fairly dazzle the eye with their white- ness. Often they would slow down and lower their eleva- tion and J expected they would alight but no they sailed past the seeming destination and were wheeling again and making for the other end of the place.” When finally they landed he described them feed- ing.’ A different style of descriptive writing is used by Taverner in his article “The Yellow-breasted Chat. A character sketch”, which was published in Bird Lore in 1906, with a drawing by Taverner of the bird during its flight song. The style here stands out as anthropomorphic and literary; completely out of keeping with the aims of present day ornithology. “The Chat is a will-o’-the-wisp, it plays hide-and-seek with you just out of sight behind the foliage, it taunts you fearlessly with its private opinion of you, your family, and all your ancestors . . .” and much more in this vein. It is intuitive, arch, deliberately playing for effect.! But we need to remember that Bird-Lore, founded in 1899, was dedicated to the study and protection of birds, and was intended mainly for Audubon Society members. Taverner’s perception of the average Audubon member at that time was coloured by Fleming’s view of them as a bunch of elderly ladies 1996 who tended to “gush” about the dear little birds.!°! Taverner was anxious to have his article and drawing published so he wrote it in his literary style. When Frank Chapman’s The Warblers of North America was published in 1907 he quoted from Taverner’s description of the chat’s flight song.!° In addition to an occasional “purple passage” Taverner’s style was marred by a tendency to be too long winded. A fellow member of the GLOC, William Brodie, expressed it neatly when he noted on one of Taverner’s frequent contributions to the Bulletin: “Friend T. may remember sugar making in Muskoka. If so he must know the necessity and virtue of boiling down.”!93 If Taverner had been trained at a university he would probably have had these tendencies towards over-writing eradicated. Instead his friends and acquaintances performed the same service. Another side of his written style can be seen from the many letters he was constantly writing to his friends. Here the style is usually colloquial, written as he might speak, often with a bubbling sense of humour, the equivalent of a twinkle in his blue-grey eyes which those who met him remembered. Percy was impeded in expressing his ideas verbally by his stammer. There was so much that “turned him on” in the natu- ral world, so much that he wanted to communicate to others, that he was forced to compensate for his stammer by setting it down at length in writing. His letters and bird journals were his way of expressing his sense of excitement and appreciation of the beau- ty of the natural world and some of the unexplained ways in which living things interacted. Swales and Taverner went together to Point Pelee for the 1907 fall migration from 24 August — 6 September. Their friend Norman Wood, from the Natural History Museum of Michigan University went with them. Saunders joined them “bringing with him Mr. J. S. Wallace of Toronto, a beginner in the bird business but an experienced camp man who proved to be quite an acquisition in more ways than one.”!°* Taverner and Swales both kept bird jour- nals. Comparing them for style and content one sees a wide difference. Swales’ method was compressed and entirely factual, and dry to read.'° Taverner’s method was fluid, responsive to people and ideas, more lively to read. During their stay they were eaten alive by mosquitos which prevented them get- ting much sleep, and the weather was very hot. Discussing the value of Point Pelee as an observation station in correspondence with Fleming Taverner wrote: “Yes Pelee would make a great station for a season’s observations . . . I think that with a very little continuous work Point Pelee could be made as classical a ground as Helegoland almost . . . If I were a good talker I think I could get some of our wealthy men to finance some sus- tained work there. Think what a small amount of money CRANMER-ByYNG: A LIFE WITH BIRDS 33 would do there. Maybe the chance will come some day.”106 Now we can take a critical look at the contents of the work. The first issue appeared in The Wilson Bulletin of June 1907 and consisted of a twelve page introduction and an annotated list of waterfowl. The authors gave credit in separate paragraphs to each member of the GLOC from whose notes the report ‘was mostly compiled, as well as to Mr. Albert Gardner whose information on various birds they found reliable. His information on waterfowl was particularly valuable because it was difficult to gath- er waterfowl data on short and unsystematic visits.!°7 A physical and ecological description of the area included information on vegetation, and a paragraph by A. B. Klugh on its Carolinian aspects. The impor- tance of Point Pelee as a funnel for migration across Lake Erie was also discussed.!°8 Taverner and Swales were fully aware that the list was based on relatively few visits. This was a preliminary survey, one that should be updated as more data could be collected. The study is valuable because it provides a base line of information against which recent information on the birds of Point Pelee can be measured. It sup- plied information on bird distribution — what birds might be found on migration with some indication of abundance, approximate dates, and sometimes loca- tion. For instance Connecticut Warblers were seen in late May and early September. In September 1905 nine individuals were seen during the first ten days, while in the first three days of September 1906 Connecticut Warblers were almost common. “They haunted the damp tangle bordering the eastern beach near Gardner’s and along the cross-road, and were still more frequently met with in the beds of jewel- weed, closely adjoining, in the open spots of the woods. By remaining quiet in such places we were able to observe this interesting species at will.”!° Other species about which there is distributional information are Golden-winged Warbler; Northern Parula; Prairie Warbler; Louisiana Waterthrush; Bald Eagle; [Northern] Cardinal; Dickcissel; and Eastern Bluebird. The descriptions of the behaviour of several species are particularly vivid and show Taverner’s powers of making a bird come to life through words. Some examples are: Sanderling; Piping Plover; Sharp-shinned Hawk to which four pages were allotted to describe its fall migration; Peregrine Falcon with a first hand description of one striking down two Blue Jays to the left and right in a flock; the carefully observed “song” of an Eastern Screech Owl; Ruby-throated Hummingbird’s feeding characteristics; and Lincoln’s Sparrow’s elusiveness. The most impressive display of Taverner’s word painting was devoted to the fall migration flight of Purple Martins at the Point in 1907.!!° The final part of the Pelee list appeared in the September 1908 issue of The Wilson Bulletin and 34 THE CANADIAN FIELD-NATURALIST contained considerable additional information. Short visits in 1908 at the beginning of May and in mid- August added further knowledge especially about migration. The authors were tempted to generalize about the “wave-like” formations of many of the migrations. Birds that are not normally regarded as gregarious came together at times in such numbers as to constitute a wave and were drawn together not by any community of interest. Rather their gather- ings were the result of each individual, acting in response to common conditions, making for the same crossing place of the lake and arriving simulta- neously. Such “waves” or “flights” were recorded in the list and dates of these were summarized at this point. For example: Sharp-shinned Hawk — 1905, 10-17 September; 1906, 15-22 September. Reactions to the completed study were muted, probably because its publication had taken place in five issues of a journal extending over a period of fifteen months. Presumably the reason why Lynds Jones, the editor of The Wilson Bulletin accepted the study and allowed it so much space was that his own research was closely involved with bird migra- tion through Point Pelee, and the journal he edited was mainly focused on the northeastern United States. To make a real impact it would have been much better if such a substantial study (76 pages) could have been published as a booklet. Taverner and Swales were aware of this and arranged to have a number of copies bound together with strong paper for presentation to some of the more impor- tant ornithologists and for the exchange of separates with other authors. However, it received a favourable, short review in The Auk by J. A. Allen, who said, among other things, that the physical and biotic conditions of the locality were described in detail, followed by an extensively annotated list of 209 species identified as occurring in this limited area. Supplementary notes followed, including comments on hypothetical migration routes. The only other review it received was in the Windsor Record Daily (evening edition) of 27 February 1909, under the headline: “Birds of Point Pelee: most interesting Ornithological Centre in Canada.” One friendly but firm critic of the work was J. H. Fleming. Writing to Taverner early in 1909 he men- tioned that there were several things he had wanted to talk over with him. “One was your recent literary efforts in which I noticed a lapse that you ought to check before you get into the habit of writing like a Sunday special. I am anxious you should keep to your good old style.”!!! To which Taverner replied: “Your criticism on my new style is severe. I was unaware of my lapse. Do you refer to the Wilson Bull. stuff? I thought I had avoided mushiness though making it light for general interest. Will have to watch out in the future.”!!? Vol. 110 Whatever criticisms can be made of his first major piece of ornithological writing, such as the fiction that a bird has a “personality”, one thing stands out. Taverner had an extremely observant eye, a remark- able ability to concentrate on a single bird or flock of birds that he was observing and to describe it vividly. With a draftsman’s sense of form and movement he had the ability to transform personal impressions into word pictures informed with feeling. Above all Taverner loved birds — their calls, songs, movements and their intricate and beautiful plumage. He used a sensitive prose in order to transform what he experi- enced so warmly himself into sharp images so that others might share these experiences with him. The adoption of an anthropomorphic viewpoint from time to time was used by other naturalists in this period — Ermest Thompson Seton was an outstanding example — and was a way of communicating with a reading public which felt at ease with such terms. Early in 1906 Fleming was working on an anno- tated list of birds known to have occurred in the Toronto area. No such list had been compiled before and it posed a number of problems, outstanding among which was the status of the Trumpeter Swan in southern Ontario. There are two species of swan native to North America; the Trumpeter and the Tundra* but it is difficult to distinguish between them with certainty. Size, weight, shape of bill and position of nostrils, the presence of a yellow spot in front of the eye in the Tundra but not in the Trumpeter are indications of species, but are not conclusive evidence. Positive identification of these two swans is provided by differences in convolutions of the windpipe in the breast bone. In the Trumpeter Swan the windpipe makes a high vertical loop over a bony hump in the sternum, while in the Tundra Swan this loop is absent.'!? At the time of which we are writing these distinguishing marks were known but few ornithologists had any first hand experience with Trumpeter Swans either as preserved specimens, or as living birds, because of their extreme scarcity.'!* Fleming and Taverner each spent many hundreds of hours searching the literature, checking old skins and examining the sterna of recently killed swans, not to mention writing to each other. The search for gen- uine Trumpeter Swans became an obsession for both men — something of a “wild swan chase”. Taverner was optimistic enough to consider that specimens of Trumpeters might still be found, and set out to exam- ine recently killed swans. His experience in taxi- dermy and in making drawings on linen (blue prints) as a draftsman now came in very useful. When he started he had little knowledge of swans, only a strong urge to prove the identification scientifically of each bird he examined. *Formerly called the Whistling Swan. 1996 « sel) Sh sar Aa CRANMER-ByYNG: A LIFE WITH BIRDS 35 Five members of the Great Lakes Ornithological Club at club’s “shack” near the end of Point Pelee, 3 October 1909. Left to right: J. S. Wallace, B. H. Swales, W. E. Saunders, J. H. Fleming, P. A. Taverner (seated on steps). (Reproduced courtesy of the Canadian Museum of Nature, number 60386.) After two years of searching the two. men had made some progress but realized that they would have to make a major breakthrough if they were to write a satisfactory study of the swans. Fleming put his hope in finding a specimen of a Trumpeter in a private collection while Taverner pinned his faith in collecting a lot more sterna. By a stroke of luck for Taverner this is just what happened. On March 15th 1908, on a misty morning with a wind blowing downstream, the water below the Niagara Falls on the Canadian side was suddenly full of struggling swans. At least 125 birds were dragged out of the water while many more were swept downstream.!!> Eventually Fleming secured thirty-three swans which were taken to Oliver Spanner’s taxidermy shop in Toronto. A series of measurements were made including weights, and Taverner took drawings of all the variations in the beaks. Taverner included in his Journal for this period a short note on each bird to add to his growing collection of swan data.!!® When they had all been skinned not one of the thirty- three dead birds was found to be a Trumpeter Swan. Fleming and Taverner had devoted a considerable amount of time over a period of nearly three years to a comparison of the two swans native to North America. In the end Fleming discovered a hitherto unknown specimen of a proven Trumpeter Swan in Toronto which eventually was added to his collec- tion, and is now preserved in the form of a skin in the Department of Ornithology at the Royal Ontario Museum.!!’ Taverner developed skill in drawing soft parts of birds and their bills. The experience he gained with Tundra Swans was useful to him in future years. Although Taverner had got the worst in his corre- spondence with J. A. Allen over the revision of the AOU Check-list he was too stubborn to give up the struggle. An editorial by J. Grinnell in the Condor in 1906 on “Better Vernacular Names” spurred him to try again.'!'® This time he was on slightly firmer ground because he and Swales were working on their annotated list of the birds of Point Pelee and were frustrated by problems of nomenclature they encoun- tered. They blamed these difficulties on the AOU Check-list for what they claimed were too many ill- considered changes. Taverner sent Grinnell an article suggesting reforms that he felt should be incorporat- ed in the revised Check-list which was being pre- pared. His article was published in 1907.'! Since it contained specific suggestions which were “in the 36 THE CANADIAN FIELD-NATURALIST air” at this time it called for an answer. This task fell to a member of the AOU committee for the revision of the Check-list, Witmer Stone, who wrote giving various reasons why Taverner’s suggestions, which had been considered by the committee in the past, had not been adopted. Most members agreed that the system needed improving but they were unable to agree on what should be done.!”° This was of no help to Swales and Taverner who had to wrestle with problems of nomenclature in their attempts to main- tain uniformity in their own list of the birds of Point Pelee. Taverner was now thirty-two years old, very much a local man who had not yet made his mark and was still not a member of the AOU, only an associate. He was lacking in knowledge of what had already been proposed and discussed as regards Check-list revisions, and he was lacking in tact. He was rather like an immature eagle standing on the edge of the nest, flapping its wings vigorously, impa- tient to fly, but not yet able to take to the air. Amateur ornithologists may not have the laborato- ry training and equipment, nor the status in the world of professional ornithologists to contribute much to nomenclature. But they can and do amass valuable collections of data in their own area when they keep records regularly and methodically over a number of years. The activities of the GLOC stimulated some of its members to keep systematic records, and to discuss in the Club Bulletin the status and distribu- tion of various species in the southwestern region of the Great Lakes. For instance useful records were kept during the six-year period 1905 through 1910 by Fleming in Toronto, Saunders in London, and by Swales and Taverner in Detroit and vicinity. Also the records of the GLOC kept for sightings at Point Pelee represented the joint efforts of its members. !*! Taverner’s own records were systematic and extensive. His manuscript Journal (in typescript from 1906 through 1910) contained descriptions of visits to Pelee and various places in and around Detroit. The Journal gives a very readable account of his activities, ideas and projects during these years. In addition he kept a list of first arrivals in southeastern Michigan and southernmost Ontario for 1907 and 1908; and a record of fall migration in the same area for 1907 with birds seen at Point Pelee marked with a P. He also kept eight volumes of “Notes and clip- pings about birds” by species. These were typed and dated; some gave a note of the place where a bird was seen or collected, and a reference to where information was published when relevant. The car- bon copy of his Journal he cut up and pasted into his “Notes and Clippings” books.!** In this way he had information on individual species quickly available, not only from his own records but also from other sources. Vol. 110 Point Pelee, as an exciting place for field studies by naturalists, received publicity for the first time when a Detroit newspaper printed an illustrated fea- ture article in its Sunday Magazine section in 1909. At the end of May Taverner had gone to Pelee with two friends from Detroit — Dr. William Newcomb whose special interest was in butterflies and moths and Bryant Walker, an attorney, whose hobby was the study of shells. The News Tribune sent a reporter and a photographer to interview them enjoying their recreation. Taverner noted in his Journal: “About ten o’clock a rig drove up with a delegation from the Detroit News Tribune composed of a reporter and a photographer. We posed for them and told them all the stories that we could think of. They left about one o’clock.”!23 Writing to Fleming in June he told him: “T took Dr. Newcomb lepidopterist and Mr. Walker con- chologist with me to the Point and they enjoyed it very much. Monday 3lst we were visited by the representa- tives of the Sunday Detroit Paper who photographed and questioned us to their hearts content. Guess they are going to give us a big splash. I have copies of all the pic- tures they took and they are very amusing and interest- ing to us as they show us in all phases of our field work both inside and out of camp.”!*4 The title of the article “The Birds’ Jumping Off Place” came from Taverner. The same phrase was used by him in his Journal of September in 1905.!*° The Detroit newspaper article contains much that is straight from Taverner and can be traced to his Journal, with some contributions from Newcomb and Walker. The reporter, Henry Richmond, explained correctly the significance of Point Pelee for seasonal migration, and the part being played by the members of the GLOC in studying how it took place. He also explained why naturalists came regu- larly to Pelee. “So attractive was this jumping-off-place of the birds found to be, that a number of years ago five naturalists established a club house there and began to make period- ic visits to the point for the purpose of collecting speci- mens, studying the habits of birds and comparing notes. These men are P. A. Taverner, of Detroit; B. H. Swales, of Grosse Island; J. H. Fleming of Toronto; W. E. Saunders, of London, Ont., and James S. Wallace, of Toronto. All these men are enthusiastic bird students and the results of their researches have added much impor- tant matter to the literature of their science. These men pursue their scientific hobby as a mere matter of recreation and without a thought of making it con- tribute to their livelihood. They are bird lovers who seek every opportunity of slipping away from the prosaic rou- tine of life to catch an esthetic uplift from a glimpse of the freedom of the air and the tree tops. And so it hap- pens that once or twice each month the week-end or a holiday finds one or more of them at the little club house half-way down Point Pelee, picking up scattering [sic] bird records and revelling in the delights of the out-of- doors.” !76 The article is filled with sound information but told in a lively enough style to interest Sunday 1996 readers in the city. A fair part of it is quoted directly from Taverner’s own descriptions — of humming- birds feeding on jewelweed and then setting out to cross the lake, or the vocal courtship of a pair of Eastern Screech Owls. At the end of the day “each man has his story to tell as they gather around the long deal table in the little club house at night and make up the notes of the day.” The photographs are good and are a valuable source of evidence of what the shack looked like, and the three men interviewed, as well as their equipment. Also included in the article are two sketch maps, one of which shows Point Pelee, Pelee Island, Kelly’s Island, and Marblehead, Cedar Point and Sandusky in Ohio. All in all this was an ambi- tious attempt to explain to non-specialist readers why so many birds use Point Pelee for the spring and fall migration. Taverner’s interests in natural history had always been wide and he would sometimes collect creatures other than birds to show to acquaintances knowl- edgeable about mammals, reptiles or insects. One day when birding in Detroit he and Swales met a man collecting insects whose name was Arthur Andrews.!*’ As a result Taverner’s alert mind became interested in moths and butterflies, and his Journal for 1908 and 1909 mentions collecting moths at night near his home, as well as at Point Pelee. Through Andrews Taverner met a few other amateur naturalists including William Newcomb and Bryant Walker. A club was started which provided Taverner with a fresh outlet for his energies. As he explained to Fleming it consisted of five members. “We meet very informally at each other’s houses and have no organization, officers or anything else. We just meet and chat as the spirit moves us. Optimistically Percy continued “We are instituting a general biological survey of S. E. Mich. territory, each in our several lines .... All are good serious workers and no dilettante need apply.”!*8 The sight of monarch butterflies massed on the trees at Point Pelee in the early fall excited Taverner’s naturalist instincts and he wanted to know why they were there at that time in those numbers. Since no one appeared to have written on this phenomenon at Pelee he wrote a note himself and sent it to the Entomological News. It was a detailed description of what he saw on visits in three consecutive Septembers. He con- cluded by saying that whether the monarchs were all migrating in the true sense of the word or not, he left to the entomologists to decide. !*? Although Taverner was very careful not to put forward any hypothesis in this note he showed what he had in mind when writing to Fleming. “Am glad you liked the butterfly paper. I thought it was worth while. It is an example of the danger of too close clinging to one line of work. We bird men are too apt to discount instinct and believe nothing than cannot be explained by experience, example or acquired knowl- CRANMER-BYNG: A LIFE WITH BIRDS 37 edge. Yet here are these butterflies shewing all the phe- nomena that we are accustomed to explain by memory and example that have certainly never travelled the road before ... I have often thought that in nature the seasonal changes are to a certain extent automatic and that there is a strong tendency towards the reoccurence when cer- tain dates are reached irrespective of the weather condi- tions. In other words, the seasons tend to run by calen- dang e Taverner had no laboratory in which to conduct experiments with birds at migration time, no univer- sity to back him with the necessary funds. His insight that seasonal changes in birds tend to keep more to the calendar than to weather conditions might, under more congenial circumstances, have led him to examine the effects of the lengthening and shortening of the days, in other words of the effects of light, on the bird’s biological mechanism. Another problem of migration interested Taverner at this time. When he was living at Port Huron in the period 1899-1902 he found a fair number of Saw- whet Owls each fall which made him wonder if this was simply a local movement at the end of the breeding season, or a larger-scale migration. However, when Saunders reported the migration dis- aster that occurred on the shores of Lake Huron north of Sarnia in mid-October 1906 and he found twenty-four Saw-whet Owls drowned, Taverner noted in his Journal that it proved this little owl was a migrant. However, further proof was required before it could be established as a fact beyond doubt, and what better place for such a record than at the tip of Point Pelee. On the weekend of 14-16 October 1910 Saunders, Taverner, Swales and Wallace were bird collecting at the Point and saw several Long- eared Owls in the red cedars. Two were shot for specimens and the contents of their stomachs exam- ined. One contained feathers of a Saw-whet Owl. Taverner and Swales started searching for Saw-whet Owls and found ten in the cedars perched close to the trunks, and six feet from the ground. At this point Taverner returned to camp to collect his camera which was a substantial one with a bel- lows and mounted on a wooden tripod. He went back to where Swales had one Saw-whet under observation. Because the bird was too high to photo- graph with only the existing tripod Taverner impro- vised additional legs which he spliced onto the orig- inal ones. The result was rickety but it enabled him to raise the camera lens above most of the leaves and twigs. By standing on tiptoe he was just able to focus the camera, and stopping down to about US 64 he took a six-second exposure. Swales tried to see how close it was possible to get to the bird when it flew, but only a few feet away, and alighted in an excellent spot for photography. The additional legs were hurriedly refastened and the camera set up again at a much closer range, as Taverner noted in his Journal: 38 THE CANADIAN FIELD-NATURALIST “.. in fact the bird was as large on the ground glass as the length of the bellows would allow me to focus. I stopped down to U.S. 128 and as the bird was in the sun gave it another six second exposure.””!?! Eventually an account of this episode by Taverner and Swales appeared in The Auk. The final para- graph of the article summarized their conclusions. : “Here, then, are records of four migrational massings of this hitherto supposed resident owl. It was too early in the season to explain their gathering as ‘winter wander- ing in search of food’, and the close tallying of all the dates point to the conclusion that from the middle to the end of October the Saw-whet Owls migrate in consider- able numbers, but from their nocturnal habits and secluded habitats while en route are seldom observed. In all probability, too, such noticeable gatherings are only to be observed in such places as at Point Pelee where a constricted migration route brings many together at one time. Long Point is another place much like Pelee in this regard, and the Lake Huron episodes likely originated in other fly lines across that body of water and of which we as yet know nothing. !32 The photograph taken in good light was remark- ably successful considering the difficulties. Taverner was so pleased with the result that he printed copies on postcards and sent them to his friends. By now Taverner was a reasonably skilled pho- tographer. He was also a skilled enough draftsman to make bird illustrations and line drawings for the book on birds of Michigan which Walter Barrows had been writing for so long. Writing to Fleming in January 1910 he said that he was working on a lot of heads and legs for Barrow’s Bulletin. In the same letter he mentioned that he had recently collected together all his old bird pictures and was mounting them on heavy brown paper and classifying them into a portfolio, and was surprised at finding he had so much good material.!** Barrows’ Michigan Bird Life was published in 1912. In the preface he acknowledged Taverner’s contributions. “To Mr. P. A. Taverner I am indebted not only for hun- dreds of field notes on Michigan birds, but for the origi- nal drawings or actual electro-types from which thirteen of the full page plates and fifty-eight of the text figures have been made, the latter including almost all the detail[ed] drawings of head, bills, wings, feet and tails used in the Keys and elsewhere.” !*4 The book ran to 757 pages of text. Taverner was in good company as regards illustrators; Ernest Thompson Seton and Louis Agassiz Fuertes were the main contributors, with Frank M. Chapman supply- ing a number of bird photographs. Taverner supplied wash drawings for American Bittern, Sora, and Hudsonian Curlew [Whimbrel] among others. The list of pen and ink drawings is formidable. Percy was modest about his ability as a bird illustrator, while Barrows said that his prices were too modest. He received $10.00 a piece for the wash drawings. Taverner told Barrows about the Saw-whet Owls at Point Pelee who envied him the experience but used Vol. 110 a photo of a Saw-whet taken by Frank Chapman which looks as if it was of a dead bird posed on a tree stump. He would have shown a much more life- like photo if he had chosen Taverner’s. If Percy was ever to become well enough known to obtain a position in a museum of natural history it was important that as many of his ornithological records as possible should be accepted and pub- lished. Publication of his records in “Birds of Point Pelee” and Michigan Bird Life was useful as a begin- ning but from the perspective of obtaining a post at the museum, nearly completed in Ottawa, it was desirable to have his Muskoka and Pelee records published in the revised edition of Macoun’s Catalogue of Canadian Birds (1909). Because his contributions to the first edition had not been directly acknowledged he had taken every occasion to send subsequent records to James Macoun, responsible for revising his father’s work. Taverner was espe- cially careful to send separates of his notes and arti- cles in The Auk and The Wilson Bulletin, as well as to remind him of their previous correspondence about his records.'*> But when a copy of the revised edition arrived it caused him a sickening disappoint- ment. In spite of all the care he had taken to send separates to James Macoun the new edition con- tained no references to Pelee at all and about half of his records published in The Auk were omitted. Taverner wrote immediately to James Macoun ask- ing why he had not used the Pelee material. The answer he gave was very lame, as Taverner explained to Fleming. That the manuscript material had been passed to him in the summer of 1908 with instructions from R. W. Brock to hurry with the edit- ing. (Brock was preparing for the opening of the Victoria Memorial Museum in 1911 and presumably wanted a new publication to impress the politicians.) That he was under great pressure from his own botanical work; that the separates were overlooked, that although copies of The Wilson Bulletin (contain- ing “The Birds of Point Pelee’) were received for some reason they were not consulted. That Macoun was sorry that these oversights had occurred but he would make it right in an addendum later.'*° It was not only James Macoun’s lame excuses but even more the offhand tone of his letter that hurt Taverner’s self-esteem when he wrote: “The oversight in the whole matter was probably due to the fact that I was using Saunders’ and Fleming’s notes for Ontario and quite forgot your list, although I remem- ber now that reprints were sent me.”!37 This provoked Taverner into sending a sharp note in reply. He was upset, he said, to find that what work he had done in the Dominion was so easily overlooked, especially when he had taken particular trouble to keep Macoun’s department informed. The fact that even the notes in The Auk were not included gave it the appearance of a personal slight. He under- CRANMER-ByYNG: A LIFE WITH BIRDS Photograph of Northern Saw-whet Owl taken by Taverner at Point Pelee 15 October 1910. Taverner’s account of the difficulties of photographing it is given in Chapter 4. (Reproduced courtesy of the Canadian Museum of Nature, number 31879). AO THE CANADIAN FIELD-NATURALIST stood the difficulty of covering such a vast territory especially when much of the work had to be through the reports of people not directly involved in zoology but a number of unfortunate errors and omissions would have been avoided if some Canadian ornithol- ogists had been consulted on matters relating to their particular spheres of work.'** Taverner had made his point and shown that although he was not well known to the Macouns, while Fleming and Saunders certainly were, nevertheless he was a serious ornithologist whose records could not lightly be overlooked. He had learned to stand up for himself through this experience, though at the risk of dis- pleasing the Macouns. After the excitement of discovering the migrating Saw-whet Owls in October 1910 Taverner’s Journal began to peter out, the last entry being in November. The Journal had been the constant recipient of his ornithological notes and observations from the time of his arrival in Detroit in the spring of 1904. It con- tained not only data on numbers of birds and their distribution but above all descriptions of what it was like to be in the field and seeing, hearing and enjoy- ing the scene. Out of many descriptions of bird observations at Point Pelee one will have to suffice. “T certainly never did hear so many birds singing in the winter as we did during this visit. When we opened the door in the morning we were greeted by a regular chorus of bird music. The most conspicuous were the [Northern] Cardinal and Carolina Wren calls which answered each other back and forth across the jungle. Next to them came the simultaneous warbling of dozens of Purple Finches and under them all was a low subdued tone composed of the combined twitterings of hundreds of [Common] Redpolls and [American] Tree Sparrows, the latter continually essaying their sharp little song. Altogether from aural evidence it was more like May than Feb.”!39 Exactly ten years later, after reading this descrip- tion in his own Journal, he was moved to write to Bradshaw Swales: “Dear Brad; Do you remember blank years ago today the chorus of bird song we heard as we threw open the shack doors at Point Pelee in the morning?’ !4° There is more than a touch of nostalgia in re-liv- ing those feelings of a moment in time. Not only did Taverner’s Journal peter out during 1910 but the Bulletin of the GLOC also began to fail. Its few members were too busy with more important things. With the active support of Fleming and Saunders there was the chance that Taverner might obtain an appointment at the newly estab- lished Victoria Memorial Museum in Ottawa. (See Chapter 5.) By early 1907 the Taverner family was settled into their own house at Highland Park, and during the next four years home life became more enjoyable for them. Where they lived was just beyond the western edge of the expanding city, and was still in partial countryside. The electric street car ran close by at Woodward Avenue so they could easily get Vol. 110 into downtown Detroit and back. By 1908 they had a telephone. At the rear of the house was a section of field which Percy and his mother developed into a garden with advice from Fleming, based on his own gardening experience in Toronto, and a great deal of hard manual labour by Percy. Fleming and Saunders sent gifts of shrubs and plants during the next few years while Percy helped himself regularly to wild plants growing in the vicinity. From now onwards gardening was to take up a certain amount of his spare time. Although he had to work at the office on Saturday mornings he was able to do gardening in the afternoon and still go birding on Sundays. Taverner’s newly discovered pleasure in garden- ing led him to an interest in botany, and he hap- pened to meet a few Detroit natural scientists who were not ornithologists but glad to meet Taverner. Informal meetings were held at each other’s houses when scientific discussions developed.'4! Taverner also found expeditions with two or three members of this small group very congenial. For instance on Sunday, 15 May 1910 he made an exploring trip with two others, Newcomb and Andrews. In a letter to Fleming he wrote: “We found a fine Canadian island set down in Oakland Co. Pines, balsams, arbutus and the best sphagnum bog I ever saw, covered with cranberries, pitcher plants, cypri- pediums [lady slippers], swamp laurel and kindred stuff ... Though I got no birds I enjoyed the changed condi- tions greatly.”!47 From 1909 onwards he mentioned in his Journal that he was collecting far fewer bird specimens, but instead his interests had expanded so that he wanted to learn more about birds in relation to their habitat. That is one reason why he began to enjoy the compa- ny of specialists in other aspects of natural science. There was much he could learn from them. His outlook on taking bird specimens for his pri- vate collection was beginning to change at this time. If he relied on a shotgun less, then he would have to rely on a better pair of field glasses. Fleming had written several times about much improved makes. Taverner took up the topic in 1909 when he wrote: “... 1am just negotiating with Wallace who can get the finest Zeiss glasses I ever saw very cheap — $35.00 through friends from the Old Country. They are prism glasses of course and the largest and brightest field I have seen. I feel every day that I need a good pair more and more, especially as my collection of commoner stuff is getting filled up and I hesitate to shoot more of them.”!* In spite of having a slightly weakened heart as a result of having typhoid fever while in Chicago , and therefore not being free to play strenuous sports, he was in good health. He seemed to have kept fit on a routine of continual work, and outdoor exercise walking. He certainly did not coddle himself. He took it for granted that he could carry equipment such as a gun, heavy camera, binoculars and collect- ing bag all day without difficulty. He seems to have 1996 felt satisfaction from the challenge of physical endurance. During his local birding he could not fail to notice the effects of housing development on the former countryside. Two entries from his Journal show that he was aware of the problem of conservation. One day he and Swales went to the Mud Flats and then continued by way of the usually productive “warbler woods”. “Alas poor warbler woods, nothing remains of it but piles of brush and great lines and stacks of cord wood. It was a sad sight to me and a worse one to Swales who TOMES it when it was far vaster than ever I saw ie Another time, while walking in the vicinity of Hamilton Boulevard, he found a good patch of hard wood, the kind that makes an excellent warbler woodland. “But alas it is about gone just as it is found. The lumbermen have been at it all winter and it is almost in ruins.”!*5 One ingredient seemed to be missing from the home life of a man now turned thirty-five — there was no suggestion that Percy had any girl friends — no indication of any love interest or sex life. This may be due to his natural reticence and the fact that no records of a personal nature appear to have sur- vived. When he died in 1947 his long-time friend W.L. McAtee was invited to write an In Memoriam notice for The Auk. For information on Taverner’s earlier years McAtee turned to the one person still alive then who could supply it — Percy’s half sister Ida. But although she mentioned their life in Detroit the information she gave was mainly about wildlife and there was no mention of Percy in relation to the opposite sex. What his sister did mention was “his wonderful care of mother and me”. Percy may not have felt confident enough to have a close relation- ship with any other woman partly because of his stammer which was always worse just when he wanted to speak fluently. Several “schools for stam- merers” in Detroit advertised their courses in the City Directories of this period, and one claimed that stammerers could be permanently cured. When he first came to Detroit in 1904 Percy had taken a course which he hoped would be a more or less com- plete cure. As he perceptively wrote to Fleming: “Stammering has been so long a part of me that I would not recognize myself without it, and a cure is hard to believe as possible.” !4° He certainly was not cured by 1910. Another rea- son may have been because of his personal appear- ance. He was over six feet tall with a dark beard and side-whiskers which gave his face a slightly elongat- ed effect. Already in the club photo of 1909 his hair was beginning to recede from the top of his head. With so much of Taverner’s energy being taken up by his various natural history interests and activi- ties it is difficult to think of him in his job earning his living as an architectural draftsman. When we CRANMER-BYNG: A LIFE WITH BIRDS 41 last heard of his work as a draftsman he was resigned to continuing it as long as necessary, but he never expected that he would become an architect, and make a success of his profession. On the other hand this was not a routine job which called for no thought and little skill. He had to be able to trans- form the sketches and notes of an architect into a series of “blueprints” drawn exactly to scale. This left him room for some expression of originality though not much. He was trained, capable and, by this time, experienced in this work which required accuracy, patience and an ability to draw. Work in the architectural business in Detroit during these years fluctuated between lay offs, part time or full employment and even overtime. It was also possible to quit a job with one firm (or be made redundant) and to find a job with another firm without too long an interval. Taverner continued working for W. E. N. Hunter, until March 1909 when he was employed by the firm of Baxter and Odell. Taverner was pleased with the new job and, as he explained to Fleming, relations with his employer were pleasant. He was very glad that he made the change. It had been so long since he had received common courtesy in the draughting room that it was quite a change.!*” By 1911 Detroit was a growing city with a flourishing economy. Thanks to Henry Ford and others it had now become the automobile capital of the USA. Its population had grown from 285 000 in 1900 to 466 000 in 1910. Transportation was good — by train, interurban electric cars, and by local street cars. The city was pushing out its boundaries, and architecturally was a challenging mixture of styles, including some of the early “sky scraper” buildings. Entertainment, both popular and cultural, was grow- ing fast as were opportunities for sport and recre- ation in Detroit and vicinity. But being an architec- tural draftsman was not Taverner’s first choice for a career. He could make a living out of it but he could never consider making it his permanent career. He compensated for his lack of commitment to his office work by an absolute commitment to the study of natural history. Only this could satisfy his deepest feelings. In the summer of 1910 Taverner reached his thir- ty-fifth year. By now the main traits of his character were established and he had developed sufficient self-knowledge to understand himself and his limita- tions. To understand him it is necessary to see him as a complex person who had managed to adapt suc- cessfully to a complicated situation. He was a com- pulsive worker who needed to be fully engaged and if he did odd jobs for any length of time felt ill at ease. In a letter to Fleming in 1907 he wrote that he did not know what had been the matter with him in the last while. He had been unable to concentrate upon anything. He thought that he had too many matters on his mind. Nothing of any consequence AD) THE CANADIAN FIELD-NATURALIST but various odd jobs, including a little lepidoptera work about the electric lights, had been another dis- traction.'*8 Although there was “nothing of any con- sequence” on his mind, subconsciously there could have been something of deeper significance such as the urge to find himself a mate or the utter incompat- ibility of combining the work of a draftsman with the work of an ornithologist. How he was able to com- bine the two types of occupation without becoming schizophrenic in these years is a mystery. It showed a considerable resilience on Taverner’s part to con- tinue doing both adequately. It showed an iron deter- mination to continue both. His ambition was to become a “scientific” ornithologist and to gain some recognition for his work in this field. He drove him- self hard in trying to achieve his ambition and seems to have indulged in few relaxations. He smoked a pipe regularly and found it soothing. Taverner thought seriously about the creation of the natural world and the place of human beings in it. Fleming asked him in a letter if he had ever read Darwin’s Journal and he replied that he had a copy of it, and that it was a book he thoroughly enjoyed read- ing over and over again.'*? Fleming also sent him two articles, one of which was an interview by someone with Thomas Edison.'°° Taverner responded by send- ing Fleming his views, and revealing something of his own thinking about religion and his critical, inde- pendent cast of mind. At the end of his long letter to Fleming on science and religion Taverner summed up what he believed. “My whole religion is this. I know nothing. I surmise a few things.”!>! In 1908 Taverner, as he told Fleming in a letter, was in the process of becoming a Freemason. This was rather a financial strain on him too. Soon after- ward he had a formal photograph taken, dressed for a ceremonial occasion, which his friends said made him look “very good”. But Taverner hoped that he did not look quite as weakly goody-good as he thought it.!°? There is no evidence that he went to church ser- vices of any religious body. His outlook on life, which he had already arrived at, had been reached by reading, and by his close experience of the natu- ral world. Although not “religious” in the usual human use of the term he felt a sense of mystery CHAPTER 5. From Amateur to Professional Although Taverner may have pushed to the back of his mind any thoughts of obtaining a position at the new museum being completed in Ottawa in 1910 his friends had by no means given up hope of seeing him in the position of ornithologist there. As the time drew near for the transfer of the Geological Survey’s collections to the museum building Taverner’s future career was again on the block. Vol. 110 about the complex interactions of the universe of which the human species was a part. Taverner was an observer and thinker who wanted to come to an understanding of life and its purpose through his own reasoning rather than to accept ready-made explanations handed down by authority. The whole direction in which his philosophy of life had been developing during the past ten years was against accepting anything on faith just because someone stated that something was so. He had a questioning nature. But this was not through any idea of his own intellectual superiority or claim to leadership. The opposite is true. He realized that his lack of formal training in biology put him at a disadvantage in nat- ural science. He knew himself well enough to real- ize that he had no aptitude to organize others or to become a leader.'*? Nor did he possess much self- confidence in his relations with other people at this time. He knew almost nothing about his own father, and to have to gloss over the fact that the name he used was not his name at birth, must have caused some psychological strain. The “A” of his given names caused him uneasi- ness while Percy fitted him comfortably enough. The secret was out in 1907 when the AOU requested all its members to write their given names in full so as to include them in full in the membership list. When Fleming received this request he told Taverner that he had sent his own, James Henry, and asked what his were.!*4 Taverner replied: “As to names I do not advertise mine — Algernon is not one to be proud of.”!>> Apparently Percy did not reply to repeated requests from the AOU Secretary, T. S. Palmer, who finally sent him a letter warning “If you do not reply I shall list you as Percy Algernon Taverner.”'°° To this Taverner replied: “Guilty as charged”.!°7 Taverner had a naturally unassuming nature as well as a strong sense of humour, of comic events or absurd situations. He was not in the slightest pompous. He did not regard himself, even in his ornithological work, so seriously that he could not make jokes at his own expense. The most outstand- ing qualities in his personality were his friendly nature and his streak of stubbornness. This is the kind of man that Taverner appears to have become by the age of thirty-four. Much had happened as regards the Victoria Memorial Museum since 1906 when Taverner first heard about a possible position there. As the result of “An Act to create a Department of Mines” passed in 1907 an administrative reorganization took place by which a Mines Branch and a Geological Survey Branch were placed under a Department of Mines headed by a minister. The 1996 Survey’s duties remained virtually unchanged; it was still responsible for “examining and surveying the geological structure and mineralogy of the country...” At the same time the Act maintained the Survey’s relationship with the Museum. “The Department shall maintain a Museum of Geology and Natural History for the purpose of affording a com- plete and exact knowledge of the geology, mineralogy and mining resources of Canada.” In addition it was charged to “collect, classify and arrange for exhibition in the Victoria Memorial Museum such specimens as are nec- essary to afford a complete and exact knowledge of the geology, mineralogy, palaeontology, ethnology, and fauna and flora of Canada’”’.! The Victoria Memorial Museum was not founded as a new and comprehensive national museum but developed out of the geological museum maintained by the Geological Survey of Canada. The collection and display of natural history specimens was only a part of its functions. As a result of the Act the Geological Survey was no longer a separate government department but was now one branch of a department of government alongside another branch (the Mines Branch). “Tt would have to compete with the other branch for a share of the departmental appropriations, and for the ear of the deputy minister and minister for decisions as to which program to pursue, and with what intensity.” This would cause problems within the museum divisions in the future. In addition, the Director of the Geological Survey, Albert Peter Low, became seriously ill with cerebral meningitis at this time. As a result a professor of geology and petrology at Queen’s University, Reginald Walter Brock, was named Acting Director of the Survey Branch at the end of 1907. When Low’s condition worsened Brock was made Director of the Survey and Acting Deputy Minister of Mines in late 1908 at the age of thirty- four. Brock had joined the Survey in 1898 and had five years’ field experience before going to Queen’s as professor in 1902. He brought a dynamic quality to the Survey which was needed at this time of reor- ganization and museum development. He was fortu- nate that the Canadian economy had begun to revive so that the annual appropriations to the Geological Survey were adequate, and in turn were adequate for building up the natural history collections and appointing new staff. In 1907, C. H. Young, an ento- mologist with the Experimental Farms Service, was appointed assistant to the curator of the museum and prepared a series of display cases illustrating the life histories of butterflies and moths. William Spreadborough was employed in 1908 to make a col- lection of the natural life of Vancouver Island for display in the future museum.’ In 1909, Jim Macoun edited a new one-volume edition of the Catalogue of Canadian Birds. Brock had instructed that the manuscript should be completed for publication CRANMER-BYNG: A LIFE WITH BIRDS 43 quickly so that it would be ready before the disruption of the move into the Victoria Memorial Museum which was expected to take place in 1910. In his hurry to complete the job Macoun failed to correct many errors in the original catalogue completed in 1904, or to include much of the most recent field data supplied by correspondents such as Taverner.* Nevertheless, it was a useful foundation from which future distribu- tional studies of the birds of Canada could begin. Brock, in his report as Director of the Survey for the year 1910, wrote optimistically about the future. “Now that the Victoria Museum is being occupied, every effort will be made to strengthen the staff.” He was determined, he wrote, to make it “a complete nat- ural history museum’”.> It was at this point that P. A. Taverner was considered for a position in the Natural History Department of the museum. In March 1910, Ernest Thompson Seton was in Ottawa and heard through John Macoun that a posi- tion at the museum might become available. Macoun mentioned Taverner’s name in that context. Seton called on Fleming in Toronto on his way home and relayed this information. Fleming immediately wrote to Macoun to ask if Taverner should go to Ottawa so that he could be seen. He then wrote to Percy with the news suggesting that he should come to Toronto over the Easter holiday, when special train rates were available, and then go to Ottawa, adding “I will be responsible for the cost of the trip.” Fleming now had the bit between his teeth. He was excited by the prospect and anxious for his protegé not to let slip any opportunity to further his own chances. He warned that “... there may not be anything in it but one never knows and any way a meeting would be well worth while if we can arrange one. I told Macoun I did not know what was on your mind.”° The scenario now became involved. James Macoun answered Fleming’s letter to his father saying that “it would be quite useless for Taverner to come to Ottawa just now” and explained how they came to speak to Seton about Taverner. ‘When he was here two years ago he asked whether there would be an opening for one of his nephews, and either my father or I introduced him to Mr. Brock and I believe he made at least a verbal application to Mr. Brock at that time. When Seton was here Monday he brought the matter up again by telling us that his nephew was now in the Field Museum at Chicago and so would not be an applicant for any opening that there would be here:;7/ James Macoun explained that he asked Seton what he thought of Taverner, but that was all that had been said or done in the matter. Meanwhile, what Macoun wrote next put Taverner’s appointment in an unsatisfactory light. “Neither my father nor I know anything at all about what new appointments will be made for the new museum, nor indeed do we know whether we would be consulted in the matter, but naturally we would both like to see the best available man appointed as taxidermist, and from 44 THE CANADIAN FIELD-NATURALIST what we have been able to learn from yourself, Seton and others it seems that Taverner would be just the man. I may tell you at any rate that we know of no one else. It cannot be very long now before some action will be taken, and if either my father or I are consulted we will say that, so far as we know, Taverner is the only avail- able man for the position of taxidermist.”® This was a curious mistake for James Macoun to make. It was made perfectly clear in 1906, when Taverner was being considered for a possible posi- tion at the future museum, that he would not have accepted the post of taxidermist. Perhaps James Macoun had forgotten this. However, this is not how Fleming and Taverner regarded it. Fleming sent Macoun’s letter to Taverner to explain why his pro- jected trip to Ottawa would be useless, and com- mented: “IT have been aware from some time that there is a cold hand somewhere and I fancy it 1s Jim Macoun’s influ- ence with his father. One of the things I did not do was to recommend you for taxidermist as Jim Macoun very well knows ...”.? The new Director of the Geological Survey, R. W. Brock, was not looking for a taxidermist at this time as the following extract from Fleming’s same letter proves. A Toronto taxidermist, Horace Mitchell,!° had seen Brock in Ottawa a few days previously to ask whether a position of taxidermist was likely to become vacant. Brock told Mitchell that there was unlikely to be any changes for a year, and did not give him much hope of a post. “However he told Mitchell they were looking for a man for the position, I think, of Curator . . . and here is the joke, Brock mentioned Seton was the sort of man they would like. I must go to Ottawa and find out really what conditions are. I don’t know Brock. But I fancy you will have to wait till some future date for anything definite. I don’t believe they know what they want and I think Jim Macoun is afraid someone bigger than himself will get it. 1 am sorry to have stirred you up but Seton’s state- ment was so straightforward that I naturally felt justified in writing to Macoun.”!! Taverner tried to guess at the cause of James Macoun’s attitude. “Perhaps the antagonism of Jim Macoun may largely be due to my not carefully concealed opinion of the List. I think I told him that it was most creditable as a botanists attempt at ornithology but that it was rather to be regret- ted that an experienced ornithologist had not been fur- ther consulted as it progressed. It may not have been politic under the conditions but if the only way I can get a position of the kind is by truckling to others I think I will stay a draughtsman. However if I can get it without losing moral independence there is nothing I would like better as you know.”!? At the same time Fleming wrote to James Macoun to say that he was mistaken in thinking that he, Fleming, had recommended Taverner for a position as a taxidermist. “T stated that he had a thorough knowledge of the subject but that he would not accept such a position. The posi- tion I consider Taverner fitted for is curator of the natu- Vol. 110 ral history section of the museum that is to be, in fact he is just the man to carry out the views of the staff and work with them and stand between them and the public.” Macoun’s reply was a frank apology and an admission of the muddle prevailing in the Geological Survey over the Museum. “It was pure thoughtlessness in writing to you as I did regarding Mr. Taverner as I of course knew the position for which you were recommending him, and we had no other in mind. The word taxidermist was a simple slip. So far as we can make out nothing at all is being done about appoi[n]tments and no[nJe of us know where we stand ourselves or what new appointments will be made ... This of course is a bad state of affairs with the Museum so near completion, but I suppose the whole trouble is that no Director of the Museum has yet been appointed and there is no one in authority.”!* When Percy told the household of Fleming’s ini- tial letter it caused “a lot of commotion”. “Mother and Ida would of course go with me if any such move were made but it would be like pulling a plant up by its roots for her to break her associations here now just as we have got so nicely established. The house and garden seem to have become part of her almost now and I must confess it would be rather a pang to me. However it would only mean beginning over again and we would soon be as much at home at Ottawa as we now are here.”! Percy, Ida and their mother had already made var- ious moves together and presumably could put down roots again. At this point Saunders added his influence in a let- ter to John Macoun stating the advantages of appointing Taverner to the museum staff. “T am very strongly impressed, more so indeed than I can tell you, with the fitness of Percy A: Taverner, Highland Park, Mich., who is a Canadian, for a position in the Survey. Besides being a keen observer, an extra good Taxidermist, he has a very artistic temperament. He is a good artist and by profession Architect. I am sure it would be exceedingly difficult for you to find another man so well equipped to assist you in the particular work there, as he is. Moreover, he is well educated and his let- ters are well composed. You have perhaps seen some of his writings in the Wilson Bulletins, but for fear you may not, I will send you in a day or two, something from it which will show you how he writes. I do not know where you could find another man so good to assist you in installing the collection in its new quarters, and I hope you will give him very serious:consideration indeed.” !® This letter was answered by James Macoun who sent a non-committal reply. “As regards Taverner, neither my father nor I know any- thing about him personally but both Seton and Fleming seem to think that he is a man whom we should have at the new museum and, though my father and I cannot do anything to forward his appointment, we have no one else in view ourselves and to that extent at least may be considered as indorsing his application.” Fleming was sorry that Taverner’s projected visit to Ottawa had been turned down, and told him“. . . it is high time you should get acquainted with Ottawa. 1996 I am seriously handicapped by you not being known in Ottawa.” After advising him to go on with his work he added: “I don’t want to be meddling in your affairs but I would like to see you in some place where your work would have a permanent value.”!® There was a break in the correspondence during the summer while John Macoun was collecting plants in Nova Scotia and Brock was visiting sever- al museums of natural history in the United States. There he could see for himself how to organize a museum and how best to display natural history specimens for exhibition. The next development came in November 1910 while Thompson Seton was on a lecture tour which included Detroit. Taverner wrote to Seton inviting him to stay the night when in Detroit. Seton declined because he had to leave for Winnipeg the same evening after giving two lectures, but added: “I hope however I shall have a long talk with you about Ottawa etc.’’!9 In fact he was able to spare Taverner only a few minutes. In writing to Fleming, Taverner reported that Seton had recommended him to Brock for the position at the museum. “He outlined just such a position as you have. A bird man, not necessarily a taxidermist but a judge of taxi- dermy and capable of doing scientific museum work. Salary $1500 a year with a rapid advance to 2500. A Canadian would be preferred . . . It has occurred to me that perhaps it has not been mentioned that I am a Canadian and some of the hesitancy may have been caused by the idea that | am an American.” Seton advised Taverner to write to Brock himself and ended the conversation by saying the position was his if he cared to go after it.2” Whether or not Seton rubbed Taverner up the wrong way by being a little patronizing, Taverner was sharply critical of Seton when mentioning his visit to Fleming: “I was disappointed in Seton’s lecture on his northern trip. His undoubted genius is so overlapped by an exalt- ed ego. With the exception of one other listener I think the whole audience went away with the idea that Preble was but a hired guide in whom he had perfect confi- dence and a very good fellow for his station.”7! Taverner now wrote to Brock asking for details of the post, mentioning that he was a Canadian citizen, and asking whether applications would be consid- ered. Brock replied that it was likely that there would be an opening “for a Naturalist and Preparator in the immediate future.” What was wanted, he said, “... 18 some one who is fond of birds and animals, knows something about them and their habits, and who is something of an artist so that he could have specimens collected, mounted in a natural position with natural sur- roundings and exhibit them tastefully in the Museum. In addition to these rather rare qualifications if he could also write well it would be a further recommendation. In accordance with the Civil Service regulations the posi- tion will be advertised and applications received. Some testimonials as regarding work and samples and speci- mens of the work done would be helpful.” 77 CRANMER-ByYNG: A LIFE WITH BIRDS 45 Taverner sent Brock’s letter to Fleming and com- mented: “This looks as though something is doing at any rate. I wonder if there will be a Civil Service examination attached. If so a language requirement would be fatal. The letter speaks of a Naturalist and Preparator. Seton said that a taxidermist was not the requirement but a crit- _ ic of taxidermy.” 7+ Fleming, who was aware of various “under cur- rents” in regard to the appointment decided to go to Ottawa “to find out just which way the wind blows”.** Before leaving for Ottawa to lobby for Taverner he wrote: “I guess any Civil Service examination will be a farce. Of course I don’t know. You are independent and I would not take the ground of seeking the position, it is the other way about, at least we will try to make them think so.”?> Fleming had the instincts of an accomplished lob- byist. Suddenly the pace quickened. In a slightly dramat- ic gesture Fleming sent a night lettergram from Ottawa. “Have talked matter over with Brock. Showed my letter to Low of four years ago favourably received and Brock’s description of position fits in nicely. Saunders has already written. I am to write too formally. Don’t think position is immediate. Macoun is favourable. Can only wait for things to move.”*° Like a journalist with good copy Fleming made the best of it and sent Taverner a four-page letter describing the interview with Brock and a talk with Macoun. He had over an hour’s talk with Brock. “He is a young man with a determined face. I should say a born director of men. I showed him my letters to Low of four years ago and he said that is exactly what we want... After a little careful guiding I got him to describe the position I wanted, outlined in much my own words . . . Brock said we cannot afford to ignore the public after all it is their museum and we have no one to take the preparing of exhibits off our hands . . . | showed him the Swan drawings, he was delighted.’ Fleming then had a session with Macoun who told him some of the administrative and political prob- lems facing the new museum; its role vis-a-vis the Geological Survey and Department of Mines, and the fact it might be placed under the Minister of Agriculture in the future, among other uncertainties. While they were talking Thompson Seton came into the room and three of them discussed Taverner’s suitability for the position. According to Fleming “Macoun got enthusiastic and told us he himself would go to Brock and tell him Taverner was his choice. Seton said you could do anything but lecture. He gave you all sorts of a character enough to turn your ego into a dirigi- ble had you heard it.” He also said that Seton knew Fisher, the Minister of Agriculture, and would write to him if necessary. Fleming wrote: “Seton has done a great deal to prepare the ground and Saunders, Brock tells me, has written a very nice letter.”?° 46 THE CANADIAN FIELD-NATURALIST Fleming again warned that he detected strong undercurrents at work, and that nothing was certain until the job was landed. He ended with some advice. “One thing is certain if you should get this position you must do us proud. I haven’t any misgivings and am cer- tain you can do the work particularly as the position has practically to be made by you.”?? So wrote Fleming to his protege with just a touch of self-satisfaction. Now it was Taverner’s turn to consider his own situation in the light of recent information, and to show gratitude. This he did in a letter to Fleming in early December. “IT have read over your letter and telegram, Saunders’ letters and thought over them both in combination with what Seton said. The first thing that strikes me is that I have a lot of good friends who are working very hard for me. If the position does come my way I hope I will make good though I fear I will have to live to a pretty high standard if all that has been inferred has been said. However | think I can do it though I realize that it will take every effort I am capable of.”*° One major problem remained. The Civil Service Commission required that positions in the Survey should go only to University men. However, Fleming reported that “Brock makes light of the matter’ >! During the meeting on | December Brock asked Fleming to write a letter to him about Taverner’s suitability for the position at the museum. This he did, though not without considerable effort, as he told Taverner. “On my return from Ottawa I wrote a long letter to Brock, and to show how much I loved you I type-wrote it. It took me a whole day to compose.”*” _ Because this letter sums up very well Percy Taverner’s interests and skills it is given in full below: “267 Rusholme Road, Toronto, Ontario. December 8th, 1910. R.W. Brock, Esq., Director Geological Survey, Ottawa. Dear Sir: In reference to the conversation I had with you on the Ist about Mr. P. A. Taverner and the curatorship of the new museum, it may be desirable to state my understanding of the requirements, as brought out in our recent interview. The opening of the new Victoria Memorial Museum will make it possible to give much more room to the exhibition of Canadian Zoology than was the case in the old building, and it will be necessary to make the very best use of the material now available in order that the public galleries may contain an instructive collection that in time will grow to be a representative collection of the natural history of Canada. The curator must have a thorough knowledge of the methods used in preparing groups and specimens, and be able to direct this work, prepare attractive labels and arrange the exhibits to the best advantage. He must under- Vol. 110 stand the requirements of a popular museum, while not overlooking the scientific, and above all must possess enthusiasm for the work. Mr. Taverner is a Canadian now practising as an archi- tect, a profession that would be of great service in arrang- ing and grouping the exhibits in a museum, he is an ornithologist of high standing, an artist of ability particular- ly in ornithology, many of his drawings being artistic as well as scientifically accurate, a very rare qualification. He has recently finished the illustrations for Professor Barrows’ Birds of Michigan, and has, I believe, begun the drawings for a monograph of the fresh water mollusks for Bryant Walker, the American conchologist; he is an excel- lent draughtsman and is able to turn his hand to the illus- trating of almost any zoological subject. This ability is par- ticularly valuable in preparing exhibition labels and reports. He is a fluent writer, using excellent English and his scien- tific papers are notable for their clearness. Mr. Taverner has done considerable field work in the region of the Great Lakes, with important results to ornithology. His researches into the fauna of Point Pelee have added several new birds to the Ontario list and extended the range of others, besides adding much to our knowledge of the problem of migra- tion. He spent some years at Ann Arbor where the museum of the university has been a pioneer in the idea of reproduc- ing birds and animals in their surroundings. He there acquired a practical knowledge of taxidermy, later he was with me in Toronto and made many casts of fish and rep- tiles. I was at that time searching for a cheaper method of reproducing the plants and foliage used in groups such as the British Museum then had on exhibition, and Mr. Taverner was of great assistance in my investigations. As there was no opening for such work in Canada he went to Chicago where he took a course in modelling in one of the Art Schools there, and I have seen several excellent animals he modelled at that time. He also followed closely the work carried on at the Field Columbian Museum and was able to get a good knowledge of the methods which at that time were revolutionary and have since been followed by all the great American museums. Another qualification Mr. Taverner has is his ability to absorb and carry out the ideas of others. I wrote Mr. Low in 1906 about the new museum as follows, ‘the new museum will require someone competent to take the re-arrangement of the collections off the shoulders of the staff, one not nec- essarily a geologist, but of sufficiently wide intelligence to give effect to the wishes of the different technical branches of the Survey in the rearrangement of the present collec- tions, also someone to take hold of the Zoological part of the museum, in fact, a curator able to make the museum attractive to the public, and yet not lessen its scientific value.’ It is very rarely so many qualifications are present in one man, combined as they are with level headedness, and I feel that with Mr. Taverner as an assistant you can make the museum attractive as well as a national one. For the last twenty years I have kept in constant touch with the advance in museum methods, and while my time has been necessarily given to the building up of an ornitho- logical collection, I have not lost interest in the subject, and I know that Mr. Taverner has kept informed as I have. Yours sincerely, J. H. Fleming ”*? Shortly after Christmas Taverner received a letter from Brock about the proposed salary if he were 1996 appointed. It was carefully worded so as not to leave Taverner with any cause for complaint. “T do not think the salary you mention would be too much to pay for the successful and efficient execution of the duties of the position, but I think it might be difficult to make an appointment at that figure. It would be easier to make an appointment at $1600, as this is the initial salary for that Division of the Service, which is the Division in which men are usually appointed for their ‘try out’. After a man has proven his ability in this Division, promotion may be made to the Division above, at which the initial salary is $2100, increasing annually $100.00 a year. As you have not had any actual experi- ence in Museum management, I think it would be most satisfactory to follow the course which I have indicated as the usual one.”*4 Fleming also received a letter from Brock about the proposed salary and commented: “It looks like business all right.” Always solicitous for Percy’s finances he then did some costing based on the fig- ure of $1600 in the first year. He estimated from information given him by acquaintances living in Ottawa that it would cost Percy $12.00 a month for a room in a private house and $20.00 a month for meals at the best apartment house restaurant, so that he could live on about $600.00 a year, but should not count on less. As Taverner was to have an interview with Brock early in January 1911 Fleming invited Percy to stay at his home in Toronto over the New Year holiday. “We may have to give you a cot in the bird room, but if you are welcome to that, I know the birds won’t mind.” He also made the usual offer to pay his fare from Detroit to Toronto and return.* While in Ottawa Taverner also met Macoun with whom he had been corresponding for the past decade. As a result of the interview he sent Brock an outline of what he thought was wanted for exhibition groups, and a set of his flower pictures to Macoun. After that all he could do was to try to wait patiently. As a full month from the time of the advertise- ment in the Canadian Gazette on February 18 had to be given for other applicants to reply, the selection of a candidate could not occur before late March. Time dragged on. Eventually a telegram dated Ottawa 22 March arrived at the the Taverners’. This looked like the moment of truth, but when Percy opened it it said: “Send formal application to Civil Service Commissioners Ottawa for position as Naturalist Preparator and assistant Curator Natural History. R. W. Brock.”°6 Brock followed this with a letter explaining that the previous application from Taverner was not exactly suitable for the position as now advertised. Another disappointment, another letter to be written, another delay. Taverner was getting a foretaste of the workings of the bureaucratic system. However Brock softened the impact, somewhat, at the end of the letter. “I may say that there is no reasonable doubt of your selection as you have no serious competitors. The CRANMER-BYNG: A LIFE WITH BIRDS 47 Commissioners, however, will defer action until the receipt of the formal application.”>” At the end of March Taverner was finally informed of the decision. Brock’s letter read: ) “Dear Mr. Taverner, It is quite safe for you now to plan to come to Ottawa, as I understand the Civil Service Commission is notifying you that your application has been successful. I am sorry that you should have been kept in suspense for so long. Yours truly, R. W. Brock’”?8 At last the suspense was over. Now the challenge of a completely new type of work with new opportu- nities and new responsibilities would begin. Their home in Detroit with its pleasant garden would have to be sold, and his mother reconciled to moving to a city she did not know. Fleming was quick to write congratulating him and offering good advice. “T am delighted that the position is at last secure. I have no fear of the result. It is a great opportunity, the opportunity that comes just once in a life-time ... No doubt you will have all sorts of troubles to contend with but it is for you to decide if you will accomplish anything and I hope at no time will you get discouraged and settle down to the routine of a civil servant which is to draw pay.”?? There were still a number of points to be settled and at least one more hurdle to be surmounted before the job was assured. The main worry was over the medical examination for entry into the Civil Service. As we know Taverner had a severe illness, typhoid, while a young man in Chicago which caused him to have a slight heart murmur.*? Naturally he wanted to know how the examination would be carried out. In his reply Brock said that it was usual to obtain a medical certificate from one of the two doctors in Ottawa nominated by the Commission. In reply to Taverner’s other queries he wrote: “We should like to have you in Ottawa as early as possi- ble to discuss matters in connection with the Museum, but I can understand that it is necessary for you to remain in your present position for a short time. I think it is better for you to come to Ottawa before visiting the outside Museums.””! For information on the nature of his work, and the extent of ornithological books in the library Taverner wrote to Macoun for guidance. He received a reply from James on behalf of his father. The Museum, he wrote, had a fair library of standard ornithological works including the Awk [Auk] and similar periodi- cals. The nature of his work in the immediate future would be decided in consultation with the Director and Macoun senior. All the natural history material remained untouched after the move from the old building and there were no display cases as yet. “T need hardly tell you that we are all delighted that you are to join our staff and, like yourself, we anticipate that we shall have a mutually pleasant and profitable time together. Your work, as I understand it, will be practical- ly independent of everything but general instructions.” James mentioned the difficulty of finding some- where to live in Ottawa as regards rent and locality, 48 THE CANADIAN FIELD-NATURALIST and offered any assistance he might want in finding immediate quarters and information about the differ- ent parts of the city. John Macoun, in a handwritten P.S. added: “Our specimens are still as they were brought here. The mounted ones were boxed up by Herring. He must take all orders from you so commence with that in your mind. Put your own ideas in shape and I will support you; in other words you will have a free hand as far as I am con- cerned . . . make your own plans and [ will help you carry them out.’ This friendly note of support from the two men who could help or hinder him most in the first diffi- cult year of his new job must have eased his anxiety somewhat. In fact Taverner might have congratulat- ed himself on coming in on the ground floor when a new museum was being organized, thus realizing his long-cherished dream to work in a museum of natu- ral history and convert his part-time recreation of ornithology into the full time profession of ornithol- ogist. But reading between the lines he might have felt that John and James Macoun were also glad to welcome him at the moment when there was a great deal of tiresome (and dirty) work to be done before the exhibits, for which the Director was vigorously pressing, could be displayed. The problem of the taxidermist, Samuel Herring, was clearly being dumped in the lap of the newest member of the . Natural History Division for him to deal with.* It was under these circumstances, and with these hopes and cares, that Taverner made the journey from Detroit to Ottawa where he arrived on the morning of the first of May. It was, perhaps, a sym- bolic date. Spring had arrived. The tedious years of drafting plans for others, the frustrations of not being free to follow one’s own bent and to show one’s potentially creative abilities were over; the journey- man draftsman was about to be transformed into a newly fledged member of the species homo ornitho- logicus. But Taverner was a modest man with plenty of common sense and a very practical turn of mind. Even if he had indulged, briefly, in grandiose thoughts or self-congratulations, an incident on the very first day of his new life in Ottawa was enough to bring him down to earth with a jolt; he ran short of money. Fortunately friend Fleming was ready to help. He explained his predicament in a letter of 1 May 1911. He had been hunting for boarding houses all day. “However I am a little strained. I found expenses more than I expected and am pretty well on the ragged edge. Salary day is May 15 ... I wonder if you would mind lending me $15.00 until salary comes along. That will fix me nicely. It is rather ignominious to be forced to borrow at the start like this especially when it is my own fault not to have come better provided.”4 However, this was merely the prelude to more serious problems. By May 10 he had still received no official notification of his appointment so he wrote to the Civil Service Commissioners for a statement Vol. 110 of his official standing. He was anxious to get his family to Ottawa as soon as possible but would not do anything until he knew how matters stood. As he told Fleming “... a week and a half of boarding house life is all I want.” Meanwhile, Macoun discovered that the accoun- tant had not received any notification of Taverner’s appointment, so he advised Taverner to see the Civil Service Commission about getting his name on the current month’s payroll. When he spoke to a mem- ber of the Commission he learned that his medical examination was not satisfactory and that the Commissioners were hesitating over his appoint- ment. This was a very disquieting turn of events which Taverner described to Fleming in a long letter written three days later.4° The reason for the medical examination, Taverner was told, was to guard against appointing anyone who might be liable to frequent attacks of sickness or prevented from work- ing for a long period. To this Taverner replied that he had always understood that if ever his heart defect acted up the result would be sudden; that either he would die of some other cause or “he would drop out suddenly and effectively at once”. Since this crisis began on a Saturday morning the Commissioner could do nothing until Monday, nor would Taverner receive any salary until the following month (mid- June). Off went Taverner to Macoun’s private house. Macoun was “greatly put out” when he heard the news and advised Taverner to look for Brock “and camp on his trail until I found him”. He also volun- teered to lend Taverner whatever money he needed to tide him over the month. “I thought that very nice of the old man after so short an acquaintance’, Taverner told Fleming. Luckily he found Brock who had already been informed. Brock explained that the Commission had made a few appointments in recent years which had turned out badly and left them with some invalids on their hands after only a short time.They dreaded the criticism that would follow if they made another such appointment. Brock said frankly that “if there was anyone else visible [and] competent to fill the position they would have turned me down immedi- ately”. This was not very flattering to Taverner but he did not complain. Instead he commented realisti- cally “but there isn’t and it is either me or no one.” Brock then explained the alternatives if the second physician also gave him an adverse medical report. Either Taverner could sign a clause to the effect that he would resign in case of serious disability, or he could have a contract with the Commission to carry out specific work only. Still in the same letter to Fleming Taverner said that he had already prepared a draft report on the state of the existing zoological specimens, and a plan for their public exhibition. He had shown this to Macoun who seemed quite impressed. Taverner remarked candidly: 1996 “He is a great field man but very little use as a museum head and J think he realizes it and wants someone who is perhaps to cover up this weakness in him and perhaps in Jim who is scheduled to take his place.” Taverner believed that if his report had a similar effect on Brock this would be important because Brock had enough power “to force me through if he can get me no other way”. Taverner was undergoing a hard introduction to the world of the Civil Service and its workings. It is hardly surprising that a few days later, in a letter replying to one from Fleming advising him not to remove original labels from any of the skins, he reported on his health. “My stomach seems all out of condition and the last two days I have been wretched. I have obtained medical advice and hope that in a few days I will get set up again. I rather imagine that the change in water is likely largely responsible.” Perhaps that is true.*’ But equally it could have been the result of a fortnight of anxiety during which his desire to get on with his challenging job was being frustrated. This, alone, was enough to give Percy an acid stomach. Yet he kept on working as CRANMER-BYNG: A LIFE WITH BIRDS 49 best he could, unsalaried and in a state of acute uncertainty. Such a situation would have been upset- ting enough for anyone who was able to express himself orally with ease. But for Taverner, with his recurrent stammer, especially when he was in a state of anxiety, it was ten times worse. That he kept a cool head and survived is a tribute to his stamina. Certainly he was helped by Fleming’s knowledge of the bureaucratic system, and by his steadfast friend- ship. Fleming’s forthcoming visit to Ottawa was reassuring. But the only assurance that could settle Percy’s peace of mind came at the beginning of June when he was informed that he should present himself before the Clerk of the Privy Council to take “the oath of allegiance”. As he told Fleming, “Now if I can only get on the next salary list without having to wait for the meeting of Parliament again all will be lovely.”*® On June 5 he sent Fleming a postcard showing the main entrance of the Museum with the words “Appointment all fixed”. At long last, as though by metamorphosis, Taverner the draftsman emerged as Taverner the ornithologist. Part III - Challenge of the National Museum 1911-1919 CHAPTER 6. Museum Development (1911-1914) When Percy Taverner began his new career in 1911 at the Victoria Memorial Museum in Ottawa (hereafter referred to for convenience as the National Museum of Canada)! he was involved in something that a museum staff does not normally experience. He was taking part in the development of a major museum almost from the beginning in a newly com- pleted building, with very limited staff and almost no equipment. It is true that a small museum containing several random collections already existed, but the sudden transition from that situation to the founding of what was to become a large modern museum was an exciting challenge to all those involved. How Taverner reacted to this situation and what he achieved during the following nine years is the sub- ject of the next four chapters. However, before going into details of the prob- lems Taverner faced and how he coped with them, it may be worth looking at the attitude in North America towards museums by 1910 and, at the same time, to relate this to the state of ornithological stud- ies. In Canada at this time there were only a few museums and even fewer with natural history collec- tions of any significance. Museums need wealthy donors or foresighted city fathers to found them as well as grants and donations to maintain them. Canada, in the nineteenth century, with its small population and struggling economy was not in a position to enjoy well established museums.” However, by the first decade of the twentieth centu- ry, with an improvement in the economy, the found- ing of the National Museum in Ottawa and the Royal Ontario Museum in Toronto was possible. The build- ing of these two museums acted as examples that stimulated the development of other museums in Canada during the twentieth century. Luckily for Taverner, he had been in touch with developments taking place in some of the major museums in the United States in the period 1890- 1910. He also had a fair idea of the state of ornitho- logical studies in the United States in this period and the way in which these studies were being carried out in some major museums there. For instance he was well aware of the emphasis put on the economic importance of bird studies in relation to agriculture (“the pest or friend” simplification). Also he was informed about migration studies that were then attracting the attention of many ornithologists, pro- fessional and amateur. Moreover, the collection of data on bird migration was given impetus at this time by the first steps in North America to develop a tech- nique of bird “tagging” (banding) in which Taverner himself played a part at the very outset.* Also related to the problem of migration was the study of bird distribution and the analysis of the data being col- lected in local bird lists. In the first decade of the twentieth century The Auk and The Wilson Bulletin regularly published faunal lists to which serious 50 THE CANADIAN FIELD-NATURALIST Canadian amateurs such as Taverner, Fleming and Saunders contributed. Other topics covered in The Auk were studies of the breeding of various species, and studies of the taxonomy of certain genera, a sub- ject in which Taverner played a minor part.* Another subject that attracted many ornithologists was bird behaviour which came to play a popular role through the growing number of “life history studies” in the 1920s onwards. One of the attractions of this form of study was that the observer could concentrate on a bird’s habits entirely apart from taxonomy. It was Charles Bendire’s Life Histories of North American Birds that led the way in studies by other amateur ornithologists which subsequently became popular.> Another subject with which Taverner was familiar through practical experience was bird photography. He was alert to the value of photographs in books by ornithologists who were also pioneer bird photogra- phers such as Frank Chapman. His close-up photog- raphy of a Common Snipe and a Northern Saw-whet Owl showed his own ability with a camera.° By contrast, studies in ornithology in Canada by the year 1910 were much less advanced. The era of exploration of the Dominion’s avifauna had barely begun. Canadian ornithology was still at the early stages of surveying its bird fauna, and much useful work could be carried out through local annotated lists. This was something that Taverner encouraged from the first years at the National Museum because he quickly discovered how much work needed doing on geographic distribution, especially in certain regions of the country.’ Migration was a subject that few ornithologists and natural history clubs in Canada had seriously begun to tackle. If keen and knowledgeable ornithologists such as Fleming, Saunders, Taverner and Swales working co-opera- tively over a period of six years in such a favoured place as Point Pelee could discover only limited information about migration there, what could isolat- ed observers elsewhere in Canada be expected to achieve? Some progress had been made in the study of nesting behaviour, and a number of large egg col- lections had been built up. A few people were in pro- cess of studying various aspects of the life history of birds, though not very systematically. No one in Canada at this time appears to have been making studies of one particular species. It would be another forty or fifty years before studies of this type became established. The first step forward in this field began when those who were knowledgeable about certain aspects of the life history of a particular bird con- tributed their knowledge to Bent’s series of Life Histories of North American Birds which began pub- lication in 1919 as Bulletins of the U.S. National Museum. However, many Canadian naturalists in the decade before 1910 kept a regular journal, or other extended record of field observations. From these records some wrote papers or notes on their observa- Vol. 110 tions which were published in the ornithological or natural history journals of that time; in The Auk, The Wilson Bulletin, The Condor, The Ottawa Naturalist, and Le Naturaliste canadien; as well as in publica- tions by learned societies and by government. Another aspect of ornithology in the early years of the twentieth century was connected with conserva- tion issues. In the United States the movement for the conservation of wildlife through preserving wildlife habitat reached prominence in the 1890s with the campaign of John Muir and others to arouse public feeling. This was the decade in which Yosemite National Park near San Francisco was established and the Sierra Club was founded. A few years later the Audubon Society, devoted to the con- servation of wildlife and the natural environment, was established, and soon became influential through its local clubs and its official magazine Audubon. In addition to Muir, another man impor- tant in rousing participation in this growing move- ment was William Hornaday, director of the New York Zoological Park from 1896 until 1926.8 As a zoologist Hornaday became well known for his warnings that the sheer numbers of a species could not guarantee its survival in the face of human killing as the dramatic decline of Bison in North America showed.’ For Taverner, living in the United States at this period and reading accounts of these issues, the conservation movement impressed on him the urgent need for action to preserve wildlife habi- tat. The remark he made in 1907 that he wished he could speak persuasively because then he would seek public support for the preservation of Point Pelee is evidence of this. In 1907 he already had the vision to realize the need to make Point Pelee into a nature sanctuary although he lacked the means to carry it out.!? In Canada at that time, wildlife issues were only just being identified. When Taverner joined the National Museum there was only one offi- cially recognized bird sanctuary in Canada and a national movement to protect migratory birds did not exist.!' However, the establishment of a Federal Commission of Conservation in 1909 showed that the problem had been officially recognized. For many years subsequently, Taverner was himself offi- cially involved in the protection of birds and their habitat through his position as ornithologist at the National Museum. When Taverner settled into the National Museum in May 1911 he had time to reflect on his change of occupation and place of work. From his sparsely fur- nished office on the third floor he looked out over the roofs of Ottawa rather than the streets of mid- town Detroit. The work that faced him was not something handed to him by an employer week by week; he now set his own tasks and priorities. On 10 June he entered his thirty-sixth year. For the first time in his life he was responsible for planning the 1996 scope and organization of his own work, and the work of one man under his orders. Fleming’s recent warning that Seton, Saunders and Fleming would expect great things from him (“you must do us proud”) was sharply in his mind. At last Percy had been given the chance that he had wanted for so long, the chance to work professionally in ornitholo- gy and to do something that he considered worth- while with his life. The responsibility was consider- able — almost daunting, because the future develop- ment of ornithology, not only at the museum but also throughout Canada, depended to a large extent on how he tackled the tasks before him during the next decade. Taverner was the first professional ornithol- ogist to be employed in Canada. It was up to him to set the right course and to generate widespread appreciation of the study of birds. Before becoming immersed in the mass of detail that would continually need his attention it was important that he should have a clear idea of what a museum was about; what the objectives and priori- ties of sound museum management should be. The basic principles were not greatly different eighty or more years ago than they are today, though the tech- niques have changed considerably. A museum is based on collections of things. For Taverner the rai- son d’étre of his work was to build up an ever expanding collection of bird specimens; birds mounted for display, and skins arranged in drawers for research. But bird specimens are fragile and very susceptible to damage. Therefore they need continu- al care to preserve them from deterioration. Yet rep- resentative and well preserved collections of artifacts or natural objects are not much use if they are seen only by their curators. The main object of any muse- um collection is that a part of it should regularly be on view to the public. But as it is only possible to display a fraction of any collection at one time the major part is preserved in workrooms and laborato- ries where it can be used for research by scholars. The problems and frustrations that Taverner faced during his first decade at the museum can be better understood by reference to three main categories — exhibition, preservation and study — which consti- tute museum collections and need continual effort and financial support. Taverner’s dilemma resulted from having to decide exactly what priorities to allot when there was insufficient staff and money to do a satisfactory job in each of the three categories simul- taneously. His first impresssion of work at the Museum was one of limited chaos, and he fired off a stream of let- ters to Fleming describing the position. “As to the museum here. Well I can not tell you very much about it yet. Every one seems to be hustling about very busily but as far as I can see no one accomplishes anything. Indeed no one knows what space they are going to have or what they are going to do. There seems to be a spirit of expectant waiting for something.””!? CRANMER-BYNG: A LIFE WITH BIRDS 5) Taverner had his hands full taking a preliminary look at the zoological collection, both of birds and mammals, and deciding on the general layout of the exhibits based on them. As he told Fleming, there were two large halls in the building allotted for zoo- logical exhibits each lighted from the sides and lined with radiators in such a way as to leave no lower wall space available. The halls were separated by a square rotunda with a gallery all round. “My present idea is to give one of these wings up to a general survey of Canadian zoology showing groups illustrating the typical forms of life in each faunal group. For instance on the south side of the room having three main groups illustrating the southern fauna of the Dominion, namely Eastern Woodland, Prairie and Mountain. Across the room have a series showing the same for the Hudson Bay, Mackenzie and the Yukon and Arctic forms.” !3 He proposed to devote the other wing to “special- problem subjects” to illustrate things such as geo- graphical distribution, seasonal changes, and protec- tive coloration. An immediate problem was the large size of some of the mammal groups. One was a group of musk oxen in the snow mounted by the firm of Ward’s Natural History Establishment in Rochester, fourteen feet square. Other animals for mounting at Ward’s included Mountain Goats, Caribou, Dall’s Sheep and Moose. He proposed to display only a small exibition in the fall because he realized how important it was to begin in the right way with a properly tested plan and not to have a lot of cases made up quickly which would have to be discarded later. His immediate problem was to design display cases of various sizes and shapes to fit the idiosyncrasies of the museum’s halls. Temporarily, at least, the need to mount a display for the public had priority. After Taverner had examined the bulk of the bird specimens he was anxious to start on them immedi- ately. He told Fleming that he wanted to re-label and catalogue the whole lot and to clean and shape all the duck skins that needed it. This, he said, would be useful work to give Herring if only he would do it.!4 But Herring was getting old, had little respect for Taverner, and no wish to work on a pile of greasy ducks. He refused to take the first order Taverner gave him, and Brock hac to intervene. When Herring eventually responded he did the job in such a way that Taverner told Brock that he was incompetent, and his employment was terminated. At this point Taverner summarized the problems facing him in a preliminary report to Brock on the vertebrate collections dated 12 May 191]. He found that the mounted material was not up to the standard of modern taxidermy work and suggested it should gradually be weeded out. The study specimens need- ed immediate attention. Many carried field labels, hurriedly made on scraps of paper, while the locali- ties in which they were collected were marked only 52 vaguely. Some labels were almost indecipherable. He advocated that a new system of labeling and cata- loguing should be used, and explained what he had in mind.'> He also emphasized the need for proper storage cases for the study collections to make them safe from pests. Since the vertebrate collections needed thoroughly overhauling a properly trained and up-to-date preparator was urgently needed. In his report to Brock he was not too hard on Herring when he wrote: “The least said about our present taxidermist the better, as it is to be remembered that he is an old man versed in the ways of fifty years ago. . .”!° All these were very practical matters; but in addi- tion Taverner needed to set down his thoughts about the future scope of the collections. What, for instance, should be museum policy on the limits of the geographical area from which birds and mam- mals should be collected? Taverner’s suggestions did not originate with himself since the matter had to some extent already been decided before he joined the museum. The Geological Survey of Canada, under which the museum functioned, was concerned with the discovering, surveying and mining of min- erals only within Canada. Therefore, objects already collected by members of its field parties for preserv- ing in its museum had come from within Canada. Brock, in the Summary Report for the museum for the year 1910 devoted one paragraph to the policy and objectives of the museum just then being orga- nized. This set out an interim policy. “The Victoria Memorial Museum is a Natural History Museum including biology, geology, and mineralogy, ethnology and archaeology. As the National Museum of Canada it is hoped that it will become the repository of all objects of scientific value found within the Dominion. To fulfil its mission, it is necessary to secure the interest and co-operation of the Canadian scientists and of the public generally... For the present, at least, the Museum is confined to Canadian material, the object being to specialize in this until it becomes in all branch- es thoroughly representative of the whole Dominion, a place where the entire natural history of Canada may be studied.”!7 In his report of 12 May Taverner followed Brock’s lead but left the door open for collecting from a far wider perspective, when he wrote: “The Museum being under the Geological Survey of Canada gives it a National character at once, nor do I think it advisable to make any great effort to extend it beyond this limit, at least for the present. The time will certainly come when it will be necessary to broaden out and give its collection a world wide character both for the education of people in general and for the benefit of future Canadian zoologists for it is evident that life is so complex in its distribution that knowledge of Canadian conditions cannot be accurately arrived at without tak- ing into consideration extralimital examples and experi- ence. [emphasis added]’!® Taverner assumed that when the right time came government would authorize the collection of mate- THE CANADIAN FIELD-NATURALIST Vol. 110 rial from beyond the limits (“extralimital”) agreed to in the early years of the museum. But a precedent had been set from the beginning and it became “cast iron”. In subsequent years, whenever Taverner rec- ommended that specimens from the whole of the North American continent should be collected his advice was disregarded. Taverner then devoted several pages in his report to the problem of how best to organize and present zoological exhibitions. He recommended the group- ing of material so as to present the chief characteris- tics of various areas of Canada illustrated by the use of “typical animals and natural accessories carried out botanically, entomologically and in every other way to the last decimal point of zoological accura- cy...”!° He said that he had read all the reports he could find on the current arrangements in some of the major North American museums as well as the British Museum, but before adopting a final policy it would be necessary to visit some of the main muse- ums in the eastern United States. In the last para- graph of his report he asked Brock to consider care- fully the various points he had raised. “T am interested in making this institution take its place among the great institutions of the world. My scientific future is wrapped up in it and if I make a name for myself it will be through it. Maybe all the points dis- cussed cannot be carried into effect at once but they form a mark to aim at and will, I think, eventually great- ly assist in the reputation and the prestige of the museum Eula tetany of all who have any connection with Lte3 Taverner was still waiting to take the oath of alle- giance on becoming a member of the Canadian Civil Service. Fleming, with his impish sense of humour, teased him by writing “... let me know when you have been iniciated [initiat- ed] someday you will rise to be a deputy minister and attend levees in Windsor uniform with a sword between your legs great are the possibilities.”7! No chance of Percy becoming pompous with such a friend — he might trip over his sword. Finally he was able to tell Fleming the news. “T took the oath of allegiance the other day so now I am a Canadian citizen again I suppose. Guess things are all right now.” Under his signature he typed “Assistant Curator” and under that he wrote “Ha! Ha!’?? In the same let- ter he described to Fleming how Prof. Macoun, Charles Young and himself had recently gone on a day’s collecting trip to Mer Bleue on the east side of Ottawa. “It is a great sphagnum bog covered with Kalmia and pitcher plants and straggling patches of tamarac and some conifer. .. The Palm Warblers were breeding there rather commonly but all found had young. It is most astonishing to see Prof. Macoun in the field. Works through swamp, thicket and meadow with all the energy of a young man. He is a wonder.””?° One evening he called on a prominent Ottawa nat- uralist, George R. White, who lived beside the 1996 Rideau River.** This was the first link in what became a large network of naturalists across Canada who would collect birds for the museum and keep Taverner posted with valuable bird information. By now Taverner realized that when he came to “figuring out detail” how much he needed to make a tour of some of the museums in the USA. He also realized the magnitude of his task at the museum. “Tt is really a full sized job I have tackled and at every turn I wish I were better equipped for it.”?> Nevertheless, his first report shows that he had a fair grasp of museum work and management. He had the draftsman’s eye for visualizing arrangements of furniture and the items to be exhibited, as well as the skill of hand to produce plans and to work with wood. When it came to mounting the exhibits he showed all-round ability and considerable resource- fulness. Early in June his visit to the States was approved by Brock and he wrote to Fleming for advice — what museums did he advise him to see, who was the man at Washington from whom he could get the most useful information and other questions.”° Soon he was in Boston taking notes, measurements and observing everything. From now until the end of the month he wrote several detailed accounts to Fleming as from one museum man to another. At Harvard, he was critical of a few particulars. For instance the museum tried to combine a scientific and a popular exhibition which, he felt, did not succeed, partly because the labels were not well designed. Some of the mounting and background painting, however, was excellent. He also learned some useful things about chemical formulae. One disappointment: he could not talk with the taxidermists. “Was told that the taxidermists were close mouthed and would not tell anything. I think, however, I see how much of it is done. Did not meet them.”27 Since Taverner was looking for a taxidermist whom he could lure to Ottawa the people at the Harvard Museum were being cautious with good reason. At the American Museum of Natural History in New York he was able to look over the Preparatory Department and to meet all its staff. As though to make up for the years when Fleming went to all the AOU meetings but Taverner was unable to afford the expense or the time away from his job, he now met some of the outstanding American ornithologists of that time including Frank Chapman and Louis Agassiz Fuertes, the bird artist. He also met William Hornaday, Director of the New York Zoological Park, who gave him permission to take photographs in the grounds of the Zoo.”* However, Taverner kept his critical faculties in face of the excellence and size of the American Museum. He reported what he had seen but that he found the museum at Brooklyn almost ideal. CRANMER-BYNG: A LIFE WITH BIRDS 53 “It is not as elaborate of course as the American but per- sonally I think it is in better taste and just what I should like to make ours. The exhibits are arranged in an order that he who runs may read — everything shows some- thing and when the point is shown plainly and clearly it stops without confusing the beholder with a great multi- plicity and reiteration.””? While. at the Smithsonian Institution in Washington he spent a whole morning with Charles Richmond who showed him, among other things, the system of book keeping and specimen-preserving in his department.*° Taverner also spent time with Wells Cooke, the authority on bird migration, who explained his card systems at length. Taverner com- mented perceptively about Cooke in his report to Fleming. “T think his greatest delight is in making out cards — one card to a note. His system is perfect but when one has made it all out I do not see when he can get time to do anything else. I understand why he is so inaccu- rate.”>! From Washington he went to visit Ward’s Natural History Establishment at Rochester to examine their stock and see their methods of mounting mammals. This firm was working, under contract to the National Museum in Ottawa, on a group of Dall’s mountain sheep. Next he went to the museum of the University of Michigan at Ann Arbor. Here Norman Wood and other friends from his Detroit years may have greeted him; the same “lovable” Percy with a warm sense of humour but now a professional with the cares, and satisfactions, of a museum department of his own under his guidance. From Ann Arbor he went to Chicago, to the Field Museum where he used to spend much of his spare time while living there in the years 1902-1904. Here he was well received by the curator of mammalogy and ornithology, W. H. Osgood, and made a careful study of their exhibits.*7 On the way back from Chicago he managed to stop at Detroit, sell his house, and bring his mother and sister back to Ottawa with him in mid-July. His next task was to write a report on his trip to American Museums. After outlining the same itinerary that he described in his letters to Fleming he said how much time the curators spent showing him round and explaining plans, methods, and in giving advice. As a result of this visit he felt that the general scheme of exhibitions he had previously out- lined need not be altered. The trip had served to crystallize his thoughts and had brought his methods into sharper outline. His most important recommen- dation was for a staff of museum heads to work under the curator in charge of natural history. Also a - good mammalogist was very necessary as well as a herpetologist “if we are to do work that will measure up to the standard set by the serious museums on the other side [of the border].’*? Now that he had visited some outstanding American museums Taverner was consciously measuring the future development of his 54 THE CANADIAN FIELD-NATURALIST own museum by their standards. A preparatory department was also needed with a good taxidermist and an assistant. He pointed out that the system of sending taxidermy work out to commercial firms did nothing to develop skilled craftsmen along scientific lines. No one could realize the great steps that had been taken in museum methods and techniques until they had seen the work being done at the most advanced museums in the U.S. But, Taverner warned, good taxidermists were scarce and hard to “draw away” from their present employment. Other problems of less magnitude, but none the less troublesome, faced Taverner in his task of orga- nizing the collections of mammals and birds in the new museum. There were only two workrooms in the basement which were quite insufficient for the preparatory work. At that time there was no heating in the basement, though it was proposed to install electric heaters, but Taverner doubted that they could supply the necessary heat. “I do not see how the work can proceed without gas. This is the only museum in the country and perhaps in the world that is not supplied with gas for the necessary heating process.” He advised that the study skins should be on the top floor away from basement dampness and near to the offices. For cabinets he recommended the new design recently developed at the American Museum which had sliding doors that “are dust, insect and fire proof’. As regards the catalogue system for verte- brates he proposed four distinct catalogues, one each for birds, mammals, reptiles and fish. Then, as spe- cial curators were appointed, the records of each department could be turned over to the proper offi- cers, while Taverner kept the bird records. Registers and index cabinets were needed to begin the record keeping. Exhibition cases should be ordered and a typewriter acquired. “The present one is my own property and badly worn and besides cannot be used for writing index cards” which, he explained, was a prime need in his depart- ment. He would shortly submit plans for dividing the upper floor into offices and specimen rooms. Finally, so that there should be no misunderstanding, he made a resumé of the points which required immedi- ate action. He signed himself “Assistant Curator”.*4 Work now began on the arranging of the bird specimens by years according to the dates of arrival at the museum. As he explained to Fleming the con- dition of the labels was very bad. Luckily he was able to make the temporary appointment of a young man by the name of Hennessey to help him with this work “until college opens”.*> The re-labelling was tedious work which involved tracing the history of the accessions already belonging to the bird collec- tion by carefully deciphering obliterated labels, searching maps for little-known localities, and research among various reports and old records pre- served in old registers and manuscript lists.*° Vol. 110 Taverner’s other main preoccupation was with his catalogue system. He abandoned the idea of a card to a species in favour of a card to a specimen because it seemed a better way of keeping track of loans, and for cataloguing distributional studies. Tired of wait- ing to have the printing he needed done by Government he found a small printing press in the building. With this he made enough cards to try out his method, sending some to Fleming for his com- ments. The name of the province was printed in the corner of the species cards. “Tam getting a large Dominion map to lay out on my work table with glass over it. I intend to mark on it all the localities represented in the collections and the routes of various expeditions. I think it will be very handy for I can then lay my species cards out on it according to their locality and thus get the distributions at a glance.”>7 This is an interesting statement because it pin- points the origin of one of Taverner’s major contri- butions to Canadian ornithology, namely the sys- tem which he devised in 1911 and maintained until his retirement in 1942 for keeping track of the dis- tribution of Canadian birds. When the system was finalized it consisted of a series of large-scale maps in a binder on a large table on which bona fide records of birds were plotted, and the information for each record was noted on cards arranged by species and filed in adjacent cabinets. There were two sets of cards: one dealing with published litera- ture, the other set with unpublished information derived from correspondence and field notes. Fleming replied to this letter with good advice on cataloguing skins and the value of a printing press because one can print the locality where a bird was collected and the name of the collector. Moreover, unlike any other ink, printer’s ink is permanent. Using a card to a species it ought not to be neces- sary to give the scientific or common name, the AOU number on the cards should be sufficient. But Fleming had forgotten that the AOU numbers might be changed from time to time, causing trouble to the user of this method.°* In answer to Taverner’s news that at long last his appointment was formally acknowledged and that he had received his first regular salary Fleming moral- ized “beyond keeping out of debt and enough ahead for sickness you can live comfortably and go on with your work.”*? It seems that Percy was still Fleming’s protegé. Meanwhile Fleming was always ready to put Percy in touch with people who might help him in building up the museum collections. Late in 1911 he wrote that he was in touch with a man named Munro in British Columbia about the possibility of getting Trumpeter Swans. “If you want anything collected at Okanagan Landing J. A. Munro will be glad to get you skins, if you want anything for groups he is open to engagements and would be intelligent as well as reliable.”4° 1996 As a result James Munro became a collector for the Museum and a lifelong friend of Taverner. This raised the subject of what was a fair price to pay for birds bought by the Museum. Percy had recently written to Fleming about the possibility of selling his own collection to the National Museum, and asked him to suggest a price. Fleming replied that fifty cents per bird was well within the range, that collect- ing birds in the field averaged $1.00 per bird.*! Eventually Taverner’s own collection of over 1000 skins was acquired by the Museum at 50¢ each. By the end of the year he notified Fleming that the regis- ters had arrived, and that he had sorted the mounted birds and the study skins into accession dates ready for cataloguing. In 1912 Taverner’s horizons expanded with the prospect of acquiring more bird specimens. By now 3900 specimens had been catalogued but more remained to be entered. The Museum still wanted almost everything. Only southern British Columbia and the international boundary to Manitoba were well represented but most of the far northern materi- al collected earlier by G. M. Dawson, R. Bell, and others had disappeared. Taverner knew that to build up the collection of specimens it would be best to start collecting systematically each summer with a zoology team organized by the Museum. One project he had in mind was to work the islands at the mouth of Georgian Bay and then across north of Lake Superior to the Manitoba line. “All this is terra incognita to us.”*? Meanwhile things were going for- ward very slowly at the Museum as though some- thing was wrong. In March 1912 Taverner informed Fleming that he had had a long “conference” with Brock. Brock seemed to be afraid to talk “right out’, and although he had good ideas he did not seem to be putting any of them into practice. Taverner sus- pected that Brock was hampered by the old staff at the Geological Survey; that he doubted their capabil- ities but hesitated to promote new people over them. Things at the museum seemed to be at a standstill, and the only thing that seemed to be developing was the crack in the tower. Taverner commented: “The crack is doing itself proud, growing and flourishing like the proverbial green bay tree. Guess as soon as spring weather comes we will lose our tower.”*? To this Fleming replied: “T hope you have taken out that accident insurance I told you to, it will be a tower of strength to you if that crack widens.” In the same letter Fleming gave Taverner some news of the Royal Ontario Museum which was now in the planning stage.** From 1912 until the end of his career in the National Museum thirty years later Taverner kept up an astonishingly large correspondence with other ornithologists, both professional and advanced ama- teurs in Canada and the United States, much of which was about technical matters. There was plenty CRANMER-BYNG: A LIFE WITH BIRDS 5) of challenging work to be done and Taverner soon took up the challenge. Writing to Fleming early in 1912 he mentioned that so far he had been concerned with getting each bird properly labelled, numbered and entered and had therefore paid very little atten- tion to “determinations”.*° This is a term in taxono- my that is applied to the process of determining to what species, or subspecies, a particular specimen belongs. In order to make an accurate determination a taxonomist may have to compare a specimen with many others in other collections. Determining the correct identification of speci- mens is one very important aspect of the work of a curator in ornithology and is a continuous process. As an example, the museum had three skins of sand- piper which Taverner considered to be the Aleutian Sandpiper, a subspecies of the Purple Sandpiper. These were in “intermediate” plumage. He explained in a letter to Fleming that the museum had some mounted Aleutian Sandpipers which would have been useful for comparison but these were in full plumage and so could not be used. He sent Fleming two skins to examine. Also in this exchange of let- ters Taverner mentioned that he had a breeding record of a Great Gray Owl from Nippissing District, Ontario, a bird which was in downy plumage. Fleming asked for more information explaining, “We don’t know what the first plumage of the Great Gray Owl is, so if you have found it you are in luck ... how many and what feathers did you get?”*° Taverner had received feathers from the wing and tail which identified it, and his informant said the bird was still largely in down. Taverner arranged to buy it for the museum for $5.00. When Taverner moved to Ottawa he left behind the birds and migration of the Great Lakes Region, with which he had begun to be familiar, and sudden- ly had to face the bird life of the whole of Canada including the Pacific and Atlantic seaboards and islands, the Canadian arctic, and the migration pat- terns between the whole of Canada and the United States. He started from very limited knowledge and needed as much help as he could get from experts. Fleming was valuable for eastern birds but less so when the intricacies of western birds were involved. What Taverner needed was an experienced ornithol- ogist living in British Columbia with whom he could correspond regularly. Meanwhile he did not let mat- ters rest for long. In a letter to Brock he set out the immediate needs of the museum such as the problem of pests eating the specimens. He hoped that the points he had explained would show Brock the need to consult with those who were best qualified as judges of their own special requirements. The muse- um building was very poorly planned because it failed to embody practical museum ideas. Such mis- takes could only be avoided in future by consulting those who used the building. He asked that an oppor- 56 THE CANADIAN FIELD-NATURALIST tunity could be given members of the staff to present their views and requirements before important mat- ters were decided. The letter contained a note by Macoun typed below Taverner’s signature endorsing Taverner’s letter.4’ A few days later Taverner had to write to Brock again, this time to put on record that his typewriter (his personal property) had broken down and was too old to be worth repairing. He asked for a new one, and in the same letter pointed out that it seemed poor policy for him to spend his time typing when a stenographer could do it more quickly and better than he could and at less expense to the museum.** In May 1912 Brock worked out a plan of organi- zation for the museum and set up a committee of all Geological Survey officers interested in the museum. The executive committee consisted of H. I. Smith, archaeologist, L. M. Lambe, paleontologist, R. A. Johnston, mineralogist and Taverner representing the biological division.” This gave him leverage to push for hiring additional staff and in July a proposed list of specialists for the biological division, including a taxidermist, was adopted.~° At the end of April Macoun, now in his eighty- first year, left Ottawa accompanied by his son Jim for Vancouver to spend the rest of his life in Vancouver Island happily collecting specimens of the flora. Macoun and Taverner said goodbye, and although they never met again they kept in touch with each other by correspondence. Taverner had come to like John Macoun as he made clear in a let- ter to Allan Brooks. “Prof. Macoun though he admits knowing little of birds is a wonderful old gentleman in many ways and since I have come to know him have had a great admiration for him:”>} In 1908 a separate topographical division of the Geological Survey had been set up to meet the need for making accurate topographical maps. Taverner discovered that members of the staff were interested in the work of the zoology department and would gladly bring back material from field expeditions if they nad a little instruction in its preparation.” To supply this need Taverner began preparing a set of instructions for the collection of zoological speci- mens as well as a method of note-taking based on the monthly notebook he had devised for himself in 1905. A few months later these instructions were produced in booklet form by the Government Printing Bureau.** In the introduction Taverner pointed out that visitors to Canada on hunting or sci- entific expeditions usually deposited the results of their excursions with their national repositories such as the Smithsonian Institution and the British Museum. He appealed to Canadians whose work or sport took them to out of the way places in the Dominion to remember the Victoria Memorial Museum as a legitimate recipient for any zoological specimens they collected but did not wish to keep as Vol. 110 personal trophies. Specimens acquired by the muse- um would be kept safely and put to the best scientific and educational use. They could also help by making notes and gathering information on the distribution and habits of Canadian wildlife, because records from localities little studied were invaluable in esti- mating the zoological resources of the country. The instructions covered birds, mammals, reptiles, amphibians and fish, insects as well as land and fresh water shells. In a general section on zoological collecting Taverner set out principles for what to collect and how to collect it, and incidentally gave the reader a useful summary of the outlook of an ornithologist and mammalogist working in a museum. For instance a good rule when collecting in localities about which little is known is take what seems most characteristic of the place, because what is recog- nized as common in many places may well be rare in a little worked locality. Certainly collect species that are known to be rare but also collect common species to give an idea of local conditions. As regards birds the most characteristic forms of any locality are the summer residents; specimens of breeding birds are essential as they give an idea of local ecological conditions better than do migrants passing through. Many species vary geographically. Thus a species found in eastern Canada may differ noticeably in coloration or size from the same species in western Canada. On the other hand the forms of a species or family which are found in a lit- tle studied locality, may be identical with those from a well studied locality, but are nevertheless valuable as showing a lack of variation in that locality. But specimens are needed as proof. To drive home his point Taverner coined an aphorism. “Never neglect the inconspicuous; striking beauty or unusual form are no criteria of scientific value.” For study collec- tions specimens were required of each species in every possible natural condition, in every plumage or pelage, worn or moulting as well as full breeding plumage or thick winter coat. Specimens of young, of moulting or changing plumages or birds, or sum- mer coated animals difficult to find, were especially wanted as the museum was weakest in this line. After stating the principles of collecting Taverner set out the principles of preparing specimens in the field and the care of skins. Again he highlighted cer- tain basic rules. For instance that no matter how rare or valuable a specimen may be, unless full data on it are attached it is practically worthless. A good label without a specimen has a certain value but a speci- men without a label is valueless. He stated exactly what information each label should contain, such as the altitude at which a bird or mammal was collect- ed because, he explained, altitude has a lot to do with distribution. The sex of the specimen should always be stated. One must determine this by dis- 1996 section. Do not put your own inferences or conclu- sions on labels, only the ascertained facts; all other material should be recorded in your field notebook. Next, Taverner went into considerable detail drawn from his own experience on the best equipment to use. “A pair of field glasses is nowadays almost indispensible to an ornithological field worker.” Modern prismatic binoculars, he said, were best for the job. Then followed a very detailed description of how to skin and “make up” a specimen down to the formula for making arsenical soap. All the informa- tion and expertise in this section was based on the first hand training he got from A. B. Covert at the University of Michigan Museum, in his many dis- cussions with Fleming, from taxidermists such as Oliver Spanner and from work in the field with Saunders at Pelee. He also learned much about basics by continual reading of Coues’ Key to North American Birds. When writing the section of the booklet dealing with mammals he had to rely on information in books, and from the advice of taxidermists who were expert in making up mammals of all sizes. The “sport” of hunting animals and having them made into trophies was widespread and Taverner himself had seen many such trophies being mounted in taxi- dermy shops in the past. For the section on insects he could use the expertise of “Bugs” Young at the Museum, while A. G. Ruthven of the Museum of the University of Michigan supplied advice on collecting reptiles, and Bryant Walker, a friend from his Detroit days, allowed him to use his pamphlet on the collec- tion of land and fresh-water shells. The writing of a booklet of this scope showed something of Taverner’s flexibility and experience, and the value of his network of friends and acquaintances. In the _ same period he prepared a field note-book for record- ing daily observations on flora and fauna. He expect- ed that this would be of considerable use because it could be kept by members of the various staffs of the Geological Survey without interrupting their field work. In his report for the year 1912 he explained that “by their use we expect to obtain exact data on comparative abundance and distribution of many species in many out of the way localities.”>4 But no plans to enlarge the collections significant- ly could succeed without a competent taxidermist. As a result of his visit to museums in the States in 1911 Taverner had heard of a taxidermist by the name of Clyde Patch who might be willing to come to Ottawa in the future, but nothing further could be done until a post existed for one. This happened in July 1912 when the Museum committee agreed that such a position should be requested. After Taverner had attended the AOU meeting at Cambridge, Massachusetts, in November 1912 he took the opportunity of going to New York, in the company of C. H. Young, and spent a week in the American Museum of Natural History and the Brooklyn CRANMER-BYNG: A LIFE WITH BIRDS Dil Museum. While there he was able to meet Clyde Patch and discuss details with him. The prospects seemed promising. Writing to Swales he explained: “In New York I think I found a jewel of a taxider- mist and expect to get him here later on in the win- ter. I need him badly. I am planning on taking him to Point Pelee next spring to make a collection for a Canadian Carolinian group, as the first of a series of geographical distribution exhibits.’”°> Clyde Patch was appointed chief taxidermist to the National Museum from | May 1913.°° In a letter to Fleming in December 1912 Taverner looked back on the slow progress he appeared to have made in organizing the zoological department since his appointment. He realized that the work of cataloguing and organizing, while extremely impor- tant, made no show at all and was appreciated by hardly anyone at the museum. However, he felt that he had made pretty good order out of chaos. “You can hardly imagine what a disgraceful order or lack of order things were when I came here.” In the same letter he mentioned another matter which was that until Clyde Patch was secured he would be unable to mount any exhibitions. He was very anxious to make a showing because he could hardly expect a raise in salary until he had shown what he could do. In the zoology section of the annual report for the year 1912 he sounded a note of urgency about the need to collect specimens before it was too late. He suggest- ed that in addition to the Geological Survey other survey and exploration parties were sent into the field, and if a naturalist could be attached to them specimens could be collected inexpensively. Taverner showed foresight when he wrote: “The Canadian country is changing rapidly from an unsettled state to that of civilization and cultivation. This is having a most profound effect upon our flora and fau- nal life, and vast changes are being brought about in our biotal [biotic] conditions. The old order is passing away, in many places has already passed, without leaving a record of its being behind. If the next generation is not to charge us with being indifferent to their interests we must improve every opportunity of making record of present conditions. The time for this work is now, for every day means some loss on the pages of our records that can never be filled.”*® Ottawa in the year 1911 was a small city of about 50 000 people. Scenically it was very attractive, part of an area being built around three major rivers and their confluence — the Ottawa, Rideau and Gatineau rivers — as well as the Rideau Canal. Beyond it in the near distance looking northwest stands the low line of the Gatineau hills — in the winter showing grey or white — but in autumn a fiery background of orange and gold. The Rideau Canal, originally built to link the town of Kingston on the St. Lawrence River with Ottawa, gives distinction to the capital city with its neat locks and the trees and flowering shrubs lining its banks.*° 58 THE CANADIAN FIELD-NATURALIST But in the spring of 1911 Taverner was in no mood to appreciate the scenery; what he longed for was a home of his own in Ottawa. After looking at the price of land, with his experience of building behind him, he decided to wait before buying a lot because prices were too inflated. As he reported to Fleming: “You remember those lots we pass[ed] on the car by the Experimental Farm? They want $900 for a 50 foot lot way out there. They are opening three subdivisions all around the city and I cannot see that the city is growing to an appreciable extent ... it looks to me like a Real Estate man’s boom and prices will drop heavily soon.” Also, on account of his mother he was not pre- pared to buy on the outskirts of town where few con- veniences as yet existed. “After having a comfortable home it would be more or less like roughing it to her and it is hardly fair to ask her at her age to try and get along without the comforts she has been used to unless there are other very great advan- tages.”°! By mid-July his mother and sister were with him in Ottawa; in August the sale of their Detroit house was completed and he began looking in earnest for a location in which to buy a lot. At this time they were living in a house immediately south of the canal near Bank Street and liked it so much that if they could find a lot in the area at a reasonable price they would buy there. As he explained to Fleming: “Tt is really the prettiest part of the city in my estimation and the closeness of the canal water makes it desirable if one wants to do any boating.”©? Fleming was anxious to know how Percy’s mother liked living in Ottawa, especially since at first she was likely to feel lonely. To encourage them he said: “T fancy once you get to know people you will all like it. There are more people of culture and intelligence in the civil service than are usually to be found in a city of Ottawa’s size and consequently there are [people] inter- ested in things other than money getting.” The Taverners found a suitable lot near to where they were renting. He told Fleming: “Tt is a small farm — 88 X 132 over near where we are now living. It has no trees on it but it is within sight of the canal and the beautiful elms at the end of the street form a vista half a block away. I have my plans all pen- cilled in and expect to start building as soon as I can get them finished and figured. All I fear now is that it will run high. If only I can build it as I have it drawn it will make a most comfortable and beautiful place ...’°+ It also promised to be a good location for birds on migration as Taverner told Fleming in October. “Where we live are a fine lot of big elms and beeches just across the road and this fall they have been filled with birds of all kinds. A flock of Whitethroats [White- throated Sparrows] found that the rigs crushed the acorns that fell in the road at the side of the house and have been feeding there on them in great numbers.” He also sent Fleming the plans he had drawn for the house, and a photo of the model he had made for it. Fleming’s mother was somewhat critical saying that it looked like a Chinese pagoda, but Fleming Vol. 110 offered several practical suggestions for improve- ments.°° Building started in 1912 and by early April the — foundation was almost “up to grade”. Percy described their excitement especially since it was his own design. “We are all watching it with the greatest interest as like- ly you can imagine. Mother takes her first look in the morning and the last at night out of her window across country at the scaffolding that marks the concrete forms and can think of little else.” Writing to Arthur Andrews, his entomological friend in Detroit, he said: “Yes, we still like Ottawa immensely. There are a very nice lot of people here that take intelligent interest in things and can talk about something besides the last baseball game... Mother and Ida are delighted with the people and the place. Ida is right in a musical crowd and Mother is realizing the dream of her life and is studying French to beat the band.” However, Percy was not much impressed by the buildings in Ottawa. “The city, too, is delightful, though architecturally rot- ten. We are right at the base of the Laurentian hills which rise just across the river and half an hour will take us right in the Canadian fauna while around to the south all is transition.” But building a house was not all smooth going. “At present I am filled with the bitter-sweet pleasures of building a house for ourselves. It is lots of fun but lots of worry as well. We have had all kinds of trouble but at last things seem to be straightening out. They started laying brick above grade today after having to tear a great lot down to put a water-proof grade course in. We have a fine big lot 88 x 134 and are planning a garden that is a garden. We are only a block from the Rideau Canale72 Mrs. Fleming had been visiting Ottawa in the summer and reported an outbreak of typhoid fever. Fleming urged Percy to take precautions to which Percy replied: “T take care of what I drink and eat. We use nothing but boiled water in our house for everything in connection with food, teeth cleaning etc., drink no water away from home unless we know that our hosts take the same pre- caution. Having done this much I see nothing else to do and try to forget that there is typhoid in the city.” With museum work and house building Percy had very little time left for correspondence other than to Fleming. But in October he managed to type a long letter to Bradshaw Swales, pleading delays in build- ing for not having written sooner to old friends. At last the house was finished and they had moved into it, though there was a lot of grading, filling and gar- dening to be done. He added a piece of family news. “Ida passed the Civil Service examinations and was installed here in the library of the museum.’’”? With a convenient and attractive-looking house of their own Percy, his mother and sister settled down to enjoy their family life and get on with the next project — a really good garden. The only cloud was that the house cost more than estimated and Percy found CRANMER-BYNG: A LIFE WITH BIRDS Sy) The Taverner house, 45 Leonard Avenue, with construction near completion in 1912. (Reproduced courtesy of the Canadian Museum of Nature, number 41307. himself under financial strain finishing the payments on it. He had anticipated a raise in salary but when that was delayed he was only just able to make ends meet.’! But Taverner was used to that problem. Another problem which Taverner had been famil- iar with for some time was how to cram into each twenty-four hours all the activities to which he was committed. His appointment at the National Museum made him drive himself very hard. He was always “up to his eyes” in work. Before each year came to an end he had to begin preparing a report for that year, covering the zoology section, for printing in the Summary Report of the Geological Survey of Canada. In November 1913 he began writing his report for that year. He had now been wrestling with the problems of the section continuously for twenty months and could feel that he had achieved consider- able progress. For Taverner the year 1913 was a year of systematization when a great part of the catalogu- ing and arranging of the old collections was complet- ed and new projects started. Particularly important to him were the people he was working with. At the top was Reginald Brock, director of the museum. Taverner had already expressed some frustration over what he considered Brock’s lack of forceful- ness. Now he complained to Fleming of Brock’s lack of communication with the representatives of the various divisions serving on the recently formed museum committee. But his good qualities out- weighed his weaknesses. Taverner criticized Brock’s handling of a stalemate among the members of the museum committee over the format and the names of museum publications. But then he wrote: “The more I see of Mr. Brock the better I like him though I think it would work for much more satisfaction should he [let us] in on his intentions a little more.’””” Taverner was not diffident in writing a letter to Brock when he wanted something, or had an idea for improving some aspect of the museum. For instance he asked for the occasional use of a stenographer. Brock sent back his letter with a hastily scribbled note on it that the museum had provided in the esti- mates for permanent stenographers. Late in 1913 Winifred Bentley was transferred to the zoology sec- tion as assistant and typist.’* In another letter to Brock on the same day Taverner raised the question of the name of the museum in a nice piece of Taverneresque logic. He suggested that in future less stress should be put on the title of Victoria Memorial Museum. 60 THE CANADIAN FIELD-NATURALIST “As I understand it that name applies only to the build- ing we are in and means no more than the ‘Jones Block’ does to a law firm having offices in it.” He argued that it was an unfortunate choice of name since there were two other museums for. which it could be mistaken, one in British Columbia and one in Australia. He suggested that authority should be obtained for using the name Canadian National Museum which would be more appealing to the national spirit of Canada. He concluded: “The inaptness of our present... name is obvious if we consider what would happen if we were moved to anoth- er building, then what would we be?”’”4 Brock wrote on the letter “Good suggestion bring it up at museum committee”, and returned it to Taverner. The name was not changed until 1927. Another letter he wrote to Brock at this time was about his salary. He said that he understood at the initial meeting before he was appointed that it was lack of actual museum experience that caused his starting salary to be placed at $1600.00 instead of $2100.00. “T think I have shown that in spite of having never been formally engaged in museum work I understand its prac- tical requirements fully as well as though I had and that this objection is therefore removed.””° Brock could not know how strapped for money Taverner was at this time. His salary increased in the following year when he was promoted in rank. Other matters on which Taverner reported to Brock as his superior were: A request from Maxwell Graham, Chief of Animals Division, Parks Branch, for his opinion on the introduction of English song birds into Canadian National Parks.’° A request from James Harkin, Commissioner of Dominion Parks, for his written opinion showing reasons for establishing wild animal sanctuaries.’’ He also reported to Brock regularly on loans of study specimens made to other institutions or individual collectors of good standing. In the case of making permanent exchanges of bird specimens between the National Museum and anoth- er institution or individual he would make a recom- mendation to Brock.’* Such exchanges were quite regularly made by museums in order to add to their collections, particularly concerning specimens they lacked. It worked well so long as the judgment of the curator responsible was always sound, and one’s own museum always got a fair exchange. The reason for sending a recommendation to a senior authority was to keep a check on the proper working of the system rather than to obtain his advice, since only the head of a department was in a position to esti- mate whether a particular exchange would be to the advantage of his own museum or not. Another of Taverner’s colleagues was Jim Macoun, acting head of the botany section. They appear to have got on well together in spite of Macoun’s dislike of Brock. The correspondence which has survived between the two men was mainly about botanical matters.”” A member of the zoologi- Vol. 110 cal division with whom Taverner was on good terms was C. H. Young the taxidermist, mainly responsible for preparation of invertebrate material (hence his nickname “Bugs” Young) and a willing worker who produced highgrade results. They worked together during Taverner’s first field trip in 1913 (see Chapter 8). The only other member of the museum staff whom Taverner appears to have respected was Harland Smith of the archaeology section of the Division of Anthropology. The year 1913 was an important one for Taverner’s future career because it was then that two long associations began that were to continue throughout his professional life. In the spring of 1913 Clyde L. Patch was appointed taxidermist in zoological preparatory work at the museum and was to work with Taverner until the latter’s retirement. Also in the spring of 1913 Rudolph M. Anderson first began to impinge on Taverner’s life when he was appointed to the museum as mammalogist. When Taverner first heard that Anderson was going with Vilhjalmur Stefansson to the Arctic again he wrote to Brock about the possibility of getting Anderson a museum appointment. Brock had already written to the Minister of the Interior and the Prime Minister recommending that the government con- tribute $25 000 and that Canada should participate to some extent in the expedition. The government decided that Canada should sponsor the expedition exclusively, and as a result Anderson was appointed.*° In reporting this promising turn of events Taverner grew quite eloquent in a letter to Fleming. “Anderson got a most remarkably fine lot of stuff last year with Stephenson [Stefansson], as an example 19 bears and over 900 bird skins. As it was entirely a foot expedition there is no knowing what he can do on this occasion with an annual ship to work from. Anderson will be more stationary than Stephenson and will work more about the base and have more facility for storing and caring for stuff. I understand that he makes up beau- tiful specimens. He has had some Museum experience in Kansas. He is the man that wrote The Birds of Iowa in 1907, but is preparing to specialize on mammals. Personally I understand from Brock he is a most delight- ful fellow. B. says he is quite taken with him and has been making extensive inquiries as to his ability with most satisfactory results.””*! Taverner was very anxious to have a mammalo- gist appointed who would take the responsibility for the mammal collection off his shoulders. He had not been trained or appointed to preserve and display mammals; the care and development of the bird col- lection was ample work for one man. The only draw- back about Anderson’s appointment was that he would not be available to start work in the museum for an indefinite time because he would be exploring and collecting in the Arctic. Taverner, meanwhile, was looking for opportuni- ties to fill the large number of gaps in the museum’s i ' Victoria Memorial Museum Building, about 1913. Viewed from the northwest looking past the building toward Elgin Street in background. The massive main entrance tower, on the north side, was subsequently removed after it start- ed to tilt and sink into the underlying clay. (Public Archives of Canada photo 9273, print courtesy of the Canadian Museum of Nature.) collection. He required species, and subspecies, not already represented in the collection in order to be able to work out the range of the subspecies. Also he needed series of various plumages, such as breeding and intermediate plumages, male and female, and if ' possible, between various juvenile plumages.*” i One piece of luck that came Taverner’s way was when Fleming presented the museum with a collec- / tion of 350 birds, most of them mounted, but some | study skins. In his Summary Report for 1913 Taverner noted that the workmanship on the mounted birds of the Fleming gift was especially good, while the collection included some rare species and some type specimens. He also pointed out that it was “par- ticularly gratifying to receive such a valuable gift from a private individual, for it is through the interest and beneficence of public-spirited citizens, only, that a national museum can attain greatness and impor- tance.” Another point of importance was that the gift was made without conditions “so that for all time the curators will be free to make the most valuable use | a Rit ca ee Ps CRANMER-BYNG: A LIFE WITH BIRDS 61 possible of the material.” Conditions, Taverner point- ed out, change during the course of time and restric- tions accompanying a gift, however wise they appear at the time, may in the long run prevent it from being used for as long as it might have been, had it been given without strings attached. “Mr. Fleming, with his knowledge of museums and museum work, has made no such mistake.’’*? Other additions came through purchase, and by collection in a summer field trip to Point Pelee (Chapter 8). Two mounted Passenger Pigeons were bought, one of them from the Toronto taxidermist H. H. Mitchell for $50.84 Another purchase was made from a private fur trader, Clement Lewis of Whitehorse, Yukon Territory, of 211 birds and 380 mammals from the Teslin Lake region near the Yukon-British Columbia boundary.*° What pleased Taverner especially about this collection was that it included several specimens of Blue Grouse collected in October 1912. When compared with the two sub- species recognized as ranging into Canadian territory 62 THE CANADIAN FIELD-NATURALIST they were seen to be somewhat different. This gave Taverner the opportunity to attempt to establish a hitherto unknown subspecies of Blue Grouse by a very detailed description of plumages. These birds were smaller and darker than Richardson’s Grouse [Blue Grouse] which they most resembled. After borrowing summer and autumn specimens of Richardson’s Grouse from the U.S. Biological Survey for comparisons Taverner wrote a description of the Teslin Lake birds as a new subspecies of Blue Grouse, from Southern Yukon Territory. He named ‘it Dendragapus obscurus flemingi after J. H. Fleming “in recognition of the value of his ornitho- logical work in Canada.”*° The proposal was accept- ed by the AOU and published in The Auk thus giving it the stamp of authority. This is an example of Taverner carrying out his duty as curator of the bird collection at the National Museum at the highest level of ornithology by determining whether any of the specimens under his care were viable new sub- species hitherto unrecognized as such. This particu- lar subspecies lasted a reasonable length of time and then was “knocked out of the list” when the AOU Committee on Classification and Nomenclature car- ried out a purge of some of the Blue Grouse sub- species by lumping them into two only.®’ Possi- bilities for acquiring new specimens for the museum were liable to occur quite unexpectedly as when the professor of botany at Queen’s University called to see him. Taverner learned that Queen’s had some of the birds brought back by Robert Bell from his Hudson Bay expedition. In addition to building up the bird and mammal collections Taverner was anxious to make the zoolo- gy section attractive to visitors and of value to the public educationally. At the same time as he gave a collection of birds to the National Museum, Fleming deposited, on loan, a number of mammals including the only two Ontario-killed cougars known to exist. Another specimen of popular interest was a wolf, one of the original wolf pack made famous by Ernest Thompson Seton in his story “Lobo, King of the Corrumpaw”. Writing to Fleming about the loan he said: “I should much like the wolf for popular exhi- bition. Anyone who has read the King of the Corrumpaw will be interested in it and I would like to make the Museum interesting from as many stand- points as possible.”** In the same letter he mentioned that the museum estimates were on the way to being passed and that $25 000 was being included for the bird range rooms and their cases. “The Dept. of Public Works hope to have it ready by July. Things do move though Government work is like the mills of the gods.”*®? Taverner was becoming more sure of himself as a museum man by now. In spite of continual delays and difficulties he had already achieved a state of order and had something to show for his efforts. His nature had not changed in spite of a permanent Vol. 110 salary. There was no slackening off. If anything he was working harder than ever and enjoying it, and even finding time to take on extra work in addition to the museum by making an ambitious garden. In May he wrote to Fleming: “Ida has her typewriter at home writing me a set of specifications for a house I am planning for one of the members of the Survey and I am taking advantage of it to get off some pressing correspondence.””? He must have been under pressure, and the message he was sending his friends was “I am desperately busy”. He used two different expressions on the same day to depict his state of busy-ness. To Brad Swales he wrote he was “as busy as the Devil in a gale of wind”, while to Fleming he said “I am chasing around like a chicken with its head off.’”?! But that was his style. He had the bit between his teeth and was galloping with it. By the beginning of 1914 Taverner had been responsible for the development of zoology at the museum for two years and eight months. He had come through a difficult and frustrating time, had survived the challenge, and was now on the way to achieving some of the immediate goals he had set himself. But the conditions necessary in order to carry out his long-term plans for the expansion of the zoological collections depended on circumstances outside of his control. These included more museum space for research and display, more members of staff, and congenial colleagues to work with. Taverner might reasonably have expected that the year 1914 would be a good year for him now that things were running reasonably well in the ornitholo- gy section. But life is unpredictable. The year started with a personal setback for Percy and ended with a catastrophe for humankind. On Tuesday 13 January 1914 the temperature in Ottawa fell a long way below zero Fahrenheit. Furnaces were stoked up to keep pace with the bitter cold. Number 45 Leonard Avenue caught on fire. Percy turned to Fleming for advice. “Just a line. Was burned out Tuesday and am in a peck of trouble now with the adjusters... The house is a bad wreck, but all walls are standing, most of the roof intact, floor weakened and the wood work thoroughly ruined, but if I get any justice from Insurance Co. hope to be in old home again by May.” His house, he said, had cost $4700.00 and was insured for $5000.00. The furniture was insured for $1000.00 which would hardly cover the loss if the damage to the piano, which Ida played, was considerable. Percy was afraid that he might have to put matters in the hands of a lawyer and asked Fleming for his advice. He assured Fleming that he had sufficient financial resources for the present and that the holder of their mortgage, Dr. Saunders, was lenient and helpful.”” Fortunately his bird books and other things were in the museum but it meant that he and his family had to take a furnished house nearby until rebuilding was CRANMER-BYNG: A LIFE WITH BIRDS 63 Early exhibit hall and visiting public at the Victoria Memorial Museum, 1912. (Reproduced courtesy of the Canadian Museum of Nature, number 22217.) | finished.*? The insurance claim was settled satisfac- torily in February. Taverner had better luck over his own position at the museum. In a letter to Brock, written in March 1914, he noted that his appointment to the museum staff read “Preparator and Assistant Curator”. The first term was a misnomer and he asked that this be altered to “Ornithologist”. As he pointed out: “My _ work is chiefly ornithological and I regard my pre- | sent direct oversight of other branches, except per- | haps general exhibition, but temporary until compe- tent experts can be appointed to assume their | charge.”** R. M. Anderson was appointed mammalo- | gist in 1913 though he did not start his work in the museum until early 1917. Another matter that | Taverner was anxious to have put right concerned his salary. He had written to Brock in January 1913 | asking him to consider promoting him. (See above.) In August, Brock notified him he had been promoted | to subdivision B of the first division.?° By 1914 a major problem, which had been grow- ing since the collections were first displayed and properly stored in the new museum, came to a head. The problem was one of space, but this in turn grew | | out of a problem of organization. The Victoria Memorial Museum was originally built to house the collections of the Geological Survey, and was part of the Survey rather than an entirely independent body as most museums are. The museum director was always a geologist. This was a flaw in organization and administration from the beginning. It suffered from a “geological fault”. It was natural, therefore, that when the new building was being occupied in the period 1910-1911 members of the drafting and ‘topographical staff also moved in, though their duties had nothing to do with museum work. However, they and their field equipment took up essential space. The museum committee had drawn attention to this situation in a memorandum to Brock in July 1912 but without success. A second memo- randum was sent to the new Deputy Minister of Mines in October 1914. It was a strong indictment of the misuse of essential space required for museum work by the inclusion of the drafting and topographi- cal staffs in the building. The memorandum reported that the west wing hall on the ground floor was being used by these staffs as a freight shed and storage room for tents and other field equipment. No valid 64 THE CANADIAN FIELD-NATURALIST reason existed for its continued occupation in this manner. On the third floor the whole west wing hall was occupied as a drafting room. The memorandum described the overcrowded conditions in the base- ment and the other three floors, as well as a litany of the deteriorating condition of specimens in the seven sections of the Biological and Anthropological Divisions crowded into the museum. The committee warned that valuable materials were becoming more and more inaccessible for want of space. The condi- tion was rapidly approaching in which it would no longer be feasible to take advantage of opportunities to secure specimens of value when offered. The doc- ument concluded with the hope that those members of the Geological Survey whose duties had no part in operating the museum would be removed and the way cleared for the rearrangement of working space for museum purposes. The final paragraph opened up broader problems when the writers stated their conviction that “under proper guidance and with a freedom of action which it has not as yet experi- enced the museum will rapidly assume the place amongst Institutions of its kind which the people of this country inténd it to occupy.””° The words “prop- er guidance and freedom of action” pointed to prob- lems from which the museum was to suffer during the rest of Taverner’s lifetime. As though this problem was not enough, in August 1914 Brock suddenly resigned. He .had been treated somewhat badly by the administration over his posi- tion. He had accepted the post of acting deputy min- ister in November 1908 on the understanding that when Low was no longer able to carry out the duties of Deputy Minister of Mines he would be appointed in Low’s place. But even when it was clear that Low had become too ill to perform his duties he was still retained in the position year after year while Brock continued to carry out the duties of deputy minister without salary for the post. Not until January 1914 was Low retired on pension and Brock became Deputy Minister of Mines. The long battle to obtain sufficient space, equipment and supplies for the new museum, as it grew, probably made Brock tired of government administration and glad to become the dean of the Faculty of Applied Science at the newly founded University of British Columbia. He was replaced by R. G. McConnell, a well known geologist with long service in the Geological Survey. He was fifty-seven years old at the time of his appointment and he served as deputy minister until his retirement in 1921. Taverner had got used to Brock and could work with him, and found him supportive and reason- able, though he criticized Brock’s policy which, he wrote, “was to throw everyone in together and let them all find their own level.”®’ Taverner now had to get used to working under a less dynamic director of the museum. In wartime, as Taverner and his col- leagues in the museum were to find, government Vol. 110 finances were minimal. The museum was still an off- shoot of the Geological Survey rather than an inde- pendent institution in its own right. Not all Taverner’s time was spent inside the muse- um working on administration and studying bird specimens. To members of a museum staff, after long months cooped up within the walls of their “mausoleum”, the precious weeks or months spent in the field collecting specimens and artifacts came with a sharp sense of release. Taverner’s first field trip took place at Point Pelee in 1913 (mid-May to mid-July) and must have given him the same kind of feeling of escape from indoor routine, as well as challenge from the job awaiting him and his associ- ates in the wide outdoors. (See Chapter 8, devoted to the collecting trips undertaken by Taverner and his staff, and by professional collectors working for the museum, in the period 1913-1919.) Evidence which Taverner saw for himself while at Pelee made him strongly aware of the danger to birds from commercial hunting and by loss of suit- able habitat. From late 1913 Taverner became seri- ously involved in a problem which was to concern him for the rest of his life — the urgent need for con- servation (see Chapter 9). A further addition was made to the permanent staff of the zoological division in May 1914 when Claude Johnson became colourist in the preparation department. His main work was making coloured illustrations of mammals and painting backgrounds for habitat groups.?® Meanwhile, new specimens were regularly being acquired by purchase, often as a result of information from Fleming who was con- tinually skimming through bird dealers’ cata!ogues. For instance Fleming wrote: “Wards of Rochester were offering a California Condor for $35.00 recently and it seems to me we will have to include this bird in the Canadian list. It might be advis- able if the skin is a decent one to get it if you felt you want to eventually have a complete Canadian collection. I paid more than this for one in England last November and this is the fourth one I have seen for sale in 20 years. They are hard to get.”?? Fleming also located another Passenger Pigeon being offered for sale by a private owner in Toronto, and notified Taverner who acquired it for the muse- um for $25.00.!°° Taverner acknowledged Fleming’s help in his 1914 report for the zoology division. “Among the most notable of the accessions is one com- posed of some 113 specimens, obtained by purchase. This consists largely of extra-limital material of peculiar interest to Canadian ornithology such as European and southern types of forms allied to Canadian varieties. In obtaining these we are grateful to Mr. J. H. Fleming, whose active efforts and advice were of great assistance to us in selecting them.”!! At last the cases Taverner needed so urgently were being delivered; six new ones for the storage of birds and small mammals, and three cases adapt- ed to zoological exhibition. Ornithological work 1996 continued in the collections with the birds being determined in their systematic order. Taverner obtained help from eminent ornithologists in the United States such as H.C. Oberholser and J. Dwight as well as Fleming and Brooks in Canada. But his lack of identified specimens for comparison held up the work considerably. For instance, when working on the Great Horned Owls Taverner came to the conclusion that there were probably three “good” (reliable) forms in Canada: a white one, a _ black one and an Ontario one. As he told Fleming, | tion of éegree and I think he goes too far. “Here is where it seems impossible to do much without comparable and identified specimens. In size there is not over half an inch difference in any of our specimens except the white Manitoba birds which are small. My opinions on our geographical races is growing more fixed than ever.””!°? What Taverner means, I think, is that he is begin- ning to find confirmation of what geographical races occur in Canada, not that he has closed his mind to any further evidence. This is a continual theme run- ning through his letters with Fleming and Bishop and with other ornithological correspondents — the danger of accepting subspecies on other people’s determinations without examining them critically. To Fleming he wrote: “I do not think there is any question of Oberholser’s earnestness nor of his ability. It is only a question as to how far the sub-division should be carried. It is a ques- 9103 CHAPTER 7. Museum Problems (1915-1919) In the spring of 1915-an article by Taverner enti- _tled “Suggestions for Ornithological Work in Canada” was published in The Ottawa Naturalist. It was a manifesto of ornithology in Canada at that time and Taverner explained what needed to be done, and the kinds of studies that amateurs could usefully undertake. Taverner started by showing some of the main gaps in our knowledge of ornithol- ogy in Canada. Our information on the geographical ' distribution of birds in Canada was fragmentary, as he showed, while hardly a typical Canadian species _ had been studied as the subject of a life history. So much needed to be done that ornithology offered a fine field for original research. The southern penin- sula of Ontario was perhaps the only area of any size that had received adequate attention. Very little was known about the bird situation in the area from northwest of Georgian Bay to the Manitoba bound- ary, while there was only scattered information of conditions in the far north, often consisting of no _ more than short lists of birds. Very little original _ work had been done on the role of birds in the econ- omy of the country. Study collections in Canada were too small to permit anything comparable to work done in the U.S. to be accomplished in the study of Canadian birds. In addition, Canada lacked trained zoologists. CRANMER-BYNG: A LIFE WITH BIRDS 65 But from this we should not infer that Taverner, Fleming and several others were personally hostile to Oberholser and others who made subspecific determinations rather often. Taverner and Oberholser wrote to each other, loaned specimens to each other’s institutions for study, and talked when they met each other. Ornithologists belonged to an international community in which exchange of information was a recognized courtesy. Indepen- dence of mind and responsible criticism of col- leagues was accepted as necessary to keep ornithol- ogy in a healthy state. If anyone had told Percy Taverner four years previ- ously that in 1914 he would be established as ornithol- ogist at the National Museum of Canada, extremely busy running his own department with exciting oppor- tunities for field studies, he would not have dared to believe him. The frustration at having to earn his liv- ing at work that was uncongenial was a thing of the past. He said so openly in a letter to an acquaintance of his bird-banding days, J. Leon Cole, who had writ- ten thanking him for a separate which Percy had sent him, and asking about his job. Percy replied: “Of course my principle work is ornithology and I find it a great relief to do one’s chosen work as a livelihood instead of sandwiched in between the grind of pot-boil- ing. Being head of a division, building up a National Institution is in many respects an enviable one and I appreciate it.”! “Thus, it seems that ornithology in Canada still has most of its history before it . . . the work that should have been done by our own people has been accomplished by naturalists from the United States . . .”? Taverner noted an apparent failure to produce any ornithologists of marked ability in spite of the teach- ing of natural history in every public school in Canada. He believed that an introduction to nature in the schools “had failed to awaken any serious interest in natural problems.” But he remained optimistic that now, when museums were being started or rejuvenat- ed in various provinces, “the time seems ripe for a general wakening of interests in zoological subjects”. In order to stimulate interest in field studies Taverner suggested two main subjects that needed regular study. They were: 1. Life histories. These could be compiled by stalk- ing birds with a camera and notebook in order to study their behaviour. Valuable information could be gathered by watching birds around one’s own home. This was something that was both interesting and enjoyable. People with more leisure and opportuni- ties might wish to study less common species farther afield. By way of suggestions for further investiga- tions Taverner listed twenty six specific questions including: migration — is the species resident or migrant? Dates of arrival and departure. Courtship 66 nest building, brooding and chick rearing — how is this carried out? By one or both birds? How were eggs and chicks protected?* The list of questions can be expanded into many other aspects of a bird’s life history. 2. Bird distribution. At present, Taverner said, the published ranges of birds in Canada “are based upon geographic probabilities, a priori reasoning or are copies and recopied, from previous writers.’’> For example, lists of southern Canadian birds gave the Hairy Woodpecker, rather than the Downy Woodpecker, as the common form, where he consid- ered it a rare winter visitor. In order to establish the Canadian ranges of our birds, Taverner said, we need skilled observers at all possible points to collect local data and specimens. There ought to be an observer and a collection in every county, someone who was keeping track of his own area and comparing and checking it with the results from adjoining stations. Provincial museums should collect local details, and all the information should then be correlated at the National Museum. “In this way we would have cooperation and series of local collections illustrat- ing intensive work throughout the Dominion.”® But, Taverner warned, all such work, to be of use, must be based on exact personal knowledge and substanti- ated in every way possible — there must be data by which to check the statements made. The local observer should study the bird life of his chosen locality thoroughly. This would involve searching old literature and compiling a local bibliography. He should connect every record with a fully confirmed specimen, either in his own collection, or else note where one is located. In determining a specimen he should refer to those with greater experience if he is at all uncertain. “In fact, to a local faunal list it is better to add a hypo- thetical list for all species whose occurrence cannot be substantiated by specimens or on equally unimpeachable evidence. A long hypothetical list is often an indication _ of careful work rather than the contrary.””” As regards collecting specimens, the student of a locality should attempt to gather a representative series of all birds of the area showing every plumage in which they occur there. Single individuals or even pairs are not enough, nor is one example of each stage sufficient. “It is only by a series of several that the average can be established.” Even aberrations such as albino or melanistic specimens are of limited general scientific interest, “the normal is a much more desirable subject of study”.’ As a good source of information on specific occurrences the local taxi- dermist is someone to cultivate, not only to secure specimens but also to see what passes through his hands. It is also worth keeping in touch with the shooting and sporting fraternity. The best way to keep a collection is as dry skins rather than stuffed and mounted. Since the object is the indefinite preservation of the specimens, with the least deterio- THE CANADIAN FIELD-NATURALIST Vol. 110 ration, skins are much preferable. The effects of dust and light, as well as insects on mounted specimens are very destructive, thus making their life a limited one. Since specimens need to be examined closely in a good light and compared with each other, properly made skins are the answer. At this point Taverner discussed the vexed ques- tion of subspecies and how far it is desirable to study them, and the current system of nomenclature. He explained the reason for studying subspecies; that species were not permanently fixed, that geographi- cal variations do occur, the extremes of which show considerable differences when compared with the type specimen. Since various stages of differentia- tion (intergrades) between the “norm” of a species and examples of extreme variations exist, this seems to indicate that these are evolutionary variations, and should therefore be regarded as “species in the mak- ing before the connecting sequence between them and the parent stock has been disrupted or broken down to form isolated species.’ He then explained the scientific method of naming birds — one term for the genus to which it belongs and another for the species. But since it seemed use- ful to give supplementary names (cognomens) to geographical variants to distinguish them from the typical form a third name was added. After dis- cussing the method of naming the species with two names (binomial) and the subspecies with three names (trinomial) he then stated his own view. “Tt is... only necessary to name subspecies . . . where special exactness is required by context or scope of con- sideration. Subspecific designation should only be based upon examinations and authoritative determination of specimens, and not upon probabilities or assumptions.” 10 Taverner considered that, although in theory innu- merable subspecies of a widely varying race must exist, “most of them are too fine for human recogni- tion.”!! In the light of his strongly expressed views on the question of naming new subspecies Taverner summed up his position at this time in a conciliatory way. The question, he said, was not whether to make new subspecies or not, but where to draw the line [emphasis added]. Whether it is in the best interests of science to name variations that only an expert, especially trained, can recognize is still to be argued. However, whether we agree with the position of the “splitters” or “lumpers” it seems best for the majori- ty to follow the lead of the American Ornithologists’ Union. “However, for the sake of uniformity it is better to err on the conventional rather than the radi- cal side and to keep as largely as possible in harmo- ny with accepted contemporary authorities.” !? On the controversial issue of whether or not natu- ralists should ever shoot birds Taverner set down the axiom that “no enduring faunal work can be accom- plished without the collection of specimens”. He argued that collecting was a necessary evil but that 1996 CRANMER-BYNG: A LIFE WITH BIRDS . 67 due regard must be given to humane principles; that a permit to collect specimens was a privilege, and _ that specimens once taken should be regarded as a _ kind of public trust. They should be kept safe from _ damage, made available for study by others, “and | finally placed in some known repository where they | will be available for coming generations of investi- | gators.”!3 He disagreed with the fear that legitimate collecting would deplete Canadian bird life and argued that the millinery trade caused far greater | destruction of birds in one year than all the collec- } | | | tions made in North America during the past fifty years had done. Taverner also dealt with the feeling _ against scientists collecting “rare birds” on the | assumption that if they were allowed to breed they _ would become more common. But this, he said, was | not proved. Birds become rare from other, more per- vasive, causes. He pointed to the extinction of the Passenger Pigeon and cited the example of netting the birds in the trees of the Petoskey breeding “rook- eries” in Michigan. In the autumn of 1878 the birds left on their usual migration in large numbers, but few returned the following spring as if some calami- ty had hit them in their wintering range.!4 At the end of his “Suggestions for Ornithological Work” Taverner returned to the wish that the provinces would establish museums that would develop into repositories for provincial data. “In the meantime we have a Dominion Museum that is prepared not only to store but to scientifically use such _ material and is slowly building up a national collection __ for future Canadian students in proportion with the growing dignity of the country it represents. It is to be hoped that the time will come when it will take equal rank with other national museums of the world, the British Museum, the Smithsonian Institute and others of like repute. To do so, however, requires the co-operation __ and sympathy of the Canadian people as a whole. No __ public institution can do all the necessary work itself but _ must rely largely in the building up of its collections and prestige upon the interest and aid of the people it repre- | sents.””15 Taverner’s manifesto is important for the light it _ sheds on his own attitude towards ornithological _ needs and problems facing him in 1915. We should not judge it by our knowledge and attitudes of today but view it in the light of the problems which faced him as ornithologist at the newly evolving National Museum of Canada. It is here that we can read Taverner’s principles most clearly expressed. His “manifesto” marks a step forward in his career, and shows him beginning to demonstrate leadership at the national level. In the spring of 1915 Taverner was thinking seri- _ ously about writing a book. He asked Fleming for advice on a plan he had worked out for a popular bird guide which might have for title “A century of common Canadian birds”. It would be limited to birds of the provinces east of the Prairies. Fleming replied that he thought the plan was excellent and sent him some helpful suggestions.!© From early 1915, until the book was published in late 1919, cor- respondence continued between the two friends on the subject. The writing of his first bird book was a major step forward for Taverner. It increased his knowledge of ornithology in Canada and his under- standing of the problem of how to present it to the public. It also increased his self-confidence while making him better known. Taverner must have been very busy during the winter of 1914-1915 because early in 1915 he pre- sented his “Recommendations for the creation of three new national parks in Canada” in a report to the Canadian Commission of Conservation. He also wrote up his investigations of 1914 on the economic status of the Double-crested Cormorant in the Gulf of the St. Lawrence. With these reports Taverner assumed an important role in the growing Canadian awareness of the urgent need to protect wildlife in specific habitats (see Chapter 9). Meanwhile field work, begun the previous year at the Gaspé Peninsula, was continued there in 1915 by Taverner together with Young, Patch and Johnson (see Chapter 8). The trip lasted nearly two months and Taverner was faced with plenty that needed his attention at the museum. He told Fleming, “Back again and very busy clearing decks to get at the bird book and Gulf of St. Lawrence report. Had a delightful trip not many specimens but got a good bird’ s- eye-view of the country and some fine moving pictures and ordinary photographs of the Bonaventure birds’’.!7 Taverner kept a carbon copy of every letter about museum matters that he sent to friends. But now that Miss Bentley was working in the department he would have to be careful what he wrote, or else keep his letters locked up. For instance, writing to Fleming he mentioned Patch. “Patch is still in Perce. Evidently he enjoys being left alone, and head of his camp. He is a queer fellow and hard to get along with. Johnson, his assistant, however, is a fine camp mate and a good fellow.”!8 Fleming warned him to be very careful what he said in his letters if Miss Bentley could read them.!? Percy saw the humour of his remark and replied in a P.S. to his next letter to Fleming: “Miss Bentley not only can read your letters but she can read mine as well — that is going some.””29 Taverner’s handwriting was moderately legible but Fleming’s was more difficult to read because many words seemed as if written in a private form of shorthand. Several other ornithological problems and matters occurred during this period. For instance Taverner’s work on determining species continued, this time on gulls and geese. He turned to Fleming for help again. “If you can separate California and Herring Gulls, let me into your secret. I am all at sea over them. I do wish also some one would tell me what a Hutchin’s Goose is. Everyone has a different theory; none of which work out 68 the same with our stuff. Though they say that in life the birds are absolutely distinct.”’2! Fleming invited him to come to Toronto for help in identifying specimens and for advice on the colour and make up of his book.” Because the National Museum did not have sufficient series of bird speci- mens for all the comparisons that Taverner needed for scientific determinations he relied much on the collections of his friends, among which those of Fleming and Brooks were outstanding in Canada. Anything that affected their collections affected Taverner. A passage in a letter from Fleming at this period must, therefore, have interested him. “T am still considering the future of my collection. I hope I am only 1/2 way through as to size. About 40,000 is what I have in mind. I would like to extend my American collection much faster than I am but two years more will mean little additional material and after that I may be so handicapped financially that I will have to forgo new material. However, let us hope for the best, and I believe the chances of getting material are likely to be good after the war. As far as the museum [the Royal Ontario Museum] is concerned I look forward to seeing it get past a purely Canadian field and well established in general natural history. | would feel much easier about the destiny of my collection.”’23 Because the National Museum was aiming to form a strong collection of the birds of Canada only and not of North America or the world, Fleming’s collec- tion was likely to be given to the Royal Ontario Museum. Taverner could hardly expect only the Canadian material in the Fleming collection to come to the National Museum. Matters were different with the Brooks collection. In June 1915 Fleming had visited Brooks’s home in the Okanagan Valley on his way back from the AOU meeting in San Francisco, while staying with Munro. Brooks had left in a hurry at the outbreak of war and Munro was doing all he could to keep his bird collection from being infested by dermestid beetles, but had run out of napthol. Fleming asked for help.24 Taverner replied that he would send some naptholin [napthol] to Munro, and added: “Brooks has arranged that his specimens come to our museum when he dies so that gives me an excuse for supplying a presentation from museum stores.”’25 Taverner should have remembered such popular sayings as not counting your chickens before they are hatched when he wrote those words. If Brooks had been killed during the war then certainly his col- lection would have gone to Ottawa. But instead he had a distinguished career in the course of which he not only won a D.S.O. but also survived active ser- vice in France. He continued to live almost as long as Percy and had ample time in which to change his mind about which institutions would finally receive his collections of birds and paintings.° Also from around this period Taverner began developing a wider group of correspondents with similar interests to his own. One such was Henry THE CANADIAN FIELD-NATURALIST Vol. 110 Mousley of Quebec Province. Mousley had emigrat- ed from Britain to Canada in 1910 and was soon studying the natural history of the area around Hatley in the Eastern Townships of Quebec. Here he began his extended field studies on the nesting behaviour of birds, especially warblers. Taverner heard of these studies when Mousley wrote to the Macouns about the Catalogue of Canadian Birds and mentioned his own observations over the past years. By late 1914 Taverner had started collecting data for his projected work on the birds of Eastern Canada, and when Mousley’s letter was sent to him to answer he took the opportunity to request from Mousley any ornithological notes he might have. Taverner said, in part: “As soon as a reasonably complete reconnaissance has been made I intend publishing the whole results for the Eastern provinces. In the meantime we greatly feel the lack of other workers for the field is broad and we can- not cover it all in anything like a complete manner.”27 Mousley responded with a copy of his notes on 110 species of birds found in the locality of Hatley. Taverner acknowledged this with thanks, at the same time apologizing for his slowness in replying. “Do not think that this seeming neglect is a measure of our interest in your work. It is but the result of covering a large field with a limited staff. Such lists as you sent are of great value to us and will be [of] inestimable value when we come to work out our distribution again.”28 Taverner’s use of the word “we” does not mean that he had suddenly assumed the royal “we” but simply that in 1914 he hoped that Brooks, Saunders and Fleming might be co-authors of the book with him. Writing to Fleming about Mousley’s work Taverner reported: “Have been corresponding with an H. Mousley of Hatley, P.Q. He seems quite a find, and reports numbers of warblers breeding and evidently knows what he is talking about. He is giving us a [Northern] Parula nest and promises us more of like stuff’.?? So began a fruitful correspondence between Mousley and Taverner which continued for a quarter of a century. With all this activity and writing Taverner began to prove himself in his post as the only professional ornithologist in Canada. But he was never allowed to develop a swollen head, even if he had been that kind of person, because Fleming was ready to poke fun at him gently as when he wrote: “You ought to keep notes of your work as you go along a sort of journal would be the thing. I don’t mean your field journal but something like lawyers keep. Mr. So and So telephone 50¢. There are so many things you do in the course of your work that you forget.”30 However, Fleming also told Taverner that during the return journey from his trip to the AOU Conference at San Francisco he had visited Macoun in Victoria. “He talked about you quite a lot is very proud of your success.”3! Macoun himself wrote to Taverner: “I am very glad to hear how well you are 1996 getting along and pleased to know that my work was not in vain.”32 The professor need not have worried. Taverner had made the grade. Early in 1916 another fire affected Taverner’s life. This time it was not his home but the Parliament buildings. His eye-witness description of the results of the fire on the museum building are worth quot- ing. “A cyclone has struck us. The fire that burned out Parliament was almost as bad to us. Any way it spread desolation and disaster our way. Fortunately I have not been moved but the exhibition and office floors had to be vacated. We tore out the big cases and piled them up as best we could. Thank goodness though the study col- ~ lections have not been disturbed and I suffered less than perhaps any one else. The Invertebrate paleontologists got hit hard and for two days and nights all the girls in the building and half the Geological staff were wrapping up specimens and packing them in boxes from the old exhibition cases on the first floor. I guess they lost a lot of stuff through breakage and unskilled handling. It was awful.” The fire occurred during the night of 3-4 February 1916, destroying the Centre Block of Parliament. The Commons started moving into the museum immediately, followed by the Senate a few days later. Not all staff were moved out of the museum. | James Macoun, who had known the Director of the museum, McConnell, for over thirty years, persuad- ed him that the Biological Division would need more time to prepare for moving. But the presence of Parliament in the building was disturbing and in some ways members of the Division were better off when they moved into a place of their own in January 1918.33 Taverner’s main work during 1916 was involved with his bird book. Because he was working hard to complete a draft of the manuscript he decided not to make a field trip himself that year. As a result he had more time for going through the collections and determining the species. His correspondence with | Louis B. Bishop in 1916 contains interesting letters between the two men on various species and the _loans of various skins. These letters, written in a _telaxed style, show examples of Taverner’s ornitho- logical work at that time. Writing to Bishop in July he said: | “Dwight was here a short while ago. Our series of gulls | rather surprised him as well it might. I think we have all the puzzles in the country, and to see practically the | genus Larus linked together by apparent intermediates is / enough to stagger one. These birds, especially the white- ' winged ones need study. When Anderson comes back this fall from the Arctic and brings the remainder of specimens I guess it is up to us to get busy on them.”34 It seems that Taverner lightened the burden of some of his regular and lengthy correspondence by | repeating passages used in a letter to one friend in a _ letter written soon after to another friend. Thus mate- _ rial in the letter to Bishop quoted above had already | CRANMER-ByYNG: A LIFE WITH BIRDS 69 been written, in part, in a letter to Fleming as fol- lows: “Dwight was here yesterday. I think he was surprised at the extent of our collections. He went over the Horned Larks and several other species and the very forms I had failed to find he regards as untenable. His ideas on them cleared up a lot of my difficulties. I was glad to have him corroborate my identifications of Wright’s and Hammond’s Flycatchers that I was pretty shaky on. I sweated on them but could not work out the wing formu- la at all.” In the same letter he mentioned the gulls known to occur in Canada, but in somewhat different wording. “Dwight looked over the gulls and is just as puzzled as I am. An intergrading of the California Gull through Kumlien’s to the Iceland taking in the Glaucous by the way and connecting with the Herring Gull is enough to cause thought. It must have some reasonable explanation but the above is the superficial appearance.”> A study of the taxonomy of many species of birds found in Canada was still a task for the future, and Taverner was the one on whom the task mainly depended. With help from a few experienced ornithologists in Canada, and more in the United States, some of the work of distinguishing among the various species and subspecies of gulls, in their dif- ferent plumages, was accomplished. But in 1916 there was a great deal still to be discovered. It was like trying to solve a complicated puzzle. Writing to Fleming at this time he said that he had been going through the Red-winged Blackbirds which he called ‘‘a badly split up mess”, and continued, “We have nearly 60 specimens but the only subspecies that I can see is arctolegus of Oberholser that is not accepted by the A.O.U.”36 In November 1916 Taverner attended the AOU meeting in Philadelphia, afterwards visiting the National Museum in Washington and the American Museum in New York to examine some of their series. After going over the gulls that he brought with him from Ottawa with Dwight at the American Museum, he left them there for Dwight’s further examination. In a letter to Fleming he said that he had come to no definite conclusions but felt that he ought to publish something on the gull question in order to call attention to the question of intergrading, and whether thayeri was a true subspecies.’ Sometimes, in his letters, Taverner gave a close friend his opinion of fellow ornithologists. When Joseph Grinnell, of the University of California at Berkeley, had spent a day at the museum, Taverner told Fleming: “He appealed to me very strongly. I liked him immense- ly. Though he is a splitter we agree on so many points in general that we got along nicely. He named the Song Sparrows and the Fox Sparrows for me. He ran foul of Oberholser’s determinations on a number of cases [of Song Sparrow subspecies] discarded rufina altogether and lumped them under morphna . . . and threw out all our Sask. and Alta. birds into juddi. H.C. O. [Oberholser] had them all melodia but they certainly are 70 THE CANADIAN FIELD-NATURALIST different from eastern birds. His wife is interested in bats and went over what we have and named them so you see I got something out of the visit . . . Grinnell was much interested in my collection of pictures especially those of fleshy parts of birds. I am certainly getting some good stuff in that line.”38 The “fleshy parts” refer to the bare parts which are quick to fade once the bird is dead, parts such as eyes, beak, feet, wattles, inflatable sacs and other soft parts. Although Taverner stayed at the museum during 1916 he arranged for Patch to spend December 1915 and January 1916 collecting in Barkley Sound, Vancouver Island. The principal object of the trip was to collect sea lions, but at the same time he made a good collection of sea birds for the museum, including Barrow’s and American goldeneyes, Surf and White-winged scoters (good series in all plumages) and Surf-birds. These were quite a sur- prise as this species had not been reported previously wintering as far north. He told Fleming that a con- signment of birds had just arrived from Patch and that he had Johnson making sketches of the feet and bills.39 In the summer of 1916 Young joined James Macoun and Spreadborough in eastern British Columbia to collect birds in the Howe Sound and Lillooet area. These two collecting expeditions helped to enlarge the holdings of birds from British Columbia. The major collection to reach Ottawa during the year was that brought by the Canadian Arctic expe- dition, though it was some time before it was fully prepared and catalogued. The good news for Taverner was that Anderson was to join the museum staff. In writing to Fleming he mentioned the mam- mal collection and said, “The Arctic Expedition has returned and I am expecting Dr. Anderson daily. You know he is to be our mammologist and it will be a relief to hand this stuff over to him.”4° Fleming was excited by the opportunity this collection pre- sented for study purposes. Writing to Taverner about a possible monograph on the birds of Labrador that W.E. Clyde Todd might some day write he remarked, a propos of Anderson’s expected report on the work of the Southern party: “It ought to be a great opportunity to get out a monograph on Arctic birds. Who will write it? Anderson I suppose.”’4! In fact Taverner was chosen to write the section on birds in Volume Two of the official reports of the expedition while Anderson was to contribute the sec- tion on mammals. Eventually the volume on geogra- phy and geology was published in 1924, and the anthropological reports in 1925. But owing to a long drawn out dispute between members of the Southern party and Stefansson, the leader of the Canadian Arctic Expedition, the volume on natural history was never written.42 Perhaps this was just as well because owing to the vastness of the area, and the lack of knowledge of the distribution of birds of arctic Vol. 110 Canada, the writing of such a monograph would have been premature. The exploration of the breed- ing grounds and the ranges of many Canadian species remained to be carried out in the 1920s and 1930s [see Chapter 14]. If Taverner had been required to cooperate with Anderson in writing a section on the birds collected by the expedition, quarrelling between the two men might well have begun at the very outset of their years together in the Zoology Section at the museum. There was plenty of time for that later. Because monographs on Canadian birds, either by categories (sea birds, waders, game birds) or by regions, were comparatively few it was all the more important to have a bibliography of what had been published including articles, local annotated lists and reports. During 1915 Winifred Bentley, general assistant in the Biological Division since 1913, start- ed work on a bibliography of the literature of the ornithology of eastern Canada and Gulf of St. Lawrence, as the beginning of a bibliography of Canadian ornithology.*? One way of measuring the value of a museum to the community it serves is by how much the public uses it. Another way is by the educational role it plays in the community. The National Museum took its first step in an educational role during the winter of 1914-1915 when it instituted a number of popular lectures given by members of the museum staff and | the Geological Survey. Illustrated lectures on topics relating to the subjects within the scope of the muse- um’s work such as glaciers, fossils, winter birds, Indians of Canada, were offered to audiences from the Normal School, the Collegiate Institute, and the Public Schools of Ottawa in an improvised lecture room at the museum. Taverner’s friend, the archae- ologist Harlan Smith, was largely responsible for organizing the lectures and gave several himself. Patch and Bentley each gave some but Taverner, because of his speech impediment, was unable to take part. Instead he was responsible for the prepara- tion of the slides required for zoology topics. In the winter of 1915-1916 the scope of the lectures was enlarged and they were continued. A useful addition to the museum’s educational function was the loan- ing of study collections to the Ottawa public schools. Specimen boxes were issued to selected schools con- taining sets of birds such as common woodpeckers; winter birds (consisting of 13 species that might be seen on a winter’s day about the city together with specimens of common winter food); examples of protective coloration of birds and mammals. A total of eleven different boxes were circulated among these schools. The zoological lantern-slide collection was also in demand by scientific and educational bodies in various parts of the Dominion on topics such as “sea birds of Bonaventure Island”, while the moving picture on the same subject and the film 1996 showing Jack Miner’s success in banding wild geese were shown to scientific and popular gatherings and were received with enthusiasm.*# Life at 45 Leonard Avenue continued pleasantly when the family moved back into their home after the fire damage was repaired. Their main preoccupa- tion at this time was the garden which had been in the making since April 1913 when Percy rented a team of horses and a scraper to level the ground. In the spring of 1915 he told Swales that he was work- ing hard on the garden before leaving for the field and added: “though what I want a garden for when I am away all summer is beyond me. Neither do I know what Mother would do without one but the work is too hard for her . . . Last year Mother entered the garden in the city competition and got second prize — with $25.”45 In the autumn Taverner began making a tennis court on the other lot which they owned but had not yet developed. Also at this time he borrowed a book from Fleming on rock gardens with the comment that this was his ideal form of gar- dening if only he had time to make one.*¢ In the spring of 1916 he took a few days’ holiday and “worked like a trooper” making a large lily tank in the middle of the lawn. He planted several fragrant water lilies (Nymphaea) and an Egyptian lotus. He did most of the work himself, digging out the soil and wheeling it by barrow onto the other lot. At the same time he continued the existing trellis work the whole length of the yard. The garden, and birds that visited it, gave the Taverner family a great deal of enjoyment. One of the first things Percy did when the garden was being laid out in the spring of 1913 was to put up a Purple Martin house. In 1915 a pair nested in it, and in April 1916 three pairs returned to the house. The garden survived the winter well because there was a heavy fall of snow in the autumn of 1915 which lasted until the spring break-up. Percy was able to boast, “In consequence I had California poppies overwinter and are coming on again from the old last years plants. How is that for Ottawa. The Xmas rose was a wonder and when the snow left was covered with great blossoms.’’47 Meanwhile the Purple Martins were a constant joy. He told Fleming, “They chatter all day every day and the house is sur- rounded by a constant flock of them.”48 Also they were interesting to watch. In June Taverner observed them carrying poplar leaves into their house. He described their actions to Fleming who told him to see if the martins were carrying leaves to their house on hot days. This might be the answer — to cool down the inside of the house.” Next spring he put up another house. He told Fleming: “Have a new Martin house to put up. It is quite an elabo- rate affair and is an attempt to put some real design into such a structure. Artistically I think it is a success. We will see what the Martins think about it. Design while CRANMER-ByYnNG: A LIFE WITH BIRDS Wl Purple Martins at martin house designed by Taverner. Photographed by Taverner in the garden of his home in Ottawa in the summer of 1916. (Reproduced courtesy of the Canadian Museum of Nature, number 41306.) not especially attractive to them should not be any deter- rent as long as other practical conditions are complied with.”50 In his daily life Percy was constantly using his hands to good advantage. He gave the impression of being self-reliant and a first rate handyman. There are various examples of his readiness to tackle a repair job, or to take on major construction work even if he had no previous experience of that particular job. Gardening and handiwork around the home seem to have been his physical exercise together with field trips to collect and study birds in the summer. In the winter he mentioned skating. Presumably the tennis court was mainly used by Ida and her friends. Percy was not a games-playing man by choice. Although Ida had a position in the museum library and by this time had made a circle of friends, never- theless her life was made miserable from time to time by acute outbreaks of eczema. Percy explained the problem in a letter to Fleming. “My sister is quite sick so just at present we cannot entertain . . . Ida is not dangerously ill but has such an acute attack of eczema and has to use such villainous looking ointments that she naturally withdraws from the public gaze.’”>! Fleming was sorry to hear this and jokingly sug- gested she might try “dew on the grass in the early morning”, though he added that this was an old English folklore idea. However, to English folklore Fleming added a medical one of his own: “But as a matter of fact constipation is responsible for most 72 things of the sort.’>? In 1917 Ida went to New York for advice and was told that her eczema might be caused by the state of her teeth. But dental treatment failed to make any permanent cure.°? In contrast to Ida’s health, Percy was fit and well during these years and full of energy both physical and mental. He was clearly enjoying his career, and his home life. The only dark cloud was caused by the war. All hope that it would be over quickly had long evapo- rated. If Percy ever had any ambitions to serve his country in the armed forces, as his friend Allan Brooks was doing, his heart murmur would have precluded him on medical grounds. This did not pre- vent him from expressing his feelings strongly to close friends. During the blackest period of the war he wrote to Louis Bishop about his disgust with American politics as they appeared to an “ex-patri- ot’, referring to his residence in the United States. “It is all very well to stick to one’s country but only when it deserves it. My accident of Canadian birth that fretted me for many years now is a more comforting thought. We will see in November.”54 But the United States delayed declaring war until 6 April 1917. Taverner fired a salvo of his own at the U.S. government in a letter to Bishop. The nation, he claimed, had grown too soft with mollycoddleism, prohibitionism, feminism, and Audubonism culmi- nating finally in pacifism and “TI did not raise my boy to be a soldier” mentality. “All are emanations of the same fundamental spirit of wrapping the unfit in cotton-wool . . . It is because I am more or less still American that [I] hate to see the nation stultifying itself.’”’55 The influence of Darwin’s Origin of Species, which he had read with strong feelings of empathy as a young man, now came to haunt him over the human condition in wartime. Part of his letter to Bishop was about war being more or less a biologi- cal necessity for the human species. This was a dan- gerous argument which he was to express again dur- ing the Second World War. However, writing to Fleming in February 1917 he was rather more per- ceptive. He commented that it looked as if the United States would enter the war “with us”. Taverner was glad because he considered that it would be a distinct loss to the world if the war fin- ished without the U. S. being directly touched by it. “The demonstration that even the extreme of diplomacy will not avoid trouble will make them more willing to assist in preventing the rise of aggressors in the future.””>° In contrast to Taverner’s attitude towards the war his new colleague in the department of zoology at the museum, Rudolph Anderson, was more belli- cose, at least on paper. Taverner had written to Fleming to tell him that Anderson would be in Toronto at the end of March 1917 when he was due to address the Royal Canadian Institute on the Canadian Arctic Expedition, and that Mrs. Anderson THE CANADIAN FIELD-NATURALIST Vol. 110 would be with him as she was also to give a lecture. They were due to go to Hamilton on the following Monday. Diplomatically Taverner suggested “It would be well, if not in opposition to your other plans, if you could have them out to your house some- time Sunday”.>7 This was the first meeting between Fleming and Anderson, and it began a correspondence between them during which Anderson expressed his feelings about the war. In a letter to Fleming in June 1917 he mentioned that Taverner was in Red Deer, Alberta, preparing to raft down the river to join the Sternbergs who were digging for dinosaurs in the badlands near Drumheller. Anderson commented “T find it pretty hard work to stick to museum work with all the war excitement this summer, both in Canada and the U.S. My personal preferences are all for flying the coop and getting into the big game”.>8 Fleming in his reply suggested, either through ignorance or with his tongue in his cheek, that since Anderson had a degree in medicine his services would be very valuable in the army. Anderson replied that he only had a Ph.D. in the, at present, unpractical sciences of zoology and animal morphology. “If I were a medical man, I could see my duty more clearly, as physicians and surgeons are badly needed everywhere. “T could get a commission as captain, or possibly major, of infantry without much trouble by going back to the U.S., into the new National Army, but it doesn’t look well to be a “hyphenate,” so I suppose I'd better repress my military desires until Canada invites me to take up the job of little buck private. The infantry is the only branch that appeals to me, as there is something personal about the rifle and bayonet that is lacking when one deals with events by machinery several miles away.’5? The physical difference between the two men was very pronounced. Taverner was tall, lean and bearded while Anderson was tall, thick set and clean shaven. Anderson had recently returned from three consecu- tive years of field work in the Arctic while Taverner had completed five years of museum work in Ottawa. Anderson’s background was university, army, foot- ball, and athletics; Taverner’s background was non- academic, non-military, and cultural rather than sporting. Anderson had already published a book on birds, Taverner was in the process of writing one. In age Taverner was one year Anderson’s senior. These two men were to-be close associates at the National Museum for the next quarter of a century.® Now that Anderson had taken charge of the mam- mals Taverner was able to concentrate entirely on the bird collections. Moreover, since Parliament con- tinued to occupy much of the museum building, staff - did not have to spend time on making their collec- tions open to visitors. As a result Taverner was able to devote much of his time to the study and classifi- cation of the enlarged collections of bird skins. The problems caused by the determination of these speci- 1996 mens occupied an increasing place in his correspon- dence with ornithological friends and acquaintances. For instance Taverner was working through the specimens of Savannah Sparrows and found a form taken on Vancouver Island which did not appear to have been described before. He told Fleming: “Either Ridgway is color blind or else there is a whop- ping new subspecies in western Canada. I see Swarth calls the Vancouver Island bird Savannah but this it cer- tainly is not.” He then described it.*! Between early February and early May 1917 ten letters between Taverner and Fleming discussed this hypothetical new subspecies without reaching any definite conclusion. Taverner, with his sardonic sense of humour, compared what they were doing to a game. “It is a great game; look wise, name everything, never retract and bluff it out and one can easily make a great name as an ornithologist. The trouble is that the splitting is too fine and when we get through we have shavings and chips and not building lumber.” When Fleming copied out Lucien Bonaparte’s description of Passerculus anthinus Taverner with- drew his tentative description of a new subspecies of Savannah Sparrow because he came to the conclu- sion that his bird was only an anthinus.®> But such an exercise in determining correct names for specimens in the museum collection was of value to Taverner. The more determinations he made now the more cer- tain he was likely to be over which forms might be considered “good”, and therefore described in litera- ture in the future. Other hypothetic forms that Taverner and Fleming were discussing at this period were subspecies of Song Sparrow, Fox Sparrow, and Merlin. But the most important development in Taverner’s career was that at last he was free to turn his atten- tion to the study of the bird life of the western provinces of Canada in the field. There is evidence that he was very conscious of his lack of first hand experience of the birds of the prairies and British Columbia. In a letter to Brooks in 1916 he explained that after he had managed to do some work in the _ prairie provinces he would then turn to the localities in British Columbia which Brooks had spoken of and added: “Perhaps I may have the pleasure of your company there”. In May 1917 he was able to leave Ottawa for Manitoba where he collected in the vicin- ity of Shoal Lake. Later he moved to the Red Deer river in Alberta, and then to Jasper National Park. While in the field large collections were made so that the birds of the west might be more fully repre- — sented in the National Museum’s collections. Meanwhile Taverner was running into difficulties over the content of his book on the birds of eastern Canada. He had intended to write a book that would be of practical use for the identification of Canadian birds but would also stimulate the reader’s interest through the appeal of birds to his aesthetic sense. But CRANMER-ByYNG: A LIFE WITH BIRDS Ws the “powers that be” thought differently, as Taverner told Fleming. “The bird book is causing me a lot of trouble. McInnes is the Acting Director of the Survey now and it is in his hands and he has not done a thing to it. He seems to think that nothing but the baldest kind of statements are allowable and would turn it into a census tabulation. Your citicisms are very mild indeed in comparison. Unless I can influence him it will be only a Reed’s Bird Guide, or less. It is not the expense he objects to and I fail to catch his standpoint at all except that he is a scien- tific purist, and thinks that nothing that is popular [is] within the dignity of the Geological Survey. I am going up to have it out with him tomorrow. Such is the way, I really miss Brock now. He at least was not fossilized.”® While Percy was enjoying his first acquaintance with the landscape and the birds of the prairie provinces his mother and sister were enjoying a visit to a lakeside cottage. In July 1917 they were invited by a friend to stay at Blue Sea Lake in Quebec province, and they fell in love with the place. The following month they returned with friends to stay at a cottage on Big Island which was loaned to them for a few weeks while the owners were away. Before they returned to Ottawa Mrs. Taverner had arranged to buy lot number 4 on the island. There were good reasons for their hasty “love at first sight” reaction. Nostalgia for the years of cottage and lakeside living at Beaumaris lay deep in their memories. However pleasant life was becoming in Ottawa, when the warmth of summer returned and their friends left for “the cottage”, they longed for a cottage of their own. The journey from Ottawa to the village of Messines, at Blue Sea Lake, took three hours or more by a sin- gle-track railway running beside the dark waters of the Gatineau River into rural Quebec. The railway ran through cuttings of pale grey granite blotched with pinkish-brown outcrops, with the river winding below, until Blue Sea Lake was reached and the train ran its scenic course beside the lake. Big Island rose steeply out of the water and was completely covered with trees. The view from lot number 4 faced south down the lake with a wooded island to left and right in the middle distance framing the view and direct- ing one’s eyes through a channel between the islands to a wooded shoreline in the further distance. On a sunny day, with only a slight breeze, the blue surface of the lake reflected the light and mirrored the large white clouds that sailed slowly above it. One had the feeling of being raised above the surface of the lake, near to the wide expanse of sky, yet contained by the horizons of low forest-covered hills. It was a view to enlarge the human spirit. When Percy returned from his expedition to the western provinces he was pre- sented with the challenge of building a cottage on the island lot which the Taverner family now owned. For some undisclosed reason Percy visited Lake Muskoka at the end of September. It was only eleven years since the Taverners had sold their cottage at Beaumaris and they still had friends there.°® Finally 74 THE CANADIAN FIELD-NATURALIST at Thanksgiving 1917 Percy first saw the location selected for their family cottage. In spite of the grim events of the war, and the col- lapse of Imperial Russia on the eastern front, 1917 had been a good year for Percy Taverner. At the beginning of 1918 the Biological Division finally had to move out of the museum to make room for forty new members of Parliament. Since the specimens were already packed this was not too troublesome, and the whole Division was able to remain together on the top floor of a large building nearby which was fireproof, an improvement on the museum building. It also had good light since the west side was made of glass. The only drawback was the lack of an elevator; the staff had to climb four flights of stairs to get to their offices. The prepara- tion shops remained where they were. Because the library remained in the museum while other Divisions were scattered in various parts of Ottawa, it was time consuming for Taverner when he needed to consult other museum members, or use the library.°7 Two welcome events in 1918 should, in Taverner’s mind, have more than compensated for being uprooted. Taverner finished the manuscript of his book and sent it to the editor and, although it was not quite what he wanted, he told Fleming that he was fairly satisfied with it. Also at this time he became a Fellow of the AOU which carried much status with it. He also felt that Gordon Hewitt was now consulting him more on matters of bird protec- tion while setting up federal regulations to enforce the International Migratory Bird Treaty. But of greater significance to Taverner, personally, at this time was the question of who was to be head of the Biological Division. James Macoun had been appointed Acting Head in 1917 which pleased Taverner well, as he confided to Fleming. “I am glad this appointment has been made. He is the right man in the place, having good executive [word illegible] and a wider outlook than any other man on the staff. I get along with him very well indeed and he is a man of influence. He has the promise of a real staff under him at an early date. The return of the Arctic Expedition has brought things to a head.’®8 Anderson now began to make a number of requests and suggestions to Jim Macoun regarding the work and personnel of the Biological Division which Macoun in turn passed on to the Deputy Minister R. G. McConnell. In one covering letter Macoun wrote: “While the duties of Acting Head of the Biological Division are not at present onerous, they are troublesome, and matters come up which cannot be dealt with satisfactorily except by the one [who] is the real responsible head of the division.’©? As though in response McConnell made Macoun Head of Division immediately. But Jim found that his increased authority did not necessarily make it any easier to mediate in internal differences. His position Vol. 110 as Curator of the National Herbarium give him more satisfaction, and his field work more pleasure than being Head of the Biological Division.”° The sum- mers of 1917, 1918, and 1919 he spent with Spreadborough as assistant, making a botanical sur- vey of the Jasper Park region. At the end of each field season Jim returned to Vancouver for a short visit with his father before returning to Ottawa.7! Meanwhile the museum staff faced a reclassifica- tion of the Civil Service involving status and salary. Writing to Fleming in the fall of 1918 Taverner explained, “We all have cards to fill and are searching the dictio- nary for words in which to express the importance of our work. I have found a number of them but as yet not found how to work them in.”72 Eventually, when the new scheme was made known in July 1919, it came as a shock. It seems that some people, when filling in the form, laid most stress on the administrative work they were perform- ing as though this carried more clout than anything else. But in doing so they failed to double guess the attitude of the revising board correctly. Taverner’s explanation was that members of the board, when set- ting the new salaries, regarded the work of adminis- tration as considerably lower in value than research work. In fact no upper limit was placed on the salary of those whose replies showed them to be mainly “research men”, while being engaged mainly in administration was rated as detrimental rather than advantageous. As a result Taverner’s salary advanced immediately to $3300.00 and was not pegged to any upper limit while Jim Macoun and Anderson both found their salaries lowered and pegged to a limit. But, as Taverner told Fleming, of course it was going to be revised.’3 Writing to Macoun on the subject Taverner gave his own opinion. “Of course the distinction made between Dr. A and myself is absurd. The ignoring of you as botanist is also a bull . . . Under it you, as Head of Division, would not have as good prospects as you should as Botanist . . . Of course I am satisfied with my status [and] am not anx- ious to have it reduced but would like to see Anderson brought up at least equal as he should be.””4 During the years 1918 and 1919 Taverner was busy with a variety of aspects of museum work, both advising on policy and building up the collections. He was required to write a memorandum for the minister, McConnell, on the need for the museum to have extralimital bird specimens in its collection for research purposes.’> During 1918 a total of 1300 new specimens were added to the collection, and work continued on preparing groups for exhibition. Taverner’s own collecting was limited to Ontario during 1918 and 1919. Work on the study and classi- fication of the collection went on unabated. Early in 1918 he worked through the juncos and kinglets; later the hummingbirds when he arranged an exchange of some with Fleming.’© Writing to Brooks | 1996 in the fall, Taverner could be pleased with the progress being made. Young had continued the pre- vious year’s work at Shoal Lake, Manitoba and had collected over 600 birds, while Spreadborough brought back some good specimens from Jasper Park, Alberta. In an optimistic vein he continued: “The museum has really gone ahead most satisfactorily though at times the indifference of our chief is madden- ing. Fortunately Macoun is now in charge of the Biological Division and if the promised reorganization goes through we have good hopes of the future.”””” His opinion of the Director of the Museum, McInnes, however, was expressed bluntly in a letter to Fleming when he wrote “Wish we had a director that had enough gumption to try and make the department useful, — at any rate a bluff at it. We surely have a King Log now. The Survey is in a very bad way. All the energy and ambition has been taken out of the staff.”78 By 1918 Taverner was in touch with a number of enthusiastic amateur ornithologists several of whom were younger men seeking a career related to the study of birds, and an outdoor life. He was fortunate to know people such as Fleming and Saunders, Harkin and Hewitt who told him of people he might find helpful as correspondents. For instance Gordon Hewitt showed him a recently published annotated list of the birds of the Wolfville region of Nova Scotia. Local faunal lists were valuable to Taverner in his task of mapping the distribution of Canadian birds.7? He wrote to the author, Robie W. Tufts, ask- ing for a copy and explaining his interest. “T am endeavoring to keep in touch with and to correlate all ornithological endeavor in the Dominion and your assistance in this direction will be greatly appreciated.”’89 From his office in the National Museum Taverner sat at the centre of an expanding network of corre- spondents who kept him informed of what was hap- pening, ornithologically, in their own areas. He could expect to receive both information and speci- mens to add to the collections. There was so much that the museum required such as specimens of nests and eggs, and nesting records of various species from a number of provinces. As an example Taverner mentioned White-winged Scoters off the coast of Labrador. He had seen thousands of them in the summer of 1915 in all plumages, both male and female, yet he knew of no nesting records.®! Another new correspondent was Fred Bradshaw, Chief Game Guardian for Saskatchewan. Taverner was in touch with him in 1919 about the possibility of coming to his province on a field trip the follow- ing year and wrote: “Would like to get enough data to prepare a local Sask. list like those of the Red Deer and Shoal Lake. Mr. Fleming is publishing one for the northern part of the Province. These local lists to my mind are very impor- tant for we are really unable to put out a complete provincial list until we have enough of the more local ones to form a basis. Saskatchewan is now the western CRANMER-BYNG: A LIFE WITH BIRDS i Province most poorly provided with such results but we hope to remedy this in the near future.”®2 Bradshaw replied that a start had been made on a bird list and enclosed a copy of the Game Guardian’s annual report for Saskatchewan.®3 In his reply Taverner suggested that in the next report it would be helpful if Bradshaw could annotate the list to indicate which birds were common and widely dis- tributed and which were stragglers, or confined to limited areas. Also he encouraged Bradshaw to obtain records on the fluctuations of grouse over a long series of years, and recommended that game wardens under him be required to make systematic reports on grouse including abundance or otherwise of food, and of the predators, bird and mammal, that fed on grouse.*4 A new contact in Quebec province was W. J. Brown of Montreal. Brown had written to Taverner early in 1919 to congratulate him.on his report on the birds of Red Deer River.5 In thanking him Taverner revealed his ideas on the importance of interesting the general public in birds. To do this the museum would need to conduct a campaign of education in order to rouse the public to prod the government into action. “As it is I fear the government reflects the public but too well. I was in hopes that we were going to establish a truly National Museum when I came here but after building a fine expensive building it laid down on the job and refused to equip it with either facilities or per- sonnel. I think however there are signs of awakening and if the few of us there make enough noise we can give the effect of a crowd. So all get behind and push.”’8° Brown responded warmly to Taverner’s letter and wrote: “After grasping the fact that there was at least one other who thoroughly recognized our shortcomings, I felt relieved, and frankly it makes cheerful reading to know that the right man is at the helm. For years I could never understand why Canadians were so inactive and disinter- ested while our neighbours to the south were progressing rapidly in this most fascinating study.”°7 With a few correspondents showing the same appreciation of what he was trying to achieve Taverner must have felt encouraged to retain his belief in the value of his ornithological work in spite of the apparent inertia of his superiors. Brown sent him some notes on various species seen in Montreal, and a promise to help in collecting specimens. This prompted Taverner to write to Brown about another of his main concerns: “Good working collections are the tools that are most lacking in Canada today. American results when pub- lished are as available to Canadian workers as to American ones and from a broad scientific standard as good. Collections however in the States are not available to us and hence until we get good series of necessary species we are still dependent upon others for any sound work. I feel it my first duty to build up these collections as working tools. I have tried to conduct an educational 76 THE CANADIAN FIELD-NATURALIST campaign incidentally but it has been up hill work over- coming inertia. | am very hopeful now that we have Mr. Lloyd taking up an agressive policy in this direction. I think he is well prepared for it and has the full backing of his department. One great advantage he has is that he belongs to a new department that is not tied up with a couple of generations or more of tradition and precedent. We are trying to get some interest stirred up in mak- ing the Ottawa Naturalist a medium of intercommunica- tion between all the Natural History Societies in the Dominion. A letter with a scheme for closer cooperation will reach the Sec. of your Bird Protection Club shortly and I hope that it will meet with general approval.”88 When Brown asked what birds were needed Taverner replied that he would be glad to have addi- tions to any series. He told Brown that the museum contained almost nothing from the Montreal region and would be glad to receive anything, bird or mam- mal, that could be procured. Whilst rarity was wel- come in a specimen, a complete collection from his important region was the ideal; nothing was too common at this stage for usefulness. Hawks and owls, being the hardest to obtain, were very desir- able. Great Horned Owls were especially welcome because Taverner was hoping, some day, to work out the geographical races into which this species had been split. But to do this he needed large series, and this would take time to obtain. “Birds actually known to be breeding are the crux of the question and whilst migrants are desirable breeding birds are the final necessity.”89 Taverner also mentioned that the museum was anxious to obtain nests and eggs as well, especially nests in situ for use in life history groups, and invited Brown to come to the museum to see the work they were doing in that direction. Taverner showed a natural ability to say the right thing when writing to people predisposed to help the ornithology section at the museum. His correspon- dence with Brown, Mousley, and others shows him taking trouble to encourage, help and advise them with the result that they in turn became enthusiastic to help the museum.?! This did not apply to his let- ters only but also to his meetings with those of like- minded interests — in spite of his stammer. The Commission of Conservation also claimed a share of Taverner’s time during the war years. One major advance which was to be of great help to Taverner during his future career was the appoint- ment of Hoyes Lloyd as ornithologist with the Dominion Parks Branch when this position was established in 1918. Lloyd was one of several appli- cants when the Civil Service Commission advertised for someone to assist the Parks Branch in the admin- istration of the Migratory Bird Act. Fleming knew Lloyd as a keen student of birds living in Toronto, and favoured his appointment to the post. He also realized the importance to Taverner of appointing the right man for the job. He told Taverner: “I feel he is the right man to cooperate with you, and would be of immense value to Harkin. Lloyd will shine Vol. 110 in investigating economic problems such as will arise in ornithology.”92 Apparently Klugh and Munro both wanted the job but Fleming was not in favour of their appointment because he considered them too combative and lack- ing the tact needed for such a difficult job. Instead he again expounded on Lloyd’s good qualities and the fact that he was a “thorough student of birds”’.%3 Taverner, therefore, was favorably disposed towards Hoyes Lloyd when he came to Ottawa to be inter- viewed by Harkin, Hewitt, and Anderson. Lloyd called on Taverner who introduced him to Anderson who in turn introduced him to the other two mem- bers of the interviewing committee. When Lloyd was the successful applicant Fleming wrote to Taverner: “T saw Lloyd last night, he appears much pleased with the way everyone treated him and you have added anoth- er admirer to your long list.”94 This is an allusion to Taverner’s ability to warm people towards him by his friendly manner. In reply to a further letter from Fleming praising Lloyd’s ana- lytical mind Taverner said he, also, thought that Loyd was the right man for the job, and that he would try to make him feel at home in Ottawa. The Taverner family invited Lloyd to their house on Christmas Day.®> So began a close friendship between Taverner and Lloyd which was to last, to the benefit of both men, until Taverner’s death. Two pleasant things occurred early in 1919 that affected Taverner. He received a letter informing him that his name would be proposed as a Colonial Member of the British Ornithologists’ Union. Thinking that Fleming had sent his name forward he wrote asking him how much it cost to become a mem- ber before committing himself and said: “I appreciate the honor and am glad to get it. Probably it is your suggestion.”°° Fleming replied that he had nothing to do with the proposal but was very glad to hear of it; that it was certainly an honor, that membership in that category was limited to ten; that it involved no sub- scription and no obligations; and the only privilege was to be able to subscribe to /bis on equal terms with members (£25 per annum).%” As a result Taverner was now a Fellow of the American Ornithologists’ Union, a Colonial Member of the British Ornithologists’ Union, and had a book on the birds of eastern Canada in the press. He had reason to be pleased with the achievements he had to show for all his hard work since his arrival at the museum in 1911. Taverner first met Fred Bradshaw, Chief Game Guardian, Province of Saskatchewan while attending a national conference on wildlife protection held at Ottawa in February 1919. When Bradshaw told Taverner that a new museum was being planned for Regina Taverner suggested to him that H. H. Mitchell, provincial naturalist and taxidermist at the existing museum, should come to the National Museum to work with Clyde Patch and observe his methods. This was part of Taverner’s perception of 1996 his role as ornithologist at the National Museum — to help provincial museums to train their staff in up to date methods. Apart from this it would be pleasant to renew acquaintance with Mitchell whom he had known from long ago when Mitchell was working at Spanner’s taxidermist shop in Toronto. It would also give Taverner another useful contact to include in the growing list of those he was in touch with across the Dominion.%8 Brooks was now back into collecting specimens for his own museum, and when he had surplus ones not yet in the National Museum collection he pre- sented one of each to it. In this way Brooks added four “Canadian” species in 1919, namely: Canyon Wren, Sage Thrasher, Brewer’s Sparrow and White- throated Swift collected by Charles de B. Green and George Gartell.°® Taverner’s own collecting of spec- imens from the western prairies would not begin on a large scale until the 1920s. In spite of not being a married man, Percy’s fami- ly life continued contentedly enough. When Ida was seeking a cure for her eczema in New York for five months in 1918, accompanied by her mother, he had to fend for himself. Being an extremely practical man and able to cook in a rough fashion he survived perfectly well but, as he confessed to Fleming, he had learned that “batching is not a comfortable life”. The Taverners had a pleasant home, which Percy had designed, in a very attractive part of Ottawa with a garden which Percy worked hard to create and maintain. As a family they entertained regularly so that Percy was able to invite his friends and acquain- tances to a meal, or offer them accommodation when they were in Ottawa. But because Ida was working in the museum library each day, and Percy was fully employed in his professional work, the running of the home fell on Mrs. Taverner who was now in her sixty-f-urth year. Perhaps his extended experience as a “bachelor” alerted Percy to the burden that his mother was carrying. Percy hoped to find someone who would come to live with them while sharing the running of the home with his mother.!0° The three Taverners were a loving and close knit family, and Percy was lucky to have such a good home life to support him in fulfilling the demands which he made on himself in his efforts to stimulate the study and enjoyment of ornithology across Canada. Looking back at the end of 1919 on the eight and a half years that Percy had served at the museum we have gained some perception of his relations with his friends and colleagues, and we know something of his affection towards his mother and sister. We know that he had an appreciation of music, and painting, and gardening, and that he had an artistic eye. We have seen the principles on which his character had developed, and how his outlook on major social problems had matured during recent years. Two examples may help to show how he felt on deep human questions. One was the issue of prohibition. CRANMER-BYNG: A LIFE WITH BIRDS dd, Both Fleming and Taverner were convivial, and enjoyed having a drink together. When he received a comic account from Fleming of spending a few days at Point Pelee with Saunders, Wallace, and Jack Miner, all strong teetotallers who tried to persuade him not to have a drink, Percy gave a serious reply which showed his feelings on the wider implications of the subject. Fleming had argued against prohibi- tion on the grounds that it was an unwarranted infringement of people’s liberties and any confisca- tion of property without compensation was scan- dalous. Although Percy liked his glass of beer, nev- ertheless he felt that the country was much better under prohibition, and was willing, he wrote, to forgo his personal inclinations for it. He never found any use for liquor as a medicine except when given under strictly medicinal conditions. Then he shifted the argument to a very topical issue. “If Winnipeg had been wet during the late strike there I think conditions would have been immensely more seri- ous. If all the soldiers could have boozed away their gra- tuities things would be bad indeed. I know they can get it but only at a high price and with difficulty . . . and with malice aforethought. On the whole I am for prohi- bition though I think that mild beer could still be allowed and that investors in the liquor business should have been reimbursed or at least been given sufficient time to readjust their works.”!0! Taverner had something more to say on the sub- ject of strikes when writing to Fleming about the reclassification of the Civil Service in 1919. “Speaking of the Civil Service though I wonder if the country realized that it has not had a raise since 1908? What other large employer can run today on a 1908 basis? It seems the only way to get just consideration is through organization and strike? . . . I deprecate strikes but they seem the only cure until employers will meet employees in an honest effort to concede what is just . . . Surely it is putting a premium on striking when only strikers receive consideration. Last year the lower divi- sions of the service received a bonus, about one fourth of what the price index indicated as a fair increase. Higher divisions received nothing — except the highest, deputy Ministers who got $1000 increase in salary, — not bonus. When the lowest and the highest get raises does it look logical to neglect the intermediate. This year we all get [a] bonus, highest in the lowest divisions and scaled down above. My bonus will be about nine dollars a month. Glad to get this but it is a mere bagatelle in com- paring prewar with present conditions. Of course I understand that we all have to stand our share of the bur- den of the war but why should more be expected of Civil Servants than of railroad and munition employes? The answer is they will stand for it, the others won’t.”!02 Taverner had quite strong opinions on social and political questions, and was capable of expressing them forcefully in writing when roused. By now the Andersons had settled into a house near the Rideau Canal, and the two families began to get to know one another. Taverner, writing to Jim Macoun in August 1919, said that he had taken 78 THE CANADIAN FIELD-NATURALIST Anderson on a visit to Blue Sea Lake to look at alter- native sites for building a cottage on his lot. While there they went to the local sawmill to buy timber, and then explored the lake in a borrowed power boat. Taverner reported to Macoun: “Anderson and I just returned from a weekend at Blue Sea [Lake]. Had a fine little visit there and I think Anderson is quite enthused over the place. Would not be surprised if he looked for a place there. He certainly opened up in camp and makes a good camp fellow.” !0 The next weekend the Taverner family came up by train and chose a site for the cottage facing south down the lake. Percy had taken a three week vaca- tion and the timber required was delivered to the nearest shore and towed across to the site. By the end of their stay, working with the help of one local man, he nearly managed to finish his summer cot- tage. The family agreed on the name “Hyla” — tree frog — for their new cottage. Writing to Fleming in mid-September he could say, with a touch of satis- faction, that the cottage was now in fair shape with the windows and doors hung, and the smokestack in place, and that he would finish it on the next week- end “so mother can occupy it next summer when I am away.” The plan, which he drew up himself, was, he said, “rather unique” since it had folding doors with glass the whole length which could be thrown open to give access to the porch. In good weather the porch and living room could be made practically one room.!% Near the end of September the Taverners’ first guests, the Andersons, arrived. The visit is recorded in the Hyla log book. “On the last day of our stay, all buckled in and put the finishing touches to the cottage. Mrs. Anderson stained the doors, and said she would think all winter about the two doors for which there was not enough stain. The partitions were all up, shelves etc. in the kitchen, the windlass for hauling up the water, stand for water barrel made, and when we left the place on Sept. 29, it was practically complete.” !5 Anderson wrote to Fleming in October about a visit he had made to Point Pelee to interview parties in a dispute over the trapping of muskrats and men- tioned that Taverner had built an artistic cottage on Big Island.!° In mid-October the Andersons them- selves bought a lot on Big Island.!0 At this point in his life Percy Taverner might con- gratulate himself on his pleasant prospects for the future with an increase in salary, and a summer cot- tage all ready for occupation next summer. But no CHAPTER 8. Field Collecting Point Pelee 1913 The first field expedition that Taverner undertook was in the summer of 1913 when with Clyde Patch and Charles Young he camped at Point Pelee for nine weeks. This was his first experience in the diffi- culties of mounting and carrying out a museum Vol. 110 sooner had he settled down to face the long winter than he had to face the worst possible news. Jim Macoun had become progressively ill and in November exploratory surgery showed that he had an advanced cancer. He died in January 1920 at the age of fifty-seven.!98 Taverner had expected that his pleasant relations with Jim Macoun, both as a friend and as head of the Biological Division, would contin- ue for many years. Now he suddenly was faced with the question of who would be appointed head of the Biological Division. The choice was between Anderson and himself, but since Anderson had a doc- torate in zoology and Taverner had no higher educa- tion the odds were weighted in Anderson’s favour. In March 1920 Dr. Rudolf Martin Anderson was appointed acting head of the Biological Division. In the two preceding chapters a fairly detailed account of Taverner’s early career at the National Museum of Canada has been given with emphasis on his work in the museum rather than in the field. It is hoped that the reader now has an idea of the working of the ornithological section. The next three chapters will focus on Taverner’s work by topical themes. Chapter 8 concentrates on the part played by Taverner in building up the bird collections and the field trips he went on himself, or organized for oth- ers to participate in. Chapter 9 outlines the role played by Taverner in protecting wild birds in the vital years of wildlife conservation in Canada between 1911 and 1919. Taverner was responsible for giving firsthand evidence, in his position as the National Museum ornithologist, on the situation at Point Pelee and at Bonaventure and the Bird Islands in the Gulf of St. Lawrence. Chapter 10 outlines the work that went into the writing of his first book which described the birds of Canada as opposed to merely cataloguing them. It is important to under- stand his objects and methods in writing this book since they set out the style and format which his sub- sequent bird books would follow. This chapter also will include information on the part Taverner played in enlarging and upgrading the Journal of the Ottawa Field Naturalists’ Club from a local journal into one designed to appeal to the interests of naturalists across Canada. (It became the Canadian Field- Naturalist in 1919.) These three chapters will bring Percy Taverner’s career to the year 1920 and the beginning of two decades of high hopes between the two world wars. expedition. The object of the expedition was to col- lect material for a Canadian Carolinian group show- ing characteristic birds and mammals in a typical Carolinian landscape.! In order to identify the plants and trees that he would need for leaves and branches in making the group he hoped to have the help of his 1996 friend K.C. Dodge, of Port Huron, who had com- | piled a list of the flora of Point Pelee as a result of visits there in 1910.2 He also hoped to go over to ' Pelee Island to verify the accuracy of Lynds Jones’ records from the Island.3 The first problem that faced Percy was uncertainty of whether the expedition would be able to leave Ottawa at all owing to the slowness of government in passing the yearly financial appropriation for the museum. Until Parliament voted the money at the beginning of the new financial year at the beginning of April everything was in doubt. Taverner wanted _ to reach Pelee early in May in order to catch the early spring migration but was delayed by ' Parliament and by problems of getting the various _ supplies required from the various departments involved.* Finally, on May 15, Taverner with Patch and Mrs. Patch (who had been engaged to do the cooking) arrived at Point Pelee in the rain with a cold wind blowing. The collecting of specimens began in earnest when Young arrived two days later. From now until the end of the expedition on 24 July Taverner kept a record in the Great Lakes Ornithological Club’s roll book kept in the shack, of birds seen or collected as well as brief notes on the day’s activities.5 Sightings began on a good note when Taverner found a large black looking duck on one of the ponds and thought it was an American Scoter [Black Scoter]. However, it was in an inter- mediate plumage without a white spot on its crown and the white on the neck obscured. Later he _ returned and collected it, when it proved to be a Surf _ Scoter (19 May).® On the same day he took a male ' Prothonotary Warbler. Both were first records for _ Point Pelee. In the last week of May they found sev- _ eral nests of Piping Plover with eggs while other nests were discovered in June. The first eggs to hatch had four young by 11 June, and Taverner took pho- tographs of them. Another bird sighting of interest was a Cory’s Least Bittern which flushed in compa- ' ny with a regular Least Bittern. They were both col- lected and the Cory’s proved to be a non breeding female. The three men pushed themselves hard since there was much collecting to be done before they had all the material they needed. They usually were away from camp soon after first light, and were often preparing specimens by the light of lamps in the evening. But not all was intensive work without relaxation. Percy had invited three entomologists of the Detroit Natural History Club whom he knew from his Detroit days, A.W. Andrews, W.W. Newcomb and W.S. McAlpine, for the last weekend in May. By chance three members of the GLOC, Fleming, Saunders and Wallace decided to use the club shack at the Point on the same weekend. This made nine men from Friday evening until Monday. It must have been great fun for everyone except Mrs. Patch who CRANMER-BYNG: A LIFE WITH BIRDS 1 did all the cooking. Apart from collecting in conge- nial company Percy had the rare opportunity to dis- cuss with other members of the GLOC the problem of preserving the beaches of Point Pelee from being carried away by an American company’s sand and gravel mining.’ One active member of the GLOC absent from the get-together was Bradshaw Swales who was recovering from an operation for appen- dicitis. To remedy this in a small way the visitors joined Taverner in writing the following “get well” letter to Swales: “We, the members of the Camp Coues Ornithological Club, having met for the ninth consecutive season, in our club quarters, together with the Detroit Naturalist Club who are visiting us for a few days, join in regretting the absence of their common member and send greetings and express sincere wishes for your speedy recovery to health and reappearance in our midst, June Ist 13. A.W. Andrews, W. W. Newcomb, Jas. S. Wallace, W.E. Saunders, J.H. Fleming, P. A. Taverner, W.S. McAlpine.” Bert Gardner, on whose land they were camping, was invited to join in signing the letter.® Early in June Taverner and Young started looking for samples of Carolinian foliage to use in their dis- play groups — such as magnolia, tulip tree and paw- paw. But while looking for plants they kept a sharp eye open for unusual birds. One day, while returning from a trip to Leamington they saw a Dickcissel perched on a post singing. When they returned two days later they found at least six pairs of Dickcissels in the field just north of the dyke at the base of the Point. They were evidently breeding in the fields there. While searching for nests they disturbed two Henslow’s Sparrows. In mid June they had more luck. They found a male and female Lark Sparrow near the Point, and a [Northern] Mockingbird sang in the cedars near to the shack one evening. Taverner was particularly elated by this because he had never heard a mockingbird’s song before. Just when the work was going really well things began to fall apart. Writing to Jim Macoun on 19 June he said that Dodge: was sick and did not expect to be able to come to Pelee until 5 July and then only for a week- end. This was unfortunate since Percy needed help from a botanist for identifications and advice on what should be included in a Carolinian group.? The following day he had more bad news. Writing to Fleming he said that Young had to return to Ottawa very recently on a family matter, which left the col- lecting party short-handed. The absence of Young indefinitely, and the failure of Dodge to come in June were blows that upset his careful plan. He did not feel that he should go to Pelee Island alone, he could not very well take Patch with him, so unless Fleming or Saunders could come with him he would have to call off the visit.!° As it turned out Fleming could not go and Saunders wired him at the last moment that he also could not go. This was a shame 80 THE CANADIAN FIELD-NATURALIST because Taverner ought to have explored Pelee Island and compared its bird population in the sum- mer with that at the Point. He never had another opportunity to see the territory where Lynds Jones made so many extraordinary sight records without corroborating them with specimens. Without Young the preparation of material for the Carolinian group took much longer than expected. Brock had instructed Taverner to have an exhibition ready for the members of the International Geological Congress showing the museum’s meth- ods and future plans.!! Taverner had hoped to leave Point Pelee at least by mid July so as to give himself sufficient time to prepare but it was 24 July when he left. Percy gave a last glimpse of the group driving into Leamington to catch the train in one of his let- ters to Fleming: “We had a great rustle getting away at all and when our wagons were at last loaded and filing into Leamington I felt like the manager of a circus on parade.”’!2 Apart from the information in Taverner’s report on the Pelee expedition (see note 1) it is worth not- ing that on his first field expedition Percy put one of his special skills, photography, to good use. He took photos of mammals such as four flying squirrels and their young, and reptiles such as the twenty-five blue-tailed [five-lined] skinks which Saunders saw while visiting the camp, and noted as “wonderfully variable in color’.!3 Percy also took various views of the scenery at Point Pelee in 1913 which have sur- vived.!4 When the party left the Point the fall migra- tion of shore birds had not yet begun in earnest and Taverner regretted the circumstances which com- pelled him to leave before the migration had start- ed.!5 But the expedition, in spite of problems, was a success. Taverner found Young to be an excellent field worker, and they got on well together, while Percy proved to be a tireless leader. As regards the bird collecting Saunders summed it up in a letter to Swales when he wrote: “The Point has done itself proud this year as it would any year if an enthusiast like Percy went down and stayed there.’’!6 Gulf of St. Lawrence 1914 (Miscou and Bonaventure Islands) With the successful completion of a field expedi- tion to the southernmost part of eastern Canada Taverner could now decide where he should go next. Because the museum collection was seriously weak in eastern specimens and comparatively better off in birds from western provinces he decided that sys- tematic collecting should begin in the eastern part of Canada.!7 The decision on where to go was not a dif- ficult one to make; it was made for him. In March 1914 Taverner heard a rumour of a movement to develop the top of the famous Percé Rock in the Cape Gaspé for sightseeing. Writing to Fleming he mentioned that the rumour included an Vol. 110 elevator to the top and a pavilion and merry-go- round.!8 A few days later, in a letter to Allan Brooks, Percy let his imagination run away, adding to the list of improvements to nature such things as summer hotels and shute-the-shutes [chute] as well. But he was serious about the implications for sea birds on the eastern coast of Canada where breeding places were becoming more restricted every year. The Rock, he said, was practically the last sanctuary of several species of birds. “T think a strong movement should be started towards the establishment of sanctuaries with this as the first one to be proclaimed.” He added that members of the civil service had their hands more or less tied by red tape in such mat- ters but those outside the service should get such movements started.!9 Meanwhile a letter was received at the museum from a resident living near Percé Rock protesting about plans to drive away the cormorants nesting on the Rock. Through discreet enquiries Taverner dis- covered that this was indeed being considered by the Fisheries Department because the cormorants were thought to be eating the salmon. fry which were being cultured in a nearby fish hatchery. Taverner did two things. He wrote to Brock, in his capacity as Deputy Minister of Mines, and warned him what was being planned and the wider consequences that might result from such action. He also told Fleming that the Fisheries Department apparently intended to have men scale the rock and break up the cor- morants’ nests. In that case, he said, “T am going to be on hand when they go up and make a survey of the place before the bloody work begins.”20 It was a challenge which Percy accepted swiftly. In late May 1914, Taverner and Young arrived in the Bay of Chaleur in the Gulf of St. Lawrence on muse- um business. Before settling in the Gaspé vicinity Taverner and Young spent three weeks collecting on Miscou Island in Chaleur Bay, New Brunswick (late May — mid June). Apart from a brief report on field work in the Gulf of St. Lawrence written by Taverner and printed in the “Summary Report for 1914’,?! there also exists a series of twelve letters from Percy to his mother and sister covering the period from the end of May to mid August.22 These letters give a vivid account not only of his bird collecting but also of the camp life and adventures of Percy and Young, and later of Frank Hennessey, in Pictou, Percé and other places. The entertaining content of these letters com- bined with Taverner’s lively style cannot be properly appreciated in a few short quotations. They deserve to be published in full as a contribution to Canadian travel literature. The first letter, dated 31 May 1914, was written from their camp on Miscou Island in Chaleur Bay, New Brunswick. The weather had turned cold and it snowed. A few days later there was a memorable a a a i ee a eee 1996 storm, graphically described by Percy, in a letter of 6 June, during which, he said, their “precious bird skins were scattered all over the place, and ham- mered into the ground and mud with the rain.” By ingenuity and determination they salvaged most of the camp equipment, and all except four of the 130 bird specimens. Another problem they had to over- come was caused by the difficulty of travel in that region in spite of local railroad and boat services. Taverner had estimated that they should arrive in Percé about mid June, but they had some difficulty | getting away from Miscou caused by weather and steamer timetable changes. Even then to travel by rail round Chaleur Bay took three days with two overnight stops, and eighteen pieces of baggage to transfer six times. As Percy remarked in a tone of resignation “Surely one has to have patience to travel about in this part of the country.”23 Unfortunately the railroad passed a few miles from Percé and they were dumped on a platform in a hamlet called Cape Cove in pouring rain at night. But the drive into Percé by cart next morning was exciting. Percy described the hilly scene as the village of Percé, dot- ted with pretty white houses came into view and the high ground of Mt. Joly above. “The next moment Percé Rock burst into view just beyond, coming out from the trees like the prow of a giant ship and forging ahead until full in view — the blue sea showing through the pierced hole* near the stern and the shaft-like column standing isolated in the rear. It was a most impressive sight .. . and around the top was a cloud of white gulls circling and hovering ... It took our breath away.”24 Before leaving Ottawa, Taverner was told by James Macoun that he ought to call on the lady from whom the museum first heard of the threatened destruction of the cormorants on Percé Rock. This is how he described his visit. “T called on a lady who now lives here — a Mrs. James — widow of an artist who built on a cliff face just oppo- site the rock, and has a most beautiful home with fine colonial interiors filled with articles of vortu — Sherraton chairs, willow ware and luster ware to burn — a Victor with fine records — two white cats — a3 2” telescope to study the birds on the rock — a bottle of Benedictine — and home made fruit cake, and a hearty welcome. She is an old lady, and informs me that the winters four nights a week are for bridge from which it is always morning when they get home.”25 A short visit to Bonaventure Island, just across from Percé village, to see the large Northern Gannet colony on the ledges of the cliffs facing the Gulf of St. Lawrence showed Percy the need for time to study them. He wrote to Brock suggesting that it would be desirable to remain in Percé all the summer to study the cormorants. The next step was to camp on Bonaventure Island in order to take photographs and make a collection of specimens. In a letter of 6 July he described making camp after an uphill climb CRANMER-BYNG: A LIFE WITH BIRDS 81 on a rugged terrain carrying half a ton of baggage. They camped on smooth green sod among spruce trees with large purple irises growing nearby. They slept on balsam boughs, and cooked on a stove fire- place, getting water from a rippling brook. But that was incidental, although very enjoyable. It was the birds that attracted them. It was an exciting chal- lenge as he described it. “To crawl to the crest of the cliff and lie on our stomach — not a practical attitude but the only way in which one could look with any easiness, and to see a thousand Gannets — birds as large as geese — pure white with cream heads and jet black pinions and long pointed tails sitting unconcernedly on their nests almost within hand’s reach — ledge after ledge of them on their nests down three or four hundred feet — the whole face white- washed with their excrement (odorous) down to the blue sea breaking in great white masses and shreds below. It was a sight indeed, and then when for every bird on the nest there was another in the air sailing in great circles past the cliff — some so close that one could see the color of their eyes, and others lower down, until the air was full of them all passing, passing, passing, in the same direction, then out over the sea and back again to pass again, like the rim of a great wheel or the center of a cyclone bearing a cloud of snowflakes with it.”’2° In the same letter he described where murres and puffins lived — in crevices in deep horizontal cracks in the cliff. Here they laid their eggs which were scattered all along the rocks without any form of nest. The parent birds had the task of keeping their young from coming into the open. Percy described how between boulders they could see many young birds “lined up like soldiers on parade against the rear wall with occasional parents standing guard before like officers. Sometimes one would suddenly make a dash out, knocking its comrades pell mell as it came, and almost into our faces”. But they had no success in finding [Atlantic] Puffins’ eggs which must have lain in the deepest part of the cliffs. At this point Taverner received a letter from Brock authorizing him to remain and study the cormorants and telling him that the Evinrude motor that he had asked for had been ordered. After spending just over a week on Bonaventure Percy began the next stage in his field work. He returned to Percé and then went to the town of Gaspé to take a look at the cormorant question from the fish hatchery and the salmon rivers. In his next letter he devoted a paragraph to the adventure he had getting from Percé to Gaspé by train. Because the train was expected to be three hours late various complications followed and he ended up by being driven in a rig by a farmer’s daughter to a town a few miles distant. Her name was Clara. This is how Percy described her: “Great strong girl, brown as a berry, beautiful hair and strong arms and dressed neatly and with open-work stockings. She knew how to drive also, and made the horse go like a man”’.27 In Gaspé he called on Commander Wakeham, in charge of the fisheries of the Gulf, whom he found 82 THE CANADIAN FIELD-NATURALIST living in “a dream of place” which he described in some detail. Wakeham had just returned from the Labrador in the Marine and Fisheries steamer Princess. The two men appear to have got on well, and Wakeham offered Taverner “every service in his power’. Percy now needed to make a study of the contents of cormorants’ stomachs in order to deter- mine the amount of salmon fry they were eating. Commander Wakeham arranged for Taverner, Young and Hennessey to be accommodated in the ample buildings of the Hatchery. Later in July they were collected at Percé by the Commander and taken in the Princess to Gaspé. In his next letter Percy wrote: “It was a most delightful trip. The steamer is like a yacht, our host was perfect and the chef a genius. We had supper on board and it was a treat. We were sorry when the anchor dropped in Gaspé Basin and our voyage was ended.”28 The assistants at the Hatchery were only too will- ing to help Taverner and his assistants to explore two cormorant rookeries in the bay and to obtain some nests. On another day they were taken to the St. John River to see how the fisheries people planted young salmon, which he described in some detail. There was one detail of particular interest when he wrote: “We saw lots of salmon in the water — big ones 20 and 30 pounds in weight, but could catch none as the river is under lease to a private American club.” They continued this pleasant life, collecting cor- morants and analysing the contents of their stom- achs, and studied their plumages.?? For young Frank Hennessey it was especially enjoyable since he was invited to all the parties and dances. By early August he could write that they had thirty cormorant skins. This was far more than they needed for specimens but they were collected in order to get the stomachs and it was best to save the skins as well. The war between Germany and various European countries including Britain, and therefore Canada, broke out at the beginning of August. Writing home on 8 August Percy alluded to it briefly and the vari- ous rumours about the German fleet, adding rather flippantly: “Perhaps the next time you hear from me will be from a German prison after being captured in the Princess.” This referred to a short visit to the Magdalen Islands (Iles de la Madeleine) and the Bird Rock. Percy was elated about going to the Bird Rock in a special steamer which he called a “luxury no other birdmen ever had — most of them had to take what passage they could get in smelly fishing schooners”. At the end of this letter he tempted fate by writing “I cannot think of any more news now, though when I get back from Bird Rock will proba- bly be loaded with it.”39 Percy should have known better. The next letter, dated 12 August, began “Another disappointment. In a way the fortunes of war. We did not get to Bird Rock.” What happened was this. They landed on Amherst Island in the early Vol. 110 morning to wait for the mail steamer from Pictou. They were given two hours to collect what birds they could. When the mail boat did not arrive they made for Grindstone Island, expecting the mail boat to catch them there. They set about skinning the twenty birds they had collected at a table between decks. At Grindstone the Fisheries officer, who was also the Customs collector, came aboard. His news was bad. He had received orders from Ottawa to allow no boats to clear for the Gulf because of a report of two German ships near the mouth of the Gulf. The Commander was not bound by this order, being on government business, and referred the decision to Taverner who did not feel like taking the responsibil- ity. Since it was not necessary to visit the Bird Rock he advised returning to Percé. As he wrote “the Bird Rock might wait”?! The remainder of the time was spent on visits to Bonaventure Island where they collected a [Black- legged] Kittiwake’s nest, and carried out more pho- tography. This time they brought a rope with them and Percy was able to lower himself through a crack deep down into the cliff where it was almost too dark to see. Lying flat on the damp ground, on rough peb- bles, and pushing his camera ahead of him, he came out on a ledge filled with puffins and got some good photographs of them at a range of ten or fewer feet. Because he appeared from inside the cliff face the birds showed no sign of fear. Young found a nestling Atlantic Puffin at the back of another crack. Percy described the scene: “The ground was strewn with unfertile, unhatched eggs of Murres, now in advanced stages of decomposition, but in he went. Two eggs broke and almost stifled him, but he got the bird, though his clothes were smeared with old egg.’’32 How did they ever get rid of the reek of rotten egg from their clothes? Perhaps they didn’t mind since they had achieved one of their prime objects in returning to Bonaventure. Another was to find a young Razorbill but they were too late; the young must have been taken off to sea by their parents. On another visit they expected to spend the night on Bonaventure in order to be ready to catch the photo- graphic light on the bird ledges. They also required some young Petrel [Leach’s Storm Petrel], a Northern Gannet’s nest, and more photographs (autochromes).33 By mid August they were packing up to leave; Percy went to say goodbye to Mrs. James and left us this pen picture: “She takes great pleasure in assuming the office of Lady Patroness of the Rock, and delights to entertain visitors to it. Her figure — that of a little old lady, slightly stooped, her arms full of packages and leading a mon- grel dog by a strap is a familiar one on the street (road) of Percé.’’34 What, then, was there to show for the work of three men during a three month field trip? What were the results of the Percé — Bonaventure expedi- 1996 tion? The first and chief result was that Taverner, as the ornithological specialist at the National Museum of Canada, had been able to form an opinion, based on research, of the cormorant question: His first impressions were given in a letter to his mother and sister as follows: “The Cormorant question is interesting. I always thought the damage they do was exaggerated but not to the extent that the evidence shows. Their effect on the salmon is almost nil. Those in tide and salt water feed almost entirely on sculpin and eels. Those in fresh must eat salmon, but one of their bitterest enemies only admits of seeing 3 or 4 Cormorants up river in a year, so you can see what that means. However I will have an interesting report to write as it touches several phases of the subject that I have always wanted to write about.’’35 His nature observations were set down in a study printed in the museum Bulletin in April 1915 entitled “The Double-crested Cormorant (Phalacrocorax auritus) and its Relation to the Salmon Industries on the Gulf of St. Lawrence”.3° This dealt with the bird’s food habits at length. His conclusion, based on data examined, was that they fed almost entirely on species other than salmon, but that they inconve- nienced fishermen locally by taking some of the her- rings out of their nets. Taverner also took this oppor- tunity to start a study of a colony of breeding Northern Gannets including a collection of his own photographs. This study was to continue the follow- ing summer and was eventually published in The CRANMER-BYNG: A LIFE WITH BIRDS 83 Ottawa Naturalist in May 1918 at a time when the whole future of Bonaventure Island was uncertain. This article represented his mature thoughts on the birds of Bonaventure Island, especially the Gannets. Finally the expedition was a valuable learning expe- rience for Taverner that did much to increase his self confidence. He matured as a consequence. He real- ized this when he wrote: “This has certainly been a season crowded with ornitho- logical experiences and I am commencing to feel like a veteran.”37 It also gave a special interest to his correspon- dence with some of his ornithological friends. Two letters exist between A.C. Bent and Taverner written in December 1914. Bent requested information about Taverner’s trip to the Gaspé and the plumage of gan- nets to be used for his Life Histories. Taverner replied that they obtained specimens in all mid sum- mer plumages, and that Hennessey made a consider- able number of water colour sketches of the bill, feet, eyes and soft parts of various species of various ages, including the colour of the mouth.38 Bonaventure Island and North Shore of Gulf of St. Lawrence, 1915 When Taverner returned from Percé in the fall of 1914 he was drawn more deeply into the urgent prob- lems of bird protection and the conservation of bird habitats in eastern Canada. While writing his report on the cormorant problem he also began writing up a. = Gannets on bird cliffs on Bonaventure Island, 1915. Photograph by P. A. Taverner. 4 (Reproduced courtesy of the Canadian Museum of Nature, number 32273.) 84 THE CANADIAN FIELD-NATURALIST his material on the gannets of Bonaventure Island, and realized that he would need to return to Percé the following summer. The object of the 1915 expedition would be to complete his study of the gannets, to take more photographs of them on the cliffs, and more especially to make a film of breeding Gannets on their ledges and in flight. Such a film would be very useful for educating the public, and rousing support for protecting the gannets at their breeding sites. The experience gained in 1914 would be turned to good account during the 1915 expedition. Taverner took with him Young, Patch and Johnson. They reached Percé at the beginning of June, and began working the woodlands and shores near the village, with frequent visits to the bird ledges of Bonaventure Island. Patch and Johnson were responsible for the preparation and mounting of exhibition material including specimens. They stud- ied habits and attitudes, took casts and photographs of rocks and accessories for reproduction in life his- tory groups. Taverner and Young gathered additional scientific information and material, taking pho- tographs with autochrome and ordinary plates, as well as taking moving pictures of the bird ledges. Late in June Taverner‘and Young left in the steamship Princess on a three week voyage along the north shore of the Gulf of St. Lawrence, courtesy of the Department of Marine and Fisheries. Patch and Johnson remained at Percé till late August.3? This is a bare outline of their field work. But fortunately Percy’s letters to his family and to Fleming bring the expedition to life. As soon as Percy got to Bonaventure Island he started taking photos in spite of poor weather — fog or smoke (bush fires) nearly every day. He took with him a graflex camera and with the use of new, fast plates he hoped to improve on the results of last year. He took what he felt were some good autochromes on the ledges and sent the plates off to the museum to be developed to find out if they were satisfactory. As he explained in a letter home: “The exposures are rather long, and with living birds it makes the result rather uncertain.” Meanwhile he was entertained by Mrs. James (he had met her the previous summer) who invited Percy, Young and Mrs. Patch to play bridge. Percy reported that Mrs. Patch had been entertained by Miss LeBrun, and was getting into the Percé society. Although it was not extensive it was not to be sneezed at. “These Jersey people are true gentlefolk”.4° On the other hand Percy was not impressed by the French inhabitants. Apparently a party of eight sportsmen went in early June to Bonaventure Island and shot birds indiscriminately all day. Writing to Fleming he gave a gruesome account of the carnage, and was clearly upset by what he had seen.*! In a let- ter to his mother he omitted such details but explained why they acted as they did. Vol. 110 “Last Sunday a party of eight sportsmen (?) went to the Island and shot all day. They were intelligent looking men too, not ignorant simple fishermen, — but French. How the local English dislike the French, and one can- not blame them. Percé is the county seat of Gaspé Co., and there are a number of French Lawyers and a doctor here. Men of obvious educational advantages, but you would be surprised at their narrowness. To a lawyer I suggested the iniquity of shooting the harmless birds, and he could not grasp any idea of wrong in it, and thought the answer There are lots of them was sufficient justification. But how could they be other than narrow? How can they imbibe modern trends of thought and ideas without reading, and they are confined to the few French Canadian publications which are notoriously under church domination. This is, I think, the real race problem in Canada — getting the French to read English, where they would have the press of all English speaking countries to draw from, and not but the one sided presentations under the control of reactionists or others as circumscribed as themselves.” Percy then gave a detailed description of how the local Catholic Church celebrated the feast of Corpus Christi. He was an interested spectator, watching at a distance.*? Commander Wakeham, who had been in com- mand of the steamship Princess in 1914, had died in the interval, but the new commander invited Taverner and Young to accompany him on the ship’s next cruise in the Gulf of St. Lawrence. They embarked at Gaspé on 27 June. The ship’s route was along the north shore of the Gulf as far as Blanc Sablon at the border between Quebec and Labrador by the Strait of Belle Isle, and close to the Atlantic Ocean. The object of the voyage was to regulate the fisheries by issuing licences, and enforcing the regulations. As the route followed the land closely there were daily opportunities for going ashore though little opportunity to explore or collect for any length of time anywhere. They reached Blanc Sablon on 30 June but found ice coming in through the straits of Belle Isle, and were not able to enter the harbour until 8 July. They returned by the same route, making a final short stop at Ellis Bay on Anticosti Island before reaching Percé on 15 July. There is a fairly detailed account of this voyage in a letter that Percy wrote to Fleming in which he said that they were unable to obtain many birds but that he got a good idea of the geography of the coast and poverty of the fishermen living there. Bird information was not his main object on this trip but rather to gather information on people in relation to birds so that he would be in a good position to push for legislation to protect the birds of the Gulf. His first thoughts on this subject were expressed in this letter, namely that the coast was a mass of islands, that it would take an army of game wardens to patrol them, that compulsory gun licences would be the only feasible method. The people, he explained, were so poor that they killed 1996 CRANMER-BYNG: A LIFE WITH BIRDS 85 f Cl] e = The wheel of the C. S. S. Princess, 1915, from the left: P. A. Taverner, Captain Chalifour, and C. H. Young. (Reproduced courtesy of the Canadian Museum of Nature, number 32351.) birds to survive, and if it was not for the abundance of eider ducks many would starve to death. In conclusion Taverner mentioned the more detailed exploration of the same stretch of coast made by Charles Townsend between the end of June and September 1915. Townsend and Taverner had met through AOU meetings, and had corresponded since 1911. He had studied the birds to the east of Blanc Sablon in 1906 and to the westward of Natashkwan [Natashquien] in 1909 and 1912. Now, in 1915, together with a botanist, Harold St. John, he explored the coast in a small schooner. Since he was working in the interest of the National Museum, but was not tied to a strict schedule, Townsend had good opportunities to make a detailed study of bird life there. His preliminary report on the expedition was included in the museum’s “Summary Report for 1915” following Taverner’s report.*4 Both reports contained remarks on the need to preserve birds in the Gulf of the St. Lawrence. 86 THE CANADIAN FIELD-NATURALIST The Prairies, the Rockies, and British Columbia in 1917 Taverner did not undertake any field work during 1916 but in 1917 he and Young were at East Shoal Lake, Manitoba, by mid May. The lake is about thir- ty five miles northwest of Winnipeg. It had been vis- ited by a number of egg collectors and ornithologists between 1867 and 1912, the best known being Frank Chapman in July 1901.45 In order to make a repre- sentative collection of birds occurring in Manitoba Taverner chose one location and worked it thorough- ly. Manitoba is the most eastern of the prairie provinces and is where the interface of prairie and eastern woodlands occurs. As such it is one of the most important areas of geographical distribution in Canada.*° In mid-June Taverner and Young went to Red Deer, Alberta, where a small boat was built to Taverner’s design capable of carrying them and their camp and collecting equipment. With an outboard motor attached they began the descent of the Red River. This entailed a journey of some 217 miles to the southernmost point in the river, well southwest of Drumheller, where they set up their final camp on 19 July near to the Geological Survey’s paleontolog- ical collecting camp where Charles Sternberg was excavating dinosaurs. They spent nearly a month of the expedition setting up camps along the way from which they made observations and collected speci- mens. Taverner left Young and continued into British Columbia while Young remained collecting in the vicinity of the camp until late September. The expedition was written up by Taverner with a six- page narrative and an annotated list of the species recorded.*7 Taverner’s next stop was at Banff where there was a small museum. Here he visited the White Museum of the Canadian Rockies and met with some mem- bers of the Alpine Club of Canada.‘*8 He then went to visit Munro at Okanagan Landing where the latter was collecting, for sale, birds, mammals, insects and plants.4? From there he went to Vancouver Island to see “Professor” Macoun, and then up the coast to Prince Rupert opposite the Queen Charlotte Islands.5° From there he took the grand trunk Pacific Railway to Hazelton where he spent a few days assessing the bird population.°! From there he con- tinued by rail to Jasper Park where he joined Jim Macoun in camp beside a small glacial lake below the peak of Mount Edith Cavell. It was here that Percy met for the first and only time the well known collector of that era — employed by the Macouns in their bird collecting — William Spreadborough.>2 Finally, for the period September 17-26 he returned to the same part of Shoal Lake as earlier to obtain an idea of the autumn conditions there and fill some of the gaps left in the spring collecting. When he reached Ottawa he had been on an extended field trip of nearly five months. It was pos- Vol. 110 sible to be away from the museum for so long con- secutively because the exhibition material was packed up and much of the building was occupied by the Senate and House of Commons. Most of Taverner’s travel was spent on collecting and note- taking, but some of his time was spent meeting peo- ple in the world of ornithology as well as museum works. He visited the Provincial Museum of British Columbia in Victoria where he took notes on the bird collection. Wherever he went he made good use of the opportunity to weigh up the value of the local topography for birds, and to meet old or make new acquaintances. He paid his last visit to John Macoun, the grand old man of Canadian botany, while in Victoria and managed to spend two weeks with his son, Jim Macoun, in the Rockies. The meeting there with William Spreadborough was one that Taverner remembered with pleasure even though he never managed to persuade Spread- borough to join him in any collecting expeditions after Jim Macoun’s death in 1920. What he wrote about those few weeks in camp in Jasper Park served as a warm tribute to William Spreadborough, the boy from Bracebridge, Ontario, whose collec- tions and observations were “the real beginnings of our national systematic collections and the basis of many scientific papers by specialists.”53 This was how Percy described Spreadborough in camp after a long day of hard work. “By day he was afield investigating the most hidden recesses of the mountains or skinning specimens, distrib- uting the by-products of his operations to the Whiskey Jacks momentarily lured from the caribou. In the dark of the evenings it was pleasant to sit with back against a shielding rock or sheltering trunk, feet extending towards the grateful camp fire, listening to William ‘reminiscing’ from his inexhaustible store of memories of travel and exploration by field and stream. He could go on and on and, being a natural story teller with a pho- tographic memory that forgot no detail, interest was always sustained. His energy was inexhaustible and no mountain was too steep or way too rough for him to face if a desirable specimen were the objective.”>4 During field expeditions Taverner was on the alert to identify every bird he saw and to record in his field notebook whatever he had identified beyond all reasonable doubt. “No record”, he wrote at this time, “is absolutely unassailable until specimens are secured and examined by competent authority”.55 Anything unusual he saw concerning such things as plumage, behaviour and migration he recorded in notebooks. When writing letters to his friends during these expeditions he discussed what he thought would interest them, not only about birds and plants but also about the people he met in his travels. His tales were mainly of birds and men. Exploring in Ontario 1918 and 1919 Taverner had intended to visit Mitchell in Regina during 1918 to discuss with him the best area in 1996 CRANMER-ByYNG: A LIFE WITH BIRDS 87 sik C. M. Sternberg’s field camp, Red Deer River, Alberta, 1917. Standing: Dr. R. L. Rutherford (University of Alberta), P. A. Taverner, C. M. Sternberg, C. H. Young, C. H. Sternberg, Dr. J. A. Allen (University of Alberta), three unidentified assistants are seated in foreground. (Reproduced courtesy of the Canadian Museum of Nature, number 40000.) Saskatchewan for an expedition in 1919, as well as _ to collect in the prairies. But he was forced to change | his plans. In a letter to Fleming he explained what had happened: “Parliament was so late in passing supply bills, you know that they never bring up an appropriation bill until the last thing when everybody is too anxious to get home to discuss them, that the western trip was not worth while. Instead of that I planned a short trip along the Rideau Canal to Kingston to get better acquainted with local conditions and relationships.’5° In a rowboat with an outboard engine with a canoe _ in tow, and accompanied by a man from the taxi- _ dermy shop, he set off on 11 June. On the day they ' left the weather turned cold and continued with rain | and wind all the time, only clearing just as they returned to Ottawa on 15 July. Since then, he told | Fleming, the weather had been roasting. They must _ have looked an odd sight, huddled up in waterproofs _ sitting low in the water in a rowboat, trying to find | birds in the rain in the middle of the summer on the _ Rideau Canal, river and lakes connecting it. The result was minimal — just over 100 species record- ed, the best bird seen a Least Bittern, and the obser- | vation that the marshes of Smith Falls were particu- larly good for hunting. As though this experience was not enough Percy spent two weeks with Clyde Patch with similar equipment exploring the Ottawa river as far as its mouth, this time in better weather but with disappointing results. Writing to Brooks he put the best face he could on the expeditions. He said that although they did not find a large number of birds they managed to fill a few holes in their local collections, and obtained considerable information on local conditions, and therefore it was worth while. One conclusion he came to may have been of slight interest at the time when he wrote: “I think there is quite a migration route up the Rideau system but none at all on the Ottawa [river] below here.”5’ All in all these boating trips round Ottawa in 1918 were something of an anticlimax after the achievements of the previous summer. In the spring of 1919 Taverner suffered the same frustration over his plans for an expedition to the west as in the previous year. By mid May he still had not left. He told Fleming that everyone was waiting for the word to go or stay; meanwhile the birds were migrating “but no one cares except us”’.°8 Eventually it was too late. Instead Taverner took advantage of 88 THE CANADIAN FIELD-NATURALIST the newly opened stretch of the Canadian Northern Railway from Cochrane westward to Kapuskasing. Accompanied by Claude Johnson he spent three weeks in Kapuskasing district making a representa- tive collection of summer breeding birds in a north- ern Ontario forest area that had recently become accessible. Kapuskasing at that time was a “model town” mainly consisting of a prisoner-of-war intern- ment camp and a Dominion Experimental Farm, and an Ontario government Soldiers’ Settlement Project. Letters from Percy to his family, if he wrote any, have not survived, but letters to friends give some details. He wrote Fleming an account of this trip dur- ing which he collected a fair number of specimens including a Richardson’s Owl [Boreal Owl]. Lincoln’s Sparrow and Mourning Warbler were found to be abundant breeders near Kapuskasing.°? Taverner used his own design of field notebook on all his field trips, and the ones for the Kapuskasing area are an example of what these were like. Thus volume 1, bound in red, contains eight pages of type- written notes, together with photos of their camp, which provide some information about the trip itself. Species lists are in part of volume | and all of vol- ume 2. These are the expeditions and shorter trips which Taverner managed to undertake between 1913 and 1919.0 There was one expedition, supported by the Geological Survey, for which Taverner was invited to find an experienced field naturalist. This expedi- tion was to be led by Charles Camsell, “Geologist in charge of Exploration” of the Geological Survey, who planned to make a canoe traverse from Lake Athabasca to Great Slave Lake in 1914. The object was to observe the geography, geology and biology along the route. Since only three canoes and seven men were being taken it would be impossible to bring out skins of birds and mammals. The naturalist chosen, therefore, would have to be capable of tak- ing reliable notes of the wildlife seen. Taverner first asked Brooks who declined, and then Munro who also declined. Taverner next asked Fleming’s advice. They came to the conclusion that there were no other Canadians with suitable qualifications for the job, and available to spend from May through September on the expedition.®! Taverner now started searching for an American naturalist with the right qualifications. His contacts in the AOU were useful in this search. Eventually CHAPTER 9. Bird Protection The origins of bird protection in North America go back to the first half of the nineteenth century when a few sportsmen realized that game birds were begin- ning to grow less numerous in their own localities. Vol. 110 Dr. Arthur A. Allen of the Cornell University Zoological Laboratory suggested a suitable man who was keen to go. This was Francis Harper who had recently obtained a degree in biology at Cornell University and was now with the Brooklyn Museum. After travelling over land from Edmonton to Athabasca Landing and down the Athabasca River to its mouth, in May and June, Camsell and party left Fort Chipewyan on the northwest shore of Lake Athabasca in late June 1914. From the middle of the north shore they started their traverse towards Great Slave Lake, by way of Lake Tazin, and the 'Tazin River to its junction with the Taltson River. From here they went down the Taltson reaching Great Slave Lake early in August, a distance of about three hundred miles. They then followed the south shore of Great Slave Lake to the small trading post of Fort Resolution. From there they paddled up the Slave River to Fort Smith, a distance of two hundred miles, in ten days. Here Harper was supplied with horses and, together with the buffalo guardian, made a six day trip into the wood buffalo country. By mid- September the party had returned to Fort Chipewyan.°? Harper wrote a biological report of the expedi- tion for Taverner who included it with his own report in the Zoology Section of the Geological Survey’s “Summary Report for 1914”. Harper noted 85 species in the region between June and late August among which the following were con- sidered by him as interesting records: Short-billed Gull [Mew Gull] nesting by the north shore of Lake Athabasca and during the traverse, a range extension southward for that time;®* Arctic Tern; and Hutchins’ Goose [Canada Goose] on the Taltson River where four young were taken; sev- eral records were received of Whooping Crane being in a locality they passed through; an estimat- ed 700 or more Stilt Sandpiper in the Athabasca delta in early June; a single Rock Wren at Fort Chipewyan in mid June. Harper collected 93 speci- mens of birds and took over 450 photos of topogra- phy and vegetation.®® For Taverner this expedition brought back useful information on the north west- ernmost part of Saskatchewan, the north eastern- most tip of Alberta, and a small part of the Northwest Territories south of Great Slave Lake. A selection of the specimens taken by the expedition was a welcome addition to the National Museum collection. By mid-century several sportsmens’ organizations had been formed in various states for the preservation of game. These were active in their state legislatures to secure laws to protect game. During the second 1996 44460.) | half of the century the fashion of decorating ladies’ | hats with bird plumes caused the millinery trade to | buy large quantities of plumes from sea birds to _ Hummingbirds as well as various species of tropical birds. By 1883, when the American Ornithologists’ | Union was founded, the need to protect non-game _ birds had become urgent. A campaign was started to _ educate public opinion, data on the plumage trade _ was collected, and work for bird protective legislation on a national scale was begun.! | The term “Audubon Society” was coined by George | B. Grinnell and first used in his magazine Forest and Stream in 1886, while at the same time the AOU orga- nized a Committee for the Protection of Birds. _ However, it took ten years to rouse sufficient support | to make the work of these two organizations effective. | In 1900 the AOU Committee decided to protect breed- ing colonies of gulls and terns by means of guards. The money with which to pay them was given by a private individual. To this action can be traced the beginning of the Sanctuary idea in America.? In 1904 a “National CRANMER-BYNG: A LIFE WITH BIRDS 89 | Taverner in an open boat with an outboard engine while exploring the Rideau waterway system between Ottawa and Kingston in search of birds in June 1918. (Reproduced courtesy of the Canadian Museum of Nature, number Association of Audubon Societies for the Protection of Wild Birds and Animals” was formed. By the first decade of the twentieth century public interest in bird study had developed rapidly with the publication of books of a popular nature such as Frank Chapman’s Handbook of Birds of Eastern North America, first published in 1895 and reprinted regularly thereafter, and Chester Reed’s Bird-Guides, specially useful for beginners, in 1906. Meanwhile the Federal Government, prompted by the AOU, had established a government office of Economic Ornithology in the U.S. Department of Agriculture in 1885 which became the Bureau of Biological Survey in 1905. This office issued reports dealing with the food habits of wild birds, their eco- nomic status and laws for their protection. These reports were available to the public on written applhi- cation and Taverner made use of this service while living in Detroit. The effects of the information con- tained in these publications on conservation groups, on government officials and politicians were consid- 90 THE CANADIAN FIELD-NATURALIST erable, and probably contributed to the first federal measure aimed to protect birds — the Lacey Act of 1900 (amended 1909). This act was designed to curb the bringing of foreign wild birds into the United States. Another act, passed in 1913, was aimed at bringing under federal control birds that migrate twice a year between different states. Eventually the principles of this act became the subject of a treaty between the United States and Canada.3 Because Taverner lived in Michigan during the decade 1900-1910 and was an active ornitholo- gist in touch with the Bureau of Biological Survey, the AOU, and an avid reader of journals and books he knew about developments in the wild animal pro- tection movement. When he came to Ottawa in 1911 he not only had up-to-date information on the sub- ject, he also had acquaintances among some of the leaders of the movement in the United States. In Canada, however, serious concern over the pro- tection of wildlife had only been developing since about 1890. During the first half of the nineteenth century the impressions one gets from bird notes that have survived show that many species were quite abundant. Charles Fothergill kept systematic notes on birds seen along the north shore of Lake Ontario between Port Hope and Toronto in the period 1816- 1840. He mentioned a total of approximately 180 species in his manuscripts, most of which were shown as relatively abundant.* During the second half of the nineteenth century some people in the more settled areas began to realize that game bird populations were in decline. One example was the Wild Turkey in south western Ontario which was said to be fairly abundant in the Lakes Erie and St. Clair regions in the 1860s. But the Ontario Game and Fish Commission in its 1872 report predicted its extinction in Ontario which, in fact, happened by 1904.5 Non-game birds were also beginning to feel the pressure of increased hunting combined with loss of suitable habitat. The best known example of this was the Passenger Pigeon which was still very abun- dant in Ontario in the 1870s but in the 1880s suf- fered a drastic decline throughout its range.° The Ontario government set up a Royal Commission to review the wildlife situation in the province. Its report, published in 1892, and based on the evidence of many sportsmen, stressed the decline in numbers of various birds and mammals, some of which were thought to be on the verge of disappearing. As a result Ontario established a Game Protection Act in 1893, while similar measures were established by British Columbia in 1895, Quebec in 1899 and Manitoba in 1900.7 At that time in Canada protection of wildlife was considered a provincial matter to be regulated, if at all, mainly in the interests of sports- men and farmers. The myth of superabundance, and the belief in the duty of government to aid the exploitation of all natural resources had moulded the Vol. 110 federal government’s attitude to wildlife. The first Dominion park to be established by the federal goy- ernment was Rocky Mountains Park at Banff in 1887. Although the motives of the government were not in creating a wildlife preserve nevertheless “the very wording of the Act went a long way towards establishing, in theory at least, the principal of wildlife protection”.’ As Janet Foster clearly shows in her study Working for Wildlife: the Beginnings of Preservation in Canada, during the twenty five years between 1887 and 1912 a few senior civil servants in departments most concerned with wildlife “turned their own goals of wildlife preservation into govern- ment policy”. When James Harkin was appointed Commissioner of Dominion Parks in 1912 he had “a clear and unfailing vision of what wilderness, parks, and wildlife signified for the Canadian people...”.9 Percy Taverner arrived at the newly opened Victoria Memorial Museum in Ottawa just at the time when the federal government, prodded by a few senior civil servants and the nucleus of a public opinion, was about to become seriously involved in wildlife conservation. His arrival was extremely opportune. In 1909 a bill was passed in Canada to establish a Commission of Conservation which would be sepa- rate from the machinery of government. Although it would report to Parliament through the Minister of Agriculture it would be responsible only to Parliament as a whole. It was intended that the Commission should be “an independent... and fully non-partisan body that would explore all questions pertaining to natural resource conservation in Canada.”!0 In 1911 Parliament established a Parks Branch under the Department of the Interior to manage and develop national parks. The appointment of James Harkin as Dominion Parks Commissioner was a sig- nificant step forward that would result in greater pro- tection of parks and wildlife throughout Canada.!! Several things occurred in 1913 to start a train of events that would result in far reaching changes in the protection of birds in the next few years. The Commission of Conservation held its annual meeting in January each year and subsequently issued an annual report. At the January 1913 meeting J. Walter Jones, who had been asked by the Commission to report on fur farming in Canada, pointed out the changed conditions in regard to nat- ural resources since the passing of the British North America Act in 1867 when natural resources were placed under provincial jurisdiction. For example, he asked, “what legislative body should have charge of migratory birds?” This was an important question since the Weeks-McLean bill was before the United States Congress. A copy of the bill was published as an appendix to the 1913 Report.!2 Soon after the Commission’s meeting Maxwell Graham, chief of the Parks Branch Animal Division, sent a memo to Harkin suggesting that the time was suitable for leg- | | 7 | | islation by which migratory birds could be protect- ~ ed.!3 Graham next wrote to James Macoun for infor- - mation that would corroborate what he had told i Harkin, and Macoun passed the letter to Taverner to _ answer.!+ Taverner was in a good position to give information on the need to regulate spring shooting ' by an international agreement. He also gave some estimate of agricultural losses in Canada because many species of insect-eating birds were being killed | legally in the United States, but recommended Graham to obtain information from the U.S. ' Biological Survey which had been collecting biologi- cal data since 1885.!5 Harkin meanwhile wrote to the | secretary of the American Game Protective and | Propagation Association about the Weeks-McLean bill saying that it was felt that Canada ought to coop- erate with the United States in protecting species of ‘birds which migrated between the two countries.!° About this time another senior civil servant in | Canada involved with wildlife, Gordon Hewitt, Dominion Entomologist with the Department of _ Agriculture, wrote to the Chief of the United States | Biological Survey. He wrote as a private citizen inter- ested in bird preservation, and asked for information on the Bill for Federal protection in the United States. He also suggested that Canada and the U.S. should | find a way to cooperate in protecting birds. As a result he was sent material relating to the Weeks- + McLean bill then before the Senate.!? Harkin now _ wrote to Fleming and Saunders asking whether there was a need to protect migratory birds in Canada. _ After his return from the Point Pelee expedition of 1913 Taverner became more and more involved in | the question of wildlife protection through the » Commission of Conservation and its secretary, James White.'§ The Commission of Conservation | was already gathering information on twelve or more } issues one of which was about the need for animal | sanctuaries on the North Shore of the Gulf of St. Lawrence. An address on this need was presented at the Commission’s annual meeting in 1911 by Lt. Colonel William Wood, a resident of Quebec province, who had firsthand experience of the situa- tion. Wood sent a copy of his address to many scien- tific journals, and to a number of prominent men 5 who realized the need for conservation of wildlife, | requesting their comments. From their replies he pre- ; pared a report which was published in the _Commission’s report of 1912.!9 This was a detailed review of the situation in the Gulf of St. Lawrence together with endorsements of several prominent | people. For example Dr. John M. Clarke, Director of | the New York State Museum, who had made repeat- ed visits to the coast of the Gaspé and the islands in | the gulf, reported that on the Magdalen Islands there | was a wide variety of shore birds during the summer _ months. These were supposed to be protected by law but the law was not in the least respected by hunters. a — ; 1996 CRANMER-ByYNG: A LIFE WITH BIRDS 91 They knew they were too far away from authority to be prosecuted. The islands, he said, were also known to students of birds from the United States who were guilty of collecting shore birds for their skins, and for their eggs, during the breeding season. On the Bird rocks one American oologist was known to have taken 369 clutches of eggs. On Bonaventure Island, where the birds were not protected by law, shooting at birds “to stir them up” was not unusual. On Percé Rock the same temptation to “the man with a gun” existed but the people of Percé, he said, were so attached to the birds that no one would ever think of killing one. Napoleon A. Comeau, author of Life and Sport on the North Shore, who had fifty years of practical experience in the North Shore area as a trapper and student of birds, maintained that it was strangers and so-called sportsmen who were respon- sible for killing birds unnecessarily, not the local people. Commander W. Wakeham of the Canadian Department of Marine, who had witnessed the destruction of shore birds and sea birds on their nest- ing grounds in the offshore islands, suggested that certain groups of islands, or even certain sections of the north coast, should be set aside as bird sanctuar- ies. He also reported on the stupidity of setting aside part of the interior of the Gaspé peninsula as a park without providing any form of protection for the birds and mammals there. Wood wrote sections of the report on laws, leasehold and sanctuaries to place the issues in perspective and set out recommenda- tions.2° Whether Taverner received a copy is not known. However, from early 1914 Taverner was involved at an official level in making investigations at Percé and Bonaventure into the effects on salmon fry of predation by cormorants and gannets.2! From this time until 1919 Taverner was in correspondence with James White on the subject of bird protection. In the fall of 1914 Taverner had a talk with Harkin about the idea of establishing a national park at Point Pelee and sent him a memorandum on the subject in December.22 At the sixth annual meeting of the Commission of Conservation in January 1915 Dr. John M. Clarke spoke about “Protection of the Sea Fowl of the Gulf of St. Lawrence”, Gordon Hewitt, Dominion Entomologist, gave a short talk on “The Protection of Birds”, while Taverner presented a memorandum on the “state of wildlife at Point Pelee, and at Percé Rock and Bonaventure Island with rec- ommendations that these three be included in the National Park system of the Dominion.” In his memorandum Taverner first described the natural aspects of Point Pelee followed by the econom- ic aspects and ownership. Lastly he discussed the wildlife aspects of the marshes where, he stated, “‘the place has been over-shot and under-protected and nothing done to attract game”. He recommended that the marshes be established as a bird sanctuary, and that the few privately owned lands on the point be acquired 92 THE CANADIAN FIELD-NATURALIST and the whole area be made a national park and bird reservation. He then described Percé Rock through the eyes of a naturalist-photographer with the innumerable gulls and cormorants on it during the breeding season. He ended with a plea for its preservation: “Such a national monument should be enrolled immedi- ately among the nationally owned wonders of the Dominion, for then, and not until then, can we be certain that its beauty will not be sacrificed to private greed, and that it will remain to be a joy and an inspiration to the people of the Dominion forever.” Finally he described the estimated 7500 Northern Gannets nesting on the rocky ledges of Bonaventure, and recommended that Percé Rock and the Birds’ ledges of Bonaventure Island be included in the national park system, and that a resident of Bonaventure Island be appointed warden at a nomi- nal salary to see that the park regulations were enforced.23 Taverner had now stated the case for pre- serving these three outstanding bird habitats. From this action various developments followed. Early in February 1915 Taverner had a long talk with Harkin, Commissioner, National Parks Branch, who told him that the Minister of the Interior had approved of his suggestion in principle. But since there was no money at that time for buying out the settlers at Point Pelee it would have to be deferred till a more favourable time.?* The next task for Taverner, Saunders and others, who wanted to make sure that Point Pelee and Percé-Bonaventure were preserved, was to bring the issue to the public through the use of films, slides and public talks. That was why Taverner, with two assistants, returned to the Gaspé in 1915. Late in that year Saunders was in Ottawa speaking before the Commission of Conservation on bird pro- tection. In a letter to Fleming, Taverner described Saunders as “a delightful speaker. He has a homely confidential man- ner that is most attractive and convincing. He made a hit. We are bringing him back in March to show our moving pictures again to the Forum and the Ottawa Naturalists. The Bonaventure Island pictures were shown the other day and I was delighted with them. They have aroused a good deal of favorable comment.’”’25 Taverner also made known his concern for bird protection on the north shore of the Gulf in his “Summary Report for 1915” which was a handier source of reference for a wider public than the Commission of Conservation report. He referred to the research of Dr. Charles Townsend, who, with a botanist, cruised along the north shore coast of the Gulf in a fishing schooner for several months in 1915. In his own report Townsend emphasized the urgent need for effective bird protection on that coast before it was too late. “Bird reservations or refuges watched over by guardians will alone save the remnants from extinction” .26 Taverner commented that the alarming conditions described by Townsend, which threatened the exis- Vol. 110 tence of the eider ducks and larger birds, were by no means exaggerated. Taverner therefore urged that steps be taken immediately to conserve these valu- able species on the north shore of the Gulf and on the Labrador coast. “It is more than an aesthetic or academic question in this region. It is more than a matter of game supply for the sportsmen. The sea fowl on the Labrador coast are almost a necessity of life to the inhabitants and furnish practically all the animal food available.”27 Taverner did not rely on written reports only but was regularly in touch by letter with James White at the Commission of Conservation. Writing from Percé in June 1915 he described to White the condi- tions on Percé Rock and Bonaventure Island, and along the north shore of the Gulf of St. Lawrence where he found the game laws a dead letter. Taverner suggested that all fishermen should have gun licences on board before a fishing licence would be granted, and that all the officers of the Fisheries Service on the coast should be made game wardens. In a letter of early September he sent White a letter and photographs he had received from Charles Townsend about the bird conditions. In the same let- ter he suggested that the Commission should call on Townsend to discuss this subject at the meeting of the Committee on Fisheries, Game and Fur-Bearing Animals in early November 1915.8 Taverner was severely handicapped in not being able to make public speeches nor chair committees. If he had not suffered from a stammer his sense of humour, wit and lack of pomposity might well have made him an effective speaker in the cause of con- servation. Instead he had to put his ideas into writing and leave speaking to others. Apart from W. E. Saunders there was another man who spoke effec- tively in support of conservation, a man who had turned from hunting birds to working for their pro- tection. The story of Jack Miner, and how he started banding ducks and Canada Geese at his family brick- works at Kingsville, Ontario, is too well known to need retelling.2? But popular accounts of his early years as a bander and advocate of private bird sanc- tuaries have failed to place Miner’s work in the con- text of the pioneering work of others in the United States and Canada. As the conservation movement developed it gave momentum to the interchange of information and cooperation among professional nat- ural scientists and amateur field naturalists in both countries. Miner started his interest in conservation through his association with a group of local sports- men. By the beginning of the twentieth century the decrease in the number of game bird species in east- ern Canada was becoming obvious. Miner’s home was situated in Essex County in southernmost Ontario near the north shore of Lake Erie between Windsor and Leamington. In 1904 this group of sportsmen organized the South Essex County Wildlife Association in an effort to prevent the Bt996 decline in the number of game birds. While Taverner, Saunders and other members of the Great Lakes Ornithological Club were compiling data at Point Pelee from 1905 onwards Miner started experi- ments in banding ducks in a pond on his family brickworks. His first bands may have been made by Taverner and given him by Saunders.3° Saunders and Miner were likely to have been in touch with each other by this time.3! Saunders first visited Point Pelee in 1882, and in the same year produced a “List of the Birds of Western Ontario”.32 The following | year he was elected a member of the AOU at its first regular meeting. By 1900 he was in touch with sportsmen and naturalists in a wide area radiating from his home in London, making friends, and pass- ‘ing on information and new ideas. Saunders and | Taverner visited the Miner home in 1909 or 1910.3 Miner put his first band on a bird in December 1909 ) when he tagged an American Black Duck which was shot in South Carolina on 14 January 1910. Miner’s major contribution to the conservation movement in eastern Canada at this time (1910-1920) _was to apply the principle of bird banding to water- fowl, in particular to Canada Geese. He was the right man in the right place at the right time because his family owned land on which there already existed two | ponds suitable for attracting and banding on a large scale. These ponds were also close to Lake Erie so ‘that the geese could fly from the lake where they _spent the night, to the Miners’ ponds, where they were fed. Miner’s experience as a hunter helped him to suc- ceed in attracting them after earlier setbacks. When he found that local hunters waited for the geese to fly to his ponds in order to shoot them, he got the family property declared a wildbird sanctuary. From 1915 he was banding Canada Geese on a large scale and his success was assured provided that he could manage to feed an ever increasing number of geese with corn. In order to raise money to buy more feed be began to give talks to groups and organizations about his work, /and the need to set up more sanctuaries for protecting _wild birds of various species, not only geese. In 1917 the government of Ontario declared the Miner farm a provincial Crown game reserve which gave it status and some financial support.24 _ Throughout the period 1915-1920 Taverner made certain that White was supplied with up to date information on the problem of bird conservation in ‘the Gulf of St. Lawrence, sending him copies of let- ters and articles by Townsend on the subject. In 1916 Taverner sent White a manuscript copy of a chapter ‘on “Bird Conservation in Labrador’ from a forth- |coming book by Townsend, called In Audubon’s | Labrador.*> In asking Townsend’s permission to jhave it published in the next Report of the Commission of Conservation he explained that _ “the Commission is a live organization and get[s] things accomplished in a manner that few others can touch. They get their stuff in most presentable shape and CRANMER-BYNG: A LIFE WITH BIRDS 93 though we regard them more or less as thieves giving lit- tle credit and stealing whatever they want in their work they certainly get results.”’36 The Canadian Commission of Conservation played a major role in bringing authorities in Canada and the United States together from 1913 onwards to discuss the possibility of the two countries signing a treaty for the protection of migratory birds, many of which were threatened with extinction. When William Haskell, legal counsel for the American Game Association, came to Ottawa in January 1914 to speak on the wildlife conservation movement in the United States he included in his talk a brief out- line of the legislative struggle for bird protection in the U.S. to that date. He emphasized the importance of the benefits that Canadians would obtain through such protection. The Commission asked the provinces to call on the Dominion government to get Great Britain and the United States to negotiate a convention for a treaty. Shortly after this the United States government sent a draft treaty for the protec- tion of migratory birds to the Canadian government for consideration. Copies of the draft treaty were ‘passed to the provinces, and all but British Columbia and Nova Scotia agreed to the draft terms. By the spring of 1915 an order-in-council was passed agree- ing to the principle of international protection for migratory birds. It took another year of compromise between the Canadian and American negotiators before the objections of provinces and states were sufficiently met. On the Canadian side it required all the energy and skill of Hewitt, Harkin and Graham in negotiation to reach a revised order-in-council that the Canadian government could accept. A Treaty for International Protection of Migratory Birds was signed in Washington by the British Ambassador on behalf of Canada and by the United States Secretary of State.” A full year was taken up by the ratification of the treaty by Canada and the United States. When members of Parliament were assured that the provin- cial governments were in full agreement, the bill passed the House of Commons and the Senate, and received royal assent on 29 August 1917. Owing to problems related to the U.S.’s entry into the First World War the bill was not passed by Congress and the Senate until June 1918 and finally became law when signed by President Woodrow Wilson on 3 July 1918.38 Protection was afforded to migratory game birds, insectivorous birds and migratory non-game birds. But raptors and crows were omitted from the act. This was an accepted thing at that time since many country people regarded such birds as vermin. But the non-protected status of hawks and owls was to cause Taverner a great deal of trouble later. Although Taverner was not directly involved in the negotiations of the treaty he was involved in a movement to educate the public in the value of birds, and the importance of protecting them through regu- lations that could be made enforceable in the 94 provinces. Writing to Swales in November 1915 he mentioned that he was trying to influence the Conservation Commission in some protection mea- sures.3? One way of arousing public interest was by showing slides and films of wildlife and habitats that carried a conservation theme. In a letter to the secre- tary of the AOU he said the museum had two films of the Gannets of Bonaventure taken by Taverner, and one film of the wild geese at Miner’s sanctuary. Taverner added that Saunders was willing to speak about them both at the coming AOU meeting at Portland, if they could be included in the program.*° The extreme slowness in putting the protection of birds into action in North America can be attributed to the trauma of the final year of the First World War. With the signing of the armistice in November 1918 the Canadian government, at the federal and provincial levels, began to take action. Early in 1918 Taverner was being consulted by Hewitt on bird pro- tection matters and the setting up of new federal reg- ulations in accordance with the treaty. In a letter to © Fleming, Taverner enclosed a preliminary copy of what was being discussed and explained: “The main idea is to reduce the open season on every- thing to 3 2 months as the treaty calls for, to keep the seasons similar as possible on all similar classes of birds and to depart from present provincial seasons as little as possible. British Columbia is the only province that did not come into the agreement and maintained its indepen- dence of action in matters of detail. It agreed only to the principle, consequently the many B.C. exceptions. In discussing it, which we have only done partly as yet, a few minor changes have been made, more clearly indi- cating the tidal regions of P.Q. and allowing Indians to kill certain species for human food. What is the worst puzzle is the regulations thrown about collectors’ shipments.” Naturally he told Fleming what was being sug- gested as regards regulations governing shipments of birds sent to collectors in Canada from abroad which could not be allowed to enter without a special per- mit.4! He asked Fleming to send his suggestions on what he would use as “scientific advice” on the issu- ing of permits. Taverner attributed the fact that Hewitt was now consulting him on bird protection matters to his new status as a Fellow of the AOU. Whether this was true or not, it throws some light on Percy’s feelings about his own position at the muse- um. An Advisory Board on Wild Life Protection had been created in December 1916 with responsibility for framing policies covering wildlife protection in Canada. It was composed of five senior members of government departments concerned with wildlife: James Harkin, Parks Commissioner; Gordon Hewitt, Dominion Entomologist, Department of Agriculture; James White, Secretary, Canadian Commission of Conservation; Rudolph Anderson, Zoologist, Geo- logical Survey; Duncan Campbell Scott, Super- intendent-General of Indian Affairs.42 There were strong reasons why Taverner should be included on the Advisory Board and several reasons why THE CANADIAN FIELD-NATURALIST Vol. 110 Anderson should, especially since much of the advi- sory work was concerned with the protection of mammals, and a knowledge of wildlife and human conditions in the Mackenzie Delta of the Northwest Territories was especially needed. There was con- cern over the decline in numbers of Muskox and bar- ren ground caribou and the effects of this decline on the livelihood of the eskimos.*? Taverner would have grounds to feel upset if Hewitt continued not to consult him on the conserva- tion of birds. However, by the end of 1917 he had begun writing to Taverner for information. His first question concerned the state of the duck population in British Columbia. Taverner answered it as best he could but had little firm information. He did, howev- er, explain the relation of the cycle of the rabbit pop- ulation to the decline in the game bird population, especially in the grouse species. When the rabbit numbers declined, he said, the large owls and [Northern] Goshawks depleted the Ruffed Grouse, Sharp-tailed Grouse and [Greater] Prairie-Chicken.“4 Hewitt also asked for Taverner’s advice on a wider subject. By this time the Advisory Board on Wildlife Protection had been able to agree to dates for open seasons on wild fowl in all provinces except British Columbia. Hewitt sent Taverner the dates suggested through discussion with Francis Kermode, curator of the British Columbia Provincial Museum, for open seasons in northern and southern B.C., and asked Taverner for his comments in the light of any records that he had on migration and other subjects. This gave Taverner opportunity to inform Hewitt of the research he was carrying out, though he regretted how little accurate information he had on the move- ments of birds in the far west of Canada. He also dis- cussed the question of whether the killing of large raptorial birds should be encouraged in order to pro- tect game birds and poultry. This was a complicated | problem in Taverner’s opinion. Based on his experi- ence in the prairies during last summer (1917), Taverner said, he was amazed at the number of gophers that Red-tailed, Rough-legged [probably Ferriginous, as Rough-legged Hawks are not in the prairies in summer] and Swainson’s hawks killed. Bearing in mind the great damage that gophers do to grain crops great care should be taken not to awaken — any more animosity than already exists against these _ raptors that prey on them. The gophers go into hiber- nation early in the fall and only then do the hawks | begin to take game and poultry. Even then, the | buteos migrate from the prairies during the winter, while the [Northern] Goshawk and the Great Horned Owl tend to move south into more settled areas dur- | ing the winter months. These, in Taverner’s opinion, were the only raptors that should be considered as doing serious harm to poultry and game, and he offered this advice: “The only safe proceeding is to educate the farmer to discriminate between helpful and harmful species. The | | / | 1996 fact of the matter is that it is as much a farmer’s bounden _ duty to learn his bird friends from foes and to know the ' weeds from his crops.” He would like very much, he said, to write a paper ‘on these large birds of prey and accompany it with descriptions and pictures by which the raptors could be easily differentiated.+ | Writing to Fleming at this time he gave his opin- lion of Hewitt, saying | “Hewitt is in many ways a good man. He has great influ- | ence and energy and now that he has shown some ten- _ dency to seek advice from proper quarters he can be of | great value. It is good that there is someone of his cali- __ bre to push things.”46 | Brooks, back from the war, wrote to Taverner in ‘September 1918 about the permit system for the young bird collector, and said there would be no young field naturalists in Canada in the next genera- ton. ' “As Dr. Merriam puts it the Universities are producing a ' school of section-slicing and tissue scratching laboratory | workers”. | Taverner replied that he was fighting for the col- ‘lector all the time, and described how permits were jezauted and the regulations that existed for those receiving them.*7 | The next move forward in wildlife protection ‘came in December 1918 when Hoyes Lloyd was jappointed “Ornithologist’” with the Parks Branch to ‘administer the Migratory Bird regulations under the ‘Convention Act (on his appointment see Chapter 7, page 76). Taverner was much encouraged by this ‘development, as he told several of his ornithological friends. When writing to Swales he mentioned the hard work he had been doing to turn the Ottawa Naturalist into a presentable publication which ‘would fill a long felt want. He continued: “We have a new ornithologist in Ottawa now, Hoyes _ Lloyd, probably you have seen his name in the Auk. He | 1s a protege of Fleming. He has come to the Dominion _ Park Branch to have charge of the enforcement of the _ Migratory Bird Convention. I guess it is about the first ' time an ornithologist has been made Game warden. I think he will be a good man too. His ornithological experience is not very great but he has the proper spirit and understands the ornithologist’s viewpoint. Behind him is a Wild Life Protection Advisory Board composed _ of two good zoologists, Anderson and Hewitt, a repre- sentative of the Indian Affairs who is amenable to reason | and the Commissioner of Dominion Parks [Harkin] who | has direct charge of the Act’s enforcement. He is broad- i] | er than most bureaucrats and has at last realized that we bird men have not got private axes to grind. All told I think it looks good. It is also good to have some one to talk bird[s] with once in a while.’’48 _ It was ironic that Taverner, who was appointed mainly on his experience as an ornithologist, should not have been given the title of Dominion ‘Ornithologist officially, but that Hoyes Lloyd, who was primarily not an ornithologist, should be called CRANMER-BYNG: A LIFE WITH BIRDS 95 “Dominion Ornithologist’”. At first his responsibili- ties were to ensure that the provisions of the Migratory Birds Act were put into effect. After a year his title was changed to Supervisor of Wild Life Protection in Canada. At this time (1918) the con- cept of ornithology was not well understood by many people. Academically ornithology meant a branch of zoology studied by a specialist on birds. However, no formal courses in ornithology were offered in Canadian universities, and only a few in the United States. Although provincial governments gave official recognition to the field of ornithology, the federal civil service did not have even an accept- ed definition of what an ornithologist was.4? Taverner was pleased with the appointment though he gave the government only faint praise for it. In a letter to Bishop he wrote: “I think Canada has made a record in appointing a bird man in sympathy with the collector and scientific proce- dures as sort of federal Game Warden. Our new man who will have the details of enforcing the Migratory Bird Convention seems to be an admirable appointment. Sometimes the government does the right thing by mis- take. Hope the promise of intelligence develops.”>9 Lloyd appointed three experienced ornithologists as Federal migratory bird officers: Robie W. Tufts in 1919 to be responsible for the Maritime Provinces; Harrison F. Lewis in 1920 for Quebec and Ontario; and James A. Munro for the four western provinces, also in 1920. Wardens were appointed to work under them. The Migratory Birds Convention Act gave a mea- sure of protection to birds during the spring and fall migrations but more specific protection was required by certain species during the breeding season. Migrating wildfowl such as geese and ducks, birds that nested in colonies, such as herons in trees and seabirds on rocky islands and cliffs all need special protection from human predation and disturbance. The idea of setting aside specific areas as wildlife sanctuaries, or reserves, where birds would learn to feel safe from attack by hunters and egg collectors was not new. The first bird sanctuary in North America had been created in 1887 in the Northwest Territories of Canada (present-day Saskatchewan) at Last Mountain Lake, fifty miles northwest of Regina. The islands and eleven miles of shoreline were with- drawn from settlement and set apart as a breeding ground for waterfowl. In the United States bird sanc- tuaries were being set up from 1900 onwards. In Canada Jack Miner’s experience in banding wild geese at his family brickfields near Leamington led him to seek provincial government support in declar- ing the area a sanctuary to migrating duck and geese.>! As regards Point Pelee Taverner had official- ly recommended setting it aside as a Dominion Park in January 1915.5? It took considerable pressure from public opinion in favour of making it a reserve before this was finally achieved. Taverner, as a civil servant, 96 THE CANADIAN FIELD-NATURALIST was in a position to influence the Conservation Commission and other civil servants, but he had to be careful what he said and wrote in public. Also his speech impediment prevented him from talking in public. Meanwhile the Essex County Wild Life Conservation Association, which Miner and other sportsmen in the Point Pelee area had founded some years previously, was active in prodding government into action. After holding public meetings and writing letters to the appropriate authorities, the Association could rightly claim that its members had persuaded sportsmen of Essex county and the local press into supporting the idea of Point Pelee as a Dominion Park and therefore a bird reserve.*3 By the beginning of 1918 Hewitt and Taverner were in correspondence, and early in May Hewitt invited Taverner to attend a meeting of the Advisory Board on Wildlife Protection in his office. The reason, Hewitt explained, was that a proposal to turn Point Pelee into a wildlife sanctuary was to be considered shortly and that, since Taverner was well acquainted with the conditions there, the Board would be glad if he could attend the meeting to give it his experience.*4 Point Pelee was proclaimed a National Park by the Governor-General the Duke of Devonshire on 29 May 1918. From this time onwards there was a regular and friendly correspondence between Taverner and Hewitt. For example, Hewitt asked Taverner to lend him some slides on Bonaventure Island for a lecture he was to give in Quebec, and thanked him warmly for the loan. In December 1918 Hewitt asked Taverner to prepare a short statement for the Advisory Board on the importance of Bonaventure Island and the Bird Rocks in the Gulf of St. Lawrence as bird sanctuaries, especially for the preservation of gannets. Hewitt explained: “T feel that you may speak with greater authority than I on account of your personal acquaintance with the islands”.>> During 1919 they were in touch over such subjects as the preparation of a French checklist of Canadian birds, and the need for officially established bird names for a French translation of the Migratory Birds Convention Act. In another letter Hewitt thanked Taverner for detailed notes and suggestions that he had given for a chapter on game birds and larger non-game birds for inclusion in Hewitt’s report on Wildlife Conservation. Taverner’s notes covered: distribution, extent to which more or less common, and comments on protection.*® Their corre- spondence ended in February 1920 with the sudden and unexpected death of Gordon Hewitt.5’7 His book The Conservation of the Wild Life of Canada was published posthumously and probably contained some of the information which Taverner gave him on request. When Point Pelee was set aside as a National Park the practice of fall duck hunting was allowed to con- tinue on the Pelee marshes in spite of their being Vol. 110 inside a game sanctuary. This caused ‘an outcry among conservation supporters, outstanding among whom was W.E. Saunders.°* His publicity against continued fall hunting drew a nasty letter from Edward R. Kerr, the Association’s secretary, who wrote that he had been keeping a close eye on the situation at Point Pelee and noted that Saunders was particularly interested in a man named Bert Girardin who had been charged before F. H. Conover, the newly appointed park superintendent, with setting traps for muskrats and Mallard ducks. “On this occasion I presume he was only collecting specimens for your good self, at whose authority I am at a loss to know. If you enjoy special privileges on this - Park it seems to me that you would not be within your rights in employing others to kill and collect specimens of birds or animal life. The question of taking of speci- mens of any kind on Dominion Parks and Bird Sanctuaries will be taken up by me with the government as it is a most unusual thing to permit and should be stopped at once.” This was the opening paragraph of an attack and was a sharp blast below the belt. Kerr followed this up with a second barrel of the same shot. “I note particularly that you are greatly enthused over and interested in the salvation of wild ducks visiting the marshes adjoining this landmark periodically and that it is your purpose to propose to the government that duck shooting be prohibited. So far as I am personally con- cerned it would make little difference to me but I have to bear in mind that upwards of one hundred and fifty good citizens of Ontario enjoy recreation with a gun and I pre- fer that they be permitted to take a fair toll of migratory water-fowl during a limited and favorable period of the year in preference to expending Canadian money to culture water-fowl to feed the stomachs of negroes and other gunners of the southern United States, Mexico and Central America. To shut off duck-shooting as an experiment would be silly. We need no experiment; we are not without experience as we have maintained in Ontario for thirty years a closed season during the spring of the year and water-fowl have decreased enormously and continuously. Of course you are entitled to your opinions and are at liberty to make any recommenda- tions you deem advisable but I need not familiarize you with the fact that your views will be strenuously opposed. Personally I honor and respect you as an ornithologist but I cannot under any circumstance honor and respect the attitude you assume respecting what the people of Essex County alone have a right to discuss. I have already placed my views before the government.>? How Saunders answered this letter is not known, but he obviously did because a week later Kerr wrote another letter to him. This was about government permitting sportsmen to continue to shoot duck on the marshes when Point Pelee was set aside as a Dominion Park and Bird Sanctuary. He said that since 1911 the Essex County Wildlife Association had succeeded in keeping a closed season on quail with the result that the quail population was on the way to recovery. During the past seven years the Association had built up a friendly public feeling on 1996 _behalf of the Advisory Board on Wild Life Protection of which Dr. Hewitt was Secretary. Kerr then explained why members of the Association were allowed to shoot duck in the fall. “At the time Point Pelee was set aside the question of inclusion of the marshes within the boundaries of the Sanctuary was discussed and the Honourable F. G. Macdiarmid, a universally recognized friend of the sportsmen, thought it unwise to deprive the sportsmen of limited duck-shooting and other recreation on the marsh- es. This matter hung fire until the association consented to the inclusion of the marshes with permission to take a small toll between October Ist and December 15th, four days per week. There was no further discussion and the suggestion was adopted.” Kerr ended his letter by telling Saunders: “T think you should have taken up with us the question of duck-shooting previous to recommending to Mr. Conover that it be abolished as no doubt the members would have liked to express themselves previous to any change.”©0 Although national parks are usually regarded as wildlife sanctuaries, in the case of Point Pelee in 1918, by special privilege, a fall duck hunt was per- mitted, and no government since has ever seen fit to stop it until 1989.¢! Taverner was lucky to be able to leave the trouble over Point Pelee to others. However, he was still active over the future of Percé and Bonaventure islands. In 1918 he completed an important study of “The Gannets of Bonaventure” which appeared in The Ottawa Naturalist, and as a result offprints were available for wider distribution. It was written in a clear style which brought alive the scenes on Bonaventure Island described by Taverner in his let- ters written during his visits there in 1914 and 1915. It was also a strong plea for the protection of those powerful and fine-looking birds, the gannets, which summarized accurately their main aspects. Although from a political point of view all was ready for estab- lishing bird sanctuaries there and on Percé in 1917, yet by May 1918 there were still problems in secur- ing titles to land on the island as Taverner explained: “There is a movement under way by the Conservation Commission to reserve this wonderful spot as a perpetu- al bird reserve under the control of Dominion or Provincial authorities, but such is the conservativeness, to call it by its mildest name, of the local population that considerable objections have had to be overcome and it is still doubtful after three years of effort, whether the plan will succeed or not. Some day the local population will realize that these rookeries are a source of attraction to strangers and too valuable a local asset to be wantonly destroyed. Until some such light breaks upon the com- munity, and awakens public opinion and a spirit of pro- tection, the senseless destruction will proceed. It is to be hoped either that the protective measures will be com- pleted or this awaking will come before it is too late.”’? Taverner also sent Hewitt a report on the urgent need to reserve Bonaventure. He stressed that the Gaspé region was becoming a tourist attraction, and CRANMER-ByYNG: A LIFE WITH BIRDS 97 that more revenue could be expected from it once it became a national reserve.®? In November 1918 Harkin recommended that Bonaventure cliffs and Percé Rock be declared sanctuaries under the Migratory Birds Convention Act. In March 1919 the Quebec legislature passed a bill designating these areas as provincial bird sanctuaries, and a few days later they were declared federal bird sanctuaries by the Dominion Government. Taverner had recently become active on the edito- rial committee of the Ottawa Field Naturalists’ Club’s journal The Ottawa Naturalist. It had meant a lot of hard work, and some lobbying before the poli- cy of the journal could be changed, and the journal issued under a new title with a wider appeal. (For the part played by Taverner in bringing about this improved journal, appealing to a wider readership, see Chapter 10.) He used his position on the editorial committee to publish articles, notes and letters about the urgent need for wildlife protection. Taverner soon enlisted Hoyes Lloyd as an author and provided him with a useful platform for putting over informa- tion on bird protection in notes on topics such as “Bird Protection and the Law” and “The Ornithological Collector and the Law”.® In return Lloyd enlisted Taverner to write on “Vanished and Vanishing Birds” in order to sound an alarm to any- one not yet aware of the extreme urgency of the situ- ation.® In a small pamphlet Taverner gave examples of birds formerly known but by 1918 extinct — Passenger Pigeon, Great Auk (formerly on rocky islands in the Atlantic), Labrador Duck and the Eskimo Curlew which, Taverner wrote, was once so numerous that they were supplied to markets by the barrelfull but which by 1918 was in all probability extinct. “These are cases where the harm is already done. No regret, skilful law framing, or law enforce- ment will ever bring them back ... ’©° To the present day reader Taverner’s remarks on the extreme scarcity of several other birds in 1918 may come as a surprise. The Hudsonian Godwit was once so com- mon that considerable bags [kills] could be made but now, Taverner said, it was so rare that scientific col- lectors were searching for stuffed specimens from old collections and out of the way places, while occurrences of living birds were important enough to warrant recording. Likewise the Wood Duck was progressing towards extinction. Only a generation ago, he wrote “it bred on nearly every slack water and overflow of our woodland streams and was the commonest summer duck within its range”. But because it did not retreat to marshes where it would be comparatively safe during the breeding season it was easy game for the amateur sportsman. In the early fall, when the shooting season started, it was numerous on “ducking grounds” but its less wary habits made it an easier target. “Unless care is taken, this, the loveliest ... of all our American ducks, will go the way of the Passenger Pigeon and the Eskimo 98 THE CANADIAN FIELD-NATURALIST Curlew.” The [Common] Eider had also been seri- ously reduced within living memory. The cause, he explained, was because it nested in great numbers on the north shore of the Gulf of St. Lawrence and in the Labrador, where dogs had cleared them from the mainland within reach of even the smallest settle- ment, while on the small adjoining islands they were easy prey to the fishermen. A comparison between the conditions reported by Audubon when he visited the area, and the situation in 1918 showed the grave effects of such wasteful usage. At that rate of decrease, Taverner estimated, it would be only a few decades before eiders would no longer breed on the coast. In addition, he said, nearly all species of game birds had been sadly reduced. The vast flights of waders, the pursuit of which once provided a recog- nized form of sport, were now so reduced as to hard- ly repay the effort of hunting. The same could be said of duck populations. Taverner then turned to the causes of this steep decline in numbers which he blamed mainly on hunt- ing. In recent years most jurisdictions within the provinces and states of America had made their own open seasons. These were short enough for each locality if the birds had to face no other, but as the birds worked their way south in the fall they moved from one local jurisdiction to another with the result that they were in a continuous open season. With the Migratory Birds Convention Act, Taverner explained, federal governments of Canada and the United States could control fall shooting both as regards length of season and numbers of birds allowed to be taken. When the birds returned in the spring, however, they had passed through this process of elimination, and those that survived were the stock on which the summer breeding of each species would depend. Spring shooting, Taverner said, was like killing cattle that have survived the winter and are about to increase, a policy no stock- man would follow, and which should be equally avoided by the sportsman. If shooters in the United States could be persuaded to restrain themselves, we in Canada would have no excuse for not fulfilling our part part of the agreement. He ended his pam- phlet with a strong endorsement: “It is to be hoped that all parts of the Dominion will uphold the authorities in their enforcement of the new game acts. If some are too shortsighted or selfish to deny themselves a little for the general good, it is well to remind them that the laws are now based upon treaty and as such form part of an international obligation which we, at least, do not intend to treat as a scrap of paper and will enforce regardless of the consequences to individuals.”’©7 In his search for contributions to the upgraded The Ottawa Naturalist Taverner wrote to Miner asking him whether he would write a description and histo- ry of his experience banding geese. The magazine, Taverner wrote, was devoted to educational and sci- entific Natural History, and efforts were being made Vol. 110 to appeal to nature students and protectionists. He suggested an article of about 1500 words, with one or two photos.®8 Miner replied on notepaper with the printed heading “John T. Miner & Sons Farmers and Drain Tile Manufacturers”, excusing himself from writing the article. He said that he was trying to fig- ure out how to make his hobby self-sustaining. He was undoubtedly protecting five times as many birds as any gov[ernment] game warden, yet he and his friends had been asking the gov[ernment] for finan- cial help ever since 1910. Last year gov[ernment] gave him $100.- when in fact it cost him over $500.- for feed during March and April. Now he was plan- ning to write a book on the value, migration and intelligence of our birds. He thought that he could sell copies of the book at the end of his lectures. So he hoped Taverner would pardon him for the pre- sent, and if Taverner would come to see his geese he would explain things more fully to him.®? Taverner replied with an encouraging letter sug- gesting that it might be possible to have Miner’s place made into some type of national park under the Parks Branch. He argued that the Parks people fed the bison, elk and deer in their parks and would be able to feed winged game under similar circum- stances. It might be worth considering. Taverner tried again to get Miner to write about the operation of his geese banding ponds. He suggested that a pre- liminary article in The Ottawa Naturalist would act as an advertisement for him, and would assist rather than hinder the publication of a book. He then told Miner of his belief that the conservation movement must reach the children. “The longer I engage in this work the more I am con- vinced that all such movements must begin with the chil- dren. It is almost impossible to teach old dogs new tricks but children are easily influenced and early impressions stick. The children of today are the grown people of ten years hence. The anti liquor movement was regarded as unheard of and absurd to the first generation. To the next. it may have seemed uncalled for but it was not new. The following one thought there was considerable to say for it and the final generation accepted it. Thus it goes in all | movements and none gain great headway except as the © juveniles mature.”7? But Taverner had no success; there is no article « from Miner in The Ottawa Naturalist, nor later n+ The Canadian Field-Naturalist. However, he did‘ have ample opportunity to talk with Miner about! his wild geese sanctuary when Miner came toc Ottawa to attend a National Conference on Game ¢ and Wildlife Conservation held in Ottawa on 18! and 19 February 1919. The Taverners had him to¢ stay at their home. This conference on wildlife conservation was co- hosted by the Advisory Board on Wild Life Protection and the Commission of Conservation. The object was to give members of the federal govern- ment a chance to meet provincial delegates, and for provincial game guardians and conservationists to_ 1996 meet each other and discuss the regulations estab- lished under the Migratory Birds Convention Act, and a number of other wildlife issues. Gordon Hewitt, who led most of the important debates, gave the opening address in which he identified three major points: the need for foresight in wildlife preservation, the need for a national effort in wildlife preservation, and the need for close dominion- | provincial co-operation. Hewitt’s address set just the right tone to inspire the delegates with an under- standing of the significance of what had been achieved already. He then spoke of the need for a nation-wide effort of all those concerned with pro- tecting wildlife to assist provincial governments’ law enforcement. “We shall never again have such an excellent opportunity”, he concluded, “of attaining, by mutual efforts, the ends for which we are individ- ually striving, as we have now”.7! Various papers were given at the conference on various aspects of wildlife preservation, and dis- cussed at some length. The idea of sanctuaries inter- ested many delegates, perhaps because this was something very practical that showed direct results, and was something that citizens could take some part in. Edith Marsh related how the owners of Peasemarsh Farm in Georgian Bay managed to per- suade the Ontario Government to declare their 300 acres of land as a wildlife preserve in 1917,72 the same year in which the Ontario Government had declared the Miner family farm a provincial game reserve. Jack Miner then related his experiences in attracting wildfowl to his sanctuary.’ Although Percy Taverner did not give a talk he certainly attended the conference where he could continue to develop a wider network of useful peo- ple. Taverner had already played an important part in the wildlife protection movement through his bird researches at Pelee and Bonaventure, his official pro- posal for making both places National Parks, the information that he gave Gordon Hewitt, and his paper on Vanished and Vanishing Birds. Through Taverner we catch a glimpse of the conference in action. Writing to Swales he said: “We have just had a most profitable convention here of the game wardens and conservationists of the Dominion discussing the Migratory Bird Treaty. Nelson was here representing the U.S. All is not going absolutely smooth- ly here as Provincial rights are quite similar to state rights and we have a few dissenting provinces. Naturally we do not want to coerce until diplomacy has failed... That historic ‘scrap of paper’ phrase comes in handy now and gives us a catch word that no one can dare question. It is the final argument when all else fails.””74 Elsewhere in the same letter he reported: “Jack Miner was here to the protection meeting and lec- tured on his geese. He certainly is a genius. Though his grammar is a minus quantity his religious interpolations boresome and his humor unsophisticated his enthusiasm, sincerity and lack of self consciousness are such that his audience go crazy over him and every one that hears him CRANMER-ByYNG: A LIFE WITH BIRDS 99 loves him and takes his lesson to heart. He had the crowd actually weeping between bursts of side splitting laughter. He is certainly a wonder and can do more for protection than any other agency I know of. He leaves a trail of enthusiasm behind.”75 Writing to Fleming on the same subject he said: “They had Jack Miner here to speak before the confer- ence on his geese. He certainly made a hit. Had to repeat his lecture in the Normal School and the house was packed and about ten times as enthusiastic as with Hornaday who spoke the night before. They are consid- ering sending him on a grand tour lecturing all over Canada. He can do more for the subject of protection, and the Convention, than any other influence I know.”’76 After the conference Miner wrote a “thank you” letter to Taverner from Union Station, Toronto, on his way home, as follows: “My dear Percee Many, many thanks for a rite good time. Please thank mother for me. I was well pleased to think I stopped over. Will you send me paper containing report (should there be one). Should I get any more owls in good health I will send you one or two... So Long from Jack Miner”?? To which Taverner replied that he was sorry there was no notice of his lecture in the next day’s paper but all the reporters were busy with Laurier’s funer- al. Taverner guessed they had covered Miner’s talk in the paper of the previous day but added: “At any rate you caught the crowd in great shape. The next day everybody was talking of it and Ottawa was divided into two classes, those who were there and those who wished they had been.”78 The reference to the owls “in good health” has a rather sad significance. By this time Miner had begun to defend the birds in his sanctuary from predators, mainly hawks and owls, by a programme of pole trapping. While he was visiting Ottawa, Taverner asked him if he would be willing to send the National Museum a few of the owls and hawks which were in reasonably good condition. They could be sent C.O.D. Since the museum still needed specimens to complete its sets of skins Taverner was glad to have some, however much he disliked the idea of owls and hawks hanging from a pole with one leg almost severed. Miner had no such feelings. He claimed that it was the will of God that any bird or mammal that killed song or game birds should be exterminated. God’s will conveniently supported Jack Miner’s will. Many naturalists were disgusted by the callous pain inflicted on owls and hawks by Miner through the use of pole traps. This became a hot issue that smouldered throughout the 1920s and 1930s, and involved Taverner in his position as Dominion Ornithologist.’ Taverner’s contribution to the struggle for wildlife conservation in the years covered by this chapter had important results for birds on Point Pelee, on Percé, and Bonaventure Islands.8° This alone was an out- standing achievement, but in addition he influenced the awareness of the general public about birds and 100 the need for protecting them and conserving their habitats. This he did by his numerous articles, and by his films, photographs and slides. His greatest influ- ence on the public at this time was through his first book, Birds of Eastern Canada. This is the subject of the next chapter. One further achievement should be mentioned. His warm co-operation with Charles Townsend led to Townsend’s survey of the summer birds of the Gaspé Peninsula which had not previously been CHAPTER 10. Birds of Eastern Canada “IT hope one of these days we may be able to issue a Canadian book with illustrations by you and make it wholly Canadian as well as wholly good”. P. A. Taverner in a letter to A. Brooks! This chapter is about two of Taverner’s concerns in the period 1912 through 1919. A major concern was to write and publish a book on all the birds normally found in Canada. Another was how to turn The Ottawa Naturalist, a modest, local naturalist periodical, into a more substantial one with Dominion-wide appeal. Taverner had been at the National Museum for less than a year when he began to consider the need for a reliable guide to the birds of Canada to replace the Catalogue of Canadian Birds by John and James Macoun. He knew very well how frustrating it was to be a keen ornithologist yet lack a field guide that was scientifically reliable, and illustrated in colour. What, then, were the books that existed in 1912 for naturalists in Canada who were seriously interested in the study of birds? If Taverner had been asked this question he would have mentioned the following: John and James M. Macoun Catalogue of Canadian Birds revised edition (Ottawa 1909).? Ernest E. Thompson [Seton] “The Birds of Manitoba” Proceedings of the United States National Museum 13 (Washington 1891) pages 457-643. Thomas Mcllwraith The Birds of Ontario 2nd edition (Toronto 1894). C. E. Dionne Les Oiseaux dela Province de Québec (Québec 1906). In a wider context Taverner would have referred the enquirer to Elliot Coues Key to the Birds of North America of which he wrote: “This is perhaps the most generally accepted authority upon American birds. It is primarily intended for the advanced student but it contains a mass of information that can be found nowhere else and is a final court of appeal to the majority of ornithologists”.* Robert Ridgway The Birds of North and Middle America U.S. National Museum Bulletins (Washington 1901—). Seven volumes were then in print. Frank M. Chapman Handbook of the Birds of Eastern North America (New York 1895). First revised edition 1911. Chester Reed Birds of Eastern North America Part I Water and Game Birds east of THE CANADIAN FIELD-NATURALIST Vol. 110 studied. In his article Townsend acknowledged the use he made of Taverner’s notes kept during the summers of 1914 and 1915, and praised the part played by Taverner in determining the Provincial government to make Percé, Bonaventure, and Bird Rock near the Magdalen Islands into reservations. Townsend saw this as a good precedent and he hoped that other reservations would be added else- where, especially along the Labrador coast where they were much needed.*! the Rockies; Part If Land Birds east of the Rockies (1905-1906). The two parts were combined into a single volume (New York 1912). A coloured illus- tration accompanied each species described. These were printed from watercoloured paintings made directly from “perfectly plumaged specimens” which, the author claimed, “faithfully represents each species in a pose commonly assumed in life”. Reed’s illustrated pocket guide was a step forward in the design of bird guides, and probably influ- enced subsequent writers of guides. SANDPIPERS (242) Pisobia minutilia (Vieill.) (Lat., very small}. LEAST. (SANDPIPER “PEEP: Feathers of upper parts edged with bright rusty; breast distinctly streak- ed with dusky. L., 6.00; W., 3.50; Wars; 27038. 51275 Range — Breeds from N, S, and Keewatin northward. Winters from southern U. S. southward. (246) Ereunétes pusillus (Linn.) (Gr., a searcher; Lat., small). SEMIPALMATED SAND- PIPER; PEEP. Slightly grayer than the last; breast indistinctly streaked. Feet with partial webs. Range — Same as preceding. (247) E. madri Cabanis WESTERN SANDPIPER. Bill averaging a trifle longer; Chiefly west of the Rockies. that of the Pectoral and Least Sandpipers. The former is the more abundant during migrations on the Atlantic coast, while the latter is much more common in the interior. Their habits are not different in any way from the more common smaller sandpipers with which they are often associated. The difference in size is quite evident when they are seen together. “Peeps,” by which name both LEAST and SEMIPAL- MATED SANDPIPERS are most often called, are by far the most abundant as well as the smallest species of sand- pipers that we have. Although of different genera, they may well be considered together, for their habits are the same, and in life they are usually associated during nesting, migrations, and in their winter quarters. As we usually see them, the upper parts of the Little Stint are quite rusty, while the back of the Semipalmated specics is inclined to grayish. The breast of the former is distinctly streaked, while that of the latter is very indistinctly so. Selection from Chester Reed’s Bird Guide (1912) vest- pocket edition. | 1996 It was clear to Taverner that the kind of book he _ wanted had not yet been written. As there was no precedent for the style and organization of the book _ he visualized it was clear that he would have to take the first step himself. This happened in 1912 when he realized that writing an illustrated guide to the birds of Canada was beyond the scope of one man. What he needed was the cooperation of someone who knew the birds of western Canada. He also needed a good bird illustrator. In his conversations with Fleming, and in their correspondence, he had often heard of Allan Brooks as a first-rate Canadian ornithologist who was also an accomplished wildlife artist. It seemed to Taverner that Brooks might be the man he was looking for, and in March 1912 Taverner wrote to him introducing himself and men- _ tioning their mutual friend Fleming.* Brooks replied _ that he hoped to meet Taverner in Ottawa in August while “at the rifle matches”.> Taverner followed with a letter outlining his needs which were for informa- tion on, and specimens of, the fauna of Canada. He also wanted to get all the information he could get from “good men” for a new edition of the Macouns’ Catalogue of Canadian Birds. The problem would be that of judging the merits of the contributors. He would want to know them personally, or to have them vouched for by reliable people. He hoped to have Brooks’ help in much of this work, and to talk with him about such matters in Ottawa. He confided that he had a larger ambition. “I hope one of these days we may be able to issue a Canadian book with illustrations by you and make it wholly Canadian as well as wholly good”.® Brooks and Taverner met _ briefly in Ottawa late in August.’ During the winter of 1912 they corresponded a few times, mainly about the possibility of collaborating on a book.*® In 1913 the two men met again in Ottawa where | Brooks was taking part in rifle matches, and they _ spent a day together partly at the Taverners’ home _and partly in the museum. In a letter to Fleming _ Taverner said he was surprised how expertly Brooks _ picked out the subspecies in a tray of birds. “He _ went over the Hermit Thrushes as fast as he could _ finger them and named them as he went, and his determinations tallied in every case with Oberholser’s”.? Taverner was pleased to have started a friendship with an experienced Canadian ornitholo- gist whose opinions on determinations he could trust. The fact that Brooks was becoming recognized in North America as an outstanding illustrator of birds was an additional advantage. The artist’s eye for _ colours and details of plumage was of special impor- _ tance in making correct determinations, and in paint- ing lifelike birds. After Brooks left Ottawa Taverner wrote two let- _ ters about the proposed book. To Fleming he wrote saying that a new Canadian bird book would have to be written in the near future. He realized the problem posed by the lack of specimens and data but, never- CRANMER-BYNG: A LIFE WITH BIRDS 101 oe theless, “... more is available now than ever was before and the time will never come when we can say that we know everything for such a work... I have been thinking deeply over the question and will think more.” Taverner asked if Fleming felt it would be necessary to write new descriptions since these had been written over and over again. Why not, he asked, copy descriptions already in use, where they were adequate, although more plumages of juvenile birds ought to be described. He was planning, he said, to start work in the fall of 1913, when in New York, by looking at the Canadian specimens in the collections of Dwight, and in the American Museum, and then in New Haven, working through Bishop’s collection. Later he would need to visit the collec- tions in Boston, and the US Biological Survey’s col- lection in Washington. The following summer he hoped to look through collections in eastern Canada. He then put forward several ideas for the format and organization of the book which he repeated in a let- ter to Brooks on the same day.!° Taverner started by telling Brooks that the subject of publishing a Canadian bird book had come up much sooner than he had expected. The English edi- tion of the Macouns’ catalogue was now out of print, and since many people had written to the museum for a copy the authorities had consulted him on how to satisfy this demand. Taverner cautioned against reprinting the old list or rewriting it because the Macouns’ list was not what most applicants wanted. Taverner then set down what he considered should be done. What he wanted was a book on the birds of Canada giving a key and descriptions of each species together with information on its distribution and, where appropriate, its behaviour. The distribution of species should be indicated by a small map opposite each one. He told Brooks that ideally he would like to have one illustration to each species. The book should aim to be popular in interest and scientific in accuracy. Such a book would be expensive to pub- lish because the demand for it would be considerable and a large edition would be essential. Since no one man could do justice to such a large undertaking, Taverner proposed that it should be published under the joint authorship of Brooks, Fleming and himself. “If Saunders would come in on it well and good, he can bring an immense amount of life history information into the project but I doubt whether he will come”. He proposed that each author should write the sec- tions that he was best informed about and receive credit from his initials at the end of the paragraph, or whatever he had written. It would take a great deal of preparation because a great deal of work would have to be done looking through several American and all Canadian collections. Taverner estimated the preparation would take two or three years. He ended the letter by asking Brooks to consider the idea and let him have his suggestions. “Also what terms you would require for picture work”.!! 102 A fortnight later, in a letter to Fleming, Taverner refined what he had recently written to Brooks. He said that although the Macoun book had a wide cir- culation it was only of use to the advanced student of birds who wanted detailed information on their dis- tribution. Taverner guessed that there was not more than a dozen men in the Dominion who could use the book to advantage since it did not contain descrip- tions, nor information on behaviour. What was need- ed, he said, was a reference book to fill the long- standing need of the less advanced bird student in Canada. This would be a large undertaking that would require the help of men experienced in ornithological study in Canada. The first problem, he pointed out, would be to devise a plan of work that would satisfy all contributors. He suggested that wherever possible a brief note on “field marks” should be included. Each species should be the focus of the information while subspecies should simply be mentioned by name. Taverner suggested a scheme of how the text could be written so that each contrib- utor received full credit for his work. He added a modest word about himself when he wrote: “Tam afraid that I could bring least to the common fund though naturally I would like to be in on it. I wonder how some such scheme as this would work out. THE BIRDS OF CANADA, by J. H. Fleming, Allan Brooks and W.E. Saunders (if the latter would come in) edited by P. A. Taverner. Illustrations by Allan Brooks”. Finally he asked Fleming what he thought of the scheme, and added “if Saunders cares to come in well and good, if not I guess we can get along without him and it will be his loss. 12 Taverner knew Fleming and Saunders well from experiences shared with them in the Great Lakes Ornithological Club. He knew how cautious Fleming was about committing himself to print. Saunders was the opposite. He spread himself widely over the field of natural history producing a large number of arti- cles and notes on birds, plants, insects and horticul- ture. This would have been enough to inhibit him from starting any major work. Taverner wisely cast himself in the role of editor. In this way he could ensure that the contributions of others would be used to produce the kind of book he felt was wanted by the kind of public he visualized. From Taverner’s viewpoint in 1913 Fleming and Saunders were his mentors in ornithology while he was their protege at the museum. He now offered them the opportunity, together with Brooks, to be the authors of this, the first handbook to be devoted to Canadian birds. Fleming’s answer was typically cautious. He took care to point out all the problems. His letter told a lot about Fleming and his attitude to anything ornitho- logical. By contrast Taverner may have appeared a little too ready to tackle problems head on, and push forward with the job. The question of writing a “birds of Canada” was a serious one, Fleming said, THE CANADIAN FIELD-NATURALIST Vol. 110 because in Canada we did not have a working series of skins, we knew insufficient of where skins were distributed, and we had yet to find out what speci- mens there were in American collections. He then stated his own position as regards collaborating with Taverner. “Personally I think you are competent to do the work except the question of subspecies which I don’t think you are interested in which is where Brooks and myself would come in. Life histories I could not manage at any price. I could do the descriptions but it is a weary task and I would require a good stenographer as well as the material to work on. I do not know if I am justified in imposing such an amount of work on myself”. However, he said, he would be glad to help who- ever wrote it.!> Brooks was more practical in his reply. He said he would do his best to help in any way, and would be willing to come to Ottawa for six months. He suggested painting four species to a plate, without backgrounds, like Fuertes’ illustra- tions in Bird Lore. The price would be about $25.00 per plate.'* During 1914 much of Taverner’s time was taken — up with the expedition to Miscou Island and the | Gaspé Peninsula, as well as with writing up the | results. Museum administration took up the rest of | his time and the book was pushed aside. Brooks | went to England in June 1914 with the Canadian | Rifle Team and was competing at Bisley when on | August 4th the war broke out. Since Brooks already | had an officer’s commission in the Canadian Militia he returned to Canada to the training camp of the | First Canadian Contingent at Valcartier, Quebec.!> | Taverner was able to talk with him before he sailed for England and the First World War. Brooks was to be away from Canada for the next four and a half years. With Brooks involved abroad and a very uncertain future Taverner now turned to young Frank Hennessey to produce coloured illustrations to accompany each species in the projected book. Although there is no indication in his correspon- dence at this time of Taverner’s disappointment at | the loss of Brooks’ collaboration on the book, never- theless he must have felt considerable frustration.!° Apart from Louis Fuertes, the outstanding bird por- traitist of his time, Brooks had begun to make his | mark in North America. Although Taverner was glad | to be able to get Hennessey to paint the illustrations, | he openly admitted that they were inferior. Anyone? comparing Brooks’ bird portraits with those byy Hennessey will understand how Taverner must have | felt. He suffered an unlucky stroke of fate just when} the prelims. of the plates for his book were needed."” Early in 1915 Taverner began work on an outline» for the book. In April he sent Fleming a draft! “prospectus” asking for his suggestions. Using the genus Sterna as an example this is what he proposed — to include in the book: A total of one hundred’ species should be included of which so far eighty- 1996 three had been chosen. He asked for advice on the remaining seventeen species to be selected. He con- sidered including a key to the orders based on line drawings of bills and feet. With the prospectus he sent some illustrations to give an idea of their size and general effect. He hoped to be able to persuade the Government printers to produce a very much bet- ter quality in printing the illustrations than the Government normally managed. On matters of print- ing and illustrating Taverner wrote with some authority due to his training as a draftsman and his experience as a bird illustrator.'* Fleming replied that Taverner’s plan for the “bird prospectus” was excellent. He sent a list of suggestions for Taverner to consider. One was about the type of language to be used. Fleming considered this the key to success, and told Taverner to bear in mind the importance of simple language when writing. “Why not consult a country schoolteacher on this point’, he wrote. He also included a list of about thirty birds that he sug- gested might be included. Under [American] Black Duck he noted that it was much commoner than the Mallard — it was more distinctly an eastern duck. Another suggestion he made was to include coloured plates of birds’ eggs. Since so many boys collected eggs, he explained, why not guide them by indicat- ing proper rules for collectors.!” Taverner first answered Fleming’s points, then restated his main aim, which was to provide a fairly complete coverage of the birds of eastern Canada, of all those that observers would be likely to see. He would prefer not to describe fall and intermediate plumages because the book was designed for an ele- mentary level of knowledge. The inclusion of detailed plumage descriptions would be more likely to discourage, rather than encourage, readers. The novice, he wrote, should learn to identify the adults in spring plumage first; other plumages would be recognized more easily later.”° In a letter to his friend Swales he said that he had just started an interesting job, a popular “Birds of Eastern Canada”, for which there was a definite demand. “The people want it and it will do more to awaken public interest in our Department than any- thing else. Popular interest is what the government scientific departments need in their business”. If the authorities would let him produce it in the way he wanted, he said, it would be “some book”, with 100 coloured illustrations and a key. He had it planned out, and a dummy made which Fleming had approved, and he was now waiting for suggestions from Saunders.”! Taverner was in the Gaspé and the Gulf of the St. Lawrence during June and July 1915 and when he returned was busy with museum matters. However, he was in touch with Fleming about Hennessey’s illustrations, and sent him copies of the first two reproductions made by the printer who would be responsible for the coloured illustrations. Taverner CRANMER-BYNG: A LIFE WITH BIRDS 103 considered them to be quite satisfactory.?* Fleming replied that Hennessey’s drawing of the Bobwhite was far ahead of the Blackburnian Warbler which he considered weak. He wanted Percy to come to Toronto during the time of the Canadian National Exhibition when railway rates would be cheaper. Fleming was quite insistent. “Better come up to get my help in identifying specimens and advice on the colour and makeup of your new bird book. There is much to talk about and to see”.** Taverner was well aware of Hennessey’s immaturity as a bird artist and wrote to Fleming that he was impressing on Hennessey the need for accuracy in drawing bills which was his main weakness at present, though his portrayal of the eyes also needed watching. But Taverner recognized the progress Hennessey had made since he was first hired on a temporary basis by the museum in 1913. As he expressed it to Fleming “one cannot expect a Fuertes or Brooks all at once. This large order, with me at his elbow, I think will be of great benefit to him”.** By mid- September, when Hennessey had to return to college to continue his education, he had completed about 60 of the illustrations and Taverner was hopeful that he would be able to finish the rest before Christmas.”° Meanwhile Taverner was in touch with Saunders about the bird book, sending him a prospectus, and asking for his opinion.”© Saunders replied making a number of suggestions, and Taverner answered them. He added: “I will be very glad of your assis- tance and accept your offer thankfully. We want to make this as good as it is possible to make and your advice will be invaluable”.*’ In October 1915 Taverner managed to meet Saunders in London and see Fleming in Toronto in order to discuss ideas about the book. He then started writing the text, species by species. For Taverner 1916 was the year of the book. He neglected everything that he could in order to keep writing. By early 1916 he had reached the shore- birds. He wrote to Fleming asking if he would be kind enough to read sections, as he wrote them, in order to catch any mistakes. To prepare Fleming for an expanded text he wrote: “It has much outgrown the first intention. But as we are getting the expen- sive pictures I feel that a little extra work in writing and letterpress is excusable” .*® Fleming replied: “I shall be glad to do all I can to help you on the bird book so send it along”.*? Taverner decided to remain in Ottawa during the field season of 1916 in order to devote most of his time to the book. This decision became an obvious one when the Parliament building caught fire in February 1916, and Parliament moved into the National Museum. As a result the exhibition halls were no longer open to the public, and consequently Taverner had less work to distract him from writing. He now worked continuously on the first draft of the book, sending chapters to Fleming and Saunders, at 104 intervals, for their comments. In February he sent Fleming three sections to read. He said this was his second set of corrections, and he felt the text was not far from the final form. He had three main ideas in writing the book, he wrote. The first was to describe each species so that the common man could recog- nize it; the second was to note its economic value; the third to arouse a feeling for the protection of birds.°° With the next section he enclosed a letter to Fleming in which he explained his plan for including subspecies. He proposed to describe only species while recognizing subspecies only “as occasion demands”, and would explain the reason for this in an introduction.*! He expected to be criticized by leading ornithologists for appearing to disregard the importance of subspecies. But he felt that it was nec- essary for him to make a stand at this time, when he was working on a guide book for the ordinary observer, especially since he expected that it would be used by a new wave of observers.*” By now progress was rapid as sections of the man- uscript moved back and forth between Taverner and Fleming. In returning another section of text Fleming found only one fault. He suggested that Taverner should avoid using uncommon words such as “presage, palatable, unimpeachable and differentia- tion”. Simpler words would be understood better.** Taverner sent a forceful reply. Certainly, he said, he wanted to avoid making the book above his readers’ heads, but he did not want to turn it into a primer. It was not advisable to avoid using good English words that any high school student should know. “Tf they do not know them’, he wrote, “there is something wrong with their educational system. The words you mention are words of just this class. If readers do not know them it is time they did. I think modern educational systems are too much on the Kindergarten system any way. Children are too much written down to and are not encouraged to develop. I have been talking this over with Mr. Slemon, the direc- tor of the Nature Study work in the Public Schools and he agreed with me. He has however promised to go over the ms. when I get it in shape and criticise the wording from a school teacher’s standpoint.”*4 When Fleming received the section of the book which included the herons and bitterns he was rather surprised to read that Taverner had included casual stragglers such as Glossy Ibis and American [Great] Egret in a book designed to introduce new observers to the field study of birds in eastern Canada. What Fleming particularly disapproved of was the wording of the entry under American Egret. Taverner had writ- ten a substantial paragraph on the effect of collecting birds’ plumes for sale in the millinery trade. After briefly describing plumage, field marks, nesting, and distribution Taverner condemned the taking of egret plumes in terms that are surprisingly strong for a sci- entific bird guide. Fleming wrote advising him not to get involved with the Audubon “outfit” and to omit THE CANADIAN FIELD-NATURALIST Vol. 110 the egret matter from the book.*> Fleming said he realized that Taverner would find some of his com- ments unpleasant, but he wrote from a genuine wish that Taverner do justice to himself. Nevertheless, he returned to the question of the kind of English he should use, and lectured him once again. In spite of saying that his duties would end when he had checked the correctness of the descriptions and ranges he then told Taverner that his use of English was deteriorat- ing; it showed a “tendency to flacidness”’.°*° This was an unpleasant letter to receive, even when it was from an old friend with the best of intentions. Taverner reacted with a long reply. He had no objection, he said, to the openness of Fleming’s criticism. What he wanted was Fleming’s sincere opinions even if he did not agree with him. Fleming need have no fear of hurting his feelings. “We do not view all things eye to eye and probably it is a good thing we do not. I think that criticism is good for us all even when we do not think it is quite justified”. As regards the Audubon people, Taverner said, he was in full sympathy with many of their objectives, or to put it the other way round they were in sympa- thy with many of his. What he objected to were their methods — their attitude towards scientific collec- tion, their exaggeration, and their sentimental slop. Next he turned to the question of the economic value of birds. Their value, he said, has been greatly exag- gerated. They are probably important as a controlling influence in the reduction of insect pests, but only as one of the influences. Probably parasites should come first with birds second or third. But they are important enough to make their economic value a practical reason for their protection. However, even if a large number of birds were to be killed not every- thing green would be eaten by the bugs. “The real reason for the protection of birds”, he wrote, “is an esthetic one, the pleasure in having them around, and becoming acquainted with them either individually or specifically”. He then returned to the trade in egrets. He had included the American Egret in the book because he felt so strongly on the matter. He wrote: “T think it is our duty to do this hence I lugged in bodily the egret in a Canadian bird book as the most striking example of the iniquity of the trade”. Taverner ended his long letter by stating that his feel- ings were not hurt.?’ By May 1916 Taverner was confident that the museum authorities would publish his book. It was now not a question of whether but of how they would print it. He took the matter seriously, and had three sets of different page specimens made in order to decide on type and the general layout. He sent a copy of each to Fleming asking for his opinion. The main question was where to place the illustrations. Taverner wanted each one to be with its own text and not several pages away from it. He also wanted it to be printed as a pocket handbook but could not get the museum authorities to agree. It was to be 1996 issued in the museum bulletin format and they would not break the size of the set. However, Taverner, whose hobby was bookbinding, said he would try to have it printed with wide margins so that the bulletin format could be heavily trimmed to reduce it to pocket size.** The last two pages of the letter were full of bird information including the news that a Caspian Tern had been sent to him from Quebec. He commented “It is our only specimen and I am glad to eet it”.?? By now Allan Brooks had been in France for two years and had been in action several times. He was promoted Major in November 1915. Taverner wrote to him regularly and his long and newsy bird letters were much appreciated by Brooks. Meanwhile _ Taverner’s sister, Ida, and his secretary, Miss Bentley, sent parcels to the major. In one letter to Taverner Brooks asked him to thank Miss Bentley for the knit- ted socks etc. In May 1916 Taverner wrote that Hennessey had just finished the last of the one hun- dred bird pictures for the book. Although some were not good, on the whole they were very satisfactory, and some were very good. Taverner then discussed the text, and some of the problems he met in writing it.4° In May 1917 Brooks sent Taverner a long letter about birds. Taverner, writing to Fleming, reported that Brooks had been collecting birds just behind the battle line “and is sending some of his specimens over to us. You cannot keep a good man down’”.*! The - Brooks-Taverner correspondence during the four _ years that Brooks was in France (1915-1918) tells lit- tle about the war and much about ornithology. _ Although there were not many letters exchanged those that exist are very interesting in content. They take the form of ornithological discussions between two thoughtful Canadians. For instance in one letter Brooks wrote about the need for a limited use of sub- species because some had different calls from the species, while others had developed anatomical and structural differences. In another letter Brooks wrote on the problem of colour in painting bird plumages. Colours, he wrote, are so largely dependent on the light in which they are seen, and amount of wear on the feathers, that colour values are largely a question of comparison.*” Several quotations from Brooks” let- ters, set in the context of when they were written, are given in H. M. Laing’s biography of Brooks.** The debate between Fleming and Taverner over whether to include subspecies went backwards and forwards through the mail like the volleying of two tenacious tennis players. When Fleming protested _Taverner’s idea of cutting out the three scientific _fames for each subspecies (trinomials) Taverner replied that he would explain the reason in the introduction. The public, he wrote, must have a - common name by which they can speak of species - without having to refer to particular subspecies. He continued CRANMER-BYNG: A LIFE WITH BIRDS 105 “When the public realize that a subspecies is only a fine splitting of little popular interest and can be disregarded except in specially technical work I think a great point will have been gained. I expect to be criticised for this stand but I am sure that I am right.’”*+ Not all was written in a rather harsh tone. Fleming admitted that he would not have made himself so objectionable had he not known that Taverner could improve on the draft he had read. Fleming admitted that no laymen, for whom it was written, would see the objections that he did, least of all the Conservation Commission. He then gave Taverner some lukewarm praise. “You deserve much credit for the work you have done. I never could have stuck it out myself and I only wish you had left innovations to some other time and place”.* Taverner did not accept this weakly, but hit back a few days later about there being room for argument in questions of policy and perhaps principles. “T think that a popular work such as this is just the medi- um to introduce a sane presentation of the subspecies question”. He said he aimed to get “the young Canadian observers started right” so that the importance of subspecies is kept in its proper proportion. “Let the splitter split all he has a mind to. That is his business as long as the other man knows that it is only subspecific and of but technical interest only ...”* Taverner’s mind was quite heated over the ques- tion of species and in October he returned to the issue in a letter to Brooks. He told him that the man- uscript was now finished. It had taken a year’s work and had been quite a load to carry. He was innovat- ing by using a different nomenclature, one that was bound to arouse criticism. In each case the AOU check-list gave the specific name of the type sub- species in the vernacular nomenclature. This, Taverner claimed, was wrong. He continued: “The idea that the species is what is left after the sub- species has been split from it is incorrect. The original specific common name should remain covering all the races while the type subspecies should be regarded as but another form with no superiority of rank over the other subspecies. I have made the common names agree with this conception”.47 The Taverner-Fleming cross-fire spluttered on through September and October, both sides repeating their arguments. In a long letter of early October Fleming made one more attempt to explain his point of view to Taverner, then ended his letter on a con- ciliatory note. “T suppose an exchange of ideas is good and I wish you luck in your controversy. I am glad we can disagree without rancour but I feel it is more a misunderstanding of what each other means”.*8 This is true. Reduced to its essence what Taverner argued was that the species is a definite unit of clas- sification, each species being sharply separated from other species, while subspecies “merge into allied forms at each point of contact’.4? Fleming, however, maintained that a species is not a definite unit, it is a 106 group in which no one member 1s of higher rank than another.>? Although the manuscript of the book was in the hands of the editor, William McInnes, by October 1916, Taverner was still not free to give all his time to other matters that required his attention. Since the beginning of the year Taverner and McInnes had been in correspondence over the content and format of the book without coming to an agreement. McInnes was a geologist of long experience whose last field work had ended in 1910. He was then assigned to scientific editing and administrative work related to the International Geological Congress of 1913. He was the general editor of the report by the Congress on Coal Resources of the World in 3 volumes. He was appointed “Directing Geologist” of the Survey in 1915, and assumed the duty of editing the publications of the Survey such as the Bulletins and Memoirs.°! McInnes had a clear idea of what was required in a publication of the Biological Survey — a plain statement of facts pre- sented in a regular format with no allowance for any innovation, imagination or literary style. McInnes was not used to authors in the Survey having any ideas on the content and style of a book, and would not make any concessions. Taverner, however, had strong ideas of what he considered would best edu- cate the public for whom he was writing. At times Taverner could be stubborn, especially when he was convinced that his ideas and methods were the best in a particular situation. This was such an occasion and he dug his heels in. Predictably McInnes dis- agreed and the verbal hassle continued throughout 1916 and into February 1917 when Taverner, writing to Townsend, mentioned that the book was still occupying a lot of valuable time, and explained why. “T cannot make the authorities above see eye to eye with me as to its style and general tone and it has been a con- tinual struggle to save it from being reduced to [a] Reed pocket book guide without the convenience of the small size of that publication”. He explained that because it was more popular in character than most official publications it was important that the life should not be knocked out of it. “Government red tape”, he added, “is tangled and slow to unravel’”.°? Taverner had worked at the museum long enough by now to know the strangling effects of red tape, but to find it strangling his own creation into which he had put some original ideas was exasperating. He refused to give in and finally Taverner and McInnes referred their disagreement to McInnes’ immediate superior, R.G. McConnell, Deputy Minister, Department of Mines. Taverner gave Fleming this information in February: “T hope I am coming out all right with the bird book. McInnes and I reached a stalemate and referred the matter to McConnell. He has advised with Macoun but is any- way more sympathetic than McInnes. I think that between Macoun and I we can keep the best part of the matter”.>? THE CANADIAN FIELD-NATURALIST Vol. 110 Things were eventually straightened out but this delayed the book for a year because Taverner was ready to leave on his field expedition to the western provinces, postponed in 1916, and therefore could not remain in Ottawa to prepare the manuscript for the printer. The revised version finally went to the printer in August 1918. Page proofs came back in the spring of 1919 and Taverner finished correcting them by early June but was afraid that a good many “bulls” [mistakes] might have been missed. He explained to Fleming that this was because “the system of the editorial office follows the policy that it knows much more than the author and he needs be consulted only occasionally. The editor makes all deter- minations and it is only accident when the author is informed of decisions and usually too late to change it. Nice isn’t it?” Taverner’s ability to write scathing comments was, by now, well developed. Birds of Eastern Canada was published early in November 1919, though dogged with troubles to the end. Writing to Brooks to tell him that a copy was on its way to him Taverner said that there had been a lot of trouble over its distribution and it was almost held up altogether, “but that is nothing for this book”. According to Taverner: “It is only a mere shadow of its former or original text. Remember this when you are disposed to criticise it, you will have lots of opportunity”.°° Before the criticism rained down a letter arrived from Louis Agassiz Fuertes compli- menting him on the book, and Hennessey on the illustrations, especially the bluebird and warblers.°° It was a welcome letter, especially in view of what Taverner told Fuertes in his reply. He said that a government official was advised by “one of the greatest bird authorities on the continent” that the warbler illustrations were inaccurate in the markings on their plumages, and that the whole edition should be withdrawn.>’ The name of this great bird authori- ty was never revealed. Taverner was cheered by a short note from Hewitt who said that such a study had been needed for a long time, and continued: “It will do an immense amount of good and I venture to predict that it will soon be out of print”. Taverner himself was not entirely pleased with the result because he had not been allowed to follow out his own ideas fully. This was all the more frustrating for him because, with his experience as a draftsman, he knew more than most people about printing and binding, photography and types of paper. As an ornithologist not trained in any professional tradition he had a fresh approach which he wanted to carry out in his first book, but which could not be fully incorporated. Also he was not satisfied with the illustrations which he had hoped would have been painted by Allan Brooks. He knew that Hennessey’s were not high grade, but he had to put a good face on the problem, and he accepted them as serviceable. There was no one else in Canada at that time, apart 1996 from Brooks, who could produce coloured illustra- tions showing the distinctive markings of one hun- dred species. Taverner was reasonably satisfied with the brief descriptions of each species he had written, though he was less than satisfied with his short sen- tences on the distribution of each species. This, how- ever, was not something that he could remedy because of insufficient information in Canada. He had incorporated as much information as he could from what local lists he had obtained from men such as W. H. Mousley and R. W. Tufts,°? and from his own distribution maps based on the museum collec- tion. Most of the information came from Taverner’s own observations supplemented by those of Fleming and Saunders and correspondence with many people. Contemporary opinions of the book by authorita- tive ornithologists were reasonably appreciative. The fullest review was by Witmer Stone in The Auk. The first half of this two-page review discussed the book from the point of view of the general reader, and began as follows: “This notable work has been prepared to meet a growing demand for a handbook that will present in concise form the more important information on the habits and distrib- ution of the birds of East Canada and keys and descrip- tions that will enable one to identify them. In providing for all these needs we think that the author has been remarkably successful.” Stone also wrote appreciatively of the illustra- tions. - “The colored plates by Mr. Frank C. Hennessey are very attractive and the postures of the birds usually good, some of them like the Kinglets rather daring in their originality. Mr. Hennessey evidently studies his birds and his paintings are his own interpretation of what he sees rather than copies of conventional attitudes. We need just such effort in ornithological illustration.” However, Stone was critical of Taverner’s attempt to dispense with subspecies in the nomenclature which he called “an unfortunate feature of this work”, because they caused some confusion in the mind of the reader. The systematic index also leads to misunderstandings when trying to carry out the author’s nomenclature.’ A one paragraph review of Birds of Eastern Canada, by Lynds Jones, appeared in The Wilson Bulletin. In his opinion Taverner’s unconventional presentation of the material in the species list seemed worth studying in spite of the inconvenience it caused by not following the 1910 edition of the AOU Check-list. He also said that the coloured plates of F.C. Hennessey “shows that we have another successful artist in the field”.°! W. E. Saunders wrote a critical review of it in The Canadian Field-Naturalist but explained that the author had included more interesting detail in the first draft than appeared in the printed version. Saunders placed the blame for this on “a mistaken editorial policy”. The reason for the elimination of so much, he said, could only have been through poor judgment or economy. This supports Taverner’s CRANMER-BYNG: A LIFE WITH BIRDS 107 statement in a letter to Brooks “it is only a mere shadow of its original text’. However, since this book was intended to be for general readers rather than advanced ornithologists its reception can best be judged by the size of the demand. The first edition was printed in November 1919. A French edition was printed at the same time. Early in 1921 a second edition was being printed, which shows that Taverner’s innovative method, far from confusing beginners, encouraged many people to take up bird study as a recreation. As an example; a schoolboy by the name of George North, living in Hamilton, was given a copy of Birds of Eastern Canada when it was published. As the writer of an “In memoriam” notice for George North put it: “From that point on, birds dominated his life”.° Another man who bought a copy when it was pub- lished, and found it very useful, was Harrison Lewis, then living in Quebec. The demand was so brisk that in 1921 Taverner was asked to produce a companion book, Birds of Western Canada. A modern evaluation of Birds of Eastern Canada in historical perspective can be gained by the fact that, when it was reprinted commercially in 1974, it was accompanied by an appreciative Introduction by W. Earl Godfrey, then Head of Ornithology at the National Museum, a position that Taverner had held until 1942. His words were so perceptive that they are quoted at length here: “A new era in Canadian ornithology began in 1919 with the publication of P.A. Taverner’s ‘Birds of Eastern Canada’. At the time it appeared, there was nothing like the book in this country. Not only did it tell the reader just what bird species he could expect to find, but also how to identify each with minimal difficulty by a short- cut method that concentrated on a few diagnostic fea- tures, or combination of features, diagnostic for each species and called ‘field marks’. Over one hundred species were depicted in natural colors and postures. Data were given on the food habits and other pertinent aspects of the bird, and these furnished the basis for a weighing of the good it does against the bad, and thus people were made aware of the economic status of each bird species. All this (and more) under hard covers sold for just fifty cents! Taverner was in many ways well ahead of his time. He grasped the value of the ‘field marks’ method of identification, not yet generally in use. He applied a ver- nacular name to the species as a whole, rather than treat each subspecies separately under names so different that species relationships were often completely lost. For this, he was rather severely criticized by some reviewers of his book. Taverner’s method, however, gradually caught on and time has shown that he was right. In fact, some 38 years after the appearance of “Birds of Eastern Canada’ the standard authority for nomenclature of North American birds, the American Ornithologists’ Union’s ‘Check-list of North American Birds’ [Fifth Edition, 1957] adopted the practice of furnishing one vernacular name for the species as a whole, and discard- ed vernacular names for subspecies altogether.”°* 108 Taverner explained, in his Introduction, how the book was laid out, and why the classification was different from that used in previous bird books. He also discussed migration, protection of birds, how to attract them, and ornithological books of that time. He explained the use of a key which was accompa- nied by line drawings of parts of a bird’s anatomy by the division’s artist, Claude Johnson. This was need- ed for keying out birds that one had collected. One feature that he had hoped to include was a distribu- tion map for each species. This was thwarted by lack of sufficient information.“ However, the quality of the reproduction of Hennessey’s illustrations in the first printing (1919) was good, and did justice to them. In the second printing (1922) they were rather less clear. A few of the best of Hennessey’s portraits were used in Taverner’s Birds of Western Canada (1926) but had to stand comparison with the majority which were by Brooks. Just a few of Hennessey’s best were included in Birds of Canada (1934). The printing of these, however, showed them as being much paler than the originals of 1919. One can com- pare the style, as bird illustrators, of the two men on pages where one portrait by each is shown. For instance Brooks’ Red-headed Woodpecker with Hennessey’s Pileated Woodpecker (plate 38); Brooks’ Western Kingbird with Hennessey’s Eastern Kingbird (plate 42); Brooks’ Clarke’s Nutcracker with Hennessey’s American Crow (plate 50). While birds painted by Brooks usually look warm with life and about to fly away Hennessey’s were static. They were painted to show the novice the main distin- guishing mark of each species, but they are not likely to fly out of the illustration. Two portraits contradict this criticism: the White-breasted Nuthatch and Brown Creeper (plate 52) do have a living quality. Unfortunately, when the plates were used again for the Birds of Eastern Canada edition in 1974 the result was disastrous as all the illustrations are drained of colour and are fuzzy. They are unfair to Hennessey’s originals. Finally, some of the species descriptions contain a paragraph of general information, drawn from Taverner’s Own experience and observations, that are still worth reading today for the insight he dis- played into the characteristics of those particular species. For instance here is his comment on the Lincoln’s Sparrow: “Though a rare sparrow it is an interesting one. It has reduced hiding in brush to as fine an art as any bird. When first disturbed it hops to a branch, where it obtains a good view, regards the intruder for an instant, and then dives into the tangle and is gone. The most diligent search thereafter gives no more than a fleeting glimpse of a brown shadow disappearing into ile nearest brush- pile’. For another example of Taverner’s comments on a species’ characteristics, here is the American Goldfinch: THE CANADIAN FIELD-NATURALIST Vol. 110 “One of the merriest of summer birds, sometimes remain- ing through the winter in the more southern parts of Canada. It is a great lover of fluffy white thistle and dan- delion seed-heads and can often by seen plucking the down, cutting off the fruiting end, and letting the airy tops float away on the wind. Its song is as pleasant as its bright appearance as it sits on some lone elevation and sings Sweet-sweet-chewit-chewit-chewit or goes speeding off through the air in a merry flock repeating their cheer- ful “Per-chic-o-pee.” The American Goldfinch, though a relative of the Old World bird of the same name, is an entirely different species, named, as the original settlers named many birds, from various fancied or real resem- blances to the familiar forms known at home.’””©° Taverner’s writings about birds, when he gave his feelings and imagination free reign, contained both scientific information, and an awareness of the living presence of the birds described, blending science with poetry. From The Ottawa Naturalist to The Canadian Field- Naturalist “We have a club here, the Ottawa Naturalists, who are much on a par with the Detroit bunch. They have nice picnics every week but they are no place for you and me. The worst of it is that they have had all kinds of direction with good men, Fletcher, Macoun, Gibson and the whole Geological Survey and that of the Experimental Farm but they haven’t evolved a single naturalist in their some twenty five years of existence. The only thing they have got is a publication that has a government grant and appears regularly and in which we can get publication at any time. I * were not for that the real students here would have let the whole organization die a natural death long ago.” P. A. Taverner to Arthur W. Andrews, 29 April 1912.%7 The Ottawa Field-Naturalists’ Club was founded in March 1879. The Club published annual Transactions for eight years based on papers deliv- ered to the Club concerning the natural history of the Ottawa locality. In 1887 the name was changed to the Ottawa Naturalist and it became a monthly publi- cation. Its scope was widened over the years to include papers by members of the Club on the gener- al natural history of Canada.°* Taverner summed up the situation of the Club as he saw it after his first year at the Victoria Memorial Museum. As a pro- moter of natural history study it was rather ineffectu- al, but it issued a journal regularly in which keen Ottawa naturalists could publish their contributions. This state of affairs continued through 1917 with Taverner, J. M. Macoun, G. Hewitt, and a few others active on the editorial committee. They were already thinking how to make The Ottawa Naturalist into a publication that could more effectively represent Canada scientifically. 6° However, early in 1918, there was a “palace revolution”, in which the old style journal was supplanted by a new style journal in an improved format and published in April 1919. | 1996 | 7 _ Fleming the gist of what happened. He wrote: “A few of us got in control and overrode the conserva- tives and when you get your next number you will not recognize it.”””° _ The announcement of the change appeared in the Foreword to The Ottawa Naturalist of April 1918 which explained what was happening, and why. | “With the appearance of this number of THE OTTAWA NATURALIST a new era in the history of The Ottawa Field- Naturalists’ Club is launched. THE OTTAWA NATURALIST __ was established in 1887 as the organ of the Ottawa Field- Naturalists’ Club. It is now in its thirty-second year and is one of the oldest natural history periodicals in North America. While modest in form and not too prepossessing in appearance, it has published many important papers and the great number of original descriptions that have appeared in its pages makes its files a necessity in | libraries of original research in biology. The time has come, however, when a local periodical of this nature is inadequate and the Dominion requires a more creditable and representative publication for the record and dissemination of the results of scientific research. THE OTTAWA NATURALIST, with its already established position, long and honorable history and scientific stand- ing, seems a logical nucleus from which such a publica- tion should be developed. Beginning, therefore, with this number, THE OTTAWA NATURALIST will appear in an improved form. The size | is enlarged and the number of pages increased. The bet- ter paper used throughout will permit of more and finer illustrations; these will improve the appearance and add interest and value to the magazine. To widen its geo- _ graphical sphere of influence a change of name to one of less local significance is being considered, but cannot, by the constitution of the Club, be effected until next wee year. __ The statement then explained that for the journal _ to reach its maximum usefulness the interests of the _ general, educational and technical public would have to be considered. It was hoped that in future teachers in the various schools of Canada would find The | Ottawa Naturalist valuable to them in teaching ele- _ mentary natural science and nature study. Members | of the editorial committee had felt for a long time _ that there was a field in Canada for a journal record- _ing noteworthy biological information as well as _ bringing pleasure and assistance to nature lovers in _ general. The proposed increase in size and improve- _ ment in appearance would place a financial strain on _ the club so an appeal was made to existing members _ to enrol new subscribers. The foreword ended with an appeal to natural history societies through Canada _to make this publication indispensable to every _ working naturalist.”! Taverner, writing to Fleming in August, explained _ their views and objectives. _ “We aim to make it a Dominion wide organ and worth while. We have to combine science with popularity on account of circulation but I do not think that will be a serious drawback. Now we want all our friends to boost . | The exact details are not known but Taverner told | | CRANMER-ByYNG: A LIFE WITH BIRDS 109 it. At the next annual meeting of the club we propose changing the name to one of less local character and appeal. We hope to make it the scientific and nature study periodical of Canada”. Writing to his friend Frank Farley, in Alberta, he discussed a friend of Farley’s who used to claim see- ing rare birds before he was familiar with the com- mon species. Such people were well-meaning enthu- siasts, Taverner said, who were usually working without contact with others of greater knowledge and did not realize their own limitations. He added: “T am in hopes that the new Ottawa Naturalist will assist in this direction [instructing them], including modern systems of zoological work and thought. Glad to hear you approve of it. I would like to get your Alta. Nat. Hist. Soc. interested in it. Perhaps we can when we change the name?”/? At that time he also wrote to Robie Tufts asking if he had seen The Ottawa Naturalist in its new form, and continued, perhaps with a touch of pride: “We have greatly improved its appearance and are endeavoring to make it nation wide in its appeal, We contemplate changing its name to something less sec- tional with the next volume and wish to make it truely representative of Canadian zoology. We think it is time that Canada took her proper place in such matters and that there is room for such a publication. As an educa- tional medium I think it is worthy of all support.””° Tufts answered that he would be glad to do what he could to make the Naturalist a success, promised to subscribe to it, and expected to be able to enrol several more people.” Taverner was busy at this time drumming up sup- port for The Ottawa Naturalist as his letters to friends indicate. Writing to Swales he said: “T have been working hard on the Ottawa Naturalist and I think we are going to make it go. It has meant lots of work though and we are not out of the woods. I think we have turned out a presentable publication and with the enlarged field before us can fill a long felt want.’’”> Taverner’s friends were encouraging in their replies. Farley suggested the name might be changed to “The Canadian Naturalist”. Taverner replied that this title had already been used and it would “ball up the bibliographies if they revived it’.”° However, to get over this problem the title was changed to “The Canadian Field-Naturalist”, while the name of the club remained ““The Ottawa Field-Naturalists’Club”.”” Not all subscribers to the journal supported the change of name. Taverner told Fleming of the event in a letter. “Last evening we changed the name of the Ottawa Naturalist to the Canadian Field-Naturalist and arranged the next volume to be.a short one running to the end of the year so as to bring the volume number to correspond with the calendar year”.’* Fleming was critical of the change. He said it was too bad the name had been changed, and that there should have been a vote taken by the members. He stated that the change in name would not help the journal.’? In his reply Taverner defended the decision. 110 “You are mistaken in believing that the change in name will not help the Ottawa Naturalist. It has helped it great- ly already. The change in form can only be supported by increased subscriptions and we have to rely mostly on Canadians for this. The jealosy [sic] of anything labeled Ottawa throughout Canada is surprising. We are not going to change anything but the name, — the volume numbers are to run along consecutively so the identity will suffer no break.” He also mentioned an index. “A general index would be a good thing but we cannot do it until we get money to publish it. I think I see this in the future’”.8° From January 1920 the number of issues per year was fixed at nine, the jacket was redesigned, and the price was increased.*! Taverner had already published a few articles, in addition to several brief notes, in The Ottawa Naturalist, before the change took place. At the beginning of 1919 his article “Bird-houses and their Occupants” was published.*? This was illustrated by a photo of a Purple Martin house with elaborate- ly designed roof and about twenty five martins roosting on it. It was built by Taverner in 1917, and was described in the article, together with two pages of scale drawings and instructions for its con- struction. The martin house was fixed to a hollow box pole set on a concrete base with a heavy weight inside running over sash pulleys at the top of the pole. In this way the house could be lowered to be cleaned, and easily raised again. In addition to the multi-roomed, colony-home for the martins Taverner showed designs for single room houses suitable for House Wren, Tree Swallow, Eastern Bluebird, Great Crested Flycatcher, Black-capped Chickadee, Eastern Screech Owl, and American Kestrel with the relevant measurements. The two pages of scale drawings by sections, as one might expect of a former professional architectural drafts- man, were neat and precise. When the issue of The Ottawa Naturalist containing the article appeared THE CANADIAN FIELD-NATURALIST Vol. 110 in January 1919 it was quickly in demand and in 1920 copies were reprinted for the Canadian National Parks Branch, with the addition of two pages of illustrations of various styles of bird hous- es.°* Two further editions were printed, as well as a French version. At the time of its publication there was nothing available on bird houses so well described and accompanied by such clear plans. But then, how many ornithologists were also trained draftsmen? It would be wrong to get the idea that Taverner took up too much space in the Ottawa Club’s jour- nal. In fact he was aware of the danger of the few active contributors on the publications committee giving the wrong impression, and said so to Fleming in a letter in 1918. “We can nearly keep it going from right here in Ottawa but we want to keep our own names out of it as much as possible so it won’t look as if we few wrote the whole thing’”.® Certainly the year 1919 was an outstanding one for the number of publications by Taverner. Apart from his article on Bird-houses in The Ottawa Naturalist, a local list, compiled by him, was printed in The Condor.*° Writing to Brooks at the beginning of 1919 he reported that three articles by J. Munro, B. Bowdish, and himself were all published in the same issue of The Auk. This, he wrote, was quite a Canadian number with a distinctly Canadian flavor and “helps to put us on the map”.*’ By now Canadian ornithology had taken a big step forward compared with its meagre position in 1911. The Great War was over, and when the New Year of 1920 arrived bring- ing in the decade of the 1920s Taverner might have looked forward to a bright future for himself, and the ornithology section of the National Museum. By now, however, approaching the age of forty-five, Taverner was experienced enough to look forward, though with cautious hope only. Part [IV — Ornithology in a Wider Perspective CHAPTER 11. Birds of Western Canada: Part 1 “T guess your reminiscences could be written from your letters to me. I have often thought what a valuable acquisition to the ornithological historian some generations hence. They will be a find for somebody.” From a letter by Taverner to Fleming, 16 March 1920.1 The decade that began in January 1920 presented Taverner with new opportunities but also new diffi- culties. He had now been at the National Museum for almost nine years and he brought with him into the 1920s considerable knowledge and experience of what was required of the Dominion Ornithologist. Not least of his qualifications was a growing net- work of friends and acquaintances in the field of ornithology. He had learned a sharp lesson in 1914 when he was unable to find a competent Canadian naturalist to accompany Camsell on his expedition into the Northwest Territories. From that time onwards Taverner was on the alert to encourage any Canadian who might become a sound naturalist capable of taking part in field expeditions arranged by the museum. He hoped that eventually he would find a few younger men with the qualities needed to lead field expeditions. In January 1915 a young man by the name of Dewey Soper wrote to Taverner from Guelph, Ontario, asking for advice on how to become a | | 1996 trained observer in natural science. He wanted to know what kind of wildlife to collect, what notes and observations to make, and what use he could make of photography. He also told Taverner something about himself. He was born near Guelph in 1893, his father was a carpenter who wanted him to follow his profes- sion, but Dewey preferred to work out of doors and earn money by farm work and trapping locally. Soper wrote in curiously stilted English.? Taverner seized the opportunity to send this young enthusiast a long letter from an experienced, older enthusiast. In this letter Taverner explained that he had received several enquiries such as Soper’s, and he decided to write a detailed reply. He wrote it in the form of an essay with the idea of working it into a paper for publishing and warned Soper “if you see most of it again in print do not be surprised”.* This material appeared in The Ottawa Naturalist a few months later.* Taverner out- lined for Soper the steps he should take on the way to becoming a good ornithologist, all of which would involve him in a great amount of work. But to the enthusiast this would be a labour of love, and if taken up as a hobby would develop an interest in nature to last the whole of one’s life. Sharing this interest with co-workers often was the basis for true friendships, and fruitful associations. Taverner concluded by say- ing that the museum would be glad to see anything he chose to send, either in the form of notes or speci- mens. Any time that he could be of any help either with advice, or in the determination of specimens he could call on Taverner. This was the beginning of a long association. In the next few years Taverner helped Soper by sending him suggestions on how to make good skins, also what to read of a scientific kind about nature. This was in reply to a serious “letter of intent” by Soper about finding his life’s work in natural sci- ence, and about throwing aside the restricting influ- ence of his vocation (farm work) and launching into the work nearest his heart. He confided frankly to Taverner his hopes for the future, and ended by writ- ing: “Now I sincerely hope I have not unduly imposed upon your time and good will in asking for so much information. As a matter of fact Mr. -Taverner you are the only gentleman with whom I might correspond on a matter of this kind.’ In the same letter Soper asked about openings in zoological work. Taverner was equally frank about the prospects and replied, “The demand for straight scientific work in Canada is very slight indeed, though the need of such work is great... It is to be hoped and expected that one of these days Canada will wake up and resolve to do her own work and not be dependent on American institutions.” He then cited his own experience in getting a job at the National Museum by working for years as an ama- teur ornithologist at some cost to himself. He was lucky to be well enough prepared as an ornithologist, when the opportunity came, to be considered the CRANMER-BYNG: A LIFE WITH BIRDS dale most promising choice. “It was largely chance and the forming of scientific connections who thought me worthy of recommendation.” He continued, “‘It is too bad that no hope can be held out from this insti- tution. We are in great need of a staff of men but till the authorities above can be made to see that, and this war is over, I can see no hope here.”° In a letter to Taverner written in 1918 Soper gave some details about himself. He was a single man, 26 years old, with considerable wilderness experience, who was making a collection of skins, and was known to W.E. Saunders.’ During 1918 and 1919 Taverner was trying to find him a position as camp- cook/naturalist with one of the Geological Survey teams on a summer expedition, but without success. In 1918 Taverner heard from Fleming about a Canadian naturalist of promise named Hamilton Mack Laing who was stationed at Beamsville, on the south shore of Lake Ontario opposite Toronto, as an instructor at the Royal Air Force school of aerial gun- nery.® Laing was born in 1883 and brought up in a pioneering farm community south of Winnipeg, had taught school at Oak Lake in southwest Manitoba, and taken art courses in Brooklyn, New York. By 1918 he had achieved a reputation as a nature writer. Taverner was already familiar with Laing’s stories in the two outdoor magazines Recreation and Outing. In addition Laing could draw well and take good pho- tographs.” Fleming used his influence to have Laing offered a job writing nature articles for the Toronto Globe which Laing did from 1918 for the next six years. Laing also came to know Hoyes Lloyd who was still working in Toronto before he was appointed to administer the Migratory Birds Convention Act. Fleming and Lloyd suggested that Laing should keep notes on bird migration while stationed beside Lake Ontario, and encouraged him to send the resulting article to Taverner as a contribution for The Canadian Field-Naturalist which he did.'° Taverner welcomed his article, and also wrote an appreciative notice on Laing’s articles in the Globe for The Canadian Field-Naturalist in which he said: “The subject of these papers cover such a range of sub- jects as “Hawks Everyone Should Know,” “The Wood Warblers,” etc. The subjects are treated in a popular, entertaining manner, in a style that more than occasion- ally warrants the term “fine writing”, sympathetically but with an absence of gush and with a good substratum of personal knowledge and common sense. We can stand many more of such popular science writers in Canada as well as elsewhere.””!! Laing spent time in the summer of 1919 exploring the Similkameen Valley and hills in the far south- west of British Columbia. He wrote enthusiastically to Taverner on the varied terrain, describing its trees, alpine flowers, birds and mammals with lists of what he recognized. He also took photos.!? In another let- ter to Taverner he described the years he spent in southwest Manitoba around Oak Lake between 1901-1914.'3 Taverner realized that he was in touch 112 with an observant naturalist with qualities that could cause him to develop into a first rate museum collec- tor. In this early correspondence with Laing Taverner took considerable time to help him with information, and sketches on how to distinguish birds such as gulls and geese by their bills, and by various stages of their plumages.'* A friendship between the two men developed that lasted until Taverner’s death. Another useful contact that Taverner made was with Frank L. Farley of Camrose, Alberta who was a friend of W. E. Saunders. In 1918 Farley met Anderson who was working in the prairie provinces on behalf of the Advisory Board on Wildlife Protection.!° After Anderson had returned to Ottawa Farley wrote to him as follows: “I have often wanted to meet Mr. Taverner and have him spend time with me on the Battle [river] when he is in the west. Will Saunders always mentions him as a fine bird man, and I like these kind of fellows. Ask him to write me for any information that he might wish as to our common birds.”’!° Taverner replied about the possi- bility of meeting him during his next trip west and added “... don’t believe all the nice things W. E. S. says of me as he is a partial judge where his friends are concerned.”!’ Farley’s next letter was about local birds and where some of them occurred on his farm. Taverner asked him about which species of blue- bird(s) occurred in his location. When Farley said that he always thought that it was the Mountain or the Western [Bluebird], Taverner asked him to col- lect a specimen for identification. Farley wrote that he had seen the first migrants on March 21st of that year and commented: “When I saw the three brave little fellows this spring I could not have killed one for a lot”, but added “I will try and bring myself to collecting one next spring and send it to you.” He next enquired about the two gulls that Anderson and he had shot and given to Taverner for identification. He had been out with Brooks and Saunders several times, he said, but neither could satisfy him whether the common gull in his location was Bonaparte’s or Franklin’s [gull]. Could Taverner determine the names of the gulls sent, and let him know what they were. He concluded “I shall be glad to hear from you often, and give information whenever I can.”!® Taverner responded warmly saying he had received Farley’s most recent letter. “I was very pleased to get it and anticipate that I have found a very helpful as well as pleasant correspondent. I see much of interest in it. | think we can probably help each other considerably.” Taverner had reason to be pleased since Farley had sent him 23 years of notes for Camrose and Red Deer as a basis for a local bird list. As regards the two gulls, Taverner wrote, the only positive criterion for distinguishing between the Herring and California Gull was the colour of the feet of the adults. On a personal note Taverner commented: THE CANADIAN FIELD-NATURALIST Vol. 110 ”You seem to have, like W.E.S., a good ear for bird notes. Now I lack that considerably and rely more on eyesight than ears.””!° In reply to questions about his list of birds seen in the Red Deer area, Farley sent useful information on such things as the breeding of Sandhill Cranes and Ospreys, among other matters.”? By now correspon- dence between the two men began to flow like maple sap in the spring. Farley gave Taverner useful information on several other matters such as conservation, and Taverner replied with information of much use to Farley. For example, on the decline in bird numbers Farley wrote: “The Swans (Trumpeters) are killed every fall and spring here, one man got three in a bunch last fall. Sandhill Cranes are getting scarcer each year, and histo- ries like what I am send [sic] you on the Pelicans might be valuable. All these birds that lay only two or three eggs are more liable to extinction than birds that lay in quantities like the ducks.””! This excited Taverner who replied: “Swans of all species are absolutely protected by the International Treaty now, but if you hear of any more Trumpeters being killed for heaven’s sakes let us get a chance to get them. We will pay at least twenty five dol- lars for Trumpeters, perhaps more if necessary if I can induce to [the] powers to shell Outs On a different topic Farley wrote about the muse- um at Camrose, and Taverner sent him information on how to exhibit birds in a museum and what kind of labels to use.” Taverner began introducing Farley to ornithology, and scientific collecting, just as he was doing for Soper and Laing. By now Taverner’s network of potential collectors, and sources of information on the bird situation in various localities, was developing well. It was greatly strengthened in 1919 when he received a letter from a man named William Rowan and was able to meet him in Edmonton in 1920 where Rowan had been appointed Assistant Professor of Biology at the University of Alberta.** On return- ing to Ottawa after his western expedition Taverner wrote to Rowan about someone living near Edmonton named Farley. Among other things he said “He is a man you should know and I advise you to open correspondence with him. He is a charming fellow and an enthusiastic, if not a deep ornithologist.””> In this way Taverner brought together two new members of his corresponding network. By the beginning of 1920 things in the museum seemed to be going well for Percy Taverner. His group of ornithological collectors had grown and held out promise for the future, his book Birds of Eastern Canada was selling very well, and the new Canadian Field-Naturalist was well received. But a cloud began to shadow his situation like a warning of troubles ahead. Jim Macoun died on 8 January 1920. Although his death had been expected for the past few months Taverner was upset. His hopes of working under the friendly administration of Macoun, as head of the Biology Division, were 1996 dashed. Taverner could hardly relish working under the supervision of Anderson who had started work- ing at the museum several years after him, when much of the hard preliminary work had been done. But there is nothing in his correspondence to show that he strongly wanted to be chief of division him- self. His lack of a university degree, his stammer, his admission that he was not good at managing others inhibited him from applying for the position with conviction. He was resigned to being under Anderson’s supervision but consoled himself with the knowledge that he would have considerably more time for his own ornithological work. Writing at this time, Fleming gave Taverner some positive advice when he said: “However, you yourself have work you like, and can plan it out yourself so you are fortunate. I do not see any reason if the direction of the department were offered to you why you should refuse.””° Taverner was not offered the post. Taverner’s change from guarded optimism about the future to one‘ of dismay can be seen in his corre- spondence with Fleming in 1920. When the Department of Mines was placed under the Minister of the Interior, Arthur Meighen, the opportunity occurred for the museum committee to approach the minister through his secretary and not through the director of the museum, William McInnes, for whom Taverner had little respect. As a result the museum attempted to strengthen its position vis-a-vis the Geological Survey, by asking for a number of changes. These changes, if put into effect, would go a long way towards “the establishment of a permanent and continuing policy”.”’ The result of these negotia- tions for a changed structure was that in December 1920 a compromise between the Geological Survey and the National Museum was made by which the museum became ostensibly a separate branch of the Department of Mines, but in reality not much was changed, and the museum had no separate existence as regard finances.** When the National Art Gallery was allotted an exhibition hall in the building Taverner asked Fleming if he would find a way of making public the museum’s situation. Parliament, he said, had voted nearly one million dollars to build the structure, it had appointed competent staff, but then had made the value of this investment questionable by failing to provide the proper equipment necessary for carrying on museum work. Instead the authorities allotted space in the museum to the National Gallery of Art for its expanding collection. The public did not appreciate the problems that the museum staff experi- enced in trying to carry out their work fully and serv- ing the public better.” Early in 1920 Taverner wrote a long and heated letter to Fleming, pointing out the weakened position of the museum committee by the death of Macoun, and now, he wrote, “Today we hear that Dr. Hewitt who might have been of considerable assistance is dead. Another thing to be con- CRANMER-BYNG: A LIFE WITH BIRDS 113 sidered is the fact that the whole museum staff now is American except myself”. Taverner asked Fleming if he could come to Ottawa and see the minister about the matter. “As Honorary Curator you have a right to say things that we in the service can not.’°? Meanwhile all that the members of the committee could do was to hope for better things when Charles Camsell was appointed Deputy Minister of Mines. He arrived in Ottawa from Vancouver to take up his appointment in June 1920. Another person, in addition to the Deputy Minister of Mines, who would have an impact on Taverner’s future career was Rudolph Anderson, who was appointed acting head of the Biological Division early in 1920. Until that time Taverner had hoped that relations between Anderson and himself would be reasonably satisfactory. But in March 1920, when Taverner could expect Anderson, as head of divi- sion, to assume a strong tone on behalf of the muse- um, Anderson kept quiet. Taverner relayed this unwelcome news to Fleming, and said: “Have been trying to arouse Anderson but he is hard to wake. Will have to get after his wife, she sees things as they are, and is a hustler and is determined that he will hustle too. Cannot make up my mind just where Anderson stands.”*! A week later Taverner told Fleming that the muse- um committee had held an indignation meeting over the reduced allocation of exhibition space, and had requested a meeting with McConnell. It was appar- ent that the Geological Survey was now indifferent towards the museum, and that Anderson could be helpful if he was prepared to take a firm line. But this was not to be, as Taverner explained. “The worst of it as it looks to me just now is that Anderson is not with us. As I said before I do not under- stand him as he does not seem to see the absolute impos- sibility of the proposal as far as a museum is concerned. He seems to think that in demanding what we regard as a minimum space in a building built for museum purposes we are unreasonable.” In his next letter to Fleming he said: “Anderson did not take part in the meeting yesterday. Do not think he is contemplating leaving [the museum]. He objects but makes no constructive suggestions. Associations with an inarticulate man is a little trying. A man who will not commit himself is always problemati- cal. Have tried very hard to win his confidence but can- not make it.”°? Taverner’s next letter was a very pessimistic one in which he said “Tt is yet to be shown that the government is intending to abolish the museum or that it really desires to use a mil- lion dollar museum building for offices.” In the same letter Taverner reported a talk with W. H. Collins, one of the leading younger geologists, who was seriously thought of as the next Deputy Minister. “His feeling towards the museum is one of academic interest but not at any expense of the Geological Survey.’>4 114 The uncertainties and frustrations experienced by Taverner during the past three months were relieved in April by preparations for the summer expedition. Writing to Fleming he said he was planning to explore an area of sloughs and ponds south of Quill Lake, Saskatchewan, which was a large expanse of uninhabited country where he expected to find cranes “and goodness knows what else”. He and Young were due to leave on | May.*> Just before leaving Ottawa Taverner informed Fleming of a more ambitious plan: “Have a new plan for the summer. After leaving Young at Quill Lake I may join Lloyd at Banff and come back with him in an auto right across the prairie Provinces, stopping at all the lakes and zig-zagging back and forth for them. This should give a magnificent opportunity for seeing the country and the birds with opportunities for stopping to collect any where along the road. I am get- ting quite enthusiastic over it. We would camp all along the poed and so have evenings and mornings for collect- ing.” Taverner and Young were disappointed with the conditions they found when they started collecting because a series of dry seasons had lowered the water levels. H. H. Mitchell advised them to try Kutawagan Lake, southwest of Quill Lake, where he joined them early in June. When Mitchell had been at Kutawagan in 1917 there was a large nesting colony of Franklin’s Gulls, but by 1920 it had been “dried out’, but Sandhill Cranes were found breed- ing throughout this district, though sparsely. When Taverner left for Banff, Mitchell, together with Young, moved to the southern end of Last Mountain Lake, near the Arm River, where they collected until early September. This area produced a good selec- tion of passerines.*7 Taverner joined Hoyes Lloyd in Banff in late July and they set out to make what he called in his Annual Report “an ornithological reconnaissance of the provinces of Saskatchewan and Alberta, for the pur- pose of gaining a general knowledge of the faunal conditions of these provinces.”** Percy wrote a graph- ic description of the early stages of the trip in Lloyd’s faithful Model T Ford, to his mother and sister. “We left Banff with our car piled high with outfit so we look like the real thing I can tell you. Water bags and tent poles tied outside — dunnage bags on each front fender beside the radiator and the rear seat piled full — we in rough clothes. Such an outfit is not too rare out here but we do occasionally excite comment on its com- pleteness.’?? Everything depended on the satisfactory perfor- mance of the Model T which was said to have been completely overhauled in preparation for the journey. It ran all right on the level and the brakes were fine, but its hill-climbing capacity was very unsatisfactory. After several wrong diagnoses nothing but a reboring job on the cylinders would do. This took two days, but after that the car went like new through cactus, sage bush, and desert till they reached Medicine Hat. THE CANADIAN FIELD-NATURALIST Vol. 110 Now they were in antelope country, and Taverner gave a word sketch of a herd of 12 antelope, ““beauti- ful creatures who galloped away from us in long springing jumps that just touched the ground at each bound”.*° In the same letter he described being shown over the range of an antelope reserve by its warden. He treated them to some fancy prairie auto driving over bumps and badger holes, so that they would have soon been lying on their backs on the ground if the car had not had a firm roof. Luckily the warden warned them that his brakes were poor, and offered them the opportunity to get. out and walk before descending into a large coulee between boul- ders at a 45° angle. They walked. So their journey continued, zigzagging from one place to another, camping the night and exploring in places such as Pokowki and Elkwater Lakes. Since Elkwater Lake was a Forest Reserve Taverner was forbidden to shoot, so he took a half day’s rest while Lloyd explored the country, and made enquiries from the local people. Lloyd, on his first trip to the west as Supervisor of Wild Life Protection in Canada, was beginning to realize the vastness of his domain, and the almost insuperable problems he had taken on. Taverner, on only his second trip to the western provinces, was a little frustrated at the lack of time in any one place to observe and collect. Lloyd, however, wanted to see as much of the different habitats as he could, so they had to keep moving. By September they were camping in places in Alberta such as Provost, Buffalo Park Camp at Wainwright, and at Vermillion. Here the superintendent of the park drove them in his car through herds of buffalo, which Percy described to his mother: “It was like the accounts of early explorers as we drove through their dusky ranks and saw a good part of 5000 buffalo scattered over the prairie as far as eye could reach.” They camped one night by the Battle River but whether they met with Frank Farley is not known. The car, now called by Percy “Henry”, was found to need a new differen- tial axle and new bearings. After that Henry was able to carry them successfully for the remaining 1000 miles of the trip.*! But before heading for home the two men drove to Edmonton to meet several people. It was here that Taverner first met his new correspon- dent, William Rowan, now assistant professor at the Department of Biology, University of Alberta. In his field notes for September Rowan wrote: “Tonight [20 September] Dr. Clarke came in at about 9 p.m. to haul me off to the MacDonald [Hotel] to meet Hoyes Lloyd and Taverner who were passing through Edmonton. I had a most delightful evening and came home about 12.30.”*? Lloyd and Taverner returned to Ottawa in October. Taverner summarized the trip in a letter to Fleming as “3500 miles of prairie work” and praised the Ford machine, saying that there were repair parts available “on the notion counter at every cross roads’ [store]”’.44 1996 Fleming also had been mixing ornithology and travel while on a visit to England, and in a letter to Taverner described some of those he met at func- tions of learned societies which he attended, also a visit to the zoological gardens, London, and enter- tainment including country house luncheon parties. Writing from Oxfordshire he caught the English country house in a few telling sentences. “Drove six miles yesterday to dinner at 2 p.m. at a famous house. Pheasant and hare, getting to be quite a gourmand, they certainly know how to live over here. If only they had central heating and baths with running hot and cold water as the Romans had in Britain 2000 years ago living would be the most comfortable in the world.”4 In mid-April Fleming was in Ottawa conferring with Hoyes Lloyd about a scheme for a major reor- ganization of the administration of the museum. Fleming told Taverner of the scheme in his hotel room instead of Taverner’s office for reasons of secrecy, and they discussed it. The plan was to create a single department of government out of the Parks Branch, Geological Survey and museum, under one executive who would be James Harkin. Taverner then went off on his field expedition to the western provinces, while Fleming and Lloyd continued to develop the plan. In writing Arthur Meighen, then Secretary of the Interior, Fleming outlined the plan, and the part assigned to James Harkin. He also explained to Meighen the reasons for the aura of secrecy under which he was acting. So anxious was Fleming about secrecy that he failed to warn Harkin in advance of the plan, so that when Meighen’s letter arrived instructing him to draw up an Act bringing the three departments under one head it came as a complete surprise to him. Early in November Fleming sent Taverner a long letter explaining the proposed act. Taverner read this draft of “An Act to Create a Canadian National Museum” thoroughly, -and sent Fleming his comments.*° An order in council was issued in November cre- | ating the post of a director of a national museum, but by the end of the year that was as far as the plan to ‘separate the museum from the Geological Survey had progressed. The post of director was given to William McInnes, but he was already a sick man when appointed. Taverner sensed that things had gone wrong when he heard that McInnes was to be the museum director, and realized that this probably ‘meant an indefinite postponement to the plan to cre- ate one department out of three. This proved to be the last chance for the museum to break away from the control of the Geological Survey and gain a mea- sure of independence.*’ As a result the museum never had a financial basis of its own. Its staff and its finances were under the control of the Geological Survey and remained so until 1947.%8 We can now turn to the writing and illustrating of Birds of Western Canada. Although Allan Brooks CRANMER-BYNG: A LIFE WITH BIRDS IS) left Canada at short notice in 1914, just when the Birds of Eastern Canada was being planned, he was lucky to return to Canada alive and in time to start work on Taverner’s next book from the beginning. Brooks arrived in Ottawa in April 1919 and went to the Taverners’ home. Writing to Fleming, Taverner described his visit with enthusiasm. “Am sorry you missed Brooks. He arrived Wednesday and left Saturday night for Vancouver where he is to be discharged. We had a four days wild dissipation of bird talk, and the more I know him the more I think of his ability both scientific and pictorial. He is one of our great men and it is only his extreme modesty that pre- vents him from being a leader. For wide and deep ornithological knowledge he has Furetes [Fuertes] backed off the boards and his personality is charming. I look forward to being in the field with him this summer. It should be a treat. Artistically I think he has advanced greatly. His contact with English artistic circles has sharpened his esthetic perceptions — a field he always excelled in. The amount of knowledge of birds that he carries in his head is astonishing as I realized when he sat down at the house one evening and drew a Chipping Sparrow from memory that is perfect in detail yet he has not seen one since 1914.” From now onwards correspondence between the two men expanded as they discussed various sub- jects of common interest. By late 1919 they had begun to exchange birds’ skins. Brooks how had a substantial collection but needed more to serve as specimens from which to paint. Brooks sent the National Museum a box of 20 skins, together with some interesting information for Taverner. He reported that from the Osooyos Valley, where he was living, to where the Similikameen hills divide there were five different zones of vegetation caused by the rise of the land from about 1200 to 8000 feet in a few miles. Within these zones he had found a colony of Sage Thrashers, noted that Brewer’s Sparrows were common although Spreadborough missed them, and that the first White-throated Swift in Canada was discovered in the crags at the top.*° Brooks had told Taverner that he required good skins for his work. In reply Taverner had said that although he preferred good specimens for the muse- um he could not be fussy. What the museum needed above all were sets of birds in various plumages.>! He sent a box of 20 skins to Brooks in return, among which were Spruce Grouse, Willow Ptarmigan, and two Stilt Sandpipers in spring plumage. In the same letter he mentioned that he had been comparing specimens of Evening Grosbeak, and discussed his tentative findings.°? Another subject on which they corresponded at this time was the possibility of Brooks compiling a provincial list for British Columbia, or at the least a local list. Taverner heard a rumour from Munro that Brooks had given up working on his Okanagan list, and took Brooks to task. “Don’t backslide like that”. “You are not getting any younger and should you drop out your work will all have 116 to be done over again — some of it cannot be repeated unless it is put in permanent form. And for goodness sake make it full. What may be an old story to you is news to us.””>? Brooks replied that he had not “reneged” on the Okanagan list, that The Condor wanted a B.C. list. His British Columbia list, when finished, would come out in a Pacific Coast publication of the Cooper Club but was delayed by his lack of records from southeast Vancouver Island. He was convinced that a provincial list was needed, even if it was somewhat deficient.°* Taverner was anxious to obtain distribution data on the birds of British Columbia for use in writing his book. He warned Brooks not to delay its completion for too long and added: “You know complete data is never obtained, and howev- er much you have you will always want more.” Brooks had also answered Taverner’s request for full details in his list by saying he would make it short and to the point. Taverner answered that he still thought full details should be given. “Brevity may be the soul of wit but ornithology is no laughing matter and I have found so many difficulties in working up localities from the brevity of notes that I hardly agree with you.”>> Correspondence between Taverner and Brooks blossomed in the period 1920-1926 because of their common ornithological interests and because of the book on which they were collaborating at that time. Taverner had several problems, though they stemmed from the central one of the correct identifi- cation of bird skins. Because of his position as ornithologist at the National Museum he was regu- larly faced with determining the identification of specimens of birds sent to him from all over Canada. This was not as straight forward as one might think, even when holding a specimen in the hand and com- paring it with trays of other specimens. The speci- men might have been badly damaged when it was killed, it might have taken several weeks or more to come by rail from a distant part of Canada. This would cause it to lose some of its colour. A satisfac- tory coloured guide to the birds of eastern or western North America still did not exist (1920). By 1921 Taverner had officially been instructed by the museum authorities to go ahead with writing a book on the Birds of Western Canada, with illustra- tions by Brooks. Taverner wrote to Brooks with the news saying that the Birds of Eastern Canada was such a popular success that a second edition was being printed, and that there would be no difficulty in arranging reasonable terms for Brooks to illustrate Birds of Western Canada.°° Far from relying on other books for accurate colour illustrations Taverner and Brooks had to make their own. Much of their correspondence in this period was concerned with getting information on distributions and making descriptions of the species and some subspecies. THE CANADIAN FIELD-NATURALIST Vol. 110 Taverner was among a few ornithologists at this time who wanted to form their own judgment on whether one group of birds was sufficiently distinc- tive from another group of the same species to war- rant it being described as a subspecies. This would involve the addition of a third name (trinomial) to the first name, which identified the genus, and the second the name of the species. A new subspecies would be proposed by a taxonomist and would be based on a group of individuals with similar charac- teristics in a geographic area. One of the main diffi- culties in proposing a new subspecies that would be accepted by the AOU committee responsible for revising the Check-list at intervals was to define the differences in colours that distinguished one group of birds from another as a subspecies. Sometimes the nuances of colour differences were very fine. In that case one had to rely as well on such things as slight variations in the measurement of the bill or wing. When Taverner was puzzled by a particular speci- men received from, say, southern Saskatchewan, which did not appear to fit the subspecies normally found in the prairie provinces, he often referred to Ridgway’s Birds of North and Middle America when he wanted to read a detailed description of several allied subspecies. But here another problem arose. Taverner was sometimes unable to reconcile the colours given in Ridgway’s descriptions of plumages with those of plumages as seen through Taverner’s own eyes. For example he described his problems over colour variations in a letter to Brooks of December 1922. Taverner had looked at 40 speci- mens of Pileated Woodpecker equally divided between specimens from British Columbia and east- ern Canada. He found variations in their plumages from deep black through slightly to decidedly brownish colours, but these colour variations did not appear to divide up by geographical regions. When he laid out a total of 111 Stellar’s Jays he found remarkable variations in the blue of the underparts, and the darker blue of the back. Some mantles appeared almost wood brown. In examining speci- mens from Vancouver Island and the coast, the Queen Charlotte Islands, and the interior the only trend he found was that the Vancouver Island birds were without a white eye spot, while the white spot was found on most of the interior specimens. He added that some day he wanted to study them with more care, and commented, “Large series showing [the] run of collecting and not selected material cer- tainly confuses the most plausible looking lay- outs.”°’ This was the story of Taverner’s career at the museum, so much that needed doing immediate- ly. It was a good thing that he was not a married man at this time when he was hustling to do the basic research for Birds of Western Canada. Taverner was unusual in stating publicly that he wanted to form his own judgment on any subspecies, unbiased by hitherto accepted authority. As he CRANMER-BYNG: A LIFE WITH BIRDS Lay Field camp, Eastend, Saskatchewan, 1921. P. A. Taverner, Alan Sampson, and H. M. Laing. (Reproduced cour- tesy of the Canadian Museum of Nature, number 53435.) expressed it, “... too many are guided by preconcep- tions, and refuse to look facts in the face’’.*8 Taverner was too frank to pretend to notice a nuance of colour when he was sure that he could not. In his continuing disagreement with Brooks over when to create a subspecies and when not Taverner some- times wrote that he could not see the colour differ- ences that Brooks declared existed. This exasperated Brooks so much that on one occasion he wrote blunt- ly, “No good talking to you of color distinctions. I really believe you are slightly color blind”.*? Taverner argued that he was not, and wrote, “I do not think I am color blind. Nobody ever admitted such a thing of course, but I have tested myself against others and cannot find any evidence of it.” In the same letter he complained that they may have misunderstood each other. “I may not always write perfectly clearly but I do not think you read any too carefully. At least you misunderstand a number of _ things I have said.” Early in 1922 Taverner wrote a letter to the lead- _ ing systematics research workers of the AOU enclos- ing a statement protesting against what he consid- ered was the debasing of the concept of genus in ornithology by the proliferation of subgenera. He claimed that the tendency towards the finer division of the genus threatened to complicate nomenclature and taxonomy until each group in a genus became the private preserve of specialists in that group. He asked recipients of the statement to read it carefully, and return it with their opinion.®! Taverner must have known that he was “sticking his neck out” reck- lessly. He might as well have saved himself the trou- ble, but that was not his nature. By the end of the year he had received only a small number of signa- tures in support of the statement. What Taverner hoped would be a strong protest to the Committee on Nomenclature of the AOU was, in fact, a fizzle and left Taverner looking a little quixotic. Brooks, who was at Berkeley, California, in December 1922 wrote that Taverner was not satisfied with trying to limit the number of subspecies. Now he wanted to limit the genera. He said that Grinnell, Director of the Museum of Vertebrate Zoology, University of California, and Swarth had discussed the matter with him, and two points struck them. One was that Taverner had only collected 14 signatures out of a total of sixty fellows, and only two were Canadians. The Washington section of the AOU, Brooks said, could start a counter petition and get more signatures than he had for his. Secondly, this multiplying of the genera was nothing new. During the past 100 years 118 one could trace its fluctuations in the form of a graph — rather like a fever chart — Brooks said. But this was not Taverner’s point. He wanted to prevent the proliferation of genera from becoming excessive. Although Taverner usually lost in his disagree- ments with Brooks, in correspondence and in person, on problems such as when to accept determinations made by fellow ornithologists, or assertions by others of the true colours of a particular plumage, nevertheless these arguments did much to enlarge Taverner’s knowledge of ornithology. The two men remained good friends, and respected each other’s sincerity. Taverner gained valuable knowledge from Brooks on such subjects as the physiological struc- ture of birds, and the nesting of various species in British Columbia. Taverner spent the season of 1921 continuing the “biological reconnaissance” of the western provinces which he had started in 1917 and 1920. He first went to Cypress Lake, Saskatchewan, where Mack Laing, his field assistant for the summer, and Taverner met for the first time.®? Next they spent a week around Eastend in the French River valley, Saskatchewan, camped at Gower Ranch, owned by a knowledgeable naturalist, Laurence B. Potter.“ Taverner then joined Hoyes Lloyd in late June, and together they worked through southern and west central Manitoba, collect- ing museum material from new districts, and investi- gating bird life in several forest reserves and bird sanctuaries in the prairie provinces. Laing went from Eastend to Oak Lake, Manitoba, where he remained collecting till the end of October. Taverner man- aged to visit Brooks at his home at Okanagan Landing, for the first time, early in September. In a letter to Laing he described his impressions with enthusiasm. “The country about Ok. Landing is very beautiful in every way, well diversified with arid flats, mountains, marsh, lake and hillside. The Bull pines are magnificent with their great red trunks, clean boles and slight under- brush. We talked over bird matters from many angles and though I already knew him fairly well I came away with still greater admiration for Brooks’ great store of ornithological knowledge. I know of no one with such a broad grasp of both closet and field information.” Brooks had suffered a setback during the summer of 1921 when a building used for storage and a work- shop burned down, destroying part of his earlier col- lection as well as part of his library and many note- books.®*’ Taverner must have discussed this disaster with Brooks while visiting him in September because in December Taverner wrote with a specification for building a bird museum room. This showed Taverner the draftsman, suggesting a six inch thick concrete wall up to the windowsill, the frame of the building above to consist of stucco on metal lath or galvanized iron. The roofing was to be made of non-inflammable material thus, in Taverner’s words, providing maxi- mum fire protection at minimum cost.®* Taverner had THE CANADIAN FIELD-NATURALIST Vol. 110 more than friendly goodwill in designing a new building for Brooks’ collection. The study skins were made by Brooks himself with noticeable ability, but the fire had been a warning and he began to think of depositing his collection in a zoological museum. Writing to Taverner in August 1921 he mentioned the subject by suggesting that he would be willing to deed his collection and his “new museum building” for cost of the building estimated at $3000, but retain- ing a life interest and pledging himself to enlarge the collection during his life time. In ten years’ time it would be a really representative North American col- lection. Californian people, he wrote, would jump at such a chance. However, he softened the prospect by saying, “but I would like the collection to stay in Canada as it is just what you need to round out your big Canadian series — enough U.S. stuff for compar- ison, and the [word illegible] and Mexican species for illustration.’ Taverner replied as best he could, but he was in a wretchedly weak position. Dr. Anderson, he said, as head of the Biological Division was the logical person to present the idea to the Director of the museum, but he was not yet back from a field expedition.’? Meanwhile Brooks was faced with a new possibility. In October his friend Harry Swarth of the University of California visited him while returning from collecting on the Skeena River in British Columbia.’' When Swarth was again at the Museum of Vertebrate Zoology, Berkeley, he wrote to say how impressed he was by Brooks’ collection. “The more I think of your collection”, he said, “the more I am impressed with its unique value, and it cer- tainly should be put beyond the possibility of damage by fire or otherwise. I have never seen such a series of water birds anywhere — not in numbers I mean but such beautifully prepared skins, such carefully selected plumage stages and such a mass of color notes.” Brooks relayed these words verbatim in a let- ter to Taverner adding that Swarth was unable to per- suade the University of California to take up the proposition that he had made to Brooks.’? Taverner quickly replied, but without any hope of achieving anything through normal channels. He said that he had discussed the question fully with Anderson, and unofficially with the Director, McInnis. As head of | the Biology Division it was up to Anderson to push it. He himself had proposed the matter, explained it | and urged it as far as developmental propriety permit- — ted. In any case nothing could be done until after the > election. Only when the new government was formed and settled could anything be done. Even then, Taverner continued, it would depend largely on the minister we drew in the shuffle. “Our present one is hopeless as he had absolutely no sympathy at all with any kind of research work that is not definitely and directly connected with material benefits. I understand he wanted to abolish the museum altogether | at one time.” 1996 Nature, 56275.) By February 1922 the Mackenzie King govern- ment was installed and estimates were being pre- pared. Taverner wrote to Brooks to advise him to send a formal proposal for deeding his collection to the National Museum to the Director, William McInnis, or better still to the Deputy Minister, Charles Camsell. If Taverner presented a proposal to Anderson, and he to McInnis, and McInnis to _ Camsell, the force of his proposal would largely be _ eliminated. To ram his point home Taverner added _ “Tt takes a dose of H. E. [high explosive] at this end _ to produce an impression at the other.” The analogy _ to a powerful emetic would have been noted by any- one reading his letter.”* It was not only the apathy of politicians towards the museum that drew Taverner’s anger, it was also the museum being ignored by the wealthy business class. Writing to Laing at this time about the complete lack of funds to purchase any- thing for the museum collections he said the situation was hopeless. “As for donations, we get so few of them that they hardly count. The American Museum is the one with tame millionaires to buy and donate _ such things. In this country people regard the govern- _ Ment as an opportunity to unload at a gain on, rather _ than a recipient of donations.” CRANMER-BYNG: A LIFE WITH BIRDS “Asoyoos Meadows” [Osoyoos] south Okanagan, British Columbia. 14 May 1922. Left to right P. A. Taverner, A. C. Brooks, T. L. Thacker, C. De B. Green. Photo by H. M. Laing. (Reproduced courtesy of the Canadian Museum of 119 As always it must have been a great relief to get away from Ottawa and the shadow of the govern- ment-business world, and be with congenial people in the natural world. In 1922 he was able to leave Ottawa unusually early and meet his assistant Mack Laing at Osoyoos Lake at the southern end of the Okanagan Valley. Here Brooks was installed at a cabin on a ranch at the beginning of May. Many years later, in writing his biography of Allan Brooks, Laing gave a humorous description of his first meeting with Brooks who was at a table skin- ning a bird. Taverner began to make the introduc- tion: “Hamilton Laing — Major B-B-B” but that was as far as he got when “I’m Brooks” the bird- skinner said, holding out his hand. “Brooks” echoed Taverner loudly.’° Brooks was on “home territory” and finding uncommon species was not much of a problem. Peregrine and Prairie Falcons on the cliffs, Williamson’s Sapsuckers in larch trees at the sum- mit of a creek, and the rare White-throated Swift. This bird was known to occur in Canada only in the southern Okanagan Valley and entailed a climb to crannies high on the face of mountain cliffs to watch them in spectacular flight around their “com- munal nesting strongholds”.’?7 On 19 May they 120 moved to the southern end of Vaseaux Lake where they remained until 15 June when they went to Okanagan Landing for a week. Taverner wrote to his mother and sister from Vaseux Lake with a lyri- cal description of the scene. “We got here Friday P.M. and it is one of the loveliest spots I ever saw. A beautiful green and level valley right at the foot of Vaseux Lake. On either hand rise great mountains, and the exit just below us is framed by a sheer 800 foot cliff, standing up like El Capitan in the Yosemite. We have another comfortable house; about it are apple trees and green, green meadows, punctuated by wonderful bull pines, with forms like Norway pines, but 150 ft. high, and with a golden pink trunk. In sight from our window I can see a great Osprey nest in the top of a tall pine. Canada Geese occupied it until the Ospreys drove them off. Out the other window is a flock of 14 Canada Geese feeding in the meadow, not 100 yards away, and absurdly tame. Yesterday Brooks and Laing saw a mountain sheep not ten minutes walk away. Birds are singing all about us. You can see what a beauty spot it is.”’* When Frank Farley arrived and a hunting friend of Brooks, George Gartell, as well as a student assistant, Alan Sampson, a certain air of light heart- ed fun, like young men on holiday, developed. For instance in a letter to his mother and sister Taverner wrote that Farley had spent a few days. “He surely enjoyed himself, and we did too. Was sorry to see him go. He has all of W. E.’s enthusiasm, and a heart like an ox.” Then Percy related how Gartett drove Farley, Taverner and Laing to White Lake in his car to walk the sagebush area in search of a Sage Thrasher. Although they failed to find one they had a fine adventure on the way back, as Percy wrote in the same letter. They stopped at a Mr. Hady’s on Dog Lake. “He has a beautiful place, all orchard behind and a lake in front. Porch covered with wistaria in bloom. He entertained us in the basement where he had barrels and barrels of cider two years old, and port wine that would bring a smile of satisfaction to a teddy bear. We were unanimous that a Province where such things could still happen was the country to live in.””? This became known as “the incident in Mr. Hady’s cellar” among the four men and their friends.°° Brooks returned home to Okanagan Landing on 7 June but Taverner and Laing camped for a few days near the summit of a 3000 foot peak to collect species not yet seen. They had considerable luck. First Taverner collected a dark form of the Pigeon Hawk [Merlin], then considered to be one of the rarer North American birds.*'! Both men knew that Brooks had been in pursuit of this dark coastal form for a long time. This is how Laing recorded it in his life of Brooks. “Tt is my considered opinion that Taverner was the luckiest man who ever packed a collecting gun afield. He turned up the most unexpected things, not by special skill but by sheer blundering luck. To THE CANADIAN FIELD-NATURALIST Vol. 110 prove my point, next day at base camp near Vaseux Lake he returned from a short walk at midday with a Dickcissel, a bird that anywhere west of Manitoba could be called a rare take. This was a new record for British Columbia!’’*? Since Brooks himself had seen neither a Dickcissel or a male dark form Merlin (Falco colum- barius suckleyi) in the Okanagan Valley it was cer- tain that he would say that the observer was wrong in claiming he had. Brooks was well known to Taverner for holding an opinion more stubbornly than anyone else — only a specimen in front of his eyes would persuade him. The two specimens, appearing suddenly on the table in front of him came with considerable force. The Major didn’t keel over but he went scarlet, and didn’t say a word.® Taverner and the others spent ten days at Brooks’ home where he maintained an unofficial private bird sanctuary except that birds of prey — owls, crows, hawks, magpies and other “undesirable birds” were shot on sight with the result that Brooks had plenty of “desirable” birds to study — almost tame and eas- ily observed nearby. Brooks had this trait in common with Jack Miner — that he alone should determine what birds should remain alive in his private sanctu- ary. While there Taverner was able to look at Brooks’ collection of several thousands of bird skins.8* Jim Munro, Federal Game officer for the western provinces, came to visit them from his home close by. Another visitor was James Harkin, Dominion Parks Commissioner, which gave Brooks and Taverner the chance to have a good discussion * about birds, and especially on the justification for the protection of wildlife, and the conservation of the habitats in which it lived. Harkin was expressing a philosophy for wildlands preservation at this period, and his meeting with Brooks, Munro and Taverner in the Okanagan Valley can hardly have been just for a day or two’s holiday.®> Since Percy was having such a stimulating time in such beautiful surroundings while his mother and sister were spending the summer at Ottawa and the cottage, which was hardly as exciting as travelling, Percy took trouble to give them any news that he felt would interest them, as well as to reassure his moth- er that he was in good health. In this way pieces of information can be gleaned from his letters. Taverner was still troubled from time to time while on field trips when he prepared his own skins using arsenic. On the present expedition he was bothered so much by arsenic that he had to give it up altogether. He explained: “Am using alum now instead. I did not like to do this, but after all with our good tight cases and liberal use of naphthalene, arsenic is only moral support. Otherwise all is going well.”°° While at Vaseux Lake he heard of someone called McAstocken who wanted to see him. Taverner had no idea who this was until one day he came to their 1996 CRANMER-BYNG: A LIFE WITH BIRDS WIL Farley; left to right, seated: Hamilton Laing, George Gartell, Alan Sampson. (Reproduced courtesy of the Canadian Museum of Nature, number 56293.) ‘camp, and turned out to be one of the Guelph boys with whom Taverner used to go bird collecting, and ‘they had a great time talking about the old days. /When in Penticton one day, while waiting for the boat, Taverner saw McAstocken again in his home town. This is how he described it: “Penticton is an ideal little place to live in. Flowers everywhere, every home has nice grounds, and covered with roses. Mountains, and a lovely lake, with a beach | where the whole town turns out in the afternoon for bathing. McAstocken has a nice little cottage, four chil- dren, two dogs, and three cayuses on which the kids scour the country for miles around. Some life.””*” In the same letter he mentioned the nearest neigh- bour at Vaseux Lake was a family by the name of Parham who were friends of Dr. Saunders.”®* On the recommendation of Brooks Taverner, Sampson and Laing continued their collecting expe- dition at Comox on the north east coast of Vancouver Island from 28 June until 15 August. While there they collected a Pelagic Cormorant, Phalacocorax pelagicus, the violet-green cormorant of the Alaskan coast and south to Washington. While it was still fresh Taverner made a colour wash drawing of its soft parts. The inside of the mouth he showed as pink to reddish colour, while the feet were solid black. This specimen is still in the collection of the National 122 Museum, Ottawa, and Taverner’s drawings on 5 x 8 inch cards, are still in good colour.*? On the way from Comox to Ottawa Taverner spent a day in Victoria looking at material in the museum, and spent the evening with William Spreadborough. Writing to Laing, after his return to Ottawa, Taverner suggested that if Laing wanted an interesting evening when in Victoria he should look him up and added, “he has seen more of Canadian wilds than most people and will talk about it by the hour.” In the same letter Taverner mentioned a talk with Thompson Seton who came to see him. Taverner wrote, “The day after I got home Seton came in and I had to give up all my time to him. He is a genius and an enthu- siast even if he has got some erratic spots. It is too bad that his methods of gaining the public ear should obscure the really good work that he has done and is still doing.”?? Each season when Taverner went on a field expe- dition he kept daily record books. He sent his notes for the summer of 1922 to Laing. Laing answered, when he had read them, that he would be dizzy for the next few days and added, “That typewriter does- n’t spell and punctuate better than it adds 6 + 3. Better get a new one.” In his notes Taverner had CHAPTER 12. Birds of Western Canada: Part II During 1923 and 1924 Taverner worked mainly on the first draft of Birds of Western Canada, while Brooks worked on the first draft of the “pictures” during 1922 and 1923. Early in 1922 Brooks sent some small examples of what he planned to do. In thanking him Taverner said they were “delightful”, and asked the authorities for approval for Brooks to make one hundred illustrations for the book. ! Taverner wrote to Brooks in September about the book and the list of his planned illustrations, enclos- ing a copy of a memorandum for him to read and add suggestions. Approval was given for 100 illus- trations at $14 each.’ Throughout 1923 Taverner worked on the manu- script without going on a field expedition. Much of his time was spent on correspondence with Brooks which included a good deal of technical, ornitho- logical detail. Writing early in 1923 he explained his plan for the book to Brooks. It was designed for the amateur ornithologist, plumage descriptions were only to indicate the main points, though the “distinctions” should be clear enough so that a species could be distinguished from any other species that it might be confused with. As Taverner explained: “I think that a perfectly accurate manual can be made this way without the confusion of elaborate details. Field marks of course [are] self- explanatory”. In addition to the illustrations by Brooks small pen and ink figures by Taverner were THE CANADIAN FIELD-NATURALIST wanted to use the word butte meaning a steep, flat | topped hill, but spelt it phonetically. Laing jokingly } ' wrote: “You really ought to spell “butte” so. | Posterity may think you mean a “beaut” of a differ- ent sort.’?! This may sound as if Laing, the former school teacher, was trying to teach Taverner the) rules of good writing. But this was not a deliberate “put down”, because in the same letter he wrote | with warm praise for something Taverner had writ- - ten in his 1922 notes. That paragraph on the song of the Yellow-headed | Blackbird was the best thing I ever saw you write. You | waxed eloquent. I copied it out and put it away. Why | don’t you ride a tall horse like that a little oftener?””” Taverner answered in the same jocular style, | when he wrote: | “As for the aspersions cast on spelling, I am above such | details. Genius refuses to be bound by such technicali- | ties — see how Shakespear spelled?” Taverner also thanked him for his kind words on > the song of the Yellow-headed Blackbird but added, “Tf you were writing for our department you would soon learn to look for small horses to ride.’’?? Laing was a professional writer who was paid to | write with a measure of imagination while Taverner + was held on a leash of red tape. planned for inclusion in the text. In the same letter | he explained that these were intended to point out the field marks of selected birds only.? A good | example of two such drawings are of the Semi- palmated and the Piping Plovers. The head and breast diagrams point out the differences quite effectively. Other of Taverner’s figures show eagles, vultures and hawks in flight as well as their heads and beaks, claws, and wing patterns. Many of © the readers of the book would collect birds for identification, and would need to distinguish their specimens by measurements and detailed compar- isons of wings, bills and feet. This was a demand- ing task and Taverner sought help from Fuertes on raptors, and from Brooks on field markings of some western birds with which he was not sufficiently familiar. Fuertes sent him six pages of notes and sketches 5 on hawk outlines.* Taverner told Brooks how valu- - able these were for the book. “He certainly has an enormous fund of detailed informa- - tion on how plumage lies. His rough sketches are beauti- ful in the certainty of their drawing”.> Writing to Laing early in 1924 Taverner said “With the advice of Brooks and Fuertes it won’t be my fault if the amateur cant identify a hawk at the first flick of its wings.”° Taverner was making every effort to devise the best ways, both by illustrations, figures, and in the organization of the text, to present the reader with an Vol. 110 ene 1996 accurate but handy manual without the confusion of elaborate details.’ Throughout 1923 and 1924 Brooks was reading sections of Taverner’s draft of the text, and com- menting on them. Both men were frank with their criticisms. Taverner told Brooks that he was glad to have criticism of his line drawings, and that Brooks would not hurt his feelings by improving his draw- ings. “T do not claim to be other than a conventional drafts- man. Pen and ink is not my medium and I do not feel at home in it”.® Brooks also criticized Taverner’s written material, especially when the meaning was not clear. In reply Taverner admitted one example and wrote “T admit not reading over what I write. In fact I know of little that is more disagreeable than reading my own stuff either in letter or in proof and I am a wretched proof reader, and constantly read the thoughts I had in mind rather than what the page says.”” After Brooks had decided on which species he wanted to paint he sent Taverner a list of those already painted by Hennessey which, he said, should be retained for use in Birds of Western Canada.'° Meanwhile Taverner was regularly returning illustra- tions to Brooks with his comments and, when neces- sary, requests to alter them.!! Both men were send- ing frank and voluminous notes to each other. As an example Brooks wrote about Taverner’s description of a Western Bluebird as “Abdomen white with a slight bluish tint’, the words “Take a look at a good male & see if this is correct. I would call it dull bluish”. At the end of this section of comments Brooks wrote “T think you have reached a very fine climax in these last two parts. Some of the habit-portraits are the best thing I think you have ever done and I predict that they will be quoted very often in the future. My sincere congratula- tions.”!? Taverner could be equally appreciative, praising Brooks’ portrait of a pair of Western Bluebirds poised against a background of a bare, curling branch which looks weighed down by the two birds perched side by side on it. They are high-lighted by a two- _ toned sky, light grey in front and a pale yellow above, suggesting an evening in early spring. By con- _ trast, a pair of Eastern Bluebirds, painted by Hennessey, are shown in the same plate, in much the same pose, but against a pale wash over an empty background. Another portrait shows a pair of Mountain Bluebirds perched on a redstone boulder with a suggestion of vegetation at their feet. In front of them is a mountain valley with a pale apricot wash | in the sky. The birds fill almost the whole picture.!* But not all their correspondence was satisfactory. They disagreed over the confidence with which even an experienced ornithologist can identify subspecies without first knowing the region from which they came. Lack of sufficient specimens, Taverner said, was largely the reason why he hesitated to use sub- CRANMER-ByYNG: A LIFE WITH BIRDS 123 specific names where there could be any doubt. He gave examples of problems over the identification of subspecies of thrushes and Song Sparrows.'* This led to a battle of words which had Taverner writing in exasperation: “You are certainly ‘sot’ in your ways. Made up your mind that I am color blind and cannot be convinced otherwise. You don’t get the other fellow’s view point even constructively.”!> Brooks hit back when Taverner would not agree with him over subspecies of geese in North America. He told Taverner that he had studied Taverner’s tables on the geese “but I absolutely doubt your ability to see anything after studying your tables ... on the geese. You are the first man I ever saw who did not see light when he saw real minima [Branta canadensis minima] — as a matter of fact (from your letter) you did but your sceptical nature at once started to nullify this impression’’.!° Taverner was challenged by enough well-known ornithologists in North America over a long enough time on the subject of not accepting subspecies read- ily on other people’s authority that he acknowledged that he was considered to be something of a maver- ick. A new Canadian ornithologist had become a correspondent of Taverner’s from 1923 onward, Lester Snyder of the Royal Ontario Museum of Zoology, in Toronto. Snyder had written to Taverner for advice on what birds to look for when collecting in the Lake Nipigon country of northwestern Ontario. In due course Snyder sent specimens col- lected there to Taverner to examine, and when Taverner returned them with his notes and identifica- tions he explained to Snyder how difficult it was to be completely certain in identifying various sub- species of the Great Horned Owl, and the Sharp- tailed Grouse found in the area north of Lake Superior. He proceeded to explain why, and to dis- cuss another species, the [Dark-eyed] Junco. He was by no means convinced that all the existing junco subspecies in Eastern Canada were viable. He cau- tioned Snyder as follows: “One thing I warn you in accepting my opinions is that I am rather of a skeptic and will not receive without quali- fication postulated subspecies that I can not demonstrate in specimens. I am regarded as more or less of a crank because I will not vouch for what I cannot see, and do not believe in jumping at geographical conclusions.”!” But although continually attacked in the “bush war’ over subspecies Taverner persisted in holding his head up. Although it must have been quite a strain to keep up Taverner’s circle of regular correspondents, he clearly enjoyed the exchange of the latest informa- tion on both birds and people. There was no formal style in his letters unless he was writing to a senior civil servant and etiquette required a standard form. Otherwise he seemed to have suited the style to the nature of the man to who he was writing. Like his friends he was a compulsive letter writer. From time 124 to time he would mention that he was writing his fourth or fifth letter for that evening and he felt worn out. But writing letters was part of his life, some- thing which required a lot of self-discipline to keep up, but something which brought much pleasure. In the days before the telephone was an easy alternative to a letter, writing a letter was like talking to a friend. The writer needed to have a clear image in his mind of the recipient, how he would respond to the subject matter and the style of the letter. In return Taverner would know what kind of letter to expect in reply. When writing to Fleming Taverner gave free reign to his humorous, whimsical thoughts, knowing that Fleming would do the same. They were about the same age. Neither had a university education, and they both expressed themselves freely and equally. His relations with Brooks were a little different, perhaps because Brooks was a few years older and had spent the whole of the war in France where he served with distinction. By the time their correspondence blossomed Brooks had already begun to make a reputation as a good bird artist, something which Taverner had the training to appre- ciate. The letters between the two men were frank and sincere; they appreciated each other, and exchanged forthright views about the AOU, and other ornithologists, but the tone of Taverner’s let- ters to Brooks rarely contained the banter and leg pulling that the letters between Taverner and Rowan, and Laing did. When you got to know him, Brooks was a good camping companion, but he held very firm opinions. If you disagreed with him you had to be prepared to stand up to him, and argue back. But to do so was a serious business, and could not easily be turned aside by a few witty or amusing phrases. When writing to Mack Laing, Taverner was on different ground, because Laing was eight years his junior, and because Taverner was a professional ornithologist, and had just written the only bird guide to eastern Canada when Laing got to know him. Fleming and Taverner were both Laing’s patrons when he began to develop a new career as a collector and writer. The relationship between Laing and Taverner was an easy one in the early years of their friendship. They both were ready to rib each other, to write at times in a light-hearted style of banter. They had plenty of jokes and fun when in the field together observing and collecting birds. They did not have to feel any sense of rivalry over who was the better ornithologist — Laing had a wider education than Taverner, and was a professional writer while Taverner was an ornithologist with long experience who also wrote well for publication. Much of their correspondence was written in a spirit of exuberant bonhomie. Here were two middle aged bachelors keeping up their spirits from time to time by a slightly artificial style of boyish humour. In the early 1920s, each may have needed the other to talk to, a congenial person with whom to share recent THE CANADIAN FIELD-NATURALIST Vol. 110 experiences and present enthusiasms, someone to buoy him up over the low points in his work, in par- ticular the absurdities and the red tape.'® Another recent correspondent of this period was William Rowan. Writing to Brooks in late 1923 Taverner said: “Rowan is a great acquisition to Canada & is going to do something worth while for Alberta. His list of 30 species of waders seen at Beaver Hills L. [Beaverhill Lake] in May is a stunner. ... Rowan is very much taken with Harold [Harrold] and not, I believe with Soper, and you see the best (& the worst) of a man when you are camp- ing with him.”!? The tone of Taverner’s correspondence with Rowan was a little different from that with other major correspondents in the period, perhaps because Rowan had a very different background and training. Son of an Irish father and a Danish mother, he attended an English Public (1.e., private) school, enrolled in London University where he graduated with a B.Sc. in zoology in 1917, and an M.Sc. in 1919. By this time Rowan had become interested in the distribution and breeding biology of birds in Britain. As a boy he had been impressed by illustrat- ed lectures given by Ernest Thompson Seton, and decided he wanted to become a naturalist in Canada. The time was opportune; the war was over, the uni- versities were appointing academics in zoology. Rowan accepted a post as lecturer in zoology at the University of Manitoba in Winnipeg, in 1919, and was appointed the following year to build up a Department of Zoology at the University of Alberta in Edmonton.”? Rowan was already in touch with Taverner in late 1919 from Winnipeg and a regular correspondence soon developed. He was glad to refer to Taverner for information on ornithological questions, especially since Taverner was working on a book about birds of western Canada. In addition, Taverner could keep him posted on what was going on in the field of ornithology, not only in Canada but also in North America, through the AOU. Above all Taverner introduced Rowan to other ornithologists at this time when Rowan was rather isolated in Edmonton. Some quotations from their correspon- dence will show this relationship. For instance, in March 1922 Rowan wrote: “Another long letter from you, for which many thanks. They are one of the things that make this crowded life worth living.” Later in the same letter he wrote “I do hope you do not mind being bombarded with my perpetual queries. I must waste an awful lot of your valuable time. It is not often one gets the opportunity of talking birds here reasonably with any one.”7! Taverner, from his point of view, was willing to spend a lot of time writing to a man of Rowan’s training and enthusiasm, who was collecting and studying birds in the prairies. This was expressed clearly by Taverner in August 1922 when he started a letter: 1996 “Dear Rowan; Bully for Rowan. Glad to see that he got so much good stuff. I foresee that one of these days Canadian Ornithology is going to come into its own. Brooks col- lecting in B.C.; you in Alta.; Mitchell in Sask.; Harrold and another young fellow of whom I see promise in Man. besides the few in Ont. and Mousley in Quebec ' makes the future look rosier than it has ever [been] before. I hope that I have been and may be in the future some little influence on its development.”?” _ That Rowan was now settled in Edmonton with ‘zoology students, and a keen group of men joined him -at Beaverhills Lake to hunt water birds was, for Taverner, a great stroke of luck. Not only did he ‘receive new records and specimens from Alberta, he ‘also got excellent information for his book Birds of Western Canada. Each man appreciated the help of ‘the other, and said so. This is Rowan writing to | Taverner near the end of 1922. “I think you are one of the best letter writers in the country. Many thanks for ‘the last which was, as always, full of interesting infor- /mation.” In the same letter Rowan mentioned that although he was fond of collecting study skins his real interest lay in problems of migration, dispersal etc. “that can be so admirably studied out here where one ‘has the ground to oneself.”’** In response to a request from Rowan about what studies had been written on birds introduced into Canada Taverner sent him infor- mation on: [European] Starling; [Ring-nécked] Pheasant; Hungarian [Gray] Partridge; Mina [Crested ‘Myna]; [Eurasian] Skylark among other species.”* _ Both men had a strong sense of humour and the ability to laugh at themselves. The topic of bad ‘spelling came up early in their correspondence and each recognized the other’s weakness. Rowan once wrote “cheque” when he meant “‘check’’.2> Writing to ‘Taverner to thank him for a letter, Rowan put the ‘matter with a touch of whimsy; “I wonder whenever I get one of your delightful letters _ which of us would win out in a competition for rotten | spelling. I used to think I was unbeatable ... ”° __ Taverner replied also with a touch of whimsy: _ “Don’t rub it in too hard on the spelling. If the Dept. were richer I would have a ten dollar a week stenograph- er to attend to those details for me. As it is my mind out- _ runs my fingers and this machine doesn’t know the first thing about spelling.” But he ended the topic on a serious note when he ‘said | “I hope that you, like me, believe that the sense is the _ important essence.””/ | Taverner’s personality was a little unconventional. He had a creative rather than a strictly “play by the tule” mind. So did Rowan who replied with a witty thrust. “T'll forgive your rotten spelling. A rival is always a good thing.”28 However, most of the Rowan-Taverner correspon- dence was about ornithology, Taverner commenting on Rowan’s records of birds collected and records of I | CRANMER-BYNG: A LIFE WITH BIRDS 125 birds made at Beaverhills Lake, and Rowan com- menting on his frustrations over not receiving per- mits to collect, as well as a strongly-worded diatribe against Harrison Lewis, editor of The Canadian Field-Naturalist, for suggesting “vermin” such as crows and coyotes should be exterminated.”? Taverner had already had some experience of Rowan’s impetuous nature when frustrated, especial- ly over matters affecting his research, and his ten- dency to send off sharp letters of complaint without thinking of the consequences. Several times Taverner had warned him to be careful. About his criticism of the vermin matter Taverner wrote: “Constructive and sympathetic criticism is helpful but look out that you do not encourage attack that may ... cast discredit on the whole movement.”*? Later Taverner wrote that Lewis would not alter his beliefs once formed. “Lewis is all right. He is very set in his ways but very conscientious and painstaking. He is open to conviction but awefully hard to convince. He will be absolutely fair according to his lights but wont compromise them one bit ... There are a great lot of his type among the bird protectionists and they can not be overridden roughshod as I fear they are the majority among the active powers. They got control of the Audubon Society a few years ago. Educational work has begun to lower their influ- ence and sanity is reappearing ...”*! About this time Rowan was in a state of anxiety over a questionnaire that all Alberta permit holders had received. If permits were cut down he would lose skins sent him from various parts of the province, which might upset his research work. “For the love of Pete”, he wrote, “if you get the chance, dig Lloyd in the ribs and get him to hustle things.”*” Taverner sent Rowan’s letter to Loyd who replied: “Dear Percy. Tell Rowan to keep his shirt on. There are dignities to uphold on both sides, but we will always do anything in reason within the constitution.”*? As a result Taverner wrote a diplomatic “soothing down” reply to Rowan’s letter, the first of a number of such letters that Taverner would be required to write.*4 In November 1925 Anderson and Taverner attend- ed the American Ornithologists’ Union (AOU) meet- ing in New York and presented an official invitation from the Minister of the Interior, Museum officials and the Ottawa Field Naturalists’ Club, to hold the forty-fourth stated meeting of the Union in Ottawa in 1926. The invitation was accepted and the date was set for October. This would be the first meeting of the AOU to be held in Canada. In addition to the program of talks, meetings and an art exhibition, planning for several other events was carried out from the beginning of 1926. For Taverner and Brooks the main event was to be the presentation of a copy of Birds of Western Canada to each AOU member attending the meeting. This meant that it was urgent to begin the process of printing the coloured plates, and Taverner wrote to the Editor-in- 126 Chief at the museum setting out what would be required. The technical language used by Taverner in explaining this shows his familiarity with terms used in art printing.*° Taverner then had a meeting, which he jocularly referred to as a seance, with the printing bureau. This he reported to Brooks in the technical terms used by engravers.*© During 1925 and early 1926 letters went back and forth between author and illustrator on the printing of the book. But these let- ters also touched on the wider purpose of the wildlife artist, and the skill required by him to make his illus- trations live. Brooks told Taverner how he had paint- ed a picture of waveys (Snow Geese) but it did not turn out as he wanted. He explained why, and what he would try to do next time. “Composition is everything. If I try it again there will be nothing but Waveys on one plane, quite big and only a solid blue sky.”37 While at the AOU meeting Taverner went to see Audubon’s original paintings of North American birds owned by the New York Historical Society. In a letter to Brooks he remarked that there was consid- erable discrepancy between the colourings of these originals and the colourings in the editions printed from them.** Brooks replied with an account of work done by Audubon’s contemporaries in France, stud- ies of dead game birds coloured on copper plate, “of exceptional merit made about that time. The trouble with Audubon’, he wrote “was his absurd method of crucifying his subject on a board; he never looked at a live bird [while doing it]”. Brooks then described the old style of reproduction, hand coloured on cop- per or stone engravings, and how it was carried out.*? Meanwhile Taverner was wrestling with the problems of printing the coloured plates of their own book as he explained to Brooks in January 1926. Originally he had expected it would take a month to run the edition but had now learned that nearly half a million impressions would be required and that these would take four months to run. The complete text was now printed but if everything went well it would be mid-June at least before the complete edition would be ready for distribution. Taverner added: “T will certainly be relieved when it is all done and off my hands. Think I will swear off making books then, especially under official auspices.’”?? At last, in March, Taverner could write that the edition was all printed, that the new paper for the coloured plates had arrived, been seasoned, and that printing was ready to start “next week”. He added some technical information for Brooks’ interest: “We have put the fear of the Lord into the [Printing] Bureau in general, I think, and within a few weeks and before the weather makes a marked change they will have four presses on the job so there will be as little time for atmospheric changes in the paper between printings as possible. When you come down to the details of this color printing one is surprised at the enormous difficul- ties that have to be surmounted. Saw a case yesterday where they turned the heat off the printing room over a THE CANADIAN FIELD-NATURALIST Vol. 110 week end and Monday morning the paper was nearly one eighth an inch out of register. Hygrometers show the Ottawa air in winter approximately bone dry while in summer it nearly approaches saturation mark yet we have to straddle these two seasons now and keep regis- ter. With four presses going, one on each color I think we can get results. I think you would be much interested if you were here and could see the job through right with the men on the machines rather than with the men in the office. It raises ones confidence in the Canadian work- man. As the plate printing will take about 3 months I see no reason why the book will not be out in June and ready for distribution by July. However it is always the unexpected that happens and this book seems to have had a hoodo so I won’t make any promises. I hope it will be ready for the A.O.U."4! Because Brooks and Taverner could not talk to each other on the phone they had to keep up their “long distance conversation” by writing an intermit- tent series of letters. Their main topic was birds, and the handbook they were working on together. But also they wrote about artistic matters, travel and mutual friends. They seem to have enjoyed their cor- respondence in these years, perhaps because they had much in common to enjoy. Early in 1924 Taverner wrote to Brooks that the Biological Board of Canada was interested in pub- lishing a fauna of the Atlantic Coast. It was to be a students’ manual and Taverner had been asked to write the bird section. His contribution would be to show the influence of the birds on marine life of the Atlantic coast. He would be able to include all the line drawings he wished. Writing to Rowan he said that his contribution would be rather more advanced in content than either his Birds of Eastern or Western Canada. It would be rather a large undertaking, and would be likely to keep him in Ottawa for the sum- mer.** However, Taverner managed to visit the seabird breeding colonies on the north shore of the Gulf of St. Lawrence where he took photographs, and a man from the Canadian Government Motion Picture Bureau, Department of Trade and Commerce, took films of the birds. In theory the trip in July should have been successful and enjoyable but in fact it was a failure. Harrison Lewis, the Chief Migratory Bird Warden for Ontario and Quebec, arranged for Taverner to come with Lewis in his boat. Let Taverner explain what went wrong in his own words in a letter to Fleming. “The boat Lewis has is a very nice little cruiser and had she been properly fitted with lockers and stowed would have been very comfortable. Lewis is a good sailor but has no more idea of keeping things ship-shape than he has of collecting. As long as his boat goes and is sea worthy, nothing else is worthy of attention. Had not even pots and pans to cook with nor nearly enough knives, forks, plates, cups or spoons to go round, so we had to eat in relays. Also thinks boiled potatoes with an oxo cube and ships biscuits without tea is an abundant feast. In fact the whole living conditions aboard were impossi- ble. The fact that all this discomfort was unnecessary was what got my goat... 1996 “One thing we did have and that was good fish. Fresh cod is my favorite fish and we had quite a lot besides salmon, trout, and capelin. The latter are delicious. Also had quite a number of lobster. However if we had not bought them from the fishermen we would not have had them. Lewis seemed to think that he was not justified in buying food on the coast when he had outfitted in Quebec. This is not a departmental ruling, but he is a funny fellow indeed.” Taverner ended the letter with the wish that Fleming and his family lived in Ottawa, something that he had said several times in the past: “I do wish you lived in Ottawa instead of Toronto but that is another thing that cannot well be helped.” In order to complete his manual of birds of the Canadian Atlantic Coast Taverner needed to add the common names in general use among fishermen and sportsmen of the marine birds. Lewis had given him the names for the north shore. Now Taverner wrote to Robie Tufts, Chief Migratory Bird Warden for the Maritime Provinces, asking for names of the south shore birds. This was a difficult letter to write because Taverner had heard, through Hoyes Lloyd, that Tufts was “somewhat put out at finding a set of Sharp-tailed Sparrow eggs that he collected in the hands of some American collector”. Since Tufts had presented this rare set of eggs to the museum Taverner had no excuse to offer. He tried to bluff it out by saying that his assistant, Young, had assured him that he had found several Sharp-tailed Sparrow nests in Nova Scotia in 1910 but none had eggs. He made the matter worse by admitting the set had been exchanged with the professional collector, Edward Arnold, for other items. He compounded the injury by saying that he was not much of an egg man, but was satisfied with the material the museum got in exchange.“ But that was not the point. The eggs were rare, the museum did not have another set, Tufts had donated them, and they should not have been exchanged. In his reply Tufts said that he was very sorry to learn that this set, the only one he had ever known to have been taken, had been removed from the Government collection. “Your utter lack of appreciation of these rare eggs was rather galling to me, I confess, and I can hear Mr. Arnold chuckling to himself now as he put this one over on you. However, the set has found its way into the hands of a particular friend of mine who greatly appreci- ates them on account of their rarity as well as because they were collected by myself, and these facts help soothe my injured feelings somewhat.”*> Taverner was still trying to justify his action in his next letter to Tufts when he wrote: “Don’t worry over those Acadian Sparrow [Sharp-tailed Sparrow] eggs. Arnold may have his chuckles but we have ours as well. From Armold and through him we have obtained several hundred sets of eggs besides a number of skins including a pair of Passenger Pigeons, — all without cash cost to us”.*° No amount of explanation could bring back Tufts’ gift, and the episode left him feeling sore. The prob- CRANMER-BYNG: A LIFE WITH BIRDS 127 lem of how decisions of exactly what specimens should be exchanged from a collection of a museum, and who should make the decisions was to arise from time to time in the future, and will be discussed in Chapter 14. A problem which was on Taverner’s mind at this time was the health of his mother. Apparently she had been “going down” for the past two years, but in the spring of 1924 she had her tonsils out. In his over optimistic way in matters of health he reported that she was now her old self again. He also said that he had suffered from “rheumatics” for the past year or so, and was now going to have his teeth out, and added “think it will fix me up again.”*” In July Martha Wiest, an old friend from Detroit, had come to the cottage to be with Taverner’s mother. When she left to fetch her son from a summer camp in mid- August Mrs. Taverner went with her to their home at 45 Leonard Avenue in Ottawa. On September Ist Ottawa friends of the Taverners, Dr. and Mrs. Charles Saunders, who had recently returned from a two year absence in Paris to study the French lan- guage, came to visit them, which Mrs. Taverner enjoyed very much. The next day Percy and his mother, with Martha Wiest and her son Karel, returned to the cottage, while Ida expected to go up the following weekend. They spent a quiet few days at the cottage but then the weather changed and on the morning of September 5th was very windy. Mrs. Taverner had an asthmatic attack and died. Percy described her death in a letter to Fleming. “T never saw her in better health and happier than she was the last week and even up to three hours of her death. However it was as easy a passing as could be imagined and she did not even suspect there was any danger until she lost consciousness. Just two weeks before it happened I managed to get a great lot of unusu- ally satisfactory pictures of her in the garden. We are thankful to have very beautiful memories of our mother. We will miss her greatly and it will throw considerable burden on my sister. We contemplate making no more changes in our living than necessary.” * Taverner also sent the news to Brooks in which he said, in part: “I suppose that you have heard of the death of our dear mother... Ida and I are closing up as best we may but there are not many men of my age that are as close to their mother as I was and we feel her loss sadly. Our consolation is however that we made her happy after a particularly bitter life and her passing was peaceful and even without anticipation.” Brooks wrote with his sympathy, and thanking for the press cutting which he was glad to have. “Your mother”, Brooks said, “spoke so little of her past, she was always so interested in the present, like all big- hearted people are.”° Mrs. Taverner was in her seventy-first year when she died. She was buried in the family plot (Section 50, Lot 63) in Beechwood Cemetery, Rockcliffe, 128 Ottawa.” At that time Percy was 49 and Ida Clare 37 years old. The family of three had remained closely together since Ida’s father had ceased to visit her mother nearly thirty years ago. They had always helped each other, and shared their resources and to some extent their friends. Ida Taverner had worked for the Order of Foresters when they lived in Detroit but when they moved to Ottawa she was able to live in a different style with leisure to develop new inter- ests. As the family found new friends Ida Taverner found herself entertaining interesting people whom Percy invited home. Their Ottawa house was consid- erably larger than the one they had moved from in Detroit, and had a style all of Percy’s own design.” The garden gave his mother a great deal of pleasure during the last years of her life. At the end of 1922 Percy had the porch glazed in, but with a moveable sash so that it could be opened in the summer. As he told Brooks: “It makes a lovely study and extra sit- ting room, half conservancy.”>’ Since 1914 Percy had put much hard work into developing an attrac- tive garden with a lily pool with gold fish and water lilies in it, and a Purple Martin house beside it. A photograph of 1923 shows the Hoyes Lloyds’ three small children by the pool, and behind it a lattice- work fence and ornamental gate with a flower border with tall hollihocks in flower in front, and the martin house seen just behind the gate.* The rock garden, for which the Taverner home became well known did not reach its zenith until the 1930s. Taverner’s idea of a holiday at this period in his life was not of lazing about but rather some activity that was different from his usual work. In one entry in the Hyla visitor’s book Ida noted: “Percy’s ardent desire to be always improving the place spent itself in building a chair with birch legs which mother says is quite beautiful.”°> The same was true of their home at 45 Leonard Avenue where Percy spent his spare time working in the garden, or in the basement making Christmas presents for his close friends. For Christmas 1922 he sent Brooks a little statue of a Gyrfalcon for which he had made a mould twenty years previously. In thanking him Brooks called it a thing of beauty.*° Another present that he regularly gave to his friends was his own Christmas card. First he made a mould of it, then he could “pull” from it the number of cards he required. Another kind of activity was to bind a book as a present. In 1925 and 1926 he was particularly active in his hobby of book binding which he explained in a letter to Laing. “Have been binding all winter”, he wrote, “and my bookcases now look quite imposing with all the paper backs properly covered with leather. It is nice work too and not difficult if you have manual dexter- ity and know how. Would be glad to show you some time.”*’ It was a psychological release from typing letters and examining skins, and gave him a feeling of satisfaction to see the results of his manual skill. As a special present Taverner bound a copy of THE CANADIAN FIELD-NATURALIST Vol. 110 Laing’s account of the expedition he made as muse- um naturalist in the C. G. S. Thiepval in 1924 from Canada to northern Japan including Alaska, the Aleutian and Kurile Islands, and Hokkaido, with some chance of collecting specimens.>* Another friend te receive a book bound by Taverner at this time was Allan Brooks. This book was the result of a collecting expedition to Atlin Lake, British Columbia, during June through August | 1924 undertaken by Brooks and Harry S. Swarth.*° During their stay there they collected ten Brewer’s Sparrows high in the mountains at the timber line. These specimens were sufficiently different to justify describing them as a subspecies of Brewer’s Sparrows. They named it Spizella breweri taverneri in honour of Percy Taverner.© As a result of their studies and collecting of British Columbia birds Brooks and Swarth compiled a provincial list which ‘Taverner had been encouraging Brooks to make ear- lier. It was published in 1925 under the title A Distributional List of the Birds of British Columbia.®! This was the book bound by Taverner as a gift to Brooks. Taverner not only obtained a copy of the provincial bird list for British Columbia which he wanted, he also had a new subspecies named after him. Another aspect of Taverner’s artistic appreciation came into flower in 1925. As a result of his mother’s will he was able to start buying the first of paintings by bird artists. A painting by Brooks at the AOU meeting at Pittsburgh in November 1924 of a Golden Eagle was for sale. Taverner wrote to Brooks asking if it had been sold yet, and how much he was asking. He explained “I am going to buy a few such things but though I do not want to compete with more afflu- ent purchasers I do want some more of yours.”® At the same time Taverner wrote to Fuertes confirming his conversation at Pittsburgh asking him to paint a picture for him on the general lines of his “Geese going south”. The size he left to Fuertes — “the best you can do for one hundred dollars”. Fuertes replied warmly: “Tam greatly delighted; nothing in the whole course of my career pleases me as much as to have my feller [sic] ornithologists order my stuff, because I know it is because they like it for the reason I like to do it, and it is the most honest-to-goodness tribute my work could or ever does get.”°+ The painting was 36 x 24 inches, and arrived safe- ly, getting through customs without any difficulty. Meanwhile Brooks replied that the Golden Eagle had been sold to a man in New York for $120. After further correspondence Brooks sent four paintings for him to choose from. “Take your choice” he wrote. “Eagle $100; Coots $75; Waveys [Snow Goose] $50; Heron”.® Taverner chose the eagle. He liked the Snow Goose next, and the heron last because he considered it “a little too chromolitho graphic, and contained at least three different pic- 1996 -tures”.© In a later letter Brooks added a P.S. “You are quite right, I think, about the Heron picture. Wish [had you to criticize as I make ’em’”.°’ The year 1924 had been a bad one for Percy Taverner. Apart from his mother’s death he had been working very hard all the year with very limited recreation. In contrast 1925 was exciting, both as regards museum studies and field work. Starting from September 1924 correspondence between Taverner and Rowan contained information about Rowan’s early research into the problems of bird migration. But first here is Taverner thanking Rowan for the bird records from Alberta that he kept send- ing him. “Now here you add Pine Warbler and the Dickens knows what. Well, we are finding out something about prairie ornithology now. It takes the resident observers to get down to brass tacks. I see that you have enough to transfer to my files to keep me busy the rest of the after- noon.””°8 Rowan had written to Taverner in early 1924 about Beaverhill Lake in the spring migration, invit- ing him to come. It was an enthusiastic invitation in which he told Taverner that he had learnt a lot from their regular correspondence of the last few years, and had derived lots more pleasure from it than he could well say. ; “Tf I have teased you a bit more than I should from time to time, that’s part of me and can’t be helped. But there are a lot of things one can’t write about that one can dis- cuss, and I need not say what a delightful treat it would be for me if you did no more than spent just a day here.” He explained that the C.N.R. passed through Edmonton, that the last stop before Edmonton was Tofield, and that every train stopped there. All he need do was to get out there. “The taxi man will meet you and take you right to the front door of the tent, 4 1/2 miles for $1.50 and no hotel expenses at the other end. You will then see the finest wader ground in the Dominion.” After telling him all the good birds he was likely to see: Hudsonian Godwits, Buff-breasted and Stilt Sandpipers, Smith’s Longspur, Ross’ Snow Goose [Ross’s Goose], Richardson’s Merlin; he ended by promising: “You will never feel sorry for it, and it will keep me going for another year or two. We get completely starved for decent company.”© Rowan clearly found Taverner a congenial compan- ion, and someone on whom to try out his theories while formulating them. Rowan’s biographer, Dr. Marianne Ainley, referred in her thesis to Taverner as “Rowan’s long-distance mentor”. She also suggested that Taverner’s comments on the reactions of birds to different migratory situations caused Rowan to insert a section in his 1926 paper on this question.” At this time an amateur ornithologist, the Reverand Gustave Eifrig, had had an article pub- lished in The Auk in which he doubted that birds migrating in spring were influenced by any physio- CRANMER-BYNG: A LIFE WITH BIRDS 129 logical stimuli.’' Rowan had been formulating a hypothesis of his own at this time to show that some species did receive external stimulus in the spring. In October 1924 he began experimenting with juncos by giving them additional minutes of artificial light after sunset. He published an account of his first experiments in the English journal Nature in April 1925. From this year onwards his letters to Taverner reported, in some detail and with obvious excite- ment, how his experimental work was progressing. He enclosed a copy of the article in Nature for Taverner and commented: “T believe this bit of work of mine is the first attempt in history to prove any of the migration theories experi- mentally.” Rowan also enclosed a copy of “an onslaught” on Eifrig’s article mainly written in July 1924, and commented “T seem to have jumped hard on Eifrig but really haven’t done so nearly hard enough. To my mind it is an inde- fensible procedure to crib a good idea from a couple of botanists, stick it onto birds, and make it apply without a moment’s thought.”7 At this point Taverner met someone who became another member of the naturalist-collectors of his western network. This was Cyril Harrold, who had come to Canada in 1914 at the age of eighteen. He was saved from joining the forces because of defec- tive eyesight, and instead spent the war years in the dangerous work of manufacturing explosives. All his spare time was spent in studying and collecting birds. After the war he came to Winnipeg with the object of making a career in ornithology. He learned the art of taxidermy, but was far more than a bird collector, and a sound education gained in England with a grounding in Latin, and a knowledge of French and German were useful in his study of ornithology. He was especially interested in the problems of migration and distribution, and was a first-rate field man.” Taverner started his field work for 1925 by a sec- ond expedition along the Red Deer River with Harrold as his assistant. Starting from Red Deer on 20 June they reached Drumheller in July where that part of the trip ended. Taverner was anxious to increase his knowledge of the raptors of that area begun in 1917. New information on distribution, behaviour and relationships was gathered. From Drumheller they went by train to a point near to Charles M. Sternberg’s camp where a group of museum men were digging for fossil remains of dinosaurs. Percy wrote an informative letter to his sister Ida, who was on a trip to Europe with a group, about their Red Deer trip. He described Sternberg’s camp, with its team of horses and a farm wagon as well as Sternberg’s prized tin Lizzie (Model T Ford) which he kept covered by a large tarpaulin — rain or shine. Apparently Sternberg was a bit downhearted at not 130 finding anything of particular interest until he found a really worthwhile specimen. Taverner thought it was a carnivorous dinosaur complete except for its skull. It was in an undivisible mass of over 4000 lbs weight; quite a challenge to extract. Taverner, in contrast, found nests of a Say’s Phoebe, and a Rock Wren which he crated with the plaster moulds he made of whole sections of cliffs to support these nests for exhibits. He told Ida that he had been mak- ing a lot of movies for Sternberg. “If they turn out well”, he said, “they should make a very good pic- ture we can call ‘Hunting Dinosaurs in the Bad Lands of Alberta’.” He also took a lot of stills and movies of birds. Conditions were not entirely com- fortable because of the weather. It was extremely hot, and the steep river banks and cliffs made it sti- fling. One night in camp there was a gale which blew off the awning fly, and then the main fly over the tent which ripped it badly. Harrold and Taverner spent all day sewing it up. Frank Farley joined them for a few days and won all hearts by his enthusiasm and energy helping to get some heavy specimens into both wagon and boat. He ended his letter with a PeS: “Getting along with my teeth, but if it were not for the experience of others would declare that one never could use them in comfort or with efficiency!””4 Taverner left Sternberg’s camp about 20 August (Harrold had left earlier) and went by train to Tofield station and then by car to Rowan’s camp at Francis Point, Beaverhills Lake. Here he met Munro and his wife, who took the taxi back. Rowan described Taverner’s arrival in his field notes as follows: “Taverner turned up about 7 o’clock and Munro and his wife went back on the taxi. He (Taverner) brought his complete equipment with him, some load, but nothing on earth necessary for comfortable collecting appeared to have been forgotten. The tent has a ground cloth which hooks around the bottom and is practically mouse proof. It can also be made mosquito and fly proof; has windows, stove pipe exit, and a door at each end which are weather proof when closed. His collecting trays can be removed from their box and hung from the ridge pole in a fly proof cover. He brought various odds and ends with him including two Great Horned Owl skins, one of which is whiter than I have ever seen although bred down the Red Deer Valley.” 7° Robert Lister, who was one of the group, added “Rowan’s own tent usually had a rope as a substitute for a ridge pole; the walls sagged; it attracted insects and ground squirrels gambolled through it at will.”’”° Thirty years later Rowan looked back on the event during a talk he gave in Calgary with the help of his field notes. He seemed, he told the audience, to have been filled with envy at Taverner’s equipment, the first decent camping outfit that he had ever seen, and a great contrast to what he and Harrold were used to. Rowan mentioned that Taverner’s equipment includ- ed bedstead and air mattress, adequate cooking uten- sils and that nothing was missing. THE CANADIAN FIELD-NATURALIST Vol. 11 \ “On the other hand, Harrold’s bed and mine were merel | blankets on the bare ground, whatever it might happen t be: our tent was borrowed, complete with two majo leaks: our cooking outfit, a single frying pan and a col. fee pot, the latter serving in turn for coffee, tea or stey as occasion demanded, with an erstwhile coal-oil con tainer for drinking water which came out of the lake o the nearest slough. Taverner carried his with him: eve) his drinking water was respectable.””” Taverner gave his impressions of Rowan whe1 writing to Ida: “He is the strangest littke Englishman you ever saw — mercurially enthusiastic and too interested in shore bird) to eat while they are present. In fact he and Harrold — due to this and economy — camp like Siwash rathe than white people, and I was glad that I brought my equipment with me. When I left I donated a lot of my worn out equipment to their camp and added consider. ably to their comfort.”””8 | Taverner spent several days there, then went tc Edmonton to spend a weekend at Rowan’s house. Ir) the same letter he told Ida: “He has a very fine wife that is an admirable correction to his mercurial nature and four dear little children.” From Edmonton he went to stay about a week with “good old Brooks” at Okanagan Landing. In March 1925 Taverner and Anderson were involved in preparations for an expedition to climb, for the first time, to the summit of Mount Logan, the. highest peak in Canada, and to measure, as accurate- ly as possible, its height. The National Museum was asked to suggest a naturalist to accompany the expe- dition, and the name of H. M. Laing was put for- ward. By an agreement with the Alpine Club of Canada Laing was asked to accompany the expedi- tion as naturalist and cinematographer. While he was training for this role Taverner sent him names of birds, some of which the museum particularly want ed to have evidence of as breeding in that region: Wandering Tattler; Surfbird; Sharp-tailed Grouse; Fox Sparrow and Timberline Sparrow [a race of | Brewer’s Sparrow, see note 60 above]. Ten moun- > taineer-scientists with the task of measuring the’ mountain, together with Laing, set off early in May, — reaching the base camp at Trail End at the head of) the Upper Chitina River, the limit of pack-horse travel where Laing remained for the next three ¢ months doing field work in the vicinity of the Chitina moraine. He returned with the climbing party ) in mid August by a partly new route. Six members of the climbing party reached the summit, and the © height of the mountain was estimated to be 19 850 | feet = 6050.28 metres. (Sixty-seven years later, in | June 1992, a larger expedition carried out scientific — work near the summit of Mount Logan and deter- mined its height to be 5959 metres.) When Taverner heard that Laing had arrived home | in late August he went to join him at Comox and | they had a great “talk-feast’” which lasted a number of days. Since Laing was expected to write three 1996 eports on his field work and collecting, it was good ‘that he could first have a talk with Taverner.” For Taverner his field work for 1925 reached an ‘enjoyable climax in his first visit to California. He ‘spent a week studying the ornithological collections ‘at the University of California, Berkeley, and Bishop’s collection at Pasadena, Los Angeles where the saw interesting specimens. He also made a num- ‘ber of personal contacts that were likely to be of value to the museum. While at Pasadena he did some | At the New York meeting of the AOU in ‘November 1925 Taverner and Anderson went, with Bishop’s permission, to visit his series of gulls in his bird house at his home in New Haven, Connecticut. ‘Writing to Bishop about the AOU meeting he noted ‘the lessening influence of the Audubonites and explained, / “So many of the old sentimental extremists have taken museum jobs and attained a better balanced view — Cleves and Harper for instance.” Bishop had a fine collection, just the kind of col- lection that they needed as a foundation for a National Museum. “T certainly wish that such a collection were available for work in cooperation with ours ... If we could only induce you to come to Canada it would be a National benefaction”.*! CRANMER-ByYNG: A LIFE WITH BIRDS ar a a - Beaverhill Lake, Alberta, August 1925. P. A. Taverner and C. G. Harrold. (Reproduced courtesy of the Canadian Museum of Nature, number 66835.) 131 ies Pe It was a nice idea, but nothing more. In addition to working on his book, field expedi- tions and keeping up a regular correspondence, Taverner still took an active part in the Ottawa Field- Naturalists’ Club. Although he was not officially the editor of The Canadian Field-Naturalist in the 1920s he wrote regularly in its issues, gave his opinion on ornithological manuscripts submitted, and edited various contributions. For example, in a letter to Brooks he wrote of the problems that editors of the journal faced. Brooks had found bad errors in a list of western birds printed in the 1925 volume and wrote to Taverner complaining.” In reply Taverner stated the problem faced by editors who knew their own area very well but could not know all other localities in the same way. Unpaid editors were hard to find. The best way would be to send all B.C. material to Brooks for his approval but that would be impracticable. “T acknowledge my slip up especially in the Chipping Sparrow which I did not notice but if you saw all the stuff that I do question and eliminate you might be [less?] surprised at things that do get by. The editor of unpaid material has his hands full’. He suggested that readers should send the editor a note calling attention to mistakes which would make authors more cautious.** Another task he undertook was to write to Camsell, Deputy Minister of Mines, about supporting the application for a government grant of $1000.00 to The Canadian Field-Naturalist. 132 Camsell had promised to do so in conversation. Taverner now set down the reasons for asking for this grant. It is a strongly argued letter which shows how Taverner regarded the role of the journal. There were five affiliated Natural history Societies at this time, he told Camsell: Quebec; Ontario; Manitoba; Alberta and British Columbia.*4 Sometimes Taverner led a bird walk in “south’ Ottawa, for members of the Ottawa Field- Naturalists’ Club. From where he lived on Leonard Avenue he could walk to Dow’s Swamp which lay in a depression between Dow’s Lake and the Rideau River (where part of Carlton University now stands). In the 1920s this area was a tangled swamp three miles from the town east of Bronson Avenue, and approached by Colonel By Drive. By using the C.P. Railway tracks it was possible to cross the Rideau River. Taverner could then walk eastward to Bank Street at Billings bridge and back to Leonard Avenue.*> With all Taverner’s work obtaining information for Birds of Western Canada, and then writing it and reading proofs, much of his time in the years 1920- 1926 must have been taken up. However, he found some time to write to his correspondents in provinces other than those of western Canada. The province of Quebec was not well represented among Taverner’s correspondents although Harrison Lewis, Chief Federal Migratory Bird officer of Ontario and Quebec, kept him informed to some extent. There are a few letters preserved between William Brown and Taverner, and several between Henry Mousley and Taverner. In 1923 a new correspondent from Quebec wrote to him. This was Gustave Langelier, who was Superintendent of the Experimental Station for central Quebec, Department of Agriculture, at Cap Rouge, Quebec City. He said that Mr. Harkin had referred him to Taverner, and asked for help in identifying birds.*° Thus began a correspondence which lasted until 1940. One of the birds that Taverner received was unusual for Quebec. When Taverner looked at the number on the collector’s licence he found that it was in the name of Mme Bernadette Langelier. Taverner wrote direct to her congratulating her on collecting a Yellow-billed Cuckoo.*’ In reply Mr. Langelier explained that he and his wife had started a small collection in 1922. He also invited Taverner to stay at Cap Rouge as his guest. In October 1923 Taverner wrote to Langelier telling him that he had voted on his name as an Associate Member of the AOU. These annual meet- ings, Taverner explained, “are very enjoyable and more like a reunion of friends than the usual dry sci- entific meeting. It is astonishing what an active inter- est is growing in bird banding. Some of the results being obtained are of immense interest and impor- tance.”** Correspondence between the Langeliers and Taverner continued whenever they had birds THE CANADIAN FIELD-NATURALIST Vol. 110 they wanted identifying. By the end of 1924 they had over 500 mounted specimens in their collection. Meanwhile Mme Langelier was making a collection of skins and by December 1925 had made over 700. Some skins they exchanged with Bent, others they sent as far away as California and Alaska. Taverner was glad to hear of the progress they were making, and stressed that for study work skins were a neces- sity. He mailed them a copy of his “Instructions for Collectors”, and said he would like to visit them and see their collection some time.*? One other ornithologist with whom Taverner kept up a steady correspondence was Arthur Cleveland Bent. Their letters were full of ornithological infor- mation and need to be read as complete letters in their proper sequence. Anyone doing so will see how much help Taverner gave Bent during the 1920s. Writing to Bent in 1922 Taverner said that he had a good deal of information at the museum but the problem was how to get it to him, so he invited Bent to come to Ottawa and see the information for him- self. “I always have a room at your disposal and should be glad to act as your host.’?? Of course this correspondence was helpful to Taverner who was part of Bent’s wide network of ornithologists. Taverner was kept informed of what ornithologists “on the other side of the line” were doing, particular- ly in taxonomy, even if he did not always agree with their findings. Bent asked Taverner to read sections of his Life Histories at this time. Writing in March | 1922 he said, of a recent letter from Taverner, that it — “impresses on my mind the fallibility of human nature and how difficult it is to try to avoid errors in © a comprehensive work like mine.””! By early 1926 there was nothing more that — Taverner needed to do for The Birds of Western Canada as it was being printed, so he left Ottawa in mid May on a field expedition to Belvedere, Alberta, about 60 miles north of Edmonton. He was joined there by Laing and Harrold. These three experienced observers studied two adjacent lakes — La Nonne and Majeau.”” They left camp in mid-June for Beaverhills Lake where Rowan and a group of his helpers were preparing to band a large number of Franklin’s Gulls. By 6 a.m. on 24 June thirteen bird- ers went into the vegetation around the lake where | the colony was nesting to start banding. The noise ° was deafening. By 9 a.m. the banders returned to camp for refreshments, and a rest, having used about 3000 of the 10 000 bands that Rowan had with him. . By 10 a.m. they were at work again but by midday they could hardly stand up so they returned to camp. The total banded was a few hundred short of 5000. Taverner, Laing and Harrold stayed in camp for a second day to bring the total bands used to 5000. | Whether this heroic effort was really wise is open to argument. At least it hit the headlines in The Edmonton Journal.” { 1996 | | | | / / : i While Taverner was in Alberta collecting and banding, the printing of Birds of Western Canada continued. Anderson wrote to Taverner in June with _the reassuring news that the job was nearly finished. “Mr. Miles is on the job every day and is keeping the bureau up to the mark. There is only one more plate ‘to run, and then he says they will send a bunch to the ' bindery, and the magnum opus will be out.”4 During the early months of 1926 Taverner, with ‘his inventive mind and nimble hands, was preparing several “stunts” for exhibiting during the AOU meet- ing in order to lighten a little the solemnness of the scientific papers.* One such “stunt” was making a small plaster auk, about 3 inches high, and inscribed CRANMER-BYNG: A LIFE WITH BIRDS 133 “Ottawa 1926”, copies of which could be made in a mould. One of these could be placed in each mem- ber’s place at the banquet during the AOU meeting. In contrast Taverner projected another “stunt” which was to make a model of a Great Auk, of twice life size, for display. When the time came for the chal- lenge of the AOU meeting Taverner was reasonably well prepared.” *<“Stunt” in the sense, not of circus stunts to thrill audi- ences, but rather in the sense of an act showing boldness of invention together with manual skill. CHAPTER 13. Meeting of the American Ornithologists’ Union, Ottawa 1926 | _ “For the first time in its history the American _ Ornithologists’ Union met outside the United States and | the forty-fourth stated meeting in Ottawa, Canada, in 1926, was a decided departure in several respects from | any of its predecessors. Through the invitation extended | by the Minister of Mines and Interior the meeting was made a semi-official affair and numerous courtesies | were extended to visiting members by the Canadian | Government. Special arrangements were made through the Immigration Department to facilitate crossing the _ border, the Museum where the meetings were held was | made a temporary bonded warehouse thus permitting receipt and reshipment of exhibit material with a mini- _ mum of tariff restriction, and the general session was / opened by an address of welcome by the Minister of Mines and Interior who also held a public reception for the members and attended the annual dinner.” T.S. Palmer, Secretary of the AOU! Much effort went into the organization and day to day running of the meeting. The committee of the AOU in charge of local arrangements consisted of: iP. A. Taverner, Chairman; Hoyes Lloyd, Secretary; R.M. Anderson. This committee kept in close con- ‘tact with the Ottawa Field-Naturalists’ Club Com- ‘mittee chaired by Clyde Patch, Ist Vice-President of ‘the Ottawa Field-Naturalists’ Club. This committee ‘was appointed by the Club to deal with financial and other arrangements for entertaining the guests. It included representatives from various provinces ‘across Canada.” Clyde Patch, as chairman of the ‘committee, informed all Canadian members of the AOU early in 1926 of the arrangements, and encour- aged them to be present. He also wrote to many ‘other Canadians, requesting funds so that the local ‘committee would be able to put on a really good ‘Meeting. The meeting began on Monday 11 October 1926 with business sessions of the Council in the elegant banquet room of the Chateau Laurier hotel. A meet- ing of Fellows and Members was held there in the evening at which reports were given to provide those present with information on the state of the American Ornithologists’ Union which appeared to be sound, judging by its membership which stood at one thousand eight hundred and fifteen. The follow- ing officers were elected for 1927: President, Alexander Wetmore; vice-presidents, Joseph Grinnell and J.H. Fleming; secretary, T.S. Palmer; treasurer, W.L. McAtee; Members of the Council: A.C. Bent, Ruthven Deane, E.H. Forbush, H.C. Oberholser, W.H. Osgood, C.W. Richmond and T.S. Roberts.* Taverner was on good terms with sev- eral of them and well known to all. The appointment of J.H. Fleming as a Vice-President was a step for- ward for Canadian representation in the running of the AOU. Also, from the list of Associates, two Canadians, W.H. Mousley and J.A. Munro, were elected members. The general business included authorization to continue work on the “Ten Year Index of the Auk” and on the “Check-list of the North American Birds”. Resolutions adopted included one opposing the general destruction of birds of prey which stated: “RESOLVED, That the American Ornithologists’ Union deplores the present tendency to wantonly destroy birds of prey as more likely to result in ultimate economic loss than in gain; and also deplores the use of the word ‘ver- min’ for these birds as tending to produce an unwarrant- ed prejudice.’4 The presentation of scientific papers occupied October 12, 13 and 14. These were held in the National Museum and were open to the public. “On entering the Museum on Tuesday morning members found on the landing of the main stairway, near the registration desk, a model of a Great Auk of heroic size, resting on three volumes of the Union: “The Auk”, “The Code”, and “The Check-list”.°> These were modeled by P.A. Taverner. On registering, each member was presented with a copy of Taverner’s handsomely illustrated and finely printed work Birds of Western Canada, provided courtesy of the Canadian Government, as a souvenir of the occasion. 134 The general sessions were opened by the Hon. Charles Stewart, Minister of Mines and Interior. A total of fifty-seven papers were given ranging from a few of 45 minutes to many of ten or fifteen minutes. One of special interest to Canadian ornithologists must have been by Edward Preble of the Biological Survey, Washington D.C. on “Canadian Field Ornithology, 1750-1900”. J. Dewey Soper, recently back after two winters in the arctic, made a brief report on his ornithological work in Baffin Island. Henry Mousley, of Montreal, spoke about his study of the home life of the Northern Parula [Warbler]; Frank Chapman gave an illustrated talk on “An Ornithological Reconnaissance in Venezuela”; F.C. Lincoln of the Biological Survey, Washington D.C. gave an illustrated talk on “The Migration of the North American Herring Gulls”. Ernest Thompson Seton attended the meeting and spoke on two topics with intriguing titles: “Conservation Gone Mad” (15 min); “Bird songs enjoyed by other species” (with lantern slides); R.E. DeLury, of Ottawa, spoke on “Banding redpolls”; Clyde Todd spoke on “The 1926 Hudson Bay Expedition” and Prentiss Baldwin of the Baldwin Research Laboratory, Cleveland, Ohio, spoke on “The sport of bird banding”. It would have been interesting to have heard how he defined bird banding as “a sport”. © While all this was in progress several exhibitions were on show. Preble’s talk was supplemented by an exhibit of the more important publications on Canadian birds, illustrated by portraits of the authors which were loaned from the Ruthven Deane collec- tion of photographs of members of the AOU.’ Several exhibits of paintings, photographs and publi- cations were installed in the halls of the museum. Of these the most important were the exhibits of bird paintings. The first formal exhibit of bird paintings at an AOU meeting was held at Washington in 1920. The catalogue of paintings and photographs at the Ottawa meeting included 444 entries representing approximately fifty artists. Thirteen artists showed photographs of birds. The exhibition contained sev- eral special displays in addition to the main one. A special one was the Robert Ridgway Exhibit contain- ing drawings, manuscripts and letters illustrating his development as an ornithologist between 1850 and 1880.8 Two special collections were devoted to bird paintings by Allan Brooks. One was of the illustra- tions made for Taverner’s Birds of Western Canada, a total of 96 originals.? The other was a collection of 32 bird paintings commissioned by Wallace Havelock Robb.!° Brooks was also represented by five water-colour paintings in the main exhibition. Another special collection was loaned by the Emma Shearer Wood Library of Ornithology, McGill University Library, Montreal. This was mainly of water-colours by European artists in the nineteenth century.!! THE CANADIAN FIELD-NATURALIST Vol. 110 The names of other well-known bird artists of that time represented were: George Lodge; Robert Bruce Horsfall; Louis Agassiz Fuertes; Charles Livingston Bull; F.L. Jaques (4 scenes on the coast of Peru). Two artists who were to make their reputation later — George Miksch Sutton (4) and Roger Tory Peterson (2 in charcoal), also exhibited. Frank Hennessey, of Ottawa, showed four paintings in oils. Claude Johnson, of the National Museum, showed seven water-colours, five being of soft parts of birds, and an oil painting of a loon. Among those exhibit- ing photographs, Mousley of Montreal showed 5 of Marsh Hawks [Northern Harrier] and 3 of [American] Woodcock; Edward Warren of Colorado Springs, 6 of a White-tailed Ptarmigan, among oth- ers. Finally, P. A. Taverner’s contribution was five photographs, field notes of colours of soft parts of birds (water-colour), and a statuette of a black Gyrfalcon. Those attending the AOU meeting cer- tainly had plenty to interest them visually. The social gatherings were more varied than usual and included a conversazione* on the first evening and the annual dinner the next evening. Informal receptions were held at the homes of R.M. Anderson, Harrison Lewis, Hoyes Lloyd, P. A. Taverner and George White on the Thursday evening which gave AOU members an opportunity to meet each other. The conversazione was organized by the Ottawa Field-Naturalists’ Club to entertain the visiting mem- bers of the AOU and was held in the museum on Tuesday evening. Hoyes Lloyd reported: “The guests were received by the Honourable Chas. Stewart, Minister of Mines, and Mrs. Stewart, Dr. Chas. Camsell, Deputy Minister of Mines, and Mrs. Camsell, Dr. W.H. Collins, Director of the Museum, and Mrs. Collins, and by Mr. C.L. Patch, Ist Vice-President of the Ottawa Field-Naturalists’ Club, and Mrs. Patch. A fea- ture of this entertainment, which appealed to the audi- ence, was the demonstration of the cleaning of eider- down by Mr. Harrison Lewis. Motion pictures were shown by the Canadian National Parks. The evening closed with orchestral music and refreshments.”’!* T. S. Palmer reported on the films shown during the meeting: “The moving pictures covered a wide diversity of sub- jects and, including those shown at the conversazione, numbered more than twenty reels. The most remarkable were Gromme’s unique views of [Common] Loons taken in Wisconsin and the scenes in the National Parks presented through the courtesy of the Canadian Parks Service. Lewis’ pictures illustrated some of the econom- ic phases of Eider Duck conservation, while Cordier’s close ups of the Water Ouzel [American Dipper] revived the discussion regarding the nictitating membrane of this bird.”! *in normal meaning of the term: “a meeting of a partly social and partly scientific or artistic kind”. 135 : A LIFE WITH BIRDS CRANMER-BYNG 1996 “BI]ODS BAON ‘OT[IAJ[OAA ‘SOOUTAOIG DUNE] UY} JOJ JOJO pl ATO WwISIPY [RIOPIY JOTYO ‘SYNL “MY ‘Sp ‘VNOIS BAON ‘OPIAJ[OM ‘SUNT “MA A SIAL “Lp ‘epeuen ‘emeno ‘wnasnypy [BLOW] PIIOIIA ‘JSISO[O}adIaH pur ISIWUApIXe] JoryD ‘yoeg “JT 'O ‘Op ‘owewG ‘owoIoY ‘ASoO[OOZ, Jo winasnyy O1IeIUC [eAoY “JapuUAS “J “J “Gp foLeIUG ‘oUOIOT, ‘ABZO[OO7, jo umnasny] OLeIUC [RAOY ‘puowA “Y ‘f ‘pp ‘oUeIUO ‘oWWOIOL ‘SuNOX “_@ “M “Ep :daqGangd ‘[eanuo] ‘ddeVID op sue ION ‘oNUdAY PIeNOIH PLZ ‘Aapsnoyy AIUD “WIAA “Zh ‘saqand) ‘[eanuoyy ‘skempley [eUOneN] URIpeURD ‘plouly pleMpy ‘[p ‘elTeNsny ‘purlsusand ‘oueqsiig ‘Jopuexa[y “{ PAYTIM ‘Op {OleIUC ‘eMENG ‘[[MUIWURH Lf “SIN “6E ‘eIqUINnjOD YsHig “sulpueyT ueseuryoO ‘syoorg uepy Jolepy ‘gE ‘oaqgand ‘feanuoyy ‘Aydinyy osmno7T ‘sal “LE ‘9aqond) “Woquie'y] 1S ‘onusay AURIS pH ‘T[LIAL “PIA “T “9¢ “Od ‘UNPIOA ‘[eanuUOy Jo yueg ‘yITUIS JaIdeN “CE ‘oeIUG ‘UOITUIRH ‘199 PUOSAS ISAM OG ‘ULUUILLIAJ] PPT SSIPY “PE ‘OLIeIUGE ‘UOIMUL] ‘oNUdAY OLIEIUGC gy ‘NouTy “D “HW "Id “€€ {OLeIUGC ‘UOIIWILH ‘199.19$ PUOIIS ISI OG ‘URUUILLIDPY UIMGO Waqoy “ZE ‘oLeUG_ ‘adoy og ‘uleyNRY 4 OD TE fOuRIU_ ‘AT[IAI]Og ‘qo “HM ‘OE Sepeurn ‘emeng ‘UINISNY] [PLIOWIIJA[ LIOJOIA “ISISO[OUWUIQ, ‘IOUIOARL “VW ‘d “67 ‘epeuRD ‘eMENO ‘UWNaSN [PIIOW I] VIIOJOIA ‘IOTUIOAR TL, BP] SSIP QZ ‘epeueD ‘emMeno ‘Yyourig syieg [euOHeN uvipeury ‘Aoyeool ‘WW ‘d SSUA “LZ {OHeIU_C ‘OOIOL ‘peroy swWjpoysny {97 ‘SUIWIA, “Hf “OT SepeuRD ‘eMLNGC “IOLIA}UT oY} Jo JUUAIedag ‘YyoueIg syieg jeUOTeN uvIpeury ‘UONd9}01g IIT PIM “Jostasodng ‘pAoyy sodop “SZ LOLeIUC “BMPNGE ‘1920S UddNC) OOP ‘PAOTT SeAOH ‘SAP “PZ {9aqGang ‘TINH ‘onusay syooiqsays [¢ ‘astyoursyeT] “TA CW SUA EZ ((laiseg [Hun ssorppe) o1eU_C ‘dR][g SULURD ‘SuNOX ‘fC pudataady “ZZ WoNoIUUOD ‘YdIMUdaID ‘UoJag UOSdWIOY,L, SoU “[Z ‘oleIU_C ‘UO\ssuUTy ‘Ysnpy ‘Ed “V ‘07 :oleIUYC ‘UOIsSUTY ‘YSNPY “_ “V ‘SII 6] ‘epeurd ‘emenoO ‘wWnasny] [BLIOWAPY PIIOIOIA ‘ASO[OIG JO UOISIAIG ‘JalyD ‘UOsIOpuYy ‘WY Jd 8] ‘epeued ‘emeno ‘UOSIOPUY “WY ‘SIAL “LI ‘epeued ‘emeyo ‘Uswo MA Jo [louNoD [euONneN ‘WopIsolg “UOSTIAA “VW “ff ‘SIA ‘OT ‘o0qGond ‘[eonUOy ‘WoqqrH ploleHy “say “ST ‘epeueg ‘emeno ‘SaqonQ) pue OLILIUGC JOJ JIdjJJO plrg AJowAS I, [elopoy JoryO ‘SIMO “4 UOSTIeY “pl LOleUG ‘“SInqsyIe[O ‘UIE Ysieuasvog ‘YsIe WPA Ssy “E] ‘epeueg ‘emeno ‘soul JO JuoUNIedag ‘uINTIeqIoY [PUOHeN ‘IsIURIOg JoryD ‘PIN “OM IC ‘ZI {oleIUGE ‘UOpuO’T ‘siapunes “q “M “LT ‘epeueg ‘eMengo ‘Winasny] [eLIOWIIA] PLIOIOIA, ‘radog Aamo ‘f ‘OI ‘ouRUO ‘eMeNC ‘anUsAY JOUR OEE ‘AIN TO “AY 6 {/OULIUE ‘“VMLNC 1980S pose [ZE ‘Joane “YW ‘_ “8 ‘oLIeUG_ ‘Uo}s3ury ‘aidneag UIMpY "L Seqoyueypy ‘sodruury ‘Adporg “] ‘OD ‘SIA ‘9 ‘eqoyurpy ‘sodruutpy ‘Adpoig “J ‘O “¢ ‘ueMoyoeyses ‘puajseq “Ianog ‘gq “J ‘p ‘eIquInjoD yYsiig ‘Surpuey] ueseuryy ‘orunjy "VL SIA ‘€ SeIquinjoD YsHiig ‘surpur’] ueseuvyKC ‘SdoUIAOIg UIDSIM JO} JOJO pig Aros] [Pope Jory ‘omunyy “VW ‘f ‘Z ‘epeued ‘emeng ‘sourlpy Jo Jaysturpy Ajndaq ‘TTasueD sopeyD “[ YS sy) 0} Yo] 9) WIZ SuIpedy “OLIRIUD ‘eMC ‘Sunda. poleis [enuuY 976] UOIUL] SISISO[OYyIUIQ UvoLIOUTY ay) Je Udye) YdesIso}oyg dnoiy ueIpeury 136 As arecord of the meeting two group photographs were taken, one of the entire membership, and the other of the Canadian members. !* The annual dinner at the Chateau Laurier Hotel was attended by about 180 guests. At each place at the tables a miniature of the Great Auk was to be found, a sign of Taverner’s handiwork. After dinner the evening was given to entertainment. “Newsboys” distributed copies of the annual publication The Auklet, with its usual satirical parodies on recent AOU happenings, controversies and topical events, especially with a local flavour according to the city in which the current meeting was being held. Here are two items from the 1926 Auklet. One is from the section headed ‘Naturalists’ guide to Ottawa”: “Visitors wishing to see large flocks of ptarmigan should take the Bank Street car to the end of the line and walk four or five miles in any direction early in the morning. If no ptarmigan are seen, the visitor will realize that the excursion should have been made at an earlier hour.” > Another spoof centred round the problem of pub- lishing any account of Canadian birds in both English and French. In The Auklet there was a note on the Wood Duck in parallel columns. The left col- umn was headed “Histoire Naturelle Officielle Pour les Provinces”. Under this was the statement “Bilingues redigee dans les deux langues par les pro- fesseurs de la faculte du college de St. Pingouinelle pour les Sourds-muets”. The right column was head- ed “Official Natural History for the French Canadian Provinces”. Under this it stated “Done Bilingually by Professors of the Faculty of the College of St. Aukie the Dumb”. The title THE Woop Duck follows with a short column of nonsense on the species. The left side column, also nonsense, is headed LE CANARD HupPe.* For the benefit of readers with a knowledge of French it is quoted in full. Il est aussi appele branchu et en anglais “Wood Duck” c.a.d. canard de bois, ce qui est la meme chose, car qu’est-ce que c’est que des branches sinon du bois. Cependant, son nom de huppee indique que c’est an aris- to: n’a-t-il pas un sein de cuivre, et des pattes d’or?, n’est il pas tout ruisselant d’ argent? Il est toutefois, toute du bois. Quelquefois appele la fiancee, mais le galant homme, naturellement, n’a pas une seule brassiere. C’ est de lui sans doute qu’il s’agit dans la chanson: Le fils du roi s’en va chassant Avec son grand fusil d’argent et plus loin — Par ses yeux sortent des diamants Et par son bec I’or et l’argent.” !° It was also acceptable to make fun of individual members, though gently, by exposing their foibles or enthusiasms. For example a Note in The Auklet said: “Members are urgently requested not to mention dahlias, irises or gladioli in the presence of W.E. Saunders dur- ing sessions as it is desired that he concentrate on birds during this meeting.”!” *posh, classy THE CANADIAN FIELD-NATURALIST A whole “genus” of members could be the subject | of fun even when the claim was completely untrue, | provided it was meant as a joke. For instance: “A feature of the meeting was the presentation to each | registered guest of a copy of Taverner’s ‘Birds of | Western Canada’. On taking these out of the country, the | ungrateful Yankees, to a man, declared them for customs | purposes as government reports of no value”. During the after dinner part of the program W. H. | Robb of Belleville, Ontario, presented Allan Brooks | with a gold medal in an eloquent address praising his | work as a bird artist. This referred in particular to the | thirty-two water colour pictures of Canadian birds on | display, and called “The Wallace Havelock Robb | collection”. An explanation of this collection from | the point of view of Brooks and Taverner, as shown in their correspondence, follows. In the summer of 1923 Brooks had written to Taverner asking for his advice as follows. “What do you know about Wallace H. Robb? He has written me a long effusion telling me he is going to devote his life and poetry to the cause of bird protection ‘after 1926’, and demanding that I supply him with bird pictures for the honor of Canada at a reduced price. Is he one of those sloppy, slushy, anti-sportsman types? Please tell me so I can guide myself in my reply.”!® In one paragraph of a letter to Brooks about various bird matters Taverner dismissed Robb as a sentimen- talist as regards nature. He wrote: “Yes, I know all about Robb, he is a sentimentalist with a very exalted opinion of himself. Much ego in his cos- mos. He is one of our thorns. Father is something very big in the G.T.R. [Grand Trunk Railway] and with the Government. He wants to turn the Naturalist into a Sunday-school leaflet affair with lots of ‘pep’. Has an uncomfortable amount of influence and hurts our badly needed subscription list.”!? Early in 1925 Brooks wrote to Taverner enclosing a letter he had received from Robb which drew a typical Taverner comment. Robb, he said, wanted to establish “a bigger and better and finer Bird-lore in Canada and run it in his own way. I think he likes jazz in bird literature. He is hopeless. I think it a case of more sail than ballast.”?? After the AOU meeting Robb wrote to Taverner sending the official copy of his address of presentation to Brooks, but complain- ing about the the derogatory style of the issue of The Auklet handed round at the dinner.”! Taverner replied in such a way as not to hurt Robb’s dignity but at the same time not to over praise him. After all, the paint- ings were by Brooks; Robb merely bought them. The paragraph that Taverner included on Brooks says something about Taverner’s sense of values as well. It is reproduced here in full. “Dear Mr. Robb; All is fair prey to the “Auklet.’ Its main duty is to pick out all our pet foibles and hold them up to public deri- sion. It is good for our souls and amusement and warn- ing to our friends. Am glad to get the official copy of the address of pre- sentation to Brooks. It made him squirm a bit for he Vol. 110 )} 1996 __ greatly dislikes public praise though he is human enough to appreciate the appreciation of friends and after the presentation was over am sure that he was gratified at the recognition while the publicity was good for his soul as well. Brooks is a very great man as well as an artist and ornithologist. He hates fuss or fullsome praise and is modest in the extreme. He has very high ideals and never compromises with them and he expects as much from his friends. He is very thorough in everything he does but has no patience with others who are less consci- entious. A perfect gentleman and genial with those he likes he can hold off others with a polished urbanity that is impassable. Your pictures have been taken down from the Art Gallery as the space was required for other material. The agreement was but for the week. I am holding them for your orders. The printed cards will be packed with them for your further use. Let me know where and when to send them. I think they were a great success. Their excellencies viewed them Monday and were much pleased with them. Te ce tg a A Ee SS Cc Sincerely, P. A. Taverner”2 | On Friday morning about sixty people, led by | Hoyes Lloyd and Clyde Patch, went on an excursion _ by motor bus to Kingsmere in the Gatineau Hills. On ‘arrival the majority of the members walked to the top of King’s Mountain to enjoy views of the /autumn foliage from the summit. The remainder of the group visited the grounds of the summer home of | the Right Honourable W. L. Mackenzie King, Prime | Minister of Canada.” On the return of the main party \ to the foot of the hill an al fresco lunch was provided of hot dogs, rolls and coffee.** In the afternoon a | party of eight left on foot for Ottawa, a matter of “some 14 km, and reported seeing eighteen species of birds. The main group walked by a pleasant route to ‘the village of Chelsea where they were met by bus for the return trip to Ottawa. The highlight of the outing was the fall colours which were at their peak in mid-October. During the weekend of 16-17 October thirty-nine people went on an excursion to Blue Sea Lake. They travelled northward along the Gatineau River by a single track railroad to Messines, in the province of Quebec. From there they were transported by boats _to Big Island in the lake. The members were billeted |in cottages belonging to the Andersons, Taverners, | Patches and Miss Maude Scott.*> Taverner referred to the excursion in his racy way in several letters to friends who had not come to the AOU meeting. _ Writing to Laing he noted: __ “Were some crowded but all good natured and happy. It was quite an experience to stroll along the trails of the | island and meet famous ornithologists at every other ”° 2 turn eo Writing to Ernest Thompson Seton he said “Our good luck and good weather held out over the weekend, and about 35 people went on the Blue Sea Lake excursion. They saw Bald Eagle, Surf Scoter, | 4 CRANMER-BYNG: A LIFE WITH BIRDS 37 Hooded Merganser, [American] Pipits, Three-toed Woodpecker etc. and went home happy, as far as we could judge.”?’ With the return of the Blue Sea Lake contingent to Ottawa the 44th meeting of the AOU came to an end. Secretary T.S. Palmer concluded his report of the meeting for Zhe Auk with an analysis of its atten- dance. A few facts presented by Palmer may still be of interest today. A total of 135 members attended the Ottawa meeting. Among these were: one of the original Founders, Charles F. Batchelder, and three Fellows elected at the first meeting: Ruthven Deane, Jonathan Dwight and W.E. Saunders. One corre- sponding Fellow from Australia, W.B. Alexander, attended. Fourteen natural history museums were represented by one or more of their members. Here followed the names of 22 Fellows (and one corre- sponding) and the 24 members present. The names of Associates were due to be published in the annual Directory of Members in The Auk for April 1927.78 Taverner, since he played a major part in organiz- ing the meeting and provided much of the high spir- ited fun (“stunts”) will be given a chance to sum up the proceedings by quoting from his letters to several friends. To Mack Laing he wrote: “The great Meeting is over and all declare it was one of the most successful ones ever held. I know we put over some stunts that have never been attempted before and everything went without a hitch. I only heard two short papers read but had the time of my young life in spite of that. The Exhibition is certainly the best the A.O.U. ever held. Conditions of display were not ideal as our facili- ties for hanging were limited but we had a broader repre- sentation than ever before,- Millais and Lodge and Wolf and Gronvold were represented from England and Larsen and Lillejefors from the Continent besides a quaint and historical collection from all over the world from the Casey Wood collection at McGill. The conversazione was quite a brilliant affair. The Minister of Mines and the Interior and his lady received and we had real music and dancing something the A.O.U. never saw before. On the landing of the main stairs we had a great big Auk modeled in plaster sitting on a pile of volumes representing the Auk, the Code of Nomenclature and the Check List, this was surrounded with palms and backed with a trophy of British, French and American flags. Also a smaller Auklet standing on the work and scolding was a secondary feature here. These aroused considerable interest and approval. We served luncheon daily in the museum with the aid of the ladies of one of the churches not a commercial caterer.”?? What made the greatest hit, Taverner said, was to present every bona fide registrant with a copy of Birds of Western Canada. Another of Taverner’s friends was unable to come to the meeting — William Rowan. This was espe- cially unfortunate because Rowan was one of the very few professionally trained zoologists working in Canada at this time, and the only one with a grow- ing reputation based on scientific research. His pres- ence at the meeting would have interested a number of the ornithologists from the U.S., as well as being 138 of considerable value to Rowan himself. It was iron- ic that one of Canada’s foremost ornithologists was unable to attend the first Canadian meeting of the AOU. This was partly because of his teaching com- mitments at the University of Alberta. He expressed his feelings in a letter to Fleming when he wrote, “Nothing that has ever happened has been so disap- pointing to me as to have to stay in Edmonton whilst the AOU is meeting in Ottawa.”*? Taverner, also, was very sorry that Rowan could not attend the AOU meeting and said so in a letter to him: “Of course was very sorry that you could not be here to enjoy all the fun and profit of the A.O.U. meeting but as we did not expect you was not disappointed. The whole affair went off very well and in a manner that was a credit to Canada and the Canadian A.O.U.s ... I object to your applying the name ‘Yankee Pie’ to the A.O.U. That Yankees outnumber Canadians in its ranks of all classes is but natural considering the relative size of the source of the two divisions. No such body could be less narrow than they are. That most of the leading ornithologists hail from the other side is because we have not raised them but they recognize talent from over here as quickly as from there. I don’t like this subtle innuendo back and forth across the line, — it is a sign of inferiority com- plex and provincialism.”>! To Louis Bishop, who also could not attend the meeting, Taverner wrote a long letter, beginning with two lines from Rudyard Kipling’s poem, Recessional (verse 2), presumably for dramatic effect. “Dear Bishop: The tumult and the shouting dies, The captains and the kings depart and I am trying to settle down to the hum drum routine again with but indifferent success. Entertaining an A.O.U. meeting is rather heady business, — but a barrel of fun. All you mentioned were here and lots more, in fact we had a large meeting in point of numbers and many were good enough to say that it was the best meet- ing ever, — but the last one is always that any way.”*? Most of his letter was taken up with ornithological questions raised at the AOU meeting. Taverner had been working for a considerable time on a study of the Red-tailed Hawk, including two forms recog- nized in the Check-list; Krider’s and Harlan’s Hawks. When Swarth read his paper at the meeting on “Evidence of the systematic status of the Harlan Hawk” Taverner told Bishop “I was forced to take issue with him.” Even after the talk, when Taverner showed his specimens of Harlan’s Hawks, Swarth showed little or no interest in them. Although Taverner was displeased by Swarth’s paper on Harlan’s Hawk he had no need to worry about the reception that Birds of Western Canada would receive from reviewers. What criticisms that reviews contained were relatively minor compared with the praise with which it was welcomed. In his review in The Condor Swarth explained that the book was designed as a popular manual, to give THE CANADIAN FIELD-NATURALIST Vol. 110 information to an intelligent public, but one knowing little of the technicalities of ornithology. “This pur- pose in the main has been excellently carried out ...” However, as Taverner explained in his introduction, the scientific name used was always binomial, while subspecies were not given a formal title and, if men- tioned, received only a subordinate paragraph at the end of the entry for each species. Swarth then gave an example: “As examples of questionable treatment, the author’s adherence to the rulings of the A.O.U. Committee on nomenclature obliges him to lump under single heads such widely diverse birds as the Slate-colored and Oregon Juncos [Dark-eyed Juncos], the several White- crowned Sparrows, and the equally diverse subspecies of Fox Sparrows and of Song Sparrows.” Swarth argued that individual scientists may regard “degree of difference” as a criterion for the separation of species but the lay observer who turns to a popular manual for information will be puzzled. Some of us, he said, who believe that the Juncos mentioned “are two distinct species, will sympa- thize with him”. Having made his point, Swarth continued: “However, I am by no means inclined to quarrel with the author in his manner of meeting the difficulties of nomenclature; quite the contrary, for in such a publica- tion as this subspecies should unquestionably be accord- ed a very subordinate position. Mr. Taverner may in fact, in this instance, ride his own particular hobby with little danger of colliding with any one. If he is in fault at all in the details mentioned — a debatable question — it lies in his attempt to follow consistently the rulings of some one accepted authority.” Swarth praised the large number of coloured plates, mostly by Allan Brooks. “The artist’s skill has brought out the salient points to be emphasized, and as the engraver, too, has done his work well, these illustrations are all that could be desired.” Swarth concluded his review by praising the vol- ume as a whole. “Too much can not be said in praise of a governmental policy that places a volume like this within reach of every one interested in the subject. It is a book for prac- | tical use, and it is also a beautiful volume, to be cher- ished in any library.”*? Witmer Stone, past president of the- AOU, reviewed the book in The Auk. After mentioning the many line cuts of heads, feet, wings etc. by Taverner, and very useful silhouettes of hawks and other birds of the air as they appeared from below, and the coloured plates, Stone wrote: “Mr. Taverner is to be congratulated upon doing for west Canada what he had already done for the eastern provinces and doing it still better. The work will enable western students to familiarize themselves with the bird life of their region and will result in the development of many an ornithologist for the future ... Mr. Taverner, as in his previous works, strives to sup- press the subspecies as much as possible and we find in the brief mention of the subspecies which is appended to the account of the species, again and again such state- 1996 CRANMER-BYNG: A LIFE WITH BIRDS 139 A professionally taken portrait of Percy Taverner framed with the words “Sincerely P. A. Taverner taken about 1927”. Percy had recently discovered that a Mrs. Fowler was living in Los Angeles, She was an elderly widow of someone named Fowler who might turn out to be Percy’s deceased father, hence the gift of his photograph to Mrs. Fowler. (National Archives of Canada, Accession number 1984-178 Hoyes Lloyd collection.) | 140 ments as: ‘the distinction between the two forms is too fine for general recognition’ or ‘the differences are so slight as to be of little popular interest.’ We quite agree with the advisibility of supressing the subspecies in such a work as Mr. Taverner has written, in the majority of cases, because as he says they are too finely drawn to concern the general public. But we must not lose sight of the fact that subspecies are not based upon degree of dif- ference but upon the criterion of intergradation and that there are many subspecies quite as distinct as many species. Indeed some of the subspecies of Song Sparrows are far more easily distinguished than are the small Flycatchers to which full recognition is accorded.””** Finally, the book received a good endorsement from Taverner’s colleague in the zoology division at the museum. Writing to a cousin in the U.S. in 1927 Anderson mentioned the publication of Birds of Western Canada, and said that for any young friends “who want to get a good illustrated bird book at a bar- gain, this is a very useful one, as it includes all birds from the Ontario border to the Pacific Coast. It is a large book, sold post paid for 75 cents in paper covers, and $1.25 in cloth.” He also said that the museum had received “any number of orders from U.S.A. Teachers and schools take a large number.’””*> Family Affairs During the years 1925 and 1926 Taverner had been working hard on field studies, and writing and re-writing versions of his book while enjoying little recreation and no holidays. It is not surprising that he did not undertake a collecting expedition in 1927, but instead went to California to see his ornithological friends there, and the birds of that region. He told a few friends that he was also going “for family rea- sons”, and would have to stay “with an elderly rela- tive in Los Angeles”. How did Percy find the address of this relative at this time in his life? He did not seem to have mentioned a relative of either his moth- er or his father in correspondence that has survived. Since Percy kept very few private letters, and certain- ly not a single one about his father, we have to fol- low up any clues that happen to appear. Presumably his mother had lost track of her first husband after her marriage to Albert Tavernier in 1881. We know nothing of what she told Percy about his father, but most likely it was a closed book between them. Certainly they were both too busy to waste time rak- ing over the past. But, after his mother’s death in 1924, Percy thought about the hardships of her early life, which he mentioned in a letter to Brooks.*° Perhaps he also thought about his own “lost” father, Edwin Fowler. However, nothing could be done by Percy himself until someone else made a move. While on a collecting expedition along the Red Deer River with Harrold in 1925, Percy received, like a bolt from the blue, a piece of information which he passed on to his sister. He wrote: THE CANADIAN FIELD-NATURALIST Vol. 110 “Got a strange and unexpected letter the other day from a man in Oak Lake, Man., asking if I were the infant he knew in Guelph 50 years ago. Asking me to visit him as I passed homeward. I do not know him but he seems to know all about me. Strange, isn’t it?”>7 Tantalizingly Percy did not give the man’s name in his letter to Ida but- presumably he showed it to her on his return home. There is no indication that he called on this stranger, although Oak Lake is only about thirty miles from Brandon which was on the Canadian Pacific Railway. It would have been easier to correspond with him. All we know is that by early 1927 Taverner was in touch with a Mrs. Fowler, liv- ing in Los Angeles. Writing to Munro about various conservation matters he casually mentioned that he and his sister were taking a trip west in the summer. “She has never been west and I have to go down to Los Angeles on personal business, so she has decided to accompany me’”.*® When Bishop heard from Taverner that he was intending to come to Los Angeles in the fall of 1927 Bishop invited him to stay. Taverner thanked him but explained that he would have to spend much of his time with an old lady relative “whom I have just discovered [living] in Los Angeles.”*? Writing to Fleming in August he mentioned that he was plan- ning to leave for California shortly, and explained the reason. “T have to go down to Los Angeles on personal business. Was going to do it next winter but it seems better not to wait. There is an old lady down there over 80 who is likely to go any moment and it is important that I see her. It is a rather strange story and some day I may tell you about it.”*° Confirming arrangements for their visit in a letter to Bishop of August he said that Ida would be com- ing with him, and that they would both have to stop with Mrs. Fowler, at 920 Gramercy Drive, Pasadena.*! Percy and Ida had an enjoyable trip wherever they went, with all kinds of courtesies extended to them. They stayed four days with the Munros at Okanagan Landing, and spent time with Brooks and his wife. In Victoria they spent one day sight seeing, including a visit to the Butchart Gardens. Taverner, as a keen gardener, considered these wonderful and wrote: “There is nothing like them even in California. Victoria is certainly the place in America to grow flowers.”*? In San Francisco he visited Swarth at the California Academy of Sciences, and Grinnell at the Museum of Zoology, University of California, Berkeley. With Ida he spent two days in Yosemite National Park sight-seeing, and they decided that it was the loveliest spot they had ever seen. In Los Angeles he spent an evening with Bishop and six of his friends. As he told it to Fleming “it was quite a little bird party”. On the way home they spent a day at Tucson. An ornithologist friend of Swarth, Joseph Mailliard, took them for a motor trip through the desert. Here they saw the unfamiliar forms of the 1996 large cactuses — Great, Barrel, Choya [Cholla] and Opuntia, as well as the wonderful coloration of the desert. Among the birds they saw were [Greater] Roadrunner, Vermilion Flycatcher, and a great num- ber of Swainson’s Hawks migrating.** It had been an enjoyable month’s holiday, travel- ling and visiting friends, for both Ida and Percy. Perhaps the most exciting event was the visit with Mrs. Fowler. At last Percy was able to ask someone who had been married to his father in more recent times all the questions that had been puzzling him during the past two years. But although Percy and Ida now knew, and although Percy may have told _ Fleming some things about his father, he left nothing on paper that tells us anything more than where Mrs. Fowler was living.** While Percy’s mother was alive there is no refer- ence that I have been able to find in any surviving correspondence to any woman friend, or of Percy having wanted to get married. After Mrs. Taverner’s death, Will Saunders, writing to Fleming to say that he had heard of the event, added, in his Puckish sense of humour, “It is too bad as it may compel Percy to get married which he does not want I sup- pose”.*> It is not clear why Saunders should have made this supposition. Certainly at that period in Taverner’s correspondence with his closest unmar- _ ried friends there was a certain amount of teasing. ' When men wrote to one another about having an | affair, or preparing to get married, they were likely to use jocular phrases and jargon that covered over the seriousness of the subject. While Laing was hav- ing a long drawn out affair with his fiancee Ethel, from 1920 through 1926, he had to put up with some chaffing from Taverner who claimed that he was saving up for the happy day when Mack and Ethel _ would finally be married. _ Brooks was the first of the bachelor friends to get _married, in April 1926, to Marjorie Holmes of Arundel, England. Laing wrote to Taverner, giving him the news, and added, “Well you’re next. I could- n't think of beating you to it. They will forget about me for a while.”*© After the AOU meeting Taverner ' wanted Brooks to stay a few days longer and look over some birds, but Brooks was too anxious to return to his wife to stay. Writing to Laing Taverner | remarked, “Who would have thought that Brooks would ever side step birds for a woman? Wonders never cease. There are still greater wonders to come.” At last Mack and Ethel were married in January 1927. The next to get married was Soper, in the spring of 1927. Writing to Fleming this is how _ Taverner described the wedding. “Soper’s wedding went off very nicely. The lady came on a week ahead and stayed with us as she had no friends here at all. The wedding was at our house also and a pretty ceremony it was. Just about the prettiest bride I ever saw. Dewey was proud as Punch of course. He is some fast worker. Met the girl when he went back CRANMER-BYNG: A LIFE WITH BIRDS 141 to Edmonton after the A.O.U. and had her in three days, stayed three days more with her and arranged for her to come on here to be married after six days aquaintance, — and seems to have made an admirable choice too.”*” In a letter to Munro of February 1927 Taverner commented on Brooks and Laing getting married. In a P.S. he added: “Soper gets married March 14th at our house to a western girl. He did it all in six days, some worker that boy.” But he warned Munro “Don’t expect me to walk the plank yet at any rate. However, I seem to be the sole survivor.’*® It was now Taverner’s turn to put up with some funmaking by his friends. In a letter to Laing about a mix up over arrangements for Laing’s field work for the summer of 1927 he added: “Yes all the world seems marrying mad. All friends and neighbours have had me married or about to be so this winter. It was cer- tainly a flattering report but it has caused more amusement than trouble.’ Although Taverner con- tinued his correspondence with his old friends with the same zest, sometimes he overdid the joke about marrying a wife and having the services of a stenog- rapher. In a letter to Laing of May 1928 he became quite eloquent on the subject. Whether Laing took it in the spirit of of light hearted banter one cannot say. But his conclusion may have had more than a grain of seriousness when he wrote: “P.S. The most interesting thing about your recent infor- mation is your wife. A typing, bird-skinning, camping and good cooking wife is a pearl without price, — are there any more at home like her? I am interested.” Did Taverner realize what he was missing more than he was prepared to admit?°° While Percy’s friends were getting married he was not left entirely “in the cold”. What eventually linked both Percy and Ida to a partner was, to some extent, their cottage, Hyla, on Blue Sea Lake. When their mother’s health was declining a friend of long standing from Detroit, Martha Wiest, came to spend her summer holiday at Hyla. Martha was a widow who taught music at a private girls’ school in Detroit. Each year after 1924 she and her son, Karel, came to Blue Sea Lake for their summer holiday. This was convenient for Percy and Ida since neither was at the cottage much in the summers of 1925 and 1926. By this time Percy had come to realize that the amount of entertaining that Ida was required to do while sharing with him their home at 45 Leonard Avenue was considerable. Although they had a good housekeeper, the strain of entertaining the guests during the AOU meeting, and working as a librarian at the museum as well, could not be glossed over. Percy was hospitable by nature and enjoyed enter- taining ornithological friends and acquaintances from across Canada, and from the United States. Letters of thanks from guests in this period continu- ally referred to their hospitality and kindness, but the amount of entertaining Ida had to perform was oner- ous. In June 1929 Ida had her birthday party at Hyla, 142 one of the guests being John McLeish, director of the Mines Branch of the Geological Survey.*! From the guest book kept at Hyla we know that John McLeish first spent a weekend at the cottage in May 1927. From that time he was a guest there from time to time, and there are a number of photographs of him with Ida. There is no information in letters which have sur- vived about the events leading to Percy’s and to Ida’s marriages in 1930, but their close friends in Ottawa, especially visitors to the cottage, must have realized what was “in the wind”. The only person whom Percy told ahead of the announcement of his forthcoming marriage was Fleming. Taverner must have asked Fleming for advice on a hotel in Washington for his honeymoon, because Fleming replied that the Burlington was the most comfort- able. Writing to Fleming on 7 March Taverner said, “T escaped any dire results in Detroit. The ring was a great success and the lady was delighted with it. Everything is fixed and I expect that [on] the 29th my bachelorhood will be smashed irrevocably.” In the next sentence he went straight into bird news when he wrote, “one day in the school grounds there saw a [Northern] Cardinal whistling in great shape. Was a pleasing sound that I have not heard since 1913 at Pt. Pelee.”>? This must have been in the grounds of the Liggett School, Detroit, where Martha taught piano from 1923 until early 1930. The school newspaper, the Gopher, under “Society Notes”, told the news of her marriage. “Friday, the last day of school before spring vacation, an exciting bit of news was quickly spreading. Not only exciting, but romantic! Mrs. Wiest, that perfect atten- dant, who firmly believes that no cold should keep you from school, or that you should ever have one, was absent! This, however, did not cause as much comment as the whispers of certain people-that-should-know to the effect that she was getting married! We couldn’t learn much about him, not even his name, but it was said that he was connected with a museum and had a Van Dyke beard. A few days later we nodded wisely as we saw in the paper that our own Mrs. Wiest, had, on Saturday, March 29 married Mr. Percy A. Taverner of Ottawa, Canada.”>? CHAPTER 14. Strains at the Centre: Percy Taverner and Dr. Anderson After Taverner had assumed his duties at the Victoria Memorial Museum in May 1911 he began to assess the collection of birds which the Geological Survey had been storing in its buildings. He soon discovered that it was hardly one to be proud of. The first thing that needed doing was to catalogue them, a major undertaking, in which Taverner was aided by Frank C. Hennessey, re-labelling specimens, veri- fying data and tracing the history of the accessions. Although a student, Hennessey had recently returned from an expedition into the Arctic as assistant natu- ~ THE CANADIAN FIELD-NATURALIST Vol. 110 Percy sent news of his impending marriage in a letter to Munro. He asked: “Did you know that Ida is going to be married? For that | matter so am I but don’t let the cat out of the bag ... until at least the first of April, — after that it will be all | over. Yes, I think I deserve congratulations. Hope I can | do as well as Brooks did.’>+ Munro passed the news to Laing who wrote in a jocular way: “Dear P.A.T.: I guess I'll have to believe it but its __—ihard. Very | hard. For a case-hardened old sinner so d- glad he was | single and not as other men are, on Dec. 9th last, you | sure have suffered a change of heart — quite a transfor- | mation in fact.” Laing ended the letter in the same humorous way as follows: “Well any how I hope you will be as happy as I am and though of course your ornithological career is ruined I congratulate you on showing that after all you are human and here’s our best wishes for happiness and good luck to you and yours.”> The wedding took place at Taverner’s house. The | invitation card was quite different from the usual | printed one. It was designed and printed by Percy | himself. It said in part: Martha Hohly Wiest — Percy A. Taverner United March 29, 1930 At their Home in Ottawa It was elaborately decorated with symbolic themes | — a thrush singing, a man with a winged helmet rid- | ing a horse among the clouds, a castle, a hill and a lake. Also some bars of music which later Laing }} deciphered as the “Ode to Joy” theme from: Beethoven’s Ninth Symphony. Perhaps Percy} arranged to play that theme on the gramaphone since he had a recording of it. It would have been just like him to have pulled off such a “stunt” — at his own wedding. The bride was fifty years old, the bride- groom was fifty four. They had known each other since 1904.°° They spent their honeymoon in Washington and Charleston.°/ ralist and artist under Captain Joseph-Elzear Bernier The reason for this voyage requires explaining. During the nineteenth century the British Navy) and the Hudson Bay Company served to substantiat« Britain’s claim to the Arctic archipelago. Canada’ | claim to the Arctic islands dates from 1880, but he concerns over American activities there began earlie’ in the century. This was the main reason for thc transfer of the archipelago from Britain to Canada 1) 1880.! Geological mapping, mineral exploration ani exploitation came first, but from the 1880s the threé 1996 to Canada’s claims to sovereignty over the Arctic islands had become clear. The Canadian government began sending expeditions to show the flag, and to stake a claim to territories not acknowledged by other countries as being under Canadian sovereignty. One way to do this was for the Dominion govern- ment to establish permanent police posts, and to resupply them by regular “Arctic patrol” voyages each summer. Another way was by sending an offi- cial expedition that might last a year or more and during which a-cairn would be built or a plaque erected. The Neptune expedition of 1903-1904 com- manded by A. P. Low of the Geological Survey of Canada was sent to assert sovereignty over Hudson Bay and the east Arctic, including a claim to Ellesmere Island.” Captain Bernier made several voyages in government service, the most important being that of 1908-1909, when he overwintered at Winter Harbour on Melville Island. Here he set up a plaque claiming the Arctic Archipelago for Canada from the mainland to the North Pole.* By 1912 Taverner realized that the museum’s col- lection was well represented by birds found from southern British Columbia to southern Manitoba. Unfortunately most of the “northern” material col- lected by G. M. Dawson, R. Bell and others had dis- appeared.* Taverner determined to build a collection of Arctic and subarctic specimens, and whenever possible bought from those returning from Arctic regions. Thus when Captain Bernier, who had been in the Arctic since 1916, returned to Ottawa in 1920, bringing with him specimens of birds collected mainly at the southeast corner of Victoria Land [Island] he sent them to the museum. Taverner, writ- ing to Fleming, listed the most desirable specimens. These were: [Red] Knot, including downy young in several stages, also eggs; White-rumped Sandpiper with downy young; Black-bellied Plover with half- grown young; white and gray Gyrfalcon; Rock and Willow Ptarmigan, both with downy young; Sabine’s Gull adult, and half-grown young.° Taverner also wrote to Bent on the same topic telling _ him that the museum wanted to purchase a lot of _ them, and asking him to give his opinion on the value of the most desirable ones.° (Bent had very recently written to Taverner asking him for data to be used in Volume I of his Life Histories in making up the egg dates for each species. Previously Taverner had sent him dates for sets of eggs in the museum’s collection of ducks, geese and swans for numbers 1 through 143. Now he wanted data on the remainder, showing locality and date for each set.) Bent and Taverner could give each other valuable help in ornithological matters.’ By far the best opportunity for the museum to expand its small collection of arctic birds happened during the Canadian Arctic Expedition of 1913- 1916, and the appointment of R. M. Anderson as zoologist to the Geological Survey of Canada. After CRANMER-BYNG: A LIFE WITH BIRDS 143 the expedition he was appointed to the Division of Biology at the museum in Ottawa where he worked until his retirement in 1946. As a result Taverner and Anderson worked in close proximity at the museum until Taverner’s retirement in 1942. They interacted on each other continuously during those years, though their family backgrounds, upbringing, educa- tion , and experiences before each joined the muse- um could hardly have been more different. With this in mind it may be useful to give a brief “flash back” to Anderson’s background until he and Taverner met at the museum in late 1916. Rudolph Martin Anderson was born in Winneshiek County, Iowa, on 30 June, 1876, of Swedish and Norwegian descent. He attended public schools and high school, graduating in 1894, entered the State University of Iowa, where he obtained a B.Ph. in 1903 and a Ph.D. in 1906. During these years he specialized in systematic zoology, compara- tive anatomy and animal morphology, while working his way through six years of university study as an Assistant Curator in the University Museum, and an assistant in the zoology department. During military training he served for five years in the University battalion, and in the National Guard of Iowa Infantry for six years. In athletics he was captain of the uni- versity track team and set two records that were unbroken for a number of years.* His interest in birds started early, and from 1893 he was publishing short papers on several species. At the age of twenty-one he produced a list of birds of two counties in lowa, which was privately printed (1897), followed by his well-received book The Birds of lowa which was in part fulfillment for his Ph.D. degree. This was a solid achievement based partly on his own observations, partly on information from a large number of contributors, and partly on writings such as Paul Bartsch “The Literature of Iowa Birds” (1899).’ His book was reviewed by J. A. Allen, Curator of Birds and Mammals at the American Museum of Natural History, in The Auk 1907; by F. M. Chapman in Bird-Lore in 1907; and by Joseph Grinnell in The Condor 1908, who said that it was the best State list to have come to his notice. “It has the stamp of scholarly workmanship. Iowa ornithologists are to be congratulated upon so satisfactory an exposition of their avifauna.” The author’s own annotations were extensive. While Anderson’s book was being published and reviewed, Taverner and Swales’ ornithological research on “The Birds of Point Pelee” began publi- cation in The Wilson Bulletin in 1907.'° In the first three chapters of this book we have seen what a poor start in life Percy Taverner received, in particular because he had no father of his own, nor any grandparents from his father’s or mother’s fami- ly. Nor did he have any brothers or sisters (except his half-sister, Ida Clare), nor any cousins. This prevent- 144 ed him having the morale that a young boy with a full quota of parents, grandparents and close relatives would normally have had at that time. By comparison Rudolph Anderson’s life was sup- ported by two families. His father was a Swedish born graduate of lowa University, and a member of the Iowa Legislature. His mother was a daughter of Nelson Johnson who immigrated from Norway, eventually becoming a pastor and farmer in Iowa. Several Anderson and Johnson uncles were of some prominence in their states. In 1913 Rudolph married Mae Belle Allstrand of Swedish descent, who had a B.A. (Iowa) and later an M.A. from Wisconsin. Percy Taverner knew nothing of his paternal grandparents, and had only the vaguest recollections of his father, if any. He never knew his maternal grandparents. He had no uncles or aunts, nor any cousins. He had no wife until later in life. He left no “family papers” when he died. By the time Anderson graduated with a Doctorate in 1906 he had already displayed a great deal of promise. Compared with Taverner, at the same age, he was in a different world of achievement. In 1906 he became assistant Commandant at Blees Military Academy, Macon, Missouri. Here he had little time for field zoology and museum work, both of which he found satisfying. However, he soon had the chance to make a turn in direction, a change which influenced his whole future career. While at the State University of Iowa in 1902 Anderson had made the acquaintance of a student, born in Manitoba of Icelandic parents, Vilhjalmur Stefansson, who moved to Harvard University the following year. When Anderson read about Stefansson’s plans to make an expedition to the arc- tic he wrote about joining him. An agreement was made, with Anderson being hired by the American Museum of Natural History as a field agent for zoo- logical collecting.''! After the “Stefansson - Anderson Expedition” of 1908-1912 returned to the United States Stefansson began organizing a major expedition into the arctic, this time by ship. When the financial support promised by the National Geographic Society of Washington and the American Museum of Natural History was insuffi- cient to support an arctic expedition on the scale desired by Stefansson, he turned to Reginald Brock, Director of the Geological Survey of Canada, which had given some limited support to the previous expe- dition. Brock secured him an interview with the Canadian Prime Minister, Robert Borden. '* The expedition was divided into a northern party under Stefansson, and a southern party under Anderson, with Stefansson in command of the expe- dition as a whole. The scientific work of the expedi- tion relating to geology, geography, anthropology and biology was to be under the direction of the THE CANADIAN FIELD-NATURALIST Vol. 110 Geological Survey. Anderson was responsible for the work of mammalogy and ornithology. As a result of this organization Anderson, and the scientific staff of the southern party, should report to the Geological Survey. But Stefansson had final authority over the division of the ships, so that in an emergency he might challenge Anderson’s powers over the south- ern party.'? One further problem was to cause ill feeling within the organization of the expedition. The department of Naval Service was to have “gen- eral direction” over the other departments of govern- ment in the expedition, including the Geological Survey, which was a branch of the department of Mines. During the expedition the scientists of the Geological Survey acted as a group, making Anderson’s position as leader of the southern party extremely difficult and Stefansson’s position even more awkward, as the showdown at Collinson Point camp, in March 1914, showed.'* Anderson was to feel the scars of this episode long after members of the expedition had returned to Canada. With this introduction of Anderson to the Canadian Arctic Expedition we can now return to Taverner at the National Museum in Ottawa. By February 1913 Taverner had heard that Anderson was going on the Arctic Expedition with Stefansson, and that J. A. Allen had recommended him highly as zoologist and mammalogist.!° Taverner wrote. to Brock calling his attention to this. In a letter to Fleming Taverner said that it would be great if Anderson could be given some museum appointment for the coming trip, “and perhaps land him here as part of our staff when he returns. Am in hopes that we can give him some sort of honorary appointment on the staff to be confirmed on his return so that he can collect for us and at the same time receive salary from the exploration fund.”!® Anderson and Taverner did not meet while prepa- rations were being made for the expedition to leave from the west coast of Canada in June 1913. Taverner wrote to him in 1914 at Bernard Harbour in Dolphin and Union Strait, where the southern party had its headquarters since August 1914, which reached Anderson in November after his return from a trip up the Coppermine River. Anderson replied with a letter dated 10 January 1915, from Bernard Harbour, giving news of his party, and his collecting zoological specimens. Anderson had sent a report on collecting done by the southern party during the first year 1913-1914 (from July to July) amounting to 212 birds and 77 mammal skins.'’ Now, in his letter of 10 January 1915, he reported good specimens of Barren Ground bears, and where they came from. Birds were not numerous, though he took about 12 specimens of Yellow-billed Loon, and some Black-throated Loon [Pacific Loon], and 2 Red-throated Loons which were rather scarce in the region of Dolphin and Union Strait. The only species of eider duck there, | 1996 he said, were Pacific [form of Common], and King Eider. He watched a nest of Semipalmated Plover “from the time the eggs were laid until the young hatched, and got some good pictures of eggs and downy young as well as the parents.” The only com- mon shore birds, he reported, were Baird’s Sandpiper. Anderson told Taverner that he intended to stay permanently with the Survey if conditions were favorable for scientific work at the museum. “Next summer will be my ninth summer in the arctic, and I have quite a bunch of notes which I ought to work up. He ended his letter “I am yours sincerely / Rudolph Martin Anderson.”!® Anderson and the southern party returned to Victoria in September 1916.'? This brought Taverner a lot of extra work preparing for Anderson’s arrival at the museum. Taverner was having great difficulty obtaining hard-wood sawdust for use with the large mammals that would be arriving, because the Mint had recently taken the museum’s regular source of supply for packing their gold coins. He also had to find office space for Anderson and his specimens, all of which were large. Writing to Brooks in France he told him: “The Arctic Expedition has returned and I expect Dr. Anderson daily. You know he is to be our mammalogist, and it will be a relief to hand this stuff over to him.” The two men met for the first time in October 1916.7 Anderson assumed his duties as zoologist, and took charge of the mammal collection at the muse- um. At the end of 1916 he was appointed to the Advisory Board on Wildlife Protection. Through membership of this Board, which met eight times in 1917, Anderson got to know several key men in the departments most concerned with wildlife. These were: James Harkin, Parks Commissioner; Gordon Hewitt, Dominion Entomologist; James White, sec- retary of the Conservation Commission, and Duncan Campbell Scott, Superintendent-General of Indian Affairs.*! Late in 1918 Anderson, together with Hewitt and Harkin, interviewed Hoyes Lloyd for the post of ornithologist with the Parks Branch, to be _ responsible, in particular, with enforcing the Migratory Birds Convention Act.” In 1917 Taverner introduced Anderson and his wife to the Flemings in Toronto.”? This gave Anderson someone with contacts in the world of ornithology, outside of government people, with whom he could correspond. During the summers of 1918 and 1919 Anderson was loaned to the National _ Parks Branch for work inspecting large areas of lake, swamp and agricultural land in Saskatchewan and Alberta reserved for wildlife sanctuaries and pro- posed for wild fowl breeding grounds.”* In telling this news to Fleming he mentioned that “Mr. Harkin is going out as far as Regina with me”. In the same CRANMER-BYNG: A LIFE WITH BIRDS 145 letter he enclosed an “accession card” notifying the birth of a daughter.”° In another letter to Fleming of 1919 Anderson mentioned that he had been down to Point Pelee on Parks Branch work and that W. E. Saunders and Dewey Soper had come for the weekend, and they stayed together over Sunday at “the shack”. Soper was just beginning a career in which he was to become an outstanding explorer-naturalist in the next decades. He was delighted to meet Anderson and told Taverner in a letter: “Dr. Anderson was with us ... he had so many interest- ing things to tell us about the north country. It may be that he will not realize just how much I enjoyed his com- pany.”2° In the same period Anderson passed various requests and suggestions to Jim Macoun, who was running the biological division during his father’s partial retirement in Victoria, British Columbia. Taverner was pleased when Jim was made perma- nent acting head of division and told Fleming “T am glad this appointment has been made. He is the right man in the place ... I get along with him very well indeed ...””7’ By mid-1919 the Taverners and the Andersons were beginning to get to know each other now that the Andersons had a house near the Rideau Canal, and Mrs. Anderson had a daughter, Dorothy Ann, one year old in August 1919. Taverner took Anderson to Blue Sea Lake for a weekend to look at possible sites for building a cottage. In writing to Jim Macoun Taverner said that they had “a fine visit” and added: “He certainly opened up in camp and makes a good camp fellow”. In October Anderson bought a lot on Big Island, quite near to the Taverners.*® Together Taverner and Anderson worked out the principles on which collecting per- mits could be issued. James Macoun, who was becoming progressively ill, died in January 1920. No worse a stroke of bad luck could have struck Taverner. His good friend “the genial, unassuming” James Macoun was gone. Taverner wrote, “Personally I feel that I have lost one of my best friends, one upon whose judgment and willingness to assist I could rely under any circumstances. Departmentally he is irreplaceable.”?? In March 1920 Rudolph Anderson was made act- ing head of the Biological Division under whom Taverner, as ornithologist, was to serve for the next 17 years. Taverner regarded the position of the museum in relation to the Geological Survey, as it developed early in 1920, as a fight for survival. But Anderson, who should have been a strong voice in his position as head of the Biology Division, kept quiet.°° Taverner tried hard to persuade Anderson to take an active part without success. Writing to Fleming in March he said that he had been trying to rouse Anderson to action “but he is hard to wake”. Taverner showed that already he had summed up Mrs. Anderson astutely when he said: 146 THE CANADIAN FIELD-NATURALIST “Will have to get after his wife, she sees things as they are, and is a hustler and is determined that he will hustle too. Cannot make [up] my mind just where Anderson stands.”?! To Taverner, Anderson may have seemed difficult to understand, and hard to rouse, but there may have been a very good reason for his attitude. We shall now take a look at some of the effects that the return of Stefansson from the Arctic were to have on Anderson and his companions of the Southern Party. Anderson had the unpleasant experience of facing Stefansson, the expedition leader, at Collinson Point Camp from December 1913. A major confrontation developed in February 1914 when Stefansson real- ized that he must challenge the “dissidents” of the southern party before they undermined his position and creditworthiness on the Arctic coast. Stefansson wrote to the deputy minister of the Naval Service, G. J. Desbarats, that he ought to bring the dissidents into line; that Anderson should attempt to carry out instructions given him by Stefansson, and that just because Anderson considered some other course bet- ter should not be considered an adequate excuse for disobedience [emphasis added]. Each side felt justi- fied in its own position and Stefansson regarded Anderson’s position as mutinous; while Anderson was under the strain of challenging Stefansson in order to protect what he considered to be the rights of the southern party. Stefansson came to Collinson Point camp again, after a visit to Herschel Island, on 8 March 1914, fully determined to “have it out” with Anderson and his party.°? After Stefansson’s return from the Arctic in the fall of 1918 the disagreement between the Canadian Naval Service, the Geological Survey and Stefansson centred round the publication of the reports on the work carried out by the expedition. Already some of the papers on the scientific results of the southern party were being written. Anderson had been given the task of writing a narrative of the journeys and field trips of the southern party. The deputy minister of Naval Service, Desbarats, wanted Stefansson to complete his report on lands explored north of Prince Patrick Island so that the government could lay down a chart of the lands he had discoy- ered. Since the southern party’s publications were progressing well it was necessary that Stefansson should carry on with his. But Stefansson had his own plans and gave priority to an immediate lecture tour, and the writing of his unofficial version of the expe- dition under the “loaded” title of The Friendly Arctic. When published in 1921 it caused a furor, especially in the Anderson camp. Mrs. Anderson put the matter forcefully: “Steffy does not realize that he is fighting the Canadian Government when he fights Rudolph ... [T]here is no room at the present time in Canada for VS and Rudolph at the same time unless Steffy reforms ... he must be taught to play fair.”* Vol. 110 A long letter from Diamond Jenness disputing many of the allegations made in The Friendly Arctic was published in the prestigious journal Science. Using documentary evidence Jenness was able to cast doubt on Stefansson’s ability to live off the land. A heated dispute developed between Jenness, Anderson, the southern party and the Geological Survey on the one side, and Stefansson with his vari- ous backers on the other.*4 Walter Brock, who had resigned in 1914 to become dean of the Faculty of Applied Science at the University of British Columbia, wrote to Camsell in 1923 about the rights and wrongs of the quarrel between the two parties. It was a judicious letter from the man under whose leadership the southern party had been planned. He suggested that only harm could come to all concerned by pursuing the matter further, “it has gone too far already ... I do not think for the per- sonal vanity or private satisfaction of one or a few indi- viduals, the officials or any others should be dragged into this dirty mess ... In the end, the individuals, the parties and the expedition will be judged on one thing only and that is the scientific results they present to the public.”* Unfortunately, a report on the zoological results of the expedition due to be published in volume 2 (mammals and birds), and written by Anderson, never materialized.*° When The Friendly Arctic was published Anderson gave vent to his feelings of anger and frus- tration by writing letters to those he could trust to take his side. One such was J. H. Fleming with whom he had been in correspondence since 1917.°7 In an angry letter of late 1921 Anderson complained that he had been taken to task in a review in the New York Times book review section of The Friendly Arctic. This was obviously by some literary hack ignorant of Arctic conditions, Anderson stated, among other things. It was a powerful piece of argu- ment.°® Anderson wrote several more letters to Fleming on the same subject and in the same tone of frustra- tion and suppressed anger. Eventually in 1924 Stefansson realized he was no longer persona grata in Canada and his later Arctic activities were carried out in the United States.*? As a result the Andersons now appeared to be on good terms with most people, although two civil servants in Ottawa were soon to feel Mrs. Anderson’s displeasure. Although the collection of birds obtained by the Canadian Arctic Expedition was a good addition to the museum, many more arctic specimens were still required. In 1923 Taverner and Anderson arranged for Dewey Soper to go to the arctic as naturalist in the annual expedition of the Canadian Government Steamer (C.G.S.) Arctic visiting Ellesmere and Baffin Islands with supplies for recently installed 1996 police posts.*? It was impossible to carry out any extended scientific investigations during the short stages of an arctic summer voyage as the ice packs were only open for a short time. To make extensive investigations and collections in zoology, botany and ethnology it was necessary for a zoologist to winter there and spend the following season on intensive field work. As a result the museum decided to send Soper to Baffin Island where the prospects of a large, unknown field for exploration were good. Soper sailed from Quebec in C.G.S. Arctic, commanded by J.E. Bernier, and landed at Pangnirtung, Cumberland Sound, on the southeast coast of Baffin Island on 22 July 1924. During a violent storm in the north Atlantic a large part of the ship’s cargo was lost, including coal. Soper decided to land at Pangnirtung on the outward voyage in case no stop could be made there on the return run.*! This was a wise decision because it enabled him to explore parts of the area with help from the Hudson’s Bay boats, and a power launch of the RCMP, and during the fall of 1924 provisions were laid down near the head of Nettilling fiord and more provisions in the spring. Soper, with Constable T. Tredgold, RCMP to accompany him on an exploratory police patrol, with Eskimos, sledges and dogs left Pangnirtung in April, arriving at Nettilling Lake about 180 miles to the west in May. The summer was spent collecting, sur- veying and photographing. Soper’s report, written in September 1925, told of his plans for wintering, and what he planned to do during the summer of 1926 when he hoped to find out something of the nesting grounds of the Blue Goose, [now considered a blue morph of the Snow Goose] which were still unknown.” This was Soper’s first arctic expedition. He arrived in Ottawa in October 1926 just in time to give a brief report on his ornithological work on Baffin Island to the AOU meeting in Ottawa. Writing to Bishop about Soper’s discoveries Taverner said that he had hoped for more gull data, but he was glad that Soper had found a colony of Kumlien Gulls [Iceland Gull] breeding, together with Glaucous and Herring Gulls on the south shore of Baffin Island.* The AOU meeting at Ottawa in 1926 ending in a weekend excursion on Blue Sea Lake was, in some ways, the climax of Taverner’s career. He was in high spirits (perhaps a little too pleased with himself) in his letters to Laing and Bishop. The greatest hit, he told Laing, was to present every registrant with a copy of Birds of Western Canada. This was substan- tial, well printed, and well illustrated — a book to be proud of. He told Bishop that entertaining an AOU meeting was “rather heady business, — but a barrel of fun”. He found it difficult to settle down to the humdrum routine again. Anderson also seemed pleased with the results of the meeting, and wrote CRANMER-BYNG: A LIFE WITH BIRDS 147 nicely about Taverner’s book.*4 But perhaps Taverner may have seemed a bit too pleased with himself. From this time onwards warnings of trouble ahead began to appear. The continuing feud between the Andersons and Stefansson appeared to be still simmering deep down. Writing to her husband from the cottage in August 1926 Mrs. Anderson mentioned carbon copies of letters about “Steffy” which he had left on the mantlepiece. She said: “You might care for my opinion on a few points.”*° Now, a fresh cause for Mrs. Anderson’s disapproval was Percy Taverner. In July 1926, Karel Wiest, who was staying with his mother, Martha, at the Taverner’s cottage, was reported to be planning to have a huge bonfire there. Mrs. Anderson sent a message to Karel that he was a guest on the island, that he was to read the regula- tions and abide by them. She also told her husband to speak to Ida Taverner about the matter. Apparently Karel did not have the bonfire.*° Another new target of her disdain was Hoyes Lloyd. Writing to her husband at the beginning of November she said: “T heard the supervisor of wildlife protection was slaughtering birds on our private property for his private collection. I was at first minded to call up his chief [Harkin] and ask him what /egal right Mr. Lloyd had to do this but on reflection called up the president of Big Island Community [Mr. Patch] and told him about it confidentially”. What she wanted was to have their island made a sanctuary, and signs posted to that effect. At the top of page | she wrote “Mr. Harkin better tell Lloyd again he is a flat-footed policeman and nothing elseng The next day she wrote again with second thoughts. “On reflection I am inclined to think our friends, the Lloyds, are doing only what P.A.T. wishes and conse- quently I would not blame them so much. I wonder how Percy or Mr. Lloyd would like it if you would take a shot gun and, armed with your permit to collect birds in Ontario, go into their back yard in Ottawa and shoot birds they brag about seeing there? ... I am sorry Mr. Lloyd is secretary of the Advisory Board & Wild life Protection. A much better secretary would be Mr. Lewis who practices what he preaches as I understand. I honor Mr. Lewis more than ever ... As to instructions from P.A.T. to Mr. Young — that seems queer. Doesn’t the chief of the Biological Division give his instructions to his own field party? I fear the help and advice of the bumptious ornithologist was not needed — or appreciated.” So Taverner was conceited and assertive, and spoke out of turn. In a letter Anderson wrote to H. P. Allstrand, an uncle of his wife, in the following year he said some- thing rather enigmatic. “My health has been fine for the past year, never better. I have been weighing consistently around 182-186 lbs for a year or more. I have not had a scrap with anybody for 148 a long time now and it is about time to knock on wood. If it keeps on this way I shall be loving all my enemies, and some of my friends say that I need a scrap occasion- ally to keep in the best form. There are two or three that I have not forgiven yet, however, but am not worrying about them.*? Taverner had an unpleasant setback at this time when he asked Laing to go with him as assistant ornithologist on a field expedition in 1927. Taverner thought that he had written early enough only to find that Anderson had already settled with Laing to come with him as an assistant mammalogist.©° This was partly Taverner’s own fault caused by the over jocular tone of his letters to Laing. This is how it started in early February, when Taverner wrote to Laing assuming that he would be too busy to go ona field trip in the summer because he and Ethel had at last got married on 19 January 1927. These were his words: “Good bye carefree bachelor days, working all summer for nothing, guess I have seen the last of you in summer field work. Golly, I hope Harrold does not desert the ship just yet.”*! This proved to have been a silly thing to have » said. Taverner had been taken off guard, and Laing’s reply showed that he was hurt by Taverner’s assumptions. “T like the way you put me, as an old married man, on the shelf regarding field work ... it relegates me to the ranks of the mouse-catchers ... you come right out and kiss me good bye.”?? Taverner knew Laing better than to expect that he would pass up field work, just because he had recently got married. That was not his style. This is how Laing’s biographer explains it. “The role of ‘the woman’ was clearly defined to Mack. Women were put on earth primarily to help men in the struggle for existence. Only after they had accepted their subsidiary or service role by getting married could they expect something approaching equal treatment or respect from men.”>? In March Taverner learned from Anderson that Laing had been hired by him for summer field work. Taverner was hurt and reproached Laing when he wrote: “Tt was rather a little shock to learn that you are thinking of going out this summer for Anderson. Think he at least might have told me of it or consulted with me again.”>** Laing replied: “Well, how in heck am I to know what you fellows are consulting about down there? Anderson asked me if I can go as usual and I said where and as you sort of kissed me goodbye when I got married, I naturally sup- posed you have side tracked me. Truth is that I suspected just the truth and feel between the upper and nether mill- stone.””>> At this point Taverner wrote to apologize for the mix up in the arrangements for the field work for the 1927 season. He explained, rather weakly: “Suppose I should have closed up immediately I found that you were willing to go out again. If you had not THE CANADIAN FIELD-NATURALIST Vol. 110 been such a bum correspondent this winter [I] probably should have. I assumed that you would want to stay home with wifie this summer. Your disclaimer was long in coming and I had no idea that Anderson was dicker- ing with you as a mouse.” He mentioned that Harrold was now going to Alaska for the California Academy of Science, and added that losing both him and Laing was the last straw that made him remain at home for the summer.°° Taverner had assumed too much, and had been taught a lesson. In the fall of 1927 he wrote to Laing: “Hope that next year you will be with me doing nice bird work again instead of hunting shrews and such ver- minous creatures.”>/ But in January 1928 he received a cordial but firm reply. Laing explained that Anderson wanted him to continue the past summer’s work on mammals along the B.C. border, and he said he would, providing that Anderson would make arrangements for Laing’s wife to come with them. He explained to Taverner that breaking up one’s home for a hundred dollars a month for six months of the year is not what it used to be. That is the reason he would skin mice and things. His wife, he said, made a good assistant and could make as good a bird skin as he could, though very slowly.°* Taverner and Laing never went on a field expedition together again, though it didn’t pre- vent them remaining friends and writing letters to each other, though less frequently. The “flare up” between Anderson and Taverner in early 1925 had soon settled down, and the few letters they needed to write to each other were quite friend- ly. But by early 1927 there were signs of coolness between them. At this point it may be helpful to explain the official position of the two men at the museum. Taverner’s title was ornithologist, Anderson’s was mammalogist, until he was made “chief of the division of biology” in 1920. This placed Anderson one step above Taverner in the civil service hierarchy. Anderson was now responsible for all returns and reports from the biological division (ornithology, mammalogy and botany). Anderson’s military training may have made him sensitive to his rank, and his duty as head of a division. Everything that Taverner wished to have done in the museum had to be discussed with Anderson first, and any let- ter.on official business would have to be approved by Anderson. If Taverner wrote an article that he wished to have published this would have to be sub- mitted first to Anderson who would edit it, correct Taverner’s spelling, and discuss its content with Taverner, if he wished. In more technical matters Anderson considered it his duty to keep a watch over administrative matters which Taverner, in his posi- tion of the museum ornithologist, considered to be his responsibility. These included the loan of study skins to other museums, and to individuals such as Fleming and Brooks who were semi-professional 1996 ornithologists. If the museum had sufficient dupli- cates of a particular species in a particular plumage, then an exchange of skins might be made with another museum or individual, thus helping to build up the museum’s collection. Just as stamp collectors may exchange their duplicates for ones they do not possess, so collectors of bird skins made exchanges. But in the case of a museum the responsibility of the curator of the ornithological section was consider- able. It required considerable knowledge and judge- ment, as well as an efficient system of registering all birds received into the collection, and all birds sent to other institutions and individuals on loan, or on exchange. Taverner started by keeping such a regis- ter entitled “Victoria Memorial Museum Register of Birds” which was kept from 1911.°? CRANMER-BYNG: A LIFE WITH BIRDS 149 Another humiliating thing for Taverner was that he had to make all applications for leave time, and for attending a conference, through Anderson and not to the director of the museum. As a result Anderson could advise the director whether he felt that Taverner should be granted any request. Also Taverner’s yearly reports on the work done by the ornithology section for the previous year for publica- tion in the Annual Report had to be submitted through Anderson. Because Taverner had been his own “boss” during the period 1911 through 1919 this was a particularly galling thing to have to do. As we have seen, Dewey Soper returned from his second arctic expedition in the fall of 1926. These two expeditions had been arranged by Anderson and Taverner jointly for the National Museum. Soper’s Joseph Dewey Soper upon return to Cape Dorset after surveys run to Nuwata across the interior of Foxe Peninsula and from there east to Ungmaluktuk Lake and south to Gordon Bay during March 1929. Reproduced courtesy of the University of Alberta Library Archives, The J. Dewey Soper Collection, 79-21-34: 1579. See W. E. Stevens and George W. Scotter. 1983. Canadian Field-Naturalist 97(3): 350-355. 150 third expedition to the eastern Arctic was sponsored by the Northwest Territories and Yukon Branch, Department of the Interior, because the National Museum could not find a permanent post for Soper, though Taverner gave him his support. There were still several species of birds whose nesting grounds in the arctic were unknown. One was the “Blue Goose” which Soper had tried hard to find on Baffin Island, in 1925-26. At that time the AOU Check- list (1910) showed the Lesser Snow Goose as having two distinct colour phases, the white and the blue. These were known to winter in Texas and Louisiana. Taverner’s friend McAtee reported blue geese abun- dant in Louisiana with white-morph scarce. In Texas blue-morph geese were known to be scarce. North American ornithologists were specially interested to discover where the breeding grounds of the “Blue Goose” were. They also wanted to discover how it managed to retain its blue genes and remain a sepa- rate species.°! In 1928 Soper returned to Baffin Island, reaching Cape Dorset in August, making his headquarters at the Hudson’s Bay Company post there. From there he made various important journeys by land and coastline, exploring in southwestern Baffin Island, with one or more Eskimo assistants and several teams of sled dogs. The stamina of Soper, the Eskimos and their dogs is vividly described in words supported by photos, in two articles by Soper cover- ing a whole year of arctic travel. When spring arrived Soper had developed consid- erable familiarity with the vast Foxe Land interior and the Foxe basin coast. He had twice been over the region in which the “Blue Goose” was thought to nest, and knew the best route by which to reach his proposed quarters for the summer. On 17 May 1929 he left Cape Dorset with five Eskimo drivers, four sledges and forty-two dogs, carrying sufficient food and equipment for a period of three months in the interior. After eight days of hard travel they reached the proposed site of their summer camp on the banks of a river near Foxe basin at latitude 65°35’ north. This they named Camp Kungovik, the Eskimo name for “Blue Goose”. Three Eskimos with all the dogs and all the sledges started on the trail back to Cape Dorset. Two Eskimos — Kavivow of Cape Dorset and Ashoona of Gordon bay remained to assist Soper. They relied on a single freighter-canoe with which to navigate Foxe basin and travel by rivers and lakes across Foxe Land to Hudson strait in August when the ice was melting — a daunting chal- lenge. The first geese passed over camp on the evening of 2 June, the next on 5 June when 34 alighted on a snow-free strip bordering the Kungovik river — twenty four of them were “Blue Geese”. By 8 and 9 June thousands of geese were crowding bare patches of tundra in the region. By late June the lowlands were rapidly being cleared of snow and breeding THE CANADIAN FIELD-NATURALIST Vol. 110 birds were withdrawing to the nesting areas. By now the migration was over, and “Blue Geese” were in the majority. Soper and his assistants searched the region on foot for nests. Finally, on 26 June a small colony of breeding geese with ten nests, eight of which were of the “Blue Goose”, were discovered. Several years previously Kavivow had tramped through this region on a caribou hunt, and had noted a place eight miles southwest of Camp Kungovik. It was here that they were now busy collecting birds and sets of eggs. On that date Soper wrote a two page letter to Taverner describing what he had seen. He said, in part: “... notwithstanding the wretched climate, I have wit- nessed some wonderful events. A book could do justice alone to the details. As a bird migration route and a breeding grounds it eclipses by far anything I have ever beheld in the past. Think of a region swarming with Blue Geese — during the migration geese of all kinds, and Brants —, and around camp to have such birds common- ly nesting as Red Phalarope, White-rumped Sandpiper, Parasitic and Long-tailed Jaeger, Black-bellied and Semipalmated Plover, Ruddy Turnstone, King Eider, Black-throated Loon [Pacific Loon], Sabine’s Gull and — a little removed, Blue and Snow Geese! Yes, indeed, it has been a great experience, especially during the migration, with thousands upon thousands of these birds and others swarming over the snow-free patches of tun- dra on their way to higher latitudes. For music, we depend as usual, upon the abundant [Snow] Bunting and Lapland Longspur, the darlings of the lonely wastes. I have added several new species to the Baffin Island list, names which I have carried around in anticipation before, blank spaces since 1923! And as for collecting, it is a paradise for northern birds. And by the way, I have among other things discovered the real Baffin Island migration route of the Purple Sandpiper — it is right past Camp Kungovik! — by the tens of thousands, com- pact flocks of hundreds, so closely ranked on the restricted areas of bare ground at time of passing that a single charge would desolate half a hundred future homes at once.” In the last paragraph he tells Taverner that as soon as the ice disappears in the southern part of Foxe Basin they will embark on the river, and begin the long voyage to the sea. “From the basin our route lies thru a partially unexplored chain of rivers and lakes thru the heart of Foxe Land to Hudson Strait. Luckily, I know more than the half of it from explorations and mapping conducted last fall, so may now lay my plans, as [I did] before leaving Dorset, to the very best advantage.” He ended this letter with the words “Adios, senor Dewey P.S. This camp is named from the Eskimo for Blue Goose.””©? This is a letter that should be known to all Canadians as an account of an epic in the exploration of the Canadian arctic, as well as the discovery and first hand study of an arctic breeding species of birds. (A photocopy of Soper’s handwritten letter is 1996 CRANMER-BYNG: A LIFE WITH BIRDS Syl A photograph received by Taverner in January 1930 from someone as a gift. It was 4 x 2 1/2 inches. Percy had it framed and gave it to his wife, Martha Wiest, after they were married in 1930. She kept it until just before her death when she gave it to her son, and just before his death he gave it to John Cranmer-Byng. (Reproduced courtesy of the Canadian Museum of Nature, number 100028.) 152 included in Appendix | at end of book). The impor- tance of the achievement of Soper, together with the work of Kavivow and Ashoona, has not been forgot- ten. A large area on the western side of the Great Plain of the Koukdjuak is now named Dewey Soper Bird Sanctuary, while a bay north of the Koukdjuak river where it flows into the Foxe Basin bears the name Taverner Bay. North of this lies a river named by Soper for the explorer and ornithologist Bernard Hantzsch, of Dresden in Saxony, who died there in June 1911.% One of the first North American ornithologists to ask to be able to see specimens of the “Blue Geese” was Frederick H. Kennard, Curator of the Museum of the Boston Society of Natural History. Kennard was gathering material on geese for a study of the genus Chen, and had requested a specimen of the “Blue Goose” for his museum.°° In 1928 Anderson was appointed naturalist to the Canadian Arctic Expedition for that year, and sailed on S. S$. Beothic from Sydney, Nova Scotia, in July.® In 1929 Taverner accompanied the annual expedition on the S. S. Beothic as naturalist, sailing from Sydney on 20 July. The reason for the annual expe- dition was that from 1903 the Dominion Government had been extending its administration northwards, and advanced police posts had been established as far north as the Bache Peninsula on Buchanan Bay off Norton Sound. RCMP officers of these posts had carried out wide-ranging patrols over formerly inaccessible country, and had collected information and species of value, which had reached the National Museum of Canada. The voyage was a circuitous one, depending to some extent on what supplies were required, and the condition of the ice. After sailing to the west coast of Greenland the S. S. Beothic went as far north as the Bache Peninsula post on Ellesmere Island, then to Devon Island and Pond Inlet, then Pangnirtung on southeast Baffin Island. A special trip was made to Chesterfield inlet in Hudson Bay for provisions and coal for the RCMP detatchment there. Taverner returned to North Sydney on 3 September after a voyage of 7800 miles lasting a month and a half. He told Fleming that he only obtained 65 specimens. He col- lected about six species that he had never taken before. The most interesting were Hudsonian Curlew [Whimbrel] and Red-backed [Dunlin] and Stilt Sandpipers from Chesterfield. The Red-backs were in an interesting plumage change which he had not seen before. He then raised the question of where the Hudsonian Curlew bred, a subject which he dis- cussed in detail in an article published in 1942.°7 Taverner summed up his experiences during this voyage in an account in The Canadian Field- Naturalist.°’ The National Film Board made a film of the expedition to the arctic on the S. §. Beothic in 4 reels containing a brief description of subject mat- THE CANADIAN FIELD-NATURALIST Vol. 110 ter, including “Shots of naturalist P. A. Taverner, and J. Dewey Soper discoverer of the nesting place of the Blue Goose”. On the return voyage the S. S. Beothic called at Lake Harbour, Baffin Island and Taverner’s Journal read: “Aug. 17 Lake Harbour — D. Soper not yet arrived from Cape Dorset.” Almost at the last moment Soper hurriedly arrived. The two men had a great opportunity for plenty of talk on the homeward journey via Chesterfield Inlet, Burwell Harbour, coast of Labrador, Strait of Belle Isle to North Sydney Harbour. Soper’s outstanding success on the Bowman Bay breeding ground of Foxe Basin, and Taverner’s opportunity to see parts of the eastern high arctic from on board ship excited his mind with the wish for new opportunities for studying more of Canada’s Arctic breeding birds on their breeding grounds. It was no surprise that Taverner planned his field work during 1930 for the arctic. Problems of trans- portation caused him to choose Churchill, Manitoba, which he said in a letter to Munro, would give him a base line from which to start, since he was anxious to work northward in future years.°? Taverner, with A. (Bert) C. Lloyd as taxidermist made camp near Churchill in late May where they were joined by Victor E: Gould, of Wolfville, Nova Scotia, as stu- dent assistant. They collected a few miles north of Churchill. Taverner left Churchill at the end of June, Gould and Lloyd in September. They made a useful collection for study and exhibition purposes includ- ing 650 birds and 68 mammal skins, and material for making habitat groups. Early in January 1931 Taverner was making plans for the coming season, as he told Laing. “I want to send Bert Lloyd up to Pond’s Inlet for a year if pos- sible. I think it is a key locality for the Arctic fauna ...” He explained that one could not get into the high arctic early enough to be able to do anything in the same season. The only way, he said, was to win- ter over in a location so as to be ready for the breed- ing season.’° But Taverner’s hopes for an expedition in 1931 were dashed because the economic situation was too serious for government to allot funds for field study expeditions from 1931 through 1934. However, an expedition organized by the Carnegie Museum of Natural History, Pittsburgh, Pennsylvania, under the direction of George M. Sutton, and John B. Semple, with Dr. Olin Pettingill and Albert Lloyd, went to Churchill. A second party, of individual ornithologists from Canada, consisting of Frank Farley (Camrose, Alberta) and Arthur Twomey, with two others spent June and part of July working the tidal flats, tundra, and bushland with their base at Churchill townsite. In 1932 Arthur Twomey, and Miss Marguerite Heydweiller of Rochester, New York, collected in the Churchill neighbourhood during June and July. As a result of 1996 the field studies carried out in this region during the years 1930-1933 Taverner and Sutton collaborated in writing an account of what was achieved. This is a substantial study consisting of ten pages of historical introduction, and the problem of the naming birds as coming from “Hudson Bay” when they had often been found farther south. A brief account of the field work carried out by Taverner with Lloyd and Gould in 1930, and by others during 1931-1933, follows. A helpful description of the country around Churchill gives an idea of the habitat there (pages 5-10). The major part of the book is taken up by an Annotated List of Species. Taverner was responsible for the account, quotations and conclusions covering the work done in 1930 and 1932; Sutton for 1931. It is embellished with 13 black-and-white illustrations, a sketch map of the country round Churchill, and an attractive coloured illustration of a Hudsonian Curlew [Whimbrel], with a downy young, by Allan Brooks.”! Early in 1930 Taverner was reassessing his posi- tion as regards field work. Writing to the Director of the Museum, W. H. Collins, about an expedition to Hudson Bay in the summer. He said that now Harrold was dead, Laing was collecting mammals for Anderson, and Soper was in the Service of the Northwest Territories, it was important that others should be trained to take their places.’? Writing to Mitchell in Regina about future field work he explained: “We have done the preliminary work in the southern part of the province, and can leave it for future develop- ments, more or less to local effort, while we concentrate on the more inaccessible areas.””* Another reassessment he needed to make at this time was concerning the American field expeditions in northern Canada. While Taverner was working in southern Canada with two or three assistants things went well. But as a result of the arctic patrol voyage of 1929 Taverner realized that Canada was at a great disadvantage as regards ornithological work in the arctic. No Canadian university had the funds to mount an expedition, and only a few museums could do field work in the far north and then only in their own province. There were no foundations in Canada to donate funds for field study in natural science. When the big financial slump hit the economy in 1929 Canada had limited resources for even the National Museum, and from the season of 1931 until July 1935, no funds were available for field expedi- tions. However, wealthy American museums such as the Carnegie Museum of Natural History, Pittsburgh, and Museum of Vertebrate Zoology, University of California were able to carry on. This was a double blow to the National Museum of Canada because, while Anderson and Taverner were unable to fund anyone to collect birds and mammals in arctic Canada, American institutions could, and several did.’* In the “Summary Report” for 1910 Walter CRANMER-BYNG: A LIFE WITH BIRDS 153 Brock had clearly stated the national objective in natural history. “For the present, at least, the muse- um is confined to Canadian material, the object being to specialize in this until it becomes in all branches thoroughly representative of the whole Dominion, a place where the entire natural history of of Canada may be studied.” Writing to W. H. Collins Taverner explained: “If we are to do serious and lasting ornithological work in the museum, it is practically imperative that we have certain series of extralimital specimens for comparison with our own. This is particularly true of arctic work where so many of our own species are circumpolar.”7° The situation was especially tiresome to Taverner at this time because he was asked to revise the galley proofs for a forthcoming “checklist of the birds of the world” organized by the American Ornithologists’ Union, the British Ornithologists’ Union and the International Ornithological Congress. But his work was handicapped by not having access to the specimens he needed for the job. This may have been the main reason why he exchanged what he believed were surplus museum specimens, when an opportunity occurred, for extralimital ones not represented in the museum. When the Ministry of the Interior decided, in 1933, that a descriptive account of the eastern arctic should be written, Charles Camsell, Deputy Minister of Mines, was approached asking whether Anderson would write on mammals, and also saying: “We would also be glad to have Mr. Taverner write up a section on birds. Due to his personal touch with the Arctic I think he would be the best person in the service to undertake this work.” The writer explained that the book “would be of value in showing that Canada has contin- ued to actively administer the area. The importance of evidence of this kind is borne out in the case of Norway vs. Denmark, before the Permanent Court of International Justice.””/ Taverner started with an eight page overview of ornithological collecting in the far north and Arctic, including that by Bernhard Hantzsch.’* Five pages were allotted to a description of landscape, climate, plants etc. Taverner wrote economically and with precision. “The land is bare, the wind-swept hills absolutely so, elsewhere covered with mosses and low-growing alpine plants. The only shrubs are dwarf pencil-trunked wil- lows raising scarcely six inches from the ground and grasses grow only in the most sheltered localities. The summers are short and with the aid of a twentyfour hour day, the hardy vegetation flowers, ripens its few seeds and prepares for the long bitter winter in weeks instead of months. Birds come before the ice has lost its hold and navigation is possible, nest immediately between snow banks, labor hectically to raise their young and are gone again in a few weeks. The difficulties of ornitho- logical work under these conditions are considerable. Birds come and breed after sled travel has ceased and before water transport is practicable. By the time the 154 first ship or boat can introduce the naturalist to a chosen ground nesting is largely over and return migrations have begun to obscure the characters of the resident pop- ulation.” Taverner wrote briefly about the effects of the ecological conditions on bird life. In the eastern arc- tic islands the short irregular season seeds are few so that seed-eaters are lacking, except for the Snow Bunting whose numbers are few per acre, and widely scattered. Ptarmigan, because they have the ability to feed on buds, twigs, fruit and bulky “roughage”, thrive. Predators and offal-feeders of hardy forms such as Gyrfalcons and [Common] Ravens, Snowy Owls and jaegers, by covering considerable ground or occupying favourable situations, find sufficient subsistence. Taverner concluded by mentioning migrational routes through the area. Most waders worked up the west shore of Hudson Bay and the hinterland; few except for the Purple Sandpiper and the phalaropes seemed to travel the eastern (Atlantic) coast. He speculated that the area of the Melville and Boothia Peninsulas, of which little was known ornithologically, were important migrational high- ways into the centre of the archipelago. In the east- ern arctic there were a number of European wander- ers (intrusives) that arrive by way of Greenland. Just how extensive these foreign influences were, Taverner said, was one of the problems of arctic ornithology that they were attempting to solve.” Although what Taverner wrote was interesting and well written it was not accepted for the book, issued by the Department of the Interior in 1934. Instead he wrote a section on “Birds of the Eastern Arctic”, a useful resumé of 113 species found in the arctic regions of eastern Canada. Baird’s Sandpiper was shown to have a high northern breeding range; the European Golden Plover to occur in Baffin Island as a migrant.®° While this was being printed Taverner received a letter from W. B. Alexander who was doing research in economic ornithology at Oxford University. Alexander had been appointed Director of the 1934 Oxford University arctic expedition to Ellesmere Island and asked whether Taverner could recom- mend any paper on birds of this area, or if not a list of birds likely to occur. This drew a reply from Taverner on how he saw the situation in arctic ornithology in 1933. In part he wrote: “There is still a great deal to be done in arctic ornitholo- gy as much to correct old errors as to make new discov- eries. The trouble is that so much of the arctic work has been done by inexperienced and casual observers that it is too bad that some one of qualified ornithological experience is not going to take part. We have learned considerable of late years but much of it goes to show how sketchy much of the old work inavoidably was. The time has come when careful and more intelligently directed work is desired in the high north.” THE CANADIAN FIELD-NATURALIST Vol. 110 Taverner told Alexander that the Ministry of the Interior in Canada was in the process of publishing a booklet for the needs of travellers and explorers in the eastern Arctic which would contain an annotated bird inventory.*! As an example of what Taverner knew in 1933, here are two items from his annotated check-list: “Ross’s Goose, Chen rossi, like a very diminutive Snow Goose hardly larger than a large Mallard Duck, is still one of the enigmas of the north, migrating through lakes Athabaska and Great Slave it disappears into the no- man’s-land of the Northwest. Territories. It may nest beyond the continental mass on Prince of Wales or Somerset islands or on the Melville peninsula and any data on it after it passes from well-known localities are desirable. Kumlien’s Gull, Larus kumlieni, though regarded by some as a hybrid between Thayer’s and Iceland Gulls this is undoubtedly a distinct species for as yet its only known nesting localities are far removed from the breed- ing range of either of these forms. The demonstrated localities for the species are, Cumberland sound, south- western Baffin island, and on the mainland across the strait near Cape Wolstenholme.” _ Taverner also reported Ross’s Gull, Rhodostethia rosea, as extremely rare. When Soper was searching for the “Blue Goose” in 1929 he took a Corn Crake (Crex crex) in Cape Dorset which Taverner consid- ered extraordinary but not impossible. In part, he wrote: “There seem to be a number of these erratics that hit Baffin Island — [Northern] Lapwing, Barnacle Goose and this Rail. The flight line is probably Iceland, Greenland and Baffin.”’°2 By 1930 relations between Anderson and Taverner were uneasy, and under the surface there were signs that not all was well. However, as long as Anderson and Taverner were able to get away from the museum for a month or more most sum- mers on a field expedition they could just about bear each other’s company for the rest of the year. But from 1931 through 1934 they were confined to the museum at close quarters because of the economic depression. Writing to Rowan in May 1931 Taverner told him, in his graphic way, of the cutback in government funds. “No field work this season, Government economising. Will be lucky if I don’t lose my job. Heads have been falling right and left in the Civil Service and no one knows who will be next.”** Camsell, Deputy Minister of Mines, wrote: “Field work in anthropology and biology, which in pre- vious years has constituted the principal activity of the National Museum of Canada, had to be postponed alto- gether in 1931-32 owing to lack of funds ... ”°4 Zaslow put the situation starkly when he wrote: “In the biological field the four specialists — Anderson, Taverner, C. Patch, and M.O. Malte, the chief botanist — were reduced to three when Malte died in 1933 and 1996 . CRANMER-BYNG: A LIFE WITH BIRDS 155 Rudolph Martin Anderson, May 1934. (Reproduced courtesy of the Canadian Museum of Nature, number J-6611.) 156 no replacement was sought for some years, the position being eventually filled by A. E. Porsild (1936). The competent museum specialists were trapped in a neglect- ed, declining part of the public service with resulting frustration and lowered morale.”®> The first indication of serious reductions to finances allotted came in a letter from the Department of Finance to the Department of Mines of August 1931 saying that it would be necessary to make curtailments in its expenditure programmes, and asking that the personal opinions of departmen- tal officers should be sought. Camsell, as Deputy Minister of Mines, was asked to make a survey of services under his control in order to find out what might be postponed or curtailed, and tabulating the result into three categories: Services that it was considered necessary to continue in full; that could be curtailed; that could be withheld dur- ing this fiscal year (1931-32). An estimate of the savings which could be made under each item should be given. With this letter from the Department of Finance was a request that an answer be sent urgently. Camsell happened to be visiting mining companies in Noranda and Timmins, and the Acting Deputy Minister, L. L. Bolton, had to relay information to Camsell as a result of his discussions with an official in the Department of Finance. One thing he learned was that this official put forward some suggestions, such as: Cancelling any proposed field work not already under way; recalling parties from the field’ forthwith; dropping proposed purchases of any new equipment; making no more purchases of museum specimens. There was no suggestion for curtailments that would result in a decrease in staff.*° At some time in the fall of 1931 Camsell probably had a meeting which included senior members of the biology division. There was an anecdote about Taverner and this meeting which was in keeping with his personality — he could be very outspoken when he wanted to put over a point. Apparently Camsell was explaining the situation, telling them that he needed to be able to assure the Finance Department that the biologists had important work to perform. He said he could not think of anything of importance for them to do since they could no longer have money for field work. Whereupon Taverner, who was acting as spokesman of the group rather than Anderson, at this point “balled Camsell out.”*’ As the world economic situation deteriorated from 1930 Anderson and Taverner found themselves struggling to fund their field projects. Eventually Parliament released sufficient money by mid-July 1935 to support 10 field projects for the museum, of which the biological division obtained money for three. Anderson was able to send out a party to col- lect mammals in the late summer and fall while their pelage was turning from breeding into winter coats. But for Taverner the bird breeding season was nearly THE CANADIAN FIELD-NATURALIST Vol. 110 over. Only some birds in fall and winter plumage could be collected while on migration. Therefore Taverner postponed a field expedition for another year.®8 Until 1935 the lack of warmth between Taverner and Anderson was noticeable but could not be called animosity. They worked in close proximity, had to consult together continually, sat on several commit- tees together, and attended various other functions in common. Taverner felt the Andersons’ disapproval though he could not know the reasons why. He guessed that they were jealous of what he called “some little recognition” that he had received. There were other reasons which surfaced from 1935-36 onwards which made Taverner’s position more than merely uncomfortable. It became unbearable. Taverner was so exasperated that he wrote to Collins, who was still director of the museum, that he found himself handicapped as the ornithologist of the museum, and put in an intolerable position. He said, in part: “This has been caused by Dr. Anderson’s recent and apparently unauthorized assumption of authority and interference that has never been exercised before over my office or over the coordinate section of Botany by him or his predecessor in office. Some of the unjustifi- able methods of this interference you are already aware Of wie Taverner argued that it had always been assumed that the ornithologist, mammalogist and botanist and other heads were duly authorized and responsible officers that knew their business and could be trusted to direct their own sections. Until such a person was proved incompetent or unworthy of trust there was no reason to hamper them by autocratic interference. He urged that the ornithological section be returned to the status that it had held for the past twenty-five years under which it had prospered and achieved some success in the scientific world.*? What had been some sparring between two men of very different temperaments and abilities now became something rougher. According to the Civil Service system and code of behaviour the man at the top of any division or department had considerable power over the next person under him, however experienced and able that person might be. Anderson certainly had the power; it depended on how he used it. The fact that the two men managed to carry on their work without any public quarrel was because they were expected to do so by the system. But it was very galling from Taverner’s position. For example he had to take anything he wrote for publi- cation in his work as museum ornithologist to Anderson to be edited. This meant that Anderson could make suggestions as to length, style, content. At times it could work to Taverner’s advantage, especially when Anderson corrected his irregular spelling. Anderson was helpful in the publication of Birds of Western Canada (1926), but when 1996 Anderson started suggesting, from 1935, that some of Taverner’s short notes on ornithological matters were merely rewritings of articles that had been pub- lished twenty or more years earlier, Taverner was not impressed.” In 1936 Taverner wrote to Laing telling him that relations between “the Dr.” and himself had been growing worse for some years and had now reached a climax. “I won’t go into the dirty details but I have had some mean tricks played on me. Lately I have been made aware that he has been writing in a most derogatory manner of me to friends. He may have done so to you and I don’t want to mix you up in private quarrels but to warn you against accepting all he may say in the matter. He seems obsessed with jealo[u]sy of me because I have received some little recognition and he has done practi- cally nothing to obtain it in his 18 years here. Even his Arctic report is still to be written. However that is enough gossip. You will probably be under his orders this year though I hope that I have not lost your services indefinitely.?! Writing to Laing again the next month Taverner mentioned various matters and then added “T don’t want to get you mixed up in any personal dis- agreement we are having here.”°” Another piece of information that Taverner included in the same letter was that there would probably be a change in the museum’s organization. “It is badly needed. The Biological Division is quite unworkable under the present circumstances. It is not my fault. I have been trying to avoid this condition for years.” A few months later, writing to Brooks about a dis- appointment he had suffered over a collecting trip by sea for the museum that had fallen through, Taverner said that he did not know why, but he added “R.M.A. is so d —- close that I know nothing that is going on beyond my own personal experience.’ In March 1936 the newly re-elected Liberal gov- ernment of Mackenzie King began a major reorgani- zation of the government. A new Department of Mines and Resources, with T. A. Crerar as minister (1935-1945), was set up. Camsell became deputy minister of this new Department. The reorganization took effect from 1 December 1936, with the minister and his deputy presiding over five large branches. Mines and Geology was headed by John McLeish, formerly director of the Mines Branch. It comprised three main divisions: the Bureau of Mines; the Bureau of Geology and Topography, which Lynch continued to head; and the National Museum of Canada, also controlled by Lynch. Taverner’s sister, Ida, had married John McLeish in March 1930. Taverner was now the brother-in-law of the head of the Mines and Geology Branch.” For some years Taverner and Anderson had been like two semi-blind men playing a game of bluff in which neither knew clearly what the other was doing. If that was all it was, the game might have CRANMER-BYNG: A LIFE WITH BIRDS layj/ ended when they both retired, as something rather comic for them to look back on. But this was not how it turned out. In April 1936 Anderson sent Camsell, Deputy Minister of the new Department, an eighteen page memorandum about the qualifications of ornithologists in the National Museum, and vari- ous other matters. This brought all Anderson’s griev- ances about Taverner to the knowledge of the gov- ernment officials.*° Taverner’s requests to Collins to be given a Division of his own finally came to fruition soon after Collins died. Writing to Brooks in April 1937 Taverner was able to say: “T suppose you know I have things straightened out here now. Am chief of my own division of Ornithology, it is only a one man department as I have no assistant — but Thad none anyway.”?” At the beginning of 1937 Taverner had an inter- view with Lynch, who by now had administrative responsibility over the National Museum (with Wyatt Malcolm under him as its effective head). In his record book Taverner noted: “Agreed that the title Chief of Ornithological Division can be used as a semi official title of my office, though not at present recognized by the Civil Service list.””* In order to grasp the seriousness of the struggle between Anderson and Taverner we need to examine two major issues that inflamed relations between the two men from 1935 onwards. The first issue concerned the proper procedure fol- lowed at most museums in the early decades of the twentieth century in making exchanges of bird speci- mens between one museum and another. It was for the head of the ornithological section to decide what specimens could be considered surplus at any one time, and what birds the museum needed in exchange. When Taverner was appointed the first ornithologist at the Victoria Memorial Museum in 1911 he learned, while visiting several museums in the United States, the usual procedure in the case of loans and exchanges. From that time onwards he was in charge of the ornithological section, and by the time that Anderson became Chief of the Division of Biology in 1920, Taverner regarded himself as well qualified to decide what exchanges should be made, without approval from Anderson. On the other hand Anderson, as Chief of Division, wanted to know what specimens Taverner proposed for exchange and to which museum. Unfortunately, in exchanging specimens in order to keep building up a collection there is no absolute standard that can be applied automatically; it has to be a matter of individual judgment. The problem was even more delicate when it concerned an exchange between a museum and an individual collector of bird specimens, such as Fleming, Brooks, Bishop and many others. By 1935, it was clear that Anderson expected Taverner to refer to him in such matters of judgment. This Taverner found unacceptable in his position of the 158 chief ornithologist at the museum. The following let- ter from Taverner to Lynch shows Taverner’s posi- tion on the subject. “This Division of the Museum, especially the ornithologi- cal section, is constantly receiving favors, especially in the way of rare specimens, from Major Allan Brooks, the artist and one of the leading Canadian ornithologists. Have just received from him four valuable additions to our collection. Major Brooks takes a great interest in the growth of our National collection and takes many oppor- tunities to fill our gaps. These are not strictly ‘exchanges’ but I have tried in the past to render the exchange of cour- tesies not altogether one sided. I have a list of his desider- ata and wherever we can spare any of these items I have supplied them. There is another side to the subject that should be considered. Major Brooks has what is probably the finest collection of “plumages” of North Americal birds extant. Through the war he had willed these speci- mens to the National Museum. I do not know his plans in this regard now but it is most advisable that we retain his interest in this museum, otherwise the collection may eventually go to some museum[s] in the States that I know have already made overtures to that effect. The question is this, — Can I handle this material for this purpose, and in other necessary exchanges to my own best judgment or do I have to leave it to some one else less familiar with our collection and less informed as to its needs, strength and the ornithological questions involved? I have hitherto had full charge of the bird col- lections and have built them up from almost nothing to the best series of northern neararctic birds that there is. It is still not complete (probably never can be made com- plete) and there are many gaps to be filled. I do not see how I can fill them in the future as I have in the past THE CANADIAN FIELD-NATURALIST Vol. 110 unless I have as free a hand as in that past or until I fail to fullfill the responsibility in a satisfactory manner. It is quite important that some decision re the distribu- tion of responsibility in the ornithological section be arrived at shortly. An early decision is urged as I have to acknowledge Major Brooks’ specimens immediately.””? A second issue that caused trouble between Taverner and Anderson was over the way in which the head of a section should deal with the results of a field expedition. One example was the Soper expedi- tion to Baffin Island 1924-1926, and more important, the Soper expedition to the same area in 1928-1929. When a field party collected birds or mammals the material was sent to the Museum to be accessioned, identified, catalogued and distributed into the collec- tions.'°? Each year’s accomplishments in field and lab- oratory work were published in the Museum’s Annual Report. It was customary for the chief of each division to be responsible for the Report which was based on information submitted by the staff of his division. These issues showed that there were substantial differences of outlook in what each believed was the best for the National Museum. Here were two strong willed men struggling for their position in the peck- ing order. By 1936 Taverner was desperate to get out from under Anderson’s control. When he had achieved this by a measure of luck, Anderson was equally anxious to have Taverner disgraced. How he tried to ensure this can be seen clearly in his memo- randum to Deputy Minister Camsell. But the Minister did not “bite’’.!°! CHAPTER 15. The Widening Field of Studies I, 1928-1936 The Matamek Experience 1928 Taverner’s widening field of ornithological studies and experiences can best be shown to have started with his research papers on the Red-tailed Hawk (1927) and the Canada Goose (1931), as well as his arctic patrol voyage of 1929. They continued to expand until curtailed by the violence of the Second World War. During his ornithological career Taverner’s field work fell into three periods. Between 1911 and 1919 his expeditions, with one exception, were in eastern Canada. From 1920-1928 the major ones were in western Canada, with two lesser ones in eastern Canada (1924 and 1928). In the years 1929-1940 he managed two major ones in the arctic and two minor ones in northern Manitoba (1936; 1937). In 1928, accompanied by Harrold, Taverner spent from early June to mid August on the north shore of the Gulf of St. Lawrence. Here they carried out field work in an unusual situation. They spent part of their time based on the residence of a wealthy American, Copley Amory of Washington D.C., at the mouth of the Matamek River where he had rented a salmon stream for the past twenty years. Luckily Taverner sent a descriptive letter about Amory’s “establishment at Matamek” to his sister Ida. Explaining the situation he said it was not a camp but an establishment. cc . A good big house and about a dozen little ones grouped about like a hen and her chicks. A great big store house is the “Factory”. We sleep in the upper story of it. A fine bed room in one side, and a regular biological lab- oratory in the other. A bright little French maid comes in daily and makes our bed and “reds” up the room, fills the pitcher and places fresh towels. Our tent is pitched just behind the Factory in its shelter, and here we work. It is often cold and raw, and our stove is a life saver ... ... The ‘Big House’ has just been all renovated. The floors are all newly scraped and waxed, and the wood- work throughout repainted. Two bath rooms, hot and cold water. A great big living room with a big fire place, a swing seat in front of it, and a table in the middle capable of seating twenty people — guess it is often filled too. Dozens of little cubby-hole bedrooms. ... ... On the rivers are numerous rapids demanding portages. There is a little sort of hut at either end of each portage and in it are a canoe and a skiff — you only have to portage yourself and chattels — canoes are ready at each end. 1996 Amory is a delightful fellow, an ideal host, and a very delicate and charming human. Mrs. Amory is here also — she is quite lame and is evidently rather an invalid and a bit petulant — sometimes more so, but Amory is a won- der of good nature and takes it with a smile.”! Although this was a difficult region to explore on account of the densely wooded coastline, Taverner reported that a collection of land birds was made con- siderably east of any locality on that coast represented in the museum collection to date. An earlier report on the Great Cormorant, called by Taverner “Common Cormorant”, breeding on the north east coast of Anticosti Island was verified, about 100 nests found, and series of specimens obtained. In the Annual Report for 1928 Taverner noted that though this once common species was enormously reduced, its situa- tion was not quite so precarious as feared. A large rookery of [Black-legged] Kittiwake was located at Gull Bay estimated at some 75 000 birds, probably the largest in North America, Taverner wrote. A small rookery of [Northern] Gannets was also verified, making four known breeding places of this species in North America — Bonaventure Island, Gaspé; Bird Rock, Magdalen Islands; Cape St. Mary’s, Newfoundland, and Gull Bay, Anticosti Island. According to Taverner: ““A number of very interesting plumage observations were made on the water birds and considerable additions to our knowledge of their life histories were made.” They also spent from 27 July until 7 August at Natashquan, but this was late in the season and they did not find anything of importance. They returned to Matamek on 8 August. Here they took a number of Razorbills changing into winter plumage with their bill sheaths already shed, and their wings flightless. A Common Murre in the same con- dition was obtained. Reporting this to Charles Townsend in a letter Taverner commented “T never heard it advanced that these birds lost the power of flight in the summer moult but this seems to indicate that like the ducks they do”. In the same letter he said that midway between Anticosti Island and Harve St. Pierre on the east shore they came across two [Atlantic] Puffins in full winter or juvenile plumage with black shaded face and small bills. “Does the Puffin ... take two or more years to arrive at maturity, and do they spend these adolescent summers at sea?” Taverner asked.? It is refreshing to find a well established ornithologist making field observations of points about which he needed fuller information — a sense of excitement at being on the edge of discovery. In the 1920s an ornithologist could not easily obtain information that he wanted. Monographs on particular species had not yet been written, or if they had were printed in journals that had volume indexes, but not cumulative ones. However, Taverner was able to ask Townsend for information because Townsend was CRANMER-BYNG: A LIFE WITH BIRDS 159 something of a specialist on sea birds having spent several summer seasons on the St. Lawrence and its estuary. By chance Taverner met Townsend briefly on the river steamer when travelling from Matamek to Harve St. Pierre. Townsend was about to be taken on a voyage along the St. Lawrence by Harrison Lewis in his capacity as Chief Migratory Bird Officer for Ontario and Quebec. Taverner referred to the period from 8-18 August as “the Matamek meeting”, though it was not a formal meeting but rather a pleasant “get together” of a few like-minded naturalists. In a letter to Bishop, sending news of his field work along the “North Shore”, he wrote: “.. Had quite a meeting of bird men at Mr. Amory’s near Seven Islands. There were Kennard, Glover Allen, Francis Allen and Oberholser. The North Shore was over- run with bird men this summer. A. A. Allen was on it with Lewis for a while. Townsend gave it the once over and Todd worked it from Natashquan eastward. Did not see the latter however.” When Taverner returned to Ottawa he sent Glover Allen some photographs and said how glad he was to have met him.° In reply Allen wrote to Taverner about a student of his by the name of Oliver Austin, Jr., who had made three trips to the east coast of Labrador, and intended to make use of material from there for inclusion in his Ph.D. thesis. He mentioned that Austin brought back two Lapwing Vanellus vanellus “which friends preserved for him ...”° In reply Taverner quipped: “Hope Austin does publish his Labrador stuff but a bird man or a man doing bird work going on three such trips and not able to prepare specimens should be skinned himself ”’.’ Altogether Taverner and Harrold collected 326 birds and 61 mammals, but no rare or intrusive species. It was not an entirely satisfactory field expe- dition. They started too late to see the full northward migration, but they explored the North Shore from Moisie Bay eastwards as far as Anticosti Island and Natashquan. The Matamec River and its tributaries provided them with good fishing.* After returning from the Gulf of St. Lawrence Taverner wrote an article in which he outlined the extent of their expedition, and explained what was understood by the term Canadian Labrador in contrast to “the true Labrador”. The main part was an annotat- ed list of birds observed.? Copley Amory organized an ambitious biological conference at his Matamek residence in 1931 from 24 July — 1 August, to discuss the significance of the “ebb and flow of animal and plant life of both land and sea, and the far-reaching effects of these cycles or fluctuations in abundance on the health, economic status, and general well being of the human population”. The Federal Government and the Quebec Provincial Government sent delegates to take part in the conference. Charles Camsell acted as chairman.!° 160 Although Taverner was not invited to the confer- ence, Amory drew on him for help in trying to estab- lish a sanctuary for eider ducks on the North Shore of the Gulf of St. Lawrence. The Taverners’ Home Life, 1930-1936 Percy and Martha were married at the end of March 1930, spent a fortnight’s honeymoon in Washington and Charleston, then settled into Percy’s home at 45 Leonard Avenue, just south of the Rideau Canal and west of Bank Street. For Percy this was no change but for Martha it was a major change of location and career, though Ottawa was not com- pletely new to her, and certainly the cottage, “Hyla’, on Blue Sea Lake was familiar to her and her son Karel. She did, however, keep possession of her house in Detroit. One of the first things they did, when settled into married life, was to buy a car — a 1929 model T Ford. Since Percy might have “a bad spell” at any time due to his heart murmur, he had never learned to drive.'! Instead Martha soon learned, and thoroughly enjoyed herself when dri- ving. Having the use of a car brought a new sense of freedom to travel to both of them. In November they motored to Cornell University to visit Karel who was studying architecture there. The next ten years of their married life were most enjoyable partly because of the holidays they were able to take in North America, sightseeing and visiting friends of Percy’s in the ornithological world. One of their earliest trips was by automobile around the Gaspé Peninsula in July 1931. He described this holiday briefly in a let- ter to Brooks of early August when he wrote: “Got back Friday afternoon from the Gaspé trip. Had a most enjoyable time. Camped some five days on Bonaventure Island with my wife, almost in sound of the bird cliffs. Spent all our days on them watching the [Northern] Gannets and photographing them.” [Nothing more about their holiday — rest of letter about collecting and photography].!* Laing and Taverner kept up a correspondence which was almost more about gardening and listening to music than about birds. Writing to Laing in 1933 Taverner said that he was working on the rock gar- den. His old rockery, which he thought was the first in Ottawa, had not been built properly. But now that everyone had rockeries, he explained, “... [have to regain my lead in some degree ... we have been scouring the country for nice mossy and worm-eaten limestone and sandstone. We run out in the evening, gath- er all the back seat of the old Ford will hold without drop- ping through, and bring them back in.”! The following summer he wrote to Laing “We have been ‘rocking’ violently all spring. The garden now is almost more rock than soil ... The result is really pretty fine. Lewis [Harrison] just sent down a great pack- age of alpine plants from the North Shore.” Taverner knew that there would be many failures. “Still”, he said, “it is much more fun establishing things from the wild than ordering from a nursery firm.”!* THE CANADIAN FIELD-NATURALIST Vol. 110 To Munro, in 1935, Taverner wrote: “Have had a great time this season with Cypripediums [orchids with pouched lip]. For the first time managed to get acaule [pink lady’s slipper] to survive to a second season and am planting a lot more with lots of peat and spruce needles in the soil to see if that would suit them. Trying the same with the Ram’s-head [C. arietinum] which as you know is pretty rare and supposed to be an even more difficult subject.”!> But it was not always the climate nor the soil that harmed one’s plants as Taverner’s heartfelt words to Laing showed. “There should be a special hell for lying nurserymen. To spend five or six precious years growing trees and then to find them untrue to name is heartbreaking.” In a letter to Soper in the fall of 1936 thanking him for some mammilaria [a species of desert cactus], for his rockery he asked him to look out for a very pretty little rock primrose that he once found growing in scree on the south side of the Cypress Hills. “Tt has a starlike pink flower rising from a whorl of leaves pressed close to the ground.” Taverner would be delighted to have a few roots. He then told Soper what he had been doing. “Have just built a rather ambitious new rockery in the garden. Have it partially planted and now all I can do is to wait for spring to see results. A number of the things I brought down from northern Manitoba seem to be doing well, — others were as could be expected failures or at least unpromising.’”’!7 In addition to gardening and entertaining friends, another thing that Percy and Martha enjoyed between them was music. This went back to the Taverner years in Detroit when Martha taught young Ida music, and the Wiests, Jacob and Martha, together with the Taverners — Ida Van Courtland, Percy and Ida Clare — took part in theatricals which involved them all in singing and making music. In the 1920s Percy and his sister Ida bought gramophone records which they greatly enjoyed playing. When Martha settled into her new role as Mrs. Taverner she started giving piano lessons to a number of children and teenagers at their home. Music at the Taverners’ blossomed and brought them a great deal of deep enjoyment, which Percy passed on to Mack Laing in his letters. Percy also wrote to other friends about their music making. In 1931 he told Rowan “Dr. Eidmann was here a few days or so and I saw some- thing of him. All Sunday he was playing four hand Beethoven symphonies and sonatas with the violin.” !® Percy, when giving Mack advice on what to buy, said: “We have a very extensive library of serious music but have had to hunt the catalogues of the world to get them. They are making a great lot of wonderful things in Germany, Italy, France and England. Our poor little Canadian catalogue is most disappointing.” He then talked about the U.S. catalogue, and its great advantage over foreign ones especially the many recordings made by the Philadelphia 1996 Philharmonic under Stokowski which were consis- tently the best orchestral records. During the winter of 1931-1932 the Taverners were lucky because a Philadelphia firm was selling off many foreign records at half price. They bought big sets of records such as Verdi’s Requiem and Beethoven’s Missa Solemnis “and a lot of other magnificent stuff. Of course now with all the tariff and super tariff and excise, dumping duties and the exchange [rate] there is no more ordering from abroad ...””)? In the same letter Taverner said he had been through the Victor list for 1931 and he recommended nine including Schubert’s Trio, Opus 99, recorded by Cortot, Thibaud and Casals; Schubert’s Seventh Symphony, and Beethoven’s Fifth Symphony. He also gave Laing friendly advice on how to work up to enjoying symphonic music. Beethoven’s nine sym- phonies, he said, were the best introduction to sym- phonic structure, and added: “You will probably recognize a great many of the themes from my old whistling as you did those of the Unfinished.”?° Another piece of information was that the manu- facturers were beginning to bring out long-playing records, and Beethoven’s 5th Symphony was now on a double-sided record. “More long playing recordings can be expected”, he said, “and will be a godsend, and will cost considerably less per composition than the old ones. I think we are on the eve of a great improvement in phonograph reproduction”, he said.! However, by 1933 they could not buy much new music because of the cost due to heavy duties. Instead, Taverner described to Laing the music-mak- ing of their own that they were enjoying. “.... Two or three evenings a week of quintet, quartet, trio and good vocal stuff which is highly exciting. The wife has just started a course in musical appreciation that is also lots of fun and considerable profit. Of course she is trying to get a six weeks regular university course in ten lessons ...”?? He added that when they finished it he hoped that some of them would know the difference between Bach and Brahms. One further letter, written in 1933 to Laing, con- tained good advice on appreciating music. Catchy tunes become tiresome with constant repetition, he said, acting more as a soporific than a spur to the mind. Instead, he wrote: “... [wish you could have attended some of our musical appreciation classes this winter and seen how these big symphonies are built up from definite and balanced plan, the best are really as much the result of a thought out architectural plan as a great cathedral and to listen to them properly is an intellectual as well as an emotional experi- encerre ... [cannot say that I know much about it but the little I do know has opened up worlds of keener appreciation to me than I knew existed. The idea that music could be anything but a sucession of agreeable sounds was a new idea to me as it is to most people. I believe no one can CRANMER-ByYNG: A LIFE WITH BIRDS 161 take such a work as this and know much about it at the first hearing or so ...” In the same letter Taverner told how, one evening, he and Martha had a special musical experience. They went with Percival Price, the carillonneur, up into the Peace Tower in Ottawa to hear some new records he had brought back from Europe. This is how he described it. “... In his wonderful little studio, close under the clock and held up high in the air with lights out and only a glow from the quarterfoil windows all around we listened to Bethooven’s Missa Solemnis, — 12 double sides, well reproduced. It was quite an experience.”?> It seems that Percy and Martha were pleased with a letter from Laing of this time. Beethoven evidently “struck home” and Percy wrote: “That “Ode to Joy’ is a haunting thing and that is what was on the wedding announcement. You were quite foxy to catch it. Only two or three others got it and they were rather advanced musicians.”’+ Early in February 1932 Taverner was ill for a fort- night. Writing to Saunders he said jauntily that it was nothing much. “Have been laid up for a couple of weeks with a nice little attack of the ‘flu’. However as it is the first time I have been really sick in thirty years do not think that I have much to complain of, especially as I am all back on the job and in good shape once again.” To Laing he explained: “I got your letter of long ago when I was flat on my back with the ‘flu’. As it was the first real illness ... in thirty years I have no complaint, especially as it all ended well.”*° But Anderson, writing to Kenneth Racey in Vancouver, put it more strongly. “Mr. Taverner has been away for [from] the office for past nine days, in bed with cold or influenza at first, bringing back heart trouble.”?” A return of his heart murmur was something that Taverner wanted to keep hidden from others, if possi- ble. When he began ‘rocking’ vigorously from 1933 onwards he seemed to want to show his friends that he was strong enough to lift chunks of rock into the car and out into the rockery, but whether he was wise to do so is open to question. About this time Taverner’s eyesight was deteriorat- ing. He first noticed it when taking photos of muskox- en on the northern patrol in 1929. Only a few out of 24 exposures were properly in focus. He had to use one pair of glasses to read the camera marks, and another to focus by. His long-distance sight was as good as ever, but he had to spend half the time putting his glasses on and off again.*® A Study of the Red-Tailed Hawk and the Canada Goose Taverner put strenuous efforts into writing his first book, Birds of Eastern Canada (1919), for a period of 162 five years, and even then realized how very much there was still to be known about the major species. When he began to collect material for his next book, Birds of Western Canada (1926), he realized that he would have to put an even greater effort into working on the taxonomy of species such as the Red-tailed Hawk, the Great Horned Owl, and the Canada Goose when coy- ering the whole of western Canada. By this time in his career Taverner knew that, as a professional ornitholo- gist, he should not be content to write brief summaries of taxonomic relationships within a species, but must undertake a monograph or two on a large species, showing its several races, extending across North America, and where they intergraded or not.’ Furthermore, competition or jealousy among ornithologists existed as early as the 1920s. To have a monograph, on a difficult genus or species, published by a respected ornithological journal, or published as a book, and then to receive some reviews that gave it a favourable reception — this was the height of accla- mation for an ornithologist in that period. By the time that Taverner had started, in 1932, to put material together for his third book, Birds of Canada, he had published two monographs that had caused him much work, thought and discussion.” Taverner began working on the Red-tailed Hawk early in 1925 and corresponding with others on its colour variations. The problem he set out to solve was: what viable subspecies of Buteo jamaicensis should be recognized by ornithologists? During 1925, Taverner and Brooks were corresponding about Harlan’s Hawk (Buteo jamaicensis harlani). Brooks had discovered, breeding in the Lake Atlin area of northwest British Columbia, Harlan’s Hawks in sufficient numbers to suggest that it was a discrete geographical variant.*! In reply Taverner said that until this time he had a “half formed conclusion that it [Harlan’s] was an extreme phase of calurus” [Western Red-tailed Hawk (B. 7. calurus)]. The only variation he could find for harlani was the marbled or mottled tail.’ By the end of 1925 Taverner told Brooks that he now expected that he could write a paper on the situation as regards the Red-tails, with coloured illustrations of the various plumages. “T intend to make them rather diagrammatic, pictures of skins instead of living birds, and all alike in outline and form so that they can be readily compared. Will show you some of these drawings one of these days to show you what an astonishing lot they are.” In the same letter Taverner wrote that Laing had collected some very important specimens which might turn out to be key specimens to the whole situa- tion.*> Also Taverner had been obtaining interesting information from Norman A. Wood, of the University of Michigan Museum of Zoology, who had a collec- tion of harlani, though they were all migrant speci- mens, mainly from Grafton, North Dakota.** While Taverner was working on his monograph he was also hurrying on with Birds of Western Canada THE CANADIAN FIELD-NATURALIST Vol. 110 for a printer’s deadline of early 1926. In this book Taverner stated that characteristic adults have brick- red tails, but western birds are so variable that colour, in the tail or any other part of the body, was an uncer- tain criterion for identification. In discussing sub- species of the Red-tailed Hawk he wrote: “The Eastern Red-tail Buteo borealis [jamaicensis] bore- alis extends westward through Ontario; but, beginning with the prairie sections in Manitoba, it intergrades, inter- mixes, and interbreeds so thoroughly with the Western Red-tail Buteo borealis calurus that there is great difficul- ty in defining the distinctive characters or saying where one begins and the other leaves off. ... Besides these two forms, borealis and calurus, there are two others recognized by the Check-list that present peculiar problems to the Canadian ornithologist — Krider’s Hawk Buteo borealis krideri and Harlan’s Hawk Buteo borealis harlani. These are so variable that it is dif- ficult to say just what are their distinguishing characters or whether after all they are not respectively just the light and dark extremes of the Western Red-tail.”*> At this point we should examine Taverner’s study of the Buteo borealis |jamaicensis| which was pub- lished in an issue of the Museum Bulletin a few months after his book.*° Because considerable materi- al had become available it seemed that this was a good time to make a critical study of the species. A large number of specimens, mainly of breeding birds, in many cases of complete families, were collected across Canada. Taverner was also able to borrow a series of migrant and winter specimens taken in North Dakota and Arkansas from the Museum of Zoology, University of Michigan, and important collections were studied at the Museum of Vertebrate Zoology, University of California, and specimens from various other university and private collections. This enabled Taverner to bring together a series of 157 skins for direct comparison. The body of Taverner’s study consisted of nine pages of discussion. He began with a general intro- duction. The species was considered by the AOU Check-list of that time to be composed of five sub- species: Buteo borealis borealis (Gmelin), Eastern Red-tail Buteo borealis calurus (Cassin), Western Red-tail Buteo borealis krideri Hoopes, Krider’s Hawk Buteo borealis harlani (Audubon), Harlan’s Hawk Buteo borealis alascensis Grinnell, Alaska Red-tail 37 Taverner pointed out that the principal characteris- tic of the eastern Red-tailed Hawk, the type form of the species, was its constancy of plumage. Over most of its range it had only a single phase [morph] which was brown. After six pages of rather complicated explanation of the differences between the forms borealis, calurus, krideri and harlani, and the problems that ornithologists studying this species encounter, Taverner summarized his conclusions: — the Red- tailed Hawk of eastern Canada was pure borealis, a bird of practically constant character. 1996 “The prairie provinces are inhabited by birds that are pre- dominantly borealis, but with calurus features, more or less common intrusion of krideri, and at least a sporadic influence of harlani towards the west. Southern British Columbia is inhabited by practically pure calurus, and the far northwest, at least centring about the British Columbia-Yukon-Alaska boundary intersection, contains birds that are predominantly harlani, but with strong intermixtures of calurus and krideri.” Five pages of bibliography up to the year 1926 fol- lowed. Last but not least were three pages of hand- painted coloured plates which explained the author’s discussion in the paper satisfactorily. Plate I depicted central tail feathers showing common variations in the principal subspecies at that date. Plate 2 showed 8 diagrammatic representations of body and tail col- orations in krideri and calurus. Plate 3 showed 8 dia- grams of characteristic body and tail colorations of calurus and harlani. Taverner’s study was reviewed briefly in The Auk by Witmer Stone. He summarized the main argu- ments and said that Taverner had “thrown much light upon a puzzling problem and we see no reason why his general conclusions are not sound”. But he questioned the advisability of upsetting the current name of the Western Red-tail. “It is not a good plan to replace a certainty with an uncertainty in nomen- clature”.** A longer, more critical review by Swarth appeared in The Condor. Swarth recently had a study of Harlan’s Hawk published, and had crossed swords with Taverner at the AOU meeting in Ottawa in 1926. (see Chapter 13) He said that Taverner had produced an excellent summary of the range of variation that exists among the more northern of the currently rec- ognized subspecies. Swarth continued: “His descriptive accounts of subspecies and individual birds, together with the helpful colored plates, form an important contribution towards an understanding of this difficult species ...” But Swarth could not accept Taverner’s conclusion — that the current five names covered two forms only, an eastern race, and a western race, to which he applied the name harlani. Swarth then discussed the Red-tailed Hawk in detail and concluded that Taverner’s treatment of the nomenclature should not be followed.*? Taverner had no expectation that his suggested changes would be followed. However, with the publi- cation of Peters’ Birds of the World volume 1, 1931, he considered that the treatment of the Red-tailed Hawks therein justified him in reviewing his previous study. Eventually his paper “Taxonomic Comments on Red-tailed Hawks” was published in The Condor in 1936. Taverner commented on a form Buteo bore- alis alascenis Grinnell that had been recognized by Peters but was not in the 1931 Check-list. He also commented on a subspecies recently described, Buteo jJamaicensis fuertesi Sutton and Van Tyne, from CRANMER-BYNG: A LIFE WITH BIRDS 163 Texas. Taverner was given the privilege of examining the type series of this race.*° The Canada Goose During all the years that Taverner was ornithologist at the National Museum he was involved in the prob- lem of unravelling the species of geese that breed in Canada. From the beginning of the 1920s Taverner was working on a book about Birds of Western Canada, and was writing to friends and acquaintances with knowledge of geese found in the western provinces of Canada. Brooks was the best source of information on western birds, especially ducks and geese. By 1923, Rowan, at Edmonton, was corre- sponding with Taverner on the Cackling Goose, a race of the Canada Goose, which shows an example of the kind of problems of identification and distribu- tion that ornithologists faced during the 1920s and 1930s.*! When Taverner’s Birds of Western Canada was published, those making a study of the species/sub- species of geese that might be seen in Canada were able to compare short summaries of information on them, together with coloured illustrations by Brooks and drawings of beaks of each by Taverner. In his account of the Canada Goose, Taverner included the subspecies Hutchins, Cackling and White-cheeked Geese (pages 107-108). Writing of the Canada Goose Branta canadensis as a species, he said: “Although few birds are as well known to the sportsman and general public, none is so little understood systemati- cally by either scientist or layman.” (pages 105-106). Taverner said that on the prairies it was being exposed to the destructive effects of civi- lization, and feeding flocks were hunted by automo- bile from one patch to another. “Its disappearance would be a serious aesthetic and economic loss”, Taverner wrote, and became quite eloquent. “Tt is a noble bird, a point of interest in any landscape. No one fails to thrill at sight of its long V-shaped flocks fly- ing over, or at the sound of its wild barbaric music com- ing down through the twilight. Domestically, the Canada Goose is a model for man. Unlike Ducks which mate for the season and then part for ever, Geese mate for life, mourn a lost mate, and are not easily comforted. Both sexes assist in the responsibilities of family life and if necessary share the supreme sacrifice in its behalf. We speak of the Goose as the personification of foolishness, but the Canada Goose is one of the most intelligent and wiliest of birds and exhibits occasional bits of strategy that are astonishing.” Taverner next produced an eleven-page study of the problems of taxonomy within the Canada Geese.*? He began by saying that recently much interest had been shown in the characteristics and complex rela- tionships of the various geographical races of Branta canadensis. Although several attempts had been made to unravel them, the difficulties posed by dealing with what he called “mixed migrant material far from their 164 distinctive distributions” had been great, and no con- vincing solution had so far been presented. Now that breeding and summering specimens of geese, brought by Dewey Soper from Baffin Island, were available in the National Museum of Canada, Taverner considered that another attempt to clear up some of the problems should be made. After studying many specimens in various collections and considering the whole subject for a number of years, Taverner put forward several points in order to eliminate confusion in distinguish- ing five distinct groups. In the field, Taverner stated, “experienced sportsmen had far less difficulty in rec- ognizing the various forms of Canada Geese than had the systematist working with specimens in the labora- tory. The differences between the living birds are shown in stance, action, voice, and in particular habits.” The material he intended to review, he explained, could be divided into the following groups, each with well-defined ranges: I. A large, light-breasted form breeding across the con- tinent. Undoubtedly B. c. canadensis. II. A large, dark-breasted form breeding on Queen Charlotte Islands and probably adjoining localities in Alaska; evidently B. c. occidentalis. Il. A medium-sized, light-breasted form breeding in the northwest. The bird hitherto generally called B. c. hutchinsi. IV. A small, dark-breasted form said to breed along the east coast of Bering sea, B. c. minima. V. A small, light-breasted form breeding in the eastern Arctics [sic]. A hitherto unrecognized race.4 At this point Taverner began a detailed analysis of the five groups starting on page 31 and continuing through page 39 of his study. This was a closely rea- soned discussion of the problems encountered in try- ing to distinguish each of the five groups from the data available to him then. Rather than attempt to pre- sent a précied version of what Taverner wrote here, I can recommend his full discussion to the reader, since only by having it before one is it possible to follow the links, or lack of them, in their complex relation- ships. Moreover, the six tables providing the dates, places and names of the species together with their measurements, that are under discussion can best be understood when they can be studied in detail. After ten pages of discussion Taverner summed up his findings as follows: “The degree of relationship between these various forms of the genus Branta is difficult to arrive at. Though the breeding specimens under review show clear enough lines of demarcation between the groups, intermediates are said to (and probably do) occur. Whether these are true intraspecific intergrades or extraspecific hybrids is not clear.’ As a result Taverner’s revision of the taxonomy of the Canada Goose in 1931 recognized the following species and subspecies: Canada Goose Branta canadensis Linnaeus Eastern Canada Goose Branta canadensis canadensis THE CANADIAN FIELD-NATURALIST Vol. 110 Western Canada Goose Branta canadensis occidentalis Lesser Canada Goose = Branta canadensis leucopaeria Cackling Goose Branta minima Ridgway Richardson’s Goose — Branta hutchinsi Richardson *° Taverner was by no means satisfied with his study of the Canada Goose. In a letter to Rowan he said: “Glad you like the goose paper. All I am uncertain about is the specific distinction of hutchinsi. I might have been better advised to have made it a subspecies of canadensis though I do think that minima deserves that (specific) sta- tus. Another question that I am not settled in my own mind about is the applicability of the name “/eucopaeria” to the light breasted middle sized bird that we have heretofore called hutchinsi. However if one waited until they determined positively every side issue of so compli- cated a subject little could be accomplished. It was too bad that I should have to ball up the nomenclature so but there was no way out of it. Richardson’s hutchinsi was not the bird that has been so called all these years.”*” The Fourth Edition of the AOU “Check-list of North American Birds” (1931) disregarded Taverner’s suggested changes to the species Branta canadensis. He was well accustomed by this time to doing his best as he saw it, and being patient. In Birds of Canada (Taverner 1934) nine pages of print and three of coloured illustrations were allotted to geese inhabiting Canada, as well as seven black- and-white drawings by Taverner. In a short entry for Ross’s Goose (Chen rossii) Taverner noted that its nesting location was unknown and information on its distribution was “problematical’”’.*® Writing to Brooks, who was in California in the spring of 1933, Taverner said that he had heard from Grinnell who told him that Phillips, Moffitt, Swarth and Brooks had dinner at Grinnells’, where they had a general discussion on the Canada Geese. Brooks and Moffitt thought there was still another goose to be named. Taverner agreed though he had no material with which to demonstrate his suspicions.” In reply Brooks agreed that there seemed to be another goose to be described and explained: “It is known to the professional goose hunters of California and occurs in flocks of birds that all show the same characters — an intermediate size between — leu- copaeria and minima with a brownish coloration and a wide white collar running to a point on the back. It has not the narrow blackish area below the stocking or white collar on the throat which is the very best characteristic of minima. Many specimens in the M.V.Z. [Museum of Vertebrate Zoology, Berkeley] & in Moffitt’s collection. The former are labelled minima by Swarth. A range will have to be found for it before it can be described.”>? Taverner was interested with Brooks’ description and the Canada Goose will be as involved as_ the Juncos. Your description of it 1996 - “brownish” is like the type of occidentalis which is not mouse gray below but distinctly and pronounced reddish. I have always regarded it as a freak bird, — it does not seem due to rust stain.”>! An important new development came when a brief paper by W. E. Clyde Todd, describing a new eastern race, was published in The Auk in 1938.>? When Taverner retired in 1942 he had been unsuc- cessful in discovering a new species of North American goose, though several of his fellow ornithologists had succeeded. However, in 1951 Jean Delacour proposed a new subspecies of Canada Goose be named Branta canadensis taverneri in his honour.® In his Waterfowl of the World he described its characteristics as follows: “Resembles B. c. parvipes, but smaller, with shorter bill and neck, and generally darker, although very variable in colour, some specimens being as pale as parvipes, others as dark as /eucopareia. It also resembles the latter closely, but it is generally larger and it has a differently shaped bill, less high near the base and broader towards the tip, with a smaller nail. The white neck ring is often lacking, and when present it is narrow or incomplete.” Under Distribution he wrote: “Breeding areas as yet poorly known. Breeds probably throughout the interior, ten to fifty miles from the coast, from the base of the Alaska Peninsula to the Mackenzie River delta. It intergrades with minima in the Wainwright area, and with occidentalis in the south-west. It certainly also intergrades with parvipes, but the ranges and intergradation of the two forms are yet to be worked out. The identification of the various populations remains dif- ficult, and until further knowledge is acquired we unite them under the name taverneri. Winters from Washington to Texas and Mexico, mostly in the large interior valleys of California.”** It was not recognized in the AOU Check-list of 1957, but it was included by W. Earl Godfrey in The Birds of Canada 1986. In this way Taverner received posthumous recogni- tion for his years of work towards unravelling the species Branta canadensis. With the temporary cessation of summer expedi- tions Taverner found that there seemed to be more routine work than ever. Writing to Brooks he com- plained “... one gets lost in detail ... Iam loaded with routine now as it is”. In the same letter he admitted writing “over a thousand letters a year”.°° Meanwhile there was plenty of ornithological research to be done in the museum even if work in the field had stopped. There were records to be kept up to date, especially concerning the distribution maps. The information they contained was often required. A letter from Leon J. Cole, professor in the Department of Genetics at the College of Agriculture, Madison, Wisconsin, to Taverner in 1932 asked if it was possible to obtain live Mourning Doves from the northern limit of their range. This, he explained, was for use in studies following the lead by Rowan, on the CRANMER-BYNG: A LIFE WITH BIRDS 165 effects of periodicity of light in relation to migration and distribution of that species.°’ In reply Taverner sent Cole a rough copy of the museum’s distribution map of the Mourning Dove, with a note that he had data on all the spots recorded on the map. For Ottawa he wrote: “The species is rather rare here ... though I think slowly increasing yearly. It is more common at Kingston. Common throughout the lower peninsula of Ontario. In Manitoba north to between the big lakes. In Sask. to Quill Lake. In Alberta not quite to Edmonton. In southern B.C. and farther north along the coast.’>8 Before discussing Birds of Canada (Taverner 1934) we can note an event that took place in 1928 when an edition of 3000 copies of sixty coloured postcards of birds was issued. According to the Museum Bulletin: “The coloured illustrations were made from plates of “Birds of Western Canada,” some from “Birds of Eastern Canada,” and some from new paintings by Major Allan Brooks. Each card contains one coloured illustration, accompanied by descriptive text, and the cards are sup- plied in sets, in carton, for sale at $1, and also loose, for sale at two for 5 cents.”>? The printing of the colours was good, and the descriptive text, including a little information on the habitat of each bird, was useful. The paintings by Brooks carried these “picture postcards” far above the usual standard of collectible cards. Their publication served as a trial run, both for the King’s printer, and for the public which would get some idea of what to expect from a new Taverner-Brooks collaboration — this time dealing with all species of birds known to occur in Canada. Birds of Canada 1934 The earliest official proposal to combine Taverner’s two books, Eastern and Western Canada, into one large one was made by Collins to Camsell late in 1928. Taverner informed Brooks that Collins had agreed to Brooks making 35 new bird pictures for a book to be called Birds of Canada at $2.00 each picture. While Brooks and Taverner were working on the text and illustrations of the new book, and much ornithological correspondence was flowing between them, a timely development was taking place in the Royal Canadian Mounted Police. An RCMP officer, stationed in the Yukon, wrote to Hoyes Lloyd in 1933, explaining that he was fond of birds, but had no one to turn to for help in identification. He had managed as best he could by using Taverner’s book “without which of course I could have done practi- cally nothing”. He added that by reading Bird-Lore he knew that Hoyes Lloyd was a bird-lover.®! In October 1934 Taverner informed Collins that he had been told that the RCMP had officially applied to the Parks Branch for bird books to guide their offi- cers in enforcing the Migratory Birds Act. They 166 were directed to obtain copies of the forthcoming Birds of Canada. In reply the RCMP said that it would supply each police post with a copy. Since there were approximately a thousand such stations manned, Taverner said that the museum could expect an order of that number. In addition Lloyd had told him that an order of about 500 copies in French and English from the Quebec government had been received. Much of the correspondence between Taverner and Brooks in these years was about the descriptive text of the Birds of Canada. Here is an example when Taverner told Laing: “T wish I could tell Wrights from Hammonds Flycatcher when I see them. Have no great confidence in the wing formula. They intergrade in this and if so why not over- lap? Probably field observation is the best criterion.”° In Birds of Canada, under Wright’s Flycatcher, he wrote that it was a little longer than Hammond’s Flycatcher, and shorter than Traill’s. Of general colour design of dusky-olive and white, like the Wood Pewee, but smaller. After a paragraph on dis- tinctions and one on field marks he concluded, say- ing: eMery closely resembling the Least Flycatcher and Hammond’s and the wing formula (Figure 398) is proba- bly the best distinction”. In 1935 Brooks was writing more about collecting skins for the museum and said that if Taverner required pelagic skins from the west coast Arthur Peake would work hard in collecting them, and added: “But for the Lord’s sake send me a catalogue of what you have, here am [ anxious to fill your gaps gratis and you will not take advantage of it! Species and plumages”.© Birds of Canada was a solid book of 445 pages, measuring 9 1/2 by 6 1/2 inches, and weighing about 3 Ibs in the 1934 edition. It is too substantial to read in an ordered sequence, and it is best dipped into at random while enjoying the many coloured portraits of birds by Brooks and Hennessey. One way of sam- pling the book might be to read a page or two of descriptive writing about the following birds which can be recommended for reading for pleasure: Ruffed Grouse 156; Whooping Crane and Sandhill Crane 168-171; Killdeer 181-182; Long-billed Curlew 189- 190; Eskimo Curlew 191; Willet 195-196; Peeps (small sandpipers) 201-202; Hudsonian Godwit 209 “a fine bird on the verge of extinction”; American Avocet 211; Wilson’s Phalarope 213; American [Eastern] Screech Owl 257; [Common] Nighthawk 268-269 — the good they do; Calliope Hummingbird 275; Black-capped Chickadee 311-312 — “their shape, a round bundle of feathers with long tail and no appreciable neck, and contrasting face marks”; House Sparrow Introduced 366-368; Bobolink 368- 369; Western Meadowlark 370; Yellow-headed Blackbird 371; Redpolled Linnet [Common Redpoll] 388-389. THE CANADIAN FIELD-NATURALIST Vol. 110 Birds of Canada was very well received by reviewers and the general public. J. Grinnell reviewed it so warmly in The Condor that this has been reproduced fully here: “A truly beautiful as well as useful book is P.A. Taverner’s new ‘Birds of Canada’ ... While in a general way a combination of the author’s previously published two volumes, Birds of Eastern Canada and Birds of Western Canada, the text has been entirely rearranged and there is much new information. The abundant col- ored illustrations are chiefly from the brush of Allan Brooks, while most of the line drawings are the work of the author. The subjects of these latter are well chosen to show important structural or field characters of species; indeed, some points are thereby brought out that we do not recall having seen portrayed or even mentioned in any other work. Cooper Club members living in the northwestern United States will find this book, we think, quite the best single volume to own as a guide to their local bird-life. General topics and ques- tions of the day are dealt with in Taverner’s well-known common-sense way; and they apply south of ‘the line’, as well as north. The book was written with the aim of stimulating ‘an interest, both esthetic and practical, in the study of Canadian birds.’ It must surely fulfill this aim "iG! The review by W. Stone in The Auk contained a few mild criticisms but otherwise referred to it as an excellent work which would do much to advance ornithological interest in Canada.®* In Britain it was reviewed by W. L. Sclater in The Ibis, who had nothing important to say since he knew nothing, at first hand, about the subject. However, the last sentence of his review was encouraging to potential readers: “Tt has, moreover, one supreme merit, it can be obtained for the trifling sum of two dollars (about nine shillings), a remarkable price considering the coloured plates, which are eighty-seven in number.”©? Taverner’s many black-and-white illustrations were also well received. One writer for a Montreal newspaper congratulated him on the illustrations throughout the book. Replying, Taverner said that he considered Brooks’ work outstanding. As regards his own he said: “Of course my little pen and inks are only hammer and tongs draughtsmanship. I do them because it is easier to do them myself just when wanted than to tell some one else just what I want.” Just at the time when Anderson was mounting a campaign to ridicule Taverner’s ornithological knowledge and ability something very heart-warm- ing happened to him. He received a letter notifying him that he had been elected Fellow of the Royal Society of Canada, and asking him to send the Secretary a note on his academic degrees. Even when about to be awarded an honour Percy Taverner was faced with the unpleasant fact that he had no academic degrees to support him. But he made the best of the situation in the following letter to the Secretary of the Royal Society of Canada: 1996 “Dear Sir:- I take great pleasure in acknowledging your notifica- tion of my election as Fellow of the Royal Society of Canada and assuring you that I appreciate the honor. Unfortunately I have no strictly academical degrees but am Fellow of American Ornithologist’s Union and Colonial Member of British Ornithological Union both being elective and of some distinction in ornithological science. I hope to be able to attend the May meeting at Hamilton to be presented in person. My full name and address as below. Sincerely,- Percy Algernon Taverner, National Museum of Canada, Ottawa, Raptor Control Taverner tried to keep on friendly terms with as many naturalists as possible because, as chief ornithologist at the National Museum, he wanted all the information and cooperation he could get from interested people. But there were a number of con- tentious issues in which he could not help becoming involved. One such was the argument, which broke out in 1923, about whether it was necessary to control “vermin” in bird sanctuaries. This was sparked off by an article by J. A. Munro published in The Canadian Field-Naturalist, followed by an editorial answering it. Two long and peppery letters appeared in a later issue — one by Rowan and one by Brooks, and two from ornithologists in the United States. These contri- butions were discussed in a second editorial.’* As a result of this opening fusillade more supporters got into the fight on either side. In the west Brooks and Rowan squared off with Munro, in the east Jack and Manly Miner, together with E. R. Kerr, wrote strong- ly-felt letters about the necessity of destroying the fol- lowing vermin: Great Horned and Barred Owls; Cooper’s, Sharp-shinned, Red-tailed, Red-shouldered, Marsh [Northern Harrier] and Sparrow [American Kestrel] Hawks. Crows at all times. Harkin, as Commissioner Dominion Parks Branch, and Hoyes Lloyd as Supervisor Wild Life Protection, together with W. E. Saunders bore the brunt of some lengthy letters from Kerr and Manly Miner.’? Taverner, although he was tryng not to take sides, also became involved when Manly Miner wrote the following, as from his father.to Harkin, as part of a long handwrit- ten letter [uncorrected]: “I would like to see Mr. Percy Taverner and Mr. W.E. Saunders go to a crows nest and a few days before young crows fly some evening shoot the young crows and exam- ine their stomach. They will not only find yokes of eggs but they will also find shells of song and insectiverous birds eggs in the crows stomach. Why I say Mr. Taverner and Saunders I don’t think Canada can produce two men who have examined any more birds stomachs than these two men and have had years of study afield.” Manly added a rather imperious P. S. without, per- haps, realizing its implications: CRANMER-BYNG: A LIFE WITH BIRDS 167 “Read this to Mr. Taverner and Gibson and then please forward to Mr. D. McDonald, Deputy Minister of Fish and Game, Toronto. I want to keep these men informed and as I have no typewriter, it will save me writing 3 more letters.”/4 Was the Commissioner to send for Taverner and Gibson to come to his office, and then read this long- winded letter to them while they earnestly listened?” The following day Kerr wrote again to Harkin, thank- ing him for his letter, and for a copy of a letter from Taverner to Harkin. Kerr said that crows were 85% predators and harmful and should be destroyed. “I am not prejudiced, but positive”. He then turned to deal with Taverner. “Like all ornithologists, Mr. Taverner waxes evasive when discussing hawks and owls during the winter months. I still claim all hawks and owls with us during the winter are predaceous and do more harm than good.” Later in the same letter he wrote that ornithologists had no interest in sports or sportsmen, therefore their writing and views had no appeal to sportsmen. “Possibly, of all of the ornithologists I have come into contact with, directly or indirectly, Mr. Taverner and Mr. W.E. Saunders seem to be the most open minded of them all, and these men may possibly make more headway among sportsmen and farmers if they will try to agree that hawks and owls of all kinds with us during the winter are los harmful than good and should be treated according- ly.” Taverner had been in touch with Manly Miner reg- ularly in the past few years, and remained an active correspondent. Writing in February 1926 Taverner began: “IT notice from a number of sources that you are killing quite a number of hawks and owls at your place this win- ter. I do not want to take up the subject of the advisability of killing all these birds for the sake of a few admittedly bad ones but I would like to ask you when they are killed if you would not send them to the Canadian National Museum here at Ottawa. It seems too bad to waste them, why not let them do as much good here as specimens to balance a little of the evil they may have done in life?” He ended the letter with a P.S. “I note a common reference to Cannible Birds. [This was a term that Jack Miner used.] ‘A bird must eat its own species to be a cannible’. An owl that eats a quail is no more a cannible ... than we are cannibles when we eat cows...”’’ Manly wrote asking if he had seen the let- ter about the crow controversy that he had written to Harkin. Taverner replied that it was an old controver- sy, that he had been anti-crow consistently, but the problem was that the crow had not been seriously investigated in Canada. “There is no exact informa- tion as to its status”, he wrote. Later in the letter he said that he had received a copy of the editorial in the Kingsville paper and continued: “Now its conclusion I personally think is correct but it is so full of errors, half statements and silences that if I had not already made up my mind as far as I can without a thorough investigation that I would class it with other floods of selfish propaganda we have been flooded with 168 since the war ‘and would incline to the other side of the question. Calling the Crow an Arch-fiend is absurd. He eats what there is to hand as is necessary to his existence, — just as we do. I suppose that some may say that you and your father are quite devilish because you prevent honest Hawks, Owls and weasels from making a living in the way the Lord intended they should.”7® This correspondence continued for another thirty letters to the end of 1926, each man trying to put his own point of view in the best possible light. At the same time Taverner and Manly wanted to remain on reasonable terms because each had things to gain from the other, as we shall see. On 20 April the Miners tagged 240 Canada Geese. Taverner, thanking Manly for his letter with this information, which reached Ottawa the next day, said: “Glad you have banded so many geese but I do wish you would use a regulation tag that could be recognized after you and your father have gone or have forgotten about the details. I think you are missing an opportunity to make good work better.” Manly’s answer to this was: “Re our tags on geese. We have our Post Office address and date on tags and then we keep a record of how many are put on at each catch and so forth. I think the regula- tion Biological Survey tags should adopt our plan because we get more or as many reports from tags that we didn’t put on which we send report on to Biological Survey [sic].””°° During the summer of 1926 Taverner was busy with field investigations in Alberta, and in the autumn with the AOU meeting in Ottawa (see Chapter 13). Their correspondence on crows, hawks and owls in relation to bird sanctuaries resumed again in November, when Taverner wrote that he approved of Manly’s policy regarding hawks in their sanctuary in winter. The Sharp-shinned, Cooper’s and Goshawks were certainly “bad actors”, and a bird or game sanc- tuary was no place for them at all. That a few of the large Red-tails and Red-shoulders should suffer acci- dentally while the others were being controlled could not be helped, but Taverner advised padding the traps’ jaws to eliminate as much damage as possi- ble.8! Taverner realized that even the Red-tails and Red-shoulders might indirectly cause some harm by the disturbance they make, but he would judge this of secondary importance. “Personally I am rather fond of birds of prey, and have a very real admiration for their bold, free ways, bravery and hardihood. These traits are admirable in men why not in birds, beasts. We can admire an enemy even though we have to control him for our own welfare. But I would not protect even hawks at too great a cost to other birds any more than I would eliminate all hawks for the sake of those birds. There is a middle course and ... in all things the middle course is the right ... way between two fanati- cal extremes. It is the hardest course to follow too.’*? Ever since Taverner had come to know Jack Miner, and soon after through correspondence with his letter- THE CANADIAN FIELD-NATURALIST Vol. 110 writing son, he had been trying to steer the Miners away from a violent attack on hawks and owls of all species. A reading of the full correspondence between Taverner and Manly in 1926 shows that although he struggled to convince Manly not to harm them, this was impossible. Jack Miner’s mind was firmly made up about the “wickedness” of hawks and owls, and his almost divine duty to exterminate them. All that Taverner could do was to try to mitigate the worst Miner excesses. Hoyes Lloyd, likewise, in his official position as “Supervisor”, Wildlife Protection, could do no better. In a letter to Taverner of December 1926 he wrote: “In connection with your coaching of the Miners on ornithological subjects you may have an opportunity some time to call their attention to the fact that the band- ing of birds with authorized numbered bands is desirable. The Miners do not hold any Permit allowing them to band birds for scientific purposes and of course no such Permit can be issued unless they use authorized official bands ... The Miners’ work has been of such a calibre that this point on banding has not been raised officially and per- sonally will not be raised officially so long as Jack Miner lives.”®? Issuing permits for collecting birds for scientific pur- poses A subject which began with the passing of the Migratory Birds Convention Act in 1917 emerged again in 1926 when Lloyd wrote to Taverner for his opinion on the advisability of reducing the limit on the number of permits issued to collectors of birds for scientific purposes. Taverner answered with a seven- page letter explaining why a proposal to reduce the limit to two birds per species should not be adopted. In his last paragraph he explained that “the scientific investigator, working for the advancement of knowl- edge, should not be treated as a suspicious suppliant for special favor”. He should not be subject to more interference, Taverner claimed, “than is necessary to establish his good faith and prevent others obtaining his privileges under false pretenses.’’** However, by 1931 dissatisfaction with activities of certain local collectors in British Columbia exploded because of what was considered lax enforcement of the regulations by those responsible for doing so. Lloyd sent Taverner a copy of a letter addressed to the Parks Commissioner, Harkin, from a William N. Kelly of Vancouver. In this way Taverner became involved in a contentious issue. The nub of the com- plaint concerned a confrontation between Kelly, rep- resenting a number of local ornithologists, and Munro in his position as Chief Federal Migratory Birds Officer for Western Canada. Kelly stated that it was well known “in this district that collectors do violate the law and the ethics of professional work in ornithology ...”. He also made the accusation that the contributions of collectors “to scientific ornithological knowledge are not commensurate with the destruction 1996 of birds taken to build their private collections.” He told Harkin that he should not expect him to act as a public prosecutor and take on himself the duties that should be carried out by the Federal Bird officer. Kelly continued that he would not have to take on these duties “if the federal bird officer was faithfully performing these duties for which he is employed and paid by other bird lovers who are not obsessed by the desire to have a large collection.” [emphasis added]. This was a dig at Munro who was known in the district to have a large private collection of speci- mens. After several more paragraphs telling Harkin very directly how his officers should perform their duties, the writer tried to make “the Ornithologist of the National Museum” appear incompetent.* Such a forthright letter caused problems at the Dominion Parks Branch. Taverner was sent copies of the correspondence by Lloyd, and replied to some of Kelly’s points as follows: “The collection of specimens is absolutely necessary for any sound study of ornithology: That no institution or private collector can possibly col- lect in all the localities that it is necessary to have speci- mens from. It will have to rely on collectors elsewhere to supply its needs. That the person who satisfies these needs is doing a pub- lic service just as much as he who supplies any other other human need and as such is entitled to a living remu- neration therefore. Whatever knowledge we have of birds of Canada, North America and the world has been almost entirely through the collecting ornithologist.” As regards the figures of the total bird mortality in B.C. given by Kenneth Racey in an article in The Murrelet, which Taverner considered authentic, and Kelly considered doubtful, Taverner produced a rather weak answer. In his final paragraph he tried to turn the tables by saying that when Mr. Kelly and his co-thinkers could add some new and original infor- mation to ornithology “we will respect his views more”.*° In a letter to Harkin, in answer to his enquir- ing about the help that collectors under the permits issued by his department had been to Canadian ormithology, Taverner replied that eighty percent of the observations made by amateur ornithologists unsupported by specimens or by familiarity with specimens contained an element of doubt. On the contrary, the work done by such collecting ornitholo- gists as, — Allan Brooks, J. A. Munro, K. Racey, H. M. Laing, G. C. Harrold, Wm. Rowan, Ronald M. Stewart, L. B. Potter, W. E. Saunders, Robie Tufts, Gus Langelier, and a host of others in Canada and the U.S. could hardly be estimated. In all these, thanks to specimens collected and to familiarity with speci- mens, all element of uncertainty was practically elim- inated. Taverner continued to give further examples, finally concluding that much current ornithological work was correcting errors made in the past by accepting evidence unsupported by specimens. CRANMER-BYNG: A LIFE WITH BIRDS 169 Taverner had made a strong case for “scientific col- lecting”’, but the gist of Kelly’s case was against com- mercial collecting being justified under the guise of permits for scientific collecting.*’ When Taverner and Laing were carrying out field work at the invitation of Brooks, in 1922, they stayed at the foot of Vaseux Lake for a short while. Their nearest neighbour was a friend of Saunders’ by the name of Parham. (See Chapter 12, notes 86 and 94.) In November 1932 Taverner received a letter which began: “Dear Sir, I have not, like my brother, the pleasure of knowing you personally, but your book, “Birds of Western Canada”, has been by my side to guide me ever since its publica- tion, and it is owing to the Introduction therein that I now write to you. On pages 16 and 17, under the headings “Bird Study” and “Permit Principles”, some very good and useful advice is given. With this your fellow bird- lovers cannot seriously disagree, even though many of us wish that the regulations allowing the killing of non- harmful birds could be more severely restricted.” The author was H. J. Parham, who came to British Columbia in the 1890s, making his home at the foot of Vaseux Lake for eleven years. When Taverner was there he had already left. The gist of his letter was that he, and others, were not satisfied with the Government’s part in the protection of birds. There was a general feeling, he said, that the Department’s representatives were far more interested in the “col- lection” than in the “protection” of the rarer birds. He realized that these matters were not connected with the Museum, but he appealed to Taverner to do all he could through the proper authorities to ensure that the Department itself should carry out the spirit of the “principles” it set out for others. He ended by saying that the warden at Vaseux Lake, S. J. Darcus, had recently been dismissed, though he was a man “who was known to put the “protection” and study of the living bird first and foremost.”** Taverner was now “put on the spot” and had to repeat what he had written to Harkin, but in much more detail, which must have taken up a fair amount of Taverner’s time. In his answer he took up Parham’s arguments point by point: Parham had exaggerated the effect that the scientific collector had on bird life in general; that ornithologists need to collect speci- mens; that the conservationist is ineffective without the special knowledge that the scientific ornithologist alone can give. He finished this six-page letter with the information that he heard from the Parks Branch why Mr. Darcus was laid off — that it was due to eco- nomic retrenchment. He ended by saying: “Tf the general public were half as interested in control- ling the unsportsmanlike hunter as they are the unscientif- ic or unethical collector most of the worries of the conser- vationist would be over.”®? Parham’s reply was sharp and swift. He repeated that the public was convinced that some of the 170 THE CANADIAN FIELD-NATURALIST Vol. 110 Number .697...............000000005. 17th. day of...Maroh..........-. 1931... DEPARTMENT OF THE INTERIOR NATIONAL PARKS OF CANADA PERMIT TO CAPTURE MIGRATORY BIRDS FOR BANDING PURPOSES (Subject to the Migratory Birds Convention Act and Regulations) Permission is hereby given to....Mr...P oho. TAVOPN eT igsciicsc- beccnotesccecestce ot dece straneneces ee pea Ornithologist, National Museum of Caneda, Ottawa,..Qntario.,. lider the authority of the Migratory Birds Convention Act to TAKE or POSSESS in the PROVINCE........ fo) etn MANO Bibieis bel actal cena specimens of each or any migratory game, migratory insectivorous or migratory non-game bird for banding purposes only. This permit is to be Sieg by. the permittee and shown on demand of any authorized game warden or police officers; : 7It dg ‘tevécable at ny time at the discretion of the Minister ‘of the Interior, and does not igi any tights td take, injure, molest, or kill birds on any ‘Dominion or Provincial game presérve lot: ‘bird sanctuary. This permit does not confer the right to take specimens of Whooping Cranes or Trumpeter Swans. This permit expires 3lst December, 1921.. Signature of Permittee For Minister oftthe Interior Under the authority of the laws of the Province........ Of MANITOBA utes I concur in the issue of this permit. Director of Game and Fisheries. Permit to capture migratory birds for banding purposes only in the Province of Manitoba to Mr. P. A. Taverner for the year 1931. Signed by Hoyes Lloyd on behalf of the Minister of the Interior, and countersigned by the Director of Game and Fisheries, Province of Manitoba. Note that this permit does not confer the right to take specimens of Whooping Cranes or Trumpeter Swans. employees of government were given, in addition to _ that these beautiful birds were never so plentiful again their salaries, “the right to collect all the rare and at Vaseux Lake. Taverner’s figures regarding area beautiful birds that they or their subordinates care to and bird population, he said, seemed to be mistaken kill, in order to set them up, and sell or trade them.” —_ for British Columbia. He gave BS) in example, two or three collectors whom “Your fears for the continuation of ornithologists are, I the Provincial government sent to the Okanagan think, groundless. I find a dozen enthusiasts today where Valley during nesting time to collect all the female there was one in the old days, and no doubt a few of the Lazuli Buntings they could. The result, he said, was younger generation will turn to the scientific side.” 1996 He said that “we” realized the need for such men as yourself and my old friend Allan Brooks to have new specimens from time to time, and we know how much you two have done through publications to make bird-study popular. “But the scientific “spade work’ has already been done, and the garden can be kept in order by a few experts.” He then went into the treatment of Darcus in more detail.?° Taverner was clearly upset by Parham’s second let- ter and said so, answering all of his arguments in detail. As regards Mr. Darcus, Taverner said that there may well have been some political influence involved since these minor posts were not under civil control. Local members often had wide powers in their appointment. “As long as the people of Canada will stand for political patronage and ‘to the victors belong the spoils system’ departmental authorities are helpless in such cases. No system of appointment is perfect and the Civil Service has its grave faults but at the worst it cannot be as bad as the political spoils system that is its only alternative. But if the people of Canada stand for it the results are their own fault.”?! On the final page Taverner tried to be conciliatory when he wrote: “You say that the spade-work has been done and there is little use for the expert any more. To the superficial observer perhaps it so appears so but to me who am studying the country as a whole and in detail it looks as if little more than the ground was scratched. What we do not know of the Birds of Canada is enormous, I am con- tinually running up against the unknown and having to dodge here and there to make up for ignorance. I do not hope that I have convinced you altogether but I expect that at least I have shown that there is something to be said on my side.””” Parham was not easily put off by answers from Taverner or Harkin. In March 1936 he sent a copy of a chapter of a book that he was writing under the title A Nature Lover in British Columbia to Harkin, Commissioner of the National Parks Bureau. Parham was told that next time a senior officer was going to British Columbia he would call to discuss the ques- tion with him. When, after a year, no one called, Parham went ahead and published the book with the chapter on saving the birds in it. Taverner’s argument that uncommon birds still needed to be collected was brushed aside, and a new menace to sea birds was included as follows: “The important facts concerning practically all our inland birds are now known; and although there is still much to be found out about bird-life, this does not call for the slaughter of any more uncommon birds. Observation, photography, and banding should now be the ruling methods. CRANMER-BYNG: A LIFE WITH BIRDS 171 Bird-lovers should also make every effort to get their Governments to deal with the constantly increasing men- ace of fuel oil upon coastal waters, for this is taking a ter- rible toll of life amongst our sea birds.””? This was a loud warning about the damage being caused to sea birds by the loss of fuel oil from tankers along Canadian coastal waters in a book published as far back as 1937. Too many oil tankers were handled irresponsibly then, and now, over 50 years later, still are. An unsigned review in the Ottawa Journal (1938) was obviously written by someone with knowledge of the controversial subject, and was in the nature of a public reply by the Parks Bureau to Parham. Two columns were devoted to explaining how bird protec- tion was carried out by the Dominion Government under the Migratory Birds Convention Act, and the regulations made under its authority. The chief migra- tory bird officer in British Columbia, the review said, was Mr. J. A. Munro who had held that office since 1920, was an ornithologist of international standing, author of many brochures and scientific studies and author of a popular handbook on birds which was dis- tributed to all the schools in British Columbia. The administration of the Migratory Birds Act came under the National Parks Bureau of the Lands, Forests and Mines Branch of the Dominion’s Department of Mines and Resources. Officials of that bureau, the review continued, had complete confidence in Mr. Munro, and were of the opinion that Mr. Parham’s criticism was unjustified. The reviewer explained the difficulties under which Munro worked. “Mr. Munro has no permanent staff. He has seasonal assistants at small salary, works in co-operation with the Mounted Police and the provincial game warden.” The remainder of the review discussed in some detail the issuing of scientific permits in relation to the law. These permits could only be granted “on the issue of a permit by the Minister or by any person duly authorized him him’. Permits were not issued by Mr. Munro or any field officer, but only by the Department in Ottawa. In the latest year for which figures were available, in all British Columbia only fifty such permits were issued, and only about 1500 specimens were reported taken by them, with each permit a printed code of principles, three sections of which were quoted verbatim. This part of the article contained useful information on the regulations cov- ering the granting of scientific permits at that time.” But it did not answer Parham’s complaint. His point was that the Department’s representatives were more interested in the collection, rather than the protection, of the rarer birds.”° li2 THE CANADIAN FIELD-NATURALIST Vol. 110 CHAPTER 16. The Widening Field of Studies II, 1936-1942 Taverner and some museums in Canada Taverner tried to keep in friendly contact with the curators and ornithologists at other museums across Canada. Francis Kermode, director of the Provincial Museum of British Columbia in Victoria, and Taverner kept up a spasmodic correspondence. But owing to the hostility of Brooks towards Kermode, Taverner had to step carefully.! In the prairie provinces there was little of impor- tance to involve Taverner in Edmonton or Calgary.’ In Saskatchewan, Taverner was regularly in touch with Hedley Mitchell, preparator and naturalist at the Provincial Museum of Natural History in Regina from 1913 onwards.* In Manitoba, a Natural History Society was formed in 1920, and the question of a permanent public museum was raised. A museum Committee was appointed, the members being B. W. Cartwright, A. G. Lawrence and Dr. H. M. Speechly.* With the sad death of Cyril Harrold in 1929 the question arose of how A. P. Harrold should best dispose of his brother’s personal collection of birds and eggs. Taverner wrote asking him what he intended to do. He replied he was planning to give the collection to Winnipeg.° Meanwhile Cartwright and Lawrence wrote an appreciation of Harrold and sent it to Taverner for publication in the Canadian Field- Naturalist. In reply Taverner said: “A. P. Harrold tells me that he is giving the collection to Winnipeg. I must confess I knocked the idea pretty hard.” This was not for any personal reason. The more reputable museums there were in the country, Taverner felt, the more cooperation they would receive, and the better they would sell the museum idea to the gener- al public which will be to their great advantage. He continued: “But a museum is a continuing and established organiza- tion more than it is a room in a building or even a set of exhibition or storage cases. We who have watched these things carefully have every justification to be pessimisti- cal over small local museums started with the enthusi- asm over some local collection. The country is strewn with their wrecks and what is worse, — the waste of specimens that might have been of good use.’”° Taverner felt that wildlife specimens, even in col- lections under University administration, had an uncertain future. Few university professors had any lasting interest in museums. He had recently seen a ghastly wreck of a rather ambitious museum at Queen’s University. Taverner would rather see Harrold’s specimens in Cartwright’s or Lawrence’s hands personally than suffer the almost certain neglect that would come as soon as Harrold’s imme- diate friends had died and the early enthusiasm was over. “Harrold himself felt just as I do about these things”, Taverner wrote, ‘“‘and had very little faith in these small amateur museums”. He then gave his frank opinion about the best way to proceed. “If Manitoba or Winnipeg is going to have a real Museum I greatly rejoice but the essence of a museum is a skilled staff and an assured future and to start off with- out them is more likely to postpone a real one than to be the nucleus of a future one.” In 1932 space for a museum was set aside in the newly-built Winnipeg Auditorium, and the Manitoba Museum Association was set up to operate the muse- um which was opened in December 1932. Dr. Speechly became President of the Association. Meanwhile Taverner was keeping a watch on the development of the museum through letters from Cartwright, until Cartwright joined Ducks Unlimited as Chief Naturalist in 1937. Subsequently Lawrence kept Taverner informed about events in Winnipeg. Writing to Speechly, Taverner said the greatest need of a young museum was for experienced curators. Every library of any pretention expected its librarian to take a course in library work, he wrote, but when it concerned a museum any one was considered com- petent to direct it. Yet running a museum was a more complicated matter, and consisted of much more than sticking specimens in cases and on shelves. Taverner made the following proposal. “Now it is proposed that we in the National Museum where we have a considerable and varied staff of experi- enced museum men, the National Art Gallery and per- haps the Archives unite in giving a short course in muse- um management, objectives and methods, for the benefit of the struggling museums of the Dominion. “he scheme is to induce the Carnegie Foundation to pay transporta- tion in whole or part of selected applicants. The course will be free. What do you think and would any of your people be interested?”® The Carnegie Foundation gave the museum two grants for cases and exhibits. In December 1939, Speechly sent Taverner a Christmas card and a note about the museum grow- ing slowly. In reply, Taverner said it took about thir- ty years for a new idea to germinate and come to fruition in the public mind. Speechly had mentioned about 2000 visitors yearly in attendance. Taverner commented that when the children of these 2000 had passed through the museum age, he could expect some reaction from the public. Taverner warned Speechly, and by extension the members of the Manitoba Museum Association and the Honorary Curators and Assistant Curators, not to lose heart. He continued: “The idea that cultural values are equally important with economic may then be in process of seeping through the general consciousness ... Getting adequate support of a museum 1s up-hill work until the value of it is grasped by the electorate of the legislators who open or close the purse-strings. Don’t lose heart for the next thirty years and don’t let the idea die or else it has to be done all over again.” In the spring of 1940 Taverner received a copy of a talk on museums, given by Speechly, and sent to 1996 Taverner from the National Gallery. After congratu- lating him Taverner turned to the state of the National Museum. “Our Museum also has been going steadily downward for many years, — almost since its inception and under present conditions there seems little hope to stay the continuing process. With all the investment in it from a purely financial standpoint it seems absurd that it should be allowed to lapse yet that finish does not seem too far away. All we can do is to hope for better things but I sadly fear that we Canadians are not museum minded and are more interested in material than in cultural things.”!° During the 1930s Taverner had given considerable thought to the role of museums in society. In 1934 he gave two radio talks on CKCO Ottawa. The first was “The study of birds as a cultural pastime” (12 January 1934); the second was “The what, how, and why of a National Museum” (6 April 1934). There appear to be no letters of comment on these talks preserved in Taverner’s official correspondence, though there may have been among his private papers which were not preserved.'! The largest ornithological collection in Canada after the national collection was that of the Royal Ontario Museum of Zoology (ROMZ) in Toronto. Taverner heard about the ROM both before and after its founding in 1912 from J. H. Fleming, but it seems that not until 1923 did a member of the ROMZ’s staff get in touch with him. Then Lester Snyder, at that time technologist in ornithology, wrote to him for advice on what birds to look for during summer field work in the Lake Nipigon area. This was a good opportunity for Taverner to develop a useful link with the ROMZ. Taverner replied that he should look out for Connecticut and Kirtland’s Warblers, Evening Grosbeaks and Sharp-tailed Grouse, among others.!” The two men first met at the AOU meeting in 1924. When Taverner realized that the ROMZ had started a regular series of expeditions to parts of Ontario he made some kind of verbal agreement that the ROMZ would undertake field work only in Ontario, and the National Museum would concen- trate on other parts of the Dominion.'? In 1926, Snyder wrote asking for advice on rear- ranging the ROMZ study collection, and how to start a registry. Taverner replied with a detailed descrip- tion of his system. In 1928 Snyder told Taverner that the Lake Nipigon faunal report was ready for publi- cation in the Proceedings of the Canadian Institute, and he hoped to have the Lake Abitibi report done in the same way. An interesting piece of information included in the same letter was that James L. Baillie was going to night school, and would be unable to collect at Long Point that summer. Instead, Snyder wanted Baillie to collect at Ashbridge’s Bay where “recent developments indicate that this will be our last chance.” In the same letter he reported that W.E. Saunders was in Toronto for a Brodie Club CRANMER-BYNG: A LIFE WITH BIRDS 173 meeting and presented the records of the Great Lakes’ Ornithological Club on behalf of its members to the ROM. Snyder added “Please accept our thanks for making our institution the depository of these valuable records.” In 1931 Taverner was working on the Great Horned Owl, and several letters passed between him and Snyder about identification of the species in the ROM collection. In one letter Taverner said that some might be intergrades between virginianus and subarcticus but could not be intergrades with occi- dentalis which belonged to the southcentral United States. He told Snyder that the National Museum had “an unusual lot of summer and nesting birds from many localities. If we had some of the gaps filled we could speak with much more authority. We greatly need sum- mer residents from Mackenzie, Keewatin, Ungava and the maritime provinces”. Snyder replied that Taverner’s letter contained “much of interest to him on the general idea of sub- specific variation.”!° In 1932 the ROM published a booklet on the hawks and owls of Ontario. and a copy was sent to Taverner who replied that he liked it, and added “Am particularly pleased with the little pen and inks of Shortt’s”.!° During 1933 Taverner examined the ROM specimens of gulls from Churchill, including “Kumlien’s” and “Thayer’s”, and wrote with his comments. The Sharp-tailed Grouse was the subject of correspondence in 1934-1935, especially on the discontinuity of range of the dark northern form.!7 In September 1939 Taverner wrote to Snyder about his trip to Berkeley and back, with Martha dri- ving, and asked the name of the Toronto man who lugged about a big telescope and tripod at the AOU meeting. Snyder replied that he was Murray Speirs, and he was studying for a Doctor’s degree at the University of Illinois.'* At the end of the year in a letter to Snyder he touched on the “splitting systema- tists” and the blind way they followed each other like sheep. As an example he pointed to the water thrushes. A series of 125 specimens amply demon- strated to him that there wasn’t such a thing as Grinnell’s Waterthrush (Seiurus noveboracensis notabilis).'? Looking back over the careers of Taverner and Snyder in the 1920s and 1930s one can understand how much they helped each other. Taverner needed an expanding museum, relatively near to Ottawa, from which he could borrow skins for his studies of the Canadian races of species such as Canada Goose; Red-tailed Hawk; Great Horned Owl; Rock Ptarmigan; and Savanna Sparrow, to mention only a few. The ROM was the ideal place. In return Snyder needed to be able to borrow skins for studying Canadian races of various species from the much larger collections in the National Museum. Snyder, especially in the 1920s, needed to be able to draw on Taverner’s wide knowledge of bird distribu- 174 tion in Canada, and wide acquaintance with Canadian ornithologists. Both men needed a corre- spondent with whom he could feel at ease and dis- cuss some of the problems of his professional work such as the administrative difficulties of museum work, the never-ending problem of correctly deter- mining subspecies, and at times just “letting off steam”. A useful symbiosis gradually developed between the two ornithologists. They had various things in common. Neither had a university degree though Snyder had majored in museum studies at the University of Iowa. Each man, towards the end of his career, suffered from a feeling of dissatisfaction with the museum in which he had spent his career.”° Another ornithologist at the ROMZ with whom Taverner had some correspondence was James L. (Jim) Baillie. Born in 1904 he was ten years younger than Snyder, joined the ROM in 1922, and began corresponding with Taverner on museum business in 1925. Writing to Taverner in 1926 he informed him that the ROMZ aimed to compile reliable breeding records of Ontario birds in order to establish the breeding range of each species in Ontario. He sent a rough plan of the proposed method.*! Taverner replied that he had about half the Canadian species mapped out, and a card index of about 40 000 specif- ic references. He suggested that Baillie should come to the National Museum in order to collect informa- tion from their records.” Early in 1940 friends of Fleming were becoming anxious about the state of his health. Taverner wrote to Snyder asking to be kept informed. He said: “Tam very sorry indeed to hear that Fleming is so low. I do not know what I would do without him. He has been my closest friend for so many years and ornithologically my chief mentor. But the situation looks alarming. Please keep me informed.” Taverner saw Fleming for the last time when pass- ing through Toronto in March. It could only be a brief visit because Fleming was very weak. He died on 27 June 1940. When Taverner heard the news from Snyder he replied, in part, “Fleming will be a great loss to us all, personally as well as ornithologi- cally.” After asking about the disposition of Fleming’s considerable library, he wrote: “Well, I guess that this marks the end of an era in ornithology as in other things in Canada.”* When Taverner was asked to write an obituary notice for Fleming in The Auk he wrote to Snyder explaining that he did not feel that he could do justice in writing a tribute to his old friend. Snyder agreed to write it.2> Taverner wrote a warm account of his early memories of Fleming for the Brodie Club’s “Fleming Night” entitled “The Yonge Street Taxidermy Shop and Point Pelee Days”.*° In a letter to Snyder he wrote that the Fleming memorial number was a very fitting tribute, but could not help showing his disap- pointment at the lack of appreciation Fleming had received from Toronto during his lifetime. THE CANADIAN FIELD-NATURALIST Vol. 110 “It was a great regret to me that he did not receive the recognition from the Royal Society that he deserved, and it was the Toronto backing that failed him. Others got in who had done much less for their science than he did ... the trouble was he did not spout enough science jargon 927 In a letter a week later he told Snyder that they also had noticed the snootiness of Toronto that he- had mentioned. Too bad Fleming was not University of Toronto though in that case he might not have been Fleming.”® A third member of the staff of the ROMZ whom Taverner came to know was Terence Michael Shortt, wildlife artist and display preparator. Taverner first heard of the brothers Angus and Terry Shortt from Cartwright in Winnipeg. Cartwright wrote about their artistic promise and sent Taverner a selection of their pictures displayed by the Winnipeg Natural History Society in 1928. Angus was then 21 years old, and Terence 17.7? Taverner replied with a four- page letter of encouragement blended with critical talk on the basics of painting birds. This letter shows that Taverner had a fair grasp of the fundamentals. He spoke of the need for a good knowledge of the feather tracts and groupings. He advised students to do systematic drawing from nature “to place down just what they see, and to learn to see what is before them”. Everything should be to the point, and the result of his own experience.*° Cartwright replied that this was just the kind of advice they wanted. “They have been working on heads and wings, eyes etc. lately and are now giving feather tracts and pattern their attention. I have lent them Dwight’s “Plumages and Moults of the Passerine Birds of New York” and they are of course getting in field experience at every opportunity. They have access to both Lawrence’s and my own specimens so we are doing all we can with the meagre material at our dispos- al to help them along.”*! The care that Taverner took in encouraging them was repaid by the rapid improvement in their work. One other museum in Canada, with a fair collec- tion of birds was the Redpath Museum in the grounds of McGill University, Montreal. The Redpath Library, a separate part of McGill University, has a superb collection of ornithological literature. Taverner first heard about the museum through Dr. Casey Wood, who asked his advice about the Emma Shearer Wood Library of Ornithology in correspondence which started in 1917 and continued irregularly into 1934. Wood was away from Montreal much of that time.*” Taverner had been in touch with the ornithologist Henry Mousley, of Hatley, in the Eastern Townships of Quebec, since 1914, and came to admire his studies of the breeding bird behaviour, and the home life of birds. In 1924 the Mousley family moved to Montreal, and in 1927 he was employed on library work in the Emma Shearer Wood Library where he remained for the next decade.** This gave Taverner a 1996 useful link with the Redpath Museum, and an incen- tive to visit Mousley there when he could manage it. Taverner encouraged Mousley to publish his research in The Canadian Field-Naturalist, and they continued corresponding through the 1930s.*4 Taverner’s Involvement with the American Ornithologists’ Union in the 1930s. During the AOU meeting at Charleston in 1928 Taverner was elected to the Council. In a letter to Snyder he said: “The Charleston trip was delightful in every way ... think the A.O.U. treat us Canadians very well indeed. I certainly appreciate the honor they paid me.’’*> Another of Taverner’s links with the AOU was his part in the very early development of bird banding in North America. In 1931 he sent Frederick C. Lincoln of the Biological Survey, Department of Agriculture, a list of the band num- bers that he had distributed in 1905, 1906 and 1908 with the names and addresses of the recipients. He did not remember why no bands were issued in 1907, but guessed that it was for lack of funds “as I was living pretty close to the edge in those days and had to get the bands laboriously cut and numbered by hand”. His record showed that some 429 bands were used, he said, in this early attempt. The system was essentially the same as that followed in 1931. Bands were issued by a central authority in single series, and records kept centrally. “As such I feel that if I am not the father of bird-banding [in North America], I am at least its grand-father’.*° Lincoln was glad to receive this information because such matters of historical interest should be properly recorded. He told Taverner that since the Biological Survey took over the work in 1920 well over 700 000 bands had been issued. With such a large number of birds wearing bands the Survey was receiving great quantities of returned information.*’ When the fourth edition of the AOU Check-list was published in 1931, it received plenty of criti- cism. Taverner’s friend, Frank Farley, criticized it on several counts, the main one being that the check-list committee did not make use of The Canadian Field- Naturalist for information on distributions. Taverner defended the action of the Committee. The making of a check-list of this size, he wrote, was a tremen- dous piece of work. To check all ornithological liter- ature for distributions within a reasonable time would be impossible. The Committee had to draw the line somewhere, and they drew it at ornithologi- cal magazines. The Canadian Field-Naturalist was a general magazine so was excluded. On the other hand the galley proofs of the list were rather widely distributed, and suggestions from a large number of recipients were requested. Fleming, Brooks and Taverner “all had a whack at them”, as Taverner put it. Many of their suggestions were adopted, in other cases they took someone else’s opinion. The Biological Survey, he said, had an enormous amount CRANMER-BYNG: A LIFE WITH BIRDS 175 of distributional data from all over America. The trouble was that they did not use enough discretion in how they employed it. “There is a bit too much Oberholser in that department” Taverner said, “and he is most credulous. A good many errors crept in that way.” As to why there was no Canadian on the Committee, “there is no Canadian competent. We have good field men, good distributionists but no real systematists”. After much more explanation along these lines, Taverner asked: “How can a Canadian bring any important assistance to a Check-list of the American section of the birds of the world when all he knows is Canadian birds and has no opportunity to learn North American birds much less those of the rest of the world.”*® An important event in Canadian ornithology in 1932 was the holding of the AOU meeting at Quebec City. This was the first and only time in the history of the AOU that a bilingual program was presented. James Chapin of the American Museum of Natural History was fluent in French, and therefore played an important part in the program, especially in the French sessions during one afternoon. The business meeting was held at the Chateau Frontenac, and the public sessions were held in Laval University. The program was printed in English and French. During the business meeting Hoyes Lloyd was elected a Fellow. The election of officers resulted as follows: J. H. Fleming, President; Dr. Herbert Friedmann, Vice President; Dr. A. A. Allen to fill a vacancy in the Council. Other officers from 1931 were re-elect- ed for subsequent years. This included Taverner as a member of Council. The program included fifty-five papers covering a wide range of subjects. Palmer, in his report in The Auk, wrote that the outstanding contribution was “Sounds as an Aid in Bird Study” by Albert Brand, illustrated with moving pictures and slides, and reproduction of bird sounds on the phonograph. Improvement in outdoor recording, the method of editing film and the process of making bird sound records were briefly explained.*? Although, due to his stammer, Taverner did not play any part in public speaking or in giving a paper, he exhibited some of his recent photography. Above all, being a sociable man with a good sense of humour, he was in the thick of the conversation with old friends and new acquaintances. One event that would have given him considerable pleasure was the evening when mem- bers of the Union were entertained by Dr. and Mrs. Gustave Langelier at their home at Cap Rouge, on the north bank of the St. Lawrence close to Quebec. Palmer wrote “.. several hours were spent in looking over the Langelier collection of birds which is rich in specimens illustrating the various plumages of game birds and other local species and also contains a number of rare speci- mens, such as the Eskimo Curlew, Spoon-bill Sandpiper, Passenger Pigeon, and Carolina Paroquet.’"? 176 On 20 October members left by motor bus for Cap Tourmente about forty miles down the St. Lawrence, to see the Greater Snow Geese which stop for sever- al weeks each autumn en route from their nesting grounds in the northeastern arctic Islands to the Atlantic coast from Delaware Bay to Chesapeake Bay. This was a great sight, a huge flock of about 9000 birds, feeding at close range, and then to see them rise, circle around, and slowly fly up the river to another feeding ground.*! Taverner writing to Bishop pronounced the Quebec meeting a success, saying that Quebec was still largely “feudal” and could do things no other section could do. As regards the elections he said: “Oberholser was decisively beaten. I do not think that it will ever be possible for him to be elected president of the A.O.U. He has every scientific right to expect it and it is logical and just that he should have it but he has made too many enemies especially in his immediate cir- cle where he should have his best friends.”*” In 1933 the AOU Stated meeting was held in New York and celebrated the completion of the 50th year of the history of the Union. Taverner, in a letter to Brooks, wrote a fair amount on the AOU and the New York meeting. Here are two examples of what he said: “The people at the A.O.U. are a mixed lot, there is a good deal of the lunatic fringe. As men get older the more “sot” they become in their ways and the more intolerant they are. Quite a number show this effect in the A.O.U. Are we getting afflicted the same way? ... Of course New York is not a happy atmosphere for the A.O.U. The brass-hat attitude of the American Museum is not conductive to mutual understanding and IJ think was particularly evident this meeting. Also I feel that the comraderie [camaraderie] that used to be the key note of the A.O.U. is not as strong as it used to be. Is this because the Old Guard is dying out and newer more effi- ciently cold blooded men are more in evidence?” The third time that the AOU met in Canada was in Toronto, from 21-24 October 1935. The headquar- ters was at the Royal York Hotel. The public ses- sions were held in the theatre and lecture room of the Royal Ontario Museum. At the business session A. C. Bent was elected President for 1936. The Secretary reported a membership of approximately 2000. The program consisted of 67 papers. On the first morning L. L. Snyder spoke on “Ontario and its Avifauna”’, and J. L. Baillie on “The Bird Collection in the R.O.M.Z.” Later in the program Fleming spoke on “The standing of Cory’s Least Bittern, with an exhibition of specimens”, and Taverner on “An outline of the Buteo borealis complex, with an exhi- bition of specimens”’.** Members of the local committee decided to pre- sent some “awards” at the banquet to outstanding ornithologists. This is how Terry Shortt described it many years later. “... IT remember that George Miksch Sutton received a huge whitewash brush. Hoyes Lloyd was given a toupé THE CANADIAN FIELD-NATURALIST Vol. 110 of the scalp of a Great Horned Owl complete with erect “horns” to cover his shiny bald head, (his comment: “I feel like the devil”). For Percy Taverner, author of The Birds of Eastern Canada, The Birds of Western Canada and The Birds of Canada, it was considered appropriate to complete the tour de force. Lester Snyder and I took a copy of The Birds of Canada to the technicians in the Museum of Geology laboratory and had them cut the book in two with their sophisticated equipment. It was cut horizontally. I made dust jackets for each of the halves entitling them “The Birds of Northern Canada and The Birds of Southern Canada. Taverner’s comment on receiving these at the dinner ran something like this — a number of reviewers have torn my book to pieces but this is the neatest job that’s been done to date.’ The edition of The Auklet for 1935 contained a spoof review of Taverner’s Birds of Canada which read as follows: “To begin with, let us preface our brief comments on this work by stating that this hurts us more than it will Mr. Taverner. The author’s book is nothing more or less than a fragrant case of plagiarism. We recall two publi- cations, one on the ‘Birds of Eastern Canada’ and the other on the ‘Birds of Western Canada’, the authors of which we do not remember for the moment, both of which have been quoted verbatim! Mr. Taverner has apparently attempted to shield this fact by shuffling the systematic arrangement of the groups and species and by inserting a few distracting illustrations. The one feature of the book which appears to be original is that it can be cut in equal halves horizontally and one then possesses the ‘Birds of Northern Canada’ and the ‘Birds of Southern Canada’.*° One person Taverner saw at the Toronto meeting was Bent, who was elected president for 1936. When he had settled into the routine of being president, Taverner wrote him a carefully thought out letter on how he saw the state of the AOU in 1935, and some suggestions as regards the future. Taverner had now been a member of Council for seven years and had every right, and perhaps duty, to offer the new presi- dent advice. The Union, he said, seemed on the point of making some radical changes, but how far they should go required some thought. They ought to keep in mind that the AOU was a scientific and not an emotional organization, nor a bird protection society. Taverner said he was pro-protection in every way but the AOU was not the proper medium for it. Also he considered the election of the position of fellow should be based entirely on scientific acquirements and not on protectionist activity. Next he gave his views on reforming the financial structure of the Union which, he said, was almost entirely in the hands of the Secretary and Treasurer and perhaps one or two other officers. “The rest of us Councillors are just dummy directors, and rubber stamp men.” He con- cluded with a touch of humour when he wrote: “All of which is respectfully submitted which is the usual form of Civil Servants addressing their superiors”.*7 Bent replied that the AOU was primarily orga- nized for the scientific study of ornithology but had 1996 now grown beyond the interests of bird collecting and systematic ornithology into several other fields, such as bird photography, birdbanding, life history studies, etc. The vast majority of the Associate Members belonged to one of those classes, or were merely field-glass enthusiasts or bird protectionists, and the Union was largely dependent on their dues for support. Bent then stated his own position: “T am a dyed-in-the-wool bird collector and I have no patience with the growing tendency to refuse collecting permits, which is forcing us into ‘bootlegging’; but what can we do about it? We are hopelessly outnumbered by the agressive protectionists and they are forcing us to the wall. The day of the private collector is about over.”*8 Late in 1936, when Taverner realized that Bent was shortly to reach his seventieth birthday, he wrote him a warm letter of congratulation at achieving so much for North American ornithology in his “Life Histories”, and signed himself “Sincerely — your friend and Appletonian”’. In response Bent said that he had invited a few local friends to help him celebrate but he wished that all the Appletonians lived nearer.*” At this point the Appleton Club deserves mention- ing. As a “club” it had no official membership and its activities varied from year to year. Ad hoc arrangements were made to meet in a separate room at times during an AOU meeting. Here they talked about birds, people and projects in a relaxed way, smoking and drinking with banter mixed with chat about AOU members not present. A word that appeared in Taverner’s vocabulary in relation to AOU meetings was “camaraderie” in the sense of friendly good fellowship among equals, and seems to have been applied especially to Appletonians. Perhaps the main quality required was to be a like- able person, someone who got on well with others. A man such as Clyde Todd did not appear to have pos- sessed this quality, perhaps because he was consid- ered to be “unsociable”. In short the Appleton Club was a useful network of men who got on particularly well together, and served a valuable function because they brought up matters which could best be discussed by a small, informal inner group in a non- competitive and non-hierarchical way.°° At the AOU meeting held at Charleston in 1937 Herbert Friedmann was elected president. After the meeting Percy and Martha drove down the east coast to Florida, the Keys, Cape Sable and the Everglades with two days at Lake Okechobee. From there they drove to Tarpon Spring, then inland through the mountains of Carolina, the Shenandoah Valley, New York State, and then via Detroit and Toronto to Ottawa. It was an exciting experience and Taverner returned home refreshed. During the months before the next meeting Friedmann made plans to put in motion “constructive leadership” in the AOU. Between December 1937 and September 1938 a series of letters passed between Friedmann and Taverner on the role and organization of the AOU. CRANMER-BYNG: A LIFE WITH BIRDS aT Friedmann had a note in the spring 1938 issue of The Auk on new ideas on systematics and the Check- list. Taverner wrote to Friedmann on this subject, saying that the AOU should be directive as well as receptive, and that there were various lines of inves- tigation that needed encouragement. For instance he suggested that certain issues of The Auk should be devoted largely to symposia on specific subjects. At the Washington meeting in 1938 Taverner asked why not have one session devoted to a particular subject that needed discussion. Why not invite papers on some controversial subject and publish abstracts of them in The Auk, in order to suggest lines of attack? Friedmann had already asked Taverner to head a subcommittee on North American faunistics. To this Taverner replied that he would be glad to serve on such a committee but doubted his ability as an administrator or organizer. Besides he was familiar only with the situation in the northern part of the continent. The Fifty-Sixth Stated Meeting of the AOU was held at the U.S. National Museum, Washington D.C., October 17-22, 1938. During the business ses- sion three members of the Council retired after “long and valuable service”: H.C. Oberholser, T. S. Roberts and Taverner. “Dr. Friedmann, Dr. L. Cole, Dr. Ernst Mayr and Mr. Taverner reported for the divisions of the Research Committee”. Among the many interesting subjects of research was a sympo- sium on the topic “The problem of the individual versus the species in bird studies”. Among those speaking were: Margaret Nice, F. Lincoln and N. Tinberg. Friedmann introduced each subject.>! By now ornithology was in a state of change. Signs of this change could be seen in 1934 in Charles Kendeigh’s paper “The role of the environ- ment in the life of birds”? In a brief review in The Auk Witmer Stone picked out the salient points. The author considered especially temperature, relative humidity, solar radiation, food, precipitation and wind, and to a less degree biotic competition and physiographic features mostly in relation to the dis- tribution, migration and abundance of birds. The physiology of the temperature of birds and their resistance to low and high air temperatures was dis- cussed, based on experiments on House Sparrows. The paper also contained a special discussion of the distribution of the House Wren (eastern form) and the reasons for limitation on its northward distribu- tion during the breeding season. In a final summary Stone reviewed the physiological processes in birds and the behaviour responses, and the factors control- ling distribution, migration, abundance, and the role of animals in ecological communities.** Taverner read books and journals regularly, and corresponded with others, exchanging ideas and expressing his opinions. He never allowed his mind to vegetate. Here is Taverner late in 1936 in full spate in a letter to Bishop. 178 “Of course one trouble with the ornithology of today is that it is in a state of change. The obvious work of the old naturalists was purely systematic, discovering and naming new forms of life. A stock taking of zoological evidence. That has largely been accomplished. The evo- lutionary juice has been fairly well squeezed out of it now, and there are ... many more would-be naturalists in the field all seeking new worlds to conquer and not knowing quite what to do. They are essaying all kinds of subjects, mostly up blind alleys. They are run through the university mill that turns out biologists not natural- ists. They are physiologists, psychologists, pathologists, geneticists and microscopists, interested in what the ani- mal is (mostly inside) but not in its relation to the out- side world. The true relation between laboratory and field has not yet been worked out.” Taverner then went on to discuss systematics in relation to ornithology. The universities, he main- tained, were wrong in thinking that systematics was a finished subject and that further work in it was unnecessary. For all ornithologists to confine them- selves to the subject would be a great thrashing of old straw — there must still be a number of skilled sys- tematists. Systematics needs to be kept up to date in zoology just as much as dictionaries are in literature. “Just now all these young graduates are running around in circles chasing their tails in the endeavor to find something to do. The College professors don’t know anything more than laboratory work with microscope and conditioned reflexes. When they direct their students to the field all they can do is to write theses on how many times a little bird wags its tail.”>* In a later letter to Bishop he said: “Ornithological methods are changing. Instead of equip- ment with field glasses and gun the present ornithologist peers through a microscope and draws graphs with long tables of how many times a little bird wags its tail. We are already back numbers.”>° Taverner had been aware for several years that any ornithological research his contemporaries were engaged in was becoming outdated. This tended to affect members of the AOU — the old guard and the young gang. In a letter to Brooks of 1938 he com- mented: “Ornithology has greatly changed since our youth, — many new fields are opened and some of the old ones neglected. That I am not interested in some of these fields does not prevent my recognizing some of their values but it is a pity that the old collecting naturalist is being so rapidly eliminated. We need not all be collec- tors but that phase should not perish from the earth.” *© Friedmann, in his capacity as president of the AOU, wrote a piece on “The role of the A.O.U. in ornithology today” which was printed in The Auk in 1938. He explained that in its early days the AOU stimulated the work of bird protection and the popu- larization of bird study which gave rise to two orga- nizations which long ago outstripped their parent — the Audubon Society and the Biological Survey. He stated that now the field of ornithological science was the AOU’s remaining role, aside from publish- ing a quarterly journal, and various editions of the THE CANADIAN FIELD-NATURALIST Vol. 110 Check-list. Friedman ended with a call for the regen- eration of the AOU.°’ The effects of these changes in ornithology were very much on Taverner’s mind at this period. Writing to Brooks he expressed it with acumen. “The A.O.U. is not what it used to be and I quite under- stand why you and Bishop and others do not care to attend regularly. All the fine old circle is absent now. There may be others just as fine but they are of a younger strain and not as compatible. Also many of them are fol- lowing strange gods and their ornithology is not what ours used to be. Since good old Kennard went the meet- ings are not what they were. We are old-timers.”>* Writing to Rowan in 19339, in reply to a letter from him in which Rowan said that he hardly had any- thing more to do with birds “nowadays”, Taverner said, among other things, “Too bad you are losing your bird field work but I sup- pose that your university work does take up all your time. Too bad that none of our universities have such general natural history courses as at Berkeley and some other American universities. University Biology of the conventional type seems eminently adapted to smother all liking for living nature that cannot be obtained in a laboratory or viewed through a microscope. But it is too bad that Grinnell died. His death was a cloud on the A.O.U. meeting and I do not know how he will be replaced. He inspired many.”°? By 1940 it was not just ornithology that had changed, it was the ornithologists as well, death was removing more and more of Taverner’s contempo- raries. Fleming’s turn to go came in June 1940. Taverner, writing to Bishop with the news, sounded quite elegiac about it: “One by one our contemporaries drop away leaving us alone amid another generation with which we have less in common. We become, not the Old Guard but the Rear Guard and perhaps camp followers. New ideals, new gods reign. Ornithology has changed since we arose, probably the development is good, but much of the con- tents of our journals are quite foreign to us and the lead- ers that are coming on know not Paul. It is inevitable.” All that Taverner could do before someone was appointed to take over his post at the museum was to finish the research that he had been working on for the past few years, and his two pocket manuals. At last, in 1936, Taverner was able to leave Ottawa early enough to carry out worthwhile field work. His first choice would have been to make an expedition north of Hudson Bay into the eastern Arctic. But for reasons of expense it was impossible. Instead he decided to make an ornithological survey along the recently completed stretch of the Hudson Bay railway from The Pas, Manitoba, to Churchill. He explained his reason for doing this in the “Annual Report of the National Museum” for 1936. The Ornithology of the Churchill area, on Hudson Bay, at the edge of the Arctic Faunal Zone, he wrote, had been well studied in 1930 by a museum party and by others since. In southern Manitoba, Taverner 1996 CRANMER-BYNG: A LIFE WITH BIRDS 179 W. E. Saunders (right) relating some experience to his old friends Taverner (left) and Fleming (Centre) at the American Ornithologists’ Union annual stated meeting at Charleston, South Carolina, November 1937. Photo by A. F. Ganier (also appears on page 18 of W. E.. Sanders Naturalist. Edited by J. R. Ruttes. 1949. Federation of Ontario Naturalists, University of Toronto Press). Reproduced courtesy of the Royal Ontario Museum Library and Archives; number 001225. 180 explained, the ornithological fauna, to the point where the Upper Austral zone influences begin to intrude on the Transition Life zone, was well known, but in the intervening territory little authentic infor- mation was available. He continued: “The object of the 1936 work was to begin a direct line of consecutive observations between these two strongly contrasted faunas. The reconnaissance was carried south to Cormorant Lake within the drainage system and the water maze of the lower Saskatchewan River”.*! Taverner was accompanied by R. W. Smith, of Wolfville, Nova Scotia, and T. E. Randall, from Edmonton. They went to Churchill by rail and started their field work from there. The following account has been compiled from Taverner’s field notes, reports and letters, and shows the names of the places they visited. Churchill, mile 510 north of the Pas, head of the railroad and port at the mouth of the Churchill River on Hudson Bay. The next camp was at Herchmer, mile 412, at the crossing of the Owl River, 21 June — 2 July. While here Randall found 23 nests of the Harris’ Sparrow which was breeding from Churchill south to Herschmer, but scarcely at all to the Limestone River farther south.® The subsequent camps were at Bird, mile 350, on the confluence of the Limestone and Nelson rivers from 3-17 July; at Ilford, mile 286, in the neighbourhood of Splitlake, 18-30 July; at Thicket Portage, mile 185, 1-31 August; and at Cormorant Lake, mile 42, 1-14 September. Although they travelled from one camp site to another by rail they did not remain by the rail- road all the time, but explored various habitats far- ther afield. The beginning and ending of the field work happened at the Pas, mile zero, on the lower Saskatchewan River, 18-19 June, and 15 September. Although they did not collect a large number of specimens, Taverner felt that the results were dis- tinctly worth while. For example, information on the comparative limitations of range of the thrushes was satisfactory. The Gray-cheeked Thrush was the only thrush observed at Churchill. Its range extended south to Herchmer. Here, Hermit Thrush were few, but at the Limestone river it was the only thrush. The Olive-backed [Swainson’s] Thrush was not noted north of Thicket Portage. The northern limit of Wilson’s Thrush [Veery] did not seem to extend north of The Pas. As regards White-crowned Sparrows, both the eastern race (Zonotrichia I. leu- cophrys), and Gambel’s race (Z. 1. gambeli) were “inextricably mixed south to the Limestone”.® In the same letter to Brooks Taverner commented further on distribution: “But I never saw birds so scarce and shy in my life as in this north. [American] Robins had to be stalked like hawks and then we were not able to get them at Herchmer. It took several days to get Song Sparrows that nested close to camp. Terns at Herchmer all seemed to be Arctics, at the Limestone all Common. The THE CANADIAN FIELD-NATURALIST Vol. 110 Solitary Sandpipers were interesting. The form at Churchill were definitely [Tringa solitaria] cinnamo- meus [cinnamomea]. Only saw one and could not take it at Herchmer did not see it at the Limestone but [7-s.] solitarius [solitaria| was common at Ilford. The boys worked very well. Randall is an inspired nest finder and collected a great number of eggs, Harris’s Sparrow, L. [Lesser] Yellowlegs, White-crowns [sparrows] and a lot of other generally considered desiderata. His skin making was not of the best but he gained experience as he went.” Taverner continued that he was disappointed they did not get any downy chicks. He particularly regret- ted not getting downy Solitary Sandpipers. They nested commonly in the muskeg at Ilford, he said, but they just did not have the luck to find any good downies of them. Nor was it a good part of Manitoba in which to be working, as Taverner explained: “We ran into insufferably hot weather for about two weeks, 98° in the shade which with mosquitoes, black and deer flies and bull-dogs [Moose flies?] made things difficult. We were fortunate in one respect however, a forest fire came straight down at us from 135 miles back and then stopped within 500 yards of our camp at Ilford.””°° Taverner left Randall and Smith at Ilford to con- tinue the work until mid-September. In the summer of 1937 Taverner engaged Angus Shortt and W. Watkins, both of Winnipeg, to contin- ue the ornithological cross-section of Manitoba from The Pas south as far as Duck Mountain. Taverner accompanied them for a month. They started at the beginning of June at Reader Lake in a sprucewood section a little north of The Pas; to Halcrow Lake south of the Pas 9-21 June; moved into deciduous and agricultural habitat near the town of Swan River 22 June-21 July; camp 4 was at Garland east of Duck Mountain 21 July-5 August; the last camp was at Singush Lake in Duck Mountain 5-27 August.°’ The main object of this field expedition was to establish the normal northern limits of range of various transi- tion species and the southern limits of northern ones. Taverner did not type an account of the 1937 field expedition as he did for the previous year and, from his scanty ornithological information and analysis, it seems that the birds, from The Pas southwards, did not impress him. The orchids, however, seem to have been numerous, some of them being rather rare. Angus Shortt, looking back fifty years to his expe- riences with Taverner, kindly sent me a resume from his field notes. In his covering letter to me he said “I remember Taverner fondly, as a knowledgeable and entertaining companion humorous, considerate and an excellent cook.”®* Two anecdotes of Taverner as a camp cook follow. On June 3rd Shortt and Watkins got up at 6 a.m. to find it very cold. Luckily for them Taverner already had the stove going full blast in the cook tent, and was preparing bacon and eggs, when there was trouble. This is how Shortt described it. 1996 CRANMER-ByYNG: A LIFE WITH BIRDS 181 Taverner photographing a Semipalmated Plover at Bird, Manitoba, on the newly constructed Hudson Bay Railway in 1936 (Reproduced courtesy of the Canadian Museum of Nature, number 81343.) “Then we had uninvited company — tent caterpillars! Attracted to the heat of the chimney pipe, they were crawling down it in scores. Of course, when they reached a hot spot, they let go and came dropping down onto the stove and into the fry pan, sizzling and popping like pop corn. Taverner was unperturbed, flip- ping them out of the pan with the egg ladle and kept on with his cooking!” Later in June they set up camp by Roaring River, where the air was cooler and drier, and free from insect pests. While Shortt and Watkins were explor- ing a poplar-willow habitat, and its wealth of butter- flies, ,Taverner remained in camp. When Shortt returned this was what he found: “Returned to camp at 12:30 and was treated to a Taverner surprise — roast woodchuck! He had shot the animal and prepared and cooked it during our absence. | was pleasantly surprised at how good it tasted, tender and rather like pork.”©? Shortt closed his note to me saying that after Taverner had left for Ottawa — “We missed him, his humour and his cooking.””° Family life of the Taverners, 1936-1942 In chapter 15 a section was included on the Taverners’ home life, based mostly on correspondence between Percy and his friends about their garden, and their enjoyment of music. Another section about their home life is included in the present chapter, drawn mainly from material from several of their younger rel- atives and friends, relating to the period 1936 until Percy’s retirement in 1942. Other material for this sec- tion has been taken from correspondence between Percy and friends, or related to me in personal inter- views, sometimes supported by tape recordings. Much of this comes under the title “oral history”, and in quantity amounts to more than can be used here.” This section begins with a paragraph by Taverner in a letter to Bishop about a week’s holiday that Percy and Martha had spent in October 1936, and continues in the same letter with information on the development of their rock garden at 45 Leonard Avenue. Taverner wrote: “The two of us spent a week up in Muskoka where I summered for so many years when I was a boy. I dont know any summering place that is more attractive than Muskoka but the great influx of enormous wealth in my old section has spoiled it ... Our old home there now replaced by a hundred thousand dollar establishment. The beautiful natural rock shorelines built over with masonry retaining walls to make beds for masses of petunias and marigolds and everything out of keeping with the atmosphere of the place ...””” “At home have been making another rock garden, hunting up good stone and bringing it in our car load by load. Have just finished it ready for planting and am beginning to wonder where or how long it will take to get enough alpines to plant it.”7* 182 One of the Taverners’ musical friends, Paul Larose, wrote about Percy’s Christmas roses, that each year he would dig through the snow carefully, where he knew that he had planted some, and bring out the most beautiful flowers for Christmas. “Percy loved his garden as he loved all things pertaining to nature.”””4 From 1937 until 1943 Taverner was busy obtain- ing choice wild plants for his garden from friends in Ottawa, at the Experimental Farm, and at the muse- um, and by exchange from his friends and acquain- tances during their travels in various places.’ Writing to Bishop about plants Taverner said “we gardeners” when he mentioned five Lady Slipper species in his garden, all doing well.’° One particular plant he wanted was Lewisia. He wrote to Brooks: “wish I had some of those Lewisia that soon will be speckling your hillsides ...””’? When Brooks did not send any Taverner wrote to Munro about exchanging wild flowers, and managed to obtain some Lewisia from him while sending Munro hepatica, bloodroot and jack-in-the-pulpit.’”* Also during the summer of 1937 Taverner was busy taking a lot of photos of wild flowers.’? By the late 1930s Taverner’s rock garden was becoming quite well known among wild flower enthusiasts and people came from distant cities to see it.°° In a letter to Laing in May 1941 Taverner noted that the work Laing was doing at his home in Comox sounded too strenuous for him. In the past, Taverner said, he was able to do such hard work. Now, when he looked at the digging he did and the rock retain- ing walls he once erected he discovered that he had to hire a man to take down his double windows and put up the screens. “But I find laziness has its com- pensations and leisure is pleasanter than lame backs.”*! Taverner’s health was “winding down”, though he preferred not to admit it, and instead blamed it on laziness and leisure, which it certainly was not. Apparently, spring in 1941 was two weeks ahead of normal time and the flowers came through “better than ever’. As regards the alpine plants, he wrote: “My rock-gardening friend who raises all sorts of alpines that no one else can grow in little pots has had to move to a location where he has no ground for even his condensed gardening and I have fallen heir to a great lot of stuff and my rockery is chock full of rarities that I hope I can hold. As an example to the difficulties of these slow growing and exacting alpine plants, I planted some seed of tulip species in 1933. They are just show- ing their first blossom. And some of these high mountain and arctic seeds don’t germinate for two or three years.”82 If Percy was the gardener, Martha was the musi- cian. Because Martha started teaching the piano after arriving in Ottawa there were a number of people with vivid memories of being taught by Martha in the 1930s and early 1940s, who helped me during the 1980s and early 1990 when I was writing this THE CANADIAN FIELD-NATURALIST Vol. 110 book. Stuart Jenness started lessons about 1933, when he was eight years old, and continued until 1944, coming twice a week for an hour after school, and also to practice on Saturday afternoons. This usually coincided with Percy Taverner being at home. With practice completed to Martha’s satisfac- tion, young Stuart might be allowed to go down to the basement to watch Percy in his workshop as he created various articles from copper. But piano prac- tice had to be completed properly first. If he was cheeky Martha might “take a swipe at his head” — in fun — but she would not tolerate any slackness — Martha had been a professional piano teacher at a private school and knew how to keep discipline.** Another piano student was Barbara Lowe (Reynolds) who became a good friend of Martha. Her future husband, Walter Reynolds, was suddenly called overseas at the end of 1939. She did not see him again until the end of the war. As a result Barbara spent much time at the Taverners’ in the war years.*4 There were a number of other piano students who became fond of Martha, and had fond memories of “Mrs. T.”, as they called her. Some remembered the sessions on musical appreciation which she gave in the living room using her piano and the phonograph (record player). About ten students, mostly of high school age, would sit around the living room while Mrs. T. explained some music to them before play- ing it on the phonograph, which was placed in the sun room next door, with the glass doors fully opened.®° It was a great occasion when a visiting symphony orchestra played in Ottawa, and the Taverners and their musical friends had the chance to hear a live orchestra and discuss its performance afterwards. One such event was in December 1938 which Percy related to his old friend Mack Laing. Radio and phonograph, he wrote, are worthwhile but when you hear a live concert you know the difference, and he explained what it meant to them in Ottawa. “We are in the semi sticks and only have such an opportunity once in about two years. Ottawa is quite snooty about its superior culture, yet the majority of our friends have never heard a real orchestra before.’*° The other kind of live music that they could enjoy was when they made their own. As Martha became known in Ottawa musical circles she gathered together several musicians, and they would hold evenings of music and conversation followed by delicious plates of food. Sometimes there were visiting musicians invit- ed as guest performers, but usually the music was performed by a trio or quartet of talented friends.*’ One of these was a good amateur cellist, Paul Larose, who spent many musical evenings at the Taverners’. About Percy Taverner he wrote: “ ... although not a musician himself he enjoyed the numerous musical evenings which were held at his home. We had much chamber music with Mrs. Taverner, 1996 and many were the visitors who profited from these musical evenings. It was on such occasions that I met people such as Dr. [Diamond] Jenness, Dr. [Erling] Porsild and other friends of the Taverners’.” ** During Percy’s and Martha’s married life their house at 45 Leonard Avenue saw much entertaining. When they were first married they employed a Scottish housekeeper who wore formal white uni- form, and did the bulk of the cooking, a housemaid to do the cleaning, a yard man to cut the grass and do the heavy work of the garden. Percy did the weeding of the rock garden and gave instructions about gar- den work. In his later years, when his heart was a lit- tle groggy, Martha would tell him when he had done enough weeding. The Taverners entertained once a week on an average, and had house guests quite often. Karel Wiest lived at 45 Leonard Avenue during vacations from Cornell University starting in 1930, and had a German shepherd puppy named Sinbad. Later, when Karel was living away from home Sinbad, or “Sinners” as he was usually called, became a regular member of the family. Because Percy had designed his own home it was reasonably comfortable by the standards of that time. There were some details of design and decoration which were unconventional, as one might expect from Percy. The most original room was the living room with its large Steinway grand piano at one end, and the fireplace at the Leonard Avenue side, complete with an inglenook [chimney corner] and padded bench and bookshelf. On the ceiling were four wooden beams holding metal leaves made by Percy and designed to hold four light bulbs, which gave the ceiling a stylish appearance. Around the walls of the living room were several paintings of birds by Brooks, Fuertes, Lodge and others. A chandelier hung from the ceiling. A room that Percy was especially pleased with was the sun room with a large desk for him to work at, a couch by the windows facing onto the garden, and a phonograph, though this could be brought into the living room when required. There were paintings round the walls, some probably by Frank Hennessey. Since Taverner worked in the sun room in the winter there was a system of forced air through ducts which had ornamental metal grills, another sign of Percy’s workmanship. Glass doors separated the living room from the sun room. Upstairs there were four bed- rooms, the visitor’s room having a small wrought iron balcony. The basement was well planned and built with superior workmanship, no surprise since Percy designed it for his own use, and watched over it being built from a rented house quite nearby. Karel kindly sketched for me a plan of the basement as it was in the 1930s which included a darkroom for Percy’s photography, and a still room. Karel said that Percy liked a glass of Scotch, and bought it by CRANMER-BYNG: A LIFE WITH BIRDS 183 the carboy. These he stored in the still room in the order of the year bought, so as always to have well- matured Scotch available. By nature Percy was con- vivial with friends, but Karel never saw him drunk, only merry in a good sense of the word. He was never a lone drinker, a habit which he disliked, but he enjoyed a drink when in pleasant company. At the AOU Stated meeting each year an evening was set aside when the men who wished to really drink, and tell stories, could get together without the company of wives. Percy had a ready wit and took an active part on these occasions. While living with his moth- er and stepfather Karel learned how to work in cop- per and pewter with Percy.®? Various things they had made were to be seen in their home, and at their cot- tage at Blue Sea Lake. One of the piano students, Barbara Reynolds, spent so much time with Martha that she became a longterm member of Martha’s “extended family” of young people in the 1930s and early 1940s. About her life with the Taverners she wrote: “There was always something going on in the house. The sun porch was often the centre of activity. I watched Percy bind old books, learned how to knit German style and listened to records. Sometimes Sinbad, the German shepherd, would groan to the music and had to be removed. The Taverners and 45 Leonard enriched my life and that of many others.”?” By the late 1930s the street layout in the Taverners’ section of Ottawa had grown considerably since building started in the 1910s. At the south end of Leonard Avenue runs the Rideau River through flat land, and liable to flooding extensively. At the north end of the avenue is Colonel By Drive and the Rideau Canal. Trees and shrubs lined both sides of the canal between Bank Street to the east and Bronson to the west where a swing bridge crossed the canal. On a summer’s evening the north side of the canal, as seen from Leonard Avenue at Colonel By Drive, had mature trees in the gardens which stood back from the gently curving canal, adding depth and variety to the greenery. The open sky above the canal and gardens, on warm summer evenings, was espe- cially peaceful. Here a procession of clouds, like mountains of white cumulus, might drift slowly by, and a male Redwinged Blackbird proclaim its territo- ry with a loud conqueree, and a silent gull or two might glide effortlessly above the canal - these were evenings to quiet the mind and rejoice the heart after the petty vexations of the day’s work. In 1937 Taverner received a letter from a young nephew of his long-time ornithological correspon- dent, Frank Farley of Camrose, Alberta. His name was Farley Mowat. Through his uncle he wrote to Taverner about his bird watching activities in Toronto. The following year he wrote asking Taverner for advice on a project that he and another naturalist, Frank Banfield, hoped to carry out in the 184 summer of 1939 in Saskatchewan, making a collec- tion of birds and mammals. Mowat asked Taverner two questions: where would be the best area to col- lect specimens, and could he recommend museums in Canada and the States where they could hope to sell them. Taverner agreed to help and a correspon- dence followed between the young student Mowat and his senior ornithological mentor until early 1940. The collecting expedition in Saskatchewan was a success, and both men had short accounts published.?! Taverner’s Last Publications In 1934 a new style of bird guide was published by an unknown man, twenty five years old. His name had a musical ring about it — Roger Tory Peterson — and the title a down to earth sound — A Field Guide to the Birds.?* This book was reviewed in The Canadian Field-Naturalist by Taverner who went straight to the heart of the subject in his first paragraph. “A book that the amateur bird observer has long been looking for and which the most seasoned ornithologist will find of value. Presented as ‘a bird-book on a new plan’ it is the culmination of a movement in popular bird literature, that has been for some time in progress, where- by bird study has been stripped of its technicalities and with no loss of real accuracy, the meat offered to the gen- eral public without a shell of scientific verbiage and pedantry. It seems difficult to see how, within its class, it could be improved upon. It is a very practical working manual of field-marks, ignoring confusing detail and con- centrating on the salient characters by which one bird can be distinguished from another under ordinary field condi- tions. It is not for the closet naturalist identifying birds in the hand who has other literature for the purpose; but, for the field worker who depends on eyes and glasses for the recognition of species, it will be invaluable. For this pur- pose it is far and away the best that we have seen.” Taverner wrote with an artist’s appreciation, and an ornithologist’s experience. He had, he wrote, tried to do something along similar lines, and could speak of the difficulties, as well as the success of the author’s accomplishment. Taverner said he had examined the book with a rather critical eye, the result of considerable experience, but had found little to object to and much to admire. He then discussed Peterson’s illustrations. “Nearly every species is illustrated either in black and white or in colour. A few of the former are pen and inks but most are wash-drawings in half-tone. They are not naturalistic representations, but are more or less conven- tionalized or diagrammatic, with confusing details omit- ted, to show the bird as it appears to the eye in the field with the critical distinctive points emphasised. The drawings are remarkable for their clear definiteness. The author-artist shows not only competent draughtsmanship and command of his art-media but knowledge and feel- ing for his subjects. In many cases in these simplified sketches he has caught the spirit as well as the outline and pattern of the various species. Many birds in juxta- position are shown in identical attitude, a proceeding THE CANADIAN FIELD-NATURALIST Vol. 110 that may not satisfy artistic yearnings but is of practical assistance to direct comparison. Particularly should be mentioned the coloured plates of sparrows and warblers; these birds are shown of small size but with a sharpness of detail that is unusual in four-colour half-tone work. For these the plate maker and the printer should be com- mended.” He ended his review with a practical note on the format of the book. “It is a light, compact little volume in waterproof leatherette binding to stand the weather and fits nicely into the side coat pocket, just the form, size and sub- stance to accompany one on a walk in a birdy neigh- bourhood. We highly recommend it and anticipate a great success for it. — P.A.T.”% Taverner’s large book Birds of Canada was pub- lished in the same year as Peterson’s Field Guide (first edition). The difference between them, both in purpose and approach could hardly be greater. But in 1939 Taverner, himself, published two pocket field guides. How this came about he explained in a letter to Brooks. “I am preparing a little field-guide book something between Reed’s pocket guide and the Birds of Canada. Doing it for the Mussons. Think it will be quite a popular amateur aid and it is a rather interest- ing job. We are using the colored plates from the bird book and a lot of new line drawings.” In fact he wrote two books in order to make them pocket field guides. These were titled Canadian Land Birds and Canadian Water Birds: Game Birds: Birds of Prey. In the Foreword Taverner explained that this small book was designed as a hand-book of Canadian land birds that would be of interest to amateur ornitholo- gists. It was not intended to be a complete check-list of the birds of Canada. He explained that instead he had included a Field Color Key of land birds based on spring males of the more characteristic or com- monest species. For instance four species begin with the words blue-gray, twenty-two begin with the color red, and continue from there so that “Red spot on crown, otherwise dull olive” is distinguished as Ruby-crowned Kinglet, and is described on page 148, with a coloured illustration of both Ruby- crowned and Golden-crowned Kinglets, and a sen- tence on their “field marks”. The Introduction is use- ful, covering such subjects as bird photography of which he wrote: “It forms a great sport as exciting and demanding as hunting with a gun, and gives more permanent and satis- factory trophies as a result”. Another subject dealt with was bird houses for colonial nesting birds such as Purple Martins, with eleven illustrations of different designs of houses and feeders.” In the same year that Taverner’s two guides were being published, Peterson’s Field Guide appeared in a revised and enlarged edition (1939). It received an appreciative review from Frank Chapman, which began: 1996 “When, at the age of five years, a book graduates into a ‘completely revised edition’ one concludes that it was well born and has passed a distinguished youth. That its future will be marked by many new editions there can be no doubt, but the present one leaves small room for fur- ther revision.””° While Peterson’s guide began at loons and con- cluded at Snow Buntings Taverner’s Canadian Land Birds began at Band-tailed Pigeon, illustrated by a very life-like painting by Brooks (size 8 cm x 6.5 cm) from Birds of Canada (1934). The banded tail was shown in detail by a line drawing from Taverner. The next species were Mourning Dove and Passenger Pigeon with illustrations showing the two species side by side and drawn to scale of 1/4 of life size. The Yellow and Black-billed Cuckoo followed with black-and-white drawings of the two species in flight on the left page and a coloured illustration of the two on the right page drawn to 1/3 scale. Four hummingbirds were included: Ruby-throated, Rufous, Black-chinned, and Calliope Humming- birds. The last two species were illustrated by black- and-white drawings only. Not all coloured illustra- tions were by Brooks; a number were by Hennessey from the colour plates in Birds of Canada. The dif- ference in style of bird illustrations shows up clearly in a colour-wash of an Eastern Phoebe by Hennessey, which has a page to itself. The posture of the bird, as it sits upright on a small branch beside a wooden bridge, is very characteristic. Some notable illustrations by Brooks show: Steller’s Jay; Clarke’s Nutcracker; Varied Thrush; as well as 19 species of wood warblers in spring, and 18 species of sparrows, finches and buntings. In his Foreword to Canadian Water Birds Taverner explained that Land Birds and Water Birds were intended as non-technical handbooks for identi- fication of practically all the birds of Canada, and most of those in the northern part of the United States. Many of the illustrations in the Water Birds volume were from Birds of Canada but two new ones were painted by Brooks, two were by Cyril Johnson, one by Ronald Smith, and one was ini- tialled P.T. In a short introduction to “Water Bird Study” Taverner discussed the decline in numbers of water birds, and the need for strong laws for the preservation of wildlife.”’” Taverner’s Canadian Land Birds was reviewed briefly by Frank Chapman in Bird-Lore. He wrote, in part: “Of all the guides to our birds, I do not recall one that seems so well designed to fit the student’s pocket and his needs as this one of Taverner’s. With the exception of two Petersonish additions, apparently by the author, the coloured plates by Brooks and Hennessey are from Taverner’s “Birds of Canada’. As a rule they occupy the upper part of the right hand page. Beneath them are paragraphs on field-marks, habitat and nest, and on the facing page a freshly written biography which rivals in CRANMER-BYNG: A LIFE WITH BIRDS 185 form and content the life histories of the earlier volume [Birds of Canada] ... There are introductory chapters on ‘Method of Study’, ‘Birds of the Garden’, a ‘Field Colour Key’, and the best of all an announcement that a companion volume on the water birds, game birds, and birds of prey will soon follow.”?> Taverner replied thanking him for his review and saying “You have always treated my books kindly and there is no one whose judgment in this particular field I regard more highly.” He continued: “T note the Petersonesque color plates but no one seems ever to have remarked the Taverneresque hawk-plates in Peterson which, however, is nothing against that talented artist-writer whose Guide I greatly admire ...”?? Taverner’s Canadian Water Birds, Game Birds, Birds of Prey was reviewed in Bird-Lore by Robert Murphy, who wrote: “Mr. Taverner, who is to all effects the official ornithol- ogist of the Dominion of Canada, here continues his series of true pocket handbooks. The present volume covers most of the birds with which sportsmen come into contact, and the author hopes that it will serve to inform them “upon the species that they seek to perpetu- ate.” By attacking his object through keys, brief and selec- tive descriptions of plumage, and a wealth of illustration in both black-and-white and color, Mr. Taverner has produced the most practical of all books for its field ... Mr. Taverner’s brief biographical notes are excellent, with an intelligent and sympathy-awakening plea for conservation running throughout.”!0° In 1940 Snyder and Shortt nearly caused bad feel- ings between Taverner and themselves. This was a matter that, if it had come to a head, might have resulted in a permanent breach of friendship to the detriment of all three ornithologists had not Taverner backed down and kept his feelings to himself. The matter arose as follows. In 1924 the Biological Board of Canada felt the need of an authoritative manual of the birds influenc- ing marine life along Canada’s shores, and Taverner was asked to prepare this. Subsequently the Board changed its mind and Taverner was told the manual was not required. He decided to enlarge its scope to cover other ornithological fields. This was complet- ed and presented for publication in 1931. For reasons of economy it was not published at that time. Since then Taverner annotated it from time to time in an effort to keep it up to date. After Taverner had been made Chief, Division of Birds, at the museum he again made efforts to get it published, and another major revision of the manuscript was carried out. A copy of the ms. entitled “Practical Manual of Water Birds, Upland Game Birds and Birds of Prey of Canada” was sent to F.C. Lynch, chief, Bureau of Geology and Topography.'°! In his introduction to the manual Taverner described it as designed as a practical means of identifying the Water, Game and allied birds of North America. 186 “.. It has been particularly aimed at the sportsman and general naturalist, it is hoped that it is not thereby dis- qualified from usefulness to the serious specialist.” The ms. followed the slow path of a document in government routine. Eventually a memorandum about the publication of a Museum Report on the “Practical Manual of Water Birds” etc., signed by John McLeish, Head, Mines and Geology Branch, called a meeting on 9 August 1938 at which Lynch, L. L. Bolton, Taverner and two others agreed that: “The Department proceed at once with the publica- tion of the English edition of Water Birds, Upland Game Birds and Birds of Prey. In due course this recommendation was sent by McLeish to Camsell. Taverner appears to have had a presentiment of things going wrong again with the manual in a let- ter he wrote to Brooks written early in 1938. He said that he had been working against time to bring it up to date on a chance of publication, but feared that he had missed out “for this fiscal year at any rate.” He hoped that he could get it published before his retirement time which was not far away. Then he made a revealing remark. “Perhaps I won’t be so sorry when that time comes, I fear the dead- ening influence of the Civil Service is beginnning to get me.”?!02 Nevertheless Taverner was surprised and hurt by the turn of events in 1940. Apparently without warning from Snyder or Shortt, Francis Kortright called on Taverner with a letter of introduction from Snyder which read as follows: “Dear Mr. Taverner: This is to introduce Mr. Kortright who is doing a nice job of producing a Duck-Goose Book. I have suggested that he should follow you in the Branta group and get your opinion on Todd’s new Ungava form. Otherwise he is following the A.O.U. (1931). ?'m sure he will appreciate any suggestions and I assure you we will appreciate it if you give him help. Regard. Yours truly Les Snyders 20? Worse was to come as this extract from a letter of April 1941 from Taverner to Snyder showed. “Kortright was in the other day and he showed me the paintings and many pen and inks that Terry ‘has made for him. Terry has done a mighty good job in these and is to be congratulated. It is going to be an outstanding piece of work but I can see how heavily he has leaned upon you and your museum. It however puts quite a fly in my ointment for it completely overlaps and renders useless the manuscript I have been working on for years but which, without all these colored plates, cannot begin to compete with it.”!4 Snyder replied that he regretted the overlap that the Kortright book caused with Taverner’s ms. but explained it by writing: “Money talks and he has done a good orderly job of compiling this book. Terry tells me there is some talk of cutting out his beautiful plates of eclipse and off- plumages which have made the book unique.” ! THE CANADIAN FIELD-NATURALIST Vol. 110 At this time Taverner was working on a paper on the distribution and migration of the Hudsonian Curlew [Whimbrel], and asked Snyder for informa- tion on the specimens the ROMZ possessed. When thanking Snyder for the information he said in the same letter, “Yes, it is something of a disappointment that my big opus should be so anticipated and overpowered by the Kortright book, but that cannot be helped. Mine was on the verge of publication several times in the past decade or so but always fell through at the last moment. Of course I cover more than ducks and geese but there is hardly enough left with them out.”!°° Taverner accepted the disappointment as best he could. He had a warm nature and was not the man to bear a grudge for long. Friendship to him was more than success, or getting his own way. He could not expect this compilation on ducks, backed by Ducks Unlimited, and supported by the ROMZ, with the superb colour portraits by Terry Shortt not to take the market. Taverner’s mistake had been to enlarge the scope of his work from water birds to include game birds and birds of prey. By 1938 wild ducks were important in the eyes of sportsmen and natu- ralists, and a book about them could be expected to sell. Compared with Kortright’s book which was forward looking in 1942 Taverner’s ms. was rather outdated by then. This extensive study was complet- ed, but unpublished when he died, and has remained so ever since. !°7 Three papers, for which Taverner had been col- lecting material for ten years or more were pub- lished just before he retired. The first, a short paper entitled “Variation in the American Goshawk’”, appeared in The Condor.'°* After examining varia- tions in the plumages of two series of specimens he proceeded to demonstrate that fineness of pattern had no particular geographical distribution, and was a process of age, not a racial character. However, a group of [Northern] Goshawks from the coastal islands of British Columbia stood out strongly from the others. They included both finely and coarsely marked birds, but all showed degrees of darkening. Taverner devoted the remainder of the paper to a discussion of the degrees of darkness in birds from Queen Charlotte Islands (5 specimens) and Vancouver Island (19), and the lack of darken- ing in birds from closely adjoining mainland locali- ties. From comparing these specimens Taverner felt that there was a recognizable strain of goshawk on the two islands distinct from Accipiter gentilis of the continental area.! Taverner then wrote “ I therefore propose Astur atricapillus laingi new subspecies. Queen Charlotte Goshawk (Named in honor of Hamilton M. Laing, who has been instrumental in uncovering the form.) Ottawa, Ontario, January 12, 1940.” 1996 Taverner’s new subspecies was subsequently accepted by the AOU Committee on Classification and Nomenclature of North American Birds. Taverner’s second paper, published in The Wilson Bulletin near the end of his life, was on a very differ- ent subject. Its title was “The Distribution and Migration of the Hudsonian Curlew’.!'° During his voyage in the Arctic in the S.S. Beothic in 1929 Taverner noted the huge numbers of Hudsonian Curlew [Whimbrel] that migrate along the north shore of the Gulf of the St. Lawrence and down the east coast of the U.S. In a letter to Fleming he raised the question of where, in the arctic, they bred. “They are practically unknown on Baffin Island but must work up into the central arctic islands. Soper did not find any in eastern Baffin Island. The great thing will be when arial [aerial] navigation is developed so that we can use it to reach this country. To gét in and out in one season will open a great field for ornithological work up there.”!!! Information on the eastern migration was consid- erable. Taverner introduced it as follows: “The Atlantic migration is more complicated and fol- lows different routes in spring and fall. The Hudsonian Curlew is powerful on the wing and quite capable of making long sustained flight. Its staple food in the north is the low-lying fruit of the subarctic barrens, particular- ly the crowberry (Empetrum nigrum) on which it gorges. Farther south the fiddler crab of the sand beaches seems to be its main food (Wayne, 1910). Its migrations are probably largely governed by the presence of these or similar foods. Where they are absent along travelled routes the birds are likely to pass over or pay only occa- sional visits in case of necessity.’ '~ In a summary Taverner restated briefly the main points of his paper, and in his final sentence said: “We suggest that there may originally have been a Mississipi Valley group connecting the two breeding areas in the north but which were recently extirpated — perhaps along with the Eskimo Curlew, with which the species seems to have been closely associated in migration.” “Canadian Races of the Great Horned Owl” (here- after G.H.O.) was the third of Taverner’s late publi- cations.!!? For most of his years at the National Museum Taverner was collecting specimens of the G.H.O. in order to work out the races found in Canada. His earliest attempt was a brief discussion of three races in his Birds of Eastern Canada. About designating subspecies, he warned that: “As these [races] intergrade with each other indistin- guishably and overlap in range in migration, exact sub- species designations should be made only with great care and, except in extreme plumages, only after com- parison with duly authenticated specimens.” CHAPTER 17. In Retirement Percy Taverner was due to retire on reaching sixty- five years of age in mid-1940. But by then the Second World War had started, and since no replacement had been found for him Taverner’s appointment was CRANMER-BYNG: A LIFE WITH BIRDS 187 Taverner’s account of the G.H.O. in Birds of Western Canada is considerably longer than in his Eastern Canada book. Here he discussed the five races recognized in Canada in the AOU Check-list (1910), with a generalized description and distribu- tion of the west Canada subspecies. Finally he made one more attempt to bring his study of the races of the G.H.O. up to date before retiring. In a letter to Laing he wrote that there was little bird news, but Allen, editor of The Auk, had accepted his Horned Owl paper with “commenda- tion”. Taverner continued: “It does run counter to many philosophies and will probably not be generally accepted, — at first. However I do think that we should let up a bit on the microscope and use a telescope more to get the general view of our specific variations in a broader field. Many cannot see the woods for the trees.”!'* When finally it was published it contained his summation of about 25 years of intermittent work on the subject. Taverner’s Birds of Canada (1934) was becoming out of date by mid-century, but there was no book of equal scope to replace it until a completely new work with a slightly different title, The Birds of Canada, by W. Earl Godfrey was published by the National Museum of Canada in 1966.'!> Thus a period of 32 years passed between the publishing of Taverner’s Birds of Canada, and Godfrey’s The Birds of Canada. In those years the great changes which took place in ornithological studies can be measured by comparing Taverner’s national-scale study of ornithology in Canada with Godfrey’s on the same scale, but undertaken by ornithologists with different training, and working with very dif- ferent equipment. The breeding distribution of the bird species shown on maps in the 1966 volume was an important innovation that Taverner would have been delighted to have used if he could have afford- ed it in 1934. John Crosby’s coloured illustrations were deliberately painted and printed to give the ornithologist a high degree of definition in a refer- ence book designed for use in library or laboratory. These bird illustrations show a clear difference from the style employed by Brooks. A comparison between the Taverner Birds cf Canada and Godfrey’s The Birds of Canada aptly demonstrates is that the study and practice of ornithology is an ongoing and co-operative pursuit. extended for another year. This was a disappointment for Percy and Martha who had been looking forward to retirement from early 1938.! Finally, Dr. A. L. Rand accepted the position of assistant zoologist at the 188 National Museum of Canada in March 1942.7 While Taverner was handing over to Rand he was also in the process of going into retirement, not a simple matter for someone who had held the same responsible posi- tion at the national capital for 31 years, and during the difficulties of wartime. If there had been no war then he could have attended the AOU meeting in 1940, with an opportunity to say good-bye to a fair number of his long time ornithological friends. But suddenly financial controls had become very stringent. Visitors on business to the States now had to supply an item- ized expense account of necessities, and were limited to that amount. In a letter to L. Bishop of July 1940 Taverner said: “Tt seems strange also to me who has bounced back and forth across the line all my life and felt equally at home either side that I am [an] alien with passport necessary, photographs and finger prints. I wonder if the old free- dom will ever return. Probably not in our time, these things have a way of sticking like pitch.” The only way that Taverner could tell naturalists in Canada and the States that he was retiring in 1942 was to put a notice in The Canadian Field-Naturalist, which he did. It was published as follows: “Tt is with real regret that The Canadian Field-Naturalist publishes the following announcement of the retirement of Mr. P. A. Taverner as Ornithologist of the National Museum of Canada. Mr. Taverner has served long and well and this journal is deeply indebted to him for his services as Associate Editor (Ornithology) over a period of many years. — Editor. This will announce to those interested that, after thir- ty-two years, I have reached the legal limit of service and have,retired from the position of Ornithologist in the National Museum of Canada. The Division of Ornithology will continue under the direction of Dr. A. L. Rand, whom I heartily recommend to correspondents, friends of the Museum and ornithologists in general. I hope they will extend to him the same support and assis- tance that has made my past labors pleasant. Future offi- cial communications should be addressed to him.’ In the last paragraph he said that he would not be dropping his interest in ornithology or in the muse- um, and would always be glad to hear from his ornithological and other friends. His name and address was printed at the end of his letter. At the same time he told some of his friends, by private let- ters, of his retirement. Here is Percy Taverner with a graphic account, in a letter to Mack Laing, of what it was like when he reached the end of his career. He said he was in “‘an awful mess” because he was due to retire shortly. “Pulling thirty years of roots out of all the stuff here [i.e. in the museum] is difficult ... and there is a thorough “change of life” ahead.” Then there was the problem of space at home for his books. “Even our home here is uncertain. We would like a milder climate and smaller quarters for us two ... yet THE CANADIAN FIELD-NATURALIST Vol. 110 what can we do or where can we go under the war restrictions. Have thought of the west coast. What are the possibilities of somewhere near you?” Next he told Laing about Rand who had started work at the museum in March and what it felt like to see a “new broom” at work, and what Rand’s views on ornithology appeared to be. One point was clear: “He is much more of a critical subspecies shark than I am but seems well balanced.” What Taverner might have said, but didn’t, was that Rand was ambitious, and determined to get his name into print quickly. Although he only started work in March he already had an article published in The Canadian Field-Naturalist of November- December 1942, and on a particularly Canadian sub- ject — Kumlien’s Gull.° In this paper Rand gave a short outline of the breeding distribution and rela- tionships of eight Nearctic gulls of the Genus Larus in the National Museum collection. The section on kumlieni was based on Soper (1928) and Taverner (1933). The relationships of the group were repre- sented diagrammatically, and a map showed their overlapping distributions. In the last sentence Rand wrote: “In preparing this paper I have had the privilege of dis- cussing it at length with Mr. P. A. Taverner.” From Taverner’s point of view he must have been only too glad to introduce Rand to the material on Larus kumlieni and its allies that he had been collect- ing for the museum since 1916, and the result of many discussions he had held with other ornitholo- gists. Under Rand’s name, at the beginning of this paper were the words: “Ornithologist, National Museum of Canada, Ottawa”. while immediately beneath Rand’s paper was the heading “Mr. P. A. Taverner Retires”, a neat handover in print from the outgoing to the incoming ornithologist at the National Museum of Canada. Answers to Taverner’s valedictory letter on his retirement were coming in through the autumn. Twenty-four have been preserved, though Taverner may have received more. An official letter from W. B. Timm, director of the Mines and Geology branch, reached Taverner “placing on record the appreciation of the Mines and Geology Branch of your long and conscientious service of over thirty-one years.” In the pompous prose of senior civil servants of that time he was patted on the back for his “laudable achievements”. The situation at the Geological Survey to which Taverner was first appointed in 1911 was a lot less pompous when the lively R. W. Brock was director. In a different style was a warm personal letter from Harrison Lewis, chief Federal Migratory officer for Ontario and Quebec, saying how much he appreciated Taverner’s help during his early years in the service. In another vein was some- thing more formal from a well known member of the AOU, and assistant secretary at the Smithsonian Institution who knew Taverner quite well, but less 1996 personally than did Harrison Lewis. This was Alexander Wetmore who wrote, in part: “.. You yourself have done a great piece of work in sys- tematizing the collection of birds in your Museum, and also in adding to them tremendously through the field work that you have carried forward. I know full well the difficulties that you have had in this and also the plea- sure the actual work has given you.” After Taverner had handed over the direction of Ornithology to Rand, and received the usual six months’ leave of absence before final retirement, he wrote a four page typewritten “Biographical Outline” of himself. This was a very modest and down to earth account of his development as a taxidermist, collector of skins, bird bander and as a student of ornithology at Point Pelee, which covered the first two pages. The other two were devoted to an outline of his work as Ornithologist at the National Museum.® When W. E. Saunders died in 1943 Snyder asked Taverner for a contribution to the Saunders’ memori- al programme that the ROMZ was planning.’ After it had been held Snyder wrote to Taverner telling him how well his contribution was received, and then turned to an account of Taverner’s own career, and asked him: “Has anyone ever urged you to write your autobiogra- phy. I’m doing it now. Perhaps the size of the volume which it deserved would be too much of a labour but let me have a few chapters — the cream so to speak.” Taverner replied in part: “I do not feel that there has been enough of general interest in my life for an autobiography. I have prepared an outline for use when the inevitable time comes for an Auk memorial which I plan to deposit with your Museum. There are so few left who know anything about my previous life that some such guide seems nec- essary if the Auk continues the “obits” of Fellows.”!° For a person in the process of retiring from a career that has taken up a major part of his adult life it is important to have interests, and perhaps a pro- ject which will keep his mind alive, and warm with a feeling of still being able to achieve something dur- ing his retirement.'' Taverner’s mind, at his time of retirement, and for a few years after, was still rela- tively sharp and forward looking, in spite of the loss of several contemporaries of whom he was fond, particularly Fleming. But Taverner had the ability to make new friends among ornithologists, and to share past experience and new interests with younger men of professional training. One such ornithologist was Ernst Mayr whom Taverner met at the AOU meeting in Washington in 1938, when they discussed new ideas concerning theories and practices in ornitholo- gy. Taverner learned from conversation that Mayr was especially interested in systematics and evolu- tion, and in a few letters between them he was encouraged to examine the subject of bird species once again. In a letter to Snyder late in 1940 Taverner showed what he had been reading on the old problem of subspecies — J. Huxley on “the CRANMER-BYNG: A LIFE WITH BIRDS 189 cline”, E. Mayr’s paper on speciation, and David Lack on speciation. Lately he had spent considerable time on the water thrushes, he wrote, but failed to make head or tail of them. “Perhaps I lack power of fine appreciation but anyway I fail as a splitter.” What he had recently read suggested to him a paper that he wrote and called “Subspecific Values”. This he sent to Snyder.'* Also Mayr suggested that Taverner should read his book Systematics and the Origin of Species." In March 1943 Mayr wrote that he had learned with regret of Taverner’s retirement from the posi- tion as Ornithologist of the Canadian National Museum. “However, I hope that this means that you will have more time to devote to the study of birds and to other pursuits in which you are interested.” Then he told Taverner that lately he had become interested in the question of the degree of overlap between bird populations. In this connection he studied with profit Taverner’s papers on the Long- tailed [Black-capped] Chickadee and the giant Red- winged Blackbird.'* But Taverner did not give his complete measurements in these papers and Mayr wondered whether he recorded them. This was unfortunate, he wrote, since Taverner’s data “‘repre- sents a particularly nice case and I would like to use them if they are available.” In the last paragraph he asked Taverner “Did you read my book and what do you think of it?” In reply Taverner explained how the graphs in both papers were made, and that the museum had the original tables from which the graphs were made, and that they would be available to him if he required them. Next Taverner discussed Mayr’s book which he had just finished reading. He praised its clarity of presentation in places where unusual technical words were used.!° During the late 1930s we saw that Taverner was discussing changes in research into ornithology, and methods of teaching it. As a result he had begun to accept this as a necessary change in methodology even if he did not feel comfortable with it. When, in 1941, he received a letter from a farm boy in his mid-teens in Alberta, who had been forced to break off schooling due to lack of funds, asking for Taverner’s advice on his chances of making a future in bird work in Canada, Taverner knew how to reply.'’ He began by saying that he received many letters of the same nature, some from young fellows who seemed to think the career of a naturalist an easy way of making a living, but few seemed to real- ize that “nowadays” more preparation than a desire was necessary for success. He went straight to the point, saying: “The time of the self-taught naturalist achieving great success is past ... Today much more is demanded than an uneducated love of nature and an urge to wile away one’s days in the woods.” 190 It was necessary, Taverner explained, to have a sound knowledge of what had been accomplished in bird study, and what awaited to be done, as well as a familiarity with modern concepts and proceedings. This was difficult enough but when complicated by the necessity of making a living at the same time it was almost “prohibitive”. He continued: “Nowadays the only practical door to a successful living through natural science is through academic education. The country today is full of well qualified college gradu- ates in line for just such positions as you probably would like, and these have such a legitimate edge upon others that one might say none other need apply.” Taverner told Beddoes that he was not saying this to discourage him but to state the conditions and to point the way forward, and that many had struggled up from the most unpromising conditions “but only through hard work, intense application to an end and through University courses”. He then pointed to one way forward: “... There is always a lack of qualified nature teachers in the schools which gives one line of progress that depending upon the individual may lead to many things. As has been said many times there is always room at the top but the competition is keen and only the most deter- mined and best qualified ever reach it.” Taverner tried to soften the harshness of his letter at the end by saying that “many have wangled [struggled?] their way through higher education against the greatest of obstacles. Others with all the ambition ... have been by force of circum- stances unable to do so, but usually where there is a will there is a way. Sometimes there is a conflict between duty to self and to family that stands in the way. These are hard cases and the outcome is up to the individual.” In his last sentence Taverner wrote: “T wish I could offer you more material assistance and advice.”!® Meanwhile the war in Europe exploded into vio- lence in the spring and early summer of 1940, putting an end to any hope that Percy and Martha had for an enjoyable retirement. However, quite unexpectedly Taverner received a letter from a woman living in a log-house on a cliff overlooking the Mattawa river in central Ontario near North Bay. She signed her name Louise de Kiriline Lawrence. Her reason for writing, she explained, was that she had been given a copy of his Birds of Canada. With the help of his book she had begun to keep records of each species she could identify positively. Her love of nature, she said, had begun as a child in Sweden. Now, in this isolated spot in the woods, with feeding trays and suet hung from branches around the log-house, she was deriving immense pleasure in observing the behaviour of the various birds around her. “May I tell of a few things I have seen?” she asked. Then she described with skill and warm feeling some of her observations in the spring of 1940. A pair of Purple Finches mating, six pairs of Rose-breasted Grosbeaks nesting in the vicinity, a Veery sitting on four eggs, a pair of Pileated THE CANADIAN FIELD-NATURALIST Vol. 110 Woodpeckers. Here, in the first of twenty-one letters that followed, were the kind of vivid observations that later made her books such as The Lovely and the Wild so enjoyable.'? But in these letters she was writing about birds in anthropomorphic terms using such phrases as the male Purple Finch’s “impas- sioned wooing of his demure little bride” and a male Rose-breasted Grosbeak singing continually near the house until she wondered “if he was neglecting his duties or whether he was a bachelor’. At the end of this first letter she apologized for taking up Taverner’s time for so long, but excused herself by saying that the only drawback of living far away from other people was that “one needs occasionally to find someone who knows more than oneself with whom one might be permitted to share impressions, crystalize observations and find guidance. Hence this letter which I trust you will not look upon as a pre- sumption of me to have written. Meanwhile your book, excellent in a practical way and delightful in a literary sense, remains to me a prized guide and friend. If ever you should pass by this log-cabin would you stop and pay us a visit?””° This letter reached Taverner at a low ebb in his career when he was wondering if the work he had done was of much value. In his reply to Louise Lawrence he expressed his thoughts frankly, and modestly. “Of course it is a great satisfaction to feel that any work one does is so thoroughly appreciated. In these days of great trial it seems that the study of little birds is futile and pickiune [picayune], but when I get such letters it makes one feel that life has not been entirely wasted, at least it has brought pleasure to some and perhaps to many more than have been moved to express it. “How much heredity and how much environment and early associations have to do with our developed person- alities is an open and controversial question. But when they both combine to turn us towards keener apprecia- tions of nature or of art it matters little which have had the predominant influence.” He promised that if he passed their house he would be happy to drop in, and invited her, if in Ottawa, “to look in on me”.”! Their correspondence had begun well. Both had stated their need, Louise Lawrence openly, Taverner less explicitly though obviously enough. In her next letter Louise Lawrence took the opportunity of an episode concerning a duck that died of a gunshot wound, while she was caring for it, to write to Taverner again. From this letter Taverner learned that Louise Lawrence had a husband who had recent- ly joined the forces, that she had a .303 rifle which she fired from her bedroom window to warn migrat- ing ducks, in the bay below her, of danger from “sportsmen” firing from their car on the bridge on the highway. When the duck died she realized that she could at least make sure of its identity. She judged it to be a female Ring-necked Duck, and learning from Taverner’s Birds of Canada that more 1996 | information about their range was needed, thought that her story about the “wingshot one” might interest | him. In the same letter she related how six Wilson’s Thrushes [Veery], each in its own tree-top near the | cabin “jingled their golden chains”. to borrow a | phrase from his book, in a matchless serenade.” She ended her letter by explaining her strong need to | communicate her impressions to a fellow being with the same feelings about birds. “I would not wish you to feel under any obligation to acknowledge this unless you be so inclined, but please, permit me the privilege to write to you from time to time about my _ fascinating neighbours” .** Who could refuse such a gentle request? Taverner himself had written about birds with a warmth of observation and a touch of sentiment just as Louise Lawrence was writing to him now. On the verge of retirement he had found another bird observer, in an isolated spot, whom he could encourage to report individual species with some degree of scientific accuracy; yet another link in his ornithological network. On her part Louise Lawrence would have the satisfaction of knowing that her bird notes were going to be read at the National Museum by someone who appreciated her efforts, and sent her encouragement. During the next few years Louise Lawrence wrote letters filled with the excitement of her discoveries, such as the arrival of a male Cardinal at her log- house in the winter of 1941, and the appearance of a Black-billed Cuckoo in the summer of that year.” Taverner answered that the identification of the Cuckoo sounded correct, and that “one gets quite a ‘kick’ out of meeting and identifying a new bird, especially when it is an unexpected addition to the local fauna”. It was not quite a new species, he said, as it was reported in a Lake Nipissing local list of 1939, but her observation did mark the farthest north extension of range in that quarter.”° Taverner then wrote in general terms about their shared enjoyment. “This is one of the great things in bird study, there is always something new to look forward to with thrills spaced evenly enough to sharpen the interest. If this gen- tle art is a relief from the sterner atmosphere that sur- rounds us it is a blessing indeed. We need such relief these days. If I have had any hand in extending it I can feel that I have been of some use in the world.”7 By late February 1942 the cardinal had started to sing and another letter from Louise Lawrence reached Taverner describing the incident. “It was strange”. she wrote, “standing in the path with waist- high snowbanks just outside my house sitting deep in drifts up to the eaves and listen to the first spring notes of a crimson bird! Is there no climax to the marvels of our northern bush?’”’® By 1942 Taverner had introduced Louise Lawrence to bird banding, and she had told him of her harrowing experience when her officer husband was taken away to be shot during the Russian revolution. Taverner read two stories of hers and liked them. He wrote: “They are CRANMER-BYNG: A LIFE WITH BIRDS 191 literature as well as good natural history and hence hardly material for ‘popular’ magazines. I should think they are Atlantic Monthly stuff. I advise you to try that.” He enclosed the editor’s address.” The correspondence of Taverner and Louise Lawrence contains a moving sequence of letters which deserve to be read, not only for their revela- tion of the development of Louise Lawrence as a serious student of birds, and a gifted writer about them, but also for the part played by Taverner in that development. In a letter written only a few months before Taverner’s death she said: “... the contact with you I count as the most important factor to my introduction into the bird world. It has been a great privilege to have known you, although I have never seen you. You led me on, step for step, and I recall with gratitude the help it was to be able to check with you my first little discoveries, the fox sparrow, the cardi- nal, the cuckoo, now all my long-established friends.’»° This was the end of a long and rugged trail for Taverner, and the beginning of a fresh and exciting trail for Louise Lawrence, both distinguished Canadian naturalists. Besides having their lives enriched by music, alpine wildflowers, interesting examples of domestic architecture and furnishings, Percy and Martha had the last few years of his life enlivened with the liveli- ness of young children. As Percy told a friend, they had recently been up to their cottage to prepare it for Karel who was bringing his family from Lansing, Michigan, the following week. He explained that Karel “has two little girls now as well as two older boys that he acquired with his wife. I expect the girls will be parked with us as our cottage is hardly a suit- able place for such small children. I see that Martha will have her hands full but she loves it.’”*! In 1941 a step-son of Karel, Corwin Ferguson, was staying at 45 Leonard Avenue for more than a year while attending school in Ottawa. Writing to a friend on service in England this is how Percy described it: “We have a small boy with us, — not an English lad as it happens [evacuees from Britain had recently arrived in Canada] but a near relative. He contains more concealed and active energy than a phonograph spring and keeps things pretty lively about the house and place. What a lad of eleven won’t think of to do is certainly unpre- dictable. So you see we are not exactly vegetating.”* Corwin was eleven years old then and he still has some clear memories of that period. He felt that Martha and Percy seemed very close, that she kept a watchful eye on his health, that by 1941 he was not allowed to climb up stairs as often as he would have liked — it was rationed. Martha also told him when he had done enough gardening for one day. But life at 45 Leonard Avenue was not austere in 1941 and 1942 — they still entertained often, and Corwin remembered house guests staying at their house even 192 in the early war years. Also people came from afar to see the rock garden at 45 Leonard Avenue while it was still in its heyday. Above all he remembered nephews and nieces, and “honorary ones” from among Martha’s piano students being invited to stay at Hyla. Here were members of the family and an extended family, as well as close friends, such as Barbara Lowe (Reynolds) and Karin Porsild (Lumsden), all of whom referred to Martha as “Mrs. T” or simply as “T”. She was the centre of the fami- ly, and many people corresponded with her regular- ly. Corwin’s memories of Percy were about his stammer which was always with him, though at times he produced spontaneous witticisms, and amusing puns; he was not “put down” by it.°? Another member of the “extended family” was Marte, Karel’s daughter and Corwin’s half-sister, who stayed with Martha and Percy for about a year and a half,while her parents, Karel and Peggy, were abroad. Marte remembered Martha as a determined woman, not to be opposed. But Percy could stop her doing something if he wished, just by quiet moral authority. Marte told how T was spanking her one evening, and she was squealing, when Percy came into the room and, in a quiet tone of voice said “that’s enough”. and it was. Martha stopped at once.*4 In the normal course of events Taverner would have retired in mid-1940. Already by the beginning of that year he was feeling discouraged. In a letter to Fleming he mentioned the death of Harlan Smith, commenting that “he was crushed by an unsympathet- ic museum and Civil Service regime and had all the heart taken out of him.” Of his own situation he said: “It takes a more rugged man than poor Smith to keep fighting under these discouragements and I am at last beginning to understand his reactions to it. Certainly I have not the enthusiasms I had when I first came, and the fight seems less worth while.”*° One quality that showed throughout Taverner’s adult life was what might be termed “stubbornness”. in the sense of extreme perseverence. A clear exam- ple of this quality can be seen in his never-ending attempts to get his “Practical Manual” published. This work had started in 1924 when the Biological Board of Canada wished to publish a fauna of the Atlantic coast as a students’ manual, and Taverner was asked to write the bird section and include line drawings.*° When the Biological Board cancelled work on the manual Taverner continued with his Birds of Western Canada until that was published. After that he was free to enlarge the scope of his “Water Bird Manual” which would not include game birds and hawks. By now the book was aimed at hunters, and would include a large number of text drawings and line cuts.*’ If Taverner had completed the manuscript by early 1930 it might have been published by the National Museum, but a new Check-list of North THE CANADIAN FIELD-NATURALIST Vol. 110 American Birds was being prepared by the AOU for publication in 1931, and it was important that his book should cover a complete section of the new List. Unfortunately for Taverner, by the time it was sent to the publishers to examine in October 1931 it was too late financially, when the National Museum had to postpone many of its expenses.** Taverner did not give up hope and when he became chief, division of birds, at the National Museum he made another effort to get the manual published. In December 1936 Taverner’s brother-in-law, John McLeish, became head of the Mines and Geology Branch of government.*? Early in 1938 Taverner presented the updated manuscript of his “Practical Manual of Water Birds, Upland Game Birds and Birds of Prey” to F. C. Lynch for publication.*° After various delays throughout the summer John McLeish, from his position as head of Mines and Geology, sent Charles Camsell, Deputy Minister of Mines and Resources a memorandum asking for approval of the publication of the “Practical Manual”.*! However, early in 1939, with the threat of war hanging over Europe, the decision to publish was delayed. At this point Taverner, if he had been less “dogged” might have given up the struggle, but he didn’t. Instead he went on persistently tinkering with his manuscript until in 1943 it was overtaken by a very different kind of book.” Eventually, as his health deteriorated, he seems to have realized that his “big opus”. as he called it in his letters, would never be published. Writing to his old friend, Frank Farley, in 1944, he said that he was working on a book that had long been under way but whether it would ever find a publisher was another matter. “It may be love’s labor lost but what the heck, it makes an objective now.’ So Taverner hung on doggedly until the end, and his last mention of his “big opus” was in his last let- ter to Laing two months before he died.™ The failure to see his “Practical Manual” published before his death after so much work was a great disappoint- ment, but perhaps Percy still hoped that it would be published on his behalf posthumously. But it was not to be, and the manuscript remains unpublished.* In addition to his disappointment over the book, Taverner received very little recognition during his retirement. However, there were a few honours given to him near the end of his life. In 1945 he was elected an honorary member of the Ottawa Field- Naturalists’ Club “in recognition of his outstanding ornithological work, and service in the interest of the Club. ... Probably he is best known to the members of the Club as author of ‘Birds of Canada’, and as long-time ornithological editor of this publication.”*° In 1946 a new race of the Purple Finch was described by A. L. Rand in honour of Taverner as follows: 1996 “Carpodacus purpureus taverneri, new sub-species. Type No. 25387, National Museum of Canada; male adult; Government Hay Camp (Park Headquarters), Wood Buffalo Park, Alberta; May 26, 1933; collector, J. Dewey Soper.” By far the brightest honour paid Percy Taverner was a posthumous one. On the front page of W. Earl Godfrey’s revised edition of his The Birds of Canada (1986) is this inscription: “To P. A. Taverner and Robie W. Tufts with apprecia- tion” Everyone who uses this edition should be glad to glance at the name P. A. Taverner on page 5. It is a living tribute to all that Taverner achieved during his 31 years as ornithologist at the National Museum of Canada. Also it is a mark of appreciation that will continue to live as long as Earl Godfrey’s book is reprinted. Taverner could not have wished for a more enduring honour. In contrast to the last years of Taverner’s life Jack Miner received ample recognition right to the end of his life and beyond. Although Taverner was long- suffering by nature and did not express his inner feelings publicly, there is a hint of being hurt when, in a letter to Snyder, he said: “I see they are making quite a splash over Jack Minor [sic] again. It is quite a joke isnt it, — ‘Canada’s famous naturalist’. One would think him the only man who ever banded a bird’’.** In contrast Manly Miner, writing to Taverner about a visit by Saunders and him to the Miner farm in 1909 or 1910, said: “Father had a great life since those dates. All the honors came to him that any one could enjoy. Yet we all helped him. I acted as secretary, made all the appointments. News releases and so forth. It all didn’t happen in a day.” Manly wrote much more, including the fact that Mr. Mackenzie King was due to spend a weekend at their establishment soon.*? One other posthumous tribute was paid to Taverner by another Canadian ornithologist, Henri Ouellet, in an illustrated memoir published in American Birds, while the 19th International Ornithological Congress, held in Ottawa in June 1986, was still sharp in people’s minds. The title was “Profile of a pioneer: P. A. TAVERNER” and immedi- ately under it was the comment “Canada’s first pro- fessional ornithologist still has a tremendous influ- ence on the study of birds in Canada today”.°° Anyone who has collected a library during their working life is faced with disposing of most of it before or during their retirement, both for lack of space and lack of use. Writing to Baillie in 1945, Taverner asked to borrow from the ROM collection an adult White-winged Black Tern [White-winged Tern], and a skull of one of the large waders to show in contrast with that of a Wilson’s [Common] Snipe. He was still CRANMER-ByNG: A LIFE WITH BIRDS 193 drawing illustrations for his “Practical Manual”. In the same letter he said that he had arranged for the dispos- al of his library — to the Quebec Provincial Museum. “T think I have made a fair deal. At least it will be kept together as a Canadian unit and be where it should be badly needed.”>! The other library that Taverner wanted to dispose of consisted of “old standards”, from classics to moderns in various editions, about 200 volumes in all. For this purpose he wrote to Angus Mowat, Inspector of Public Libraries, Department of Education, Ontario, asking if some country school or similar repository would appreciate them. He added that he would gladly donate them.** Angus Mowat’s reply began: “T have heard so much about you, and for so long, from my bird-crazy son that it came as a surprise and, I may say, a great pleasure too, to have a letter from you this morning.” Mowat then suggested dividing them into suitable lots for distribution among some of the small and struggling libraries in the north country. Any titles which they found unsuitable for that purpose “would be turned over to Frontier College whose people are always glad to receive books of any kind.” In the final paragraph we see father talking about son with a touch of pride in his pen. “Farley [Mowat] is busy with his university course and somewhat to my surprise is working like a nailer at it. Apparently his trip to Saskatchewan last summer was profitable, although not from the financial point of view, and he still hopes to make some arrangement with the people out there which will enable him to make that province his own field. In fact he already talks about it with a most proprietary air.” Angus Mowat ended his letter: “Thank you very much for your letter and please accept my personal regards.” Throughout Taverner’s years of work at the National Museum he was often frustrated by some problem or other that should have been put right but was not, or should never have gone wrong in the first place. To discuss this problem satisfactorily would require considerably more space than can be devoted ‘to it here. It should be written as a separate paper, and printed as a comment on the National Museum, rather than as an integral part of Taverner’s life. It was only one year before he died that Taverner sent W. B. Timm, chief of the Mines and Geology Branch, a memorandum on the Status of the National Museum of Canada.** In the summer of 1946 Taverner had written to an old friend, Dr. George Prey, and asked his advice on the possibility of spending the next winter in Florida, and the problems it would entail. The answer he received was not encouraging. In the fall he applied for accommodation but without success. As Taverner wrote “... everywhere we applied we found crowded.” 194 The winter dragged on, only broken by short visits from old friends. One good omen was that in February 1947 an addition to the staff of the Biological Division of the museum reported for duty — W. Earl Godfrey. But by then Taverner was too ill to exert himself much in conversation and Godfrey only managed to meet him briefly once.» Although Taverner was unable to talk at length on serious subjects he could still tap out letters on his old typewriter. Both A. E. Porsild, who was living with the Taverners, and an old friend of theirs, Mrs. Jean York, had told him that an enthusiastic bird- watching schoolboy might write to him for advice.°° As a result there followed a friendly correspondence between the senior ornithologist at the National Museum in Ottawa and Henrik Deichmann, a schoolboy living near Saint John, New Brunswick. Deichmann wrote at the end of January 1947 telling Taverner that he had owned a copy of his Birds of Canada since the previous year which had made a great difference to his bird study. He was now read- ing it through for the third time, he said. He described where his family lived — in a little valley with a river running to the north, a high bluff in the east covered with trees, many stunted and twisted over the rocks. Looking westward were farms and woods. To the south lay the city of Saint John where on foggy days they heard the fog horn faintly. The letter ended as follows: “Tam the boy Mrs. York was telling you about. I am twelve years old and a terrible speller so I keep running to my mother and ask “How do you spell this or that?” My mother says she wishes I could look things up in dic- tionary as eagerly as I do in the ‘Birds of Canada’.” He enclosed a few of his bird sketches because he was so glad that Taverner had written Birds of Canada. He signed himself “Your friend Henrik Deichmann” Taverner replied that he was glad to receive Henrik’s very interesting letter. That he had heard of him from Mr. A. E. Porsild “who knows some of your family”. He continued: “It is very flattering and a great satisfaction to learn that my work and books have been so usefully employed and have given pleasureable information to others. It is to you youthful inquirers upon whom we have to depend to carry the work forward as we retire from the field.” In the next paragraph he wrote: In your letter I see a very great promise. You write well and I should say have considerable powers of observa- tion. I hope you will cultivate them both. Your pictures also show promise and whether they are drawn from original subjects or are copies indicate that you are learning to control your pencil which is the basis of draughtsmanship. Keep it up and develop all sides of your possibilities.” But when he wrote about the many pathways through modern ornithology this may have seemed somewhat daunting to a twelve year old, though 57 THE CANADIAN FIELD-NATURALIST Vol. 110 Taverner’s conclusion was clear enough when he wrote: “But the basis of all is field observation and wide reading on biological and allied subjects.” Taverner also said that he would be very glad to see his bird count records, or any other results or ques- tions that he cared to send him. “If I can be of any further help, call on me.” *8 In thanking Henrik for this letter Taverner wrote: “T received your last letter with the account of your meeting with the Pileated Woodpecker, the sketch and the year’s list of birds. They are all very interesting and if you continue to develop along these lines there should be a considerable future for you in wild life or natural history subjects. What you have accomplished apparent- ly without special help is most encouraging but that is all the more reason for supplying all the guidance possible and even demanding more of you than from others less gifted. Therefore you must take it as meant if I seem to be severely critical. I will first take up your list. It is a common and almost universal error for the unex- perienced amateur to see strange, unusual and rare birds. The mere fact that the amateur is restrained from collect- ing his suspects* supplies no correction to vizional [word coined by Taverner] mistakes to teach caution in identification or restrain[s] misplaced enthusiasm. That you have fallen into this error is quite understandable but must be guarded against. The greatest qualification of a naturalist is his reputation for caution and reliability, that he does not jump at conclusions without unmistakable evidence and understates rather than overstates his observations. In the first place I would say that while occasional birds may turn up in almost any unexpected place, they are extremely rare, and their occurrence can be admitted only on the most perfect evidence which usually means the production of specimens. Anyone who sees or reports any number of freak occurrences is ipso facto suspect.” Taverner continued by mentioning a list of birds near Henrik’s vicinity in George Boardman, “Catalogue of Birds found in the vicinity of Calais, Maine, and about the Islands at the mouth of the Bay of Fundy”.*? It was also reprinted with annotations privately under the title of “The Naturalist of Saint Croix”. Taverner advised Henrik to look for this in the Saint John library, and to study it carefully to see what was likely to be seen and what was improbable in his locality. Taverner then went into detail concerning species that Henrik had claimed to have seen: “Gyrfalcon. The seeing of a single gyrfalcon in your ‘neighbourhood would be sensation enough granting that identification were well established. It would not be greatly improbable to see a single and not much out of the way should you see one || times.’ ... Ring-necked Pheasant. I suppose your 123 birds repre- sent individuals repeatedly recorded. I am surprised at *birds that an amateur suspects to be of a certain species but is not certain. 1996 CRANMER-BYNG: A LIFE WITH BIRDS 195 Taverner in retirement with his favourite dog Sinbad, taken in 1944 or 1945, surrounded with many framed pictures of birds. This is the house which he designed in 1912 and where he died in 1947. (National Archives of Canada, Accession number 1984-1978 Hayes Lloyd collection.) the Hungarian [gray] Partridge and would like to know how successful its introduction seems to be.” Ten other species were mentioned briefly. It was surprising to see the [Northern] Cardinal in this loca- tion, he wrote, though “it has certainly been extending its northwest range elsewhere.” The Common and Red-breasted Mergansers, Taverner said, were difficult to separate in life, except for adult males. “Our records of them based upon eye identifications of females are in the utmost confusion.” “Don’t be disheartened at this list of criticism’. he wrote, “It is only to be expect- ed in a beginner and it will give you an objective this summer in finding out what you really did see.” By the way of praise Taverner wrote about Henrik’s experi- ence with the woodpeckers: “It shows a good use of language and a feeling for words. It is too good a start not to be cultivated. Keep it up. I wonder who is your teacher.” In closing he urged Henrik to get in touch with Mr. Squires of the Saint John Museum, [though by a slip of the mind Taverner typed Mr. Spiers instead of Squires], and added — “I am sure he would be glad to see and help you, and personal contacts in chosen fields are always inspiring.” This was Taverner’ s last letter in their correspondence.*! Henrik was very excited and happy to receive Taverner’s letter: 196 “Tt is the best letter I have ever had. I am going to be stricter with myself and work harder, as I want to be a good naturalist. I have read your letter many times over. I think I almost know it by heart.” He then explained a little more about his home life. “As we live away from town-amusements my two sisters and I spend a lot of our time writing stories and illustrat- ing them. ... In about three weeks the river will open up so that the ferry can run again.” Then he planned to go to town on his bicycle and spend Saturdays at the museum. “T think Mr. Squires will let me look at the books and specimens in there.” He ended his letter with a perceptive statement. “Your letter has made me think of many things. Thank you again for it.”©? Deichmann’s last letter to Taverner, enclosing his spring arrival list, was dated May 2nd 1947. With this letter the correspondence between Percy Taverner and Henrik Deichmann ended. Will Saunders had died in June 1943. Now, in January 1946, another of Taverner’s long time ornithological partners, Allan Brooks, died. Taverner wrote to his widow, Marjorie, with sympathy from Martha and himself. Marjorie replied with a warm letter in which she wrote, among other things: “You and he were such close friends and your names being together in that Canada wide almost family Bible — The Birds of Canada, is a wonderful memorial & we are so proud it is so.” Early in March Percy composed a two-page letter to one of his closest friends, Mack Laing. It was a good natured letter from one naturalist to another, both of whom had shared many interests for over a quarter of a century. “I wish I had your free flowing pen also a littlhe more gumption with correspon- dence” was how he started his letter. He explained that in his present existence things flowed on so uneventfully that he had little to say. The weather seemed to be the one unfailing topic. It had been a comparatively mild winter up until recently, he said, and then three feet of snow dropped on them in a two-day storm. Everything was bogged down and even after a week of clearing they were only partial- ly dug out. The entrance to their front door he described as “a snowy canyon”. while the piles of snow were so high that they could not see a car entering or leaving on their garage driveway. Percy then gave Mack a few snippets of news that they could share. The first was about Erling Porsild of whom he wrote: THE CANADIAN FIELD-NATURALIST Vol. 110 “This winter we took in Porsild the botanist who could find no other accomodation in this crowded city and has been a very pleasant house mate. He takes care of a lot of oe little household jobs that I am unable to attend to.” The next was about someone who had known Percy’s mother and Percy’s early years. This is what he said: “T have had a long letter from Mrs. Sharman of Oak Lake. Of course I never met her personally but she seems to be the only one I know of who knows anything of my early days and forbears. She must be a remarkable personage, at her age to retain such interests and clear faculties. Her writings and letters are those of a young woman. Strange too how different lifelines cross. Not only did she know my mother as a young woman but also is closely connected with other people who fortu- itously had a great influence on my life.””°® Later Percy wrote a little about the Museum, when he said that it was “stumbling along in its same uncertain way.” They had some hope that the new Deputy Minister, Hugh Keenlyside might introduce a more enlightened policy. “As it is, we are threatened with loosing Rand to Chicago or the British Museum. Rand after five years here and highly competent has only a minor classifica- tion and is yet labeled ‘temporary’ .””° Finally Taverner, the author who never gave up, made a last report on his Practical Manual when he wrote: “There are some prospects of my big opus my Practical Manual of the Water Birds etc. seeing print. The Museum may handle it, if not the Arctic Institute is also interested. But things are still hazy.”°° Prosaically Percy mentioned in the final paragraph that Munro and Cowan [Ian MacTaggart ] were in Ottawa the previous week attending the Provincial game conference.” Percy signed the letter, as usual to friends, P.A.T. Under his initials Mack Laing later wrote the one word Finis 7° By early May spring had come, early flowers were Opening, trees were putting on their first tender leaves, and early warblers were arriving. As Percy lay in bed his mind began to wander. When Martha came into the room he motioned her to come over to him. He hummed part of a melody which was in his mind but which he was unable to finish. Martha hummed the full melody to him. Soon afterwards, with the melody in his mind, his eyes closed and Percy Taverner died.’! (1996 CRANMER-BYNG: A LIFE WITH BIRDS Selected Bibliography: A complete bibliography was compiled by W. L. McAtee. 1949. Ornithological publications of Percy Algernon Taverner 1895-1945. Annual Reports of the National Museum of Canada 1939-1947. National Museum of Canada Bulletin 112: 102-113. It includes 299 items, encompasing comments and book reviews as well as research papers. _ Taverner P.A. ‘| i] Me 1904 1905 1906 1907 1911 1914 1915 A Discussion of the origin of migration. Auk 21: 322-333. The tagging of birds. Bulletin of the Michigan Ornithological Club 5 (2): 50-51. Suggests alu- minum bands be issued by a central record-keeping office. A hyperlaken migration route. Bulletin Michigan Ornithological Club 6: 3—7. Along north shores of Lakes Huron and Superior. The origin of Kirtland’s Warbler. Ontario Natural Science Bulletin (Guelph) 1:13-17. Ornithology a science. Wilson Bulletin 17: 1234. Tagging migrants. Auk 23: 232. First tag supplied by Taverner returned from a distance — value of method. The Yellow-breasted Chat. Bird-Lore 8: 131-33. Illustrated with original wash drawing of Song Flight. The Yellow-breasted Chat in Michigan. Wilson Bulletin 18: 17-21. A tagged flicker. Wilson Bulletin 18 :21—2. First return of a band from some distance. Remarks on the summer birds of Lake Muskoka, Ontario, with B. H. Swales. Wilson Bulletin 18: 60-8. ’ Trinominials. Ontario Natural Science Bulletin 2: 16-17. How far to go in splitting is the question. The new check-list. Condor 9: 55—6. Suggests pro- viding names for specific groups and the subordina- tion of subspecies. Identifications. Ontario Natural Science Bulletin 3: 23-5. Necessary to collect specimens in order to correct identifications. The birds of Point Pelee, with B. H. Swales. Wilson Bulletin 19 1907: 37-54, 82-99, 133-153; 20 1908: 79-96, 107-129, 1 map. Some raptorial migrations in southern Ontario. Ottawa Naturalist 25: 77-81. Point Pelee records for four species. Notes on the migration of the Saw-whet Owl. Auk 28: 329-334. A new subspecies of Dendrapagus (Dendrapagus obscurus flemingi) from southern Yukon Territory. Auk 31: 385-8. Geological Survey Museum work on Point Pelee, Ontario. Ottawa Naturalist 28: 97-105. Object of field work was to collect material for a Canadian Carolinian group showing characteristic birds and mammals in a Carolinian landscape. Recommendations for the creation of three new national parks in Canada. Commission of Conservation (Canada) 6th Annual Report, Appendix 3: 303-310, 2 plates. The notable bird resorts: Point Pelee, Ontario; Percé Rock and Bonaventure Island, Quebec. The Double-crested Cormorant (Phalacrocorax auritus) and its relation to salmon industries on the Gulf of St. Lawrence. Canada Geological Survey, Museum Bulletin 13, 24 pages 1 plate, 1 table. 1918 1919 1920 1921 1926 1927 1928 1930 197 Suggestions for ornithological work in Canada. Ottawa Naturalist 29: 14-18 and 21-28. The hawks of the Canadian Prairie Provinces in their relation to agriculture. Canada Geological Survey, Museum Bulletin 28, Biological Series 7, 18 pages 4 colour plates, 7 figures. Trinomials and current practice. Condor 20: 213-16. The Gannets of Bonaventure Island. Ottawa Naturalist 32: 21-6, 3 photos. The birds of Shoal Lake, Manitoba. Ottawa Naturalist 32: 137-144, 1 photo; 157-164, 1918, and Canadian Field-Naturalist 33: 12—20, 1919. The birds of Red Deer River, Alberta. Auk 36: 1-21, 4 plates; 248-265, | map. The summer birds of Hazelton, B.C. Condor 21: 80-6, | photo. Bird-houses and their occupants. Ottawa Naturalist 32: 119-126, 3 plates. Republished by the Parks Branch of the Department of the Interior in 1920 and other years. Birds of Eastern Canada. Memoir 104, Geological Survey of Canada, Ottawa. 297 pages, 50 colour plates, 68 figures. Vanished and vanishing Birds. Dominion Parks Branch, Migratory Bird Leaflet 5, 8 pages. A plea for conservation. The Scoters and Eiders. Canadian Field-Naturalist 34: 41-4, 3 plates. Shows diagnostic characters. The Evening Grosbeak in Canada. Canadian Field- Naturalist 35: 41-5. Scientific advice for wild life conservationists. Canadian Field-Naturalist 40: 105-6. Birds of Western Canada. National Museum of Canada, Bulletin Number 41, Biological Series No. 10 Ottawa, 380 pages 84 colour plates, 315 figures. Hungarian Partridge vs. Sharp-tailed Grouse. Canadian Field-Naturalist 41: 147-149. Food com- petitor. A study of Buteo borealis, the Red-tailed Hawk and its varieties in Canada. National Museum of Canada Bulletin Number 48, Biological Series 13, 21 pages, 3 colour plates, 1 map. Cory’s Least Bittern. Auk 45: 204—5. Judges it a good species. Ornithological investigations near Belvedere, Alberta, 1926. National Museum of Canada 1926 (1928):84-104. A study of the Canadian Races of the Rock Ptarmigan (Lagopus rupestris). National Museum of Canada, Annual Report 1928:28-38, 1 map, 11 tables. Bird notes from the Canadian Labrador, 1928. Canadian Field-Naturalist 43: 74-9. Some zoological aspects of the Canadian Arctic Expedition of 1929. Canadian Field-Naturalist 44: 25-7, 1 plan. Some Canadian Birds. Canadian Geographical Journal 1(1): 31-2, 17 colour plates. Notes on a selection of four-coloured plates by Brooks in Birds of Eastern and Western Canada. 198 1931 1932 1933 1934 1935 1936 1937 THE CANADIAN FIELD-NATURALIST A study of Branta canadensis (Linnaeus) the Canada Goose. National Museum of Canada Bulletin Number 67 1931: 28-40, | plate, 5 tables. Based on breeding or summering specimens. Review of check-list of North American birds, 4th edition 1931. Canadian Field-Naturalist 46: 23-5. A partial study of the Canadian Savannah Sparrows with description of Passerculus sandwichensis campestris, new subspecies of the Prairie Savannah Sparrow. Proceeding Biological Society Washington 45: 201-205. William Spreadborough, Collector, 1856-1931. Canadian Field-Naturalist 47:39-41, one plate (por- trait). A study of Kumlien’s Gull (Larus kumlieni Brewster). Canadian Field-Naturalist 47: 88—90. The birds of Churchill, Manitoba, with G. M. Sutton. Annals of the CarnegieMuseum 23: 1-83, 13 plates, 1 map. Birds of the Eastern Arctic in “Canada’s Eastern Arctic”. Department of the Interior, Ottawa, pages 113-128, 6 photos, | map. Birds of Canada. National Museum of Canada Bulletin Number 72, Biological Series 19 Ottawa 445 pages, 87 colour plates, 488 figures. Continental land masses and their effect upon bird life. Condor 37: 160-162, 2 maps. Lesser size of winter range a limiting factor. Variability in size of gulls. Condor 37: 215-216. Probably due to food supply. Taxonomic comments on Red-tailed Hawks. Condor 36: 66—71, | map. Discussion and key. The fundamentals of the duck situation, 1936. Proceedings North American Wildlife Conference Senate Committee Print, 74th Congress, 2nd Session Washington D.C. pages 523-525. Less shooting is necessary. Bird populations, past and present, a personal expe- rience. Bird-Lore 39: 289-292. They have decreased — speculation on causes. Birds vs. Poison Sprays. Auk 54: 200. 1938 1939 1940 1941 1942 1944 Vol. 110 Birds of Canada, octavo, 445 pages, 173 coloured plates, 488 black-and-white illustrations, Musson Book Company, Toronto. Photographic copy of original edition. Adventures in bird photography. Canadian Geographical Journal 16: 264-283, 24 photos with explanatory notes. Canadian water birds, game birds, birds of prey. A pocket field guide. Musson Book Company, Toronto, 293 pages nearly 100 coloured plates, many drawings. Small octavo. Canadian land birds. A pocket field guide. Musson Book Company, Toronto, 279 pages 113 coloured plates, many drawings. Small octavo. The Red-winged Blackbirds of the Canadian Prairie Provinces. Condor 41: 244-246, 3 graphs. Suggests eliminating subspecies Agelaius phoeniceus arc- tolegus. The role of the predator. Canadian Field-Naturalist 53: 88-90. It has a necessary role in the working of nature. Predators — human and wild. Bird-Lore 42: 49, 3 plates. Wild predators play a sanitary role. Canadian status of the Long-tailed Chickadee. Auk 57: 536-541, 2 figures. Variation in the American Goshawk, including description of Astur atricapillus laingi, new sub- species. Condor 42: 157-160. The nesting of Ross’s Goose Chen rossi. Canadian Field-Naturalist 54: 127-130. James Henry Fleming, 1872-1940. Canadian Field- Naturalist 55: 63-4 1 plate (portrait). The distribution and migration of the Hudsonian Curlew. Wilson Bulletin 54: 3-11, 1 photo, 1 map. Canadian races of the Great Horned Owls. Auk 59: 234-245. Memories of William Edwin Saunders, 1861-1943. Auk 61: 345-351, Plate 3 (portrait). 1996 CRANMER-BYNG: A LIFE WITH BIRDS 199 Appendix 1: # LAT-65 35 su 1929. + CAMP KUNGOVIK WEsT Coast BAFFIN J6iAND * J Dewey SOFER ¢ DEPARTMENT OF THE INTERIOR CANADA NORTH WEST TERRITORIES AND YUKON OTTAWA JUL 1 8 1929 pe Le ae fe wi Gis 022 pe Zn ee 200 THE CANADIAN FIELD-NATURALIST Vol. 110 DEPARTMENT OF THE INTERIOR CANADA NORTH WEST TERRITORIES AND YUKON OTTAWA a an LP 1.7 = 7 ef Se . / nm < oss L —” << roo r) ot fy Ue te Cao 9 Petar. Stee: 27 ete Che ets OTL ge eae oe ail | | 1996 Appendix 2: CRANMER-BYNG: A LIFE WITH BIRDS 201 Source: National Archives of Canada. MG 30 B40 Vol. 15 File I. Also: Department of the Interior, North West Territories and Yukon, file 6652. Minutes of the meeting held to discuss the matter of the final disposition of the Baffin Island Natural History Collection. A meeting was held in the Department of the Interior, Ottawa, on 20 November 1929, to discuss the final disposition of the Baffin Island Natural History Collection. Those present were W. W. Cory, Deputy Minister, Department of the Interior; O. S. Finnie, Director, N.W.T. and Yukon Branch; J. Harkin, Commissioner of Dominion Parks; C. Camsell, Deputy Minister of Mines; P. A. Taverner, Ornithologist; J. D. Soper. The collection consisted of mammals and birds with their nests totalling 512 catalogue entries. Twenty-two bird skins were of the “Blue Goose”. The collection also contained eleven valuable sets of eggs of the “Blue Goose”. During the meeting the following recommendations were made: (1) That the entire collection of Baffin Island specimens shall go to the National Museum of Canada to be unpacked and stored. (2) That the National Museum shall receive one set of Blue Goose eggs, and that one set each be pre- sented to the British Museum, and the National Museum of the United States. It was further recom- mended that the department of the Interior retain one set of Blue Goose eggs, with a further suggestion that sets be laid aside for the Banff and Jasper Museums and for a possible future museum at Fort Churchill. (3) That the National Museum of Canada be pre- sented with the remaining specimens. In assuming ownership of this material, the National Museum will undertake to prepare, in a suitable manner, all specimens which are now in an unfinished condi- tion, and make them available for study as soon as possible. (4) That such specimens and sets of eggs present- ed to the National Museum of Canada, as may be regarded by it as duplicate, or surplus material, may be disposed of in exchange with other museums for desirable study material not now contained in the National Museum or is inadequately represented in the National collection. (5) That Mr. Soper and Mr. Taverner be entrusted with the unpacking and storage of the Baffin Island collection within the National Museum of Canada. (6) That no material in the Baffin Island collec- tion be disposed of by the National Museum until Mr. Soper has pursued the study of it to his satisfac- tion, in relation to the preparation of scientific papers dealing with the fauna of Baffin Island. (7) That the National Museum be provided with a copy of Mr. Soper’s field catalogue covering the nat- ural history specimens from Baffin Island, to be used in checking in the specimens, and as an aid to proper cataloguing of this material in the national register devoted to such acquisitions. The final paragraph was about arrangements for recovering cases from Camp Kungovik, where they were stored, sometime during the next year (1930), and having them taken by Eskimos with dog teams to Cape Dorset for loading on S. S$. Nascope. 202 Appendix 3: End Note 97 of Chapter 14 refers. THE CANADIAN FIELD-NATURALIST Vol. 110 From: R. M. Anderson To: Dr. Charles Camsell Memorandum Subject: Status of Division of Biology, National Museum of Canada This memorandum consists of 17 typed foolscap pages of 23 numbered items each with an underlined heading, and one page of a Summary of Con- clusions. Here follow some titles selected from this long and rather repetitive memorandum. Anderson’s main points were clear enough: (9) Recent flagrant disregard of rules, orders, and customs ‘Further than that, I stated that [am myself a qual- ified ornithologist, working on birds professionally, and am now working on an important ornithological report, and wish pertinent specimens to be available, and to know where they are.” (10) Present lack of discipline is becoming a pub- lic scandal “The bird collection of the National Museum for a considerable time has been used more for the gratifi- cation of personal vanity and as a factor in A.O.U. (American Ornithologists’ Union) politics, than for the advancement of ornithological science in Canada.” (12) Should the museum keep up policy of Divisional responsibility? “The chief apparent reason for intrigue in the Division of Biology, and indirectly affecting other sections of the National Museum, has been to get a separate Division of Birds established, a purely one- man affair, as Mr. Taverner has always worked strenuously, directly or in an underhanded manner, to keep anybody from getting on the Museum staff who has any knowledge or interest in scientific ornithology.” (14) Qualifications of ornithologists of National Museum These paragraphs attempt to show that Taverner’s education was so elementary that “much of Mr. Taverner’s ornithological publications would hardly have been worthy of the Department if he had not had a qualified scientific ornithologist back of him.” (page 6) Continuing, Anderson showed that Taverner’s training in science did not even reach junior matriculation as understood in Canada. He also had a disability in speaking in public (page 7). (15) Ornithological qualifications of the Chief of Division 2 paragraphs of his c.v. in the field of ornithology. “... there seems to be no just grounds for supersed- ing his directional capacity in ornithology at the National Museum of Canada.” (16) Inconsistency shown in present agitation for changes “ _.. largely through the efforts of the Chief of the Division the Department of the Interior allowed the specimens from Mr. Soper’s last two Baffin Island expeditions to be “deposited” in the National Museum of Canada for safety until Mr. Soper could return and work up the reports. Mr. Taverner, as a supposed friend of Mr. Soper received some of his data and all the specimens for examination in a priv- ileged capacity, skimmed the cream from Soper’s investigations and published them under the name of Taverner without informing Soper about the fact, although Soper was working in Ottawa at the same time. Mr. Taverner also began to peddle the Soper collections away without consulting either Mr. Soper or the Chief of the Division. Soper’s two reports have not yet been published.” (page 10) [But see Appendix 2 which shows the official arrange- ments made for the disposition of the Baffin Island collection. Soper and Taverner were present at this discussion and agreed to the recommendations made. | (19) Exchanges, gifts or loans of scientific speci- mens should be checked by a reviewing officer as with other financial transactions involving Govern- ment property. (20) Active endeavours to disrupt the National Museum organization “Mr. Taverner has for at least twenty years been one of the most persistent critics and trouble-makers on the museum staff. Every chief, director, acting director, or other official above him has been consis- tently knocked, plotted against, and undermined as far as his limited influence goes. The administration of the National Museum has been receiving undesir- able publicity from one end of North America to the other, as I know very well from personal knowledge and reliable communications. Mr. Taverner has never had enough scientific training to carry on 1996 advanced work, and with not enough to keep him busy, has had plenty of time for intrigue.” (21) Lack of control of publications by members of | the staff “Recently in looking over certain technical publi- | cations by members of the staff of the Division of | Biology, I noted that two major articles were pub- | lished during 1935 as ornithological contributions, and that both of them were rank plagiarisms.” | Anderson gave their titles as A. “Continental Land Masses and their Effects upon Bird Life” by P. A. | Taverner in The Condor 37, Number 3: 160-162 (1935); (page 13). B. “Variability in size of gulls” by _ P. A. Taverner in The Condor 37 Number 4: 215-216 (1935). The whole of his paper, Anderson said, was cribbed from a lengthy dissertation by the late ' Bernard Hantzsch entitled “Beitrag zur Kenntnis ger | Vogelwelt des nordostlichsten Labradors’, Journal fur CRANMER-BYNG: A LIFE WITH BIRDS 203 Ornithologie, Leipzig, Volume 58, 1909, page 324.” Anderson explained that “Our departmental ornitholo- gist would not have lifted it directly from the original German text, but it happened that M. B. A. and R. M. Anderson had taken the trouble to translate and anno- tate the whole paper, the same being published in seri- al form in The Canadian Field-Naturalist, January 1928 to March 1929.” The section from which Anderson accused Taverner of cribbing appeared in translation in this journal in May 1929, Volume 42 under the section about the Glaucous Gull. At the end of the Summary of Conclusions Anderson signed the memorandum: Respectfully submitted R. M. Anderson Chief of Division of Biology National Museum of Canada 204 THE CANADIAN FIELD-NATURALIST End Notes Abbreviations AOU American Ornithologists’ Union BOU British Ornithologists’ Union CMN Canadian Museum of Nature GSC Geological Survey of Canada MTRL Metropolitan Toronto Reference Library NAC National Archives of Canada NMC National Museum of Canada PABC Provincial Archives of British Columbia ROM Royal Ontario Museum pter 1 Cha Ip 6. 10. Birth Certificate of Percy Algernon Fowler. Registration No: 1875-05-022716. Date of birth: June 10, 1875. Place of birth: Guelph. Name of father: Edwin Fowler. Name of mother: Emily E. Buckley. Place of birth of father and mother: not recorded. Father’s occupation: Principal. Mother’s occupation: not recorded. The age of each parent was not recorded. . Information from baptismal certificate preserved at St. George’s Anglican Church, Guelph. Date of bap- tism: 7 July 1875. His mother’s name was given as Emily Ellen Buckley, his father’s occupation as school teacher, and their place of residence as Guelph. . P.A. Taverner, “Biographical Outline”, compiled 1942. Canadian Museum of Nature (CMN) Percy Taverner Ornithology Archive, hereafter CMN. W.L. McAtee “Percy Algernon Taverner, 1875- 1947” The Auk 65 (1948): 85-106. McAtee obtained family information from Taverner’s own “Biographical Outline” and from reminiscences sent him by Taverner’s half-sister, Mrs. John McLeish. I have not been able to trace this letter, or letters. It is possible that any information about Taverner’s par- ents and first years was destroyed by McAtee since the Taverners remained silent about Percy’s father. . Carolyn Cox Toronto Saturday Night 4 November 1944. Madge Macbeth, in column entitled “Over My Shoulder”, Ottawa Citizen 12 July 1958. This was the first of four articles. She was born on 22 February 1854 at Prestwick, Lancashire. Her mother was Mary Heyward Buckley, nee Hind. In the Chicago directories of the period 1870-1874 several other Buckleys were shown who were born in England, but none called George. An index of death notices in all ten English-language Chicago newspapers, 1837-1889, failed to show any George Buckley. Information from a Chicago directory of 1871, which contained a partial listing of the U.S. Census of 1870, gives no other given name for Edwin Fowler. Hugh Douglass “An Account of Some of the Private Schools of Guelph, 1827-1900”. Unpublished ms.1960, Public Library, Guelph. The author was president of the Guelph Historical Society in 1960. Letter from Registrar’s Office, University of Oxford, 9 July 1984. City of Guelph Assessment Rolls, 1874. Also Guelph Land Registry Office, 1874, plans 175/6 lot no. | book 7. Vol. 110 12. A new marriage act came into effect on 1 July 1874. Instead of a marriage licence, which required the publishing of the banns of marriage, a certificate could be obtained by one of the parties swearing an affidavit that there was no legal cause to prevent the solemnization of the marriage. Since Emily Buckley was under twenty-one years of age she had to swear an affidavit that her father and mother were dead and that there was no one with authority to given consent to the marriage. Province of Ontario, C.6, An Act Respecting the Solemnization of Marriages, 1874. The Guelph Daily Herald published an official notice about the new marriage certificates issued by authori- ty at the Guelph Divisional Office. It stated that no bondsmen were required. Educational advertisement printed in the County of Wellington Atlas, published 1877, as quoted in Douglass “An Account’, page 10. Greata Mary Shutt The High Schools of Guelph (Guelph: The Board of Education for the City of Guelph 1961), page 92. There is a photograph of the house used by Fowler’s Academy on page 93. Douglass “An Account” page 10. Murray D. Edwards A Stage in Our Past (University of Toronto Press, Toronto, 1968). Taverner to H.M. Laing, 8 March 1947. Provincial Archives of British Columbia (hereafter PABC) Laing Papers. Fanny A. Sharman to H.M. Laing, 2 July 1947. PABC, Laing Papers. Chapter 2 1. A stock company was a resident company the mem- bers of which would put on a variety of plays during the season, often with well known actors and actress- es coming to play the star roles. Emily was a member of the “ballet” which referred to the “extra” men and women members of the company who played walk- on parts as required. Ida Van Cortland, an address given to the University Women’s Club of Ottawa about 1918. Taverner Collection, Arts Department, Metropolitan Reference Library, Toronto (hereafter MRLT). The advantage of training with a stock company was the variety of parts played and broad range of experience gained. With a touring company a smaller repertoire of plays was presented and so members of the company played fewer parts. Based on Kathleen Fraser “Theatrical Touring in Late Nineteenth-Century Canada: Ida Van Cortland and the Tavernier Company 1877-1896”. Unpublished Ph.D. Thesis (University of Western Ontario 1985). The Company alternated between Halifax and St. John’s, Newfoundland. His family lived in Toronto and Hamilton in the 1870’s. Young Albert was encouraged to get a degree in civil engineering, but he preferred to spend time on amateur college productions. For an explanation of why the son spelt the family name as Tavernier, see note 8 below. Notice in the Dramatic Mirror, 25 June 1881. “Albert Tavernier, of the Madison Square Theatre Company, 1996 17. 18. and Ida Van Cortland, of W. H. Powers’ Galley Slave Company were united in marriage by the Rev. Henry Wosse, at his residence ... on the evening of June 11.” Kathleen Fraser, personal communication, 24 May 1984, quoting Dramatic Mirror of 25 June 1881. . W.J. Florence was the founder of the Order of the Shrine. Hence The Mighty Dollar was connected with the Masonic Order. Fraser, Thesis, 21. See note 2 above. Same 23. McAtee “Taverner” 94. For professional reasons Albert always spelt his name with an i, but the rest of the family spelt it without. Percy later dropped the 7 in his name. Writing in 1940 to another Taverner in Alberta he explained that the name Taverner was only his by adoption and that he had no genetic claim to it. On the question of how the name should be pro- nounced he wrote, “Our family tradition is “Ta-vern- er.” PAT to Jessie Taverner, 16 April 1940. Taverner Collection, Theatre Department, Metropolitan Toronto Library. P. A. Taverner “Biographical Outline” 1, see Chapter 1 note 3; McAtee “Taverner” 86-87; 94. See Chapter 1 note 4. Reproduced in a newspaper article. Madge Macbeth the Ottawa Citizen 9 August 1958 in column called “Over My Shoulder”. Unfortunately it has not been possible to find the original photograph from which this was made. At that time it had probably belonged to his sister Mrs. Ida McLeish. For a detailed account of the Taverniers’ Company see Fraser chapters 2 and 3. Itinerary for Maritime Provinces 1883-84 in Fraser pages 186-192; Ontario and Great Lakes 1884-1985 in Fraser pages 195-200. . Fraser, Thesis, page 105-106. . McAtee page 94. . Same reference. 16. Purchased from the Crown by a grant of 19 December 1877. This stated that Joseph William Taverner, of the City of Toronto, in the County of York, Professor of Elocution, purchased for sixty-one dollars a tract of land of fifty-six acres composed of Gibraltar Island, in Lake Muskoka. Document in Land Registration Office, Bracebridge, Ontario. This was a deed of sale between Henry W. Taverner, gentleman, of San Francisco; Joseph E. Taverner, piano tuner, of Mount Vernon, N.Y.; Mary Ann Taverner, widow, of San Francisco; Isabel Power; Frances Graham; Theresa Lindeman; Clara Preston and Florence Knill. The last five were married women whose husbands signed the document with them. Presumably these five were daughters of Joseph William Taverner, while Mary Ann was a widowed daughter-in-law. The five hundred dollars would be divided among these eight people. See note 50 below for Fraser’s analysis of the account books kept by the Tavernier Company. Albert and Ida started a guest-book (also referred to as a log-book) for the cottage at Point Coo-ee on Gibraltar Island. This was their first entry. Percy had just turned 11 years old. 19. Ontario Birth Certificate no. 1887-05-040130. She was named Ida Clara. Clara was the name of Albert’s sister in San Francisco, but the family spelt it Clare. CRANMER-BYNG: A LIFE WITH BIRDS 20 26. 28. 29! 30. Chile Nn Her mother’s name was given as Ida Fowler. Places of birth of mother and father were not recorded. Albert Tavernier’s occupation was shown as “sales- man” and not actor. A prejudice against the acting profession still existed at that time. The Knill family was well known in the area around Markham and Stouffville in the nineteenth century. Information supplied by the Markham Museum. Fraser, Thesis, chapter 3. Fraser, Thesis, pages 108-109. Percy had suffered from a stammer in his speech from as early in his life as he could remember. It must have been a hindrance to him because as soon as he came to Detroit to live in 1904 he took a course at a school for stammerers though without much suc- cess. His stammer remained with him all his life. P. A. Taverner to J. H. Fleming, 7 December 1904. Royal Ontario Museum Archives, Percy Taverner Papers, hereafter ROM. Information obtained from Anna Clinton History of Ann Arbor Public Schools (Ann Arbor 1951); Lela Duff Pioneer School. Some Chapters in the Story of Ann Arbor High School (Ann Arbor 1958). On the back is written “Family photo — 1889 — Percy 14 — Ida 2 — Gibson — Ann Arbor”. Presumably this was written by Ida Van Cortland. There is another version of this group with Ida looking away from the camera, but with Percy smiling a little. Taverner Collection, Theatre Department, Metropolitan Toronto Library. There is a very clear black-and-white photo of the school taken by R.M. Scadin at this period and reproduced in Art Work of Washtenaw County (Chicago 1893). Copies of the Annual Catalogue and the Annual Year Book (Omega) covering this period are preserved in Michigan Historical Collections, Bentley Historical Library, University of Michigan. Ann Arbor High School, Catalogue 1891 page 11. Ridicule by direct imitation of his stammering would be the usual way, but also by a sharply spoken ques- tion such as “What’s your name boy?” “What does your father do?” When the unfortunate boy tried to stammer the answer it would provoke a burst of laughter. If he remained silent or failed to say any- thing coherent he might get a slap on the cheek with the back of a hand. From seeing this happen at junior school. Taverner “Biographical Outline” 1, CMN. Robert Henry Wolcott studied at the University of Michigan where he received a B.S. degree in1892, and an M.D. degree in 1893. He followed a career in biology and in summers of 1893 and 1894 was hired to make a survey of fish in Michigan waters. In the following year he joined the Department of Zoology at the University of Nebraska where he continued his grad- uate studies. Obituary in The Auk 52: 130-131, 1935. Taverner “Ornithological Notes” books 1-4 (May 1892—November 1903) 4. ROM. Same, 5-6. There was already a contradiction show- ing in Percy’s notes between his scientific desire to learn to identify each species accurately by collecting specimens for study “in the hand”, and his artistic appreciation of their plumage and movements. This contradiction was never fully resolved. 206 33. 34. 35) . It is possible that the school reassembled late that fall Sif 38. 30% 40. 41. 42. 43. 44, THE CANADIAN FIELD-NATURALIST In using Taverner’s notes the bird names he used at that time have been retained in order to show what he called a particular bird then. To aid the reader the names of species given in the current American Ornithologists’ Union Check-list (AOU) have been added. At this time Percy was still in the process of learning the check-list names in place of some ver- nacular names he had grown up with. He still had not seen one by 1906 when he heard from Norman A. Wood, taxidermist at the Museum at Ann Arbor, that a Sycamore Warbler, the colloquial name for the Yellow-throated Warbler, had been col- lected. Taverner to N. A. Wood, 29 May 1906. ROM. Fraser, Thesis, page 108. on account of an exceptionally late harvest, and would then make up the last weeks at the end of the school year. Most children in that period were required to help with the harvest, and the summer holiday might continue late into the fall. Taverner “Ornithological Notes” 25 April 1894, ROM. For an account of the building of the Royal Opera House, Guelph, and its management under Albert Tavernier see Wayne Fulks “The Royal Opera House”, Guelph Historical Society Publication 22: 49-69 (1983). A shorter version of Fulks’ article was published in Theatre History in Canada 4: 41-56 (1983). Taverner “Ornithological Notes”, Guelph 3 May 1895, ROM. Log-book of cottage at Point Coo-ee, Gibraltar Island, 1895. I am grateful to Mr. David J. McLeish, who possesses the log-book, for letting me read it . Note 18 above refers to the origin of the log-book. Same 1895. Carolyn Cox “Name in the News” Saturday Night, Toronto 4 November 1944 wrote: “Before graduating from High School Taverner went back to Guelph where his stepfather had taken on the management of the New Opera House...” This implies that he later returned to Ann Arbor in order to graduate. This would have been possible in the first half of 1895, but from June 1895 onwards he was at Beaumaris and after that on tour with the Company. A disastrous fire at Ann Arbor High School on 31 December 1904 destroyed all the student records. Nor does any copy of the student year book for 1893-1894 exist in Ann Arbor libraries — the year that Percy should have graduated. There is no sign of his name in the student Year Book for 1894-1895. Therefore the precise meaning of the words “before graduating from High School” are uncertain. Fraser, Thesis, pages 272-274. Taverner “Ornithological Notes” Winnipeg 21 and 24 October, 1895, ROM. The Greater Prairie-Chicken was reported to have been fairly common in southern Manitoba in the 1890s. W. Earl Godfrey The Birds of Canada (Ottawa 1986) page 163. Taxidermists were more likely to call game birds by hunters’ names than by ornithologists’ names at that time, and it could have been a Sharp-tailed Grouse. However, they were useful people for Percy to visit whenever the train drew into town. Because taxidermists had specimens of locally killed birds in their shops local hunters congregated there as well, and exchanged bird news. 45. 46. 47. 48. 49. 50. Sill 52) a3: Vol. 110 Taverner “Ornithological Notes”, Huron, South Dakota. 13 December 1895, ROM. Taverner “Ornithological Notes” Preface, page 2 note added under date 3 February 1896, ROM. Taverner “Ornithological Notes”, made at Ann Arbor 8 February 1896, ROM. Other ornithologists and wildlife artists have found fault with Audubon’s paintings for being stiff, unrealistic, disproportionate, and such reasons. But it is important to remember that he was painting for patrons who would pay well for large-scale representations of birds they could never hope to see. Audubon had to work under great difficulties in the field collecting specimens from which to paint. Taverner “Ornithological Notes”, Guelph 6 May and 4 June 1896, ROM. Fraser 277, quoting from Dramatic Mirror 8 August 1896. From Kathleen Fraser’s analysis of the account books kept by the Tavernier Company it is clear that what money the Company made by touring was lost in Albert Tavernier’s two ventures as an actor-manager in Jackson and Guelph. The Taverniers’ weak finan- cial position in 1895, when leasing the Royal Opera House, Guelph, was clearly losing money, may have prevented Percy from entering the University of Michigan, even if he had the necessary qualifications. The accounts also show numerous loans from Ida to Albert without any record of being repaid. Fraser page 110. It may also have led to the document signed by Albert Tavernier deeding Gibraltar Island to Ida Van Cortland in 1900. See Fraser “Theatrical Touring”, page 110. Fraser, same, page 10 note 14. These were the property of a Mr. Lucas, comprising about thirty skins among which Percy mentioned: Moa; Kea; Maori Hen; Huia. He also mentioned that Lucas owned “a fine illustrated work by Sir W.L. Buller which describes their habits very minutely and graficaly” [graphically]. Taverner also made pen sketches of the difference between the bills of the male and female Huia. Taverner was twenty-one years old and Mcllwraith was seventy-two. In his “Ornithological Notes” under that date he wrote that MclIlwraith mentioned a bird he called a Bay-winged Bunting and another he called the Bay-winged Sparrow. These, Taverner said, were local names for the Dickcissel and the Vesper Sparrow. Chapter 3 I. i) This was probably James Grand who bought an island near to Gibraltar subsequently called Grandview. Joyce I. Schell The Years Gone By. A History of Walker’s Point and Barlochan — Muskoka 1870-1970 (Bracebridge 1970) page 65. . James Henry Fleming, 1872-1940, only son of James Fleming of Aberdeen, who came to Canada from Scotland and established a pioneer seed-growing business on a three-acre plot at Yonge and Elm Streets. J. H. Fleming was educated in Toronto at the Model School and at Upper Canada College. His interest in building a study collection of birds began in 1886 when he was fourteen years old, ten years before he met Taverner. There is no biography of ; : ' ; 1996 Nn oon 10. Wake IW 13% 14. Fleming, but a useful short source exists in The J. H. Fleming Memorial Program, November 5, 1940 The Brodie Club (Toronto 1940) 29 pages typescript. ROM, Brodie Club Papers. Also L. L. Snyder “In Memoriam: James Henry Fleming” The Auk 58: 1-12 (1941). . Taverner “The Old Taxidermy Shop and Point Pelee Days” in The J. H. Fleming Memorial Program, ROM, Brodie Club Papers, November 1940. . Taverner to L. L. Snyder, 4 October 1940, CMN. On Lambe see obituary Globe and Mail Toronto, 5 December 1941; J.L. Baillie to W.L. McAtee, 8 October 1947. University of Toronto, Fisher Rare Book Library, Baillie Collection. . Taverner to Snyder, 4 October 1940, CMN. . T.M. Shortt “The Fleming Collection” in The Fleming Memorial Program, ROM. For example Fleming had two specimens of the Large-billed Finch (Geospiza magnirostris) taken on Abingdon Island, in the Galapagos in 1897. Charles Darwin had taken the type specimen from nearby Charles Island. Shortt 7, . Taverner “Old Taxidermy Shop ...” page 27. . A complicated story to unravel. Taverner wrote to W.F. Webb, editor of The Museum, announcing that he had taken a Wheatear at Beaumaris on 25 September 1894 and that it was identified as such by R. Ridgway. This was published in The Museum 2: 16 (1895). Covert wrote to Taverner that he had sent it to Ridgway who identified it, and that he had deposited it with the Smithsonian. Covert then wrote to R. H. Wolcott at the University of Nebraska that he had taken a Wheatear near Ann Arbor on 4 October, and that an account of this would be appearing short- ly. A.B. Covert “The Wheatear in Michigan” The Nidiologist 2: 42-43 (1894). For a review of the status of the Wheatear in Ontario see Jon C. Barlow “Status of the Wood Ibis, the Fulvous Tree Duck and the Wheatear in Ontario” The Canadian Field-Naturalist 80: 185 (1966). . Taverner, letters in The Gravenhurst Banner 28 October and 10 November 1896. Taverner “Old Taxidermy Shop ...” page 26. Taverner’s sister, writing to McAtee after his death, mentioned his sailing on Lake Muskoka. “He was wonderfully adept at small boat handling, could liter- ally sail anything. When in his early ’teens, he con- verted a little 14-foot row-boat of his into a sail-boat, decked it over, put in a centre board ... and went out in it in any gale.” McAtee Auk 65: 97 (1948). Taverner to [Sir] Peter Scott, 22 September 1938. CMN. Taverner “Ornithological Notes” 7 December 1896; also 23 May 1898. Percy told it to his wife Martha, who told it to a long- time friend, Mrs. Jean York, when she was a widow living in Ottawa. Mrs. York to author in a tape recorded interview, Ottawa, 16 September 1984. Taverner received instruction in taking photographs from W.A. Anderson who had a cottage in Beaumaris. The only photo of Percy’s to survive from this period was pasted on the outside of the notebook and shows a girl of about sixteen, wearing a short sleeved summer dress with cotton stockings. She is seated on a rock beside a lake with her feet in CRANMER-ByYNG: A LIFE WITH BIRDS 15. 16. ie . Ernest Seton-Thompson to Taverner, 18 March 1900. INS), 207 the water. She is looking directly at the camera, and is close enough to show a slight smile. ROM Taverner Papers. His mother’s address was given in the log-book of their cottage in 1899 as Port Huron. There is reason- able proof that Percy was in Port Huron at least by March 1899 because at the meeting of the Michigan Ornithological Club in mid-April “Mr. Percy Tavernier, Port Huron, Michigan” was elected to active membership of the club. Since he arrived in Toronto at least by mid-December 1898 it seems very probable that he joined his mother and sister in Port Huron for Christmas. There were three architects in Port Huron at the turn of the century, the most important being John C. Kaumeier who had designed many of the principal buildings since 1875. George L. Harvey was not well known. His office was at 201 Huron Avenue, an easy walk from where Percy boarded. At this time the pop- ulation of Port Huron was around 20 000, while Sarnia, across the river, was about 7000 and growing. McAtee, The Auk 65: 95, 1948). ROM. Clipping from American Ornithology sent to Taverner by A. G. Lawrence of Winnipeg. Lawrence to Taverner, 30 October 1942, CMN. . John Macoun Catalogue of Canadian Birds (Ottawa 1900-1904). Published in three parts. Part. I Water Birds etc. (1900); Part II Birds of Prey etc. (1903); Part III Sparrows, Warblers etc. (1904). 21. John Macoun to Taverner, 15 July 1900. ROM. The 30. Wilson’s Petrel was recorded by Taverner in his “Ornithological Notes” 18 December 1897. It belonged to Dr. Cornell of Gravenhurst and was found dead or killed at Gull Lake, Muskoka “a few years ago”, ROM. . William Dutcher to Taverner, | December 1902, ROM. Dutcher was treasurer of the AOU at that time. Fleming was responsible for nominating him. . U.S. census returns St. Clair County, Port Huron Ward, St. Clair Street, number 513. . McAtee, The Auk 65: 95. (1948). . Ida Van Cortland “Address given to the Ottawa Drama League” about 1918, page 10. Arts Department, MTRL, Taverner Collection. . Taverner “Biographical Outline” page 2, CMN. . Letter Taverner to Fleming, 9 April 1904 on notepa- per with printed heading, Independent Order of Foresters, ROM. . There were four “Courts” of the Independent Order of Foresters listed in the St. Clair County Directory, 1899-1900. It is possible that Ida Van Cortland start- ed working for the Foresters while living in Port Huron at some time between 1897 and 1902. In the agreement drawn up between Albert and Ida Tavernier dated February 1900, in which he released to her and her heirs all his rights to the Island of Gibraltar, his address was given as Detroit, and hers as Port Huron. . Letter addressed to Taverner at Port Huron dated August 1901, ROM. Taverner to Fleming, 11 December 1902, in which he said that he was attending night classes in water-color drawing at the Chicago Institute of Art “this winter’, 208 Dil . Same, 18 February 1904, ROM. 3: THE CANADIAN FIELD-NATURALIST while last winter (1901-1902) it was sculpture, just for recreation, ROM. A notebook of 34 water-color sketches has survived from the period 1902-1905. The subjects are wild flowers, butterflies and insects. Property of Mrs. Marte Kent. I am grateful to Mrs. Kent for allowing a set of coloured reproductions to be made for presentation to the Taverner collection in the Archive of the ROM. . Taverner notebook “Systematic Bird List and Migration Notes” 1893-1900, ROM. . Taverner “Biographical Outline” 2, CMN. McAtee Auk 65: 96 (1948). Taverner “Ornithological Notes” 3 May 1903, ROM. Same 15 March, ROM. Same 12 April, ROM. . Same 3 May, page 140. Then follow 7 more pages of sustained writing on the last of the Passenger Pigeons, how and why they would never be able to breed again. It is a moving threnody on the disappear- ance of this once prolific species. ROM. . Same 3 May, page 147, ROM. Dr. J. A. Allen (1838-1921). Curator of Birds and Mammals at the American Museum of Natural History from 1885. A founder of the American Ornithologists’ Union in 1884 (incorporated 1888). Special concern for problems of nomenclature and updating the AOU’s Check-list of North American Birds. F. M. Chapman (1864-1945). Assistant to Allen at American Museum and Curator of Birds. Distinguished museum-builder and educator. Editor, Bird Lore, author Handbook of Birds of Eastern North America 1906 and later editions. C. H. Merriam (1855-1924). Chief Division Ornithology and Mammology United States Department of Agriculture 1886; chairman AOU committee on migration and geographical distribution. Dr. T. S. Palmer (1868-1955). Assistant Chief to Merriam 1896-1902. Worked for bird protection through legis- lation; collected information on wildlife conservation. Secretary AOU 1917-1937. Taverner “Ornithological Notes” 3 May 1903, 151, ROM. . Same 8 June, pages 163-168, ROM. . Same 27 June, ROM. . Information from Frank M. Chapman Autobiography of a Bird Lover (N.Y., 1933) page 173. . Fleming to Taverner, 26 May 1903, ROM. . Taverner “Ornithological Notes”, 24 October 1903, 183-190, ROM. . Taverner to Fleming, 8 November 1903. Taverner said he would like to put them into the form of a paper to be read at the AOU meeting being held at Philadelphia from 17-19 November 1903. ROM. . J. A. Allen to Taverner, | November 1903, ROM. . Fleming to Taverner, 20 November 1903, ROM. . T.S. Palmer to Taverner, 25 November 1903, ROM. J. A. Allen to Taverner, 20 December 1903, ROM. This was a frank letter which might have silenced a less persistent man than Taverner. Allen himself had put forward the same ideas as hypotheses ten years previously in The Auk 10: 48 (1893). Same, 10 January 1904, ROM. The Auk 21: 322-333 (1904). The article served to bring Taverner to the notice of the readers of The Vol. 110 Auk. He sent a copy of it to A. R. Wallace in England and received a post card back which read, in part: “I agree with you generally, but I do not believe in any instinct in the matter. My latest view is given in my Studies Scientific and Social volume 1, page. 502, ‘The Migration Instinct’. Wallace to Taverner, 31 August 1904. (This card was discovered in a copy of the 1898 edition of Wallace’s Malay Archipelago given to Erling Porsild after Taverner’s death. I am grateful to Porsild’s granddaughter, Jennifer Lumsden, for this information.) Professor William Rowan of the University of Alberta was carrying out experiments on bird migration in the 1920s which culminated in his important book The Riddle of Migration (1931). In a short overview entitled “Fifty Years of Bird Migration” he mentioned Taverner’s discussion in his article of the reason why birds leave what appears to be a satisfactory environment and take the risk of travelling north to breed. “His discus- sion is lucid and full, and present-day opinion would largely agree with him.” At the end of his article Rowan acknowledged that his own indebtedness to Taverner would be difficult to estimate. In F. M. Chapman and T.S. Palmer editor: Fifty Years’ Progress of American Ornithology 1883-1933. (Lancaster, Pennsylvania. Pages 51-63.) See also Taverner to A. Wetmore, 20 January 1932, CMN. 54. T.S. Palmer to Taverner, 25 November 1903, CMN. 55. Taverner “Journal of Bird Observations”, 28 November 1903, 15-18. ROM. 56. Taverner to Fleming, 8 November 1903, ROM. 57. Taverner to Fleming, 8 November 1903; Taverner to Fleming 9 April 1904, ROM. 58. Taverner to Fleming, 8 November 1903; 20 January 1904, ROM. 59. J. Dwight, Jr., to Taverner, 22 February 1904, ROM. 60. Taverner to Fleming, 20 January 1904, ROM. 61. N. A. Wood to Taverner, 24 January 1904, ROM. Wood became curator of birds at the museum in LOW: 62. See H.M. Mayer and R.C. Wade Chicago: Growth of a Metropolis (Chicago 1969). Chapter 4 1. Taverner Journal of Bird Observations, 6 April 1904, ROM. 2. N.A. Wood “Discovery of the Breeding Area of Kirt- land’s Warbler”; and C.C. Adams “The Migration Route of the Kirtland’s Warbler’, Bulletin Michigan Ornithological Club 5: 3-13; 14-21 (1904). Charles Adams was Curator, University of Michigan Museum. 3. Taverner to Fleming, 9 April 1904, ROM. 4. P. A. Taverner “A Discussion of the Origin of Migra- tion” The Auk 21: 322-333 (1904). 5. P.A. Taverner “The Origin of the Kirtland’s Warbler” Ontario Natural Science Bulletin (Guelph) 1: 13-17 (1905). For status in early 1980s see R.B. Payne A Distributional Checklist of the Birds of Michigan (Ann Arbor 1983). 6. An announcement by Barrows asking for information for inclusion in the Bulletin on the birds of Michigan which he was compiling was printed in the Bulletin Michigan Ornithological Club (March 1904) page 26. 1996 1S 16. 30. Bile a2. . Taverner Journal, 1 May 1904, ROM. . Taverner Journal, 30 May 1904, ROM. . Taverner Journal, 16 April 1904, ROM. . Same as 9. . Taverner to Fleming 7 December 1904, ROM. . Taverner to Fleming 26 June 1904, ROM. . Taverner to Fleming, 4 May 1905, ROM. . A copy of the program of the Saint-Saens Concert held in the Light Guard Armory, Detroit, on the evening of 12 December 1906 with Martha Hohly’s name on it has been preserved in the Music and Drama Department of the Detroit Public Library. In the Detroit City Directory for 1902 Martha Hohly is listed as “teacher of music, Michigan Conservatory of Music”. Ida C. Tavernier is listed as “music teach- er” for the first time in the 1907 Directory. Fleming to Taverner, 8 February 1906, ROM. The Ben Greet Players performed Twelfth Night in the Elizabethan manner at the Detroit Opera House on 31 May 1905. Report in the Burton Special Collection, Detroit Public Library. Taverner writing to Fleming 22 March 1907 mentions “great feast of Shakespearean production” they had been enjoying. “A succession of good things have come on top of each other and we felt that we could not miss them. Among them was Annie Russell in Midsummer Night’s Dream. This was the most satisfactory perfor- mance I ever saw given. The setting was good and the mechanical effects, lights etc. were almost perfect without once overshadowing the kernal of the work.” . Taverner to Fleming, 13 April 1909, ROM. . Karel Wiest, Taverner’s step-son, in personal com- munication at Detroit January 1984. . Taverner Journal, 2 October 1904, ROM. . Taverner Journal, 5 August 1905, ROM. . Taverner Journal, 27 April 1905, ROM. He only saw a Broad-winged Hawk and two Chimney Swifts but it was the thrill of getting out of doors again that count- ed, a feeling of the promise of birds to come. . Taverner to Fleming, 30 November 1905, ROM. . E. Coues Key to North American Birds (Salem 1903) Sth edition. . J. A. Brashear to Taverner, 27 October 1904, ROM. Eventually Swales gave him a good pair. . Fleming to Taverner, 20 June 1909, ROM. Percy accepted them gratefully. . Taverner to Fleming, 18 August 1906, ROM. . W.E. Clyde Todd to Taverner, 8 August 1907, ROM. . Taverner Journal, 27 April 1905, ROM. . Taverner to Fleming 20 December 1911; 24 February 1912, ROM. His sister wrote after his death that “in Detroit his room was lined with bird cases which he made himself. He had quite a large collection ...” W.L. McAtee The Auk 65: 96 (1948). Taverner to Fleming, 18 October 1905; 16 December 1905, ROM. The use of borax as a preservative came in later. The layperson who is interested to know a little more about the collecting of birds, making of skins, sexing, labeling, and care of a collection will find a short out- line of the subject in F.M. Chapman Handbook of Birds of Eastern North America, Second revised edi- tion (1931) New York. Dover Publications reprint 1966. J. H. Fleming “The Unusual Migration of Brunnich’s Murre (Uria lomvia) in Eastern North America” CRANMER-BYNG: A LIFE WITH BIRDS 33. 34. 39! 36. i. 38. 50. Se 5 209 Proceedings of the Fourth International Ornithological Congress pages 528-543 (1905). Taverner “A Hyperlaken Migration Route” Bulletin of the Michigan Ornithological Club 6: 3-7 (1905). Most of Taverner’s scanty records for Ontario came from the Toronto vicinity. Fleming to Taverner, 3 December 1904, ROM. On Nelson’s Sharp-tailed Sparrow see P. A. Taverner Birds of Western Canada National Museum of Canada, 1926, page 288. Taverner to Fleming, 7 December 1904, ROM. For a more detailed account of the GLOC see J. Cranmer-Byng “The Great Lakes Ornithological Club. The Origin and Early Years, 1905-1911” Ontario Birds 2: 4-12 (1984); J. Cranmer-Byng “The Bulletin of the Great Lakes Ornithological Club, 1905-1909” Ontario Birds 3: 45-54 (1985). On the landscape see J.G. Battin and J.G. Nelson Man’s Impact on Point Pelee National Park (National and Provincial Parks Association of Canada, Toronto 1978) chapters 1-2; 8, pages 90-92. On the Chestnut Oak see R. C. Hosie Native Trees of Canada, Canadian Forestry Service, Ottawa (1967) page 190. A note by Saunders stated: “At Point Pelee Aug. °82 saw perhaps 50 all told in flocks of up to 10, rushing up and down through the woods. Shot only one which I have yet.” Saunders Papers, December 1902, ROM. A fuller account given by Saunders is printed in Taverner and Swales “The Birds of Point Pelee” The Wilson Bulletin 19: 91 (1907). . Taverner Journal, 13 May 1905, ROM. . Same, page 109. Taverner wrote that they crossed to the east shore on this page but when he again wrote “east” shore on the next page he then crossed it out and wrote “west”, which is correct. . Taverner Journal, 14 May 1905, pages 110, ROM. . Same, page 113. . Taverner smoked “T & B” tobacco. Saunders, a non- smoker, enlivened the monotony of the dreary march to Leamington from time to time by the remark, “Come now, let’s have a smoke”. P. A. Taverner “Will Saunders — Field Naturalist” in R. J. Rutter, editor, W. E. Saunders Naturalist (Toronto 1949) page 13. . Raymond Brothers, Awnings, Tents, Flags, Sail Covers, London, Ontario, to P. A. Taverner 25 August 1905, ROM. . Taverner Journal, 4 September 1905. . Taverner Journal, 5 September 1905, ROM. Ontario September 9th they found “quite a flock of Cardinals. Indeed this bird seems to be quite common and well established on the Point.” . Taverner Journal, 6 September 1905, ROM. . Taverner Journal, 8 September 1905, ROM. . As quoted from Lynds Jones in P. A. Taverner and B.H. Swales “The Birds of Point Pelee” The Wilson Bulletin 19 38-39; 45-46; 48-49 (1907). Letter Lynds Jones to the members of the Great Lakes Ornithological Club 16 December 1905, pre- served in papers of the GLOC, ROM. Taverner to Saunders, 27 September 1905. GLOC papers, ROM. Taverner in (1904). “Notes and News” The Auk 21: 410 210 59. 60. 61. 62. 70. THE CANADIAN FIELD-NATURALIST . Fleming Journal, 24 September 1905. ROM, Fleming Papers. . Taverner “Tagging Migrants” The Auk 23: 232 (1906). . Taverner Journal of Bird Observations (September 1904-July 1905) listed on last page of the book, ROM. . L.J. Cole “The Tagging of Wild Birds as a Means of Studying their Movements” The Auk 26: 143 (1909). . Same 137-143. . W.E. Saunders The Auk 27: 221-212 (1910). It was in 1915 that Miner began the bizarre practice of fix- ing biblical texts to the bands of geese. Taverner to Fleming, 5 November 1905, ROM. Although by early 1906 Taverner had read a number of sources on taxonomy and nomenclature, and had read Coues carefully, his knowledge of taxonomy was elementary. His arguments were developed from what Fleming told him in their discussions. Fleming could have presented a better case to Allen but was careful not to involve himself too deeply in the argu- ment; he might need the support of Allen in the future on his own account. Taverner to J. A. Allen, 30 March 1906, ROM. J. A. Allen to Taverner, 13 June 1906, in reply to Taverner’s letter of 10 June, ROM. The law of priori- ty is that the first name properly published is the valid name for any taxonomic category such as family, genus, species. The starting point for the rule is | January 1758. Thus a zoologist may describe and name what he considers a “new species” only to find that it has been described and named in literature ear- lier. In that case the earlier name stands. A proposed statute of limitations would, if ever agreed on, limit the length of time that an original name in the litera- ture would remain operative. . Indenture dated 21 May 1906. Registered at Brace- bridge 25 May. Preserved in Land Registration Office, Bracebridge, Ontario. . Taverner Journal, 16-18 June 1906, ROM. . Signed copy of “Option to buy 55 Elmurst Avenue” dated 16 May 1906. ROM Taverner Papers (Sc 51 Box 3). . Fleming to Taverner, 13 September 1906, ROM. For statute of limitation see note 83. . Fleming to Taverner, 23 September 1906, ROM. . Taverner to Fleming, 27 September 1906, ROM. . These difficulties were forcefully explained by Macoun in a letter to C. H. Merriam, John Macoun to C.H. Merriam, 2 March 1906. Typed copy in Canadian Museum of Nature, Ornithology, Taverner Papers. Albert Peter Low (1861-1942). Geologist with the Geological Survey of Canada. Commanded expedi- tion to the Arctic 1903-1904. Director of Geological Survey 1906; Deputy Minister of Mines 1907. Retired 1913. . Fleming to Taverner, 6 October 1906, ROM. . Taverner to Fleming, 9 October 1906, ROM. . Fleming to Taverner, 14 October 1906. Fleming to Low, 13 and 17 October give more information on what had been in Fleming’s mind. ROM Fleming Papers. A. P. Low to Fleming, 15 October 1906. Fleming to Taverner 16 October 1906 enclosing Low’s letter. ROM Fleming Papers. 74. 16). 76. HE 78. 79. . The Wilson Bulletin 20: 107 (1908). A second 81. 82. 83. Vol. 110 | Taverner to Fleming, 31 October 1906. ROM GLOC | Collection (xerox copy of original only). | Fleming’s father had established a seed growing busi- _ ness on a three-acre plot at Yonge and Elm Streets, — Toronto, in 1836. Fleming had experience in horticul- — ture and this accounted for his interest in plants and | his success in growing them in his own garden and — green house. See L. L. Snyder “In Memoriam James | Henry Fleming” The Auk 58: 2-3 (1941). His office was in the Chamber of Commerce Building at the corner of State and Grisworld. There was a good view from there which Taverner made use of. In his Journal for 20 August 1906 he noted: “Today about 5 p.m. I was looking out of the window of our office in the Chamber of Commerce when I saw a flock of about fifteen Purple Martins passing to the southwest. They were flying straight along with the usual aerial evolutions and were evidently migrat- ing. See note 46. It was found in the cedars on the west shore near the end of the Point on 21 May 1906. Fleming “Chuck- will’s-widow and Mockingbird in Ontario” The Auk 23: 343-344 (1906). Taverner and Swales “The Birds of Point Pelee” The Wilson Bulletin 19: 135 (1907). The Wilson Bulletin 20: 86 (1908). [Northern] Mockingbird was taken in 1909 by Bert Gardner and sent to Saunders who reported the news to Taverner. Saunders considered that this second record from Point Pelee was significant, and that its status should be reconsidered. The Wilson Bulletin 19: 151-153 (1907). Saunders recorded one seen on 24 and 26 April 1909 in GLOC’s “Birds Observed at Point Pelee” ms. record book Volume 1, 55, ROM. Two Bewick’s Wrens were taken by Saunders at Pelee on 15 April 1917. W.E. Saunders “The Status of Bewick’s Wren in Ontario” Canadian Field-Naturalist 33: 118 (1919). Bert Gardiner (Anglicized Gardner) was of French Canadian descent. He knew Point Pelee intimately and had a good knowledge of its wildlife. Taverner obtained useful notes from him on breeding Wood Duck, and Snow Goose in November, as well as study specimens. In 1908 a hut with screened doors and windows was built to Taverner’s design on his property. Taverner Journal 16 October 1908, ROM. . Taverner to Fleming, 22 March 1907, ROM. . Taverner to Fleming, 12 October 1907. As from 1983 the name Sparrow Hawk was changed to American Kestrel in the AOU Check-list. . Taverner to Fleming, 22 March 1907. . Swales to Fleming, 7 May 1907, ROM. . This permit is preserved in Taverner’s papers at the ROM and is dated 9 March 1906. . Taverner to Fleming, 12 October 1907, ROM. . Saunders to Taverner, 8 November 1906, ROM. The birds were White-winged Crossbills. . Taverner Journal 23 March 1907, ROM. On 5 October he noted a covey of Bobwhites near his home in Highland Park. . Taverner Journal 21 November 1906. . Fleming to Taverner, 22 October 1907, ROM. 1996 «94. 107. 108. 109. 110. HUE 112. 3: 114. 113), Taverner, Journal 25 May 1906, ROM. Woods claimed to have seen White-rumped Sandpipers with- out having written a description of the plumage. But when Taverner himself saw a White-rumped Sandpiper on the same mud flats a few days later he retracted his suspicions. Taverner Journal 2 June 1906. . Swales to Fleming, 18 November 1906, ROM. . Taverner Journal 11 August 1906, ROM. . Taverner to Fleming 22 March 1907, ROM. . Taverner Journal 5 May 1906, ROM. . Taverner Journal 2 June 1906. . P. A. Taverner “The Yellow-breasted Chat. A Character Sketch” Bird Lore 8: 132 (1906). Black- and-white drawing of bird in song-flight included. A colour wash drawing of a male Yellow-breasted Chat by Taverner was printed in Walter Barrows Michigan Bird Life Michigan Agricultural College Lansing (1912). (On Barrows see note 134.) The framed origi- nal is preserved in the Bird Range, Museum of Zoology, University of Michigan, Ann Arbor. . Fleming to Taverner 26 May 1903, ROM. . F.M. Chapman The Warblers of North America. Coloured illustrations by L. A. Fuertes and B. Horsefall, 3rd edition (New York 1907). . Note by William Brodie in GLOC Bulletin of 12 July 1906. ROM, GLOC Papers. . Taverner Journal 24 August 1907, ROM. . Swales manuscript “Bird Journal” Museum of Zoology, University of Michigan, Ann Arbor (extracts in ROM, GLOC Papers). . Taverner to Fleming 12 October 1907, ROM. Taverner was thinking of how much information on migrating birds could be gained if a bird banding sta- tion could be established at Point Pelee. It could be as valuable for information on migration as the German island of Heligoland in the North Sea where a suc- cessful banding station had recently been established. Taverner had the vision to realize what could be done if the finance was available. A few years later at Kingsville, near Leamington, Jack Miner owned the land and had the opportunity to start a successful bird banding station. Taverner and Swales “Birds of Point Pelee” Wilson Bulletin 19: 38 (1907). Same 48. Taverner and Swales Wilson Bulletin 20: 93-94 (1908). Same 79-82. Fleming to Taverner, 2 January 1909, ROM. Taverner to Fleming, 11 January 1909, ROM. This is clearly illustrated by line drawings in W. E. Godfrey The Birds of Canada (Ottawa 1986) page 72-73. On the status of the Trumpeter Swan in North America see W.E. Banko The Trumpeter Swan (Washington, 1960); F.C. Bellrose Ducks Geese and Swans of North America (Harrisburg, 1976); in east- ern Canada H. G. Lumsden “Pre-settlement Breeding Distribution of Trumpeter, Cygnus buccinator, and Tundra Swans, C. columbianus, in Eastern Canada” The Canadian Field-Naturalist 98: 415-424 (1984). J.H. Fleming “The destruction of Whistling Swans (Olor columbianus) at Niagara Falls” The Auk 25: 306-309 (1908); Fleming “The Niagara Swan Trap” The Auk 29: 445-448 (1912). CRANMER-ByYNG: A LIFE WITH BIRDS 116. 117. 118. 119. 211 Taverner Journal 26 March 1908; Taverner “Notes on Swans” ROM. Catalogue number 35001; gift of J. H. Fleming. The label on its leg shows that it was collected at Lake St. Clair 1873. J. Grinnell “Better Vernacular Names” The Condor 8: 154 (1906). The Condor was a quarterly journal pub- lished by the Cooper Ornithological Society of California. Swales was a member of the club and knew Grinnell personally. P. A. Taverner “The New Check-List” The Condor 9: 55-56 (1907). 20. W. Stone to Taverner, 12 April 1907, ROM. 21. Two folio books called “Birds Observed at Point 128. 129. 130. SI Pelee”. Volume | ran from May 1905—September 1912. Volume 2 began in 1913, ran until 1923 and then very spasmodically until the last entry of December 1927. ROM, GLOC Papers. . These “Notes and Clippings” are of some value his- torically for records of numbers of species such as Piping Plover, Volume 3, pages 278-283; Eastern Bluebird, Volume 8, pages 765-766, ROM. . Taverner Journal 31 May 1909, ROM. . Taverner to Fleming 16 June 1909, ROM. . Taverner Journal, 8 September 1905, ROM. . Henry J. Richmond “The birds’ jumping-off place” The Detroit News Tribune, magazine section, 27 June 1909. . Arthur W. Andrews 1866-1950. Born Goderich, Ontario, lived in Detroit, Michigan from 1900 until his death. A fine furniture maker by profession, an entomologist by avocation. The Andrews collection of Michigan Coleoptera is preserved in the Museum of Zoology, University of Michigan. Obituary notice in Annual Report of the Museum of Zoology, University of Michigan, 1950. Taverner to Fleming, 16 June 1909, ROM. P. A. Taverner “Migrating Butterflies” Entomo- logical News (1908) pages 218-220. Taverner described how they came to the extreme tip of the Point at the rate of about three a minute on fine days and launched directly out over the lake taking a line that would carry them directly across the open lake. “At times when there was a little wind their proceed- ings were a little different. As soon as they reached the end of the heavier timber they kept well down in the shelter of the dense red cedar and juniper growth as far as that went, and then crossed to the lee shore, and dropping down to near the water’s edge, proceed- ed along in the cover of the sand dune until the first breath of wind was encountered when they gradually rose in the air and started over the lake on the usual course. The exactitude with which they followed each other was remarkable. One could stand between two red cedars where they crossed to the shore and ninety per cent of all butterflies would pass within striking distance of a net.” An explanation of the why and how of the monarchs’ migration is set out in a pam- phlet published by Parks Canada, available at the Visitor Centre at Point Pelee National Park. Taverner met it as a problem to be mulled over; to us it is a phenomenon to be marvelled at. Taverner to Fleming, 13 April 1909, ROM. Taverner Journal, 14-16 October 1910, ROM. THE CANADIAN FIELD-NATURALIST . P. A. Taverner and B.H. Swales “Notes on the Migration of the Saw-whet Owl” The Auk 28: 329- 334 (1911). 133. Taverner to Fleming, 20 January 1910, ROM. 134. Walter B. Barrows Michigan Bird Life, Michigan Agricultural College Special Bulletin, 1912. Barrows was professor of zoology and physiology at the Agricultural College from 1894 until his death in 1923. 135. Taverner to Fleming, 20 January 1910, ROM. 136. Taverner to Fleming, 6 February 1910, ROM. 137. J.M. Macoun to Taverner, 2 February 1910, NAC, Geological Survey, Taverner Papers. R.G. 132, Volume 34, file 488. 138. Taverner to J.M. Macoun, 6 February 1910. NAC, Geological Survey, Taverner Papers. R.G. 132, Volume 34, file 488. 139. Taverner Journal 22 February 1909, ROM. Compare letter Taverner to Fleming, 27 February 1909, ROM. The episode was retold by Taverner in his contribu- tion to the W. E. Saunders Memorial Night organized by the Brodie Club of Toronto, March 28 1944. Printed in R.J. Rutter, editor, W. E. Saunders, Naturalist, Federation of Ontario Naturalists, Toronto, 1949, page 17. Also in Taverner’s “Geological Survey Museum Work at Point Pelee, Ont.” The Ottawa Naturalist 28: 89-99 (1914). 140. Taverner to Swales, 22 February 1919, CMN. 141. Contains 33 typed pages. Present cover and handwrit- ten title supplied by Archive staff, ROM. 142. Taverner to Fleming, 17 May 1910, ROM. A more detailed account of the expedition is contained in his Journal for 15 May 1910. On Arthur W. Andrews see note 127. Dr. William W. Newcomb was a Detroit attorney and amateur lepidopterist. 143. Taverner to Fleming, 16 June 1909, ROM. 144. Taverner Journal, 28 April 1907, ROM. 145. Taverner Journal, 17 May 1908. 146. Taverner to Fleming, 7 December 1904, ROM. 147. Taverner to Fleming, 7 February 1909, ROM. 148. Taverner to Fleming, 26 July 1907, ROM. 149. Taverner to Fleming, 20 January 1910. 150. Fleming to Taverner, 7 October 1910, ROM. 151. Taverner to Fleming, 22 October 1910, ROM. 152. Taverner to Fleming, 19 April 1908, ROM. It also showed that his hair was beginning to recede. 153. Taverner to Fleming, 31 October 1906, ROM. 154. Fleming to Taverner, 9 March 1907, ROM. 155. Taverner to Fleming, 22 March 1907, ROM. 156. T.S. Palmer to Taverner. Whereabouts of letter not known. 157. Taverner to T.S. Palmer. Also not known. Chapter 5 1. The Act of 1907 is reproduced in W. H. Collins “The National Museum of Canada” in Annual Report for 1926, pages 60-70. Canada Department of Mines; National Museum of Canada. Ottawa, King’s Printer, 1928: and discussed in Zaslow, Reading the Rocks, The Story of the Geological Survey of Canada 1842- 1972. Published by the Macmillan Co. of Canada Ltd. Toronto in association with the Department of Energy, Mines and Resources and Information Canada, Ottawa, pages 257-258. 2 Zaslow, page 259. Vol. 110 William Andrew Waiser, The Field Naturalist: John Macoun, the Geological Survey and Natural Science. Toronto University Press, 1989. Waiser, The Field Naturalist 178-179. For a strongly critical view of the Catalogue see letter Harrison Lewis to Taverner, 15 November 1919 and Taverner’s reply 19 November 1919, CMN. Waiser, Field Naturalist 183. Fleming to Taverner, 23 March 1910, ROM. James Macoun to Fleming, 24 March 1910, ROM. The nephew was Stuart Logan Thompson 1885-1960. James Macoun to Fleming, 24 March 1910, ROM. Fleming to Taverner, 29 March 1910, ROM. . See Chapter 7, note 101. . Fleming to Taverner, 29 March 1910, ROM. . Taverner to Fleming, 30 March 1910, ROM. Taverner’s letter to James Macoun of 6 February 1910 is in the Public Archives of Canada, Geological Survey, Taverner Papers, RG 132, Volume 34, file 488. . Fleming to James Macoun, 31 March 1910, ROM. . James Macoun to Fleming, 4 April 1910, ROM. . Taverner to Fleming, 30 March 1910, ROM. . W.E. Saunders to John Macoun, 14 April 1910 PAC, Geological Survey, R. G. 132, Volume 33, file 473. . James Macoun to W.E. Saunders, 18 April 1910 PAC, Geological Survey, R.G. 132, Volume 33, file 473. Saunders sent a copy to Fleming. ROM, Fleming Collection. . Fleming to Taverner, 6 April 1910, ROM. . Seton to Taverner, 15 November 1910, ROM. . Taverner to Fleming, 21 November 1910, ROM. . Taverner to Fleming, 21 November 1910, ROM. 22. R.W. Brock to Taverner, 25 November 1910, ROM. The idea of exhibiting birds and mammals in life-like positions in their natural habitats had recently taken root in some of the museums in the United States. Brock would have seen such displays in the summer of 1910 when he visited several museums in the United States to see for himself how to run a leading museum. . Taverner to Fleming, 28 November 1910, ROM. . Fleming to Taverner, 24 November 1910, ROM. . Fleming to Taverner, 30 November 1910, ROM. . Fleming to Taverner, night lettergram, | December 1910, ROM. . Fleming to Taverner, 2 December 1910, ROM. . Fleming to Taverner, 2 December 1910, ROM. Saunders’ letter to Taverner on the subject was dated 30 November 1910. . Fleming to Taverner, 2 December 1910, ROM. . Taverner to Fleming, 7 December 1910, ROM. He also said he was sending to Brock proofs from the printing plates of the drawings for Barrow’s forth- coming book, and some separates of his articles, adding “Perhaps it is best to strike while the iron is hot.” . Fleming to Taverner, 2 December 1910, ROM. . Fleming to Taverner, 18 December 1910, ROM. . Fleming to R.W. Brock, 8 December 1910, CMN. . R.W. Brock to Taverner, 24 December 1910, ROM. . Fleming to Taverner, 28 December 1910, ROM. . R.W. Brock to Taverner, telegram 22 March 1911, ROM. . R.W. Brock to Taverner, letter 22 March 1911, ROM. 1996 45. 46. 47. 48. . R.W. Brock to Taverner, 29 March 1911, ROM. . Fleming to Taverner, 3 April 1911, ROM. . Taverner, writing near the end of his life to a friend from his Ann Arbor days, described his heart trouble as “an old mitral murmur”. Taverner to George Prey, 28 June 1946, CMN. . R.W. Brock to Taverner, 3 April 1911, ROM. . Taverner to James Macoun, 12 April 1911. James Macoun to Taverner, 18 April 1911, ROM. . Samuel Herring, taxidermist at the Biological Survey since 1884, retired 1912, died 1919. . Taverner to Fleming, 1 May 1911, ROM. His mother had been unwell and faced the possibility of having an operation. It is possible that by this time she had resigned from her job, and would not be earning. In his anxiety to leave her sufficient money until she and Ida could join him in Ottawa he had left himself too little, but was reluctant to ask her to send him money so soon. Taverner to Fleming, 10 May 1911 (handwritten), ROM. Taverner to Fleming, 13 May 1911 (handwritten), ROM. Taverner to Fleming, 18 May 1911, ROM. This could have been true. There had been outbreaks of typhoid in past years, and fresh outbreaks occurred in subse- quent years. Taverner to Fleming, 4 June 1911, ROM. Chapter 6 i, On The name of the museum was officially changed to the “National Museum of Canada” on Sth January, 1927. The name was officially changed again to the “Canadian Museum of Nature” on Ist July 1990. . One exception was the Redpath Museum in Montreal. Founded in 1882 in a beautiful building built as a gift to McGill University, it contained several collections including one of bird skins. Taverner first heard about the museum through Dr. Casey Wood. See Chapter 13 note 11 for more detail. . For the early development of bird banding in North America see F.C. Lincoln “Bird banding” in Fifty Years’ Progress of American Ornithology 1883-1933 (Lancaster, Pa. 1933) pages 65-87 (hereafter Fifty Years’ Progress). Published by the American Ornithologists’ Union and edited anonymously by F. M. Chapman and T. S. Palmer. . See chapter 4 , pages 30-31, note 63. . U.S. National Museum Bulletin, 2 volumes 1892- 1995. Charles Emil Bendire was a retired army offi- cer who had gathered much information on birds dur- ing his long military service in remote parts of the west. For development in the study of life histories see Herbert Friedmann “Advances in Life History Work” in Fifty Years’ Progress (see note 3 above). . The snipe was printed in the Osprey; the owl in the Auk. On early bird photography Alfred O. Gross “History and Progress of Bird Photography in America” in Fifty Years’ Progress. (See note 3) . For regions specified by Taverner as requiring studies of bird distribution see Taverner “Suggestions for Ornithological work in Canada” discussed in Chapter 7, pages 2-3, notes 2-13. . Author of Extermination of the American Bison (1887); American Natural History (1904) and other influential writings. CRANMER-BYNG: A LIFE WITH BIRDS 9. 10. Ill 19. 20. 2A Pips 23. 24. 7o\k3} See also Janet Foster Working for Wildlife: the Beginning of Preservation in Canada (Toronto 1978) which gives information on the Canadian background to the decline in numbers of wildlife at this time. See Chapter 4, note 34. The first official bird sanctuary in North America was created in what was then the Northwest Territories (present day Saskatchewan) northwest of present day Regina. This was at Last Mountain Lake where some islands and shoreline at the north end of the lake were set apart as a breeding ground for waterfowl. After the passing of the Migratory Birds Convention Act in 1917 the reserve was officially named Last Mountain Lake Bird Sanctuary. . Taverner to Fleming, Wednesday ... (no date given) but Fleming’s annotation on the letter shows that it was written in May 1911, ROM . Taverner to Fleming, 9 May 1911, ROM. . Taverner to Fleming, 10 May 1911, ROM. . Report Taverner to Brock, 12 May 1911, pages 1-2, CMN. . Taverner to Brock page 8. . Canada, Sessional Paper 1911, “Summary Report of the Geological Survey of Canada for 1910” [hereafter Summary Report]. . Report Taverner to Brock, 12 May 1911 pages 2-3, CMN. Marianne G. Ainley in her study From Natural History to Avian Biology: Canadian Ornithology 1860-1950 argues that Taverner made a decision in 1911 that he was to regret later. But neither Taverner nor Brock had the authority to make decisions about major questions of policy such as the scope of the ornithological collection. There was considerable infighting over funding while the museum was being organized and Brock had to be very careful what moves he made in relation to the Geological Survey. Members of parliament, who had the power of grant- ing appropriations for the museum, had considerable appreciation of mining but the level of their apprecia- tion of natural science was still unknown. (Unpublished Ph.D. thesis McGill University 1985) page 117. Taverner to Brock, page 4. (See note 18) Taverner to Brock, page 9. (See note 18) At the end of his report Taverner suggested that it would be well worth while to send him to see what was being done in other museums in order to avoid making mistakes. Brock himself had visited several museums in the United States the previous summer and supported Taverner’s suggestion. Fleming to Taverner, 6 June 1911, ROM. Taverner to Fleming, 10 June 1911, ROM. Taverner to Fleming, 10 June 1911, ROM. For more detail on the trip to Mer Bleue on 6 June see Taverner, in a typewritten note of 6 June 1911 ina file entitled “Notes on Plants, Animals, Insects etc. 1907-1911” (ROM Taverner Papers). Also note on another walk at same place -15 June 1911. Charles H. Young, an entomologist with the Experimental Farms Service, joined the Natural History Branch of the museum in 1907 and worked under the Macouns’ supervision collecting and preparing invertebrate material. Taverner to Fleming, 18 May 1911, ROM. George Rivers White (1856-1927) lived in Ottawa on the banks of the Rideau River from where he collected 29: 26. 27. 28. 29: 30. 31. 39. 40. Al. THE CANADIAN FIELD-NATURALIST many bird specimens and eggs. See In memoriam notice in The Canadian Field Naturalist 43: 103-104 (1929). Taverner to Fleming, 13 May 1911, ROM. Taverner to Fleming, 4 June 1911, ROM. Taverner to Fleming, 19 June 1911, ROM. Taverner to Fleming, 29 June 1911, ROM. Taverner to Fleming, 29 June 1911, ROM. The Brooklyn Museum is part of Brooklyn Institute of Arts and Sciences, well known for its collections of Egyptian and oriental art. The Children’s Museum (1899) contains a natural history collection. Taverner to Fleming, 30 June 1911, ROM. Charles W. Richmond followed Ridgway as curator of birds at the U.S. National Museum and was responsible for the thorough card catalogue of names in the bird room. Taverner to Fleming, 30 June 1911, ROM. Wells W. Cooke, member U.S. Department of Agriculture, Biological Survey, Washington from 1901. Amassed migration and distribution data with help of a net- work of observers. Taverner supplied him with infor- mation on water birds of the Great Lakes region from 1905. . Taverner to Fleming, 25 July 1911, ROM. . Report Taverner to Brock, 27 July 1911, page 1, CMN. . Taverner to Brock, page 4. (See note 33) . Taverner to Fleming, 16 August 1911, ROM. Frank C. Hennessey b. 1894. Accompanied Captain J. E. Bernier, Canadian navigator and explorer, on an exploration trip to the Arctic, as naturalist and artist. Wintered at Melville Island 1908-09. See Colin S. Macdonald, compiler, 1968 A Dictionary of Can- adian Artists 2 volumes (Ottawa) pages 427-429. The arranging of bird specimens by years according to dates of arrival at the museum resulted in a Victoria Memorial Museum Register of Birds. Specimen num- ber 1 was a Ruff taken at Toronto in May 1877 — it was listed as received in December 1911. . Canada Sessional Papers 1912 “Summary Report for 1911”, page 374. . Taverner to Fleming, 16 August 1911, ROM. . After Taverner’s retirement the distributional maps were not kept up by his successor A. L. Rand, but when Earl Godfrey followed Rand as zoologist in 1947 he brought the system up-to-date. Fleming to Taverner, 21 August 1911, ROM. Fleming to Taverner, 2 November 1911, ROM. James A. Munro. Born Manitoba 1884. As a boy col- lected birds eggs and made birds skins in the Toronto area. Spent much time with Dr. William Brodie, biol- ogist at the Ontario Provincial Museum. Made a study of migration along the Toronto waterfront which was published in the Ottawa Naturalist in 1911. Moved to Okanagan Landing, B.C. in that year where he started collecting birds, eggs and mammals for sale commercially. In 1915 the British Columbia Provincial Museum hired him to collect for them in the Okanagan Valley. For his later career as Dominion Wildlife Officer see James L. Baillie “In Memoriam: James Alexander Munro” The Auk 86: 624-630 (1969). Fleming to Taverner, 20 December 1911, ROM. Fleming wrote to Brock with the same suggestion. apy, 56. ail 58. 59. 60. 61. 62. 63. 64. Vol. 110 Fleming to Brock, 20 December 1911, CMN. The museum bought six Trumpeter Swans from Munro for $15.00. . Taverner to Fleming, 24 February 1912, ROM. On material collected earlier by G. M. Dawson and R. Bell and others see Chapter 14, note 4. . Taverner to Fleming, 6 March 1912, ROM. . Fleming to Taverner, 10 March 1912, ROM. Sheds light on Fleming’s ideas and interests in zoology vis- a-vis the ROM. . Taverner to Fleming, 24 February 1912, ROM. . Taverner to Fleming, 24 February 1912, ROM. . Taverner to Brock, 2 February 1912, CMN. . Taverner to Brock, 13 February 1912, CMN. . Taverner to J. M. Macoun, 20 May 1912, CMN. . Taverner to J. M. Macoun, 30 July 1912, CMN. . Taverner to Brooks, 12 April 1912, CMN. . Taverner to Fleming, 6 March 1912. Also Taverner writing to Arthur Andrews, mentioned that members of the Geological Survey, “who have been all over the Dominion from the mouth of the Mackenzie to Nova Scotia, when they can be got to talk can tell enough experiences of pioneer life to make Jack London green with envy.” Taverner to Andrews, 29 April 1912, CMN. . P. A. Taverner Jnstructions Regarding the Collection of Zoological Specimens for the Victoria Memorial Museum (Ottawa, Government Printing Bureau 1912). . Canada, Sessional Papers 1913, “Summary Report” for 1912, page 441. Taverner to Swales, 26 December 1912, CMN. Clyde Louis Patch (1887-1952). Grew up in Ohio and Michigan, apprenticed to a commercial taxidermist in Washington, D.C. Employed as taxidermist by the Biological Survey, U.S. Department of Agriculture, and by the American Museum of Natural History, New York. See obituary by Hoyes Lloyd “Clyde Louis Patch” The Canadian Field Naturalist 68: 124- 126 (1954). Taverner to Fleming, 5 December 1912, ROM. Canada, Sessional Papers 1913, “Summary Report for 1912”, page 443. On what to read for an impression of Ottawa at this time: John H. Taylor Ottawa: An Illustrated History (Ottawa, 1986) Taverner to Fleming, 10 May 1911, ROM. He con- fessed: “A week and a half of boarding house life is all I want and I want a home here as soon as I am per- manently settled.” Taverner’s stammer was probably worse under the stress of his new job and new acquaintances. It would have been frustrating not to be able to speak fluently to his fellow lodgers and he would have felt at a disadvantage. Taverner to Fleming, 4 June 1911, ROM. Taverner to Fleming, 16 August 1911, ROM. Fleming to Taverner, 2] August 1911, ROM. Taverner to Fleming, 21 September 1911, ROM. . Development had recently been started in an area bounded by the Rideau Canal to the north, the Rideau River to the south, Bank Street to the east and Bronson Avenue to the west. This area became known as Ottawa South, and the street where the Taverners built was named Leonard Avenue. The Ottawa Street Directory 1911 shows which houses were built and which lots were vacant at that time. 1996 BW: 80. 81. 82. 83. . Taverner to Fleming, 17 October 1911, ROM. . Taverner to Fleming, 2 November 1911, ROM. . Taverner to Fleming, 3 April 1912, ROM. . Taverner to Arthur Andrews, 29 April 1912, CMN. . Taverner to Fleming, 6 August 1912, ROM. Outbreaks of typhoid occurred in Ottawa from time to time and a cholera epidemic occurred in 1917. The drinking water was the suspected source. Dr. Charles Camac of the College of Physicians and Surgeons, New York City, spoke to the Committee on Public Health of the Conservation Council on the subject of “The Epidemics of Typhoid Fever in the City of Ottawa” in 1912. . Taverner to Swales, 23 October 1912, CMN. . Taverner to Mrs Hinton, CMN. . Taverner to Fleming, 11 February 1913, ROM. . Taverner to Brock, 19 February 1913, CMN. She remained for thirty years. . Taverner to Brock, 19 February 1913, CMN. . Taverner to Brock, 20 January 1913, CMN. . Taverner to Brock, 6 February 1913, CMN. Not a sensible idea. . Taverner to James Harkin, 19 March 1913, CMN. . Taverner to Brock, 17 March 1913, CMN. The exchanging of specimens between one museum and another, or with a private collector, although a routine matter, is one that sometimes causes trouble. The decision as to what should be exchanged in return for what specimen(s) is a personal judgement, and may cause personal animosity between museum staff. For example correspondence with Jim Macoun while Taverner was collecting at Point Pelee between 15 May and 18 July 1913. In one letter Jim Macoun asked Taverner to send him specimens of a rare species of violet from the Leamington area (15 May 1913). Taverner did send some and they were planted in the Experimental Farm. J. M. Macoun to Taverner, 26 May 1913. PAC RG 132 Volume 34 file 488, and CMN. Zaslow Reading the Rocks pages 319-320. (See chap- ter 5 note 1.) Taverner to Fleming, 27 February 1913, ROM. Taverner had not yet met Anderson and was not aware of what was happening at a higher level. Brock discussed with Taverner the idea of obtaining an appointment for Anderson at the museum on 28 February. Taverner’s letter to Fleming is interesting in the light of later relations between the two men and the large part which Anderson was to play in Taverner’s subsequent career. For Anderson’s own career see Chapter 14. When examining a problem concerned with a particu- lar species an ornithologist is likely to require a series of plumages for comparison purposes. It is often pos- sible to trace a gradual change in measurement and/or morphological characteristics as specimens are com- pared that have been taken from regions a consider- able way apart. A change can be seen from smaller to larger, or darker to lighter. This is known as a cline. Populations of birds at either end of a cline may be strikingly different though they are linked by popula- tions that grade imperceptibly one into the other. These populations are said to intergrade. Canada, Sessional Papers 1914, “Summary Report 1913”, 7-8; page 352 under additions to the zoologi- cal collection. A type specimen is a particular speci- CRANMER-BYNG: A LIFE WITH BIRDS 84. 85. 86. 87. Die 92. 93. 99. M5) men of a bird, usually in the form of a museum skin, to which the scientific description of a species or sub- species refers. The “type specimen” is the skin for which the species or race was originally described. Such specimens are labelled with this information and can be re-examined in museum collections. This concept aims to preserve the stability of a taxon. Fleming was made an honorary curator of the National Museum of Natural Science in 1913. Taverner to Fleming, 11 February 1913, ROM. Horace H. Mitchell, born England 1868. Came to Canada as a young man and worked in Oliver Spanner’s taxidermy shop. Hired by the Saskatchewan Provincial Museum of Natural History at Regina in March 1913. A skilled craftsman who preferred exhibiting groups of birds in their natural habitat rather than single specimens. Mitchell held the post at the museum at Regina for twenty years, retiring in 1933. For further information see Annual Reports of the Game Commissioner, Department of Agriculture, Province of Saskatchewan. Taverner to Fleming, 5 and 19 December 1912; 14 March 1913. Teslin Lake is situated on the Yukon- British Columbia boundary. Taverner to Fleming, 19 December 1912. See P. A. Taverner The Auk 31: 385-388 (1914); also P.A. Taverner, Canada, Geological Survey, Museum Bulletin Number 7. December 1914, pages 1-4. Fleming’s Grouse was included in the fourth edition of the AOU Check-list of North American Birds 1931. It was also included in Taverner’s Birds of Canada 1934 and subsequent editions. But it was omitted from the fifth edition of the Check-list 1957. . Taverner to Fleming, 27 February 1913, ROM. . Taverner to Fleming, 27 February 1913, ROM. . Taverner to Fleming, 11 May 1913, ROM. This was for Dr William H. Collins. Born 1878; appointed to the Geological Survey 1905; married in 1908. His house was on Rosedale Avenue close to Taverner’s home. It was built to Taverner’s design in 1913-1914. Personal communication from Mrs Ann Whitmore, daughter of W. H. Collins. Taverner to Swales, 5 May 1913, CMN, “as busy as the devil ... “; Taverner to Fleming, same date “... like a chicken with its head off’, ROM. Taverner to Fleming, 16 January 1914, CMN. Dr Saunders was presumably W. E. Saunders’ brother, Charles Saunders. Taverner to Louis Bishop, 20 January 1914, CMN. Dr Louis B. Bishop M.D. of New Haven, Connecticut. Correspondence between Taverner and Bishop began in 1912 and continued regularly until 1940. . Taverner to Brock, 2 March 1914, CMN. . Brock to Taverner, 25 August 1914, CMN. . Memorandum to R.G. McConnell, Acting Deputy Minister of Mines, by the principal officers of the Geological Branch engaged in the work of the muse- um. Transmitted 16 October 1914. . Taverner to Fleming, 5 October 1914, ROM. . Claude E. Johnson. Mainly employed as colourist in the preparatory department, in the coloured illustra- tions of mammals and in painting background for habitat groups. Fleming to Taverner, 13 February 1914, ROM. It was bought, along with various other specimens such as a female and a juvenile Spectacled Eider; Fleming to 216 100. 101. 102. 103. 104. THE CANADIAN FIELD-NATURALIST Taverner, 10 April 1914. Taverner to Fleming, 17 April 1914 said: “The extralimital stuff is most desir- able and just what we want.”, ROM. Fleming to Taverner, 29 June 1914; Taverner to Fleming, 19 September 1914, ROM. Canada, Sessional Papers 1915, “Summary Report 1914”, page 154. Taverner to Fleming, 5 January 1914, ROM. Taverner to Fleming, 17 April 1914, ROM. Taverner to J. Leon Cole, 5 December 1914, in reply to letter from Leon Cole of 30 November 1914, from the College of Agriculture at the University of Wisconsin. Taverner’s statement that he was head of a division is not’accurate. The division was that of biology. Taverner was head of the sections of mam- mals and ornithology. Chapter 7 Ite 28. phe), 30. Sie ee a BONE SSCHNAAKWH P. A. Taverner “Suggestions for Ornithological Work in Canada” The Ottawa Naturalist 29: 14-18; 21-28 (1915). Same page 15 Same page 16 Same page 16-17 Same page 18 Same page 18 Same page 21 Same page 22 Same page 22 . Same page 23 . Same page 23 . Same page 24 . Same page 24-55 . On the last great rookery near Petoskey, Michigan, see P. A. Taverner Birds of Canada (Ottawa 1934) page 253. . Taverner “Suggestions for Ornithological Work” page 27. (See note 1) . Taverner to Fleming, 12 April 1915; Fleming to Taverner, 14 April 1915. Royal Ontario Museum Archives, Percy A. Taverner Papers and J.H. Fleming Papers, hereafter ROM. . Taverner to Fleming, 10 August 1915, ROM. . Taverner to Fleming, 25 August 1915, ROM. . Fleming to Taverner, 30 August 1915, ROM. . Taverner to Fleming, . Taverner to Fleming, 25 August 1915, ROM. 2. Fleming to Taverner, 30 August 1915, ROM. . Fleming to Taverner, . Fleming to Taverner, 29 June 1915, ROM. . Taverner to Fleming, . For a biography of Brooks see Hamilton M. Laing 31 August 1915, ROM. 18 November 1915, ROM. 14 July 1915, ROM. Allan Brooks: Artist Naturalist British Columbia Provincial Museum, Victoria, 1979. . Taverner to Henry Mousley, 27 November 1914, Canadian Museum of Nature, P. A. Taverner Ornithology Archive, hereafter CMN. On Mousley see Marianne Ainley “Henry Mousley and the Ornithology of Hatley and Montreal, 1910-14” Tchebec 11: 113-134 (1981). Taverner to Mousley, 22 February 1915, CMN. Taverner to Fleming, 14 July 1915, ROM. Fleming to Taverner, 22 November 1915, ROM. Fleming to Taverner, 24 June 1915. Dr. W.A. Waiser, author of The Field Naturalist. John Macoun, 64. Vol. 110 The Geological Survey, and Natural Science, Toronto, 1989, commented as follows on this sen- tence in Fleming’s letter: “Macoun did not like hav- ing younger people working under him because they were unreliable and liable to cause him trouble. His praise of Taverner was more of a compliment than it first appears to be.” William Waiser, letter to author, June 1986. 2. John Macoun to Taverner, 7 March 1915, CMN. . W.A. Waiser, Field Naturalist, 197-198. (See note 31) . Taverner to L. Bishop, 5 July 1916, CMN. . Taverner to Fleming, 24 June 1916, ROM. . Taverner to Fleming, 19 July 1916, ROM. . Taverner to Fleming, 30 November 1916, ROM. . Taverner to Fleming, 7 June 1916, ROM. . Taverner to Fleming, 22 January 1916, ROM. For details of Patch’s trip see Canada, Sessional Paper 1917, “Summary Report for 1916”, page 353. . Taverner to Fleming, 5 October 1916, ROM. . Fleming to Taverner, 11 September 1916, ROM. Todd’s book was finally published in 1963 by the University of Toronto Press under the title Birds of the Labrador Peninsula and Adjacent Areas. 2. Zaslow Reading the Rocks 324-325 (see chapter 5 note 1); Richard J. Diubaldo Stefansson and the Canadian Arctic (Montreal) 1978, pages 188-206. Canada, Sessional Paper 1916, “Summary Report” for 1915, page 249. . Canada, Sessional Paper 1917, “Summary Report” for 1916, pages 9-11; 344-346. . Taverner to Swales, spring 1915, CMN. . Taverner to Fleming, 14 September 1915, ROM. . Taverner to Fleming, 12 May 1916, ROM; Taverner to Brooks, 15 May 1916, CMN. . Taverner to Fleming, 12 May 1916, ROM. . Taverner to Fleming, 25 May 1916, ROM. . Taverner to Fleming, 24 June 1916; Fleming to Taverner, 2 July 1916, ROM. . Taverner to Fleming, 7 June 1916, ROM. . Taverner to Fleming, 11 June 1916, ROM. . Taverner to Fleming, 29 November 1917, ROM. An X-ray showed abcesses at the roots of what seemed good teeth. . Taverner to Louis Bishop, 5 July 1916, CMN. . Taverner to Louis Bishop, 6 March 1917, CMN. . Taverner to Fleming, 6 February 1917, ROM. . Taverner to Fleming, 24 March 1917, ROM. . Anderson to Fleming, 30 June 1917, ROM. . Anderson to Fleming, 4 July 1917, ROM. . For a more detailed treatment of Anderson’s back- ground see Chapter 14. . Taverner to Fleming, 6 February 1917, ROM. . Taverner to Fleming, 26 February 1917, ROM. . Fleming to Taverner, 29 April 1917; Taverner to Fleming, 6 May 1917, ROM. Charles Lucien Bonaparte, a nephew of Napoleon I, tackled the description of the North American avifauna in his American Ornithology during an eight-year stay in America (1822-1828). He is generally acknowledged to have been a foremost ornithologist of his time. Bonaparte’s Gull was named in his honour. For the contemporary status of anthinus see Godfrey, Birds of Canada 1986, page 524. Taverner to Brooks, 14 August 1916, CMN. 1996 65 66. 1) 80. 81. 82. . Taverner to Fleming, 6 February 1917, ROM. Taverner to Fleming, 29 September 1917, ROM. Percy sent a post card from Bala Falls to tell Fleming that he was spending the weekend at Muskoka, and that he expected to be returning mid-week and would stay a while in Toronto between trains. The only explanation I can think of for Percy’s visit to Beaumaris was to ask the advice of one or two of his contacts there on building a cottage, perhaps to obtain some plans and measurements. . Taverner to Fleming, 12 January 1918; 30 January 1919, ROM. . Taverner to Fleming, 5 April 1917, ROM. . J.M. Macoun to R. G. McConnell, 22 February 1918, NAC. I am grateful to Professor W. A. Waiser for sending me this quotation. . Waiser Field Naturalist, pages 198-199. (See note 31) . Waiser Field Naturalist, page 199. (See note 31) . Taverner to Fleming, 18 September 1918, ROM. . Taverner to Fleming, 21 July 1919, ROM. . Taverner to J. M. Macoun, 15 July 1919, CMN. . Taverner to Fleming, 31 December 1918, ROM. . Taverner to Fleming, 27 February 1918; same to same, 11 December 1918; Fleming to Taverner, 16 December 1918, ROM. . Taverner to Brooks, 24 October 1918, CMN. . Taverner to Fleming, 26 February 1918, ROM. A King Log is a roi faineant, a king who rules in peace, but never makes his power felt. An allusion to the Greek myth of the frogs who asked Jupiter for a king. At first Jupiter threw down a log to govern them, but they complained at so spineless a king. In response Jupiter sent them a stork, who ate them all up with gusto. Luckily for Taverner and his colleagues they never had to live under a King Stork. James Harkin b. 1875 Ontario. Parliamentary corre- spondent with Montreal and Toronto newspapers. Private Secretary to Clifford Sifton, Minister of Interior in Laurier’s government. Dominion Parks Commissioner 1911. Gordon Hewitt b. 1885 England. B.Sc. in zoology Manchester University, Ph.D. 1909; lecturer in Economic Zoology; Dominion Entomologist Canadian Dept. of Agriculture 1909. Robie W. Tufts “Notes on the birds of the Grand Pre region, King’s County, Nova Scotia” Transactions of the Nova Scotian Institute of Science 14: 154-199 (1917). Taverner to Robie Tufts, 28 February 1918, CMN. Tufts was born 1884 Wolfville, Nova Scotia. Educated Acadia University. Keenly interested in field observations of birds. Taverner to Fleming, 24 March 1916, ROM. It is of interest to note that sixty-one years later when the Breeding Bird Atlas of Ontario was published, although more than 70 000 Black Scoters were seen summering off the northern coast (Hudson Bay region) of Ontario not a single confirmed breeding record existed for Ontario. Taverner to Fred Bradshaw, 24 October 1919, CMN. The list by Fleming referred to by Bradshaw is: J.H. Fleming “Birds of northern Saskatchewan and north- ern Manitoba collected in 1914 by Capt. Angus Buchanan”, The Canadian Field-Naturalist 33: 109- 113 (1919). CRANMER-BYNG: A LIFE WITH BIRDS 90. 91. 92: 03) 98. 99. 100. 101. 102. 103. 104. 105. 217 . Bradshaw to Taverner, 28 October 1919, CMN. . Taverner to Bradshaw, 6 November 1919, CMN. . W.J. Brown to Taverner, 29 January 1919, CMN. P. A. Taverner “The Birds of the Red River, Alberta” The Auk 36: 1-21; 248-65 (1919). In his reply of 5 February 1919 Taverner explained to Brown that in the more southern localities of Canada changes in the countryside were proceeding very rapidly and he therefore wanted to record the birds before they van- ished for ever. . Taverner to W. J. Brown, 5 February 1919, CMN. . Brown to Taverner, 11 February 1919, CMN. . Taverner to Brown, 15 February 1919, CMN. . Taverner to Brown, 25 February 1919, CMN. Taverner did considerable taxonomic research on the Great Horned Owl. For comparison see P. A. Taverner Birds of Eastern Canada (1919) and Taverner Birds of Canada (1934) under Great Horned Owl. Also Chapter 16, pages 411-412 and note 105. Taverner to Brown, 25 February 1919, CMN. For instance Taverner to Fleming, 12 March 1918, in which he wrote that Mousley was giving the museum a lot of fine warbler nests in situ, and good for exhibi- tion groups. Fleming to Taverner, 1 October 1918, ROM. Writing to Taverner on 2 December 1918, soon after Lloyd had been appointed, Fleming said that he had an ana- lytical mind that would be of great service in ornithology, and that Lloyd was in the forefront of young chemists. Fleming to Taverner, 13 October 1918, ROM. Lloyd collected bird skins in the 1900s and published his first ornithological paper in 1917. H. Lloyd “Ontario Bird Notes” The Auk 34: 457-460 (1917). Also in conjunction with J. H. Fleming he wrote “Ontario Bird Notes” The Auk 37: 429-439 (1920). Articles by Lloyd on “Birds of Ottawa” began appearing in The Canadian Field-Naturalist in September 1923. . Fleming to Taverner, 29 November 1918, ROM. . Taverner to Fleming, 27 December 1918, ROM. . Taverner to Fleming, 14 January 1919, ROM. . Fleming to Taverner, 16 January 1919, ROM. The BOU honoured several North American ornitholo- gists at its annual meeting in 1919 including D. L. Stejneger who became an honorary member, and Dr. Joseph Grinnell and Outram Bangs became foreign members. The Ibis, Journal of the British Ornithologists’ Union. Founded in 1859. Still one of the most prestigious English language ornithological journals. Named after the /bis family of birds. Taverner to Fleming, 25 February 1919; Fleming to Taverner, 24 March 1919, ROM, reported that Mitchell had recently been to see him. Fleming told him that he hoped Patch would tell him everything possible when in Ottawa. Taverner to J. M. Macoun, 6 September 1919, CMN. Taverner to Fleming, 10 October 1918, ROM. Taverner to Fleming, 8 October 1919, ROM. Taverner to Fleming, 21 July 1919, ROM. Taverner to J. M. Macoun, 6 August 1919, CMN. Taverner to Fleming, 17 September 1919, ROM. Entry in log book of Hyla cottage 28 September 1919. I am grateful to Mr. Corwin Ferguson of Detroit for lending me the Hyla log book to take notes from. 218 106. 107. 108. Cha 1 i) 24. THE CANADIAN FIELD-NATURALIST Anderson to Fleming, 11 October 1919. ROM Fleming Papers. Anderson to Fleming, 27 October 1919, ROM. Owners of lots on Big Island were nearly all museum people. Waiser The Field Naturalist pages 199-200. (See note 31) pter 8 Taverner to Swales, 26 December 1912, ROM; P.A. Taverner “Geological Survey Museum Work on Point Pelee, Ont.” The Ottawa Naturalist 28: 97-106 (1914). . Taverner “Journal” 28-30 May 1910, pages 330-331, ROM. . Taverner to Swales, 23 October 1912: Taverner to Brooks, 8 December 1912, CMN. . Taverner to Fleming, 5 May 1913; 11 May 1913, ROM. There were rules for which department should buy what commodities, and how to order them. . Entitled “Birds observed at Point Pelee. The Property of Camp Coues”. It was volume 2, and began in 1913, ROM. . This specimen is preserved in the National Museum #6797. It is an immature male molting into summer plumage. . J.G. Battin and J.G. Nelson Man’s impact on Point Pelee National Park (Toronto, 1978) pages 89-90. . Taverner to Swales, 6 June 1913, CMN. . Taverner to James Macoun, 19 June 1913, CMN. He also wanted advice on how to use a tree such as the redbud which was not yet in flower. . Taverner to Fleming, 20 June 1913, ROM. . Taverner to Fleming, 5 August 1913, ROM. In the same letter he said that the exhibit “was a frost. Hardly a soul looked at it”. . Taverner to Fleming, ROM. . Saunders to Swales, 29 June 1913, CMN. . Collection of 13 black-and-white photos approxi- mately 3 x 5 inches with brief captions. Some were sent by Taverner to friends as Christmas greetings December 1913. ROM archival material. Some of Taverner’s Point Pelee negatives of 1913 are pre- served in the Canadian Museum of Nature photo- graphic collection. . P.A. Taverner The Ottawa Naturalist 28: 98 (1914). . Saunders to Swales, 29 June 1913, CMN. . Taverner to Swales, 26 December 1912, CMN. . Taverner to Fleming, 14 March 1914, ROM. . Taverner to Brooks, 18 March 1914, CMN. . Taverner to Fleming, 17 April 1914, ROM. . Canada, Sessional Paper 1915, “Summary Report for 1914”, pages 158-159. . Typed copies in ROM Archives, Percy Taverner Papers. . Taverner to Mrs. Ida Taverner, dated Bathurst 14 June 1914, ROM. This date is probably wrong. Percy explained that from Miscou to Percé by sailboat could take as little as four hours, but all the boats were busy fishing at that time and no reliable arrangement could be made. The rail journey round Chaleur Bay was several hundreds of miles. Taverner to Ida Taverner, 21 June 1914, ROM. The same view, on a sunny summer’s day, can have the same effect on the visitor to Percé now as it did on Taverner then. 25 26 Vol. 110 . Taverner to Ida Taverner, 21 June 1914, ROM. . Taverner to Ida Taverner, 6 July 1914, ROM. There is nothing fanciful about this description. To the pre- sent day visitor the masses of wheeling birds above the cliff on Bonaventure Island can be equally as inspiring. 27. Taverner to Ida Taverner, 18 July 1914, ROM. 4A. 45. 46. 28. Taverner to Ida Taverner, 30 July 1914, ROM. . Taverner to Ida Taverner, 30 July 1914, ROM. Percy explained that just at this period, and the next few weeks, the cormorants would be coming into a partic- ular plumage phase. Such birds were needed to com- plete the museum’s series of plumages. . Taverner to Ida Taverner, 8 August 1914, ROM. The Bird Rocks (Les Rochers des Oiseaux) lie about twenty miles north of the Magdelens. The larger one is ared sandstone hillock, with a lighthouse on it, and a population of around 12 000 birds in total in 1914- 1915. A smaller bird rock nearby is lower and less dramatic but also supported a large number of nesting seabirds. J. J. Audubon passed great Bird Rock on 14 June 1833 on his visit to Labrador, and described what he saw in his Journal of that date. Harrison F. Lewis “Some Canadian Auduboniana” The Canadian Field-Naturalist 47: 162-172 (1933). . Taverner to Ida Taverner, 12 August 1914, ROM. . Taverner to Ida Taverner, 12 August 1914, ROM. This part of the letter was written a few days later but included under this date. . Taverner to Ida Taverner, 12 August 1914, ROM, later section. . Taverner to Ida Taverner, 12 August 1914, ROM, later section. . Taverner to Ida Taverner, 8 August 1914, ROM. . Canada, Geological Survey Museum Bulletin 13, 1915. . Taverner to mother and sister, 8 August 1914, ROM. . A.C. Bent to Taverner, 2 December 1914; Taverner to Bent, 5 December 1914, CMN. From the begin- ning of the correspondence between Taverner and Arthur Cleveland Bent the subjects they discussed were interesting and the information they gave was detailed. These letters on plumage change in gannets was no exception. . Canada, Sessional Papers 1916, “Summary Report for 1915”, pages 260-262. . Taverner to mother and sister, 11 June 1915, ROM. . Taverner to Fleming, 12 June 1915, ROM. . Taverner to mother and sister, 11 June 1915, ROM. . Taverner to Fleming, 4 July 1915, ROM. He described the landscape with the eye of an artist but economically. “The moss in fact tends to cover every stone & bare place growing [word illegible] and in all colors of gray through green and brown to dull red. The color of the landscape at first glance is dull sage green but close observation shows wonderful color everywhere — gray, green, red and violet on every hand and a [two words illeg.] makes a kaleidoscopic effect I have seldom seen equaled”. See note 39 above. Frank Chapman Camps and Cruises of an Ornithologist (New York 1908) pages 315-336. Taverner to Fleming, 26 July 1917, ROM; P.A. Taverner “The Birds of Shoal Lake, Manitoba” The Ottawa Naturalist 32: 137-144; 157-164 (1919); 33: 1996 47. 48. 49. 50. Sls oy, Se 54. SS: 56. Dille 12-20 (1919). This is an annotated list of the birds of that locality from observations and collections made in 1917 and 1918 which Taverner considered to be a fairly complete and representative list. No letters from Percy to his family appear to have survived for 1917. But see Taverner to L. Bishop, 12 October 1917, CMN. P. A. Taverner “The Birds of the Red Deer River, Alberta” The Auk 36: 1-21; 248-265, 1 map (1919); Taverner to Fleming, 26 July 1917, ROM. Taverner to W. E. Saunders, 11 October 1917, CMN. For a short note on the Whyte Museum see Official Directory of Canadian Museums and Related Insti- tutions. Canadian Museums Association (yearly). Taverner to Munro, 9 October 1917, CMN. In this newsy letter Taverner told Munro that he was home again after five months in the field, and described his itinerary after leaving him in July. His letter con- tained one cryptic reference when he wrote: “Do not let my unfortunate appearance at Okanagan Landing disturb you. It was one of those unavoidable acci- dents. I am delighted to hear that Mrs. Munro recov- ered.” His “unfortunate appearance” was probably due to the use of arsenic while preparing bird skins in the field. From Munro’s reply of 3 November 1917 it is clear that Isabella Munro was critically ill. She died in 1919. Taverner to Fleming, 20 August 1917 (from Prince Rupert), ROM. Informing him of the birds he has been seeing. Also contains a brief reference to “Munro’s tragedy”. Taverner had stayed four days at Alert Bay, Cormorant Island, off Vancouver Island. Apart from birds, the village had a major collection of totem poles. He obtained a lot of good plumages and made colored drawings of bills and feet while there. P. A. Taverner “The Summer Birds of Hazelton, B.C.” Condor 21: 80-86 (1919). P. A. Taverner “William Spreadborough — Collector 1856-1931”, Canadian Field-Naturalist 47: 39-41 (1933). P. A. Taverner, see note 52, pages 39-41. P. A. Taverner, see note 52, pages 41-42. Brooks, writing to Taverner on 25 September 1919, men- tioned that Spreadborough did not identify Brewer’s Sparrow. Taverner replied: “I am not surprised that Spreadborough misses Brewer’s Sparrow. You know that he is only a shanty man”. It was surprising, Taverner explained, that with only guidance from the Macouns Spreadborough managed as well as he did. He had no one to guide him in the finer points and only collected the obvious. Certainly he got few rare species, Taverner said, “but if it had not been for him we would not have much of a collection now and our knowledge of Canadian birds would be smaller than it is even allowing for the inaccuracies that have crept in through some of his statements.” Taverner to Brooks, 6 October 1919, CMN. P. A. Taverner “Birds of Shoal Lake” page 140. (See note 46) Taverner to Fleming, 25 July 1918, CMN. Taverner to Brooks, 7 October 1918, CMN. Also Taverner to L. Bishop, 30 July 1918, CMN; Taverner to Fleming, 18 September 1918, ROM. For informa- tion on his field notebooks see note 60 below. CRANMER-BYNG: A LIFE WITH BIRDS 58 59. 60. 61. 62. 63. 64. 65. 66 219 . Taverner to Fleming, 21 May 1919, ROM. What Taverner meant by “waiting for the word to go or stay” was waiting for Parliament to grant appropria- tions to the museum for field expeditions. Taverner to Fleming, 21 July 1919, ROM. For details of contents of all Taverner’s field notes from 1913 to 1937, both the handwritten and the later typewritten ones, see the Catalogue “Field notes in ornithology collections” compiled by Michel Gosselin 1992, CMN. Taverner to Fleming, 14 March 1914; Fleming to Taverner, 19 March 1914, ROM. An account of the expedition was given by Charles Camsell in chapter 31, “Taltson River Exploration” in his autobiography Son of the North (1954). Francis Harper, Canada: Sessional Papers 1915 “Summary Report for 1914”, 159-163. Harper later published accounts of results in 1931. “Physiographic and faunal areas in the Athabasca and Great Slave Lakes region”. Ecology 12: 18-32 (1931). “Amphibi- ans and reptiles of the Athabasca and Great Slave Lakes region.” The Canadian Field-Naturalist 45: 68-70. Taverner gave the name American Mew Gull as an alternative name for the Short-billed Gull in his Birds of Western Canada pages 57-58 (1926). Taverner described this goose in Birds of Western Canada as Branta canadensis hutchinsi, a subspecies in between the large Honker (B. c. canadensis) and the small Cackling Goose (B. c. minima). Normally the Hutchin’s Goose was found breeding on the Barren Grounds and the Arctic Islands, a long way north of the Taltson River. . See under “Accessions” in the “Summary Report for 1914”, page 155. Chapter 9 ile oo T. Gilbert Pearson “Fifty Years of Bird Protection in the United States” in Frank Chapman and T.S. Palmer, editors, Fifty Years’ Progress of American Ornithology 1883-1933 American Ornithologists’ Union (Lancaster, Pennsylvania, 1933) pages 199- 200. . Pearson “Fifty Years” pages 201-202. (See note 1) . Pearson pages 208-210. (See note 1) . These bird observations were arranged according to the AOU Check-list (1931) and written up with notes based on Fothergill’s manuscripts by R. Delamere Black “Charles Fothergill’s Notes on the Natural History of Eastern Canada, 1816-1837” Transactions of the Royal Canadian Institute 20: 141-168 (1934). Fothergill’s ms. notes are preserved in the Fisher Rare Book Library, University of Toronto (MS 140 v 22). . See Hugh R. MacCrimmon Animals, Man and Change: Alien and Extinct Wildlife in Ontario (Toronto 1977) pages 27-28. Also M. D. Cadman et al., editors, Atlas of the Breeding Birds of Ontario (Waterloo, 1987) page 142. . Margaret H. Mitchell The Passenger Pigeon in Ontario (Toronto, 1935) pages 129-139. . Janet Foster Working for Wildlife: the Beginnings of Preservation in Canada (Toronto, 1978) page 10. . J. Foster Working for Wildlife page 26. (See note 7) . Foster Working for Wildlife page 13 (See note 7) 220 10. 15! . Foster Working for Wildlife 130. (See note 7) L7: DD: 23: 24. 25): THE CANADIAN FIELD-NATURALIST Foster (see note 7) pages 38-40; For a valuable account of the Commission of Conservation see Michel F. Girad “The Commission of Conservation as a Forerunner to the National Research Council 1909-1921”. Scientia Canadensis 15, Number 2, 1991 (published by Canadian Science and Technology Historical Association) pages 19-40. . Foster (see note 7) pages 77-78. 12 13. Quoted in Foster (see note 7) 127-128. Foster (see note 7) pages 128-129. The material which follows in the next pages is discussed more fully in Foster 128-138. . Maxwell Graham to James Macoun, 20 March 1913, Canadian Wildlife Service Records Migratory Bird Protection. Taverner to Graham, 22 March 1913, CMN. C. Gordon Hewitt born in England 1885. Studied Manchester University. Doctor of Science. Appointed Dominion Entomologist, Canada, 1909. For his role in the protection of wildlife in Canada see Janet Foster Working for Wildlife pages 136-138 and index. Obituary in The Canadian Entomologist 52: 96-97 (1920). . On James White see Michel Girard “Commission of Conservation” (see note 10) pages 23-39. As Chief Geographer of the Dominion, White had the first Atlas of Canada published. . “Supplement to an Address on Animal Sanctuaries in Labrador by Lt. Colonel William Wood, F.R.S.C.” Published by the author in pamphlet form 1912. See Foster Working for Wildlife 180-185. (See note 7) . Foster Working for Wildlife 182-183. (See note 7) . Taverner’s findings were published in “The Double- crested Cormorant (Phalacrocorax auritus) and its Relation to the Salmon Industries on the Gulf of St. Lawrence”. Geological Survey Canada. Museum Bulletin 13: 1-16 (1915). Taverner to Harkin, 14 December 1914. Harkin to Taverner, 21 January 1915, CMN. From this time onwards Harkin consulted Taverner fairly often. The memorandum contained photographs of the gan- net ledges on Bonaventure Island taken by Taverner. Taverner did not speak at the meeting in January per- haps because of his stammer, or because he was not sufficiently senior to be asked. Taverner to Fleming, 10 February 1915, enclosing a copy of his memorandum to Harkin, ROM. Taverner to Fleming, undated, but see on same sub- ject Taverner to Swales, 3 November 1915, CMN. Saunders’ talk, “Bird Protection in Canada”, was given before the “Committee on Conservation of Fish, Birds and Game of the Commission of Conservation”. It was published in the Report of the Commission of Conservation (1916) pages 117-119. . Charles Townsend “Preliminary Report of an Expedition along the North Shore of the Gulf of St. Lawrence” in Canada, Sessional Papers 1916, “Summary Report for 1915”, pages 262-263. . Taverner, Summary Report (1915) pages 261-262. . Taverner to White, 3 September 1915, CMN. . See James M. Linton and Calvin W. Moore The Story of Wild Goose Jack: the Life and Work of Jack Miner, Toronto 1984. Also Janet Foster Working for Wildlife (see note 7), mentions Miner’s achievements 30. Shile Bs 33. 34. 35. 36. Vol. 110 in relation to the contributions of others to bird con- servation. There is a trail which may lead to proving this to be more than a supposition. But I need time to research the ramifications of the relations between W.E. Saunders — P.A. Taverner, and Saunders — J. Miner in the years 1905-1906. See Taverner “Memoirs of William Edward Saunders 1861-1943” The Auk 61: 350 (1944). W.E. Saunders and J.A. Morden Canadian Sportsman and Naturalist 2: 183-187; 192-194 (1882). Also with Morden “The Ornithology of Western Ontario” Canadian Sportsman and Naturalist 3: 243 (1883). For Morden’s obituary notice see William E. Saunders “John A. Morden 1859-1919” Canadian Field-Naturalist 51: 108 (1937). Manley Miner to P. A. Taverner, 6 March 1946, CMN. For information on W. E. Saunders as a natu- ralist in southwestern Ontario see William W. Judd Early Naturalists and Natural History Societies of London, Ontario (London Ontario 1979) pages 57-96. James M. Linton and Calvin W. Moore The Story of Wild Goose Jack; the Life and Work of Jack Miner (Toronto 1984). Taverner to White, 28 March 1916. Correspondence between Taverner and White is preserved in CMN, P. A. Taverner Ornithology Archives under “Commis- sion of Conservation”. Taverner to Townsend, 28 March 1916, CMN; also letter 31 March 1916 in which Taverner stated “The worst of it is those people get things done, and one cannot ignore them without losing important assis- tance in good work. It is annoying, however, to find that one has to tie them up like sharpers before trust- ing them with anything”. Taverner implied that the Commission were apt to use other people’s pho- tographs and writings without acknowledgement. . This paragraph is based on Janet Foster’s account in her book Working for Wildlife (see note 7) pages 127- 144. The text of the Convention is printed on pages 225-235. The Act was to be cited as The Migratory Birds Convention Act. . Foster Working for Wildlife 144-146. (See note 7) . Taverner to Swales, 3 November 1915, CMN. . Taverner to John H. Sage, 23 October 1916, CMN. Taverner had received some praise from Hewitt in 1915 who said “In the matter of education very satis- factory work has been carried on through the efforts of the Departments of Agriculture and Mines, through Mr. Taverner in the Geological Survey ... ” C. Gordon Hewitt “Conservation of Birds and Mammals in Canada” in proceedings of a meeting of the Committee on Fishes, Game and Fur-bearing Animals November | and 2 1915, pages 144-145. This was not much praise but it was some slight recognition. . Taverner to Fleming, 30 January 1918, ROM. . Foster Working for Wildlife 161-163. (See note 7) . Foster (see note 7) 169-172 on information from Anderson. . Taverner to Hewitt, 27 December 1917, CMN. . Taverner to Hewitt, 18 January 1918. . Taverner to Fleming, 16 February 1918, ROM. . Brooks to Taverner, 10 September 1918; Taverner to Brooks 7 October 1918, CMN. | 1996 48. 49. 64. 2? Taverner to Swales, 28 December 1918, CMN. Foster Working for Wildlife page 159. (See note 7) Information based on an interview (1971) which Foster had with Harrison F. Lewis who was appoint- ed Federal Migratory Bird Officer for Ontario and Quebec in 1920. . Taverner to Bishop, 16 January 1919, CMN. . Foster Working for Wildlife 179-180. (See note 7) . See note 23 above. . Foster Working for Wildlife (see note 7) pages 125; 194-197; also the monthly magazine Rod and Gun in Canada (1917). . Hewitt to Taverner, 9 May 1918, CMN. . Hewitt to Taverner, 13 December 1918; 18 December 1918, CMN. . Hewitt to Taverner, 31 January; 27 February; 11 March 1918; 4 February 1919, CMN. . Hewitt died of pleural pneumonia on 29 February 1920. See Foster Working for Wildlife 209-210. (See note 7) Obituary by Norman Criddle in The Canadian Field-Naturalist 34: 174-176 (1920). . On this subject see NAC Records. Point Pelee National Park January-June 1919. . Edward R. Kerr to W.E. Saunders, 8 May 1919, CMN. . E.R. Kerr to W. E. Saunders, 16 May 1919. . Duck hunting was ended at Point Pelee National Park by order of the Federal Environment Minister, Lucien Bouchard, on 6 June 1989. The press release issued by the Minister (reference PR-HQ-089-24) also con- tained a one page background on “Duck hunting at Point Pelee National Park”. 2. P. A. Taverner “The Gannets of Bonaventure Island” Ottawa Naturalist 32: 21-26 (1918). . Taverner to Hewitt, 13 December 1918. Taverner’s expectation, that if the birds of Bonaventure and the town of Percé were protected, the townspeople would benefit considerably by the tourist trade, was con- firmed later by a letter to Taverner from a local resi- dent. Hoyes Lloyd “The Ornithological Collector and the Law” The Canadian Field-Naturalist 33: 93-94 (1919); “Permits to Collect Birds for Scientific Purposes in Canada” The Auk 36: 621-623 (1919); P.A. Taverner Vanished and Vanishing Birds Dominion Parks Branch Migratory Bird Leaflet (1919). Also in French edition. . Taverner Vanishing Birds 3. (See note 65) . Taverner Vanishing Birds 4. (See note 65) . Taverner Vanishing Birds 8. (See note 65) . Taverner to J. Miner, 14 November 1918, CMN. . J. Miner to Taverner, 4 December 1918, CMN. . Taverner to J. Miner, 12 December 1918, CMN. . C. Gordon Hewitt “The Need of Nation-wide Effort in Wild Life Conservation” National Conference on Conservation of Game, Fur-bearing Animals and other Wild Life. Under the Direction of the Commission for Conservation in co-operation with the Advisory Board on Wild Life Protection, February 18 and 19, 1919 (Ottawa 1919). Edith L. Marsh, local historian and naturalist, died Peasemarsh Farm, Thornbury, Ontario 10 July 1960. Author of Where the buffalo roamed (Toronto, 1908); Birds of Peasemarsh (Toronto, 1919); With the birds (Toronto, 1935). CRANMER-BYNG: A LIFE WITH BIRDS Ws 74. 80. 81. Dapaih Jack Miner “Attracting Wild Fowl” in National Conference on Conservation of Game ... (Ottawa, 1919) pages 82-89. (See note 72) Taverner to Swales, 22 February 1919, CMN. The phrase “a scrap of paper” was used by the German Chancellor, Theobald Von Hollweg, at the outbreak of war between Great Britain and Germany on 4 August 1914. When interviewing the British Ambassador in Berlin he said “Just for a word — ‘neutrality’, a word which in wartime has so often been disregarded, — just for a scrap of paper Great Britain is going to make war.” Taverner knew the significance of this phrase to the public at that time. . Taverner to Swales pages 3-4. About Jack Miner see note 73. . Taverner to Fleming, 25 February 1919, ROM. . Miner to Taverner, 21 February 1919, CMN. . Taverner to Miner, 25 February 1919, CMN. . In the early 1930s, when the future wildlife artist Terry Shortt had recently joined the Royal Ontario Museum, he was given the task of preparing, as spec- imens, some of the owls and hawks sent to the muse- um by an arrangement with Miner. To a young man with a tremendous respect for birds of prey this was a horrible experience. Handling Red-shouldered and Cooper’s Hawks with badly mangled legs brought on a bitter feeling. To hear killing of this nature justified because of a hawk’s “cruelty” to other birds was to him the result of gross ignorance of the natural order. “Human morals have no part in the rhythms and bounds of the natural world.” (Terry Shortt personal communication to author September 1986.) As one example of these results see George H. Harrison, editor, Roger Tory Peterson’s Dozen Bird- ing Hot Spots: A guide to the 12 best locations in North America for amateur bird watching (New York 1976). Illustrated black-and-white. See Harrison, author, “Point Pelee: Funnel to the North” and “Gaspe: Seabird Bastion”. New York, 1976. Surveys of seabirds on the North Shore of the St. Lawrence and the Gaspé have been continuous since. The most recent accounts are: Gilles Chapdelaine and Pierre Brosseau. 1991. Thirteenth census of seabird popula- tions in the sanctuaries of the North Shore of the Gulf of St. Lawrence, 1982-1988. Canadian Field- Naturalist 105(1): 60-66; Gilles Chapdelaine and Pierre Brousseau. 1992. Distribution, abundance, and changes of seabird populations of the Gaspé Peninsula. 1799 to 1989. Canadian Field-Naturalist 106(4): 427-434. Charles W. Townsend “Notes on the Summer Birds of the Gaspe Peninsula, Province of Quebec” The Canadian Field-Naturalist 34: 78-80; 87-95 (1920). Taverner gave additional information when writing to Fleming: “Townsend on my advice is spending his honeymoon at Percé. He is delighted with it, the scenery, the people and the birds. Has added several species to the list and is going to work up the list. Have turned over my notes to him so that work will go on record, as it should have done before.” Taverner to Fleming, 5 August 1919, ROM. When Townsend’s first wife died in 1917 he gave up his medical work and devoted himself to bird study and travel. In 1919 he married his sister-in-law. 222 Cha 1 Ds 3: 4. 5 2 13; 14. iWays 16. NNN WN ABWhN N Oo THE CANADIAN FIELD-NATURALIST pter 10 . Taverner to Brooks, 12 April 1912, CMN. Reviewed by W.E. Saunders in The Ottawa Naturalist 23: 225-228 (1910). Elliot Coues Key to the Birds of North America 5th edition, 2 volumes (1903). Taverner to Brooks, 19 March 1912, CMN. . Brooks to Taverner, 31 March 1912, CMN. Brooks was keen on military rifle shooting. See Marjorie Brooks “Allan Brooks — A Biography” The Condor 40: 15 (1938). . Taverner to Brooks, 12 April 1912, CMN. . Taverner to Fleming, 29. August 1912, ROM. Taverner wrote that he was delighted with his first meeting with Brooks. “He is all you have said of him and I am delighted to meet him and have a good bird talk with someone that is of some use. That he is a Canadian is so much the better”. Also, 3 October 1912 in which Taverner said he was most favorably impressed with Brooks. “He is quite a man and I am glad to have a reliable one out there near the coast”. [of British Columbia]. . Taverner to Brooks 24 September; 8 December 1912. Brooks to Taverner 17 November; 14 December 1912, CMN. . Taverner to Fleming, 3 September 1913, ROM. . Taverner to Fleming, 5 September 1913, ROM. . Taverner to Brooks, 5 September 1913, CMN. By “picture work” he means “‘illustrations”’. Taverner to Fleming, 23 September 1913, ROM. Fleming to Taverner, 29 September 1913, ROM. In short, Fleming excused himself from collaboratiing in the project. He was only interested in writing a schol- arly book for the serious collector which would take years to complete. Fleming realized that there was an urgent demand for a popular book but felt that Brooks and Taverner could write it between them with some help from himself when needed. Brooks to Taverner, 16 October 1913, CMN. Hamilton M. Laing Allan Brooks: Artist Naturalist British Columbia Provincial Museum, Victoria, page 106 (1979). Taverner to Fleming, 2 September 1914, ROM. Taverner told Fleming that Brooks’ eyesight and hearing were wonderful. He was forty-five years old. Taverner added “Too bad he had to get into this mix up but of course he is just the type of men that are wanted for the front just now”. . The first set of prints of Hennessey’s illustrations for Taverner’s book (1919 edition) were by far the best. The plates for the second printing (1922) were less well coloured and clear. . Taverner to Fleming, 12 April 1915, ROM. For Taverner as a bird illustrator see Chapter 4 above, note 134. . Fleming to Taverner, 14 April 1915, ROM. . Taverner to Fleming, 17 April 1915, ROM. . Taverner to Swales, no date, but from internal evi- dence it was written in the spring of 1915, CMN. . Taverner to Fleming, 25 August 1915, ROM. . Fleming to Taverner, 30 August 1915, ROM. . Fleming to Taverner, 31 August 1915, ROM. . Fleming to Taverner, 14 September 1915, ROM. . Taverner to Saunders, 13 April 1915, CMN. 27. Taverner to Saunders, 17 May 1915, CMN. 33. 34, 35: Vol. 110 . Taverner to Fleming, 10 January 1916, ROM. . Fleming to Taverner, 20 January 1916, ROM. . Taverner to Fleming, 20 February 1916, ROM. . Taverner to Fleming, 25 February 1916. See Introduction to Birds of Eastern Canada pages 1-7. . Taverner to Fleming, 25 February 1916. Taverner wanted to guide the new wave of bird-watchers, who | would be starting soon after the Great War, by the | use of a reliable system. He wished to introduce them to the study of ornithology by showing them how to recognize species first. Later, as they became more experienced, they could study the fine points needed for distinguishing subspecies if they wished to. Taverner to Fleming, 29 February 1916, ROM. Taverner to Fleming, 14 March 1916, ROM. Although Taverner might respond quite sharply at first when his ideas were questioned he usually modi- fied them somewhat, later. Fleming to Taverner, 16 March 1916, ROM. Taverner explained that the egrets’ plumes are grown ~ only in the breeding season so that they can be obtained only in the vicinity of a breeding egretry. He described how they were taken: “The plume hunter usually hides in the rookery and with a small-calibre rifle shoots the birds one by one until the flock is exterminated. The plumes are torn from the bodies which are left to rot on the ground. The remaining young starve in the nests above ... A few years ago the waters of Florida and the Gulf states were made beautiful with the forms of these immaculate birds; today they have almost lost one of their greatest attractions as the birds are approaching extinction”. P. A. Taverner Birds of Eastern Canada page 83 (1919). In contrast James Macoun advised Taverner that his book was just the place in which to make such appeal. J. M. Macoun to Taverner, 27 March 1916, CMN. . Fleming to Taverner, 16 March 1916, ROM. . Taverner to Fleming, 24 March 1916, ROM. . With a paper covered book it was quite normal at that period for the owner to get it bound. The volume could be trimmed to size at the time of binding. Taverner would usually bind his own copies. . Taverner to Fleming, 25 May 1916, ROM. . Taverner to Brooks, 15 May 1916, CMN. . Taverner to Fleming, 8 May 1917, ROM; Brooks to Taverner, 14 April 1917, CMN. About collecting, making up skins, and sending boxes of skins to Taverner. . Brooks to Taverner, 30 March 1918; 10 September 1918, CMN. . See note 15 above. . Taverner to Fleming, 5 July 1916, ROM. . Fleming to Taverner, 19 July 1916, ROM. . Taverner to Fleming, 24 July 1916, ROM. . Taverner to Brooks, 5 October 1916, CMN. . Fleming to Taverner, 5 October 1916, ROM. . Taverner to Fleming, 3 October 1916, ROM. . Fleming to Taverner, 5 October 1916, ROM. . Zaslow Reading the Rocks page 290. (See chapter 5, note 1) . Taverner to C.W. Townsend, 18 February 1917, CMN. . Taverner to Fleming, 6 February; 14 February 1917, ROM. 1996 66. 67. 68. 69. 70. . Taverner to Fleming, 9 June 1919, ROM. . Taverner to Brooks, 6 November 1919, CMN. . Louis Fuertes to Taverner, 28 November 1919, CMN. . Taverner to Fuertes, | December 1919, CMN. . Gordon Hewitt to Taverner, 25 November 1919, CMN. The first edition in English went out of stock in 1921 and was reprinted in 1922. . Taverner to R. W. Tufts, 28 February 1918 asking for a copy of his article “Notes on the birds of the Grand Pre region King’s County, Nova Scotia” Trans- actions of the Nova Scotia Institute of Science 14: 154-199 (1917): Tufts to Taverner, 3 March 1918, CMN. . The Auk 37: 147-149 (1920). . The Wilson Bulletin Number 112: 101 (1920). . Robert Curry “In Memoriam: George Webster North (1910-1983)” Ontario Birds 2 (1984). . W. Earl Godfrey Introduction to Taverner’s Birds of Eastern Canada Coles Canadiana Collection (Toronto 1974). . Taverner to Fleming, 24 March 1917, ROM. . P. A. Taverner Birds of Eastern Canada 177 (1919); 187 (1922). In the Coles edition entitled Taverner’s Birds of Eastern Canada (1974) the species Lincoln’s Sparrow is also shown on page 187. P. A. Taverner Birds of Eastern Canada 168 (1919); page 178 (1922). In his Bird Journal of August 1906 Taverner noted his attempt to translate a goldfinch’s song in human words: “I interpreted one of the little chatters of the goldfinch and think that I got it pretty close. It was flying over and as it passed it said plain- ly — “sweet, sweet, sweet, chew it, chew it”. It is my maiden attempt”, ROM. This was Arthur W. Andrews the amateur entomolo- gist in Detroit. Chapter 4, note 127 reference. W.J. Cody and B. Boivin “The Canadian Field- Naturalist and its Predecessors” The Canadian-Field Naturalist 68: 127-132 (1954); this is about the Transactions of the Ottawa Field-Naturalist Club 1880-1887 (2 volumes); and The Ottawa Naturalist 1887-1919 (32 volumes). Transactions were largely speeches and reports delivered at soirees, it was not a journal; D.F. Brunton “Additions to the Docu- mentation of the Publication History of the Canadian Field-Naturalist and its Predecessors”. Canadian Field-Naturalist 100: 423-426 (1986). Taverner to R. B. Horsfall, 14 March 1916, CMN. Taverner to Fleming, 10 August 1918, ROM. “A few of us” referred to “the gang” consisting of Taverner, Jim (J.M.) Macoun, and Clyde Patch. They each guaranteed a number of new memberships to back up the argument for this change (and reduce opposition on financial grounds). J. M. Macoun underwrote 100 memberships; Taverner underwrote 25; Patch under- wrote 25. . “Foreword” The Ottawa Naturalist 32: | (April 1918). . Taverner to F.L. Farley, 10 December 1918, CMN. . Taverner to R. W. Tufts, 11 December 1918, CMN. . Tufts to Taverner, 20 December 1918, CMN. By March 1919 the membership list had doubled from March 1918 to 540. Taverner had added 35 new members by then. . Taverner to Swales, 28 December 1918, CMN. . Taverner to Farley, 1 January 1919, CMN. . The new title came into effect with the publication of Volume 33, number | in April 1919. CRANMER-BYNG: A LIFE WITH BIRDS 78. 79. 80. 84. 85. 86. 87. Cha 1 BWP . H.M. Laing 223 Taverner to Fleming, 18 March 1919, ROM. The change was “moved by Mr. J.M. Macoun, seconded by Mr. Taverner, that the name of the Club publica- tion be changed to the Canadian Field-Naturalist”. Fleming to Taverner, 24 March 1919, ROM. Taverner to Fleming, 10 April 1919, ROM. When Taverner wrote that a general index would be a good thing, and that he thought he saw this in the future, he was very farsighted. It was finally published in 1979- 1980. (I am grateful to Daniel Brunton for this infor- mation.) . The Canadian Field-Naturalist 33: 1 (April 1919). The jacket was designed by the Geological Survey artist, C. E. Johnson. . The Ottawa Naturalist 32: 118-126 (1919). . Letter Taverner to Fleming, 11 April 1917, ROM, in which he described the new martin house which he was about to put up. “Artistically”, he wrote, “I think it is a success we will see what the Martins think about it.” I wish to thank Gerry Bennett of the Richmond Hill Naturalists for his gift to me of a copy of Taverner’s article printed in the form of a 15 page pamphlet by National Parks branch (3rd edition 1922). Taverner to Fleming, 23 August 1918, ROM. Taverner “The Summer Birds of Hazelton BC” The Condor 21: 80-86 (1919). The Auk 36: 1 (1919). P. B. Philipp and B.S. Bowdish “Further Notes on New Brunswick Birds”; J.A. Munro “Notes on some Birds of the Okanagan Valley, British Columbia”; P. A. Taverner “The Birds of the Red Deer River, Alberta” 36 (2): 248-265. pter 11 . Taverner to Fleming, 16 March 1920, ROM. Fleming replied the following day, and said (in part): “Curious our ideas should coincide about each others letters. I have kept yours carefully. Someday we will edit them and fail to find a publisher”. . J.D. Soper to Taverner, 13 January 1915, CMN. . Taverner to Soper, 18 January 1915, CMN. . P. A. Taverner “Suggestions for Ornithological Work in Canada” The Ottawa Naturalist 29: 14-18; 21-28 (1915). Also Chapter 7 above, notes 1-15. . Soper to Taverner, 6 February 1917, CMN. . Taverner to Soper, 18 February 1917, CMN. . Soper to Taverner, 23 November 1918; also 15 February 1919, CMN. . Fleming to Taverner, 16 December 1918, ROM. Laing’s first visit to Fleming was in mid-June 1918. Richard Mackie Hamilton Mack Laing: Hunter- Naturalist (Victoria, British Columbia, 1985) Chapter 6. (Hereafter Mackie Laing: Hunter-Naturalist.) . Mackie Laing: Hunter-Naturalist pages 70-72. (See note 8) “Lake-shore Bird Migration at Beamsville, Ontario” The Canadian Field-Naturalist 34: 21-26 (1920). . P.A. Taverner “Hamilton M. Laing” The Canadian Field-Naturalist 33: 99-100 (1919). . Laing to Taverner, 18 August 1919, CMN. . Laing to Taverner, 19 June 1919, CMN. For example Taverner to Laing, 22 March 1921, CMN. . Foster Working for Wildlife pages 161-163; 175-176. (See chapter 6 note 9) 224 16. 35! THE CANADIAN FIELD-NATURALIST Farley to Anderson, 18 November 1918, CMN. Francis La Grange Farley, born St. Thomas, Ontario 24 February 1870. Came to Red Deer, Alberta, in 1892 where he homesteaded before moving to Camrose, Alberta, in 1907. Lived there for the rest of his life. With encouragement from W.E. Saunders, Anderson and particularly Taverner he became an authority on the birds of Alberta. Obituary by J.D. Soper Auk 67: 218 (1950); see also W.K. Salt and A.L. Wilk The Birds of Alberta (Edmonton, 1958) page 51. . Taverner to Farley, 2 December 1918, CMN. . Farley to Taverner, 6 December 1918, CMN. . Taverner to Farley, 10 December 1918, CMN. . Farley to Taverner, 22 December 1918, CMN. . Farley to Taverner, 13 March 1919, CMN. 2. Taverner to Farley, 19 March 1919, CMN. . Taverner to Farley, 28 April 1919, CMN. . W. Rowan to Taverner, 21 November 1919, CMN. On Rowan see Marianne Ainley “Canada’s First Avian Biologist” Picoides I: 6-8 (1987). He met Taverner and Hoyes Lloyd on 20 September 1920 in Edmonton. See Rowan’s journal for evenings of 20 and 21 September. . Taverner to Rowan, 29 October 1920, CMN. 6. Fleming to Taverner, 20 January 1920, ROM. I have been unable to find any letter from Taverner to Fleming on whether he had any ambition to become chief of the Biological Division. 27. Taverner to Fleming, 23 January 1920, ROM. . Zaslow Reading the Rocks 353-357. (See chapter 5 note 1) . Taverner to Fleming, 18 February 1920, ROM. See also Lovat Dickson The Museum Makers: The Story of the Royal Ontario Museum (Toronto 1986) 36-39. . Taverner to Fleming, | March 1920, ROM. . Taverner to Fleming, 16 March 1920, ROM. . Taverner to Fleming, 22 March 1920, ROM. For information on Anderson’s career to this point see Chapter 14 below. . Taverner to Fleming, 24 March 1920, last paragraph, ROM. . Taverner to Fleming, 25 March 1920, ROM. Collins was Director of the Geological Survey from 1920- 1936, and concurrently acting Director of the Museum from 1925-1936. William Henry Collins b. 1878; graduate University of Toronto 1904; appoint- ed to Geological Survey 1905; married 1908. His home on Rosedale Avenue was built to a design by Taverner in 1913-1914. Personal communication from Mrs. Ann Whitmore, daughter of W. H. Collins. Taverner to Fleming, 16 April 1920, ROM. Taverner’s optimism was based on a letter from H. H. Mitchell when he was planning a trip to the prairies the previous year. . Taverner to Fleming, 30 April 1920, ROM. . H.H. Mitchell’s report as Provincial Naturalist in F. Bradshaw’s report as Chief Game Warden in “Sixteenth Annual Report of the Department of the Province of Saskatchewan for year ending 30 April 1921. Regina 1921. . Canada. Department of Mines Annual Report Year Ended 1921 page 27. . Percy Taverner to Mrs. Ida V.C. Taverner and Miss I. Taverner, 6 August 1920 (written from Elkwater Lake, Cypress Hill, Alberta), CMN. 40. 41. 42. 46. 47. 48. 49. 50. Sil Vol. 110 Taverner to his mother and sister, 7 August to 15 September 1920, CMN. Taverner to his mother and sister, 12-15 September 1920, CMN. Two anecdotes about Taverner on this prairie trip in the model T Ford were told me by Elizabeth Lloyd, who heard them from her father. While Hoyes Lloyd was driving the Model T Ford in the prairies, with Taverner in the passenger’s seat, they began approaching a level crossing. Each agreed to look out in case of an oncoming train on his side of the tracks. Hoyes Lloyd called out “all clear”, while Taverner was stuck on the sound “heh-heh-heh” as a train approached on his side. Lloyd sensed there was something wrong by the urgency in Taverner’s voice, and brought the car to a sudden stop just as the train rumbled over the crossing, and Taverner shouted “heh-heh-here she comes Lloyd”. One other story relayed by Elizabeth Lloyd about her father and Taverner on a camping trip is worth retelling. While Lloyd drove the Ford, cranked it by hand, and saw to its needs, Taverner did the camp cooking, which included buying food and milk. One evening they camped near a farm which Lloyd had visited previously. He knew that the farmer stuttered, and realized that if Taverner went to the farm to buy milk, and the two men started to talk, there might be trou- ble, even a fight. So he made an excuse for going to buy the milk himself, leaving Taverner to guard the Ford car and their equipment. William Rowan “Field Notes” 20 September 1920, University of Alberta Archives, William Rowan Papers Number 69-16-761. I am grateful to Dr. Marianne Ainley for giving me this reference. . Taverner to Fleming, 28 October 1920, ROM. . Fleming to Taverner, 31 October 1920, ROM. . Fleming to Taverner, 21 November 1920, ROM. The first three pages of this letter read rather like a film script. Because of his anxiety over security Fleming had not written to Taverner while he was in Manitoba “in case the letter went astray”, and it was left to Hoyes Lloyd to explain the scheme to him when they met at Banff in July 1920. Taverner to Fleming, 29 November 1920, ROM. Zaslow Reading the Rocks (see chapter 5, note 1) page 353 right column to page 354 left column, with a photograph of William McInnes. Zaslow Reading the Rocks (see chapter 5 note 1) pages 354-357. In these pages Morris Zaslow gives some perceptive insights into the effects of the subor- dination of the museum under the Survey. Taverner to Fleming, 10 April 1919, ROM. Nevertheless Brooks was very conscious of the valu- able time he had lost during the war. When Taverner and Fleming tried to persuade him to join them at the AOU meeting in New York he declined because, he explained, “I have lost 5 precious years — some thing, at my time of life. | am going to quit rifle shooting, big game, everything but bird work for the future ...”. (Brooks to Taverner, 13 October 1919, CMN.) Brooks to Taverner, 25 September 1919, CMN. Taverner to Brooks, 6 October 1919, CMN. Orni- thologists needed to have sets of birds in various plumages from downy young to fully mature male and female specimens in pre-nuptial moult, breeding 1996 Se a 58. Sy). 60. 61. 62. 63. 64. plumage, post-nuptial moult and winter plumage. Without such a range available for comparisons ornithologists could make misidentifications. See Christopher Leahy The Birdwatcher’s Companion pages 580-584. Bonanza Books, New York, 1982. P. A. Taverner “The Evening Grosbeak in Canada” The Canadian Field-Naturalist 35: 42-45 (1921). Taverner discussed a western and an eastern form, and attributed the spread of Evening Grosbeaks east of Lake Superior to the “recent” spread of the Mani- toba Maple (Acer negundo Linnaeus), the winged seeds of which the birds ate. This article was referred to 65 years later in the Atlas of the Breeding Birds of Ontario (University of Waterloo Press, 1987) page 502. Taverner to Brooks, 24 October 1919, CMN. In the same letter he said “If you leave publication altogeth- er to the half baked you cannot complain when mis- takes creep in. I think it is up to you to correct many of the popular, or the unpopular British Columbia mistakes.” . Brooks to Taverner, October 28 1919, CMN. . Taverner to Brooks, 6 November 1919, CMN. . Taverner to Brooks, 30 September 1921, CMN. . Taverner to Brooks, 6 December 1922, CMN. When Taverner wrote “111 Stellar’s Jays’ he did not mean that the National Museum had that number of speci- mens in its trays. The majority would be on loan from collectors such as Brooks and from other museums. For subspecies of Stellar’s Jays in the 1920s see Taverner Birds of Western Canada pages 257-258; for the 1980s see Godfrey The Birds of Canada (1986) page 382. What Taverner meant by saying that “too many peo- ple are guided by specific preconceptions” probably refers to people who expect a subspecies to be found in geographical areas, and therefore call specimens taken from that area what they expect them to be. They do not face the facts of a careful study of a specimen’s measurements and plumage and compar- isons with a series of other specimens. Brooks to Taverner, 17 December 1922, CMN. Taverner to Brooks, 27 December 1922, CMN. “Slightly colour-blind” is not a meaningful term. Perhaps a more accurate measure of colour percep- tion would be to use relative terms such as below average, average, above average. Taverner compounded his challenge by writing a let- ter to the Editor of The Auk enclosing a copy of the statement entitled “The Genus Debased” together with the signatures of 28 leading systematic research workers of the A.O.U. Some minor revisions were suggested by four of these. Also sent to the editor was a copy of Taverner’s covering letters accompa- nying the statement “The Genus Debased”. All three letters were published in The Auk in the Correspon- dence section, under the signature of P. A. Taverner. The Auk 40: 177-80 (1923). Brooks to Taverner, 17 December 1922, CMN. This was almost one year after Taverner had circularized his statement to leading systematic research workers of the AOU. Mackie Laing: Hunter-Naturalist (see note 8) pages 80-81. See L. B. Potter “Notes on Birds in Southwestern Saskatchewan” The Canadian Field-Naturalist 36: CRANMER-BYNG: A LIFE WITH BIRDS 65. 66. 67. 68. 69. 70. ile We. Ue 74. i). 76. le 78. Ds) 94-95 (1922). This is a short notice on the valley of the Frenchman River, and the Sage Grouse, Rosy Finches, and wintering Horned Lark he had observed there. Mackie Laing: Hunter-Naturalist (see note 8) page 81-82; Taverner to Rowan, 21 October 1921, CMN. Taverner to Laing, 20 September 1921, CMN. Taverner and Brooks worked out a draft list of birds they considered should be illustrated in Birds of Western Canada while Taverner was visiting Brooks. Marjorie Brooks “Allan Brooks — A Biography” The Condor 40: 16 (1938). Taverner to Brooks, 8 December 1921, CMN. Brooks asked Taverner to draw up plans for a building 25 x 18 yards that could be easily heated, have a working window to give a good northern light, a work bench, and windows to face N.E. and E. (Brooks to Taverner, 16 January 1922). Taverner sent the plans of the building, with specifications, on 3 February 1922. Brooks to Taverner, 3 August 1921, CMN. Taverner to Brooks, 30 September 1921, CMN. He reported that Anderson was still on an expedition col- lecting in New Brunswick with Young. Harry Schelwald Swarth (1878-1935). Fellow of the AOU. Curator of Birds, Museum of Vertebrate Zoology, Berkely, California 1908-1913, 1916-1927; Curator Department of Ornithology and Mammology, California Academy of Sciences, San Francisco 1927-1935. Obituary in The Auk 54: 127-134 (1937). The Skeena River rises in the northern interior of British Columbia, flows S.W. for 58 km and into the Pacific Ocean at Prince Rupert. Brooks to Taverner, 26 November 1921, CMN. In his letter Brooks copied verbatim the words Swarth used about Brooks’ museum. Taverner to Brooks, 8 December 1921, CMN. The Minister of Mines from 1920-1921 was Sir James A. Lougheed, lawyer and Conservative leader in the Senate from 1906-1921. Taverner to Brooks, 17 February 1922, CMN. There was little improvement, if any, during Taverner’s lifetime. Taverner to Laing, 7 December 1921, CMN. H. M. Laing Allan Brooks: Artist Naturalist (British Columbia Provincial Museum, 1979) page 2. Laing described collecting in 1922 with Brooks and Taverner in Chapter | “The Okanagan”. Laing Allan Brooks pages 3-4: Taverner Birds of Western Canada, Page 241 on the White-throated Swift; Godfrey The Birds of Canada (1986) on the distribution of the White-throated Swift in the 1980s. Taverner to Mrs. Ida V.C. Taverner, late May 1922, CMN. For a good photo of Brooks, Taverner, Farley, Laing, Gartell, Sampson at Vaseux Lake, see Laing Allan Brooks (note 76) page 4. . Taverner to his mother, 4 June 1922, CMN. . Mackie Laing: Hunter-Naturalist 85. (See note 8) . Taverner Birds of Western Canada 1926 pages 208- 209. . Laing Allan Brooks page 6. (See note 76) . For a fuller account see Laing Allan Brooks pages 5-6. . At Brooks’ home Taverner was able to look at his collection of skins in the specimen cabinets which, 226 85. 86. 87. 88. 89. 90. 91. 92. 93: THE CANADIAN FIELD-NATURALIST according to Laing, showed the stamp of perfection on every tray. A photo taken around 1924 shows three trays of specimens, and the caption “The Brooks Collection set high standards in the quality and uniformity of its specimens”. From Laing Allan Brooks pages 158; 160. (See note 76) On Harkin’s philosophy for wildlife preservation see Foster Working for Wildlife pages 216-219; 222. (See chapter 6 note 9) Taverner to Mrs. Ida V.C. Taverner, 4 June 1922, CMN. Taverner to his mother, 17 June. McAstocken was an engineer on the Kettle ValleyRailroad and offered to take Taverner on the engine to Hope, which McAstocken said was “a very wonderful bit of rail- road run”. A cayuse was a western Indian pony. This was Dr. Charles Saunders of the Experimental Farm. The man referred to was J. H. Parham, an expatriate Englishman and gentleman fruit-rancher. Taverner was to hear from him again in the 1930s. Phalacrocorax pelagicus. Specimen no. 123. The species was described by Taverner in Birds of Western Canada pages 72-73. In 1909 Seton published Life Histories of North American Animals, in 2 volumes, about the fifty-nine species he had found in Manitoba. Later he began revising and expanding these volumes with much new information. This was his main occupation in 1922 when he visited Taverner at the National Museum. The main theme of his lectures now was on the need for conservation of wild creatures. The first volume of his new work Lives of Game Animals was published in 1925. Taverner and Seton had much in common to discuss. Laing to Taverner, 6 January 1923, CMN. “A beaut” — slang. Abbreviation for “a beauty”. Laing to Taverner, 6 January 1923, CMN. Taverner to Laing, 3 February 1923, CMN. Chapter 12 ii Taverner to Brooks, 6 February 1922, CMN. 2. Taverner to Brooks, 27 September 1922, CMN. 3. Taverner to Brooks, 14 March 1923, CMN. In Birds of Eastern Canada a paragraph was given under the heading Distinctions while a few lines were given to Field Marks. As an example see Bonaparte’s Gull and how to distinguish it from other eastern gulls, and from the [Black-legged] Kittiwake. Under Field Marks he wrote “Size, black hood in summer adults, white on forward edge of wings, and bill and feet colour on other plumages make the most valuable field characters for recognition in life” (page 54). In Birds of Western Canada a good example is Franklin’s Gull where the same plan is followed, and a paragraph is given on how to distinguish it from the Bonaparte’s Gull. . Louis A. Fuertes to Taverner, 27 January 1924, CMN. In a two-page letter he enclosed sketches and tracings from some of the material in his portfolio. On a separate sheet he made drawings “as the out- lines seem to me’. But these, he said, were mere sug- gestions, and added: “You said a lot when you remarked that no two people see alike, nor fasten on the same details as field-recognition marks. Its the ‘cut of the jib’, and a very subtle thing, that enables EONS COk IEG Vol. 110 the old hand to spot species of hawks, or ducks ... it shouldn’t be expected that outlines, or the written word, will take the place of that good old teacher — hard work and experience.” . Taverner to Brooks, 5 February 1924, CMN. . Taverner to Laing, 12 February 1924, CMN. . Taverner to Brooks, 14 March 1923, CMN. . Taverner to Brooks, 3 August 1923, CMN. . Taverner to Brooks, 2 September 1923, CMN. . Brooks to Taverner, 5 January 1923, CMN. . Taverner to Brooks, 16 October 1923, CMN. Brooks had painted a Marbled Godwit in breeding plumage. Taverner said that the base of the beak should be bright orange, and the legs almost black. He sent the plate back for alterations which Brooks carried out. . Brooks to Taverner, 25 November 1924, CMN. (Plate XXXI by Brooks’ reference). This contains hand written notes by Brooks on suggested changes, and additions. Under Ruby-crowned Kinglet Brooks wrote, “I find the interrupted eye-ring of this bird an arresting field mark”. He illustrated this with a minia- ture pencil-drawing of the bird’s head showing the interrupted eye-ring clearly. In just a quick sketch Brooks achieved an unmistakable kinglet head. (Brooks reference number 989.) Brooks’ note of praise for Taverner’s “habit-portraits” comes at the end of this part. . Taverner to Brooks, 22 February 1924, CMN. . Taverner to Brooks, 3 August 1923, CMN. . Taverner to Brooks, 16 October 1923, CMN. “Sot”, a word sometimes used by English people as a joke on the Scots pronunciation of the word “set”. . Brooks to Taverner, 8 January 1924, CMN. . Taverner to L.L. Snyder, 12 February 1924, CMN. The Third AOU Check-list of 1910 contained four species and fourteen subspecies of the genus Junco. . For an example of his sense of fun see Taverner’s let- ters to Laing of 25 November and 18 December 1924, CMN, on the subject of government accoun- tants. Laing had submitted his accounts for the sum- mer of 1924 somewhat late, with a letter of explana- tion to Taverner. Taverner replied that it was no good sending him a letter of explanation since he had no pull with the accountant, and continued: “Accountants are important fellows. I think they are stuffed with red-tape inside instead of the usual guts”. . Taverner to Brooks, 26 November 1923, CMN. . Marianne G. Ainley “William Rowan: Canada’s First Avian Biologist” Picoides 1: 6-8 (1987). . Rowan to Taverner, 13 March 1922, CMN. . Taverner to Rowan, 23 August 1922, CMN. . Rowan to Taverner, 8 November 1922, CMN. . Taverner to Rowan, 15 May 1922, CMN. . Rowan to Taverner, 8 March 1922, CMN. On the subject of introduced birds Rowan wrote: “Thence onwards I hope to keep a systematic cheque on fur- ther movements ...” . . Rowan to Taverner, 23 January 1923, CMN. . Taverner to Rowan, 2 February 1923, CMN. . Rowan to Taverner, 21 February 1923, CMN. . A battle of words in The Canadian Field-Naturalist, 1923-1924, was sparked by J. A. Munro’s article “The Necessity for Vermin Control on Bird Sanctuaries”, followed by an editorial comment from Harrison Lewis, and “letters to the editor”. See Chapter 15, note 72. 1996 30. 36. iE 38. 43. Taverner to Rowan, 19 January 1924, CMN. Presumably the movement for wildlife preservation at that time. . Taverner to Rowan, 17 March 1924, CMN. . Rowan to Taverner, 27 March 1924, CMN. . Lloyd to Taverner, no date, probably early April 1924, CMN. . Taverner to Rowan, 21 April 1924, CMN. . Taverner to L.L. Bolton, 31 December 1925, CMN. Bolton was assistant deputy minister, Department of Mines, with F.J. Nicholas, editor of publications of the Geological Survey Branch under him. Zaslow, Reading the Rocks page 353. (See chapter 5, note 1) Taverner to Brooks, 31 December 1925, CMN. “Seance” here in the sense of “a meeting for exhibi- tion or investigation of spiritual phenomena” which shows Taverner’s sceptical sense of humour. Brooks to Taverner, 23 May 1925, CMN. Taverner to Brooks, 18 November 1925, CMN. Audubon died in 1862. His widow, Lucy, had little money left and offered most of his original drawings to the New York Historical Society for five thousand dollars, but eventually accepted two thousand, which was more than anyone else would offer her. In this way the Society preserved the originals for future generations to see. Information in Alexander Adams John James Audubon: a Biography (New York and Toronto 1966) page 470. . Brooks to Taverner, 18 January 1926, CMN. . Taverner to Brooks, 18 January 1926, CMN. . Taverner to Brooks, 13 March 1926, CMN. In the same letter Taverner made suggestions for improving the [American] Golden Plover illustration. . Taverner to Brooks, 22 February 1924, CMN; Taverner to Rowan, 22 February 1924, CMN. The Biological Board of Canada was established in 1912; it ran permanent biological stations at St. Andrews, New Brunswick and Nanaimo, British Columbia. In 1937 the Biological Board became the Fisheries Research Board. This manual, under various titles, was to engage Taverner’s attention, from time to time, until the end of his life. Taverner to Fleming, 25 August 1924, ROM. Also Taverner to Rowan in which he related the trip, and discussed cameras. . Taverner to Tufts, 19 April 1924, CMN. . Tufts to Taverner, 12 May 1924, CMN. This was a hard hitting letter. Although Taverner put as good a face on it as possible Tufts’ injured feelings were only “somewhat” soothed. For a outline of the latter’s career see W. Earl Godfrey, 1984. A tribute to Robie Wilfred Tufts, 1884-1982. Canadian Field-Naturalist 98(4): 513-518. . Taverner to Tufts, 11 June 1924, CMN. . Taverner to Fleming, 25 August 1924, ROM. . Taverner to Fleming, 23 September 1924, ROM. . Taverner to Brooks, 17 September 1924, CMN. . Brooks to Taverner, 24 September 1924, CMN. . Iam grateful to Daniel Brunton of Ottawa for this information. . A photograph of their house in Detroit taken in 1906 exists, and one of their house in Ottawa when it was almost complete in the fall of 1912. A coloured photo of the same house taken from much the same angle in 1988 shows no'‘major change. CRANMER-BYNG: A LIFE WITH BIRDS 53 54. 55. 59) 60. 61. 62. 69. 70. ile UW 73% A . Taverner to Brooks, 6 December 1922, CMN; Taverner to Laing, 28 December 1922, CMN. I am grateful to Miss Elizabeth Lloyd for giving me a copy of this photograph. The date 22 August 1923 is written on it in ink. Note by Ida Taverner in Guest Book at Hyla, Blue Sea Lake Book I (1917-1924) dated 30 October 1923. Property of Corwin Ferguson who kindly loaned it to me. . Brooks to Taverner, 26 November 1922, CMN. . Taverner to Laing, 12 January 1926, CMN. . Taverner to Laing, 11 December 1925, CMN. On this expedition see H.M. Laing “Birds Collected and Observed During the Cruise of the Thiepval in the North Pacific, 1924” in Canada, National Museum Bulletin Number 40, 1925; Mackie Hamilton Laing pages 88-96. (See chapter 11 note 8) On H. S. Swarth see Chapter 11 note 71. Atlin Lake is at the northern boundary of British Columbia and Alaska. See Laing Allan Brooks pages 148-156. (See chapter 11 note 8) It was given the common name of Timberline Sparrow and described by Swarth and Brooks in The Condor 27: 67 (1925). Its range was given as breed- ing at high altitudes in the Atlin district of northwest- ern B.C. Also as occurring in southeastern B.C., and Madison County, Montana, in migration. A.C. Brooks and H. S. Swarth “A Distributional List of the Birds of British Columbia” Pacific Coast Avifauna 17: 1-58 (1925), Cooper Ornithological Club; Taverner to Brooks, 15 December 1925, CMN. Taverner to Brooks, 25 November 1924, CMN. In the same letter he encouraged Brooks to paint larger pic- tures once in a while, and added “Size may not impress the discriminating critic but it does the mass- es and it is the masses who buy.” . Taverner to Fuertes, 25 November 1924, CMN. . Fuertes to Taverner, 10 December 1924, CMN. . Brooks to Taverner, 6 May 1925, CMN. . Taverner to Brooks, 13 May 1925, CMN. . Brooks to Taverner, 4 June 1925, CMN. . Taverner to Rowan, 17 September 1924, CMN. For Rowan’s notes on birds found at Beaverhills Lake 1924-1925 see Robert Lister The Birds and Birders of Beaverhills Lake (Edmonton, 1979) pages 51-60. Taverner got valuable records for use in Birds of Western Canada from Rowan in these years. Rowan to Taverner, 13 February 1924, CMN. Marianne Gosztonyi Ainley From Natural History to Avian Biology: Canadian Ornithology 1860-1950 222-225. Thesis submitted in partial fulfillment of the requirements of the degree of Doctor of Philosophy McGill University 1985. Gustave Eifrig “Is Photoperiodism a factor in the migration of birds?” The Auk 61: 439-444 (1924). Rowan to Taverner, 14 February 1925, CMN. See Marianne Ainley “Rowan vs Tory: Conflicting Views of Scientific Research in Canada, 1920-1935” Journal of History of Canadian Science, Technology and Medicine 12(1): 11-12 (1988) on difficulties caused by Tory’s attitude to his ornithological research. Cyril Guy Harrold (1895-1929). Obituary, with pho- tograph, by B. W. Cartwright and A. G. Lawrence The Canadian Field-Naturalist 43: 132-133 (1929); Robert Lister The Birds and Birders of Beaverhills 228 74. Tey 76. ies 78. 19: 80. 81. 83. 84. 85. THE CANADIAN FIELD-NATURALIST Lake (Edmonton 1979) pages 31-34; 44-68 (quoting from Rowan’s field notes). In 1925 Harrold was paid the minimum rate for a junior zoologist which was $105 per month, until it was raised to $110. Anderson to Taverner, August 1925, CMN. The name of the lake was spelt as Beaverhills by Rowan, Lister and others. The name had been changed to Beaverhill — singular — before Lister’s book was published in 1979. Taverner to Ida Clare Taverner, 15 August 1925, ROM. (written from Red Deer River). Sternberg had a long and distinguished career as a paleontologist at the National Museum of Canada: see tribute by Loris S. Russell. 1982. “Charles Mortan Sternberg, 1885- 1981”. The Canadian Field-Naturalist 96(4): 483- 486 and bibliography compiled by Richard Gordon Day The Canadian Field-Naturalist 96(4): 487-489. Lister Beaverhills Lake (see note 73) page 165-166, quoting Rowan’s field notes. Lister (see note 73), page 166, but quoting Lister’s own words. Rowan, talk at Calgary 1955. Copy of relevant text kindly sent to me by Marianne Ainley. Taverner to Ida Clare, 12 September 1925, ROM. Siwash. A derogatory term 1) as noun. North American Indian 2) as verb. To camp out, travelling light and using only natural shelter. H.M. Laing The Canadian Alpine Journal 15: 99- 114 (1925); H.M. Laing, P.A. Taverner, R.M. Anderson “Birds and Mammals of the Mount Logan Expedition, 1925” Canada, National Museum Annual Report for 1927, pages 69-71; H. M. Laing and P. A. Taverner “Notes on Birds collected and observed in Chitina River Region, Alaska” Annual Report for 1927, pages 72-95. Total number of birds collected was 203. Laing shot nearly 4000 feet of film which was edited by Allen Carpe, representative of the American Alpine Club, into a film The Conquest of Mount Logan. On the 1992 Mount Logan expedition see article by Michael Schmidt, expedition leader, with coloured photographs by Pat Morrow, entitled “To the Top” Canadian Geographic Sept/Oct 1992, pages 22-35. The text reads well, the photos are superb. Canada Department of Mines, National Museum Annual Report for 1925, page 45. Taverner to Bishop, 23 October 1925; 18 November 1925, CMN. . Brooks to Taverner, 18 January 1926, CMN. The article was by W.H. A. Preece “January Bird Notes from Mount Tolmie, Victoria, B.C.” 39: 175-176 (1925). The fact that House Wren and Chipping Sparrow were listed for January but no mention was made of Seattle Wren [Bewick’s Wren] and Tree Sparrow tells its own tale. Taverner to Brooks, 10 February 1926, CMN. The Editors of the journal were Harrison Lewis 1922- 1925; F.J. Nicholas 1925; G.A. Miller 1925-1928; Douglas Leechman 1928-1938. Taverner was an Associate Editor (for ornithology) until 1942, when he was succeeded by A. L. Rand. Taverner to Camsell, 5 December 1922; 6 January 1923, CMN. Joyce Reddoch “Dow’s Swamp” Trail and Land- scape 12: 133-141 (1978), Ottawa. I am grateful to Danie! Brunton for bringing this article to my atten- 86. 87. 88. 89. 90. ik. 9D. 93° 94 95 Ch 1 Mn & WwW 6. Vol. 110 tion, and for supplying information from his own knowledge on the early history of the Ottawa Field Naturalists’ Club. See also C. H. D. Clarke. 1965. “A | requiem for Dow’s Swamp”. Canadian Field- Naturalist 79(1): 1-3. Gustave Langelier to Taverner, 7 May 1923, CMN. Taverner to Bernadette Langelier, 18 June 1923, CMN. Taverner said that this was a considerable extension of range for that species “as the nearest Canadian record I have for it is Montreal where it is only accidental”. A sight record, he said, “would always have left a strong element of doubt in the question”. Taverner to Gustave Langelier, 17 October 1923, CMN. Taverner to Langelier, 4 December 1925, CMN. Taverner to Bent, 15 March 1922, CMN. Taverner said that he would need to see various species in vari- ous plumages at the museum, and also discuss the distribution of species in Canada with Taverner’s maps and files in front of him. In a letter of 22 March 1922 Taverner told Bent that he had about half the Canadian species plotted. Bent to Taverner, 20 March 1922, CMN. During 1923 Taverner wrote 12 letters to Bent. P. A. Taverner “Ornithological Investigations near Belvedere, Alberta, 1926”, Canada Department of Mines, National Museum Annual Report 1926 (1928) pages 84-104. This is a list of birds observed, with brief annotations. The Connecticut Warbler was noted as a common breeding bird; several nests were found. For a description of the work there see letter Taverner to Bishop, 5 August 1926, CMN. A one-page account by H.E. Mullet appeared in The Edmonton Journal 24 July 1926. It was reproduced, in part, by Lister Birds of Beaverhills Lake pages 61- 66. Rowan wrote his own account to explain the rea- sons for the mass banding, and the difficulties the banders experienced in William Rowan “Banding Franklin’s Gulls in Alberta” The Wilson Bulletin 39: 44-49, 1927. . Anderson to Taverner, 19 June 1926, CMN. . Taverner to Fleming, 19 March 1926, ROM. apter 13 . T.S. Palmer “The Forty-Fourth Stated Meeting of the American Ornithologists’ Union” The Auk 44: 73-84 (1927). Hereafter “T.S. Palmer Report”. It includes a full transcript of the programme of events. The Honourable Charles Stewart M.P. was both Minister of Mines and the Interior from 1921-1936 with the exception of a short period in 1926. . Hoyes Lloyd “The 44th Stated Meeting of the American Ornithologists’ Union” The Canadian Field-Naturalist 40: 189-191 (1926). Hereafter “H. Lloyd. Report on meeting”. . T.S. Palmer “Report” (see note 1) pages 74; 84. . T.S. Palmer “Report” (see note 1) pages 74-75. . T.S. Palmer “Report” (see note 1) page 75, which was accompanied by a plate (IV) showing the Great Auk perched on the three volumes. T. S. Palmer “Report” (see note 1), pages 79-82, lists all the papers presented by title, together with the name and address of the speaker and his institution. . Ruthven Deane’s collection of portraits of North American ornithologists numbered over one thou- 1996 sand, and was presented to the Library of Congress in 1934. It was deposited in the Division of Fine Arts, and was provided with a card index which gave basic data on each person. See In Memoriam notice for R. Dean in The Auk 52: 1-14 (1935). . These were listed, with a brief introduction, in a cata- logue entitled EXHIBITION OF BIRD ART Held in connection with the 44th Meeting of the AMERICAN ORNITHOLOGISTS’ UNION VICTORIA MEMORIAL MUSEUM For information on items under “Historical Collection loaned by the Emma Shearer Wood Library of Ornithology, McGill University Library, Montreal”, see that catalogue pages 2-3. . Exhibited in advance of the AOU meeting at a con- versazione of the Professional Institute of the Civil Service of Canada held in the museum in mid April 1926. More than eight hundred guests, from the tech- nical services of government and from both Houses of Parliament, with their wives, attended. . These formed part of a larger collection of Brooks’ paintings commissioned by Robb which were later presented by him to the Royal Ontario Museum. They are currently preserved in the Department of Ornithology there. They measure 9 X 12 inches and appear to have been painted in a rather dramatic style with noticeable colour effects in the background, as though designed as people pleasers. Or perhaps Brooks suited these paintings to Robb’s romantic nature. See page 10 of the catalogue (see note 8) of the exhibition where the name of each species is given. . This library was started by Dr. Casey Albert Wood (1856-1942) who had studied medicine at McGill University, Montreal. His first evidence of an interest in birds was a paper of his on the eye and eyesight of birds published in Ophthalmology in 1907. This work culminated in his book The fundus oculi of birds ... ; a study of comparative anatomy and physiology in folio, illustrated by 145 drawings and 61 coloured plates, 1917. Dr. Wood joined the AOU in 1917 and was elected a member in 1921. In 1920 he founded and endowed the Emma Shearer Wood Library of Ornithology, based on his own ornithological library, which he presented to McGill University. It was named in honour of his wife Emma Shearer of Montreal. For further informa- tion on Dr. Wood see S.L. McAtee The Auk 59: 611- 12 (1942); Eleanor MacLean History of the Blacker- Wood Library (typewritten, May 1988) 8 pages. As librarian of the Blacker-Wood , Eleanor MacLean has written an excellent account of its founding and con- tinual extention in the 1920s and 30s. I wish to thank Ms. MacLean for giving me a copy of her paper and for showing me round the Blacker-Wood library of Biology. Correspondence between Casey Wood and P.A. Taverner is preserved in the Blacker-Wood Library under Taverner, and in the Taverner papers at the National Museum, Ottawa under Casey Wood. . H. Lloyd “Report on meeting” page 190. (See note 2) . T.S. Palmer “Report” page 76. (See note 1) . A copy of the group photograph of 48 Canadian AOU members at the 1926 meeting in Ottawa is pre- served in the ROM Library, Archives section, togeth- CRANMER-BYNG: A LIFE WITH BIRDS D290) er with a numbered list of each member’s name, type- written. Title: “Key to the Canadian Group Photo- graph taken at the A.O.U. 1926 Convention, reading from left to right”. . The Auklet 2 (1926) page 14. . The Auklet (1926) page 16. . The Auklet (1926) page 15. William Edwin Saunders, “Will” or “W.E.” to his close friends, came from a family with interests in entomology, botany, horticulture and experimental agriculture. His father was largely responsible for the first Federal Experimental Farm being established in Ottawa in 1886, and was appointed Director. One brother, Charles, became famous for his plant-breeding exper- iments over many years which led him into develop- ing Marquis wheat. Another brother, Percy Saunders, a professor of chemistry, became an authority on peonies, irises and phlox. W.E. was the naturalist of the family. Taverner wrote a warm appreciation of him when he died. See The Auk 61: 345-351 (1944). . Brooks to Taverner, 28 July 1923, CMN. . Taverner to Brooks, 3 August 1923, CMN. Taverner was referring to The Canadian Field-Naturalist when he wrote the words “our badly needed subscription list”. Wallace Havelock Robb (1888-1976) was born in Belleville, Ontario, son of William Doig Robb, an official of the Grand Trunk Railway Company. He was interested in bird life from boyhood, and in 1921 he joined Herbert K. Job, the representative of the American Audubon Society, on a photographic expe- dition to the Magdalen Islands, in the Gulf of the St. Lawrence. The success of this trip diverted Robb away from business, in which he had done very well, and turned him towards poetry and birds. Robb could afford to pay a first-class wildlife artist to produce 32 bird paintings to exhibit at the AOU meeting in Ottawa, and in this way bring himself into the lime- light. He was ambitious to establish “a bigger and better and finer Bird-lore in Canada and run it in his own way” (See note 20). Robb’s first book of poems The Quill and the Candle; Poems of Birdland in Canada was published by Ryerson Press in 1927. Here is an example of a verse: ‘I dreamed of a Birdland Garden fair/With a lily lagoon and a dragon-fly;/And a mystical Heron was standing there/As I went by’. Here is a second: ‘A lone loon calls,/And rocky walls/Return the mournful sound./He grieves away The dying day; / And darkness creeps around.’ (Robb’s poetry was very similar to that known in England in the years ca. 1917-1926 as “late Georgian verse”. Critics scorned it as being “flabby” — it was slack in technique, false in its simplicity and shallow in feeling, among other weaknesses. There is no full biography of Robb, but George F.G. Stanley read a paper before the Kingston Historical Society at which the poet was present. See George Francis Gilman Stanley The abbe of Abbey Dawn. The Kingston Historical Society 1970. . Taverner to Brooks, 7 February 1925, CMN. . Robb to Taverner, mid-October 1926, CMN. One obvious cause of Robb’s complaint of the “derogato- ry style” of The Auklet was an item with the title “A Poem by Walter Hardrock Rod, Poet and Naturalist”. 230 Nw N THE CANADIAN FIELD-NATURALIST This poem was distinguished by the repetition ad nauseam of the refrain “Are you there?” “Are you there?” The Auklet 2 (October 1926) page 22. . Taverner to Robb, 21 October 1926. This letter shows Taverner as an experienced letter-writer. In two sen- tences he made clear the functions of The Auklet. It was to check AOU members from taking themselves too seriously, and becoming stuffy and pompous. Then, in two paragraphs, he drew an understanding sketch of Brooks — his nature, traits and character. Without directly criticizing Robb he was able to deflate him neatly. Whether Robb had enough per- ception to realize that his fulsome oratory in present- ing this commercially sponsored award to Brooks was irksome is impossible to tell. Taverner gave Robb just a glimmer of praise for his collection, but no more. 23. In 1926 this consisted of Kingswood, Mackenzie 24. 2) 26. 2: 28. 29) 30. Bil 82. King’s summer cottage beside Kingsmere Lake, built in 1903 while King was still a civil servant. In 1928 King moved to Moorside, the showplace of his estates. The names Kingsmere Lake and King Mountain were not named after Mackenzie King but were in use as early as 1905. Taverner to Laing, 23 October 1926, CMN. On Big Island see chapter 7, page 156. Taverner to Laing, 23 October 1926, CMN. Taverner to Thompson Seton, 21 October 1926, CMN. T. S. Palmer The Auk 44: 82-84 (1927). Taverner to Laing, 23 October 1926, CMN. Writing to Fleming on 28 October Taverner explained: “The smooth working of everything was due to Lloyd’s executive ability and Patch’s work. Most of the ideas were mine but I could not have carried them out so well had it not been for the able assistance I had from the above and the Department.” Rowan to Fleming, 3 October 1926, ROM. See M.G. Ainley “Rowan vs Tory: Conflicting Views of Scientific Research in Canada, 1920-1935” Scientia Canadensis 12 (1): 3-21 (1988). This article explains why Rowan thought that it was unwise for him to attend the AOU meeting. Taverner to Rowan, 22 October 1926, CMN. Also Rowan to Taverner, 2 November 1926 asking “Why take my joke about the Yankee Pie so seriously?” Taverner to Bishop, 21 October 1926, CMN. Kipling wrote this at the time of Queen Victoria’s Golden Jubilee celebration in 1897 to warn his fellow coun- trymen not to be boastful and drunk with the power of the British Empire. The third verse reads: ‘“Far- called, our navies melt away/ On dune and headland sinks the fire/ Lo, all our pomp of yesterday/ Is one with Nineveh and Tyre/ Judge of the Nations, spare us yet/ Lest we forget — lest we forget.” . H.S. Swarth The Condor 29: 84-85 (1927). . W. Stone The Auk 44: 126 (1927). . Anderson to Ada Johnson, 18 July 1927, NAC, Manuscript Division, MG 30 B40, R.M. Anderson Papers, Volume 4, page 3. . See Chapter 12 note 49. . Taverner to Ida Clare Taverner, 15 August 1925, CMN. . Taverner to Munro, 25 February 1927, CMN; Taverner to Bishop (same subject), 1 March 1927, CMN. 39. 40. 56. Sif: Vol. 110 Taverner to Bishop, 9 May 1927, CMN. Taverner to Fleming, undated, ROM. Fleming wrote on it in pencil “Aug 1927”. Taverner also mentioned a photograph of himself which he had taken for the old lady. . Taverner to Bishop, 16 August 1927, CMN. . Taverner to Fleming, 4 October 1927, ROM. . Taverner to Fleming. For a description of their visit to Yosemite National Park see Taverner to Laing, 25 November 1927, CMN. . From the fact that Taverner was staying with a rela- tive, and that his own name at birth was Fowler, we can deduce that his father had, at some point, married Mrs. Fowler. . Saunders to Fleming, 17 September 1924, ROM. . Laing to Taverner, 15 April 1926, CMN. . Taverner to Fleming, | April 1927, ROM. Also at the wedding was Richard S. Finnie, at that time starting his career of arctic explorer and writer, and Hughie Margetts of the RCMP who was Dewey Soper’s best man. All three shared a hair-raising experience during a voyage in C.G.S. Arctic, Captain Joseph E. Bernier, taking Soper to Pangnirtung, Baffin Island, in 1924. See Chapter 14. It was a great occasion for all three when they met again in the home of Percy and Ida Taverner in Ottawa. The bride was Carolyn Freeman of Wetaskiwin, near Edmonton. See Richard S. Finnie “In Memoriam: J. Dewey Soper 1893-1982”. The Musk-Ox 31: 80-82 (1982). University of Saskatchewan, Institute for Northern Studies, Publi- cation Number 2. . Taverner to Munro, 25 February 1927, CMN. . Taverner to Laing, 21 April 1927, CMN. . Taverner to Laing, 12 May 1928, CMN. . IT was in touch with David McLeish of Toronto and Beaumaris who is a relative of John McLeish. . Taverner to Fleming, 7 March 1930, ROM. . The Gopher, newspaper of the Liggett School, Volume 8,Number 7, 30 April 1930, page 3. For Martha Hohly Wiest see Chapter 4. . Taverner to Munro, 7 March 1930, CMN. . Laing to Taverner, 4 April 1930, CMN. The refer- ence to “on Dec. 9th last” is to a letter of that date which Taverner sent to Laing. He must have been feeling “down in the dumps” when he wrote it, because what he meant to be light- hearted banter had a jaundiced quality about it. I am grateful to Miss Elizabeth Lloyd for showing me a copy of the Taverner wedding card which her par- ents preserved. Information in letter Taverner to Brooks, 28 March 1930, CMN. Chapter 14 1. Shelagh D. Grant Sovereignty or Security? Govern- ment Policy in the Canadian North. (University of British Columbia Press, Vancouver 1988) pages 5-6. . See Zaslow, Reading the Rocks, pages 173-175. (See chapter 5 note 1) The expedition wintered at Cape Fullerton on the northwest side of Hudson Bay. In the summer of 1904 the Neptune explored in Baffin Bay, and landed members of the expedition at Cape Herschel, Ellesmere Island. Here a document, taking formal possession of it for the Dominion was read, and the Canadian flag raised and saluted. See photo- 1996 1D graph page 174, and map showing the routes of the Neptune, and places where stops were made. Collections of rocks and fossils were made, as well as of northern birds, eggs and nests, and biological material. In his “Summary Report for 1913” Taverner reported that the Zoological Division had received by exchange from the Department of Marine and Fisheries, the birds and eggs collected by A. P. Low during the voyage of the S.S. Neptune in 1904. . See Zaslow, The Opening of the Canadian North, 1870-1914, Toronto/ Montreal, 1971; pages 264-267. The wording of the plaque is shown on page 267. A photograph of the tablet on Melville Island, with the ship’s company standing in front of the Union Jack and the Dominion Flag, is printed in an article on “Arctic Sovereignty” in The Canadian Encyclopedia Hurtig 2nd Edition 1988, Volume 1, page 113. There is only one person sitting down at the front of the photo wearing a school cap. Presumably this is Frank Hennessey. On Bernier see Zaslow Opening of the Canadian North pages 261-277. . Taverner to Fleming, 24 February 1912, ROM. There is some corroborative evidence of this. Thomas Mcllwraith wrote to John Macoun of the Geological Survey, at its Sussex Street headquarters, where the small museum was housed, a long letter about the importance for Canada to establish a national muse- um where exhibits of her natural productions could be properly classified and arranged. He gave Macoun some very forthright advice: The position of Naturalist, to which he had been promoted (1887) was well deserved, Mcllwraith said, “but you are responsible for the condition of the museum [empha- sis added], and the country will look to you to have it in such shape as will not bring reproach on us from strangers from other countries while visiting the capi- tal.” (pages 1-2). Near the end of this letter he said: “I would advise you by all means to stay at home and get the museum hammered into the shape you would like to see it.” (page 6). But that is exactly what Macoun was not prepared to do. He continued mak- ing collecting expeditions until nearly the end of his life and Taverner had taken over the care of the birds and mammals from him. On the state of the museum accommodation of the Geological Survey see Waiser, The Field Naturalist. John Macoun, the Geological Survey, and Natural Science (Toronto, 1989) pages 120-142. Near the end of his life Macoun began dic- tating his autobiography. Speaking about the years 1882-1884 he said: the skins of birds, that I brought, were spread out by me on a table ... where the draughting group was, and, after a time, were placed in long drawers that were in the old Museum, and, in the course of time, with Dr. Bell’s specimens of birds, collected on Hudson’s Bay, were destroyed almost wholly by insects, and the remainder burned.” [emphasis added]. John Macoun, Autobiography of John Macoun, Canadian Explorer and Naturalist 1531-1920. Ottawa Field Naturalist Club, 1922, pages 219-220. I am grateful to Daniel Brunton for information on this subject. . Taverner to Fleming, 30 December 1920; and 15 January 1921, ROM. . Taverner to Bent, 15 January 1921, CMN. . Bent to Taverner, 2 January 1921; Taverner to Bent, 12 March 1921, CMN. CRANMER-ByYNG: A LIFE WITH BIRDS 8. 13: 14. IISy. 238i National Archives of Canada, R. M. Anderson Papers “Personal Record Card”. MG 30 B 40, Volume 13, File 8 and 10. These typewritten Personnel Record Cards were composed by Anderson; file 8 was pre- pared about 1920, consisting of one page of his mili- tary record and one page of family information. F.10 was prepared in 1931 for the office of the Minister of Mines, W. A. Gordon. It was revised and copied out in October 1936. This contains considerable informa- tion about his antecedents, including his mother and her family, and several of his uncles, one page about his academic training, one about his post graduate experiences, and several pages of detail about his executive and administrative experience from 1908 to 1927. Anderson’s papers in the National Archives at Ottawa are extensive, and can be searched with the help of Finding Aid Number 340 which mentions the main subjects of 39 volumes. In contrast Taverner, and his widow, preserved no personal files, and no “Personnel Record Cards”, while no collection of Taverner Papers exists in the National Archives of Canada. . Rudolph Martin Anderson The Birds of Iowa. A Thesis submitted to the Faculty of the Graduate College of the State University of Iowa, in partial fulfillment of the requirements for the degree of Doctor of Philosophy. Proceedings of the Davenport Academy of Sciences, Davenport, Iowa, Volume 11 March 1907, pages 125- 417. Paperbound 292 pages. It recorded 355 species and subspecies known to occur more or less regularly in the state, plus 44 species of casual occurrence. Consult The Literature of lowa Birds: A Complete Record of the Writings of the Birds of Iowa. By Paul Bartsch, Iowa University, 1889. . On the “Birds of Point Pelee” see Chapter 4 above, pages 65-67. It was reviewed by J. A. Allen in The Auk 26: 98-99 (1909). . For information on Anderson’s first experience of the arctic (1908-1912) see Richard J. Diubaldo Stefansson and the Canadian Arctic McGill-Queen’s University Press, Montreal, 1978, chapter 3, especial- ly 39-42, and note 27. I wish to acknowledge here the use I have made of Richard Diubaldo’s valuable account of the part played by R. M. Anderson in arc- tic exploration with V. Stefansson from 1908 onwards. Also Vilhjalmur Stefansson My Life with the Eskimo New York Macmillan 1912; 1913 and 1945. On Anderson’s second thoughts in the spring of 1913 about going to the arctic with Stefansson again see Diubaldo page 95. . On the preparations for the Canadian Arctic Expedition see Diubaldo chapter 4, especially pages 62-68 (See note 11). Also Trevor H. Levere “Vilhjalmur Stefansson, the Continental Shelf, and a New Arctic Continent” BJHS 21 (1988) pages 238- 240; Zaslow Reading the Rocks pages 319-325. (See chapter 5 note 1) Diubaldo (see note 11) pages 75-82 provides details of events leading to the confrontation between Stefansson and Anderson at Collinson Point in March 1914. See Diubaldo (note 11). Dr. Joel Asaph Allen (1838-1921) was a founder of the AOU and President for the first seven years of its existence. Editor of The Auk for 28 years, and editor 232 16. 20. Dili 30. THE CANADIAN FIELD-NATURALIST of three editions of the AOU Check-list. Curator of Birds and Mammals at the American Museum of Natural History from 1885 until his death. Taverner to Fleming, 27 February and 3 March 1913, ROM. . Canada, Sessional Paper 1913, “Summary Report of for 1914”. R.M. Anderson: “Report on the Canadian Arctic Expedition 1913-14”, pages 163-165. Diamond Jenness, anthropologist with the Southern Party, collected about 50 birds in 1914, including [Northern] Pintail, Harlequin [Duck], Red Phalarope, Parasitic Jaeger, Peregrine Falcon, Sabine’s Gull etc. mainly from Barter Island, northern Alaska. I am grateful to Stuart Jenness for this information. . Anderson to Taverner, 10 January 1915 (from Bernard Harbour, Northwest Territory). CMN., P. A. Taverner Archive. On Barren Ground bears see page 3 of letter. On a “distinct population known as the barren-ground grizzly” see A.W.F. Banfield, 1974, The Mammals of Canada, University of Toronto Press, pages 310-311. . Anderson wrote a report on the work carried out by the Southern Party which was published in 1917. See R.M. Anderson “Recent Exploration on the Canadian Arctic Coast” The Geographic Review 4(4): 241-266 (1917). In the final paragraph he wrote: “We were well loaded down when we left Bernard Harbour on the evening of July 13, 1916. We made a quick and easy voyage out: Baillie Island, June 24; Herschel Island, July 28; Point Barrow, August 8; and Nome, August 15.” Here their schooner was left to be sold while the men and specimens travelled via the Alaska and British Columbia Inside Passage to Victoria and Seattle. Taverner to Brooks, early October 1916, CMN. Foster Working for Wildlife pages 161-164 (See chapter 6 note 9). Duncan Campbell Scott, poet, short story writer and civil servant. Joined the federal Department of Indian Affairs in 1875, became deputy superintendent in 1913 and held that post until his retirement in 1932. 2. See Chapter 7 above, page 163; Foster pages 159- 164. (See chapter 6 note 9) . See Chapter 7 above, page 153. . National Archives of Canada. R.M. Anderson “Papers, Personnel Record Card” MG 30 B40, Volume 13, file 10. Anderson made a preliminary reconnaissance of this large area in 1917. See Anderson to Fleming, 4 September 1918, ROM. . Anderson to Fleming, 4 September 1918, ROM. . Anderson to Fleming, 27 March 1919, ROM. Also Soper to Taverner, 12 April 1919, CMN. . Taverner to Fleming, 5 April 1919, ROM. Also see Waiser The Field Naturalist pages 198-200. (See chapter 5 note 2) . Their house in Ottawa was at 58 Driveway, and their cottage was built on Lot 18, Big Island, Blue Sea Lake. . See Chapter 7, page 157. For more detail see Waiser The Field Naturalist 199-200 (see chapter 5 note 2). The quotation is part of a longer one given by Dr. Waiser on page 200, from a letter by Taverner to Brooks. For an account of the situation in the museum in 1920 see Chapter 11. 31: Bye, Oo Ww 34. 35. 3h 38. 39. 40. 41. 42. Vol. 110 Taverner to Fleming, 16 March 1920, ROM. The next few pages have been based largely on Richard Diubaldo, Stefansson and the Canadian Arctic. In a letter to his wife, Mae Belle, from the Arctic, Anderson wrote that Stefansson had caught him at a very unfavourable time to make any compro- mise with him. Stefansson had just brought him some mail which contained her letter telling him the sad news that their baby son had died three days after birth. As a result Anderson felt “hard and bitter”. He felt quite guilty when he thought of her lonely condi- tion. “I had counted on the boy being some company for you,” he wrote. (Letter Anderson to Mrs. Ander- son, 28 June 1914, as quoted in Diubaldo (see note 11) page 99.) . Letter Mrs. Anderson to Mrs. A. Allstrand, 5 January 1922, quoted in Diubaldo (see note 11) page 196. Also see Diubaldo pages 188-205 for details on Stefansson’s return to Canada in 1918, the publica- tion of The Friendly Arctic, and Jenness’ long letter in the journal Science challenging many of Stefansson’s statements in his book. Diubaldo (see note 11), pages 200-204. Letter Brock to Camsell, 24 April 1923, quoted in Diubaldo (see note 11) page 205. . In 1919 Anderson was writing reports on arctic mam- mals, and part of a report on arctic birds, but volume 2 of the reports on the Arctic Expedition (mammals and birds) to be edited by Anderson, never material- ized. As late as 1940, in an article in /bis, Allan Brooks deplored the parsimony of the Canadian Government in not publishing Anderson’s Arctic report. Taverner, in reply, wrote: “In this case at least it is not the government’s fault but Anderson’s. In spite of the Department pressing him to do so he has never presented a report for publication. He edited all the other reports of the expedition but had never pre- pared one for himself. It is quite inexplicable.” Zaslow Reading the Rocks (see chapter 5 note 1) pages 324-325 has given one answer, Diubaldo in Stefansson (see note 11) page 195 another. To this day Anderson’s action seems “quite inexplicable”. Until late 1921 his letters to Fleming had mainly been about wildlife and conservation, but in 1985 I discov- ered that the Fleming collection in the Royal Ontario Museum contained typewritten letters from Anderson to Fleming which give a good idea of how very upset Anderson felt about Stefansson’s account of the Arctic Expedition. Anderson to Fleming, 9 November 1921, ROM. In this, and subsequent letters, the differences between the standards, attitudes, perhaps also the natures of Anderson and Stefansson begin to show clearly. The word “fake” or faker” in this series of letters is one that seems to have a special meaning. It occurs in his letters again later in his life. It has the connotation of a dishonest person, someone who should not be trusted. Diubaldo (see note 11) page 209. His excellent chap- ter “Conclusion” runs from pages 209-215. On this see Taverner to Rowan, 20 April 1923, CMN; Taverner to Brooks, 4 July and 16 October 1923, CMN; Brooks to Taverner, 25 August 1923, CMN. Taverner to Fleming, 25 August 1924, ROM. P. A. Taverner “Annual Report” 1925, page 46-48; Taverner to Brooks, 24 October 1925, CMN; Brooks to Taverner, 3 November 1925, CMN. 49. 60. 61. 62. 63. 64. . Taverner to Bishop, 21 October 1926, CMN. . See Chapter 13. . Mae Belle Anderson to Rudolph Anderson, 10 August 1926. NAC. R. M. Anderson Papers. MC 30 B 40 Volume 4, file 2. . M.B. Anderson to R.M. Anderson, 6 July 1926, NAC. Mrs. Anderson spelt Karel’s name “Carl”. M.B. Anderson to R.M. Anderson, 1 November 1926, NAC. . M.B. Anderson to R.M. Anderson, 2 November 1926, NAC. During the AOU meeting Taverner played a major part, especially with the gift of a copy of Birds of Western Canada to each AOU member. Mrs. Anderson may have been indignant at this because her husband did not appear to have shone particularly, and did not have anything published to display. Anderson to H. P. Allstrand, 19 August 1927, CMN. His height, according to his certificate of naturaliza- tion, when he became a Canadian citizen, was 6 feet. When he wrote “knock on wood” what was he intending? . Laing’s side of the matter is given in Mackie H. M. Laing (see chapter 11 note 8) pages 94-96. . Taverner to Laing, 8 February 1927, CMN. . Laing to Taverner, 28 February 1927, CMN. . Mackie H. M. Laing (see chapter 11 note 8) pages 101-102. . Taverner to Laing, 31 March 1927, CMN. . Laing to Taverner, 7 April 1927, CMN. . Taverner to Laing, 21 April 1927, CMN. Also Mackie H. M. Laing (see chapter 11 note 8) pages 110-111. . Taverner to Laing, 6 October 1927, CMN. . Laing to Taverner, 27 February 1928, CMN. . Canadian Museum of Nature, P. A. Taverner archive. As well as the original, a microfilm of this Register of Birds also exists. I am grateful to Dr. W. Waiser for bringing my attention to this Register, and for loaning me his own microfilm copy. J. Dewey Soper “Discovery of the Breeding Grounds of the Blue Goose” The Canadian Field-Naturalist 44: 1-5 (1930). A.C. Bent “Life Histories of North American Wild Fowl” Order Anseres (Part 2). United States National Museum, Bulletin 130, Washington 1925. Bent stated what little was known of the breeding ground of Blue Geese in 1925 on pages 178-180. For detailed study see F. Cooke “Genetic Studies of Birds — the Goose with Blue Genes” Acta XIX Congressus Inter- nationalis Ornithologici, Volume I, Ottawa 1986, pages 189-214. Of special interest to Soper’s discov- ery see pages 208-211. J. Dewey Soper “Adventuring in Baffin Island” Canadian Geographical Journal 1: 191-206 (1930), with 16 excellent photographs. Soper to Taverner, 18 July 1929, CMN, from Camp Kungovik, West Coast Baffin Island Latitude — 65° 35! 1929. See Appendix | for photocopy of original letter complete. On B. Hantzsch see R.M. Anderson “The Work of Bernard Hantzsch in Arctic Ornithology” The Auk 45: 450-466 (1928). Hantzsch River and Taverner Bay are marked on sketch map in D. Soper Canadian Geographical Journal, (3): 194 (1930). Anderson CRANMER-BYNG: A LIFE WITH BIRDS 65. 66. . Taverner to Fleming, 25 September 1929, ROM. 68. 69. 1B 76. pif 78. i): 80. Mae) Headland is marked on sketch map in D. Soper The Canadian Field-Naturalist 44: 2 (1930). Kennard to Soper, 10 October 1929, CMN. Soper replied to Kennard, 29 October 1929, that the authori- ties might dispose of some of the “Blue Goose” mate- rial to other institutions, but he could not be certain. At that date the whole collection belonged to the Department of the Interior. For information on Kennard’s work on the geese of the genus Chen see A.C. Bent “In Memoriam: Frederick Hedge Kennard 1865-1937” The Auk 54: 345-346 (1937). The even- tual ownership of the collection was decided at a meeting in November 1929. See Appendix 2. R. M. Anderson Annual Report 1928 pages 10-11. Taverner kept a Journal of his voyage. Volume | has an itinerary, volume 2 analysis of species. On Taverner’s study of the migration routes of the Hudsonian Curlew [Whimbrel] see Chapter 16 below. P. A. Taverner “Some Zoological Aspects of the Canadian Arctic Expedition of 1929” Canadian Field-Naturalist 44: 25-28, 1 plate (1930). Taverner to Munro, 7 March 1930, CMN. Taverner mentioned the new Hudson Bay railway from The Pas to Churchill. . Taverner to Laing, 9 January 1931, CMN. . Percy A. Taverner and George M. Sutton The Birds of Churchill Manitoba. Annals of the Carnegie Museum 23: 1-83, 13 pls 1 map (1934). . Taverner to W. H. Collins, 19 February 1930, CMN. . Taverner to H. H. Mitchell, 6 February 1930, CMN. . Albert Lloyd, who went with Taverner to Hudson Bay in 1930, continued in subsequent years under Clyde Todd and became a permanent member of the Carnegie Museum staff. Arthur Twomey, a student of Rowan at the University of Alberta, did field work at Hudson Bay in 1930, then attended the University of Illinois where he received his Ph. D. He later took part in expeditions to the Hudson Bay area under the auspices of the Carnegie Museum. This material based on Marianne G. Ainley From Natural History to Avian Biology: Canadian Ornithology 1860-1950, Ph.D. dissertation, McGill University 1985, pages 133-136, which I gratefully acknowledge. Canada, Sessional Paper 1911 “Summary Report for 1910”. Taverner to Collins, 6 December 1929, CMN. Deputy Minister of the Interior H.H. Rowatt to Deputy Minister of Mines Charles Camsell, 27 June 1933. National Archives of Canada, Charles Camsell papers. If Anderson was shown the letter he was not likely to have been pleased to find Taverner referred to as “the best person in the service to undertake this work” i.e., to write about birds of the eastern Arctic. Translated from the German by L. H. Neatby as My Life Among the Eskimos. Institute of Northern Studies, University of Saskatchewan, Saskatoon, 1977. Taverner “The Birds of the Arctic Islands”. Unpublished typescript dated 1933. CMN. P.A. Taverner archive. This is a 16 page “Introduction”, but contains no list of birds. Canada’s Eastern Arctic. Its History, Resources, Population and Administration. Assembled by W.C. Bethune for the Northwest Territories Council and 234 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. THE CANADIAN FIELD-NATURALIST issued by the Department of the Interior, Ottawa, 1934. Taverner’s contribution is on pages 113-128. W.B. Alexander to Taverner, 25 November 1933, CMN.; Taverner to Alexander, 6 December 1933, CMN. Taverner to Fleming, 11 December 1929, ROM. The Corn Crake (Crex crex) formerly known as “casual or straggler” in the autumn along the east coast of North America, now considered as “accidental”. The Audubon Society Master Guide to Birding. See National Geographic Society Field Guide to the Birds of North America 1983, page 98 and illustration. On a Barnacle Goose taken near Cape Dorset, August 1924 see P. A. Taverner “Some Recent Canadian Records”, The Auk 44: 217-228 (1927). Taverner to Rowan, 22 May 1931, GMN. Department of Mines, Annual Report 1931-2, pages 1-2, as quoted in Zaslow, Reading the Rocks (see chapter 5 note 1) page 361; also pages 373-374. Zaslow Reading the Rocks pages 356-357. Letter Watson Seller, Assistant Deputy Minister, Department of Finance to Charles Camsell, Deputy Minister of Mines, 18 August 1931, NAC, Camsell Papers, RG 45, Volume 49; L. L. Bolton to W. Sellar, 21 August 1931; L.L. Bolton to Charles Camsell, 22 August, 1931, NAC, Camsell Papers. The language in which these letters were written is a shining exam- ple of civil service gobbledygook, well worth includ- ing in an anthology of examples of that genre. Personal interview with Hugh S. Bostock, 17 September 1984, and tape recording. Bostock was a member of the Geological Survey Canada 1926- 1965. Stationed southern B.C. until 1930; Yukon and the Territory 1931 until 1965. Promoted Assistant Chief Geologist 1950. Author of Pack Horse Tracks. Geological Survey of Canada 1979. For biographical information see “Foreword” by D. J. McLaren, and “Author's Preface”. Although Bostock could not have been at Taverner’s confrontation with Camsell he could well have heard of the episode from some- one else. The sight of Taverner, agitated and stam- mering, while “balling out” the Deputy Minister of Mines, must have been an experience to remember. Letters Taverner to Laing, 30 May 1935, CMN; Taverner to Laing, 30 April 1936 where this is explained more fully. Draft letter Taverner to Collins, undated, unsigned. Even if a complete version of his was never sent Taverner had written other letters to Collins at this period. The correspondence between Collins and Taverner in the years 1926-1936 was quite a large one. From 1935 onwards Anderson carried out a major attack on Taverner — on paper. Although Taverner guessed at some of the things Anderson said about him, he had no clear idea of their content or extent. Much of the information used in the following notes has come from Dr. and Mrs. Anderson’s correspon- dence which was donated to the National Archive of Canada by members of their family after their death. (Finding Aid N 340) It was just as well that Taverner died without knowing the contents of these letters. In addition several revealing letters from Anderson to Laing, written between 1935-1936 are in the Public Records of British Columbia. I am grateful to Richard Mackie, author of Hamilton Mack Laing: 91. 92. 93% 94. 95. 96. Wile 98. 99. 100. 101. Cha 1. Taverner to Ida Clare Taverner, 10 June 1928, CMN. ) 3 4. Vol. 110 Hunter Naturalist (see chapter 11 note 8) for bringing these letters to my notice. Taverner to Laing, 30 April 1936, CMN. By this time some of Taverner’s friends were warning him that Anderson was writing in derogatory terms about him. Taverner to Laing, 18 May 1936, CMN. This was wishful thinking on Taverner’s part because Anderson had already written several letters to Laing expressing his contempt for Taverner, not so much in terms of a personal disagreement, but rather because Taverner was not up to his job as the museum’s ornithologist, as well as more unpleasant insinua- tions. I do not know the extent of truth or untruth in these statements. Taverner to Laing, 18 May 1936. What exactly Taverner meant by this is unclear. Taverner to Brooks, 26 September 1936, CMN. For a fuller account, with a reorganization chart and a photograph of John McLeish, see Zaslow Reading the Rocks (see chapter 5 note 1) pages 377-381. For a brief biography of him see Chapter 17, note 41. Memorandum: Status of Division of Biology, National Museum of Canada. From R. M. Anderson, Chief, Division of Biology, National Museum of Canada. To Dr. Charles Camsell, Department of Mines. NAC, MG 30 B40 Volume 14, file 20. See Appendix 3 where the main points of Anderson’s memorandum have been briefly noted. Taverner to Brooks, 20 April 1937, CMN. This was his “Record Book” started in 1931 and end- ing at his retirement in 1942. In this hand written log book Taverner kept a note of anything he wanted to have a note on. For 1936 there were 6 pages; 1937, 30; 1938, 12 pages. Michel Gosselin of the Ornithological Section found this Record Book among Taverner’s papers. CMN, P. A. Taverner Archive. Taverner to Lynch, 13 July 1935, CMN. This letter was based on a letter from Taverner to Brooks dated 27 June 1935 in which he told Brooks very clearly that he (Brooks) was too generous in donating speci- mens to the museum. Taverner said that he felt guilty in accepting all Brooks’ gifts to the museum. Anderson’s position on the matter of exchanges of specimens was different, and was made clear in a memorandum on “Exchanges and Gifts of museum specimens” dated 28 April 1936 and signed R. M. Anderson. On the subject of the preparation of specimens for storage see R.M. Anderson Methods of collecting and preserving vertebrate animals National Museum of Canada 1931; fourth edition 1965, Biological Series number 18, Bulletin number 69. Anderson devoted chapter 4 to methods of collecting and preserving birds. See Appendix 3. pter 15 National Museum of Canada Annual Report 1928, page 12. (Bulletin number 62). Ottawa, 1929. Taverner to Charles Townsend, 4 September 1928, CMN. Taverner to Bishop, 6 September 1928, CMN. Glover Allen was Curator of Mammals, Museum of 1996 NY), 20. . Taverner to Laing, 2 January 1935, CMN. . See note 36 below. . Taverner to Brooks, 18 February 1932, CMN. . Brooks to Taverner, 15 July 1924, CMN. . Taverner to Brooks, 7 February 1925, CMN. This let- Comparative Zoology, Harvard; Francis Allen, Publisher with Houghton Mifflin Co., Boston; Frederick Kennard was a specialist on geese of the genus Chen, and Harry Oberholser, United States National Museum. In addition to those staying at Amory’s, Arthur Allen of the Cornell University Zoological Department, was on the north shore for a while working with H. F. Lewis who, in his official post, had the use of a boat. Todd worked the “Canadian Labrador” from Natashquan eastward. See note 9 for explanation of the term Canadian Labrador. . Taverner to Glover Allen, 7 September 1928, CMN. . Glover Allen to Taverner, 26 September 1928, CMN. For Northern Lapwing see note by Taverner in The Auk 46: 231 (1928). Also Taverner The Auk 46: 223 (1929) on the Razorbill. . Taverner to Glover Allen, 16 October 1928, CMN. . P.A. Taverner “Bird Notes from the Canadian Labrador, 1928”, The Canadian Field-Naturalist 43: 74-79 (1929). Taverner acknowledged that Harrold assisted him. Writing to Frank Bradshaw after Harrold’s death, Taverner praised his tireless work. . P. A. Taverner “Bird Notes from the Canadian Labrador, 1928” 74-79 (1929). The Canadian Field-Naturalist 43: . Copley Amory to Taverner, 18 September 1931, CMN. For a report on the conference by R.M. Anderson see NMC Annual Report 1931 (Bulletin Number 70) Ottawa 1932, pages 11-12. . Taverner to Laing, 28 November 1931, CMN. . Taverner to Brooks, 3 August 1931, CMN. Also Taverner to Rowan, 8 July 1931, CMN. . Taverner to Laing, 29 June 1933, CMN. . Taverner to Laing, 3 July 1934, CMN. . Taverner to Munro, 17 June 1935, CMN. . Taverner to Laing, 13 November 1936, CMN. . Taverner to Soper, 27 October 1936, CMN. . Taverner to Laing, 6 October 1931, CMN. Dr. Eidmann was attending the Matamek Conference in 1931. Writing to Fleming, Taverner said that he was a forest entomologist from Munich, “of very broad interests and quite delightful”. (Taverner to Fleming, 24 September 1931, CMN.) Taverner to Laing, 22 April 1932, CMN. The “Unfinished” refers to Schubert’s symphony of that name. Other melodies that Taverner used to whistle, while doing field work, were from the “Ode to Joy” in Beethoven’s Ninth Symphony, and from Brahms’ Symphony No. 1, which he considered one of the finest symphonies. . Taverner to Laing, 22 April 1932, CMN. . Taverner to Laing, 18 January 1933, CMN. . Taverner to Laing, 13 April 1933, CMN. . Tavermer to Laing, 1 May 1933, CMN. . Taverner to Saunders, 18 February 1932, CMN. . Taverner to Laing, 22 April 1932, CMN. . Anderson to Kenneth Racey, 11 February 1937, CMN. ter contains two pages of details on calurus and har- lani as Taverner understood them then. CRANMER-BYNG: A LIFE WITH BIRDS 33% 34. 35. 36. Silk 38. 39: 40. 41. D335) Taverner to Brooks, 31 December 1925, CMN. Important in showing how to illustrate birds diagram- matically. Taverner to Brooks, 7 February and 31 December 1925, CMN. Taverner had known N. A. Wood at Ann Arbor while a student there. See Chapter 3. P. A. Taverner Birds of Western Canada 1926 ad revised edition 1928 page 193; also Taverner’s Birds of Western Canada reprinted by Coles Publishing Company, 1974, same page. P. A. Taverner A Study of Buteo borealis, the Red- tailed Hawk, and its varieties in Canada. Canada Department of Mines, Museum Bulletin Number 48, 1927. With 3 pages of colour illustrations of tails. Taverner Study of Red-tailed Hawk (see note 36) page 3. Witmer Stone The Auk 45: 244-245 (1928). This refers to Taverner calling the western race harlani, and regarding it as identical with the melanistic phase of calurus, but giving harlani precedence over it for reasons of priority, because the name harlani was published earlier than the name calurus. H. S. Swarth The Condor 30: 197-199 (1928). Presumably at the Museum of the University of Michigan since he acknowledged such a courtesy in his paper. Also this subspecies was described by Van Tyne in Occasional Papers of the Museum of Zoology Number 321, University of Michigan 1935 page 1. Buteo harlani was mentioned briefly in the Thirty-Second Supplement to the Check-list page 106. Mentioned in The Auk 90: 411-419 (1973). Also see R.S. Palmer (Editor) 1988 Handbook of North American Birds Volume 5. Yale University Press, New Haven, pages 100-105. Taverner to Rowan, 19 January 1924, CMN. The first two pages are about the Cackling Goose, and how Brooks distinguished it from Hutchin’s Goose. Taverner did not feel that A. C. Bent himself was well informed on Hutchin’s Goose: “It is a case where intimate field knowledge has to supplement closet experience”, Taverner wrote. Brooks “certainly had superior field knowledge of the Cackling and Hutchin’s”. . Birds of Western Canada page 109. . P. A. Taverner “A study of Branta canadensis (Linnaeus) the Canada Goose (Based upon breeding or summering specimens) Illustrations. Plate I Branta canadensis, the Canada Goose. Figure 1. Typical bills of the races of the Branta canadensis group.” (Bulletin National Museum of Canada, number 67, pages 28-40, 1931.) . “A study” (see note 43) pages 31. “A study” (see note 43) pages 39-40. “A study” (see note 43) page 40 — at end of study. Authorities since Taverner’s study generally treat the Cackling and Richardson”’s Geese as subspecies, not separate species. . Taverner to Rowan, 6 October 1931, CMN. . Birds of Canada page 86. Its nesting ground was not discovered until 1938. . Taverner to Brooks, March 1933, CMN. . Brooks to Taverner, 14 June 1933, CMN. . Taverner to Brooks, 28 June 1933, CMN. . W.E. C. Todd “A new eastern race of the Canada Goose” The Auk 55: 661-662 (1938). The reasons 236 53% 54. 55. 56. D7: 58. 59. 60. 61. THE CANADIAN FIELD-NATURALIST why Todd discovered a subspecies of the Branta canadensis from eastern Canada that was different from the one already recognized by Taverner were explained by Todd on page 661. He said that Taverner’s study was based on breeding specimens, but since he apparently had no breeding specimens for eastern Canada and Newfoundland his remarks on typical canadensis referred to examples from inland and western localities. But the Canada Geese which migrate along the Atlantic coast and breed in Newfoundland and Labrador were not the same as those which migrate through the interior and breed on the east coast of Hudson Bay. “The former are light-colored birds; the latter by comparison are dark-colored”. Todd then determined the type locali- ty of the original canadensis and then separated the Branta canadensis interior as a subspecies (page 662). Jean Delacour American Museum Novitates 1537, page 7 (1951). Colusa, California. Same. Delacour’s B. c. taverneri was not recognized in the Fifth AOU Check-list of 1957. Godfrey (1986) page 81 under Subspecies. Number (10) B.c. taverneri Delacour. R. S. Palmer, 1976 as above, did not recognize it but made one mention of “some so-called taverneri’” under the Lesser Canada Goose Branta canadensis parvipes. Taverner to Brooks, 26 July 1933, CMN. Leon Cole to Taverner, 6 April 1932, CMN. They were first in touch with each other, over bird banding, in 1909. Cole was the first President of the American Bird Banding Association. Taverner to Cole, 20 April 1932, CMN. In writing to Taverner at the National Museum Cole addressed him as “Dominion Ornithologist”. Taverner was never given that title officially. However, from January 1937 Lynch agreed verbally that the title “Chief of Ornithology” could be used unofficially. See Taverner’s handwritten “Zoological Record Book” 8 January 1937. National Museum of Canada Annual Report 1929 17 (Bulletin Number 67) Ottawa, 1931. There were two boxes of postcards. Series A number | containing 50 cards, “Common Loon to Song Sparrows”. Series A number 2 containing 60 cards, “Eared Grebe to Eastern Bluebird”. Collins to Taverner, 5 February 1929, CMN. Taverner to Brooks, 7 February 1929, CMN. On 4 February 1933 Taverner wrote to Brooks that the book seemed to be nearing completion, and that he had prepared a lot of additional line drawings for it. C.B. Tidd to H. Lloyd, 25 March 1933, signed Sergeant C.B. Tidd, RCM Police, i/c Mayo Detatchment, CMN. The Taverner book he referred to was Birds of Western Canada. . Taverner to Collins, 3 October 1934, CMN. . Taverner to Laing, 3 July 1934, CMN. . P. A. Taverner Birds of Canada, page 294 under Wright’s Flycatcher. . Brooks to Taverner, 27 August 1935, CMN. . National Museum of Canada, Bulletin Number 72. $2.00 in cloth, $1.50: paperback. . Joseph Grinnell, The Condor 37: 179-180 (1935). The very lifelike photo, which caught Taverner look- ing directly at the camera for once, rather than turn- 68. . William L. Sclater The Ibis 5 Thirteenth Series: 688 70. WB: 74. fey 70. OE 78. Vol. 110 ing his head to one side as he usually did, was proba- bly taken by a colleague at the 47th Annual Meeting of the AOU at Philadelphia in October 1929. The original photograph was sent by the taker and on the back of the original Taverner had written “received January 7, 1930”. Martha Taverner kept the original until her death, when her son Karel Wiest had it. He kindly gave it to me when I interviewed him in Detroit only a few weeks before he died. Witmer Stone The Auk 52: 333 (1935). (1935), British Ornithologists’ Union, London. Taverner to Ernest Ingersoll, 2 April 1935, CMN. Ingersoll wrote a natural history column in the Montreal Family Herald and Weekly Star. . Taverner was not the only one of the well-known members of the AOU of his generation who did not have any academic degrees. 2. J. A. Munro “The Necessity for Vermin Control on Bird Sanctuaries” The Canadian Field-Naturalist 37: 148-149; (1923): Editorial “Bird Sanctuaries” (same issue) pages 149-150. Letter to Editor on “Necessity for Vermin Control ...” from Wm. Rowan, January 1924, The Canadian Field-Naturalist 38: 30-33 (1924); A. Brooks letter to Editor, pages 33-34; Charles Townsend letter to Editor, page 35; Editorial “Control of Predatory Birds and Small Mammals” The Canadian Field-Naturalist 38: 35-36 (February 1924). On Kerr see Chapter 9. Jack Miner’s eldest son, Manly, acted as his father’s secretary. Since he did not use a typewriter he had to write a mass of letters. Manly F. Miner to Harkin, 15 February 1926, CMN, and signed by Manly. Arthur Gibson, Dominion Entomologist. ” averner’s position at this time was stated clearly in a letter to Harkin of 9 February 1926, CMN. Briefly he said that considerable destruction of hawks and owls was going on at a great many game farms in the winter “and there seems no way to stop it, except educa- tion.” E. R. Kerr to Harkin, 26 February 1926, CMN. The letter-head on the notepaper read: “The Jack Miner League”. What Kerr meant when he wrote that Taverner and Saunders “may possibly make more headway among sportsmen and farmers” is not quite clear. Headway in what? Jack Miner’s philosophy (or religion) was strongly homocentric, one in which human terms were used to judge the actions of birds. Taverner’s arguments were sound, but Miner did not accept the linguistic defini- tion of “cannibal”, only his own. Taverner to Manly Miner, 22 February 1926, CMN. Arch-fiend was an echo from the Old Testament in English. It was ignorant to apply it to crows and other raptors. Taverner to Manly Miner, 29 April 1926, CMN. Manly Miner to Taverner, 30 April 1926, CMN. . Taverner to Manly Miner, late 1926, CMN. Taverner to Manly Miner, 12 November 1926, CMN. . Hoyes Lloyd to Taverner, 15 December 1926, CMN. Lloyd made an interesting point in his letter — that the Miners did not have a permit allowing them to band birds for scientific purposes because they did not use authorized bands. Since that time legal band- 1996 84. 93% ing has increased substantially in Ontario and contin- ues to flourish. Taverner to Lloyd, 30 December 1925, CMN. Taverner obtained full returns of permit holders in Ontario for the year 1924. This showed that of 56 permit holders in the Province, 47 took no specimens at all and that the remaining 19 took 315 birds. Taverner commented that 315 birds killed for scien- tific purposes from widely scattered localities in the Province of Ontario in twelve months was a trifle. It was nothing in comparison with a dozen or more minor causes of bird mortality that were too trivial to cause remark: — “overhead wires, glass windows, railroad trains ... while the common cat undoubtedly kills several thousand times the number without any sign of uneasiness from our game departments, and forest fires destroy millions of birds without being considered.” . William N. Kelly to Harkin, 6 March 1931, NAC; Lloyd to Taverner, 18 March 1931 with enclosures, CMN. Kelly was a consulting engineer. . Taverner to Lloyd, 20 March 1931, CMN. . Taverner to Harkin, 3 October 1931, in reply to Harkin’s letter to Taverner of 1 September 1931, CMN. . H.J. Parham to Taverner, 3 November 1932, CMN. Taverner to Parham, 11 November 1932, CMN. . Parham to Taverner, 24 November 1932, CMN. . Taverner to Parham, 15 December 1932, CMN. The term “local members” usually refers to “the local member of Parliament in a district”. . The controversy over whether some collecting of bird specimens for scientific studies was necessary, or whether all collecting was quite unnecessary, as Parham argued, was still alive in the 1980s. An event occurred beside the Ottawa River in October 1982 when a small shore bird was “collected” in order that a group of birders could identify it without any doubt, thus solving the disagreement over what species it was. Information on the “collection” of this bird was published in the newsletter of the Toronto Field Naturalists of December 1982 (Number 352, pages 6—7). Readers were invited to write to the newsletter with their views, and also directly to Dr. Henri Ouellet, Curator of Birds, National Museum of Natural Sciences, Ottawa. Dr. Ouellet replied in detail in a subsequent issue. His main point was that “a small percentage of species or individuals in cer- tain populations cannot be identified positively with- out a specimen; this applies currently ... because individual variation, even in the common species is not fully known and understood.” Dr. Ouellet appended a list of 13 titles by ornithologists on vari- ous aspects of this contentious issue. (Toronto Field Naturalist Number 355, April 1983, pages 32-34.) H. J. Parham A Nature Lover in British Columbia. H. F. Witherly Ltd., London 1937. Chapter 33 “Save the Birds” page 236, paragraphs | and 2. In para- graph 3 Parham wrote: “In southern British Columbia we have a few arid valleys and a narrow humid coastal strip in which are found, in very limited numbers, birds known to few, if any, other parts of Canada. “Some of these — in the Okanagan particularly — are the Canyon and Rock Wrens, Lazuli Bunting, CRANMER-BYNG: A LIFE WITH BIRDS 94. Ps Cha iL, N 237) Dickcissel, Brewer’s Sparrow, Bobolink, Long-tailed Chat [Yellow-breasted Chat], Sage Thrasher and probably Sage Sparrow, Williamson’s Sapsucker, Wilson’s Phalarope and Burrowing Owl.” The Ottawa Journal 9 February 1938. Parham to Taverner, 3 November 1932, paragraph 2, CMN. pter 16 Francis Kermode joined the British Columbia provin- cial museum, Victoria, in September 1890, and served as director from 1904 until he retired in 1940. Taverner and Kermode corresponded spasmodically from April 1914, and Taverner visited the provincial museum in 1922. As long as Taverner and Brooks were good friends there was no chance that Kermode and Taverner could be on good terms. There is no biography of Kermode, but there is some information on the 50 years of his service in the B.C. provincial museum in R. W. Campbell et al. The Birds of British Columbia Volume I, page 19. Published by the B. C. Museum, Victoria, 1990. A provincial museum was founded at Edmonton, Alberta, in 1905 consisting of mounted birds and mammals, historical photographs and miscellaneous artifacts, with a staff of two or three unqualified civil servants. Since then the Alberta Museum has come a long way. Although not spacious for a provincial museum it is housed handsomely. The exhibits are attractively arranged. For Horace Hedley Mitchell see Ron Borden “An Institution that Matters” Liaison: Saskatchewan's Heritage Review May 1986 page 5-6; Annual Reports of the Chief Game Guardian, Government of the Province of Saskatchewan, Department of Agriculture, Regina; Robert D. Symonds “Personal Recollections of some early Saskatchewan Naturalists” Blue Jay 24: 2-6 (1966). Taverner had known Mitchell since 1912. . Dr. H. M. Speechly. A country doctor who called himself a “humble roadside naturalist”. But he was very much more than that. See Robert Symonds, “Personal Recollections” Blue Jay 24: 2-3 (1966). For the Manitoba Museum see A. M. Davidson, A. Simpson, H. G. Lawrence et al. Natural History Society of Manitoba 21st Anniversary Bulletin 1920- 1941. Albion Press, Winnipeg, 1942, pages 56-59. A. P. Harrold to Taverner, March 1929; Taverner to A. P. Harrold, March 1929, CMN. Taverner to Cartwright, 4 April 1929, CMN. Taverner to Cartwright, 25 April 1929, CMN. Taverner to Speechly, 11 January 1937, CMN. One suggestion was to interest Angus Shortt in taking a course in museum management. Taverner to Speechly, 26 December 1939, CMN. Taverner to Speechly, 3 April 1940, CMN. This was a theme that Taverner continued to reiterate until the last months of his life. . Snyder heard the second of these talks, and wrote to Taverner congratulating him and thanking him for sending him a copy of it, which he was glad to have on file. The ideas were sound and beautifully expressed (Snyder to Taverner, 8 June 1934, CMN.). On the title page of the talk was printed: “Information for Radio Announcer”: 238 THE CANADIAN FIELD-NATURALIST “Mr. P. A. Taverner has been ornithologist to the National Museum of Canada since 1911. He is the author of several standard works on the Birds of Canada, and is a frequent contributor to journals devoted to natural history. He is a Fellow of the American Ornithologist’s Union, and his researches and contributions have given him an enviable reputa- tion among the ornithologists of the world.” . Taverner to Snyder, 7 May 1923, CMN. When. Snyder and party returned from the Lake Nipigon area, and the specimens collected had been prepared, Taverner was informed. He asked that they should be sent to the National Museum so that he could exam- ine them carefully and compare them with specimens in the museum. In a four page letter he explained the difficulty in distinguishing between specimens of closely allied races such as in the Sharp-tailed Grouse, the juncos, and others. In this matter of extreme care he told Snyder “... I am regarded as more or less a crank because I will not vouch for what I cannot see, and do not believe in jumping at geological conclusions”. Taverner to Snyder, 12 February 1924, CMN. . Lam grateful to Ross D. James, Curator, Department of Ornithology, Royal Ontario Museum, for his infor- mation on this subject. . Snyder to Taverner, 2 February 1928, CMN. . Taverner to Snyder, 9 April 1931, CMN; Snyder to Taverner, 13 April 1931, CMN. . Taverner to Snyder, 18 March 1932. L. L. Snyder The Hawks and Owls of Ontario, CMN. Illustrations by T. M. Shortt. Published by Royal Ontario Museum of Zoology, University of Toronto Press 1932. Revised edition 1947. The introduction to this edi- tion, page 16, concluded with these words. “It is a sad commentary to make, but shortly after the appearance of the original edition of this booklet [1932] , which was intended to aid in a better understanding of car- nivorous birds, all protection was removed from all hawks and owls in the Province of Ontario. This was a retrograde step. By 1947 protection had been restored to ‘Ospreys and eagles’. . Taverner to Snyder, 7 January; Snyder to Taverner, 2 February; Snyder to Taverner, 26 April 1935, CMN. Snyder described a new race of Sharp-tailed Grouse Pedioecetes phasianellus campisylvicola in 1935. University of Toronto Studies, Biological Series 4a. . Snyder to Taverner, 25 September 1939, CMN. Snyder’s letter says more than I have written here about its contents. . Taverner to Snyder, 6 December 1939, CMN; Snyder to Taverner, 12 January 1940, CMN. . On Snyder’s career see C. Long and J. C. Barlow “In Memoriam: Lester L. Snyder” The Auk 103: 809-811 (1986). . Baillie to Taverner, 13 January 1926, CMN. For a biography see Lise Anglin Birder Extraordinaire: The Life and Legacy of James L. Baillie (1904-1970) Toronto Ornithological Club and Long Point Bird Observatory, 1992. . Taverner to Baillie, 15 January 1926, CMN. . Taverner to Snyder, 7 February 1940, CMN. 4. Taverner to Snyder, 8 July 1940, CMN. 25. Taverner to Snyder, 28 September 1940, CMN. He explained that the war situation seemed to drain him 35): 36. 39: 40. Vol. 110 of all inspiration and that he would have to cal! for help. “In Memoriam: James Henry Fleming” by L. L. Snyder, The Auk 58: 1-12 (1941). Taverner to Snyder, 4 October 1940, CMN. . Taverner to Snyder, 15 March 1941, CMN. . Taverner to Snyder, 21 March 1941, CMN. . B. W. Cartwright to Taverner, 19 October 1928, CMN. Taverner to B. W. Cartwright, 14 November 1928, CMN. 1. Cartwright to Taverner, 14 November 1928, CMN. See Chapter 13 above, note 11. . See Marianne Gosztonyi Ainley “Henry Mousley and the Ornithology of Hatley and Montreal, 1910-1946”. Tchebec 11: 113-134 (1981), especially pages 125-127. For instance Taverner to Mousley, 3 July 1937, CMN. It starts revealingly “As you say we have , oe nera ol Renn usr nese a aeeneRe es front cover Taverner family photo taken at Ann Arbor, Michigan, 1889. Percy age 13, Ida Clare age 2......................... 7 The taxidermy shop of Oliver Spanner and Company at 358 Yonge Street, Toronto. .........................0... 13 Taverner preparing a skin in the Great Lakes Ornithological Club “shack” at Point Pelee, 30 May 1909. ............. 3] Five members of the Great Lakes Ornithological Club at club’s “shack” near the end of Point Pelee, 3 October 1909. Left to right: J. S. Wallace, B. H. Swales, W. E. Saunders, J. H. Fleming, ap Nm hay ernens (SEa tedvonysteps) macyaisaterorecuaie cask mesynusva eee eias satin Sees Areeuegie fo aeianeim cite eyawtenestee shesisueaeueteesueunscnetae 35 Photograph of Northern Saw-whet Owl taken by Taverner at Point Pelee 15 October 1910. ..................0..... 39 The Taverner house, 45 Leonard Avenue, with construction near completion in 1912. ...................2..22005. 59 Wistonia Memonell MW ineciimn Bini aliner ello ne ONE 5 Gesc aacandcaaueeomeaddd occa cond onoeendo ose Sea acoasds cos 61 Early exhibit hall and visiting public at the Victoria Memorial Museum, 1912..................0 000.002 s eee eee 63 Purple Martins at martin house designed by Taverner. Photographed by Taverner in the garden of his home HM Ottayapimetheys ummMacmo tall ONme ray csmicayae aire esse ce sre donate. cuits: omy chey arsine auc statue meucleeiaedheh oasredsieie ues 71 Gannets on bird cliffs on Bonaventure Island, 1915. Photograph by P. A. Taverner. .................0-00000 20200 ee 83 The wheel of the C. S. S. Princess, 1915, from the left: P. A. Taverner, Captain Chalifour, and C. H. Young........... 85 C. M. Sternberg’s field camp, Red Deer River, Alberta, 1917. Standing: Dr. R. L. Rutherford (University of Alberta), P. A. Taverner, C. M. Sternberg, C. N. Young, C. H. Sternberg, Dr. J. A. Allan (University of Alberta), three unidentified assistants are seated in foreground. ................ 87 Taverner in an open boat with an outboard engine while exploring the Rideau waterway system between Ottawa andeainestonumseatch otibirdsunmume LOS ys eens sees) cals eels ee -rererae s eye saa) sieve clei ie ei eleven eye) nie 89 Selectioniirom\@hester Reed’s Bird Guide (1912) vest-pocket edition. .... 2.222... sec eee cet ee tee eee 100 Field camp, Eastend, Saskatchewan, 1921. P. A. Taverner, Alan Sampson, and H. M. Laing. ...................... 117 “Asoyoos Meadows” [Osoyoos] south Okanagan, British Columbia, 14 May 1922. Left to right P. A. Taverner, [MECSBTOoOksslpvlethackerE MDE IB Greener Wa miciaiaic a siete Sauer ues ive odo aire) Wy = ST onene tess. race der itera lie sel Sys emeeee 119 Vaseux Lake, Okanagan, British Columbia, 31 May 1922. Left to right, standing: Brooks, Taverner, Frank Farley; left to right, seated: Hamilton Lang, George Gartrett, Alan Sampson. ................0 000.000.000.000 00. 121 BeavyermilWwakey Alberta, August 1925. PA. Taverner and'©,,G. Harrold: 2.2... 22.225. ee ee ee 131 Canadian Group Photograph taken at the American Ornithologists’ Union 1926 Annual Stated Meeting, Oita am Ontariowmnmeeyrrrie tamer kere or wee cnt meter dyes, Buin iae cla een ata ar aha okies Ravel neychnak oe mn GS So aici es 135 A professionally taken portrait of Percy Taverner framed with the words “Sincerely P. A. Taverner PAK ened DOUta O25 ampeeter yc Annus tanars cnet nioeuamt ann Gem ntyens tee qda iene cucta lenete a) cieevermus ome eh cimanie tare are 139 Joseph Dewey Soper upon return to Cape Dorset after surveys run to Nuwata across the interior of Fox Peninsula and from there east to Ungmaluktuk Lake and south to Gordon Bay during March 1929. .................... 149 A photograph received by Taverner in January 1930 from someone as a gift. ... 22... eee 151 Rudo phe lantinpAn Gerson Vay OSA ets cy espe Meer cue apeeet crete yal exayreuis eh Sieh Seay meen opesene = fo serena sieMtenel say slejsstesclatuaiato eis ete 155 Permit to capture migratory birds for banding purposes only in the Province of Manitoba to Mr. P. A. Taverner for the year 1931. Signed by Hoyes Lloyd on behalf of the Minister of the Interior, and countersigned by fhewDirectonofiGame and)Fisheries; Provinceot Manitoba 25. 4. sss ase ee ee he ee ee ee eee 170 W. E. Saunders (right) relating some experience to his old friends Taverner (left) and Fleming (Centre) at the American Ornithologists’ Union annual stated meeting at Charleston, South Carolina, November 1937. ........ 179 Taverner photographing a Semipalmated Plover at Bird, Manitoba, on the newly constructed Hudson Bay Railway LTS ORR Ae SN Re WP Hire Sy tena etry dea te ised mse al ede en Pee eae hicase Acari ateiicras smal Mend suse awa reper ara none 181 Taverner in retirement with his favourite dog Sinbad, taken in 1944 or 1945 at 45 Leonard Street. .................. 195 A Life with Birds: Percy A. Taverner, Canadian Ornithologist, 1875-1947 JoHN L. CRANMER-BYNG | Preface and Acknowledgments Introduction Part I Early years Chapter 1 Percy Fowler Chapter 2 Percy Taverner Chapter 3 Getting a start in life Part II Apprenticeship Years Chapter 4 Ornithologist-in-the-making Chapter 5 From amateur to professional Part Ill Challenge of the National Musuem Chapter 6 Museum development 1911-1914 Chapter 7 Museum problems 1915-1919 Chapter 8 Field collecting Chapter 9 Bird protection Chapter 10 Birds of eastern Canada Part IV Ornithology in a wider perspective Chapter 11 Birds of western Canada Part I Chapter 12 Birds of western Canada Part II Chapter 13 Meeting of the American Ornithologists Union, Ottawa, 1926 Chapter 14 Strains at the centre: Percy Taverner and Dr. Anderson Chapter 15 The widening field of studies I, 1928-1936 Chapter 16 The widening field of studies IH, 1936-1942 Chapter 17 In retirement Selected bibliography of Taverner’s published writings Appendix 1. Letter from Soper to Taverner, 18 July 1929 Appendix 2. Meeting on Baffin Island Natural History Collection Appendix 3. Memorandum from Anderson to Camsell End notes Index About the author: John L. Cranmer-Byng Editor’s acknowledgments List of Figures Mailing date of the previous issue 109(4) : 22 February 1996 ISSN 0008-3550 THE CANADIAN FIELD-NATURALIST Volume 110, Number 1 1996. } | | 1 253) | 254) Inside Back Cover’ The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 110, Number 2 April-June 1996 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patron His Excellency The Right Honourable Roméo LeBlanc, P.C., C.C., C.M.M., C.D., Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Anthony J. Erskine Don E. McAllister Hugh M. Raup Irwin M. Brodo Clarence Frankton Stewart D. MacDonald Loris S. Russell William J. Cody W. Earl Godfrey Verna Ross McGiffin Douglas B.O. Savile Ellaine Dickson C. Stuart Houston Hue N. MacKenzie Pauline Snure William G. Dore George F. Ledingham Eugene G. Munroe Mary E. Stuart R. Yorke Edwards Thomas H. Manning Robert W. Nero Sheila Thomsen 1996 Council President: David W. Moore Ronald E. Bedford Jeff Harrison Vice-Presidents: Michael Murphy Fenja Brodo Cendrine Huemer Lee Cairnie Ann MacKenzie : : William J. Cody Patricia Narraway Recording Secretary: David Smythe Erangis RuGaok Frank Pope Corresponding Secretary: Eileen Evans Ellaine Dickson Tom Reeve , aT Colin Gaskell Jane Topping Treasurer: Gillian Marston ipiGences Chrgcrayncr Christine Hanrahan Ken Young Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. For information on Club activities telephone (613) 722-3050. 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. Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211 Copy Editor: Wanda J. Cook Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1JO | Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9 Associate Editors: Robert R. Anderson Robert R. Campbell W. Earl Godfrey Warren B. Ballard Brian W. Coad William O. Pruitt, Jr. Charles D. Bird Anthony J. Erskine Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor with the exception of book reviews which should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $23 per calendar year. Libraries and other institutions may subscribe at the rate of ) $38 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $23 includes a subscription to The Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $5.00 to cover postage. ° 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. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Date of this issue: April-June 1996 (June 1996). Back Numbers and Index Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, and Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager. Cover: Portion of the shoreline of Loyst Lake in Lennox and Addington County, Ontario, showing open rock barrens dominated by Common Juniper, Juniperus communis, and other shrubs including Bear Oak, Quercus ilicifolia. Photograph taken in July 1994 by V. R. Brownell. See article on plants at their northern limits in the granite bar- — rens area that includes this locality by V. R. Brownell, C. S. Blaney, and P. M. Catling, pages 255-259. i The Canadian Field-Naturalist — | Volume 110, Number 2 fase April-June 1996 Recent Discoveries of Southern Vascular Plants at their Northern Limits in the Granite Barrens Area of Lennox and Addington County, Ontario VIVIAN R. BROWNELL!, C. SEAN BLANEY2, and PAUL M. CATLING! 12326 Scrivens Drive, R.R.3, Metcalfe, Ontario KOA 2P0 227 Dunnett Boulevard, Belleville, Ontario K8P 4M7 Brownell, Vivian R., C. Sean Blaney, and Paul M. Catling. 1996. Recent discoveries of southern vascular plants at their northern limits in the granite barrens area of Lennox and Addington County, Ontario. Canadian Field-Naturalist 110(2): 255-259. Aristida dichotoma (Poverty-grass), Eleocharis engelmannii (Englemann’s Spike-rush), Lindernia dubia var. anagallidea (False Pimpernel), Quercus ilicifolia (Bear Oak), and Rotala ramosior (Branched Toothcup) are disjunct in the granite bar- rens of Lennox and Addington County. Quercus ilicifolia occurs in dry, open scrub communities developed on shallow soil over granite rock, and is an addition to the native shrub flora of Canada. The other taxa were found within the annual fluc- tuation zone of open, biotite-rich metasedimentary rock shorelines, and were previously known from the Carolinian zone of extreme southwestern Ontario. The nearest populations of Quercus ilicifolia and Aristida dichotoma are about 230 km to the southeast in Oneida County, New York State. The closest extant populations of the other species range from 490-575 km to the southeast in New York State and/or 370 km to the southwest in the Carolinian region of southwestern Ontario. The disjunctions are attributed to a combination of warmer microclimate due to the abundant open rock surfaces and restricted specialized habitats of open and strongly fluctuating shorelines in the barrens region. All of the species are rare in both Ontario and Canada and are also rare in many of the adjacent states. Key Words: Aristida dichotoma, Poverty-grass, Eleocharis engelmannii, Englemann’s Spike-rush, Lindernia dubia vat. anagallidea, False Pimpernel, Quercus ilicifolia, Bear Oak, Rotala ramosior, Branched Toothcup, Ontario, disjunc- tion, rare species, phytogeography. Many vascular plants present in southwestern The Lennox and Addington rock barrens is a region Ontario are absent from the central portion of the characterized by numerous lakes, wetlands and more- north shore of Lake Ontario, but present near the far eastern end of the lake (Cody 1982; Brownell et al. 1994). Others are unique in Ontario to the east- ern Lake Ontario region. Some of these are found on the limestones, sands and clays of the Lake 2+ Ontario Lowlands (Beschel 1970), while others occur slightly further east on the Frontenac Axis, a region of rugged granitic and metamorphic rocks : connecting the main portion of the Canadian Shield and the Adirondack Highlands. Among the well- known examples of these southern species at their northern limit on the Frontenac Axis of the Kingston region are Pitch Pine (Pinus rigida Miller), Shining Sumac (Rhus copallina L.), Stemless Yellow Violet (Viola rotundifolia Michx.), and Deerberry (Vaccinium stamineum L.). Recent field studies of the Lennox and Addington FIGURE 1. Southern Ontario showing the northern barrens, a region further to the west, but still within ne nto Carme iniein orto (@etnee ts), WNORL GE Sidi aeuih oF Proce Edgiard recent interpretation - see Allen et al.1990), the 3 Canadian Shield region (diagonal hatching, provid- County (Figure 1), have revealed another concen- ed by E. Haber) and the approximate region of the tration of southern species. Lennox and Addington barrens (shaded triangle). 255 256 or-less open oak woodland as well as open rock bar- rens. The open areas of exposed rock, often extending as steep slopes right to the shorelines, are a distinctive feature, and the prominence of exposed rock is clear in satellite images. The area is the easternmost extensive section of granitic rock barrens in southern Ontario. A roughly triangular area, the Lennox and Addington barrens (Figure 1), extends from West Sheffield Lake in Hungerford Township of Hastings County northeast to Mountain Grove including the top half of Sheffield Township in Lennox and Addington County, and the southern third of Kennebec Township of Frontenac County, thence southeast to Fifth Depot Lake includ- ing the northwestern corner of Hinchinbrooke Township and the southern portion of Olden Township in Frontenac County, then west across the north side of Beaver Lake north of Tamworth to West Sheffield Lake (Figure 1). This area is mapped, although not precisely, by Chapman and Putnam (1972, 1984), as “bare rock ridges and shallow till”, and is not connect- ed to the Kaladar rock barrens area to the north and west. Here we report on and discuss disjunct occur- rences of southern species in the Lennox and Addington barrens. Methods Specimens documenting the following reports are preserved in the Agriculture Canada herbarium in Ottawa (DAO) with some duplicates at the University of Michigan (MICH). Detailed location data required for research purposes may be request- ed from the Ontario Ministry of Natural Resources offices, Natural Heritage Information Centre, or herbarium curators. Significant Southern Disjunctions Aristida dichotoma Michx., Poverty-grass Approximately 35 plants occurred along the shore of Puzzle Lake (Catling & Brownell 20569; Catling and Brownell s.n. [sine numero = without number] 12 August 1994 and at least 200 along the shore of Sheffield Long Lake (Catling & Brownell s.n. 8 October 1994; Catling and Brownell 20530). The only species of Aristida previously reported from eastern Ontario is the provincially rare A. longispica Poir. (Three-awn), which is locally abundant in the sandy pannes of Presqu’ile Provincial Park (Catling & Brownell s.n., DAO). Poverty-grass is very simi- lar, but differs in having the central segment of the lemma awn spirally twisted instead of bent, and in having longer lower glumes 6-8 mm long. Although documented from two railway yards in southwestern Ontario (Catling et al. 1978; Dore and McNeill 1980), A. dichotoma was not admitted to the Ontario rare plant atlas (Argus et al. 1982—1987) or to the list of rare vascular plants of Canada (Argus and Pryer 1990), because it was not known from a natural plant community. However, the recently discovered dis- THE CANADIAN FIELD-NATURALIST Vol. 110 jJunct sites in the Lennox and Addington barrens are in natural communities strongly dominated by native species and including many rare and restricted native species (see below and V. R. Brownell 1995. A bio- logical inventory and evaluation of the Puzzle Lake Area of Natural and Scientific Interest. Draft report prepared for the Ontario Ministry of Natural Resources, Tweed District, Tweed, Ontario. 98 pages + 3 maps). They are the only known native occurrences of this species in Canada. Associates along the open, biotite-rich metasedimentary rock shoreline (Figure 2) near the high water mark includ- ed Bulbostylis capillaris (L.) C. B. Clark and Agrostis scabra Willd. The plants occurred in the very shallow soil in small rock crevices. The nearest potentially natural site is apparently about 230 km southeast in the vicinity of Syracuse, New York on open, sandy soil (Catling and Brownell, personal observations; Paine 1865). This Atlantic coastal plain species is documented from only one location in Michigan (Voss 1972), but is known from 29 counties in the southeastern portion of New York State (New York Flora Association 1990). Eleocharis engelmannii Steud., Engelmann’s Spike-rush ' This species was first reported from Ontario in 1987 from four locations in Essex and Norfolk Counties (Ball 1987). Currently, two sites are known in Essex County and five sites have been document- ed in Norfolk County (D. A. Sutherland 1995, per- sonal communication). These closest populations of E. engelmannii occur about 370 km to the southwest of the Lennox and Addington County barrens. All of the sites have small populations, usually with less than 30 plants. On the shore of Puzzle Lake (Catling & Catling 20566), it occurred with Lindernia dubia var. anagallidea in cracks on flat shoreline rocks below the high water level. This population included 34 plants within an area of 50 m?. Other occurrences in southwestern Ontario have been on wet, open, sandy or muddy shores. In the United States, E. engelmannii is a widespread species, but is rare in several states, including adjacent Michigan, Ohio and New York (Ball 1987). It is also rare in the provinces of Manitoba, Saskatchewan and Alberta, and is consid- ered imperiled in Canada (Argus and Pryer 1990). In New York State, only one extant population, consist- ing of about 30 to 40 plants, is known (S. Young 1995, personal communication). This site is in Suffolk County approximately 575 km southeast of the Lennox and Addington County barrens. Lindernia dubia (L.) Pennell var. anagallidea (Michx.) Cooperrider, False Pimpernel This variety, known previously from southern British Columbia (Straley et al. 1985) and, recently, from the Carolinian region of southwestern Ontario 1996 BROWNELL, BLANEY AND CATLING: SOUTHERN PLANTS IN GRANITE BARRENS 257 FiGuRE 2. Open rocky shoreline habitat of Aristida dichotoma and Quercus ilicifolia in Lennox and Addington County, Ontario. Photograph taken in August 1994 by V. R. Brownell. (Oldham 4507, TRTE; Heagy 1993; Sutherland 1987; Sutherland 9351 (DAO); Goodban et al. 1994; and specimen at UWO, Oldham, personal communi- cation), is considered rare in Canada (Argus and Pryer 1990). On the open shores of Puzzle Lake (Catling & Catling 20567, 20599a; Catling & Brownell s.n. 12 August 1994), it occurred in rock cracks that were submerged during the early part of the year. Two populations were discovered each including approximately 50 plants. Associates includ- ed Eleocharis engelmannii and Rotala ramosior. This is a widespread taxon in the United States, ranging from New Hampshire to Washington, and southward from Florida to California (Pennell 1935, map 31). In Michigan it is considered rare. The New York Natural Heritage Program does not track this taxon despite its relatively restricted historical occur- rence in only six counties (S. Young, personal com- munication, 1995; New York Flora Association 1990). It is distinguished from Lindernia dubia var. dubia by its long peduncles, relatively short leaves and open flowers. With the largest leaves 10 mm long or less and pedicel/bract ratios exceeding 1.6, as well as brownish-yellow seeds ca. 0.3 mm long, the plants from Puzzle Lake are the most extreme in the direction of var. anagallidea yet collected in eastern Canada. The rank of variety has been consid- ered appropriate due to intergradation with L. dubia in the upper Mississippi valley (Cooperider and McCready 1975; Cooperider 1976). The nearest locations appear to be those in southwestern Ontario and those in Ulster County, New York State (New York Flora Association 1990) which are both about 370 km away from the newly discovered disjunct stations. Quercus ilicifolia Wang., Bear Oak The discovery of this shrub at several locations within an area of approximately 8 km? in the Lennox and Addington barrens, represents an addition to the flora of Canada. Locations include Little Gull Lake (Brownell & Catling 20492; Catling & Brownell s.n. 12 August 1994), Loyst Lake (Blaney s.n. 1, 2 July 1994; Brownell & Catling 20602), Puzzle Lake (Catling & Brownell s.n. 12 August 1994), Sheffield Long Lake (Catling & Brownell 20518) and Wheeler Lake (Brownell, Larson & Blaney s.n. 23 September 1994). The total population size within this area is estimated to range from 800 to 1000 plants, with individual sites, separated by a few hundred metres to a few kilometres, having a maximum of 300 plants. The plants occur in dry, open scrub commu- nities developed on very shallow soil over biotite granite rock, and are most often on or near to shores. The most frequent associate is Common Juniper (Juniperus communis L.). 258 Bear Oak is largely confined in North America to the Atlantic coastal plain and northern Appalachian region (Little 1977). The reports of Q. ilicifolia from “Take Huron” (Gibson and Macoun 1875a, b) are evidently based on a 1874 collection by J. Gibson (MTMG). The specimen is considered to have an incorrect label (Ambrose et al. 1987). The nearest stations to the Lennox and Addington barrens are the remnants of the Rome pine plains about 230 km to the south at the eastern end of Oneida Lake in New York State (Paine 1865; C. J. Sheviak, personal communication), which are also disjunct from the main range. Rotala ramosior (L.) Koehne, Branched Toothcup This plant is critically imperiled in Canada (Argus and Pryer 1990). It occurs in southern British Columbia and was found for the first time in Ontario in Norfolk County in 1984 (Oldham and Sutherland 1987). It is unlikely to be extant in southwestern Ontario, however, since the two areas have been converted into a corn field and pastureland (D. A. Sutherland, personal communication, 1995). At Puzzle Lake (Catling & Brownell 20599) and Sheffield Long Lake (Catling & Brownell s.n. 2 August 1994, October 1994), it occurred in rock crevices along the open shore near the water line and had been under water for at least several weeks dur- ing the spring and early summer. Associates includ- ed Cladium mariscoides (Muhlenb.) Torrey, Eleocharis elliptica Kunth and Lindernia dubia var. anagallidea. Thirty-three plants were seen along the shore of Sheffield Long Lake and over 50 occurred along the shore of Puzzle Lake. The nearest extant site is about 490 km to the southeast in Putnam County, New York State (New York Flora Association 1990; S. Young, personal communication, 1995). It is ranked as imperiled in New York with seven extant sites known, and it is considered rare in Michigan. Discussion The prominence of rock in this area undoubtedly contributes to a warmer microclimate than is experi- enced by surrounding forested regions. The lack of forest over much of the landscape probably also con- tributes to the relatively broad annual fluctuations in water level. Three species, Eleocharis engelmannii, Lindernia dubia var. anagallidea and Rotala ramo- sior appear to often occur in similar habitats and sometimes in association with each other in north- eastern North America. These habitats are usually open, periodically wet sand. The openness of the landscape, presence of numerous open shorelines, and low nutrient substrates are habitat conditions not represented to the same degree in surrounding regions in Ontario. Limestone and glacial till sub- strates exist a few kilometres to the south and extend THE CANADIAN FIELD-NATURALIST to the Lake Ontario shore and around the west side | of the lake. Granite barrens are much more restricted | further to the east on the Frontenac Axis (Chapman | and Putnam 1972, 1984). Consequently, we attribute | the disjunctions to a combination of warmer micro- climate due to the abundant open rock surfaces and restricted specialized habitats of open and strongly fluctuating low nutrient shorelines in the barrens region. Possibly these species were more widespread during the middle Holocene warm and dry interval 8—4 thousand years ago which was followed by tem- perature decrease over the past four thousand years © and southward shifts of southern forest elements (Anderson 1995). The nearest populations of Quercus ilicifolia and Aristida dichotoma are about 230 km to the southeast in Oneida County, New York State. The closest extant populations of the other species range from 490-575 km to the south- east in New York State and/or 370 km to the south- west in the Carolinian region of southwestern Ontario . Acknowledgments M. J. Oldham and D. A. Sutherland of the Natural Heritage Information Centre in Peterborough, Ontario, kindly provided information on Eleocharis engelmannii, Lindernia dubia var. anagallidea and Rotala ramosior in southwestern Ontario. C. J. Sheviak and R. J. Mitchell of the New York State Museum provided information on the distribution of Bear Oak from the museum data base and card files. Steve Young of the New York Natural Heritage Program in Latham was consulted regarding the cur- rent status of some species in that state. The Ontario Ministry of Natural Resources provided funding to V. R. Brownell for a study of the Puzzle Lake area. D. Cuddy, B. Larson, and T. Norris provided valu- able comments. E. Haber provided a Quikmap file for the illustration of the Canadian Shield. Literature Cited Allen, G. M., P. F. J. Eagles, and S. D. Price. 1990. Conserving Carolinian Canada. University of Waterloo Press. 346 pages. Ambrose, J. D., P. W. Ball, and G. E. Waldron. 1987. Rare species of Fagaceae. 1 page in Atlas of the rare vascular plants of Ontario. Part 4. Edited by K. M. Pryer and G. W. Argus. National Museum of Natural Sciences, Ottawa. Anderson, T. W. 1995. Forest changes in the Great Lakes region at 5—7 ka BP. Géographie physique et Quaternaire 49(1): 99-116. Argus, G. W., K. M. Pryer, D. J. White, and C. J. Keddy Editors. 1982-1987. Atlas of the rare vascular plants of Ontario. 4 parts. National Museum of Natural Sciences, Ottawa. Argus, G. W., and K. M. Pryer. 1990. Rare Vascular Plants in Canada. Canadian Museum of Nature, Ottawa. 191 pages + maps. Ball, P. W. 1987. Eleocharis engelmannii Steud. 1 page in Atlas of the rare vascular plants of Ontario. Part 4 Vol. 110. \\ 1996 Edited by K.M. Pryer and G. W. Argus. National Museum of Natural Sciences, Ottawa. Beschel, R. E., A. E. Garwood, R. Hainault, I. D. Macdonald, S. P. van der Kloet, and C. H. Zavitz. 1970. List of the vascular plants of the Kingston region. Fowler Herbarium, Queen’s University, Kingston, Ontario. Brownell, V. R., P. M. Catling, M. J. Oldham, and C. S. Blaney. 1994. New distributional records in relation to the phytogeography and floristic diversity of the eastern Lake Ontario region. The Michigan Botanist 33: 53-65. Catling, P. M., A. A. Reznicek, and J. L. Riley. 1978. Some new and interesting grass records from southern Ontario. Canadian Field-Naturalist 91: 350-359. Chapman, L. J., and D. F. Putman. 1972 [reprinted 1984]. Physiography of the eastern portion of southern Ontario. Ontario Department of Mines and Northern Affairs, map 2227. Chapman, L. J., and D. F. Putnam. 1984. Physiography of southern Ontario. Ontario Geological Survey Special Volume 2. 270 pages + map P2715. Cody, W. J. 1982. A comparison of the northern limits of distribution of some vascular plant species found in southern Ontario. Le Naturaliste canadien 109: 63-90. Cooperider, T. S. 1976. Notes on Ohio Scrophulariaceae. Castanea 41: 224-226. Cooperider, T. S., and G. A. McCready. 1975. On sepa- rating Ohio specimens of Lindernia dubia and L. ana- gallidea (Scrophulariaceae). Castanea 40: 191-197. Dore, W.G., and J. McNeill. 1980. Grasses of Ontario. Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario. 566 pages. Gibson, J., and J. Macoun. 1875a. The botany of the eastern coast of Lake Huron. Canadian Journal of Science, Literature and History 14: 88-100. Gibson, J., and J. Macoun. 1875b. The plants of the east- ern coast of Lake Huron, and their distribution through the northern and western portions of British North America. Canadian Journal of Science, Literature and History 14: 635-657. Goodban, A. G., J. S. Pringle, and A. E. Heagy. 1994. The vascular plant flora of the regional municipality of BROWNELL, BLANEY AND CATLING: SOUTHERN PLANTS IN GRANITE BARRENS 259 Hamilton-Wentworth, Ontario. Hamilton Region Conservation Authority, Ancaster, Ontario. Heagy, A. E., Editor. 1993. Hamilton-Wentworth natural areas inventory. Volume 2: site summaries. Hamilton Naturalists’ Club, Hamilton, Ontario. Little, E.L., Jr. 1977. Atlas of United States trees. Volume 4: minor eastern hardwoods. Forest Service, United States Department of Agriculture, Miscellaneous Publication 1342. 17 pages + 230 maps. New York Flora Association. 1990. Atlas of the New York State Flora. Edition 1. New York Flora Association, The New York State Museum Institute, Albany, New York. Oldham, M. J., and D. A. Sutherland. 1987. Rotala ramosior (L.) Koehne. | page in Atlas of the rare vascu- lar plants of Ontario, part 4. Edited by K. M. Pryer and G. W. Argus. National Museum of Natural Sciences, Ottawa, Ontario. Paine, J. A., Jr. 1865. Catalogue of plants found in Oneida County and vicinity. 18th Annual Report on the State Cabinet. University of New York. Pages 53-192. Pennell, F. W. 1935. The Scrophulariaceae of eastern temperate North America. Monograph of the Academy of Natural Sciences of Philadelphia 1: 1-650. Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. The Rare Vascular Plants of British Columbia. Syllogeus 59, (National Museum of Natural Sciences, Ottawa, Ontario). 165 pages. Sutherland, D. A. 1987. Annotated checklist of the plants of Haldimand-Norfolk. Pages 1-152 in The natural areas inventory of the regional municipality of Haldimand- Norfolk. Edited by M. E. Gartshore, D. A. Sutherland and J. D. McCracken. Norfolk Field Naturalists, Simcoe, Ontario. Svenson, H. K. 1939. Monograph studies in the genus Eleocharis - V. Rhodora 41: 1-19, 43-77, 90-110. Voss, E. G. 1972. Michigan Flora. Part 1. Gymnosperms and Monocots. Cranbrook Institute of Science Bulletin 55 and University of Michigan Herbarium. 488 pages. Received 20 March 1995 Accepted 20 July. 1995 Additions and Range Extensions to the Vascular Plant Flora of the Northwest Territories, Canada WILLIAM J. CODY Centre for Land and Biological Resources Research, Agriculture Canada, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Cody, William J. 1996. Additions and range extensions to the vascular plant flora of the Northwest Territories, Canada. Canadian Field-Naturalist 110(2): 260-270. Seventeen taxa are reported new to the flora of the Northwest Territories north of latitude 60°N, plus nine new to the District of Keewatin north of latitude 60°N (excluding the islands of southern Hudson Bay and James Bay), one to the | District of Mackenzie, and 29 range extensions within the District of Keewatin, 12 range extensions within the District of Mackenzie, two deletions from the Northwest Territories and 12 comments on distribution since previous publications on the flora. Key Words: Additions, range extensions, vascular plants, Northwest Territories. Since the publication of //lustrated Flora of the Canadian Arctic Archipelago (Porsild 1964) and Vascular Plants of Continental Northwest Territories, Canada (Porsild and Cody 1980) contin- ued field and taxonomic studies have revealed the presence of additional taxa in the Territories that were unknown to the authors at the time of publica- tion. These include both introduced and native taxa. The preparation of a manuscript on the rare plants of the Northwest Territories (McJannet et al. 1995) spurred the examination of a mass of material that had accumulated since the publication of Porsild and Cody (1980). The present paper reports these taxa and also a series of range extensions of taxa that were known from the Territories but are now known to be much more widespread. In addition, several taxa which have been reported elsewhere, two of which were unfortunately missed by Porsild and Cody (1980), are reported here in order to bring these additional taxa together in one place. Synoptic list by Northwest Territories status Taxa new to the Northwest Territories north of latitude 60°N (17) Alisma plantago-aquatica var. americana Allium fistulosum Arabis holboellii var. secunda lyrata pinetorum Botrychium simplex spathalatum Carex prairea Danthonia spicata Descurainia incisa ssp. incisa Epilobium hornemanii Tsoetes lacustris Poa pseudoabbreviata Potamogeton obtusifolius Senecio ogotorukensis Veronica longifolia Viola selkirkii Taxa new to the District of Keewatin north of latitude 60°N (9) Botrychium lunaria Cerastium regelii Deschampsia brevifolia Draba norvegica Gymnocarpium jessoense ssp. parvulum Minuartia dawsonensis Moneses uniflora Utricularia minor Viola renifolia var. brainerdii Taxon new to the District of Mackenzie (1) Deschampsia paramushirensis Range extensions of taxa in the District of Keewatin north of latitude 60°N (29) Astragalus eucosmus Betula nana ssp. exilis Calamagrostis purpurascens Caltha palustris var. arctica Chrysanthemum integrifolium Draba crassifolia Dupontia fisheri ssp. psilosantha Equisetum fluviatile Erigeron elatus Galium trifidum Harrimanella hypnoides Juncus arcticus Luzula wahlenbergii Lycopodium clavatum var. monostachyon complanatum Pedicularis hirsuta Petasites frigidus ssp. frigidus frigidus ssp. palmatus Sagittatus Pyrola minor Ranunculus gmelinii . pallasii sabinei 260 1996 Ribes glandulosum Rubus idaeus Triglochin maritimum Utricularia intermedia Viburnum edule Woodsia alpina Range extensions of taxa in the District of Mackenzie (12) Arabis arenicola Botrychium lunaria Draba lactea Oxytropis viscida Papaver mcconnellii Phegopteris connectilis Salix petiolaris Sibbaldia procumbens Silene acaulis ssp. acaulis f. albiflora Solidago graminifolia var. major Subularia aquatica ssp. americana Viola epipsila ssp. repens Deletions of taxa from the flora of the Northwest Territories (2) Danthonia intermedia Saxifraga tenuis Comments on taxa in the flora of the Northwest Territories (12) Adoxa moschatellina Braya glabella ssp. purpurascens Descurainea sophioides Erigeron alpiniformis Monarda fistulosa ssp. menthifolia Pleuropogon sabinii Podistera macounii Potamogeton subsibiricus Ranunculus cymbalaria Rorippa crystallina (Nasturtium crystallinum) Salix raupii sphenophylla Annotated list by family LYCOPODIACEAE Lycopodium clavatum L. var. monostachyon Hook. & Grev., Common Club-moss — KEEWATIN: pro- tected south slopes and ridges, in willow thickets, very local, small colonies, Griffin Lake, 61°17’N 98°47°W, K.L. Reading s.n., July-August 1990 (DAO).* Previously known in southern mainland District of Keewatin from a single locality (Porsild and Cody 1980; Cody and Britton 1989). Lycopodium complanatum L., Ground Cedar — KEEWATIN: protected south slopes sand ridges, in willow thickets, very local, small colonies, Griffin Lake, 61°17°N 98°47°W, K.L. Reading s.n., July- August 1990 (DAO). Previously known in southern mainland District of Keewatin from only two localities (Porsild and Cody 1980). *Herbarium acronym (Holmgren et al. 1990) Copy: VASCULAR PLANT FLORA OF THE NORTHWEST TERRITORIES 261 ISOETACEAE Isoetes lacustris L. (I. macrospora Durieu), Quillwort — MACKENZIE: Travaillant Lake, 67°41°N 131°48’W, Strange & MacDonell s.n., 30) July, 198s, (CAN), | (determined, iby D. M. Britton). This site is about 3100 km northwest of sites in northwestern Ontario known to Cody and Britton (1989); there are, however, intermediate sites in northwestern Saskatchewan: Stony Rapids, Fond du Lac River, 59°16’N 105°46’W, G. Argus 487-63, 30 July 1963 (CAN) (determined by D. M. Britton; this latter site was included in the rare plants of Saskatchewan (Maher et al. 1979) and it is also known from Hidden Bay of Wollaston Lake, Collins Bay of Wollaston Lake and northeast of Cantara Lake on the south shore of Lake Athabasca (all determined by D. F. Brunton-V. L. Harms (personal communication, 1995). New to the Northwest Territories north of 60°N latitude. EQUISETACEAE Equisetum fluviatile L., Water Horsetail — KEE- WATIN: solitary small tundra rock pond, abundant population, east of South Bay of Griffin Lake, 61°17’N 98°47 W, K.L. Reading s.n., 5 August 1990 (DAO). Known from only one other locality in the south of mainland District of Keewatin (Porsild and Cody 1980). OPHIOGLOSSACEAE Botrychium lunaria (L.) Sw., Moonwort — KEE- WATIN: alders and open white spruce/tamarack parkland, sandy-wet, east side of South Bay of Griffin Lake, 61°17’N 98°47°W, K. L. Reading s.n., 5 August 1990 (DAO); wet Arctostaphylos ground cover in open tamarack parkland, local, common where found, west of South Henik Lake, 61°22’N 975530 Wight Reading sun). 6) sulys 990: MACKENZIE: along portage trail, Bloody Falls, Coppermine River, photo by K. J. Hebden, 2 August 1980 (DAO). The District of Keewatin sites are intermediate between a site at Churchill, Manitoba and Fort Smith in southern District of Mackenzie; new to mainland District of Keewatin; the District of Mackenzie site represents a northeastward extension of the known range of some 550 km from the south shore of Great Bear Lake (Porsild and Cody 1980; Cody and Britton 1989). Botrychium simplex E. Hitche., Simple Grape-fern — MACKENZIE: Great Bear Lake, Atacho Point, 66°N 121°30’W, A.E. & R.T. Porsild 3468A, 24 August 1928 (CAN) (determined by W. H. Wagner). This is a northward extension of the known range of some 900 km from a site near the North Saskatchwan River in central Alberta (Cody and 262 Britton 1989; Packer 1983). New to the flora of the District of Mackenzie and the Northwest Territories north of 60°N latitude. Botrychium spathulatum W.H. Wagner — MACKENZIE: Nahanni National Park, near base of “Beehive Mountain”, 61°33’N 125°23’W, G. W. Scotter 24197, 10 July 1977 (DAO). This specimen which is a paratype of Wagner’s new species (Wagner and Wagner 1990) is the only record yet known from the Northwest Territories. ASPIDIACEAE Gymnocarpium jessoense (Koidz.) Koidz. ssp. parvulum Sarvela, Nahanni Oak-fern — KEE- WATIN: very rare, one site only on a N-facing low scarp, Culleton Lake area, 61°20’N 97°40’°W, K. L. Reading s.n., late July 1991 (DAO). New to mainland District of Keewatin; to the south it is rare in northern Manitoba; Harms et al. (1992) knew this species from 20 localities in northern Saskatchewan south to approximately 55°N latitude and considered it borderline vulnerable in that province. Phegopteris connectilis (Michx.) Watt (Dryopteris phegopteris (L.) Chr. Thelypteris phegopteris (L.) Slosson), Long Beech Fern — MACKENZIE: drainage beds under boulder fields in shade of large willow and alder bushes, rare, vicinity of Daring Lake, 64°52’N 111°37’°W, McNair & O’Brien 2, 11 August 1994 (DAO); not common, Eastern Great Slave Lake Region, Porter Lake, 61°42’N 108°02’W, Ovenden & Rowe 240, 6 July 1977 (DAO); common in “trickle irrigation” bouldery drainageway, same locality, Ovenden & Rowe 683, 13 August 1977 (DAO). This rare species was first reported from the Northwest Territories by Cody and Porsild (1968) on the basis of a collection from the Canada Tungsten Mine area adjacent to the Yukon Territory border. Cody et al. (1979) reported a second locality in the Nahanni National Park area of the southern Mackenzie Mountains. The Porter Lake collections cited above were plotted by Cody and Britton (1989) but no reference was made to the locality and habitat in the text. The Daring Lake locality which is the most northeasterly yet reported in the District of Mackenzie lies some 350 km north northwest of Porter Lake. To the south it has been collected at the east end of Lake Athabaska in Saskatchewan (Porsild and Cody 1980; Cody and Britton 1989) and seven additional localities mapped by Harms et al. (1992*) who considered it vulnerable in that province. Woodsia alpina (Bolton) S.F. Gray, Alpine Woodsia — KEEWATIN: “Pebble Beach” Lake, 62°22’N *See Document Cited Section THE CANADIAN FIELD-NATURALIST Vol. 110 97°30’ W, south of Yathkyed Lake, K. Reading 85-5, 28 June 1985 (DAO). This is the third collection of this taxon from cen- tral mainland District of Keewatin (Porsild and Cody 1980). Korol (1992) reported it from coastal District of Keewatin in the vicinity of Rankin Inlet. POTAMOGETONACEAE Potamogeton obtusifolius Mert. & Koch. (P. porsil- diorum sensu Porsild and Cody (1980) pro parte), Blunt-leaved Pondweed — MACKENZIE: in shal- low water, Mile 12.75 Mackenzie River- Yellowknife Hwy., Thieret & Reich 7858, 20 July 1961 (DAO); in small lake, Mile 38.3S Mackenzie River- Yellowknife Hwy., Thieret & Reich 8359, 2 August 1961 (DAO); in shallow water at disturbed edge of small pool, Mile 45.5S Mackenzie River- Yellowknife Hwy., Thieret & Reich 7947, 23 July 1961 (DAO); KEEWATIN: west of Bissett Lake, K. L. Reading s.n., 31 August 1982 (photo DAO). This is a more or less circumboreal species which in North America occurs from Newfoundland to British Columbia and southward. It is new to flora of Northwest Territories. Potamogeton subsibiricus Hagstr. (P. porsildiorum Fern.) — MACKENZIE: Anderson River Delta, T.W. Barry 411, 18 August 1961 (DAO, CAN); brackish lagoon, Liverpool Bay, Nicholson Island, TOON Q9OWS ALE? & RIT: “Porsild 28/6. lis= 16 August 1927 (CAN); shallow stagnant pool in muskeg, Mackenzie River Delta, East Branch, 68°40-55’N, A.E. Porsild 7242, 11 August 1934 (CAN); in shallow sluggish stream, Mackenzie Delta, East Branch,. Hansen’s Creek, approx. 68°10’°N, A.E. Porsild 7364A, 2 October 1934 (CAN); Aubry Lake, 67°20’N 126°25’W, Riewe & Marsh 237, 18 July 1976 (CAN). This taxon is known in the District of Mackenzie only in the extreme northwest. Specimens collected by Thieret and Reich north of Yellowknife mapped by Porsild and Cody (1980) as P. porsildiorum have been revised to P. obtusifolius (see above). SCHEUCHZERIACEAE Triglochin maritimum L., Arrow-grass — KEE- WATIN: rare, single colony in wet hollow in tundra, “Nocamp Lake’, 62°21°N 97°267W.; south of Yathkyed Lake, K. Reading 85-37 s.n., 9 July 1985 (DAO); rare, very local, Cullaton Lake area, 61°20’N 97°40’W, K.L. Reading s.n., 1991 (DAO). These are the third and fourth collections of this taxon in southern mainland District of Keewatin (Porsild and Cody 1980). ALISMATACEAE Alisma plantago-aquatica L. var. americana J. A. Schultes (A. triviale Pursh), Water-plantain — MACKENZIE: in mud, shallow pools near water- falls, Hanging Ice (Tethul) River junction with 1996 Talston River, 60°34’N 112°13’W, M. Anions s.n., 9 August 1980 (DAO). Cody (1956) reported this taxon from Fort Fitzgerald, in northern Alberta as a northern exten- sion of the known range northward from Chipewyan on Lake Athabaska (C.C. Loan 205 (DAO)); the specimen cited above is a further extension of the known range of some 90 km; new to the flora of the Northwest Territories. POACEAE (GRAMINEAE) Calamagrostis purpurascens R.Br., Purple Reedgrass — KEEWATIN: 10 km NW of town on moist tundra, Rankin Inlet, 62°49’N 92°05’W, J. B. Korol 176, 21 July 1988 (DAO). This collection was reported by Korol (1992) as C. deschampsioides Trin., a species which is much more delicate and has a lax, pyramidal panicle. Calamagrostis purpurascens was known to Porsild and Cody (1980) from a single collection in south- western District of Keewatin some 300 km SSE of Rankin Inlet. Danthonia spicata (L.) Beauv. ex Roemer & J.A. Schultes (D. intermedia sensu Scotter & Cody 1974), Poverty Oat Grass. Scotter and Cody (1974) reported D. intermedia as new to the District of Mackenzie on the basis of a collection by Scotter in Nahanni National Park along trails near hotsprings, 61°15’N 124°03’W, (12285A, 26 June 1970 (DAO)). This and nine additional specimens from the park were revised to D. spicata by Darbyshire (Darbyshire and Cayouette 1989), thus deleting D. intermedia from the flora of Continental Northwest Territories and replacing it with D. spicata. Deschampsia brevifolia R.Br., Hairgrass — KEE- WATIN: Rasmussen Lowlands, slope with low hummocks, on the west side of Iglulik Lake where the Inglis River drains out, 68°497°18.5”N 92°26°04"W, V. Johnston 63Al1b, 1 July 1994 (DAO). This is an extension of the known range south- southwest of some 100 km from a site plotted by Porsild and Cody (1980) on the southeastern side of Boothia Peninsula. It is the first record for mainland District of Keewatin. Deschampsia paramushirensis Honda (D. pumila (Trin.) Ostenf.) — MACKENZIE: Mackenzie River Delta, Richards Island, collected on a dry lakebed, lake drained in 1978, L. Ovenden 2257, 8-15 August Osa: Porsild and Cody (1980) mapped the distribution of this species (sub D. pumila) in North America as known to them. McLachlan et al. (1988) give the range and habitat and discuss it (sub D. para- mushirensis) in relation to D. caespitosa. The speci- men cited above is intermediate between a site on the arctic coast of Alaska at about longitude 157°W Copy: VASCULAR PLANT FLORA OF THE NORTHWEST TERRITORIES 263 and sites on the arctic islands of the District of Franklin and in central mainland District of Keewatin. It is the first known collection from the District of Mackenzie. Dupontia fisheri R.Br. ssp. psilosantha (Rupr.) Hultén, Tundra Grass — KEEWATIN: Rasmussen Lowlands, low, flat, wet, numerous small ponds, polygonal ground approx. 10 km E of Inglis Bay, 68°36°55°N 93°14°28"W, V. Johnston SAI, 17 June 1994 (DAO). This collection represents a northwestward exten- sion of the known range of some 500 km from a site at Rankin Inlet plotted by Porsild and Cody (1980). To the west it is known from the southeastern end of Victoria Island in District of Franklin and from the vicinity of Bathurst Inlet in northwestern District of Mackenzie. Pleuropogon sabinii R.Br., Semaphore Grass This species was first reported from mainland District of Keewatin by Cody et al. (1989) on the basis of a collection of George Scotter in the vicinity of Wager Bay. Korol (1992) extended the known range southward to Rankin Inlet. Poa pseudoabbreviata Roshev. — MACKENZIE: dry graminoid tundra, Northern Richardson Mountains, 68°03°31”N 135°57°57°W, Loewen & Staniforth 93-285A, 12 July 1993 (DAO). Cody et al. (1990) reported this species as new to the flora of Canada based on specimens collected by C. E. Kennedy in the British Moutains in Ivavvik (Northern Yukon) National Park and Cody (1994) has reported an additional collection from that area. The specimen cited above extends the known range southeastwards some 175 km to northwestern District of Mackenzie. CYPERACEAE Carex prairea Dewey, Prairie Sedge — MACKEN- ZIE: in woods in wet bog, 3.3 mi. S and E along Mackenzie Hwy. (from Liard Ferry), 61°04’N 121°11’W, N. A. Skoglund 708, 11 June 1973 (deter- mined by J. H. Hudson) (DAO). According to the map in Packer (1983), this is an extension of the known range northward from cen- tral Alberta. New to the flora of the District of Mackenzie and the Northwest Territories. This species has also recently been reported new to the Yukon Territory on the basis of a collection by H. Raup in the extreme southeast (Cody 1994). JUNCACEAE Juncus arcticus Willd., Arctic Rush — KEEWATIN: Rasmussen Lowlands, low and flat with many ponds, some hummocks, some elevated dry areas, near Chantrey Inlet, 67°14°08.6”N 95°10°11.9"W, V. Johnston 47B2, 3 July 1994 (DAO). This is a northward extension of the known range in mainland District of Keewatin of some 264 300 km from sites plotted by Porsild and Cody (1980). Luzula wahlenbergii Rupr., Wahlenberg’s Woodrush — KEEWATIN: Rasmussen Lowlands, hummocks mixed with rock rubble and scattered frost boils, east of the upper reaches of the Castor and Pollux river, 68°12’46”"N 93°17°21”"W, V. Johnston 36A7, 26 June 1994 (DAO); low and flat with many ponds, some hummocks and some elevated dry areas, near Chantrey Inlet, 67°47’°08.6”"N 95°10°11.9"W, V. Johnston 47A4, 3 July 1994 (DAO); wet hum- mocky plain with lots of frost-heaved rock, east side of the Paquet River, 68°12’N 93°16’W, V. Johnston 440-2-3, 5 August 1993 (DAO). These collections, which are intermediate between a site on the Arctic Coast at the east end of the Queen Maud Gulf and a site on the southeast side of the Melville Peninsula, are the northern most yet found in the District of Keewatin. LILIACEAE Allium fistulosum L., Welsh Onion — MACKEN- ZIE: living plants collected by Gilles Patenaude and Joe Mackenzie on dry gravelly S-facing slope 20 miles SE of Rae-Edzo and | mile inland from Great Slave Lake in August 1989 and grown in a garden at Yellowknife by Patenaude. In June 1992 transplanted to Sooke, British Columbia where he took up residence. Herbarium specimen (in seed) collected by Patenaude, 19 June 1994 and sent to Ottawa for determination and preservation (DAO); living plants sent to Ottawa September 1993 by Patenaude and grown in garden by Cody - specimen in flower, (Cody 35605, 23 June 1994 (DAO). On 27 October 1994 Patenaude advised me by phone that there were about 50 plants observed at the site southeast of Rae- Edso where the living plants were collected. The same day Joseph Mackenzie, a native employee of the Department of Renewable Resources at Rae- Edso, told me that this onion plant was growing at the site of old Fort Rae and that he also knew of additional sites on nearby islands. The suggestion was made that this onion had been introduced by members of a mission or by employees of the Hudson Bay Company. New to the flora of the Northwest Territories. SALICAEAE Salix petiolaris J. E. Smith (S. gracilis Anders.), Meadow Willow — MACKENZIE: well-drained gravel, Mackenzie Delta wellsite Gulf-Mobil Siku E- 21, 69°00°29"N 133°36’54”W, D.W. Smith 48, 11 August 1980 (DAO). The above collection represents a northern exten- sion of range of some 500 km from Norman Wells. Salix raupii Argus, Raup’s Willow Argus (1986) reported this species as new to the Northwest Territories on the basis of specimens from THE CANADIAN FIELD-NATURALIST Vol. 110 southern Nahanni National Park and Fisherman Lake, District of MacKenzie. It was described by him (1974) on the basis of collections from adjacent to the Alaska Highway in northeastern British Columbia and has since also been found at Fern Lake (57°45’N 124°47’W) in British Columbia and at Nose Mountain (54°30’N 119°32’W) and Kakwa Falls (54°05’N 119°40’W) in western Alberta (Argus 1986). It should be searched for elsewhere in the region. Salix sphenophylla Skvotsov, Wedge-leaf Willow — MACKENZIE: moist tundra, Cape Dalhousie, 70°13’N 129°40°W, W.J. Cody 13129, 31 July 1963 (DAO) (det. G.W. Argus). This specimen was mapped and cited by Argus (1973) but unfortunately S. sphenophylla was over- looked by Porsild and Cody (1980). BETULACEAE Betula nana L. ssp. exilis (Sukatsch.) Hultén (B. glandulosa sensu Porsild & Cody 1980 pro parte), Dwarf Birch — KEEWATIN: Rasmussen Lowlands, low and flat with many small ponds, near Chantrey Inlet, 67°14’08.6”"N 95°10°11.9"W, V. Johnston 47A5, 47A6, 3 July 1994 (DAO). This is a northward extension of the known range of some 300 km from sites plotted by Porsild and Cody (1980). CARYOPHYLLACEAE Cerastium regelii Ostf. — KEEWATIN: wet soil with occasional drier tussock, Rasmussen Lowlands, approx. 4 km south of Dewline, 68°45’°10”"N 93°25°39"W, V. Johnston 462-3-3, 7 August 1993 (DAO). New to mainland District of Keewatin. Porsild and Cody (1980) knew this species, which is com- mon throughout the Canadian Arctic Archipelago, from only two sites on the Canadian mainland: Tuktoyaktuk and Melville peninsulas. The location cited here is south of the Boothia Peninsula. In addi- tion to the Tuktoyaktuk Peninsula locality C. regelii is now also known from the Melville Hills region in northern District of Mackenzie (Cody et al. 1992). Minuartia dawsonensis (Britt.) House, Rock Sandwort — KEEWATIN: Cullaton Lake area, 61°20’N 97°40’ W, K.L. Reading s.n., 1991 (DAO). : Not previously recorded from mainland District of Keewatin; to the west the nearest sites are at the east end of Great Slave Lake in District of Mackenzie, to the southwest, the north shore of Lake Athabaska in Saskatchewan, and to the south in the vicinity of Churchill, Manitoba (Porsild and Cody 1980). Argus (1966) also reported it from the Hasbala Lake area in extreme northeastern Saskatchewan (sub. Arenaria stricta ssp. dawsonensis). Silene acaulis L. ssp. acaulis f. albiflora Hartz, Moss-Campion — MACKENZIE: near Burnside 1996 River, 66°51’N 108°20’°W, G.W. Scotter 96603, 7 July 1992 (DAO). The type specimen of this white-flowered form was collected in Greenland. It is of rare occurrence across northern Canada. RANUNCULACEAE Caltha palustris L. var. arctica (R.Br.) Huth., Marsh Marigold — KEEWATIN: Rasmussen Lowlands, low and flat with numerous small ephemeral ponds, approx. 5 km NW of mouth of Inglis River, GsesorlirouN) 93°36°22:7° WV: Johnston 63A5, 10 July 1994 (DAO). This is the most northeasterly locality yet known in Canada. It has been found some 150 km to the south southwest in District of Keewatin and to the west on King William Island in District of Franklin (Porsild and Cody 1980). Ranunculus cymbalaria Pursh, Northern Seaside Buttercup — KEEWATIN: Chesterfield Inlet, Dutilly 6602W, 16 aoit 1938 (QFA). This record for mainland District of Keewatin which was reported by Blondeau and Cayouette (1987) was an extension of the known range of some 550 km northward from the vicinity of Churchill, Manitoba. It has since been reported by Korol (1992) from Rankin Inlet, about 60 km to the south- west. Ranunculus gmelinii DC.,- Yellow Water Crowfoot — KEEWATIN: Rasmussen Lowlands, low and flat peat, approx. 5 km NW of the mouth of Inglis River, 68°387°17.5”N 93°36'22”W, V. Johnston 63A6, 10 July 1994 (DAO). This collection represents a northward extension of the known range of some 500 km from sites in central District of Keewatin (Porsild and Cody 1980). To the west it is known from the southeastern end of Victoria Island in District of Franklin and Rankin Inlet in northwestern District of Mackenzie (Porsild and Cody 1980). Ranunculus pallasii Schlecht., Pallas’ Buttercup — KEEWATIN: Rusmussen Lowlands, low and flat with many ponds, near Cantrey Inlet, 67°14’08.6"N 95°10°11.9"W, V. Johnston 47A1, 3 July 1994, low- centred polygons, at south end of Murchison Lake, 68°05’43.8"N 92°39°22.8"W, V. Johnston 3, 1994 (DAO); in peat in turfy centre of high-centred poly- gons, approx. 15 km SE of Kinngaarjuit Hill south of the Murchison River, 68°24'13.2”N 93°05’°15.8°W, V. Johnston 5I1A1, 5 July 1994 (DAO). These collections represent a northwestward extension of the known range of some 650 km from Chesterfield Inlet (Savile and Calder 1952), and an eastward extension of the known range of some 800 km from a site west of Melville Sound adjacent to Coronation Gulf in District of Mackenzie. Korol Copy: VASCULAR PLANT FLORA OF THE NORTHWEST TERRITORIES 265 (1992) reported an additional site in the District of Keewatin from Rankin Inlet, some 60 km southwest of Chesterfield Inlet. Ranunculus sabinei R.B., Sabine Buttercup — KEE- WATIN: Fly Camp 3, 65°53’N 98°34’E, Baker Lake area, S. Smith 204, 20 July 1983 (DAO). This collection represents an extension of the known range southward from the northern coastal region of some 225 km. PAPAVERACEAE Papaver mcconnellii Hultén, McConnell’s Poppy — MACKENZIE: crustose lichen on stone talus, Northern Richardson Mountains, 68°05’N 136°22°17°W, Loewen & Staniforth 93-006, 4 July 1993 (DAO). Hultén (1968) knew this endemic species from only two localities in the Yukon Territory. It is how- ever much more frequent there (Cody 1994) and its status as a rare plant in that area should be reviewed. Previously known in the District of Mackenzie from a single collection on the Yukon-Mackenzie border at 67°33’N 136°12’W by J. A. Calder (Cody and Porsild 1968). BRASSICACEAE (CRUCIFERAE) Arabis arenicola (Rich.) Gelert — MACKENZIE: on steep gravelly slope above lake, vicinity of south end of Contwoyto Lake, 65°45’N 111°15’W, J. W. Thieret 9408, 21-24 August 1962 (DAO); dry sandy areas, beaches, vicinity of Daring lake, 64°52’N 111°37’°W, McNair & O’Brien 31, 4 August 1994 (DAO). The collections cited here extend the known range of this arctic species of eastern North America for- merly known to occur in the District of Mackenzie to longitude 110°W (Porsild and Cody 1980). Arabis holboellii Hornem. var. secunda (Howell) Jepson (A. holboelii var. retrofracta sensu Porsild & Cody (1980) pro parte), Holboell’s Rockcress — MACKENZIE: shallow soil in hollows of rock, Yellowknife near Giant Mine, Cody & McCanse 2352, 30 June 1949 (DAO); shallow soil in crevice of rocky hillside, Yellowknife, Joliffe Island, 62°27'N 114°23’W, Cody & McCanse 2479, 7 July 1949 (DAO); exposed sand slope of roadside, 24 miles south of Lower Hay River, 60°40’N 116°00’W, W.H. Lewis 322, 13 June 1951 (DAO); dry sand-ridge, exposed, Moraine Point, Great Slave Lake, 61°36’N 115°38’W, W. H. Lewis 464, 29 June 1951 (DAO). This taxon, which occurs in the Gaspé Peninsula, Quebec, central Ontario, Saskatchewan to British Columbia and southern Alaska, is occasional in south- ern Yukon Territory north to Dawson; it has not pre- viously been reported from the Northwest Territories. Arabis lyrata L., Lyre-leaved Rockcress — MACKENZIE: pure sand by roadside, 1 mile west 266 of Seven Mile Lake, 28 miles west of Fort Smith, Cody & Loan 4665, 22 July 1950 (DAO) (deter- mined by G. A. Mulligan) previously determined as A. lyrata L. var. kamchatica Fisch. and A. kamchati- ca (Fisch.) Ledeb. Arabis lyrata is a North American species which extends from Ontario to Alberta, barely entering southwestern District of Mackenzie while A. kam- chatica is an amphiberingian species which extends eastwards across Alaska and the Yukon Territory to the Mackenzie Mountains of western District of Mackenzie and the east side of Great Bear Lake and southward through British Columbia to southwestern Alberta and the state of Washington. Arabis lyrata is new to the flora of the Northwest Territories. Arabis pinetorum Tidestrom (A. holboelii var. retrofracta sensu Porsild and Cody (1980) pro parte) — MACKENZIE: in shallow soil of crevice in igneous rock, Yellowknife, Cody & McCanse 2085, 11 June 1949 (DAO); dry sand ridge, Moraine Point, Great Slave Lake, 61°36’N 115°38’W, W. H. Lewis 463, 29 June 1951 (DAO); shallow humus over rock, Fort Reliance, 62°13’N 109°10’W, W. J. Cody 14739, 12 August 1965 (DAO) (determined by G. A. Mulligan). This taxon which is found from Manitoba to British Columbia, Yukon and Alaska and south into northwestern United States has not previously been reported from the Northwest Territories. Braya glabella Richards. ssp. purpurascens (R.Br.) Cody (B. purpurascens R.Br.), Low Braya — KEE- WATIN: Rankin Inlet, J.B. Korol 156 (SASK). Korol (1992) reported this circumpolar subspecies as new to mainland District of Keewatin. The nearest site on Southampton Island is about 300 km to the northeast. Descurainia incisa (Engelm. ex Gray) Britton ssp. incisa (D. richardsonii (Sweet) O.E. Schulz ssp. incisa (Engelm. ex Gray) Detting), Tansy-mustard — MACKENZIE: rare in heavy soil by airstrip, Norman Wells, Cody & Gutteridge 7462a, 22 July 1953 (DAO) (determined by G. A. Mulligan). This taxon was growing with D. incana (Bernh. ex Fischer & C.A. Meyer) Dorn (Cody & Gutteridge 7462b) (D. richardsonii (Sweet) O.E. Schulz). According to Rollins (1993) it occurs in “mountains and foothills, Alberta to New Mexico, west to California, north to British Columbia”. The northernmost collection seen from British Columbia was one mile west of Borolder Creek on Pine Pass-Dawson Creek Highway, approx. 55°37’N 121°58’W, Calder & Kukkonen 27007, 14 July 1960 (DAO). It is probably intro- duced at Norman Wells. Descurainia sophioides (Fisch.) O.E. Schulz, Northern Tansymustard — KEEWATIN: vicinity of Rankin Inlet, K. Reading 94-47, 1994 (DAO). THE CANADIAN FIELD-NATURALIST Vol. 110 This is an amphi-Beringian species which in the District of Mackenzie extends eastwards to the east end of Great Slave Lake and in the northeast, south- western District of Franklin and is disjunct to eastern Baffin Island and northeastern Manitoba. The locali- ty cited above which is the only one known in the District of Keewatin was previously reported by Korol (1992). Draba crassifolia Graham, Rocky Mountain Whitlow-grass — KEEWATIN: talus below scarp W of 30-Mile Lake, 63°40’N 97°10°W, K. Reading s.n., 1 September 1982 (DAO). Previously known in the District of Keewatin from a single collection in the central region just south of the site reported here. Draba lactea Adams, Milky Whitlow-grass — MACKENZIE: open sandy areas, top of esker, vicin- ity of Daring Lake, 64°52’N 111°37°W, McNair & O’Brien 32, 22 August 1994 (DAO) (determined by G. A. Mulligan). This circumpolar species in the District of Mackenzie was previously known adjacent to the Arctic Coast and Richardson and Mackenzie moun- tains and southward into northern British Columbia where it is rare. The collection cited above repre- sents an extension of the known range of some 280 km southwest from the head of Bathurst Inlet and some 360 km southeast from the vicinity of Coppermine. Draba norvegica Gunn. — KEEWATIN: Cullaton Lake area, 61°20’N 97°40’W, K.L. Reading s.n., 1991 (DAO). New to mainland District of Keewatin; Mulligan and Cody (1968) reported the presence of D. norvegica near the east end of Great Slave Lake, a disjunct occurrence from Southampton Island, James Bay and the west coast of Quebec of some 1600 km; the collection cited here is intermediate between these two areas. Rorippa crystallina Rollins, Asiatic Cress This taxon was described by Rollins (1962) on the basis of specimens collected by John Thieret and Robert Reich along the Yellowknife Highway north- west of Fort Providence. It is now considered to be an introduction belonging to the genus Nasturtium and G. A. Mulligan has made the transfer to that genus, Nasturtium crystallina (Rollins) Mulligan (Mulligan and Cody 1995). Subularia aquatica L. ssp. americana Mulligan & Calder, Awlwort — MACKENZIE: sandy shoreline, common, Tsu Lake, 60°35’N 111°52’W, M. Anions s.n., 17 August 1980 (DAO). This species is known in the District of Mackenzie from only four other localities: Great Bear Lake, (Porsild 1943), Indin Lake, Yellowknife (Cody 1956) and LeHaise Lake. 1996 SAXIFRAGACEAE Ribes glandulosum Grauer, Skunk Currant — KEE- WATIN: very rare in rock scarps, Cullaton Lake area, 61°20’N 97°40’°W, K.L. Reading s.n., late August 1991 (DAO). Not previously known from mainland District of Keewatin, but known to the south in adjacent Manitoba and to the west in southern District of Mackenzie Porsild and Cody, 1980). Saxifraga tenius (Wahl.) H. Sm., Slender Arctic Saxifrage. Korol (1992) reported this species as new to Continental District of Keewatin. His collections from Rankin Inlet [47 (SASK) and 57 (SASK, DAO] have since been revised to S. nivalis L., a cir- cumpolar, arctic species which was mapped by Porsild and Cody (1980) from five localities in mainland District of Keewatin. ROSACEAE Rubus idaeus L. s.1. (R. strigosus Michx.), Wild Raspberry — KEEWATIN: in willow thickets along tundra streams, very local, north of Griffin Lake, 61°17 N 98°47°W, K.L. Reading s.n., 2 August 1990 (DAO). Previously known in the District of Keewatin from only two localities, one at the south end of James Bay and one on the mainland near the locality cited here (Porsild and Cody 1980). Sibbaldia procumbens L., Sibbaldia — MACKEN- ZIE: in mossy mats on lakeshore, vicinity of Daring Lake, 64°52’N 111°37°W, McNair & O’Brien 374A, 12 August 1994 (DAO). This is a circumpolar species with large gaps in distribution. In the District of Mackenzie it is fre- quent in the Mackenzie Mountains but east of the Mackenzie River was previously known from a sin- gle collection at the west end of Great Bear Lake and then disjunct to southern mainland District of Keewatin (Porsild and Cody 1980). FABACEAE (LEGUMINOSAE) Astragalus eucosmus Robins., Eligant Milk-vetch — KEEWATIN: single colony on old raised lakebed gravel flat, “Nocamp” Lake, south of Yathkyed Lake, 62°21’N 97°26’ W, K. Reading 85- 17, 15 July 1985 (DAO); small colonies “Pebble Beach” Lake 62°22’N 97°30’ W, south of Yathkyed Lake, K. Reading 85-26, 17 July 1985 (DAO); vicinity of Meliadine Lake, 63°01’°N 92°10°51°W, K. Reading 94-26, 1994 (DAO); rock outcrop, 5 km NW of town of Rankin Inlet, 62°49°N 92°05’ W, J.B. Korol 171, 19 July 1988 (DAO, SASK). Korol (1992) reported the first collection from mainland District of Keewatin. The additional col- lections cited above demonstrate that it is more widespread in the Territory. Copy: VASCULAR PLANT FLORA OF THE NORTHWEST TERRITORIES 267 Oxytropis viscida Nutt. s.l. (O. glutinosa A.E. Porsild), Sticky Locoweed — MACKENZIE: near “Fishing Creek”, Bathurst Inlet, 66°40’N 108°51°W, P.M. Burt s.n., 9 July 1992 (DAO). This collection represents an extension of the known range of some 650 km NE from the SW side of Great Bear Lake (Porsild and Cody 1980). VIOLACEAE Viola epipsila Ledeb. ssp. repens (Turcz.) Becker, Dwarf Marsh Violet — MACKENZIE: along a small stream with a cover of alder and willows, near the Burnside River, 66°20’N_ 109°22’W, G.W. Scotter 96601, 1 July 1992 (DAO). This collection represents an extension of the known range NE from the south side of the Horn Plateau of some 750 km and ESE from the Richardson Mountains of some 1150 km. Viola renifolia Gray var. brainerdii (Greene) Fern., Kidney-leaved Violet — KEEWATIN: sandy open forest and wet Salix thickets along stream, Cullaton Lake area, 61°20’N 97°40’°W, K.L. Reading s.n., early July 1991 (DAO). New to the District of Keewatin; to the south this taxon is known from the vicinity of Churchill, Manitoba (Porsild and Cody 1980). Viola selkirkii Pursh ex Goldie, Selkirk’s Violet — MACKENZIE: Kraus Hotsprings, 61°14’N 124°02°W;S:S: Talbot sn... 23 July 1975) (CAN, photo DAO) (determined by J. G. Packer); rapidly drained bedrock substrate in Carex-Dryas alpine community, Gibson Ridge, Chick Lake, 65°53’N 128°05’W, D. Gubbe 201, 23 July 1973 (CAN). New to the flora of the Northwest Territories north of 60°N latutide. ONAGRACEAE Epilobium hornemanii Reichenb., Hornemann’s Willowherb — MACKENZIE: hidden in stand of Epilobium near creek, Hot Springs, one mile south of Canada Tungsten Mine, Mackenzie Mountains, 61°58’N 128°15’°W, K.W. Spicer 1506a, 23 July 1967 (DAO). This taxon is widespread in southern Yukon Territory (Cody 1994) and was expected to be found in adjacent District of Mackenzie (Porsild and Cody 1980). APIACEAE (UMBELLIFERAE)) Podistera macounii (Coult. & Rose) Mathias & Const. (Ligusticum macounii Coult. & Rose; L. mutellinoides auth. non (Crantz.) Willar, Macoun’s Lovage. Welsh (1974) reported this tiny umbelliferous plant for northern Yukon and the adjacent Richardson Mountains in northwestern District of Mackenzie, on the basis of collections made by him in that area. It was unfortunately not included in Porsild and Cody (1980). The basis of this report is 268 MACKENZIE: on Jurassic Bug Creek Sandstone, West Cache Cr., 68°20’N 136°26’W, S.L. Welsh 12012, 20 June 1973 (CAN, BYU, photo DAO); ridge top in alpine tundra along West Cache Creek, 68°16°N 136°22’W, Welsh & Rigby 12039, 20 June 1973 (BYU, photo DAO). PYROLACEAE Moneses uniflora (L.) Gray, One-flowered Pyrola — KEEWATIN: in white spruce copses on tundra, pro- tected places, deep moss, very local, small colonies, north of Griffin Lake, 61°02’N 98°12’W, K.L. Reading s.n., 15 August 1990 (DAO). New to mainland District of Keewatin. Pyrola minor L., Lesser Wintergreen — KEE- WATIN: riverside white spruce thickets, deep moss or grass, not common, Griffin Lake, 61°17’N 98°47°W, K.L. Reading s.n., 10 August 1990 (DAO). Previously known in mainland District of Keewatin from a single nearby locality; to the west, three localities near the Mackenzie-Keewatin border were plotted by Porsild and Cody (1980). ERICACEAE Harrimanella hypnoides (L.) Coville (Cassiope hyp- noides (L.) D.Don), Moss Heather KEEWATIN: SSW of Bissett Lake, 63°45’N 95°30’°W, K. Reading s.n., 28 August 1982 (DAO); snowbank habitat on east shore of lake, “Big Bird” Lake, 62°17’N 97°36’ W, south of Yathkyed Lake, K. Reading 85- 22, 16 July 1985 (DAO); mud along stream gorge, Fly Camp 2, 65°43’N 98°53’W, Baker Lake area, S. Smith 163, 170, 14 July 1983 (DAO). Porsild and Cody (1980) knew this species in the Continental Northwest Territories from three locali- ties, two in southeastern mainland District of Keewatin and one in the Thelon Sanctuary in eastern District of Mackenzie. The specimens cited here are intermediate between these two areas. LAMIACEAE (LABIATAE) Monarda fistulosa L. ssp. menthifolia (Grah.) Fern., Wild Bergamot — MACKENZIE: rocky exposed northwest-facing river bank, 3.5 miles up the Sainville River from Arctic Red River, 133°50’N 66°30’W, R. Shotton s.n., 23 June 1972 (UBC, CAN). This taxon was reported as new to the Northwest Territories by Straley (1986) on the basis of the col- lection cited above. SCROPHULARIACEAE Pedicularis hirsuta L., Hairy Lousewort — KEE- WATIN: dry to moist tundra, Fly Camp 2, 65°43’N 98°53’W, Baker Lake area, S$. Smith 149, 13 July 1983 (DAO); dry to moist tundra, Fly Camp 3, 65°53’N 98°34’E, Baker Lake area, S. Smith 189, 18 July 1983 (DAO). In mainland District of Keewatin this species was only known to Porsild and Cody (1980) from adja- THE CANADIAN FIELD-NATURALIST Vol. 110 cent to Queen Maud Gulf in the north and Hudson Bay in the east. Veronica longifolia L., Speedwell — MACKENZIE: along sidewalk, disturbed ground, small wooded area, Fort Smith, 60°00’N 111°53’W, M. Anions, s.n., 4 September 1980 (DAO). This is a garden escape which is apparently rare in western Canada, but is folhd established more fre- quently in eastern Canada. New to the flora of District of Mackenzie and Continental Northwest Territories. LENTIBULARIACEAE Utricularia intermedia Hayne, Flat-leaved Bladderwort — KEEWATIN: rare - wet fen - one site only, Cullaton Lake area, 61°20’N 97°40’ W, K.L. Reading s.n., July 1991 (DAO). Previously known in mainland District of Keewatin from only two sites in the southern part. Utricularia minor L., Small Bladderwort — KEE- WATIN: north of east end of Griffin Lake, 61°17°N 98°42’°W, K. L. Reading s.n., 1990 (DAO). Previously known in the District of Keewatin only from the south end of James Bay. The nearest recorded sites are north of Lake Winnipeg in Manitoba, south of Lake Athabaska in Saskatchewan and at the east end ef Great Slave Lake in the District of Mackenzie (Porsild and Cody 1980). It is also known from the Nero Lake region north of Lake Athabaska, LaLocke, Wathaman Lake west of Reindeer Lake, and Trade Lake on the Churchill River in northern Saskatchewan (V.L. Harms, personal communication, 1995). In the northern part of its range this species is nearly always sterile, as is the present collection, and easily overlooked. RUBIACEAE Galium tridifum L., Small Bedstraw — KEE- WATIN: local, Cullaton Lake area, 61°20’N 97°40’W, K. L. Reading s.n., 1991 (DAO). Previously known in the south of the District of Keewatin from only two other localities (Porsild and Cody 1980). CAPRIFOLIACEAE Viburnum edule (Michx.) Raf., Low-bush Cranberry — KEEWATIN: locally common, Cullaton Lake area, 61°20’ W, 97°40’W, K.L. Reading s.n., 1991 (DAO). Previously known in mainland District of Keewatin from a single locality west of Cullaton Lake (Porsild and Cody 1980). Adoxaceae Adoxa moschatellina L., Moschatel — MACKEN- ZIE: Liard River Valley, east bank on terrace slope just above mouth of Kotanellee River, W.W. Jeffrey 134, 10 July 1959 (CAN); wet silt under Salix back from lake shore, Fisherman Lake, 60°20’N 123°47’W, S.M. Lamont FL23, 7 June 1973 (DAO); 1996 growing on lowest lake terrace in a moist area beneath Salix, Fisherman Lake, 60°20’N 123°48’W, S.M. Lamont FLA84, 21 June 1974 (DAO). The specimens cited above are the basis for the District of Mackenzie locality mapped by Porsild and Cody (1980). Hultén and Fries (1986) mapped a second locality south of Great Slave Lake adjacent to the Slave River wk:ch has been repeated by McJannet et al. (1995). The basis for this record has not been found but it might possibly be a misplace- ment of a site in the Caribou Mountains west of Wood Buffalo National Park at approximately 59° latitude in northern Alberta (Lee et al. 1982). COMPOSITAE Chrysanthemum integrifolium Richards., Entire- leaved Daisy — KEEWATIN: moist tundra, Fly Camp 3, 65°53’N 98°34’W, Baker Lake area, S. Smith 195, 20 July 1983 (DAO). Porsild and Cody (1980) knew this species in the extreme north of mainland District of Keewatin. The specimen cited here is from a site some 200 km to the south in the interior. Erigeron alpiniformis Crong., Fleabane — FRANKLIN: dry west facing slope on middle basin of lakes, Ogac Lake, Frobisher Bay, .A. McLaren 113, 1962 (CAN). Cronquist (1947) knew this taxon, which he sepa- rated from E. uniflorus and E. alpinus, only from Greenland at the time he described it. It is new to Baffin Island and the Northwest Territories and was mapped without comment by McJannet et al. (1993). Erigeron elatus (Hook.) Greene — KEEWATIN: Cullaton Lake area, 61°20’N 97°40’W, K.L. Reading s.n., 1991 (DAO); Padley River [Padlei River 61°53’N 96°42’W], W.C. Giissow s.n., 9 August 1932 (DAO). Previously known in mainland District of Keewatin from a single site west of Cullaton Lake (Porsild and Cody 1980). Petasites frigidus (L.) Fries ssp. frigidus, Sweet Coltsfoot — KEEWATIN: ubiquitous, Cullaton Lake area, K.L. Reading s.n., 1991. Previously known in the District of Keewatin from only one locality in the extreme northwest. Petasites frigidus (L.) Fries ssp. palmatus (Ait.) Cody (P. palmatus (Ait.) Gray), Palmate-leaved Coltsfoot — KEEWATIN: ubiquitous, Cullaton Lake area, 61°20’N 97°40’W, K.L. Reading s.n., 1991 (DAO). This is the third collection of this taxon in south- ern mainland District of Keewatin (Porsild and Cody 1980). Petasites sagittatus (Banks) Gray, Arrow-leaved Coltsfoot — KEEWATIN: ubiquitous, Cullaton Lake area, 61°20’N 97°40’W, K.L. Reading s.n., 1991 (DAO); “Pebble Beach” Lake, 62°22’N Copy: VASCULAR PLANT FLORA OF THE NORTHWEST TERRITORIES 269 97°30’ W, south of Yathkyed Lake, K. Reading 85- 24, 17 July 1985 (DAO). These are the second and third collections of this taxon from southern mainland District of Keewatin. Senecio ogotorukensis Packer (S. conterminus sensu Hultén (1968) pro parte), Groundsel — MACKEN- ZIE: Mackenzie Mountains, top of steep eroding west-facing slope, Mirror Lake, east side of Carcajou Range, 64°52’N 126°55’W, Cody & Brigham 20404, 3 July 1972 (DAO). Cody (1994) extended the known range of this taxon into northern Yukon Territory; the collection cited here extends the known range into western District of Mackenzie. New to the flora of the Northwest Territories. Solidago graminifolia (L.) Salisb. var. major (Michx.) Fern., Goldenrod — MACKENZIE: rocky shoreline: Tsu Lake: 6093 50N) “hiss 2 Ww: D. Campbell, s.n., 19 August 1980 (DAO). To the south this species has been reported about Lake Athabaska in both Saskatchewan and Alberta (Raup 1936 - sub var. camporum (Greene) Fern.). Elsewhere in the District of Mackenzie S. graminifo- lia var. major 1s known only in the Mackenzie River valley between Fort Simpson and Fort Wrigley (Porsild and Cody 1980). Acknowledgments I thank Ken Reading, Toronto, for the many speci- mens he gathered during his wanderings in the District of Keewatin, Vicky Johnston, Canadian Wildlife Service, Environment Canada, Yellowknife, for her contributions from the Rasmussen Lowlands, District of Keewatin, Johanne Tungilik, Northwest Territories Renewable Resources, Yellowknife for making the specimens collected by Brenda McNair and Chris O’Brien at Daring Lake, District of Mackenzie, available for my examination, Gerald Mulligan for his determina- tions of Cruciferae specimens and Jacques Cayouette and Vernon Harms for their careful reviews of an earlier manuscript of this work. Documents Cited (marked with * in text) Harms, V. L., P. A. Ryan, and J. A. Haraldson. 1992. The rare and endangered native vascular plants of Saskatchewan. Manuscript prepared for the Saskatchewan Natural History Society. Literature Cited Argus, G. W. 1966. Botanical investigations in North- eastern Saskatchewan: the subarctic Patterson-Hasbala Lakes Region. Canadian Field-Naturalist 80(3): 119-143. Argus, G. W. 1973. The Genus Salix in Alaska and the Yukon. National Museums of Canada, Ottawa, Publications in Botany, Number 2. 279 pages. Argus, G. W. 1974. A new species of Salix from Northern British Columbia. Canadian Journal of Botany 52: 1303-1304. 270 Argus, G. W. 1986. Salix raupii, Raup’s Willow, new to the Flora of Alberta and the Northwest Territories. Canadian Field-Naturalist 100: 386-388. Blondeau, M., and J. Cayouette. 1987. Extensions d’aire dans la flore vasculaire du Nouveau-Québec. Le Naturaliste canadien 114: 117-126. Cody, W. J. 1956. New plant records for Northern Alberta and Southern Mackenzie District. Canadian Field-Naturalist 70: 101-130. Cody, W. J. 1994. The Flora of the Yukon Territory: additions, range extensions and comments. Canadian Field-Naturalist 108(4): 428-476. Cody, W. J., and D. M. Britton. 1989. Ferns and Fern Allies of Canada. Agriculture Canada, Research Branch, Publication 1829/E. 430 pages. Cody, W. J., S. J. Darbyshire, and C. E. Kennedy. 1990. A Bluegrass, Poa pseudoabbreviata Roshev., New to the Flora of Canada, and Some Additional Records from Alaska. Canadian Field-Naturalist 104: 589-591. Cody, W. J., and A. E. Porsild. 1968. Additions to the Flora of Continental Northwest Territories, Canada. Canadian Field-Naturalist 82: 263-275. Cody, W. J., G. W. Scotter, and S.S. Talbot. 1979. Additions to the Vascular Plant Flora of Nahanni National Park, Northwest Territories. Le Naturaliste canadien 106: 439-450. Cody, W. J., G. W. Scotter, and S.C. Zoltai. 1989. Vascular plant flora of the Wager Bay Region, District of Keewatin, Northwest Territories. Canadian Field- Naturalist 103(4): 551-559. Cody, W. J., G. W. Scotter, and S.C. Zoltai. 1992. Vascular Plant Flora of the Melville Hills Region, Northwest Territories. Canadian Field-Naturalist 106: 87-99. Cronquist, A. 1947. Revision of North American species of Erigeron, north of Mexico. Brittonia 6: 121-302. Darbyshire, S. J., and J. Cayouette. 1989. The Biology of Canadian Weeds. 92. Danthonia spicata (L.) Beauv. in Roem. & Schult. Canadian Journal of Plant Science 69: 1217-1233. Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum Part I: The Herbaria of the World. New York Botanical Garden, Bronx, New York. 193 pages. Hultén, E. 1968. Flora of Alaska and Neighboring Territories. Stanford University Press, Stanford, California. 1008 pages. Hultén, E., and M. Fries. 1986. Atlas of North European Vascular Plants North of the Tropic of Cancer. 3 Volumes. Koeltz Scientific Books, K6nigstein, Federal Republic of Germany. 1172 pages. Korol, J. B. 1992. The Vascular Plant Flora of Rankin Inlet, District of Keewatin, Northwest Territories. Canadian Field-Naturalist 106(3): 342-347. Lee, P.G., R.A. Ellis, and P.L. Achuff. 1982. Vegetation and flora of the Caribou Mountains, Alberta. 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Packer, J.G. 1983. Flora of Alberta. Second Edition. University of Toronto Press, Toronto. 687 pages. Porsild, A. E. 1943. Materials for a flora of the continen- tal Northwest Territories of Canada. Sargentia 4: 1-79. Porsild, A. E. 1964. Illustrated Flora of the Canadian Arctic Archipelago, (Second Edition, revised). National Museum of Canada Bulletin 146. 218 pages. Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Continental Northwest Territories, Canada. National Museum of Natural Science, Ottawa. 667 pages. Raup, H. M. 1936. Phytogeographic studies in the Athabaska-Great Slave Lake Region. I. Catalogue of the Vascular Plants. Journal of the Arnold Arboretum 17: 180-315. Rollins, R. C. 1962. A new crucifer from the Great Slave Lake area of Canada. Rhodora 64: 324-327. Rollins, R. C. 1993. The Cruciferae of Continental North America. Stanford University Press, Stanford, California. 976 pages. Savile, D. B. O., and J. A. Calder. 1952. Notes on the Flora of Chesterfield Inlet, Keewatin District, N.W.T. Canadian Field-Naturalist 66: 103-107. Scotter, G. W., and W. J. Cody. 1974. Vascular Plants of Nahanni National Park and Vicinity, Northwest Territories. Le Naturaliste canadien 101: 861-891. Straley, G. B. 1986. Wild Bergamot, Monarda fistulosa (Lamiaceae), New to the Northwest Territories. Canadian Field-Naturalist 100: 380-381. Wagener, Jr., W. H., and F.S. Wagner. 1990. Notes on the Fan-Leaflet Group of Moonworts in North America with Descriptions of Two New Members. American Fern Journal 80: 73-81. Welsh, S. L. 1974. Anderson’s Flora of Alaska and Adjacent Parts of Canada. Brigham Young University Press, Provo, Utah. 724 pages. Received 11 April 1995 Accepted 12 August 1995 Ecological Replacement of the Deer Mouse, Peromyscus maniculatus, by the White-footed Mouse, P. leucopus, in the Great Lakes Region CHARLES A. LONG Departments of Biology and Wildlife Management, and Museum of Natural History, University of Wisconsin—Stevens Point, Stevens Point, Wisconsin 54481 Long, Charles A. 1996. Ecological replacement of the Deer Mouse, Peromyscus maniculatus, by the White-footed Mouse, P. leucopus, in the Great Lakes Region. Canadian Field-Naturalist 110(2): 271-277. The possible ecological replacement of the boreal, long-tailed Forest Deer Mouse Peromyscus maniculatus gracilis, by the White-footed Mouse, P. leucopus noveboracensis, in Wisconsin was studied by analysis of the zoogeography (parapatric distribution and relict populations of P. m. gracilis in the south and on islands), and by surveying relative population abun- dance on Washington Island, where competition probably commenced in 1987, upon the arrival of P. leucopus. In Wisconsin, the geographic range of P. leucopus is expanding northward, and P. maniculatus gracilis has vanished in the south and on the Door Peninsula. The White-footed Mouse thrives in brushy and dry forest habitats (often nesting in ground burrows), and the Forest Deer Mouse seems both adapted to high tree branches and cavities, to cold weather, and to boreal (conifer, mixed conifer) forests. On Washington Island, after an initial increase of P. Jeucopus and the concomitant near extinction of P. m. gracilis, both species have coexisted at low to medium densities during the mild winters of the past three years. Key Words: Peromyscus leucopus, White-footed Mouse, Peromyscus maniculatus gracilis, Forest Deer Mouse, Wisconsin, ecological replacement, islands. The exclusion of one organism from the niche of another is difficult to demonstrate (Connell 1983; Schoener 1983). Both east and west of Lake Michigan, the White-footed Mouse, Peromyscus leu- copus, and the Forest Deer Mouse, P. maniculatus gracilis, formerly occupied almost parapatric (non- overlapping) geographic ranges. In Wisconsin, the Forest Deer Mouse occupied the boreal areas sup- porting spruce, fir, and mixed conifer-deciduous forests, whereas the White-footed Mouse inhabited brushy prairies, oak savannas, forest edge, and river bottoms north to the contact. A “tension zone” or ecotone, separates these boreal and southern (austral) areas (Curtis 1959; Stearns and Kobriger 1975; Long 1974). Balsam Fir (Abies balsamea), White Spruce (Picea glauca), Eastern Hemlock (Tsuga canadensis), Paper Birch (Betula papyrifera), Sugar and Red Maple (Acer saccharum and A. rubrum), White Cedar (Thuja occidentalis)and Beech trees (Fagus grandifolia) are found in northern lower, wetter, areas. On higher ground are White Pine (Pinus strobus), Oak (Quercus rubra) and Balsam Fir. Oak (Q. macrocarpa)-maple savanna, Jack Pine (P. banksiana), birch woods, hickories (Carya cordi- formis, C. ovata), Sycamore (Platanus occidentalis) and White Pine forest characterize southern Wisconsin. The two Peromyscus species seemed adapted to different zones, and contrasting climates and vegetation apparently resulted in their north- south juxtaposition (Hooper 1942; Long 1974). Miller (1893: 63) suggested that owing to human modification and land use, P. leucopus ranged north- ward, approaching the Forest Deer Mouse in New York and western Massachusetts. Hooper (1942) found the same kind of dispersion and replacement in lower Michigan where P. m. gracilis was confined to northern counties, and P. leucopus had moved northward up the entire Lower Peninsula. Thus, eco- logical replacement is a pattern in all three separate and approximately concurrent northward invasions. The similar habitat requirements and close resem- blance of the two long-tailed species have been noted often (Long and Long 1993; Horner 1954; Wolff 1985). In addition to occurring with P. m. gracilis in the latter’s habitats, P. leucopus thrives where P. m. gracilis is seldom or never found (e.g., sandy brushland, beaches, old field, rights of way, hedge rows, prairies (Getz 1961; Long and Long 1993; Long 1974, 1978; Jackson 1961). However, P. leu- copus does not dominate in swamps and northern latitudes. Although both long-tailed species show similar patterns of thermogenesis and accumulation of brown fat (e.g., Zegers and Merritt 1988; Lynch 1973) and in some states show little loss of mass in winter, in Wisconsin at the northern edge of its range P. leucopus shows a significant loss of mass and/or heavy mortality in severe winters which are charac- terized by cold temperatures and lack of snow cover (resulting in deep frost) (Long 1973). Pierce and Vogt (1993); Tannenbaum and Pivorun (1988); and Wolff and Durr (1986) found P. maniculatus superi- or to P. leucopus in winter adaptations including tor- por, larger nests, larger hoards of food, regressed sexual organs, and lower food consumption. 271 Did: ‘Le cor "AND LA FAYETTE THE CANADIAN FIELD-NATURALIST Vol. 110 Ficure 1. Map of the region. Left: Upper boldly dashed boundary, geographic range for Peromyscus leucopus, based on recent records (black dots). Dotted boundary, range for the same species based on Jackson’s records prior to 1961 (open circles). The fine-line boundary is that for Peromyscus maniculatus gracilis. It has shown little change in his- toric times except a somewhat reduced range. Right: Door Peninsula, Green Bay, Lake Michigan, and the isles in Green Bay. Black dots and open circles as above. Bull’s-eye dots represent occurrences of P. m. gracilis throughout the islands and northward, but not on the Door Peninsula. After P. leucopus established itself on Washington Island (in 1987) and subsequently increased its densi- ty dramatically (Figure 1), apparently at the expense of P. m. gracilis, the two kinds have been studied intensively. This study examines the interrelations of two formerly parapatric long-tailed Peromyscus now inhabiting the same woodlands. Materials and Methods Museum special snap traps baited with a mixture of oats and peanut butter were set in linear series of 25-50 stations (two traps at each) at intervals of 10 m. Trapping was carried out 20 years. Each line was left out only one night, but often the line was re-run in nearby transects, on successive nights. Most specimens were preserved for future identification, but where abundance was high, only counts were made. The number of mice caught was adjusted over 100 trap nights. Sampling was usually during the autumn, occasionally in spring. Mice in hand were identified by tail length, ear length, vibrissae length, tail pencillate or not, and (in most specimens) by skull shape and length (see Long and Long 1983). Most (approximately 275) preserved specimens, except three examined from the United States National Museum (Natural History), were placed in the University of Wisconsin-Stevens Point Museum collection. Habitat types are defined as follows: 1. Sand dunes with beach grasses, spreading junipers (Juniperus sp.), legumes (Lathyrus mar- itimus), legumes, dwarf iris (/ris lacustris), wil- lows (Salix sp.) and other shrubs sparsely cover- ing the sand. 2. Old field-fence row. Tall grasses and forbs, including field composites, milkweed (Asclepias syriaca), vetch (Vicia villosa), brambles and saplings (occasionally planted Red Pine, Pinus resinosa) growing along field-stone fences. 3. Maple-beech forest with White Cedars, Oak, Fir, Paper Birch and Hemlock interspersed, and a ground flora of ferns, raspberries, young maples and beeches, chokecherries (Prunus virginiana), birches, sparse grasses, numerous rocks and rot- ted stumps and logs. Lorraine Andersen, certified by the U.S. Weather Service, supplied the weather data collected by use of standard meteorological methods. In addition to monthly records, the last five winters were compared by use of average temperatures, snowfall and snow cover. Results and Discussion Invasion of Washington Island Members of the United States Biological Survey collected mammals on Washington Island, Lake Michigan, in 1917. These workers found P. m. gra- 1996 LONG: DEER MOUSE AND WHITE-FOOTED MOUSE IN GREAT LAKES REGION 273 40 ” f= ae QO 30 z oO © 20 = S = IO 2 < Oo 0 1975 1985 1987 1989 1991 1993 1984 1986 1988 1990 1992 1994 Forest Deer Mouse—s— White-footed Mouse —e— FIGURE 2. Annual abundance of two long- Washington Island, Lake Michigan. cilis but no P. leucopus on this 14 000 acre (5600 ha) island. After eleven years of collecting only P. m. gracilis on Washington Island, P. leucopus first appeared in 1987 (Long and Long 1993). One year later it had apparently spread several miles to Boyar’s Bluff from the Pedant’s Lane forest, both similar beech- maple-oak-hemlock-cedar and birch assemblages on the north side of the Island. In subsequent years the White-footed Mouse was caught on sand dunes, along hedges and stone fences, in bird houses, and in old fields where P. m. gracilis had never been taken. In 1992, P. leucopus showed up on Hemlock Drive and at Sand Dunes beach across the Island from where it was first found. In numbers (Figure 2) and in diverse habitats (Table 1) it had increased. Concomitantly the num- bers of P. m. gracilis steadily decreased, and it was not taken at all in some dryer forest habitat where it formerly thrived (Table 1, Figure 2). Since 1987, a total of 168 P. leucopus were collected mostly in beech-maple forest, compared to 67 P. m. gracilis. Hooper (1942) reported P. m. gracilis thrives best in dense upland beech, maple, yellow birch, hard- woods, and also in bogs, brush, jackpine, dune heath, and conifer swamps (Dice 1925). P. leucopus in Michigan prefers oak-hickory woods with a diverse understory (Allen 1938; Burt 1940; Getz 1961). Batzli (1977) found P. leucopus in wet lowlands in Illinois. Long (1968) and Hoffmeister (1989) report- ed P. leucopus from brushy prairie in central Illinois. Long (1973) found them abundant in jack pine-oak tailed Peromyscus from 1976-1994, on savanna, hazelnut groves, white pine forest and wooded ravines in central Wisconsin. Howard (1951) found White-footed Mice vulnerable to cold winters, perhaps giving an ecological edge in the boreal areas to P. m. gracilis. Long (1973) found that deep frosts (resulting from thin snow cover) practi- cally extirpates P. leucopus in central Wisconsin, but the populations recover by the subsequent fall. The two species are now parapatric there. Stah (1980) found that both species prefer to, nest above ground level, in tree cavities and hollows but that P. leucopus shifts to ground nesting when com- peting with P. m. gracilis. In an earlier study, Stah (1978) found P. leucopus to be more aggressive. The Forest Deer Mouse may be better adapted to the arboreal nesting. At certain seasons when seeds and fruits, numerous arthropods, and acorns (cached by squirrels as well as mice) are available on the ground, P. leucopus seemingly out competes P. m. gracilis. The latter is superior (having well-devel- oped cheek pouches) in hoarding food in summer or winter, an advantage unless P. /eucopus steals from these caches (Pierce and Vogt 1993). Analysis of stomachs of 26 specimens taken at Pedant’s Lane indicated that P. leucopus survived the winter in the vicinity of oaks by feeding on acorns on the ground, and also ate arthropods, beech nuts, maple and grass seeds, and pits of choke- cherries. Subsequent trapping (1994) suggested adult P. m. gracilis with young were feeding on acorns from the same oaks. Clark (1972) working mostly in northern Clark County, in hemlocks and hardwoods 274 TABLE |. Percent of Washington Island Peromyscus species found in three habitats, 1975, (N = 12), 1984-1986 (N = 99), 1987-1990 and in 1991-1994 (N = 235 mice). Sand Old —=Beech-Maple Dunes Fields Forest 1975 P. leucopus 0 0 0 P. maniculatus 1) 0 100 1984-1986 P. leucopus 0 0 0 P. maniculatus 0) 0 100 1987-1990 P. leucopus 10 5) 55 P. maniculatus gracilis 0 0 29 1991-1994 P. leucopus Pall 11 50 P. maniculatus gracilis 0) 0 17 (68 percent of his specimens of P. leucopus) found in P. leucopus, that 22% of the volume was com- posed of green plants and 63% seeds, whereas in P. m. gracilis, 47% was green plants and 44% seeds. Wolff et al. (1985) when comparing the subspecies P. m. nubiterrae with P. leucopus found seasonal differences in proportions of green plants and anthropods eaten but similarity over the total yearly diet. P. maniculatus ate more arthropods in summer - and green vegetation in autumn, and P. leucopus ate more green vegetation in summer, more arthropods in autumn. Some fruits, seeds, acorns and other available for- est foods are occasionally found in the stomachs in both species. White pulpy material was usually minced acorn meat, but some of it was the white material in eggs of Carpenter Ants (Camponotus her- culeanus). In several July-taken mice, both species were feeding on Carpenter Ants. Jaws, parts of chiti- nous exoskeletons, one jaw with muscle attached, and one thin brownish covering of a pupa were observed, intermixed with the white minced pulpy material. If these mice habitually feed on Carpenter Ants, not only are they competing for a forest resource, but the ant colonies can provide food for these mice in tree cavities in most months of the year. Weather As cold winters supposedly favor P. m. gracilis and not P. leucopus, the possible correlation of snow and cold temperature and abundance of mice on Washington Island was examined (Table 2). On Washington Island, the climate in the past five years seemed mild. It was milder than the 1951 through 1980 normals. There was less water in the swamps in 1994, but in March there were only two inches (50 mm) less of snow cover (Table 2). From 1989 THE CANADIAN FIELD-NATURALIST Vol. 110 through 1993, snow cover varied from 0 to 20 inches (50 cm) in winter. In 1993-1994 there was no more snowfall and snow cover than usual, and the mean temperature was the least of the past five years. There was a low of —24° F (= —31°C). This “hard winter,” which historically was judged also to be “mild,” may have had little effect on the Peromyscus populations of either species. The effect of cold which would be expected to be adverse for P. leuco- pus may have been mitigated by the snow cover. The greatest discrepancy in abundance was in 1991 (Figure 2). That year P. m. gracilis almost disap- peared from our samples and P. leucopus peaked. Since then P. m. gracilis seemed to be holding its own though low numbers are evident for both species. A very cold winter hypothetically might favor a recovery of P. m. gracilis back to former abundance and depress P. leucopus. In contrast, a very mild winter might be expected to favor P. leu- copus, although in Kansas, where winters are milder, temperature was not important to the abundance of P. leucopus (Kaufman et al. 1995). In northern Wisconsin, a Kohn Winter Severity Index (WSI) (McCaffery 1994) was based on the number of zero or below zero °F days and 18-inch (45 cm) snowfalls, was calculated. Data were collected from 32 Department of Natural Resources stations. The winter of 1993-1994 was only the third time Lake Superior froze over in 20 years. However, the number of subzero nights was little different (41 to 42 in 1993-1994) than usual. The mean WSI was 48, which was classed as “mild”, the fourth mild rating in the past five years. Snow depths varied in the 15-17 inch (380-430 mm) range, with crusted conditions in some places. Deep snow is adverse to deer, but the deer herd was in good condition (McCaffery 1994). P. leu- copus actually benefits when the snow cover is deep (Long 1973). Thus, climatic pattern in northern Wisconsin confirms the local findings on Washington Island. The winters have not been severe. What if winters were severe? Wolff and Durr (1986) and Wolff and Hurlbutt (1982) found that P. maniculatus tended to nest high in hollow trees. Only three nest sites were underground whereas 36 were in trees. In P. leucopus 43 nests were under- ground and 36 were in trees but P. maniculatus aver- aged higher nest height in trees (7.4 m to 4.3 m in mean height). P. leucopus often moved to under- ground burrows in winter. Madison (1984) found that P. leucopus at the northern limits of its geo- graphic range is vulnerable to thin snow and deep frost in large part because it often dwells in under- ground nests. This may also account for its avoid- ance of wet areas such as swamps and marshes. The warmest winter I observed during this study was during a time when P. leucopus populations declined. More extreme temperature fluctuation than observed in recent years may affect the relative abundance, especially a combination of thin snow 1996 LONG: DEER MOUSE AND WHITE-FOOTED MOUSE IN GREAT LAKES REGION DS TABLE 2. Winter weather data for Washington Island, 1989-1994. Temperature in degrees C. Snow in inches (and mm). Index explained in text. Departure Snowfall/ Winter Date High/Low Mean Temp Norms 1951-1980 Cover Low November 1993 10.6/-8.9 7 -0.8 6/1 (152/25) December 7.2/-20 -2.7 1.2 2/1 (50/25) January 1994 2.2/-31 -9.0 -0.9 28.5/20 (723/507) February 5/-8.3 -10.5 -3.5 12/16 (305/406) March 12.2/-13.8 -1.1 0.8 9/3 (228/76) Means 1994 -4.3 -0.6 11.5/8.2 (292/207) -31 November 1992 10/-10 1.7 -0.8 6.5/2 (165/50) December 4.4/-16 -2.8 2: 9/3 (228/76) January 1993 7.8/-17 -5.8 DES 8/5 (202/126) February 6.1/-20.5 -7.3 -0.1 6/6 (152/152) March 15/-15 -0.9 1.1 3/5 (76/126) Means 1993 -3.0 0.7 6.5/4.2 (165/104) -20 November 1991 11.1/-10 0.8 Le7/ 5.5/3 (139/76) December 5.6/-15 -2.7 i) 12.5/8 (317/202) January 1992 5.6/-21 -4.8 3.2 10.5/3 (317/76) February 3.3/-21 -3.4 3.6 11/5 (279/126) March 10/-15 -1.8 0.5 9/6 (228/152) Means 1992 1.3 2.1 9.7/5 (246/126) -21 November 1990 16.7/-5 4.3 1.8 0/0 (0/0) December 2.2/-20.5 -2.9 0.9 10.5/6 (166/152) January 1991 2.8/-25 -8.5 -0.6 9/7 (228/177) February 8.9/-21.7 -4.5 2.6 5/6 (126/152) March 11.1/-12.2 -0.3 2.0 3/4 (76/101) Means 1991 -2.4 1.3 5.5/4.6 139/117) -25 November 1989 11.1/-14.4 -0.1 -2.6 8/3 (202/76) December 3.3/-25 -7.6 -3.8 22.5/10 (570/25.4) January 1990 5/-15.5 -3.2 4.8 13/10 (303/25.4) February 5.6/-18.3 -5.4 EY 7/8 (177/202) March 13.9/-16.1 -0.2 Dell 0/5 (0/126) Means 1990 -3.3 0.4 10.1/7.2 (254/180) -25 and cold temperature (Long 1973). However, in cen- tral Wisconsin the penetration of soil frost is signifi- cantly deeper than anywhere in the state (Zimmermann 1991), but P. leucopus is the most abundant small mammal there. Dry soils and savan- na plants such as Jack Pine and Beaked Hazelnut (Corylus cornuta) allow P. leucopus to thrive, but P. m. gracilis seems excluded from sand habitats. Rate of Replacement When two species have similar ecological require- ments such as food and nest sites, and similar mor- phology, then both species should increase when con- ditions are optimal and likewise both should decline when conditions are unfavorable. When one species cycles up and the other down, then superior niche exploitation might explain the difference (Figure 2). One species probably does not take over another’s habitat in a few years (Scott and Deuser 1992; Mellhopp and Lynch 1978). Fitch (1979) showed that P. m. gracilis can invade and select open ground habitats such as lichen-prairies and woodlots. On Washington Island, open habitats have apparently never been filled by P. m. gracilis. Although replacement of P. m. gracilis by P. leucopus on Washington Island may not take place soon, the results in the year 1991 suggested it might in the long term (Figure 2). Wolff (personal communication: unpublished manuscript) concludes P. leucopus gains a numerical advantage over deer mice in summer and times of food abundance. Deer mice have higher survival in winter and in times of food shortages, but fluctuating conditions are frequent enough to permit coexis- tence, in the short term. Such fluctuations probably delay ecological replacement. Biogeography (see Figure 1) In northern Clark County, and adjacent areas in Taylor, western Marathon, and southern Price coun- 276 ties, Clark (1972) found no evidence for ecological replacement. Both species were abundant except that P. leucopus was rare in wet habitats. Schmidt (1931) found that both species in nearby western Clark County preferred the same habitat, mesic hemlock- hardwoods, but P. leucopus occurred a little south- ward of P. m. gracilis. Both species in Clark County ate the same general foods (“green plants and seeds’’), suggesting competition for food (Clark 1972), but there was no evidence given of resource limitation. Jackson (1917) mentioned a southern relict popu- lation of P. m. gracilis occurring in the Sheboygan Marsh, which probably persisted there because the marsh had cool, moist habitats. Such habitats are not preferred by P. leucopus (Long 1974; Clark 1972). Jackson (1961) implied that P. m. gracilis had been eradicated there, something easier to speculate than to prove. However, two collections have been made, and only P. leucopus were taken. A second relict population has been found in Manitowoc County and it too seems surrounded by populations of P. leuco- pus, with no connections to the P. m. gracilis north- ward (Figure 1). These small populations might be southward range extensions instead of relict populations left after the main range withdrew northward. The other evidence, however, indicates a northward invasion by P. leucopus into northern Wisconsin and Upper Michigan, only recently reaching as far as Lake Superior (Long and Long 1993) and Upper Michigan (Baker 1983). The parallel movement in Lower Michigan (Hooper 1942), and an interesting pattern of distribution in Door County, Wisconsin, and near- by habitats of Upper Michigan (Long 1978) provide sound evidence of ecological replacement of P. m. gracilis by P. leucopus (Figure 1). On the Door Peninsula (Figure 1), a huge boreal thumb of land jutting into Lake Michigan, and sepa- rated from Upper Michigan by the waters of Green Bay, there are no records of boreal P. m. gracilis in spite of intensive collecting since 1910. Available habitats are everywhere, permitting other boreal mammals such as the Red Squirrel (Tamiasciurus hudsonicus), Northern Flying Squirrel (Glaucomys sabrinus) and Red-backed Vole (Clethrionomys gap- peri) to thrive in the beech-maple and fir-white pine forests. Apparently P. leucopus has been long established on the peninsula. Robert Howe and T. Erdmann of the University of Wisconsin-Green Bay showed me skulls of P. leucopus from a Door County cave. These remains were associated with other non-boreal mammals, and were dated at approximately 600 years BP, certainly no earlier than 1000 years. The Forest Deer Mouse is abundant offshore in Green Bay (Chambers Island) and north of the ship passage called Door of Death (Porta des Morts), on Detroit Island (recent specimens of P. m. gracilis taken there by Thor Purinton and myself), THE CANADIAN FIELD-NATURALIST Vol. 110 Washington Island, Rock Island, St. Martin’s Island, and Big Summer Island (see Long 1978). Either P. leucopus replaced P. m. gracilis on the Door Peninsula and has been unable to cross the waters to the nearby islands, or P. m. gracilis has colonized all these islands without successfully reaching the Door Peninsula. Acknowledgments The University of Wisconsin Personnel Development Committee Research Grant for the sum- mer of 1992, provided help for this study. A joint grant, Partners for Wildlife, of the National Wildlife Association, Wisconsin Department of Natural Resources Bureau of Endangered Species, and the Stevens Point Isaak Walton League provided support in the summer of 1994. 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Received 16 June 1995 Accepted 17 October 1995 A Contribution to the Biology of the White-beaked Dolphin, Lagenorhynchus albirostris, 11 Waters off Newfoundland DONG JIN HAt!, JON LIEN2*, DAWN NELSON?, and KRISTINA CURREN? ' Academica Sinica, Institute of Oceanology, 7 Nan-Hai Road, Qingdao, Shandong, China 2 Ocean Science Centre and Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland A1C 5S7 3 Whale Research Group, Memorial University of Newfoundland, St. John’s, Newfoundland A1C 5S7 *Please direct correspondence to Lien Hai, Dong Jin, Jon Lien, Dawn Nelson, and Kristina Curren. 1996. A contribution to the biology of the White-beaked Dolphin, Lagenorhynchus albirostris, in waters off Newfoundland. Canadian Field-Naturalist 110 (2): 278-287. Twenty-one ice entrapments, nine strandings and three incidental captures in fishing gear of White-beaked Dolphins (Lagenorhynchus albirostris) 1979-1990 in Newfoundland waters are summarized. The results of autopsies from two ice- entrapped groups include body measurements, age estimates and stomach contents. Body-length criteria for physical maturity were determined from the degree of fusion of the bones in the flippers. On average, male White-beaked Dolphins become physically mature at a length of 281 cm and an age of 13 years; females reach physical maturity at an average body length of 261 cm and an age of 16 years, but there was considerable variation in size at sexual maturity. Key Words: White-beaked Dolphin, Lagenorhynchus albirostris, ice entrapment, stranding, by-catch. Little is known of the biology of the White-beaked Dolphin, Lagenorhynchus albirostris (Gray 1846). It is one of two species of Lagenorhynchus found in the North Atlantic; the second is the Atlantic White- sided Dolphin (Lagenorhynchus acutus). Ranges of the two dolphins are similar, although White-beaked Dolphins are more northerly animals (Evans 1980; Leatherwood et al. 1976). In northwest Atlantic waters, White-beaked Dolphins are known to sum- mer in the Davis Strait and off Labrador; they can be found as far south as Cape Cod, Massachusetts, in the winter and spring (Leatherwood et al. 1976; Alling and Whitehead 1987). A summer population off southern Labrador of 3500 individuals and an annual mortality of 10% from hunting and incidental bycatch has been estimated (Alling and Whitehead 1987). This paper presents information on the ice entrap- ments and strandings of White-beaked Dolphins in Newfoundland which occurred between 1979 and 1990 and the results of autopsies on several ice- entrapped groups. Method Reporting Network Reports on strandings, captures in fishing gear and ice entrapments of cetaceans in Newfoundland were received via a widely advertised, toll-free, province- wide telephone line. Several agencies, including the Department of Fisheries and Oceans, the New- foundland and Labrador Department of Fisheries, the Canadian Coast Guard and the Royal Canadian Mounted Police, referred reports of cetaceans to the entrapment and stranding network. A full description of the network is available in Lien (1994). When reports of ice entrapments were received, reliable local authorities were contacted to investi- gate the report and supply additional information. In most cases, crews were dispatched to the sites of the entrapments to verify the reports and to assist in res- cue efforts or autopsies. Entrapments were routinely photographed. Autopsied Specimens White-beaked dolphin specimens for autopsy were obtained from two ice entrapments (Table 1). The first entrapment occurred in Grand Bay, Port-aux- Basques (47° 34’N, 59° 04’ W) on 9 March 1982, and the second at Point Verde, Placentia Bay (47° 14’N, 54° O1’W) on 25 March 1983 (Figure 1). Additional animals from the latter entrapment were found on 26 March less than 1 km from the original entrapment location; these animals were included with the Point Verde sample. The entrapments were initially described in the annual reports of the entrapment and stranding network (Lien et al. 1982; Lien et al. 1983). Port-aux-Basques Ice Entrapment Little is known about the ice entrapment which occurred in Grand Bay, Port-aux-Basques. Pack ice moved toward the shore with southwesterly winds and covered the bay on 9 March 1982. Several beached animals were observed by a hunter and reported that evening. Initial examinations were done on 10 March at which time all of the animals were dead. Examinations were completed on 11 March. Because of the extremely heavy pack ice on and near the shore where the animals died, it is not cer- tain that all trapped animals were found. Thirty-nine dolphins were found beached at low tide on 10 March; one additional animal had been taken for 278 1996 ar FiGuRE 1. Locations of ice entrapments and strandings of White-beaked Dolphins (Lagenorhynchus albirostris) in Newfoundland (1979-1990). (* indi- cates ice entrapment; # indicates stranding) food the previous evening. The severe ice conditions made complete examination of all animals impossi- ble. Twenty-two of the dolphins were examined in HAI, LIEN, NELSON, AND CURREN: WHITE-BEAKED DOLPHIN Pays) detail while incomplete data was gathered as possi- ble on the remaining 17 animals. Sex was deter- mined for 34 animals. Point Verde Ice Entrapment Pack ice moved inshore at Point Verde near mid- night on 24 March 1983; the ice entrapment of the dolphins probably occurred at that time. Investigators arrived at the site on 25 March. A total of 59 dolphins were initially visible in the pack ice; 39 of the animals were already dead. It is likely that additional animals had been killed in the entrapment as there was a great deal of blood on ice where dol- phins were not visible. At the time of the crew’s arrival, live animals were being crushed and squeezed by heaving ice. Ice maps of Placentia Bay showed no water nearby like- ly to remain open; therefore, 20 of the live dolphins were killed by shooting. In addition to the animals at Point Verde, nine dolphins in ice were located offshore by a helicopter. Seven of these animals were dead. An estimated 80- 100 dolphins and a single Minke Whale (Balaenoptera acutorostrata) were found swimming in a small lead of open water in Placentia Arm, about | km from Point Verde. The total number of White-beaked Dolphins trapped in ice in the area of Point Verde was in excess of 150 individuals; the number of animals [J+ FIGURE 2. Measurements (in mm) of ice entrapped animals. 1) body length, tip of upper jaw to deepest part of fluke notch; 2) tip of lower jaw to centre of anus; 3) tip of lower jaw to centre of genital slit; 4) tip of lower jaw to centre of umbilicus; 5) tip of upper jaw to top of dorsal fin; 6) tip of upper jaw to leading edge of dorsal fin; 7) tip of upper jaw to anterior insertion of right flipper; 8) tip of upper jaw to anterior edge of blowhole; 9) tip of upper jaw to centre of right eye; 10) tip of upper jaw to angle of gape; 11) tip of upper jaw to apex of melon; 12) centre of eye to centre of eye, over top of head (not shown); 13) right flipper width at insertion; 14) right flipper length, tip to anterior inser- tion; 15) dorsal fin height; 16) dorsal fin base; 17) fluke span, tip to tip; 18) girth at anus; 19) girth at anterior dorsal; 20) girth at eye; 21) girth at axilla. 280 THE CANADIAN FIELD-NATURALIST Vol. 110 Radius Phalanges Humerus Uina Metacarpals Carpus FiGuRE 3. Pectoral fin measurements from x-rays. that escaped or were killed beneath the ice is which included the dolphins entrapped in ice at unknown. The animals at Point Verde and Placentia Black Point and Little Barasway, Placentia Bay on Arm were likely part of a larger group of dolphins _ the same date (Table 1). TABLE 1. Ice entrapments of White-beaked Dolphins (Lagenorhynchus albirostris) reported in Newfoundland (1979- 1990). Date Location Details of entrapment 1982 10 March Port aux Basques 40 dead; 35 examined 1983 19 February Chapel Arm, Trinity Bay 3 alive, disappeared as ice receded. 22 February Bareneed, Conception Bay 5-8 alive; dispeared as ice receded. 24 February Western Bay, Conception Bay 20-30 alive; disappeared as ice receded; some believed 25 February 26 February Adam’s Cove 1 dead; examined. 3 March Sunnyside, Trinity Bay 3-4 alive; disappeared. 6 March Old Shop, Trinity Bay 8 alive; disappeared. 8 March Whiteaway, Trinity Bay 8-10 alive; died in ice, not examined. 24 March Fairhaven, Placentia Bay 15-20 alive; 6 collected alive; rest disappeared as ice receded; not examined. 24 March Mt. Carmel, St. Mary’s Bay 4 alive; disappeared as ice receded. 25 March Point Verde, Placentia Bay 68 alive; 20 killed; remainder died in ice; 26 examined. 1984 25 March Placentia Arm, Placentia Bay 100 alive; pressure on ice; 26 found dead on beach; 14 examined. 25 March Black Point, Placentia Bay 20-30 alive, disappeared. 27 March Little Barasway, Placentia Bay 1 dead 1987 8 April Lawn, Placentia Bay 8-12 alive in ice for 24 hrs.; observed; disappeared - believed dead; not examined; 4 found 4 July. 22 April Little Bay East, Fortune Bay 2-3 in ice alive, disappeared as ice receded. 11 April Hant Harbour, Trinity Bay 6 dead; not examined. 1989 20 February Torbay 3 alive, disappeared in ice. 1990 6 February Jamestown 6-8 dead; not examined 3 March Rencontre East 6-8 dead; not examined Ochre Pit Cove, Conception Bay to have died not examined. 20-30 alive; disappeared as ice receded, some dead, not examined. 1996 Hal, LIEN, NELSON, AND CURREN: WHITE-BEAKED DOLPHIN 281 TABLE 2. Strandings of White-beaked Dolphins (Lagenorhynchus albirostris) reported in Newfoundland (1979-1990). Date Location 20 May 1983 Point Lance 13 September 1985 Point Leamington, Notre Dame Bay 2 July 1986 Point La Haye 25 August 1986 Stephenville 14 September 1987 Lewisport 31 March 1988 St. Shotts 1 March 1989 20 July 1989 27 July 1989 Point Lance Sandy Point, Port au Port Black Bank Beach Autopsies on 26 dolphins were performed at Point Verde on 25 March. Investigators returned on 26 March but ice had scoured the beach clean overnight and no animals remained. Seventeen of the 24 dol- phins trapped in Placentia Arm were also examined. All autopsies were completed by the afternoon of 26 March. Field Examinations The harsh field conditions under which the dol- phins were examined limited the collection of data. Complete morphometric data were taken for each animal when possible (Tables 4 and 5, Figure 2). Stomach contents, the lower jaw, a flipper and the sex organs were also collected for laboratory exami- nation. Laboratory Examinations The ages of dolphins were estimated from longitu- dinal and cross-sections of the teeth. Two large teeth were removed from each jaw. One tooth was cut into longitudinal sections and the other cross-sectioned at the gum line. The sections were polished to a 25 micron thickness and decalcified with a 10% formic acid soak. The sections were then stained for 40 minutes in Mayer’s haematoxylin, rinsed, dehydrat- ed in alcohol, mounted on slides and examined with a 40 power microscope with ground illumination. The flippers from animals for which teeth were also available were frozen and x-rayed. The x-rays were examined to determine the degree of fusion of the bones (Figure 3). The bones examined for fusion included the humerus, radius, ulna, and the bones of the five phalanges. Fusion of the humerus, radius and ulna were rated on a five step scale with step Details of entrapment 7 dead; not examined. 2 dead; not examined. Dead female + foetus; died in birth; examined. 3 dead; believed same animals caught by fisherman; not examined. 3 animals stranded alive; pulled from beaches; disappeared; not examined. Male stranded alive; killed; examined. Dead; not examined. 4 alive; pushed out; disappeared. Dead; believe one from 20 July; not examined. one representing no fusion and step five representing complete fusion. The number of steps in the fusion scales for the phalanges varied with the number of bones examined in each phalange; for each case, the first step represented little or no fusion and the last step complete fusion. Non-linear regressions for body length and age, and body length and flipper length were performed separately for males and females. The average body lengths corresponding to physical maturity for each sex were entered into the regression equations and average ages of physical maturity calculated. The stomach contents were weighed and sorted. Identification of food items and parasites were identi- fied by comparison with the otolith and parasite col- lections at the Memorial University of Newfoundland. Results and Discussion Ice Entrapments Ice entrapments of White-beaked Dolphins occurred regularly in years when pack ice was heavy and was widely distributed over most of the coast- line of Newfoundland (Table 1, Figure 1). Twenty- one ice entrapments were reported from 1979 to 1991 involving a total of 320 to 350 animals. Known mortality as a result of ice entrapment was about 55%. However, actual mortality may have been higher as 17 to 21% of the dolphins disappeared in the pack ice, and it is likely that some of them died. Only 24 to 28% of the dolphins reported ice entrapped were known to have survived. Ice entrapped groups of dolphins ranged in size from two to three individuals to several hundred ani- TABLE 3. Incidental entrapments of White-beaked Dolphins (Lagenorhynchus albirostris) in inshore fishing gear reported in Newfoundland (1979-1990). Date Location Details of entrapment 21 August 1987 30 March 1988 25 June 1988 Frenchman’s Cove, Fortune Bay Holyrood, Conception Bay Bauline, Conception Bay Dead; caught in groundfish gillnet. Dead; caught in herring gillnet. 2 dead; caught in codtrap. Vol. 110 LOC 3 ¢ W 9tdd ove — © d cedd LOOT 8 7¢ 4 pedd O0cC SE W eedd 6SIC d ce dd CvOC. --C d [edd W O0€dd 81ec W 6¢cdd EEC” 968. Vel LEL 097 309e, 91C 896. SOI 61v:= Eh. CC. ,P9C- -6Lc- -90h “S68 68C1 “S68 VL. “CLE. C681 Cc W 8cdd Ocel LS8 8Sel 8LL COS Mal8S" s7VGee L8c 2 COL Seyi LO SCE 98 = SOE" Coy Vl6s. CLE 4 coo. = CSo IS ool (6 d Lcdd c8cl «$68 = 067TI COL OLV = 205 -VVG. Ol he SLE = Ve eS 3666. 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S d ldd IZ OC 61 81 LI SIE SI vl el cl at Ol 6 8 L 9 ¢ v € (G I asy XE #CI 282 ‘ZT dINBLJ Ul payeuinsa oie sapoo syuoWIaInsevayy ‘sonbseg-xne-110g wo suTydjog payeeq-ay A, peddenus-sor Jo sade pur syuouloinsvay] ‘pf ATAV 283 Hal, LIEN, NELSON, AND CURREN: WHITE-BEAKED DOLPHIN 1996 IC 0c 61 81 Ov9 O19 099 Or OS9 OSt OLE OOr LI O87 OIV Or OLT O87 O87 OI OIC OSt Ore OLI 061 OIC cl OSL O&e OLY O8L Oe Ore O&e vl Orl 00C 061 Orl OST OCI el Cl Il 09C OVC OIC OSC OVC 00c O6I OIC Ol 00€ OLT OCE Ove OST OSc 6 Ore O€e O87 09¢ Ore OLT OLT O8¢ 8 Ovr OIV OCS O09V OOr 06€ 06E [E 0s9 O€8 00cI 0901 O9L O9L 098 9 ¢ O10 OS? 0c8 OS6 OcOl Or8 OLL O18 v € Oc9T OcLI OScl 06LI OS9T OOTT OrIl Oecl Cc O6L OSC Or6 09ST OLLT O87 O9VT 068 O87 OLI? Oc8I 000C O87 060¢ O8SC OrLI Ol6l O9LI OLTT OST? 0997 O18? OSIT OOLT OE9C O6LI O18I Or0T 0C6 OE9T OO 06SC Otre O87 OST OSsl OSOE OSC OSLI 0891 O6LI I cl el co Dt asV — SESuu Ses Sass es Fy fy = MS Seka k hhh Same Be Sea Sa xoS wa #1 ‘ZT ANBLJ Ul poyeulnso ore sopoo juowoinsvopy ‘Avg eNUsOR][g Wo sulydjog poyeeq-a1ty AA, poddenus-ool Jo sade pojeujso pur s}usWoINsvIyA] “CS ATAVL 284 Body length (cm) 160+ T T T T an = ce) 2 4 6 8 10 12 14 Age (years) FIGURE 4. Relationship between age and body length in male White-beaked Dolphins (Lagenorhynchus albirostris). mals. Occasionally, single ice-entrapped dolphins were also reported; these were probably the rem- nants of unobserved group entrapments. Similarly, early spring discoveries of groups of relatively fresh dead dolphins (e.g., the May 1983 stranding at Point Lance) were probably the result of unobserved ice entrapments (Lien et al. 1983). Strandings Strandings of White-beaked Dolphins which were not known to have been entrapped or driven ashore by ice are shown in Table 2 and Figure 1. A total of nine strandings were reported; five of these involved groups of animals. There were three strandings of live animals. None of these dolphins were measured or autopsied. Entrapments in Fishing Gear Very few White-beaked Dolphins were reported as incidentally caught in fishing gear (Table 3) but these data probably under-estimate actual catches. The inci- Pectoral fin length (cm) 160 170 180 190 200 210 220 230 240 250 Body length (cm) FIGURE 6. Relationship between body length and flipper length in White-beaked Dolphins (Lagenorhynchus albirostris). THE CANADIAN FIELD-NATURALIST Vol. 110 Body length (cm) Te T T T 4 6 8 10 12 14 Age (years) FIGURE 5. Relationship between age and body length in female White-beaked Dolphins (Lagenorhynchus albirostris). dental capture of dolphins by inshore fishermen in Newfoundland and Labrador is common (Lien et al. 1994); however, since dolphin entrapments generally cause little gear damage and the dolphins are fre- quently eaten, fishermen have little motivation to report these catches (Lien et al. 1994; Lien 1994). Observations of Ice-entrapped Dolphins Many of the dolphins from both the Port-aux- Basques and Point Verde ice entrapments had severe scrapes and cuts from the ice. The beaks and heads of many dolphins had been abraded when the head was pushed between ice pans into the air to breathe. Bloody froth was typically found in the lungs. Six ani- mals had liver ruptures, serious abdominal bleeding or broken ribs. One had a deep gash on its side and a rup- tured stomach. These injuries were caused by crushing between ice pans. Except for the animals which were euthanized by gunshot, the immediate cause of death for most of the dolphins was not apparent. Pectoral fin length (cm) 30 PALE aT { RNY | Se MOnoMiaei4 LielentS Age (years) FIGURE 7. Relationship between age and flipper length in White-beaked Dolphins (Lagenorhynchus albirostris). 1996 (a) Hat, LIEN, NELSON, AND CURREN: WHITE-BEAKED DOLPHIN 285 (b) Degree of ossification of radius 220 Body length (cr) (d) Degree of osstication of phalange | 240 Body length (cm) FIGURE 8. Relationship between the degree of fusion in the flippers and body length in male White-beaked Dolphins (Lagenorhynchus albirostris). The bones in the flippers are labelled as follows: (a) humerus; (b) radius; (c) ulna; (d) phalange I. Morphometric Measurements and Estimated Ages Measurements taken from the ice-entrapped dol- phins are presented in Tables 4 and 5. Longitudinal tooth sections provided the most reliable means of estimating age (Tables 4 and 5). A narrow layer of uniform dentine in the section ended in a distinct clear line; this was identified as the neonatal line (GLG). The first postnatal GLG was wider than sub- sequent layers which were regularly spaced but declined in width toward the pulp cavity. Between 15-17 GLGs were found in the teeth before the pulp cavity was filled; these layers were more difficult to “read”. GLGs beyond the first postnatal layer were assumed to be deposited annually. The ice-entrapped dolphins at Port-aux-Basques were primarily young animals with lengths ranging from 189-241 cm (n= 31, mean length = 213 cm, s.d.= 11 cm) and estimated ages from two to seven years (n = 34, mean age = 2.9 years, s.d.=1.1 years). Most of the dolphins were two to three years of age. The group of dolphins entrapped at Point Verde ranged in length from 168 to 305 cm (n = 37, mean length = 229 cm, s.d. = 39 cm). Thirteen of the Point Verde animals were one year old; the remain- ing fifteen were adults more than six years old (n = 15, mean age = 11.9 years, s.d. = 2.6 years). The dolphins grew quickly prior to three years of age (Figures 4 and 5). Males showed a different growth pattern from females in that they grew at a faster rate and tended to be larger than females after reaching about three years of age. Morphometric data for the two sites were combined and non-linear regressions between age and length were performed for both sexes. The results were as follows: Females: Body length (cm) = 191 * [Age (years) ]®!!2 (r7=0.81, df=36, p<0.01) Males: Body length (cm) = 177 * [Age (years) ]9-!79 (r2=0.96, df=18, p<0.01) Figures 6 and 7 show the relationships between flipper length, body length and estimated age. Using morphometric data from both ice-entrapment sites, non-linear regressions were performed between flip- per length and body length, and flipper length and age. The results are listed below: Flipper length (cm) = 35.5 * [Age (years) ]®!7¢ (r2=0.67, df=26, p<0.01) Flipper length (cm) = 0.128 * [Body length (cm)]!.°8 (r2=0.88, df=26, p<0.01) 286 Degree of cssilication at humerus Degree of ossification of uina THE CANADIAN FIELD-NATURALIST Vol. 110 Degree of ossification of racius Degree of ossfication of phalange | ication of phalange II! Degree of ossii FIGURE 9. Relationship between the degree of fusion in the flippers and body length in female White-beaked Dolphins (Lagenorhynchus albirostris). The bones in the flippers are labelled as follows: (a) humerus; (b) radius; (c) ulna; (d) phalange I; (e) phalange II; (f) phalange III. Four of the 24 female dolphins from Point Verde were pregnant. The youngest of these animals was one year old and 174 cm long; as this is improbable it is likely an error in field recording. The remaining three were at least seven years old and 245 cm long. None of the female animals at the Port-aux-Basques stranding were pregnant. There was insufficient data to determine the average body length and age corre- sponding to sexual maturity. The degrees of fusion of all of the bones in the flippers increased with age for both sexes (Figures 8 and 9). The humerus was the first bone in the flip- pers to show complete fusion, followed by the radius and ulna, and finally by the five phalanges. Since the phalanges were the last bones in the flippers to fuse completely, physical maturity was defined as the average body length corresponding to the fusion of these bones. By this criterion, four male and four female animals were considered physically mature. On average, male White- beaked Dolphins become physically mature at a length of 281 cm (s.d. = 2.9 cm), which corre- sponds to an age of 13 years; females dolphins achieve physical maturity at an average body 1996 length of 261 cm (s.d. = 6.2 cm), corresponding to an age of 16 years. These results also indicated sexual dimorphism in White-beaked Dolphins. A t-test showed that mature males had significantly greater body lengths than female animals (t = 5.86, df = 6, p<0.01). Stomach Contents Stomachs from all animals in the Port-aux-Basques entrapment were infested with worms (Anisakis spp.). Contents of the stomachs of 20 Port-aux-Basques. ani- mals were examined; the stomachs of the remaining animals were empty. The mass of food in the stom- achs varied from 10.5-446.0 g with a mean mass of 122 g. Most of the animals (90%) had otoliths and bones from cod (Gadus morhua) in their stomachs although there was no flesh or other indications of recent feeding. In general, the largest otoliths came from the stomachs of the largest animals. A few of the stomachs (10%) contained remnants of crab (Chionoecetes opilio) and seaweed. Four stomachs (20%) contained fish hooks from trawls which are commonly used in the area to catch cod. Stomachs from the Point Verde animals were not collected. Literature Cited Alling, A. K., H. P. Whitehead. 1987. A preliminary study of the status of white-beaked dolphins, Lagenorhynchus albirostris, and other small cetaceans off the coast of Labrador. Canadian Field-Naturalist 101(2): 131-135. Hat, LIEN, NELSON, AND CURREN: WHITE-BEAKED DOLPHIN 287 Evans, P.G. H. 1980. Cetaceans in British waters. Mammal Review 10(1): 1-52. Leatherwood, S., D. K. Caldwell, H. E. Winn. 1976. Whales, dolphins, and porpoises of the Western North Atlantic, a Guide to Their Identification. NOAA Technical Report NMFS CIRC-396. Lien, J., J. Dong, L. Baraff, J. Harvey, and K. Chu. 1982. Whale entrapments in inshore fishing gear during 1982. Preliminary Report to Fisheries and Oceans Canada, September 1982. 36 pages. Lien, J., S. Staniforth, L. Fawcett, R. Vaughan, and J. Dong. 1983. Whale and shark entrapments in inshore fishing gear during 1983. A Preliminary Report to Fisheries and Oceans Canada, 23 September 1983. 36 pages. Lien, J. 1994. Entrapments of large cetaceans in passive inshore fishing gear in Newfoundland and Labrador (1979-1990). Pages 149-158 in I.W.C. Special Issue Number 15, Gillnets and Cetaceans. Edited by W. Perrin, G. Donovan, J. Barlow. International Whaling Commission, Cambridge. Lien, J., G. B. Stenson, S. Carver, and J. Chardine. 1994. How many did you catch? The effect of method- ology on by-catch reports obtained from fishermen. Pages 535-540 in I.W.C. Special Issue Number 15: Gillnets and Cetaceans. Edited by W. Perrin, G. Donovan, J. Barlow. International Whaling Commis- sion, Cambridge. Received 31 March 1993 Accepted 22 January 1996 Identification of Greater Scaup, Aythya marila, and Lesser Scaup, A. affinis, Ducklings COLLEEN H. NELSON 318 Wildwood Park, Winnipeg, Manitoba, R3T OES, and Manitoba Museum of Man and Nature, Winnipeg, Manitoba R3B ON2 Nelson, Colleen H. 1996. Identification of Greater Scaup, Aythya marila, and Lesser Scaup, A. affinis, ducklings. Canadian Field-Naturalist 110(2): 288-293. Both relative size and the nail-width described by earlier authors will distinguish the Greater Scaup (Aythya marila) from the Lesser Scaup (A. affinis) at all ages. These criteria are most useful if the age of the scaup is known. In this study, an attempt was made to obtain a nearly constant ratio that would identify the two species with some degree of certainty regardless of age, sex, or condition. Checked on live specimens from Canada and on museum specimens from North America, this ratio was found to be 90% effective. Key Words: Greater Scaup, Aythya marila, Lesser Scaup, Aythya affinis, ducklings, identification. Both Palmer (1976) and Todd (1963) described ways of distinguishing between Greater Scaup and Lesser Scaup. Although both ways are correct, they work best if the exact age of the specimen is known. The exact age of specimens in the field, however, is seldom known. Therefore, while the work of both Palmer (1976) and Todd (1963) has been confirmed in this study, the search continued for a ratio that would separate ducklings of Greater Scaup from ducklings of Lesser Scaup at all times. Materials and Methods Employing a steel caliper, I used the chordal mea- surements of Baldwin et al. (1931) and the sexing method of Hochbaum (1942). In 1972, I made a measured pencil drawing, 2x life size, of the bill of each species each day, from inidividuals chosen at random; a selection of these drawings comprises Figure 1. All birds were weighed each day until the capaci- ty of the triple-beam balance was reached. Thereafter, the birds were weighed on appropriate Pesola balances. A database was made for the eight Manitoba Greater Scaup and the 22 Manitoba Lesser Scaup. All measurements of weight or mass are in grams and all linear measurements are in centimetres. Unless otherwise indicated, all Greater Scaup dis- cussed herein belong to the subspecies, nearctica Stejneger, following Banks (1986). Results The size differences described by Palmer (1976) are indicated in Table | by increased weight or mass (WT) and length of exposed culmen (EC). Todd’s (1963) nail-width character is expressed in Table 1 as a percentage of bill-width: WN/WB. Although the percentages do not decrease as dramatically as in goldeneyes described by Nelson (1993a), they do eventually decrease, going from 0.46 (age A) down to 0.30 (age E) in the Manitoba Greater Scaup and from 0.43 (age A) down to 0.27 (age E) in the Manitoba Lesser Scaup. Subsequently, various ratios were tested. A three- part ratio, (WN x LN)/HBB, proved useful in distin- guishing Greater Scaup from Lesser Scaup. The ratio was used to separate scaup of both species from all areas (Nelson 1993b). The ratio is usually 0.3 or above for live Manitoba Greater Scaup and usually below 0.3 for live Manitoba Lesser Scaup (Table 2). This ratio was checked on live Manitoba and Saskatchewan Lesser Scaup in 1974 and 1977, and on live Northwest Territories Greater Scaup in 1983 (Table 3). These data differed significantly between Lesser Scaup and Greater Scaup, with a confidence level of 90% or more (t-test). Table 4 includes live, known-age scaup of both species and of separate sexes. The identification ratio was fur- ther checked on museum specimens of estimated age (Table 5). Discussion There is a size difference in some Aythya species. Usually, males are larger than females (Hochbaum 1944; Weller 1957), the opposite is true in the scaup. This difference becomes apparent when ducklings are about 10 days old. However, when the sexes are lumped, all Greater Scaup are larger than all Lesser Scaup (Table 4; Palmer 1976), including age A birds. Scaup of both species show considerable individu- al size variation whether or not the breeding ranges of the two species overlap. It is important to note that scaup of both species were larger in 1974, 1977, and 1983, than either species was in 1972. This vari- ation sometimes means that birds of both species may be larger in one location than in another. The 288 1996 NELSON: IDENTIFICATION OF SCAUP DUCKLINGS 289 GREATER SCAUP _ Aythya marila nearctica LESSER SCAUP Aythya affinis 3 37da. Ficure 1. Comparison of bills of ducklings of Lesser Scaup and Greater Scaup from 1 to 40* days. 290 TABLE 1. Comparative measurements of live, known-age captives from Manitoba (1972). Live birds are lumped as follows: age A = days 1-5, age B = days 6-10, age C = days 11-15, age D = days 16-20, age E = day 21 and older. EC = exposed culmen; WN = width of nail; WB = width of bill; and WT = weight or mass. Species Origin Parameter THE CANADIAN FIELD-NATURALIST Age Sample Range A. m. nearctica Churchill, Manitoba EC 40 39 40 37 TL. > 40 40 40 40 79 = oo 40 40 40 39 79 40 40 40 40 80 A. affinis Shoal Lake, Manitoba E 82 719 76 74 154 86 80 76 68 163 = MONOWPHMOAMSZMonwyeZmoanwero 86 80 76 71 155 85 80 76 74 165 1.50- 1.96 17510139 Di 2.86 2.622) 3123 2.95- 4.36 0.47- 0.60 01534") 0158 0153-4) 10:58 0.54- 0.61 OTs) 0.72 0.42- 0.54 0.34- 0.47 0.32- 0.40 0.30- 0.35 0.24-. 0.33 37.30- 64.20 56.90 - 112.90 86.40 - 181.80 124.30 - 286.30 174.20 - 707.20 125-170 S327 2:20 1.86- 2.70 Deo 1D.05 2.60- 4.11 0.34- 0.50 0.41- 0.52 0.41- 0.54 0.42- 0.56 0.44- 0.58 0357053 0.32- 0.43 0.26- 0.37 0.25- 0.34 0.23- 0.31 26.00 - 53.40 37.80 - 122.90 57.20 - 190.90 106.60 - 273.80 165.20 - 553.50 Mean ei: 2.06 2.50 297 3.50 0.54 0.55 0.55 0.57 0.62 0.46 0.41 0.36 0.32 0.30 47.49 7915 121.2] 193.97 337.16 1.46 1.83 2.28 Pept 3:27 0.42 0.46 0.47 0.48 0.51 0.43 0.38 0.33 0.29 0.27 Sly) 69.54 121.75 184.46 327.89 S.D. 0.110 0.140 0.170 0.150 0.350 0.036 0.012 0.013 0.018 0.033 0.024 0.027 0.021 0.012 0.012 6.030 13.240 23.810 38.650 129.550 0.103 0.156 0.188 0.387 0.342 0.034 0.030 0.030 0.030 0.027 0.032 0.028 0.024 0.020 0.018 6.033 19.636 26.396 33.810 98.952 Vol. 110 | SE: 0.017 0.026 0.027 0.024 0.040 0.006 0.002 0.002 0.003 0.004 0.004 0.004 0.003 0.002 0.002 0.953 2.093 3.764 6.111 14.484 0.011 0.017 0.022 0.050 0.027 0.004 0.003 0.003 0.003 0.002 0.003 0.003 0.002 0.002 0.001 0.654 2195 3.028 3.930 7.703 1996 NELSON: IDENTIFICATION OF SCAUP DUCKLINGS 20 TABLE 2. Identification ratio, (WNxLN)/HBB, for live, known-age captives from Manitoba (1972). LN = length of nail and _ HBB = height of bill at base. Ages as in Table 1. difference between the two species can often be rather small. Acknowledgments I thank B. Batt, R. Bromley, R. Cole, R. Evans, D. Hatch, L. King, M. McNicholl, L. Simmons, J. Stoudt, and P. Ward for collecting birds and eggs for me. I thank Delta Waterfowl and Wetlands Research Station for rearing and laboratory space and for housing and I thank R. Jones for loaning a triple- beam balance. I also thank I. Grymonpré for caring for my youngest child, and D. Donald for inking Figure 1. I thank various family members, especially Species Origin Parameter Age Sample Range Mean S.D. S.E. _ A. m. nearctica Churchill, Manitoba (WNxLN) /HBB A 40 0.30 - 0.38 0.34 0.020 0.003 B 40 0.29 - 0.35 0.32 0.015 0.002 C 40 0.28 - 0.34 0.31 0.014 0.003 D 39 0.27 - 0.34 0.30 0.016 0.003 E 76 0.28 - 0.35 0.31 0.016 0.002 A. affinis Shoal Lake, Manitoba (WNxLN) /HBB A 66 0.18 - 0.34 0.26 0.032 0.004 B 72 0.21 - 0.31 0.26 0.018 0.002 E 76 0.21 - 0.30 0.25 0.019 0.002 D 74 0.21 - 0.28 0.24 0.017 0.002 E 160 0.21 - 0.28 0.25 0.014 0.001 C. Nelson and J. Nelson, for providing transportation facilities and I thank B. Kessel, B. McPherson, B. O’ Malley, and D. Trauger for helpful consultations. I thank F. Cook, A. Erskine, H. Ouellet, and an anony- mous reviewer for suggestions to improve this manuscript and I thank M. Abrahams, L. Armstrong, B. Harnish, and G. Hochbaum for statistical calcula- tions. I thank O. Adegunju, L. de Castro, C. Clifford, L. Phillips, and R. Wolfe for transcription and com- puter work and I thank the Manitoba Museum of Man and Nature for office space and for borrowing specimens on my behalf and housing them. I thank the following institutions for extending these loans TABLE 3. Identification ratio, (WNxLN)/HBB, for live, known-age captives from Manitoba, Saskatchewan, and the Northwest Territories (1974, 1977, 1983). Ages as in Table 1. Species Origin Parameter Age Sample Range Mean SD! S.E. A. m. nearctica Great Slave Lake, Northwest Territories (WNxLN) /HBB A 34 0.32 - 0.46 0.40 0.030 0.005 B 22 0.33 - 0.42 0.37 0.027 0.006 C 19 0.34 - 0.44 0.38 0.027 0.006 D 12 0.33 - 0.43 0.38 0.032 0.009 E 25 0.34 - 0.47 0.40 0.038 0.007 A. affinis Minnedosa and E. Meadows, Manitoba; Moose Jaw, Saskatchewan (WNxLN) /HBB A 73 0.20 - 0.37 0.30 0.039 0.005 B 87 0.20 - 0.32 0.27 0.027 0.003 e 52 0.22 - 0.34 0.27 0.028 0.004 D 55 0.22 - 0.32 0.27 0.018 0.002 E 36 0.24 - 0.32 0.28 0.019 0.003 292 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 4. Identification ratio, (WNxLN)/HBB, for live, known-age captives, separate sexes, from all locations, all years. Ages as in Table 1. Species Origin Parameter Age Sample Range Mean S.D. S.E. A. m. nearctica, males Churchill, Manitoba (WNxLN) /HBB A 42 0.30 - 0.45 0.36 0.035 0.005 B 36 0.29 - 0.41 0.33 0.031 0.005 (e 35 0.28 - 0.41 0.32 0.034 0.006 D 32 0.27 - 0.42 0.32 0.031 0.005 E 65 0.28 - 0.45 0.33 0.042 0.005 females A 32 0.30 - 0.46 0.37 0.040 0.007 B 26 0.30 - 0.42 0.34 0.032 0.006 Cc 24 0.28 - 0.44 0.33 0.045 0.009 D 19 0.28 - 0.43 0.33 0.047 0.011 E 39 0.29 - 0.47 0.34 0.048 0.008 A. affinis, males (WNxLN) /HBB A 88 0.18 - 0.37 0.29 0.045 0.005 B 84 0.22 - 0.32 0.27 0.021 0.002 Cc we 0.21 - 0.33 0.26 0.024 0.003 D 65 0.21 - 0.30 0.25 0.021 0.002 E 169 0.21 - 0.31 0.25 0.022 0.002 females A lel 0.18 - 0.38 0.28 0.035 0.005 B 75 0.20 - 0.32 0.26 0.026 0.003 Cc 60 0.21 - 0.34 0.26 0.030 0.004 D 64 0.21 - 0.32 0.26 0.024 0.003 E 104 0.21 - 0.39 0.26 0.026 0.002 or for allowing me to examine specimens: American Nature, Winnipeg; Canadian Museum of Nature, Museum of Natural History, New York; Carnegie Ottawa; Royal Ontario Museum, Toronto; U.S. Museum, Pittsburgh; Delta Study Collection, National Museum, Washington, D.C.; zoological Portage la Prairie; Manitoba Museum of Man and museums of the universities of Alaska at College; TABLE 5. Identification ratio, (WNxLN)/HBB, for museum specimens from Alberta, British Columbia, Manitoba, Northwest Territories, Ontario, Québec, Saskatchewan, Alaska, and North Dakota. Ages as in Table 1. Species Origin Parameter Age Sample Range Mean S.D. S.E: A. m. nearctica British Columbia, Manitoba, Northwest Territories, Ontario, Québec, Alaska (WNxLN) /HBB A 15 0.31 - 0.47 0.39 0.040 0.010 B 12 0.26 - 0.38 0.34 0.032 0.009 C i 0.27 - 0.36 0.32 0.030 0.011 D 6 0.26 - 0.37 0.32 0.037 0.015 E No data A. affinis Alberta, British Columbia, Manitoba, Northwest Territories, Saskatchewan, Alaska, North Dakota (WNxLN) /HBB A 34 0.21 - 0.38 0.32 0.039 0.007 B 15 0.15 - 0.33 0.25 0.043 0.011 € 12 0.17 - 0.37 0.25 0.061 0.018 D 2 0.19 - 0.29 0.24 0.071 0.050 E 7 0.20 - 0.32 0.27 0.037 0.014 1996 Michigan at Ann Arbor; and Wisconsin at Green Bay. Literature Cited Baldwin, S. P., H.C. Oberholser, and L. G. Worley. 1931. Measurements of birds. Contribution number 17 from the Baldwin Bird Research Laboratory, Scientific Publications of the Cleveland Museum of Natural History, volume 2. Banks, R. C. 1986. Subspecies of the Greater Scaup and their names. Wilson Bulletin 98: 433-444. Hochbaum, H. A. 1942. Sex and age determination of waterfowl by colacal examination. Transactions of the North American Wildlife Conference. Pages 299-307. Hochbaum, H. A. 1944. The Canvasback on a prairie marsh. Wildlife Management Institute, Harrisburg, Pennsylvania: Stackpole Company and Washington, D.C. NELSON: IDENTIFICATION OF SCAUP DUCKLINGS 293 Nelson, C. H. 1993a. The identification of Barrow’s Goldeneye, Bucephala islandica, and Common Goldeneye, B. clangula americana, ducklings. Wildfowl 44: 178-183. Nelson, C. H. 1993b. The downy waterfowl of North America. Delta Station Press, Deerfield, Illinois, and Portage la Prairie, Manitoba. Palmer, R. S. Editor. 1976. Handbook of North American birds. Volumes 2 and 3: Waterfowl. Yale University Press, New Haven, Connecticut, and London. Todd, W. E. C. 1963. Birds of the Laborador Peninsula and adjacent areas: a distributional list. University of Toronto Press, Toronto, Ontario. Weller, M. W. 1957. Growth, weights, and plumages of the Redhead, Aythya americana. Wilson Bulletin 69: 5-38. Received 9 November 1993 Accepted 28 November 1995 Seed Age-Germination Relationships in Plains Rough Fescue, Festuca altaica subspecies hallii J. T. ROMO Department of Crop Science and Plant Ecology, University of Saskatchewan, 51 Campus Drive, Saskatoon, Saskatchewan STN 5A8 Romo, J. T. 1996. Seed age-germination relationships in Plains Rough Fescue, Festuca altaica subspecies hallii. Canadian Field-Naturalist 110(2): 294-297. To test the relationship of germination with storage time seeds of seven collections of Plains Rough Fescue (Festuca altaica Trin. subsp. hallii [Vasey] Harms) ranging in age from 3 to 91 months were germinated at 10 and 20°C in 0.0 or -0.69 MPa of water stress for 400 degree-days. With the exception of one collection, total germination of all collections declined linearly as seeds aged. When seeds were 80 to 91 months of age germination was nearly nil. One collection appar- ently had an after-ripening requirement as germination increased initially as seeds aged, then declined. Degree-days to 50% of final germination increased linearly as seeds aged; across the seven collections about 1.2 to 1.9 more degree-days were required to reach 50% of final germination for each month of storage. Until optimal storage conditions are identified, Plains Rough Fescue seeds should be sown the first or second spring after harvest to take advantage of maximum seed vigor. Key Words: Plains Rough Fescue, Festuca altaica Trin. subsp. hallii (Vasey) Harms, restoration, seed vigor, water stress, fescue prairie. World Wildlife Fund and the Prairie Conservation Action Plan identified the grassland dominated by Plains Rough Fescue (Festuca altaica Trin. subsp. hallii [Vasey] Harms) as one of the most threatened ecosystems on the Canadian Prairies (Anonymous 1986). Less than 5% of the prairie dominated by Plains Rough Fescue remains, primarily as small remnants (Grilz and Romo 1995). Even though high priority has been placed on protecting Fescue Prairie, effective conservation strategies will likely also involve restoration. Restoration may range from reintroduction of processes such as fire and grazing to controlling exotic species (Grilz and Romo 1995) and revegetating sites that previously supported this ecosystem. One of the key issues of restoration by seeding is the need to develop an understanding of seed and seedbed ecology of Plains Rough Fescue. This long- lived perennial grass seldom produces seed (Johnston and MacDonald 1967; Toynbee 1987), thus annual harvest of seeds may not be possible and seeds will have to be stored for future use. Restoration specialists will likely have to rely on col- lecting large amounts of seed when they are pro- duced and storing them for use at later dates. Germination of Plains Rough Fescue is severely restricted by water stress, but seeds germinate over a broad range of temperatures (Romo et al. 1991; Grilz et al. 1994). The changes in vigor of Plains Rough Fescue seeds with aging are, however, unknown. The prairie in Canada dominated by Plains Rough Fescue (Festuca altaica Trin. subsp. hallii [Vasey] Harms) is one of the most threatened ecosystems. One of the most important considerations in restor- ing Fescue Prairie is the fact that this perennial grass infrequently produces seeds. The purpose of this research was to document aging effects under labo- ratory conditions on the germination of Plains Rough Fescue collected in central Saskatchewan. Materials and Methods Seeds of Plains Rough Fescue were collected in central Saskatchewan at four sites in 1987 and three in 1988. The 1987 collections were from the Strawberry Hills, Allan Hills, Radisson and Kernen Prairie while the 1988 collections were made at Sonningdale, the Eagle Hills and Kernen Prairie. All collections were made within 100 km of Saskatoon; Romo et al. (1991) and Grilz et al. (1994) provided site descriptions where the collections were made. Seeds were collected from several thousand plants at each site when seeds began to shatter from the inflo- rescences. After collection, the seeds were cleaned, 200 lots of 50 seeds were counted, and placed in paper envelopes in darkness in the laboratory. Temperatures in the laboratory ranged from 20-22°C -and relative humidity ranges from 10-30%. Seeds collected in 1987 were germinated 11, 21, 30, 42, 54, 68, 76 and 91 months after collection while the 1988 collections were tested at 3, 10, 19, 31, 43, 57, 65, and 80 months after harvest. A solution of polyethy- lene glycol (PEG) (molecular weight 20 000) was prepared to depress osmotic potentials to -0.70 MPa; the osmotic potential of this solution, determined with a Wescor vapor pressure osmometer, averaged - 0.69 MPa (SE + 0.01). This osmotic potential was chosen because it appears to be near a threshold in germination of Plains Rough Fescue (Romo et al. 294 Total Germination (%) 1996 1991; Grilz et al. 1994). Distilled water was used as a control (0.0 MPa). Seeds were incubated in closed petri dishes on filter paper disks that were moistened by adding 7 mL of water or PEG solution. These petri dishes were enclosed and sealed in clear polyethylene bags to prevent desiccation and incu- bated in darkness at 10 or 20°C for 400 degree days (base temperature = 0°C). These temperatures were chosen because they are near optimal for germina- tion of Plains Rough Fescue (Romo et al. 1991). Germinated seeds were counted and removed at 2- day intervals. A seed was considered germinated when the plumule and radicle were both 5 mm long. A randomized complete block design with 50 seeds in each of 4 replicates was used with tempera- ture and osmotic potentials factorially applied within collections and test dates. Within collections and test age, data of total germination and degree-days to 50% of final germination were analyzed with a fac- torial analysis of variance and the mean across both temperatures and levels of water stress was calculat- ed (Petersen 1985). The mean germination and degree-days to 50% of final germination at both tem- perature and levels of water stress was chosen as the index of vigor because it represents potential germi- & 80 MH i 60 5 x o wn 40 2 20 ; 0 0 20 40 60 80 100 Seed Age (Months) 100 § G £ 80 E g : 5 60 = ; : E x o & «0 2 3 oS . 5 20 7 8 is e [=) 40 60 80 j 100 Seed Age (Months) 0 20 Romo: SEED AGE-GERMINATION IN PLAINS ROUGH FESCUE 295 nation over a range of conditions to which seeds would be exposed to in the field. Standard errors for the means were determined and confidence limits were calculated. Regression analyses were used to describe relationships between seed age and total germination and degree-days to 50% of final germi- nation with the best fit regression equation being selected (Snedecor and Cochran 1980). All statistical analyses were conducted at P< 0.05. Results Total germination of seven of the eight collections declined linearly as seed aged (Figures | and 2). The Allan Hills collection apparently had an after-ripen- ing requirement for germination increased initially and then declined (Figure 1). Goodwin et al. (1995) also reported that germination of Idaho fescue (Festuca idahoensis Elmer) seeds increased during the first six months after collection and declined thereafter. The after-ripening requirement in the Allan Hills collection of Plains Rough Fescue may be attributed to seeds being collected before full seed maturity, or to a genetically fixed trait or both. The procedures used in the current study do not, howev- er, allow isolation of causal factors. Total Germination (%) r=} 3 in] o Degree—days to 50% of final germination i] o 5 t £ 80 E = & rf z 2& 60 om i 3 : 3 é 40 wn 3 2 : 5 0 20 40 60 80 100 Seed Age (Months) FiGurE 1. Regression lines for total germination (%) and degree-days to 50% of final germination (base temperature is 0°C) for the Allan Hills, Kernen Prairie, Strawberry Hills and Radisson collections of Plains Rough Fescue seeds collected in central Saskatchewan in 1987. Each symbol (solid squares for total germination and open squares for degree-days to 50% of final germination) is the mean of 16 replicates and vertical bars are 95% confidence limits. Total Germination (%) Degree—days to 50% of find germination 0 20 80 100 Kernen Prairle—1988 Total Germination (%) Degree—days to 50% of final germination 0 20 40 60 80 Totel Germination (%) Degree—days to 50% of find germination 0 20 40 60 Seed Age (Months) FIGURE 2. Regression lines for total germination (%) and degree-days to 50% of final germination (base tem- perature is 0°C) for the Eagle Hills, Kernen Prairie and Sonningdale collections of Plains Rough Fescue seeds collected in central Saskatchewan in 1988. Each symbol (solid squares for total germination and open squares for degree-days to 50% of final germination) is the mean of 16 replicates and verti- cal bars are 95% confidence limits. Germination was essentially nil at 91 and 80 months for the 1987 and 1988 collections, respec- tively. Sixty-seven to 94% of the variability associat- ed with total germination was accounted for by seed age (Table 1). Degree-days to 50% of final germination in all THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Regression equations describing correlation of seed age with total germination and degree-days to 50% of final germination for seven collections of Plains Rough Fescue. Year and Collection Regression Equation R? Total Germination (%) 1987 a Allan Hills Y%=37.5+0.76X°-0.014.X? 0.67 Kernen Prairie Y=80.4-0.92X 0.80 Strawberry Hills Y=72.8-0.84X 0.82 Radisson Y=82.7-0.97X 0.83 1988 Eagle Hills Y=69.1-0.85X 0.88 Kernen Prairie Y=26.1-0.35X 0.90 Sonningdale Y=56.8-0.67X 0.94 Degree-days to 50% of final germination 1987 Allan Hills Y°=210+1.78X 0.76 Kernen Prairie Y=2344+1.19X 0.61 Strawberry Hills Y=2214+1.47X 0.71 Radisson Y=203+1.68X 0.74 1988 Eagle Hills Y=2074+1.72X 0.84 Kernen Prairie Y=228+1.86X 0.87 Sonningdale Y=219+1.69X 0.84 *Y is total germination (%). >X is seed age in months. °Y is degree-days to 50% of final germination (base tem- perature is 0°C). collections increased linearly as seeds aged with 61 to 87% of the variation accounted for by seed age (Figures 1-7, Table 1). For each month that seeds were stored, about 1.2 to 1.9 degree-days were required to reach 50% of final germination. When first tested, the 1987 collections required 221-247 degree-days to reach 50% of final germination while ~ the 1988 collections required 212-224 degree-days. By contrast, on the last germination test seeds required 342-372 and 345-377 degree-days to reach 50% of final germination in the 1987 and 1988 col- lections, respectively. Discussion Optimal storage conditions for seeds varies among species (Mayer and Poljakoff-Mayber 1978) and the effects of aging varied among seeds collections of Plains Rough Fescue. Results of the present study illustrated that, with the exception of one collection, vigor of Plains Rough Fescue seed during germina- . tion declines as seeds aged from three to 91 months under laboratory conditions. This is consistent with the conclusion of Romo et al. (1991) that seeds of Plains Rough Fescue do not have an after-ripening | | requirement. Kearns and Toole (1939) also reported _ that seeds of Chewings Fescue (Festuca rubra var. commutata Gaud.) tended to loose their viability 1996 quickly. Creeping Red Fescue (Festuca rubra L.) retained seed viability poorly compared to several species of grass while Tall Fescue cv ‘Alta’ (Festuca arundinacea Schreb.) produced seedlings after 19 years of storage (Hull 1973). - Research is needed to identify the best combina- tion of conditions for retaining viability of Plains Rough Fescue seeds in long-term storage. Until optimal requirements for seed storage of this native perennial are identified, the following general rec- ommendations should be considered. Seed moisture content should be reduced to 5 to 7% and seeds should be stored at temperatures below -18°C (Bass 1973). If seeds can not be stored under low humidity and temperatures, they should be planted the first or second spring after collection to exploit seed vigor and fulfill the requirements for germination (Romo et al. 1991; Grilz et al. 1994) and seedling growth (Smoliak and Johnston 1968). Literature Cited Anonymous. 1988. Prairie conservation action plan: 1989-1994. World Wildlife Fund, Toronto, Ontario. Bass, L. N. 1973. Controlled atmosphere and seed stor- age. Seed Science and Technology 1: 463-492. Goodwin, J. R., P. S. Doescher, and L. E. Eddleman. 1995. After-ripening in Festuca idahoensis seeds: Adaptive dormancy and implications for restoration. Restoration Ecology 3: 137-142. Grilz, P. L, and J. T. Romo. 1995. Management consid- Romo: SEED AGE-GERMINATION IN PLAINS ROUGH FESCUE DOF] erations for controlling smooth brome in fescue prairie. Natural Areas Journal 15: 148-156. Grilz, P. L., J. T. Romo, and J. A. Young. 1994. Comparative germination of smooth brome and plains rough fescue. Prairie Naturalist 26: 157—170. Hull, A. C., Jr. 1973. Germination of range plant seeds after long periods of uncontrolled storage. Journal of Range Management 26: 198—200. Johnston, A., and M. D. MacDonald. 1967. Floral initia- tion and seed production in Festuca scabrella Torr. Canadian Journal of Plant Science 47: 577-583. Kearns, V., and E. H. Toole. 1939. Relation of tempera- ture and moisture content to longevity of Chewings fes- cue seed. United States Department of Agriculture, Technical Bulletin Number 670. Mayer, A. M., and A. Poljakoff-Mayber. 1978. The ger- mination of seeds. Pergamon Press. Toronto, Ontario. Petersen, R. 1985. Design and analysis of experiments. Marcel Dekker, New York, New York. Romo, J. T., P. L. Grilz, C. J. Bubar, and J. A. Young. 1991. Influences of temperature and water stress on ger- mination of plains rough fescue. Journal of Range Management 44: 75-81. Smoliak, S., and A. Johnston. 1968. Germination and early root growth of grasses at four root-zone tempera- tures. Canadian Journal of Plant Science 48: 119-127. Snedecor, G. W., and W. G. Cochran. 1980. Statistical methods. The Iowa State University Press. Ames, Iowa. Toynbee, K. 1987. Prolific flowering year for plains rough fescue at the Kernen Prairie. Blue Jay 45: 142-143. Received 6 June 1995 Accepted 3 October 1995 White-tailed Deer, Odocoileus virginianus, Summer Dispersion Areas in Ontario Jim D. BROADFOOT!:2, DENNIS R. Vorct! and TIM J. BELLHOUSE? ‘Ontario Ministry of Natural Resources, Southern Terrestrial Ecosystems Research Section, Research, Science and Technology Branch, PO Box 5000, Maple, Ontario L6A 1S9 ?Present address: Broadfoot Consulting, RR # 1 Midland, Ontario L4R 4K3 3Ontario Ministry of Natural Resources, Central Region Science and Technology, PO Box 3070, North Bay, Ontario P1B 8K7 Broadfoot, Jim D., Dennis R. Voigt, and Tim J. Bellhouse. 1996. White-tailed Deer, Odocoileus virginianus, summer dis- persion areas in Ontario. Canadian Field-Naturalist 110(2): 298-302. Between 1983 and 1989, 168 White-tailed Deer (Odocoileus virginianus) were radio-tracked as they migrated out of nine deer yards in Ontario. For four of these yards we tested the hypothesis that summer dispersion areas should be 10 X larger than their associated deer yards since winter range deer densities are about 10 X summer range densities. We found that summer dispersion areas were 13 X larger than their winter yards in terms of forested area, and that this expansion factor was not different form the hypothesized value (P > 0.830). Our results suggested that a reasonable approximation of the size of a summer dispersion area could be achieved by multiplying yard area by 10, and correcting the result to account for forest cover. The ability to predict the size of a summer dispersion area from yard area is important for evaluating the effects of changes to winter and summer range habitat, over or under harvest of deer, and changes in natural deer mortality rates. The relative ease of defining winter yard boundaries simplifies the use of this spatial relationship in management. Key Words: White-tailed Deer, Odocoileus virginianus, dispersion, radio-tracking, Ontario. White-tailed Deer migrate between summer and win- ter range in areas of severe winter weather (Severinghaus and Cheatum 1956). In spring, most deer leave winter range (deer yards) and travel to summer home ranges dispersed throughout the surrounding landscape (Nelson and Mech 1981). The resulting dis- tribution defines a summer dispersion area for a popu- lation of deer wintering within a particular deer yard. A literature review suggested that there were con- sistent differences in deer density between winter and summer range in northern regions where deer migrate. Winter range densities of 6 and 61 deer/km2 were reported, but most were 20 to 30 deer/km? (Table 1). In contrast, summer range densities of between 0.3 and 11 deer/km? were reported, with half between 0.5 and 4 deer/km?2 (Table 1). We cal- culated the midpoints of the range of reported winter and summer densities as 25 and 2.5 deer/km’, respectively. This indicated that typical winter range densities were 10 X greater than summer range den- sities. Because of this 10 X difference in density, it follows that deer should spread out in the summer over a dispersion area which is about 10 times larger than the yard they wintered in, in order for the popu- lation to achieve typical summer densities. In this paper we use data collected in Ontario to test the hypothesis that summer dispersion areas are about 10 times larger than their associated deer yard. Study Areas and Methods Deer were captured in nine yards throughout Ontario during January through March 1983 to 1986 using modified Clover traps, dart guns or rocket nets. Deer were radio-collared and were located at least weekly using aircraft or ground tri- angulation. Migration distance was measured as the straight line distance between winter capture location of each deer or each doe/fawn group, and the geometric cen- tre of all its summer locations. Differences in migra- tion distance among yards were assessed using ANOVA and Duncan’s multiple range test. Yard sizes were estimated from radio-locations and aerial and ground surveys. Bias in migration direction was tested using Rayleigh’s procedure (Batschelet 1981) for yards with 2 20 deer radio- collared. Summer dispersion area was estimated only for yards with 2 20 radio-tagged deer since the relationship between area of summer dispersion and sample size reached an asymptote at 18 deer. The total area of summer dispersion was measured from a minimum convex polygon and excluded areas of non-deer range (water, cities, towns). We calculated forest area from the percent forested cover in the total area surrounding each yard. We calculated multiplication factors for each yard as: summer dis- persion area (for total land area and forest area) + yard area. The Student’s-f distribution was used to assess the probability that the average of our multi- plication factors was different than our assumed value of 10. We calculated the power (1 - 8) of this test, its ability to detect a real difference, following a procedure outlined by Snedecor and Cochran (1980). 298 1996 BROADFOOT, VOIGT, AND BELLHOUSE: DEER DISPERSION AREAS IN ONTARIO 299 TABLE 1. Summer and winter population densities of migratory populations of White-tailed Deer in North America. Reference Range Density(deer/km?) Cumming (1961) Winter 8.5-46.3 Henderson, et al. Winter 21.9-24.5 (1969) Munroe (1972) Winter 21.4 yard core Mansell (1974) Winter 35 Summer 4 King (1976) Summer 1.44 - 2.05 Schmitz (1990) Winter 18-20 yard periphery 24-26 yard core Fryxell, et al. Summer 1.3-7.0 (1991) Nelson and Mech Winter 39-45 Jonvick yard (1981) 16-23 Kawishiwi R. and Garden L. yards Nelson and Mech Winter 5.8-16.9 (1986) Summer 0.3-0.7 Fuller (1990) Summer 4-10 Cumberland and Summer 1.1-2.9 Boer (1991) Banasiak (1977) , Winter 27.3-61.0 Summer 7.7-10.6 aOntario Department of Lands & Forests, Parry Sound District, bOntario Department of Lands & Forests, Parry Sound District, Results Between 1983 and 1989, 168 deer from nine Ontario deer yards, were radio-tracked during at least one migration. Deer from the Loring yard migrated an average of 36 km which was farther than for any other yard (P < 0.0001, Table 2). Migration distance did not differ among the other yards (P > 0.05, Table 2) which together averaged 11 km (SD = 5.2, n = 92). Only four yards had enough deer radio-collared to assess direction bias and to determine the size and shape of their summer dispersion areas (Figure 1 a, b). Deer from three yards demonstrated bias in the direction they travelled from their yards (P < 0.005: Loring 286°; Copelands 100°; Minesing Time of Year Estimation Method Location mid winter Pellet group survey Ontario mid winter Pellet group survey Ontario mid winter Pellet group survey Ontario mid winter Pellet group surveys Ontario Pre-hunt Not provided Ontario Pre-hunt Pellet group surveys Ontario mid winter Pellet group survey Ontario Pre-hunt Cohort analysis Ontario mid winter Aerial Survey Minnesota Aerial survey mid winter Aerial survey Minnesota early spring Aerial survey Pre-hunt Aerial survey Minnesota Pre-hunt sex/age/kill New Brunswick early winter Pellet group Maine Pre-hunt adjusted pellet group Unpublished Report, 1969. Unpublished Report, 1972. 10°). No direction bias was evident in the migration movements of deer from the Wood yard (P = 0.186). Summer dispersion areas were 23 times larger than their associated deer yard, in terms of total land area (SD = 17.6, n = 4) (Table 2). Correction of total land area for percent forest cover in the area sur- rounding each yard (Loring 95%, Wood 93%, Copelands 40% and Minesing 40%) resulted in dis- persion areas which were, on average, 13 times larg- er than their yards (SD = 6.7, n=4) (Table 2). Neither the total area nor forest area multiplication factor was different from our hypothesized estimate of 10 (P>0.370 and P > 0.830 respectively). Both TABLE 2. Characteristics of White-tailed Deer migration from several yards in Ontario 1983-1989. Distance Yard x (SD) Maximum n Loring 36 (19.3) 97 76 Wood 11 (4.3) 22 25 Copelands 13 (4.6) 25 20 Minesing 8 (2.6) 15 20 Borden 11 (5.2) 22 8 Cookstown 7 (1.7) 9 5 Saratoga 13 (8.1) 26 6 Maitland 10 (4.2) 14 4 Bayfield 7 (7.6) 28 4 Summer Dispersion Area(km?) Yard Total Forest Area (km?) Area (Multiple*) = Area (Multiple) 525 5548 (10.6) 5270 (10.0) 30 569 (19.0) 529 (17.6) 12 586 (48.8) 234 (19.5) 12 153(12.8) 61( 5.1) 10 —b — 5 we au 10 — — 6 as pes 9 ie wie aMultiple = Summer Dispersion Area + Yard Area. >Summer Dispersion Area not calculated owing to small sample size. 300 hypothesis tests had low power (1-8 = 0.1074, total area; 1-8 = 0.0301, forest area). Discussion Our data do not lead us to reject our hypothesis that summer dispersion areas are roughly 10 times larger than their associated deer yards. However, it is important to consider that, owing to low sample size and large variance, the ability of out tests to detect differences was low (low power). Swift (1946) reported that the winter range area of White-tailed Deer in northern Wisconsin was 10% of summer range. Moen (1978) suggested that in areas of deep snow, winter range is often 10 to 20% of summer range. Nelson and Mech (1987) is the only other study that reported similar data on a yard-specific basis. They presented data for four Minnesota deer yards, two of which had a sufficient number of radio-tagged migrating deer to compare with our results. The Garden Lake and Isabella deer yards had summer dispersion areas (considered demes by the authors) which were 23 and 30 times larger respec- tively. In their study, “the summer range distribution for deer from each yard was measured from the polygon formed by connecting the outermost sum- mer ranges of individual deer” (Nelson and Mech 1987). It appears that their data do not necessarily represent a minimum area polygon, nor were they corrected to account for area of non-deer range (lakes, rivers, settlements, etc.) or forest area. Nevertheless, our results do compare reasonably well with total land area data from their study. The location of deer yards across regional land- scapes remains relatively constant over time spans of 10 to 30 years and perhaps longer, provided that deer are not extirpated by some mortality factor like preda- tion (Mech and Karns 1977). However, at a finer landscape scale, yard size may vary greatly from year to year. This yearly variation occurs because deer restrict their movements to smaller areas as snow depth increases (Hepburn 1959) and as deer popula- tion size declines (Broadfoot, unpublished data). Since average conditions represent those which deer are exposed to most frequently over time, we think that they best define conditions under which migra- tion traditions develop. Therefore, when defining yard area, it is important to use data collected during win- ters having average snow depths and deer abundance. White-tailed deer population density can change quickly (McCullough 1979). These changes could affect the size of summer dispersion areas if spring migration from yards represented a “random walk” and deer established summer home ranges in the first unoccupied and suitable piece of the landscape. Clearly spring migration is not a “random walk”, since deer show strong fidelity to their summer home ranges (Nelson and Mech 1981; Tierson et al. 1985) with migration traditions passed from mother to off- spring (Hamerstrom and Blake 1939; Nelson and THE CANADIAN FIELD-NATURALIST Vol. 110 Mech 1981). Because of the strong tradition of deer migrating to specific areas within a summer disper- sion area, it seems that summer dispersion area boundaries should be relatively insensitive to short term (5 to 10 years) changes in population abundance. Since the multiplication factor we derived using forest area was closer to our assumed value than the multiplication factor derived for total land area, for- est area may be significant in this relationship. The most accurate depiction of this relationship would probably emerge when the area of summer disper- sion is expressed in terms of both the quality and quantity of deer habitat available on summer range. Therefore, it may be more accurate to say that sum- mer dispersion areas are roughly 10 times larger than their associated yards in terms of deer habitat avail- able in the surrounding landscape. For example, a deer yard measuring 20 km? with surrounding land- scape offering 80% forest cover would have a sum- mer dispersion area measuring 250 km? (20 x 10 + 0.8). The ability to predict the size of summer disper- sion areas is significant in resource management. It allows managers to anticipate the spatial extent of activities which affect deer while they are wintering in yards (eg., loss of habitat through development, high winter mortality due to severe winter weather, effects of supplemental or emergency deer feeding, etc.). It also allows managers to determine where harvest should be allocated to achieve yard specific deer population goals. Although it seems we have a simple “rule of thumb” to determine the size of a summer dispersion area, depicting their shape is not a simple matter since deer show significant bias in the direction they migrate from yards (Sparrow and Springer 1970; Verme 1973; Hoskinson and Mech 1976; Nelson and Mech 1981). In some situations deer are prevented from moving in a particular direction by large water bodies, cities, and other barriers. In our study, deer showed migration direction bias in the absence of any apparent barriers to their movement. We speculate that the spatial distribution of adjacent deer yards may influence the shape of summer dispersion areas. Acknowledgments We thank the many people who helped collect field data and specifically: J. Abott, S. Emes, D. Gilmore, P. Gormley, and B. Kraft for capturing and radio-tag- ging deer, V. Ewing, K. Marionetts, F. McKay, M. Malhiot, S. Nevard, and P. Smith for both capturing and radio-tracking migrating deer and J. Almond, M. Nietfeld, and D. Smith for radio-tracking. Special thanks are extended to J. Hamr, W. Lintack, and B. Thomas for their outstanding contributions to trap- ping, radio-tracking and the logistics of these activi- ties. G. Henry and R. Warren helped compile data. Portions of this study were funded through coopera- tive arrangements with the Northern Ontario Tourist 1996 BROADFOOT, VOIGT, AND BELLHOUSE: DEER DISPERSION AREAS IN ONTARIO 301 o_ Sa EES as Lake Nipissing = Ne ; 7} Lake Huron f N eee 10 Km Adjacent Yards pre vse x Lake Simcoe NW 10 Km l od Adjacent Yards FicurE 1. Distance and direction of migration movements of White-tailed Deer from Loring (1a), Wood, Copelands, and Minesing (1b) deer yards in Ontario 1983-1989. Each closed circle depicts the summer home range location of an individual deer. The outer boundary defines a minimum convex polygon. Adjacent deer yards shown as solid filled polygons. 302 Operators, the Ontario Federation of Anglers and Hunters, and the Nottawasaga Valley Conservation Authority. This is Southern Terrestrial Ecosystems Research Section contribution number 95:05. Literature Cited Banasiak, C. F. 1977. I-167 deer density level studies. Maine Department of Inland Fisheries and Game, Performance Report W-67-R-8. 6 pages. Batschelet, E. 1981. Circular statistics in biology. Academic Press, Toronto. 371 pages. Cumberland, R. E., and A. H. Boer. 1991. Physical characteristics of an insular white-tailed deer population. Transactions of the Northeast Section of the Wildlife Society 48: 126-133. Cumming, H. G. 1961. Spring deer yard surveys and the outlook for deer hunting in 1961. Ontario Fish and Wildlife Review 1: 21-29. Fryxell, J. M., D. H. J. Hussell, A. B. Lambert, and P. C. Smith. 1991. Time lags and population fluctuations in white-tailed deer. Journal of Wildlife Management 55: 377-385. Fuller, T. K. 1990. Dynamics of a declining white-tailed deer population in north-central Minnesota. Wildlife Monographs 101: 1-37. Hamerstrom, F.N., and J. Blake. 1939. Winter move- ments and winter foods of white-tailed deer in central Wisconsin. Journal of Mammalogy 20: 206-215. Hepburn, R. L. 1959. Effects of snow cover on mobility and local distribution of deer in Algonquin Park. Ms. thesis, University of Toronto, Toronto. 55 pages. Hoskinson, R. L., and L. D. Mech. 1976. White-tailed deer migration and its role in wolf predation. Journal of Wildlife Management 40: 429-441. King, D. R. 1976. Estimates of the white-tailed deer pop- ulation and mortality in central Ontario, 1970-1972. Canadian Field-Naturalist 90: 29-36. McCullough, D. R. 1979. The George Reserve deer herd: population ecology of a k-selected species. The Uni- versity of Michigan Press, Ann Arbor. 271 pages. Mansell, W. D. 1974. Productivity of white-tailed deer on the Bruce Peninsula, Ontario. Journal of Wildlife Management 38: 808-8 14. Moen, A. N. 1978. Seasonal changes in heart rates, activi- THE CANADIAN FIELD-NATURALIST Vol. 110 ty, metabolism, and forage intake of white-tailed deer. Journal of Wildlife Management 42: 715-738. Mech, L. D., and P. D. Karns. 1977. Role of the wolf in a deer decline in Superior National Forest. USDA Forest Service Research Paper NC-148. North Central Forest Experimental Station, St. Paul, Minnesota. 23 pages. Nelson, M. E., and L. D. Mech. 1981. Deer social organi- zation and wolf predation in northeastern Minnesota. Wildlife Monographs 77: 1-53. Nelson, M. E., and L. D. Mech. 1986. Deer population in the Central Superior National Forest, 1967-1985. United States Department of Agriculture, Forest Service, North Central Forest Experiment Station, Research Paper NC- 217: 1-8. Nelson, M. E., and L. D. Mech. 1987. Demes within a northeastern Minnesota deer population. Pages 27-40 in Mammalian dispersal patterns. Edited by B. D. Chepko- Sade and Z. T. Halpin. University of Chicago Press. Chicago, Illinois. Schmitz, O. J. 1990. Management implications of forag- ing theory: evaluating deer supplemental feeding. Journal of Wildlife Management 54: 522-532. Severinghaus, C. W., and E. L. Cheatum. 1956. Life and times of the white-tailed deer. Pages 57—186 in The deer of North America. Edited by W. P. Taylor. The Stackpole Company, Harrisburg, Pennsylvania. Snedecor, G. W., and W. G. Cochran. 1980. Statistical methods. The Iowa State University Press, Ames, Iowa. 507 pages. Sparrow, R.D., and P. F. Springer. 1970. Seasonal activity patterns of white-tailed deer in eastern South Dakota. Journal of Wildlife Management 34: 420-431. Swift, E. 1946. A history of Wisconsin deer. Publication 323, Wisconsin Conservation Department, Madison, Wisconsin. 96 pages. Tierson, W. C., G. F. Mattfeld, R. W. Sage, and D. F. Behrend. 1985. Seasonal movements and home ranges of white-tailed deer in the Adirondacks. Journal of Wildlife Management 49: 760-769. Verme, L. J. 1973. Movements of white-tailed deer in upper Michigan. Journal of Wildlife Management 37: 545-552. Received 14 July 1995 Accepted 23 October 1995 Berry Consumption by the American Robin, Turdus migratorius, and the Subsequent Effect on Seed Germination, Plant Vigour, and Dispersal of the Lowbush Blueberry, Vaccinium angustifolium D. R. CROSSLAND and S. P. VANDER KLOET Department of Biology, Acadia University, Wolfville, Nova Scotia BOP 1X0 Crossland, D. R., and S. P. Vander Kloet. 1996. Berry consumption by the American Robin, Turdus migratorius, and the subsequent effect on seed germination, plant vigour, and dispersal of the Lowbush Blueberry, Vaccinium angusti- folium. Canadian Field-Naturalist 110(2): 303-309. The American Robin, Turdus migratorius, is a major frugivore of the Lowbush Blueberry, Vaccinium angustifolium. Laboratory observations established that young robins were capable of rapidly consuming large numbers of berries. After each experimental feeding, excreta were collected, seeds extracted and sown. Germination of excreted seeds was reduced by approximately 17% compared to control seeds. Corroborative evidence was obtained from subsequent field studies in 1985, 1989, and 1993 where the germination success of V. angustifolium seeds from 157 robin excreta collected along the edges of four Nova Scotia blueberry barrens showed a 24% decrease in germination compared to those of control seeds. In 1993 in vitro studies, seeds extracted from fresh fruit were free from pathogens except for a few yeasts; seeds extracted from rotting fruit were largely infected by Botrytis, Aureobasidium and Cladosporium spp., but seeds extracted from robin faeces were invariably infected by bacteria and Mucor spp. as well as 17 additional pathogens. Key Words: American Robin, Turdus migratorius, Lowbush Blueberry, Vaccinium angustifolium, feeding, seed germina- tion, plant vigour, dispersal, pathogens, Nova Scotia. Frugivorous birds and berry-bearing plants are often highly interdependent. Therefore, seed disper- sal by avian frugivores is regarded as a quasi- mutualistic process from which both birds and plants benefit (Snow 1971; Snow and Snow 1988; Cook 1982; Herrera 1982). Many birds depend on the nourishment of the pulp in fruits, while the necessity among plants is to attract seed dispersers to complete a critical phase of their reproductive cycle (Herrera 1982). The pulp in fruit represents an energetic cost for the plants but an attractive reward to birds (Morton 1973; Stapanian 1980). The seeds are a reproductive necessity to plants but are an energy cost for birds in terms of digestive processing (Snow and Snow 1988; Stapanian 1980). In general, birds select fruit with low seed to pulp ratio (Izhaki 1992). Consequently, whenever a bird consumes a berry, it performs its role of the interaction by dropping the seeds anywhere away from the parent plant. Whether it goes to another plant of the same species for a later feeding is irrelevant or even disadvantageous for the first visited plant, as seeds and seedlings often suffer disproportionate mortality near parent plants (Howe 1986). Therefore, dispersal facilitates the coloniza- tion of a new habitat and aids gene flow (Levin and Kerster 1974). Indeed, dispersal may also lessen the risk of seed predation by moving seeds away from the parent plant where their availability is highly pre- dictable (Herrera and Jordano 1981; Janzen 1969). One such berry-bearing plant is the Lowbush Blueberry, Vaccinium angustifolium Aiton. This species is a North American endemic restricted to acid soils and is most commonly found in open areas where it fruits from June to September (Vander Kloet 1988). According to Stiles (1980) the berries are high in sugars and low in lipids and persist a rel- atively short time on the plant. More specifically, Usui et al. (1994) found that 100 g of blueberries contain 1.47 + 0.34 Kceal of fat, 2.12 + 0.18 Kceal of protein and 43.43 + 5.73 Kceal of glucose for a total energy estimate of 52.5 Kcal/100 g. Vander Kloet and Austin-Smith (1986) found that the pulp of each berry contains about 250 calories and the ratio of seed energy to pulp energy is about 1:15. Avian dispersers of blueberries include grouse (Tetraonidae), catbirds and thrashers (Sturninae, Muscicapidae), towhees (Emberizinae, Emberizi- dae), and thrushes (Turdinae, Muscicapidae) (Martin et al. 1951). Indeed, Wheelwright (1986) found that Vaccinium berries were one of the 10 most common fruits in the stomach contents of the American Robin. Furthermore, some avian taxa pass seeds through their digestive tracts in a completely viable state while others kill some or all of them. The latter are classed as seed predators, the former as seed dis- persers. However, the effect of passing Vaccinium seeds through an avian gut is not known, although Krefting and Roe (1949) found that passing blueber- ry seeds through mammalian guts reduced germina-_ tion by at least 20%. Among the numerous avian consumers of Vaccinium angustifolium berries, the American Robin Turdus migratorius L. is considered the most important in eastern Canada (Eaton 1957; Vander 303 304 Kloet and Austin-Smith 1986). Hence, the purposes of this study were (1) to feed ripe blueberries to robins in order to assess the effect of passage of seed through the gut on seed viability and seedling growth of V. angustifolium. (2) To compare the results from the captive trials with observations from wild popu- lations of robins foraging on V. angustifolium. (3) To examine and identify pathogens on seeds separated from excreta and from fresh and fallen berries which might retard germination. Materials and Methods (1) Captive bird trial: Provenance and Maintenance of Birds A nest containing four, approximately 2-week- old, American Robins was obtained from the Gaspereau Valley, Kings County, Nova Scotia, on 11 June 1984, under Environment Canada permit Number 0759. The birds were kept in an indoor aviary (2m?) until they had adequately mastered flight, then were retained in a larger outdoor 3 x 3 x 4 m aviary. The robins were held in captivity 34 days and were released upon completion of the experiments. The birds remained healthy throughout the experimental period. During captivity, they were fed a highly proteinaceous diet approximately 15 times per day using blunt forceps and a small spatula (Formulae in Crossland 1986). Provenance and Maintenance of Plants Fifty 4-year old plants of V. angustifolium grown from seed collected at 40 loci in eastern North America from West Virginia to Newfoundland (see Vander Kloet 1978, for details) were transplanted to 14 cm clay pots in 1983 and kept in cold frames until 3 April 1984 when they were brought into the Acadia University Greenhouse in order to force early flowering so that the ripening of fruit would coincide with the fledging of the robins. Once anthesis occurred, plants were placed en masse outside for several sunny and warm days in early May so that the native pollinators had an opportunity to cross-pollinate them. After anthesis, plants were returned to the greenhouse and watered daily, in order to obtain optimum berry development. Berries ripened in about 45 days. Prior to being placed in the aviary, we picked 300 berries for con- trol, 1.e., 6 from each plant. Seeds were washed from each berry and air-dried; only the plump seeds were counted from each berry because Bell (1957), Darrow (1941), and Vander Kloet (1978) among oth- ers observed that small imperfectly formed seeds are not viable. Vaccinium angustifolium has direct ger- mination, i.e., seeds germinate best immediately after dispersal (Hall et al. 1979). Feeding Experiments Five blueberry plants were put into the aviary so that the robin’s feeding habits could be observed. THE CANADIAN FIELD-NATURALIST Vol. 110 The number of ripe berries on each plant was record- ed in order to calculate how many berries were con- sumed within a given time. Sixteen feeding experiments were conducted from 19 June to 6 July, 1984. Blueberry feedings were completed each morning between 09:00 and 09:30 ADT. The floor of a separate aviary was cov- ered with paper so that the faeces could easily be recovered. Then one robin, arbitrarily chosen from the four, was fed 5 to 10 berries using blunt forceps and left in the aviary. Several changes in procedure were made as the birds matured and became more wary. Instead of isolating a bird in a second aviary for blueberry feedings, it was left with the others. Faeces containing blueberry remains were easily distinguished by their characteristic blue color. Still later, when the robins were no longer receptive to hand-feeding, a large dip net was used to capture a robin, which was then put in a separate aviary where blueberries had been set out previously. The berries were eaten rapidly once the bird was left undisturbed. A minimum of 2 h was allowed for defecation after each feeding trial before excreta were collected. Germination Trials All excreta from each feeding trial were collect- ed; then 50 plump seeds were extracted with for- ceps, and placed on top of a 1:1 mixture of sand and peat in a 9 cm plastic pot. Pots were then placed in a misting chamber under a regimen of 16 h light: 8 h dark and ambient temperature. Corresponding control pots were set up after each feeding experiment. These pots contained 50 seeds of V. angustifolium extracted from fresh blueberries as outlined above. In all, 18 control and 18 excreta pots were set out from 19 June to 6 July 1984. Each pot was observed daily for emergence of the first radicle, first cotyledons and first leaf. On 17 September 1984, the total number of germinants per pot was recorded. The small seedlings were removed from each pot after counting, leaving only the most robust seedling to develop. On 3 October, another seedling count was made to record later germinating seeds; these were also removed. Data Analysis Total number of germinants per pot was record- ed after two months and subjected to a one way ANOVA to determine whether seed ingestion by robins affected the viability of seeds compared to seeds extracted from fresh fruit. Because of unequal variances and non-normal distributions, non-para- metric Mann-Whitney U-tests were performed at each stage (day of appearance of first radicle, first cotyledons and first leaf) to determine whether ger- mination times of robin-ingested seeds differed sig- nificantly from those of controls. 1996 (2) Field Studies: Collection and germination In order to corroborate our experimental data, 157 robin excreta were collected during the 1985, 1988, and 1993 fruiting seasons along the edges of four Nova Scotia blueberry barrens, viz., at the Canne Barrens, Sherwood, Lunenburg County, at the Powell Barrens, Gore, Hants County, at the Biorachan Barrens, 5 km NE of Earltown, Colchester County, and the Parrsboro barrens just east of the town, Cumberland County. These excreta were collected in the following manner. Whenever we observed a small flock of robins feeding in a blueberry barren, we followed, using binoculars, the birds to perching and ablution sites where fresh excreta were collected and individ- ually placed in new 18 ml nalgene vials. Under a dis- secting microscope, well filled, plump seeds of V. angustifolium were removed from each faecal pellet and counted. Lots of 50 seeds were either sown directly on a 1:1 mixture of sand and peat, or the 50 seeds were mixed into a 2 cc faecal-bolus, which was then placed on a 1:1 mix of sand and peat: the latter method simulated field conditions with the fae- cal pellet intact, the former after the faecal pellet had been reworked by wind and rain. For control, several hundred blueberries were picked from areas where robins had been foraging. Berries were chopped for 30 sec in a Waring blender, then the pulp was washed away, and the seed dried, after which samples of 50 plump seeds were separated and sown as described above. Plump seeds of V. angustifolium were separated individually from excreta collected during August 1993 from the Parrsboro barrens, in a positive-pres- sure laminar-flow transfer hood, to 15 ml test-tubes, half of which were filled with about 10 ml half- strength Bacto Potato Dextrose Agar to encourage fungal growth, the other half with blueberry barren soil-agar to simulate field conditions. Similarly, 100 fresh berries and 100 fallen berries were collected from these barrens and individually placed in 18 ml neoprene vials. One plump seed was removed from CROSSLAND AND VANDER KLOET: ROBIN EFFECT ON LOWBUSH BLUEBERRY 305 each berry to a 15 ml test-tube in a positive-pressure laminar-flow transfer hood with scalpel and forceps that were flame sterilized between each transfer. For every treatment 25 to 27 test-tubes were set up and then incubated for 4 months in a growth chamber set at 16 h light at 21°C and 8 hr dark at 15°C. Test-tubes were checked daily for fungal growth and seed germination; mycelia observed growing into agar were looped out and transferred under ster- ile conditions into petri dishes with dextrose agar, incubated and allowed to sporulate so that each fun- gus could be identified if possible. Agar prepara- tion: (1) One-half strength PDA — Add 19.5 g Bacto Potato Dextrose Agar (Difco laboratories, Detroit, Michigan, USA) to 500 ml of distilled H,0 , heat to boiling to dissolve completely. Dispense 10- 11 ml into each test tube. (2) 50% soil extract agar. Add 12 g of Bacto-Agar to 500 ml of distilled H,0. Heat to dissolve completely, then add soil extract which had been prepared as follows: measure 200 ml blueberry barren soil into a 2000 ml beaker, top up to 1000 ml with distilled water and stir; filter mix- ture through 8 layers of cheesecloth. Next dilute the filtrate by one-half with distilled H,0. Add 500 ml of soil extract to the melted agar. Dispense 10-11 ml into each test tube. (3) water agar. Add 12 g Bacto- Agar to 500 ml distilled H,0. Heat to dissolve com- pletely. Dispense 10-11 ml into each test tube. Results Observations of Feeding Habits and Assimilation Rates (1984 experiment) Robins readily accepted blueberries at all times. A single robin was capable of consuming 10 or more berries consecutively, swallowing each whole. Plants presented to four young robins, bearing 40 to 50 ripe fruits each, were picked clean by the birds in 2 to 3 minutes. Seed Germination and Seedling Development (1984 experiment) Ripe berries contained 16 + 5 plump seeds (n = 300) and, of these, 92% germinated; in contrast 76 % TABLE |. Percent germination of Vaccinium angustifolium: American Robin excreted seeds vs. seeds from fresh fruit (1984-1993). Seed provenance Seeds from fresh fruit 1984 experimental 92 n= 16; 800 seeds/treatment 1985 Canne & Gore barrens 54 n= 17; 850 seeds/treatment 1989 Gore & Biorachan barrens 58 n= 13; 650 seeds/treatment 1993 Biorachan & Parrsboro barrens Sy n= 10; 500 seeds/treatment Seeds extracted Seeds in from excreta excreta F P, 76 — 24.186 < 0.0001 39 21 11.489 < 0.0001 44 Dali 7.848 0.0001 41 28 3.316 0.05 306 THE CANADIAN FIELD-NATURALIST Vol. 110 of the seeds in faecal pellets germinated (Table 1). This represents a 17% decrease in germination of robin excreted seeds. The observed differences in on ae germination success between faecal and control q piety groups differed significantly (Table 1). s eran a at Rates of first radicle emergence did not differ sig- 3 i 282 nificantly between faecal and control groups (Mann- o © HOH H Whitney U = 364.5, P > 0.10). Similarly, there were a Q es y 2 a SA RENTS no significant differences (P > 0.10) between germi- =| 3 E nation rates of first cotyledons (U = 361.5) or first as ees ya leaves (U = 360.1) in either group (Table 2). ra cS HH HI o a o é ,. 6 Pa = te Sh Seed germination of field samples collected in 1985, D 1989 and 1993. ON a Although open-pollinated berries, collected from ce blueberry barrens in 1985, 1989 and 1993 contained e. e seeds that had lower germination rates than those s on M6 00 j : g vane S DQ HoH +H]. obtained from experimental plants of V. angustifoli- s es ay P nel etaiee alee ewe S| 2 um (Table 1), the proportions that germinated in each 5 || 5 5 treatment did not vary substantially from the experi- ic 3 > a ye a mental seed populations, and indeed treatments were o . “¢ : . o eo oy St Nort ra significantly different for each of the sampling years 4 |e eg = (Table 1). In general, a 24% decrease in germination % || S ales b d in seeds extracted from robin scats and Slice onels was observed in s aS 5 se HH + s a 53% decrease in germination when the seeds were Bie =| Saas = sown in a scat bolus compared to germination rates a 3 piven alti for seeds taken from fresh fruit (Table 1). Seb | beans & pS NO) That the observed difference in germination rates ea HHS S gs g E = oot ee] > is due to wilting at the cotyledon stage rather than a ps = failure to germinate is illustrated in Table 2. 2 a Whether the seed source is wild or from experimen- 5) 3S ; § Bn tal crosses, the number of days needed for radicle 5 aa o | emergence is not significantly different in any treat- Z 0 anal s 8 8 y y eal § Ss HHH] S ment or source except perhaps when the seed is —_ o 5 . . 2 S=AS] 8 sown within a faecal bolus. Only in 1985 were the Me _ . y, . . . |S Beet ey seeds sown in a faecal bolus tardy in producing radi- |= 2 Ge Smte Tat cles. Similarly the emergence of cotyledons is not aia 2 Sore iaAiROnIUly Gutterent £61 allineotinents econ laut | aoe moon! & significantly different for all treatments except again S : z for the 1985 seeds sown in a faecal bolus. calla Been f= Significant divergence occurs only with the emer- = 2 3 Sn Aa 2 gence of the first leafy shoot. oN | onthe Du OES = In all treatments for all years, blueberry seeds q an He os began to germinate in about two to three weeks, but = a | = = 2 ; 4 = ou ee Sake losses due to damping off were widespread and per- § BR HH 41 § nicious in pots that contained seeds in excreta or S SaaS] & extracted from excreta. Should the first germinatin Oo < o g S L seedlings keel over at the cotyledon stage, emer- = S ence of the primary leaf will necessarily be delayed S 5 een Pp y y ye = 3 until a replacement cohort at the cotyledon stage is ~ Bp . . y S OS established. Hence the delay in primary leaf devel- cs eae a opment in pots that had their seeds set out in a faecal S S pa ize bolus (Table 2) and which eventually accounts for = aan a a. s the 17 to 53% decrease in successful germination in s "a, 00. OH the faecal treatment (Table 2). =| 5 o> Fa . . : s : 6 seg 3/5 Seeds of V. angustifolium incubated in 1993 on o) Ae ° 3 Sie PDA agar, a medium designed to elicit maximum io 8 5 3 Be Z growth of pathogens, had significantly different suc- ‘a ag SEO) Si|5. cess in germination: 89% of the seeds extracted from = OZ 4443 fresh berries germinated successfully; 52% of those 1996 CROSSLAND AND VANDER KLOET: ROBIN EFFECT ON LOWBUSH BLUEBERRY 307 TABLE 3. Number of Vaccinium angustifolium seeds germinating on soil agar or on potato dextrose agar (PDA). Seeds were either extracted from fresh fruit, rotting fruit or from robin faeces collected from the Biorachan Barrens and the Parrsboro Barrens during August 1993. Seed Source Fresh Berries Number of germinating seeds: on soil agar 24 >on PDA 24 Number of seeds not germinating: on soil agar 3 :on PDA 3 Legend: PDA treatment x? = 26.301; P(x’, 99, = 13.815). Soil agar treatment x? = 5.327; P (x75.9; = 5.991 NS). extracted from rotting fruit germinated; but only 22% of those seeds extracted from robin excreta (Table 3). However, parallel samples of seeds incu- bated on blueberry barren-soil agar showed no such trend (x? = 5.327; DF, 0.05 = 5.991). Seeds from both fresh and rotting fruits had similar germination success (89-93%) but seeds extracted from excreta had a success rate of 75% on soil agar, again a decrease of 18% in germination of robin excreted seeds as noted in the previous germination trials (Table 3). Fifteen pathogens were isolated from the three seed sources cultured on the two media. Similar assemblages were isolated from both media but growth was so slow on the soil agar that the pathogens did not overwhelm the germinating seed, either extracted from fresh fruit or from rotting fruit. Only on ten seeds extracted from scats did we observe a mixture of bacteria and fungal hyphae so thick that it killed the germinating seed on soil agar. In contrast, seeds extracted from fresh fruit were free from infection in both PDA and soil agar. In 54 vials, only 12 occurrences of yeasts (Saccharomyces spp) were isolated (Table 4). Seeds from rotting fruits had at least nine groups of pathogens including bacteria; Botrytis spp., Aureobasidium spp. and TABLE 4. Number of pathogens isolated from Vaccinium angustifolium seeds incubated on soil agar and potato dex- trose agar. Seeds were either extracted from fresh fruit, rot- ting fruit or from robin faeces collected from the Biorachan Barrens and the Parrsboro Barrens during August 1993. Fresh __ Rotting Robin Seed Source Fruit Fruit Faeces Bacteria — 6 pay? Saccharomyces 12 7 6 Botrytis — 35 1 Aureobasidium — 44 6 Cladosporium — 38 19 Mucor = — 46 Epicoccum — — 16 Fusarium — — 5 Taxa with one occurrence _ 4 iil Rotting Berries Robin Faeces 25 39 13 6 2 13 12 21 Cladosporium spp. were especially common (Table 4). Seeds from excreta had the highest diversity of pathogens with bacteria and Mucor spp. the most prevalent (Table 4). Discussion Germination of Vaccinium angustifolium seeds consistently decreased by approximately 17% after they had passed through the robins’ gastrointestinal tracts. This corresponded to the findings of Krefting and Roe (1949) who observed a 20% reduction in germination rate of blueberry seeds that had passed through the guts of three rodent species. This reduc- tion in germination rate may in part be due to bacte- rial and Mucor infection. In vitro, bacterial colonies and fungal hyphae were observed growing on all testa which had been extracted from robin faeces collected in the field in contrast to seeds extracted from fresh ripe fruit where only a few yeast infec- tions were found on the testa. Moreover, V. angusti- folium seeds are small, weighing only 31 + 5 mg/100 seeds (Crouch and Vander Kloet 1980). Therefore, the plant actually invests little energy in the produc- tion of individual seeds. Pritts and Hancock (1984) found that V. angustifolium populations allocate on average only 21% of total plant biomass to fruit and seed production most of which is destined for berry pulp, racemes, and pedicels. Vander Kloet and Austin-Smith (1986) estimated that the total calorific value of all seeds in a single berry is only 16 + 2 calories compared to 246 + 62 calories for the pulp. Thus if the energetic investment per seed is minimal, it can be argued that the destruction of some seeds by ingestion is not a major loss to the parent plant, just part of the cost of dispersing seed away from - parent plants. Ingestion of blueberry seeds by robins had no effect on vegetative development. We have no evi- dence that the robins’ digestive processes signifi- cantly help soften or partly digest the seed coat, thereby accelerating germination (Table 2). Short- passage times may account for the limited effect of ingestion by robins. Generally, the bulk of seeds was defecated within 35 minutes after feeding. 308 It can be argued that there are several benefits of Lowbush Blueberry consumption by the American Robin. They move seeds away from the parent plant, thus decreasing local seed densities and increasing the chance that some seeds will be deposited in an area suitable for germination. During the dispersal process, testa are scarcely abraded while in the robins’ digestive tract (Crossland 1986). In addition, our observations of the robins’ feeding habits indi- cated that they were capable of consuming large quantities of berries in a short time. This is advanta- geous as the ripened berries should be dispersed before they fall to the ground beneath the parent plant where, according to Janzen (1970), seedling mortality is often much greater than at more distant sites. Indeed, seeds from rotting fruit had a large array of bacteria and fungi on their testa, especially Botrytis, Aureobasidium and Cladosporium species, but their presence scarcely impeded germination except on PDA media, which of course is designed to elicit maximum fungal growth. Such contamination is to be expected when the seed source is from small-seeded summer fruits such as V. angustifolium which, according to Cipollini and Stiles (1992), have few if any fungal defenses against spores settling on and germinating on berry skins, consequently not only does the fruit rot readily but the testa are infected as well but apparently not to the detriment of successful germination. The behavior of robins may also decrease disper- sal efficiency. Wesley et al. (1986) found that higher seedling densities were found at edges of blueberry barrens where birds perched after feeding: 2.5 + 0.79 seedlings/m? along edges compared to 1.1 + 0.48 seedlings/m? within the barrens. Findings of Holthuijzen and Sharik (1985) and Hoppes (1987) similarly indicated that large quanti- ties of seeds are dispersed under perch sites close to foraging areas. Stiles and White (1986) also argued that dispersal is quite local as thrushes and catbirds which fed on berries in thickets rarely moved into adjacent open fields and birds of open fields rarely visited the adjacent forest. The small but persistent adverse effect that robin dispersal has on V. angustifolium seed germination is nevertheless rather surprising given the large pulp reward accruing to the birds. Indeed, these results corroborate the analyses of Jordano (1995) who found that seed dispersal syndromes are not entirely interpretable as current adaptations to seed dis- persers. Rather the blueberry may be an adaptation to dispersal agents now long extinct. Acknowledgments We wish to thank P. Austin-Smith for his con- structive criticisms of the text. This study was sup- ported by NRCC grant Number A9559. THE CANADIAN FIELD-NATURALIST Vol. 110 Literature Cited Bell, H. P. 1957. The development of the blueberry seed. Canadian Journal of Botany 35: 139-159. Cipollini, M. L., and E. W. Stiles. 1992. Relative risk of fungal rot for temperate ericaceous fruits: effects of sea- sonal variation on selection for chemical defense. Canadian Journal of Botany 70: 868-877. Cook, R. E. 1982. Attractions of the flesh. Natural History 1: 21-24. Crossland, D. R. 1986. Seed dispersal of the lowbush blueberry, Vaccinium angustifoliu. Aiton, by the American Robin, Turdus migratorius L. B.Sc. Honours thesis, Acadia University, Wolfville Nova Scotia. 33 pages. Crouch, P. A., and S. P. Vander Kloet. 1980. Variations in seed characters in populations of Vaccinium sect. Cyanococcus (the blueberries) in relation to latitude. Canadian Journal of Botany 58: 8490. Darrow, G. M. 1941. Seed size in blueberry and related species. HortScience (Proceedings of the American Society for Science) 38: 438-440. Eaton, E. L. 1957. The spread of blueberry seeds through manure and by migrating robins. Hort Science 69: 293-295. Hall, I. V., L. E. Aalders, N. L. Nickerson, and S. P. Vander Kloet. 1979. The Biological Flora of Canada. 1. Vaccinium angustifolium Ait., Sweet lowbush blueberry. Canadian Field-Naturalist 93: 415—430. Herrera, C. M. 1982. Seasonal variation in the quality of fruit and diffuse coevolution between plants and avian dispersers. Ecology 63: 773-785. Herrera, C. M., and P. Jordano. 1981. Prunus mahaleb and birds: the high efficiency seed dispersal system of a temperate fruiting tree. Ecological Monographs 51: 203-218. Holthuijzen, A. M. A., and T. L. Sharik. 1985. The red cedar (Juniperus virginiana L.) seed shadow along a fence line. American Midland Naturalist 113: 200—202. Hoppes, W. G. 1987. Pre- and post-foraging movements of frugivorous birds in an eastern deciduous forest wood- land, USA. Oikos 49: 281-290. Howe, H. F. 1986. Seed dispersal by fruit eating birds and mammals. Pages 123-190 in Seed Dispersal. Edited by D. R. Murray. Academic Press, London. Izhaki, I. 1992. A comparative analysis of the nutritional quality of mixed and exclusive fruit diets for Yellow- vented Bulbuls. Condor 94: 912-923. Janzen, D. H. 1969. Seed-eaters versus seed size, number, toxicity and dispersal. Evolution 23: 1—27. Janzen, D. H. 1970. Herbivores and the number of tree species in tropical forests. American Naturalist 104: 501-528. Jordano, P. 1995. Angiosperm fleshy fruits and seed dis- persers: a comparative analysis of adaptation and con- straints in plant-animal interactions. American Naturalist 145: 163-191. Krefting, L. W., and E. I. Roe. 1949. The role of some birds and mammals in seed germination. Ecological Monographs 19: 269-286. Levin, D. A., and H. W. Kerster. 1974. Gene flow in seed plants. Annals of the Missouri Botanical Garden 68: 223-253. Martin, A. C., H. S. Zim, and A. L. Nelson. 1951. American wildlife and plants. McGraw-Hill, New York. 500 pages. 1996 Morton, E. S. 1973. On the evolutionary advantages and disadvantages of fruit eating in tropical birds. American Naturalist 107: 8-22. - Pritts, M. P., and J. F. Hancock. 1984. Independence of life history parameters in populations of Vaccinium angustifolium (Ericaceae). Bulletin of the Torrey Botanical Club 111: 451-461. Snow, D. W. 1971. Evolutionary aspects of fruit-eating by birds. Ibis 113: 194-202. Snow, D. W., and B. Snow. 1988. Birds and berries: A study of an ecological interaction. T. and A. D. Poyser, Calton, U.K. 268 pages. Stapanian, M. A. 1980. Second International Congress of Systematics and Evolutionary Biology. Abstracts. The University of British Columbia, Vancouver. 441 pages. Stiles, E. W. 1980. Patterns of fruit presentation and seed dispersal in bird-disseminated woody plants in the east- ern deciduous forest. American Naturalist 116: 670-688. Stiles, E. W., and D. W. White. 1986. Seed deposition pat- terns: influence of season, nutrients and vegetation struc- ture. Pages 165-177 in Frugivores and seed dispersal. Edited by E. Estrada, and T. H. Flemming. Academic Press, London. 322 pages. CROSSLAND AND VANDER KLOET: ROBIN EFFECT ON LOWBUSH BLUEBERRY 309 Usui, M., Y. Kakuda, and P. G. Kevan. 1994. Composition and energy valves of wild fruits from the boreal forest of northern Ontario. Canadian Journal of Plant Science 74: 581-587. Vander Kloet, S. P. 1978. Systematics, distribution and nomenclature of polymorphic Vaccinium angustifolium. Rhodora 80: 358-376. Vander Kloet, S. P., and P. J. Austin-Smith. 1986. Energetics, patterns and timing of seed dispersal in Vaccinium section Cyanococcus. American Midland Naturalist 115: 386-396. Vander Kloet, S. P. 1988. The genus Vaccinium in North America. Agriculture Canada, Ottawa, Ontario. 201 pages. Wesley, S. L., N. M. Hill., and S. P. Vander Kloet. 1986. Seed banks of Vaccinium angustifolium Aiton on man- aged and unmanaged barrens in Nova Scotia. Le Naturaliste canadien 113: 309-316. Wheelwright, N. T. 1986. The diet of American robins: An analysis of U.S. biological survey records. Auk 103: 710-725. Received 17 July 1995 Accepted 13 October 1995 Group Hunting Forays of Wintering Northern Harriers, Circus cyaneus: An Adaption of Juveniles? THOMAS BOSAKOWSKI!2 and DWIGHT G. SMITH? 'Department of Biological Sciences, Rutgers University, Newark, New Jersey 07102 2Present address: Beak Consultants, Inc., 12931 Northeast 126th Place, Kirkland, Washington 98034-77115 3Biology Department, Southern Connecticut State University, 501 Crescent Street, New Haven, Connecticut 06515 Bosakowski, Thomas, and Dwight G. Smith. 1996. Group hunting forays of wintering Northern Harriers, Circus cyaneus: an adaption of juveniles? Canadian Field-Naturalist 110(2): 310-313. The social interactions of a wintering population of Northern Harriers (Circus cyaneus) were studied in the Hackensack Meadowlands tidal marshes in New Jersey. Juvenile harriers were numerically dominant in the population and participated in hunting groups significantly more often than adult males or females. Group sizes varied from 2-4 birds (total groups observed = 66 duets, five trios and two quartets). The occasional inclusion of adults in a foraging group was usually the result of juveniles following the adult, presumably for the parasitic benefits of grabbing flushed prey or exploiting high- yield foraging patches. Our observations suggested a non-territorial wintering harrier population documented by observa- tions of 3-6 different individuals frequently hunting the same 12-ha area each day as well as random use of four study quadrats (3-ha) by individuals of all sexes and ages. Territorial defense was observed in only one adult female, which infre- quently attempted to defend a territory. Overall, the prevalence of group-foraging behavior is consistent with the general lack of winter territory in this population. Key Words: Northern Harrier, Circus cyaneus, group hunting, foraging, following, winter ecology, Hackensack Meadowlands, New Jersey. In general, territoriality is lacking or only weakly exhibited in local wintering populations of the Northern Harrier and Hen Harrier (Circus cyaneus) which often gather in large communal roosts from which they fan out to forage over a wide area (Watson 1977). Territoriality or intraspecific agonis- tic behavior were not observed during hunting forays in the course of intensive studies conducted in Michigan (Craighead and Craighead 1956), the Netherlands (Schipper et al. 1975), or Great Britain (Watson 1977). However, another study in the Netherlands (Boedeltje and Zijlstra 1981) reported that eight adult female harriers defended winter terri- tories in a population ranging from 30 to 60 harriers, but that territoriality was absent among male and juvenile harriers. Bildstein and Collopy (1985) also reported frequent and predictable agonistic encoun- ters between harriers (called escorting flights) in Florida and South Carolina wintering populations dominated by adult females. In this paper, we report a study of the social behavior of hunting harriers in a winter population numerically dominated by juveniles. We also include several indices of spatial distribution to document the extent of aggregative versus territorial hunting in this communal-roosting species. Methods The study was conducted in the Hackensack Meadowlands district of Bergen County, northeast- ern New Jersey. The major habitat on the study area is a palustrine tidal marsh dominated by Common Reed (Phragmites communis). Harriers also fre- quently foraged over adjacent overgrown landfills covered with tall grasses, weeds, and occasional small saplings. A total of 159 h of observation of foraging harriers was made during 51 days in the winters of 1986-1987 through 1990-1991. Snowfall was rare and typically melted in a few days. Harrier numbers and behavior were roughly similar each winter. Observations were conducted at 12 different stations in the study area. In one area where most of the observations were made in 1987-1988 (35 hours), we used a stationary pick- up truck as a blind and noted the use of four adjacent quadrats (3-ha each) of habitat which were divided by a road and a creek. In most cases we could dis- criminate between males, females and juveniles (unsexed). On some occasions, poor lighting, dis- tance, or brevity of observations prevented distin- guishing females and unsexed juveniles, and these were recorded as “brown birds” (after Watson 1977; Bildstein and Collopy 1985). A few birds had distinc- tive plumage coloration or were missing primaries, which aided in individual recognition. In winter 1988-1989, two harriers were caught in bow-nets baited with live Starlings (Sturnus vulgaris). These harriers were banded, marked with tail streamers, and released at the site of capture. Observations were recorded according to the fol- lowing criteria: (1) group hunting — 2 or more birds hunting within clear view of each other (< 80 m) with no apparent territorial boundary between them or agonistic behavior. (2) escorting flight — aggressor 310 1996 chasing bird out of a territory or birds flying in tan- dem with an aggressive gesture during, before, or after the flight. (3) intraspecific attacks — aggressor attacking a bird with non-aggressive or neutral atti- tude. (4) piracy - stealing or attempting to steal prey from another bird; (5) interspecific aggression — attacking other species of raptor. Each category was only assigned once per observation, but up to all five categories could be used to describe a single group. To avoid pseudoreplication of observations, we watched each foraging group until it left the area or disbanded. Results Group-hunting forays (n = 78) accounted for 80.2% of social interactions among harriers. Group sizes varied from 2—4, with twos most commonly seen (66 times), followed by trios (5 times) and quar- tets (2 times). Overall, the duration of group encoun- ters was generally brief; usually less than 3 min., but occasionally up to a maximum of 45 min. In most cases it was not possible to observe exactly when a specific bout started and ended, thus statistics on duration were not accurately determined. The extent of involvement in group-and solitary-hunting forays differed among adult males, females, and juveniles (Table 1). Adult males and females occurred less fre- quently in groups than did juveniles (Fisher Exact Test, p = 0.006 and p = 0.067, respectively). Competitive interactions were characterized by interspecific aggression with other raptors (n = 16), intraspecific attacks (n = 14), escorting flights (n = 4), and piracy (n = 2). Intraspecific aggression occurred at the same rate as group hunting in adults, but only rarely between juveniles or between juve- niles and brown birds (Table 2). Juvenile harriers were involved in most group hunting, followed by brown birds, and most of these brown birds were probably juveniles (using the ratio of identified juve- niles to females, 69% in Table 1). Usually the non- aggressive group interactions of adults involved tol- erating the presence of a following juvenile (para- sitic relationship), although possible altruistic (parental/familial) ties could not be excluded as fam- ily histories were not known. The use of space by foraging harriers in a four- quadrat area was studied in winter 1987-1988. Harrier use of space did not deviate from a random BOSAKOWSKI AND SMITH: GROUP HUNTING OF NORTHERN HARRIERS 311 TABLE 2. Social interactions stratified by sex and age in wintering Northern Harriers in northeastern New Jersey. Group Hunting Male Female Juvenile Brown Male 1 1 2 0 Female — 0) 11 5 Juvenile = — Di 11 Brown — — — 21 Intraspecific Aggression Male Female Juvenile Brown Male yD) 0 D 0 Female — D 7 2 Juvenile — — D} 1 Brown — — — 1 distribution, even when stratified by sex and age (Table 3), thus supporting the general trend of non- territoriality observed in all years. On several occa- sions, as many as three harriers were seen hunting simultaneously within a single quadrat. On days with observations lasting 3 hours or more (n = 7), we were able to identify individually from 3 to 6 differ- ent harriers (mean = 4.4) hunting within the same 12-ha grid-zone that day which further demonstrates the non-territorial, overlapping nature of harrier for- aging in this tidal marsh. In winter, 1986-1987, an adult female attempted to defend an area; four escorting flights and one interspecific attack were observed, but despite her efforts there were frequent intrusions by many other harriers (Table 3). In late winter 1987, two marked juveniles and a single male that had been captured and released were observed in the same area in close proximity in two of three subsequent sightings. Discussion Wintering juvenile harriers frequently participated in group-hunting flights in which the relationships could be considered either mutualistic or parasitic. The social relationships between members of a for- aging group were cooperative to the extent that very little agonistic behavior resulted despite relatively close individual distances. Adult participation in these social foraging groups was the result of tolerat- ing close following by juveniles. Although social TABLE 1. Extent of group and solitary hunting forays in wintering Northern Harriers in northeastern New Jersey. Male Female solitary 20 Si; group 4 17 total 24 54 % grouped* 16.6 31.5 Juvenile Brown Total 62 45 164 51 37 109 113 80 273 45.1 46.2 40.0 *the distribution of grouped and solitary harriers differed significantly by sex/age classsification (x2, df=3, p < 0.02). S12 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 3. Use of space by hunting Northern Harriers in winter with regard to sex and age in a four-quadrat grid zone in northeastern New Jersey. Area A B C D Totals juvenile 14 10 27. 23 74 female 20 11 12 13 56 male 6 7 5 4 22 brown 10 3 8 S) 26 totals 50 31 52 45 * *the rows and columns of the 4x4 contingency table were not significantly different (y2= 13.052, d.f. = 9, p = 0.16). foraging groups are common in birds, mammals, and fish (Morse 1980), such behavior is rather infre- quently described for Falconids (Newton 1979; Bednarz 1988; Ellis et al. 1993) which are generally viewed strictly as solitary hunters after family groups disperse in late summer. In this study, group-foraging behavior was largely restricted to juvenile harriers or to juveniles follow- ing adult females. This behavior occurred in a win- tering population dominated by juveniles and where winter territoriality among harriers was rare. These conditions were contrary to a study in South Carolina and Florida (Bildstein and Collopy 1985) that reported frequent agonistic encounters as the result of adults (primarily females) trying to defend their territoriés and to prevent piracy by less numer- ous invading juveniles. Boedeltje and Zijlstra (1981) reported that winter territoriality occurred only in adult females but other investigators found no evi- dence of territoriality within wintering groups of har- riers (Craighead and Craighead 1956; Schipper et al. 1975; Watson 1977). All studies considered, winter territoriality appears to be relatively uncommon and restricted almost entirely to adult female harriers, thus providing no conflicting field evidence against a group-foraging hypothesis for juveniles. Watson (1977: 92) stated that it is not uncommon for two or three hunting harriers to be in view at the same time in autumn or winter in Great Britain, when voles are abundant. Dickson (cited in Watson 1977: 92) observed up to five harriers together hunt- ing in two adjacent 34-ha kale fields. While no pre- vious investigators have made specific reference to cooperative-hunting effects between harriers, Watson (1977: 93) reported two interspecific obser- vations of a Merlin (Falco columbarius) and a harri- er hunting closely together. In these observations, both raptors apparently benefitted from the associa- tion by catching birds that were fleeing from the other associate. This association is analogous to a well-known group-foraging technique known as “beating” (Morse 1980; Ellis et al. 1993). Dinsmore (1973) has shown that Cattle Egrets (Bubulcus ibis) can increase their foraging efficiency more than three-fold when attending cattle than when foraging alone. Even in the absence of cattle, Wiese and Crawford (1974) have observed cooperative beating by several Cattle Egrets advancing abreast through the vegetation. The latter observation does not differ functionally from our observations of aerial hunting forays by groups of harriers. Although the mutual and parasitic advantages of beating may be important for hunting juvenile harri- ers, we were unable to prove its success in the field because prey captures were rarely observed in the tall reed stands (2—3 m). However, on one occasion, two juvenile harriers hunting in parallel formation simul- taneously struck at a prey. One hawk was apparently successful as a prey squeal was heard and the harrier remained for 11 min., presumably feeding. With this strategy, inexperienced juveniles can increase their encounter rate with potential prey as well as take advantage of prey that are already startled and which might be less aware of the second following predator. We also postulate that juveniles group more often with other juveniles than with adults because: (1) juveniles might be less adept hunters (Newton 1979) so two or more might expose or flush more prey than each foraging individually, (2) adults are sometimes territorial (Boedeltje and Zijlstra 1981; Bildstein and Collopy 1985) and are, therefore, less likely to toler- ate a following parasitic juvenile. In addition to the effect of beating, group foraging may facilitate location of high-density-prey patches as harriers use foraging by other harriers as cues to locate food (Ekman and Hake 1988). Thus, the habit of winter communal roosting by harriers (see Watson 1977 for a review) may have a central role in facili- tating group hunting cooperation. Ward and Zahavi (1973) proposed that communal roosts may serve as information centers on food-source locations. In this regard, Watson (1977) speculated that harriers return- ing to the roost at dusk with a visibly full crop might reveal the direction of better prey areas [high-yield foraging patches] or that the “hungry” might follow the “full” hawks the next morning. Thus, parasitic relationships may have developed when juveniles fol- lowed closely behind an adult, apparently to take advantage of flushed prey or to “steal” the location of high-yield foraging patches. 1996 Bednarz (1988) recently reported on cooperative hunting groups of Harris’ Hawks (Parabuteo unicinc- tus) which are mainly sit-and-wait predators. These hawks showed a concerted effort to catch a single large prey item (Sy/vilagus spp.) which was eaten by the group. In contrast, the harrier is an on-the-wing predator that preys on smaller prey species (voles, _ songbirds: - Craighead and Craighead 1956; Watson 1977) which are consumed individually. Hence, it fol- lows that the cooperative group-hunting flights of har- riers are less highly structured because prey caught are only of advantage to the successful individual. Literature Cited Bednarz, J.C. 1988. Cooperative hunting in Harris’ Hawks (Parabuteo unicinctus). Science 239: 1525-1527. Bildstein, K. L., and M. W. Collopy. 1985. Escorting flight and agonistic interactions in wintering Northern Harriers. Condor 87: 398-401. Boedeltje, G., and M. Zijlstra. 1981. Territorialiet, bio- type- envoedselkeuze bij de Blauwe Kiekendief Circus cyaneus in de winter. Limosa 54: 73-80. Craighead, J. J., and F. C. Craighead, Jr. 1956. Hawks, owls, and wildlife. Stackpole Publishing Co., Harris- burg, Pennsylvania. BOSAKOWSKI AND SMITH: GROUP HUNTING OF NORTHERN HARRIERS SII} Dinsmore, J. J. 1973. Foraging success of cattle egrets, Bulbulcus ibis. American Midland Naturalist 89: 242-246. Ekman, J., and M. Hake. 1988. Avian flocking reduces starvation risk: an experimental demonstration. Behavioral Ecology and Sociobiology 22: 91-94. Ellis, D. H., J. C. Bednarz, D. G. Smith, and S. P. Fleming. 1993. Social foraging classes in raptors. BioScience 43: 14—20. Morse, D. H. 1980. Behavioral mechanisms in ecology. Harvard University Press, Cambridge, Massachusetts. Newton, I. 1979. Population ecology of raptors. Buteo Books, Vermillion, South Dakota. Schipper, W. J. A., L. S. Burma, and P. H. Bossenbroek. 1975. Comparative study of hunting behavior of winter- ing Hen Harriers Circus cyaneus and Marsh Harrier Circus eruginosus. Ardea 63: 1—29. Ward, P., and A. Zahavi. 1973. The importance of cer- tain assemblages of birds as ‘information centres’ for food finding. Ibis 115: 517-534. Watson, D. 1977. The Hen Harrier. T. & A.D. Poyser, Berkhamsted, England. Wiese, J. H., and R. L. Crawford. 1974. Joint “leap- frog” feeding by ardeids. Auk 91: 836-837. Received 26 July 1995 Accepted 14 November 1995 The Dense-leaved Pussy’s-toes, Antennaria densifolia (Asteraceae: Inuleae): An Addition to the Vascular Flora of British Columbia JERRY G. CHMIELEWSKI Department of Biology, Slippery Rock University, Slippery Rock, Pennsylvania 16057 Chmielewski, Jerry, G. 1996. The Dense-leaved Pussy’s-toes, Antennaria densifolia (Asteraceae: Inuleae): An addition to the vascular flora of British Columbia. Canadian Field-Naturalist 110(2): 314-317. The first records of Antennaria densifolia from southern British Columbia are cited. The identity of previous citations from Montana are questioned. The British Columbia specimens are all pistillate and as such may represent the polyploid, apomictic form of the species. Basal leaf length, involucre length, and the nature of the tips of the upper cauline leaves may best be used to differentiate between A. densifolia and A. pulvinata. Future collections may indicate that A. densifolia has not only a more southern distribution but that it also occurs in the intervening mountainous area between southern British Columbia and the southern Mackenzie Mountains, the latter initially considered to represent the southern limits of the species. Key Words: Antennaria densifolia, Antennaria pulvinata, Dense-leaved Pussy’s-toes, flora, range extension, British Columbia. Renewed interest in conservation biology over the past two decades has resulted in the widespread recognition that the natural diversity of ecosystems must be maintained. Because rare plants and animals are an important part of this diversity, they and their habitats should be protected (Douglas et al. 1981). Proper identification of these taxa is mandatory if they are to be used to support arguments relative to the protection of an area because of its unique natu- ral diversity. Antennaria densifolia A.E. Porsild (Dense-leaved Pussy’s-toes) was described as having minute, densely congested, obovate to oblanceolate, or obtuse basal leaves and as such was recognized as a well-marked species of the Alpinae group (Porsild 1945). Basal leaf morphology could readily be used to distinguish A. densifolia from both A. compacta Malte and A. cana Fernald & Wiegand. Although the arctic A. densifolia superficially resembles the Cordilleran A. pulvinata Greene, the two species dif- fer with respect to the possession by the latter of larger heads, paler bracts, and hispid achenes (Porsild 1945). Antennaria densifolia was initially considered endemic to the upper eastern slope of the Mackenzie Mountains in the Northwest Territories. The species is now known to also occur in the Ogilvie Mountains and southern Richardson Mountains of the Northwest Territories and Yukon Territory (Porsild and Cody 1980). Antennaria den- sifolia was said to typically inhabit dry, turfy lime- stone screes (Porsild 1945). Porsild (1975) subsequently described a second short-leaved arctic species, A. ellyae A.E. Porsild, which could be distinguished from A. densifolia by its spreading, leafy, stolon-like branches and by its twice-as-tall flowering peduncle that terminates in an elongated, cymose inflorescence. Chmielewski and Chinnappa (1990) considered A. ellyae to be conspe- cific with A. densifolia, noting that the minor differ- ences in vegetative morphology merely reflected a growth response to local environmental conditions. Using a taxonomic species concept, Bayer (1989a) concluded that A. aromatica Evert and A. densifolia could be considered discrete species because they are morphologically distinct. The two species were readily distinguished by the presence of stalked glands in A. aromatica and the complete lack of these glands in A. densifolia. Three additional char- acters, the presence or absence of flat, scarious, lin- ear-lanceolate tips at the ends of the upper cauline leaves, basal leaf length, and phyllary length, could also be used in concert to separate the species, but not as reliably as the former character. The most sig- nificant aspect of this study, however, was that Montana collections were identified as A. densifolia, representing a disjunction of approximately 1850 km south of the nearest population in the Northwest Territories. Chmielewski (1993) used gross morphology of type and non-type material, the Pearson product- moment correlation coefficients presented by Bayer (1988), the protologues of A. aromatica and A. pulv- inata, the fallibility of using glandularity as a diag- nostic character, and the results of canonical variates analysis to conclude that A. aromatica and A. pulv- inata are conspecific. The latter specific epithet was deemed the legitimate name for the species. Bearing in mind that my understanding of A. pulv- inata has changed over the past decade to include A. aromatica, this investigation initially evaluates mor- phological affinity between A. pulvinata and A. den- sifolia. Conclusions based on this evaluation are then 314 1996 used to reassess the identification of individuals from the single disjunct population of A. densifolia reported by Bayer (1989a), as well as assess the identification of several small-leaved British Columbia collections. Materials and Methods Mature, predehiscent specimens of Antennaria densifolia (N=69), A. pulvinata (N=153), the dis- junct Montana population of A. densifolia (both pis- tillate and staminate specimens of Bayer, DeLuca and Lebedyk, MT-725, RM530380, and Lackschewitz 4611, MONTU73034) and the small leaved individ- uals from British Columbia (N=5) were borrowed from ALTA, CAN, CAS, CM, COLO, DAO, F, GH, LEA, MO, MONTU, MOR, MT, NDG, OS, PH, RM, SLRO, UAC, UB, UC, and US (Holmgren et al. 1990) for inclusion in the phenetic study. Qualitative characters used to identify specimens of A. densifo- lia and A. pulvinata prior to analysis included habit, degree and type of pubescence, color, shape, and texture of phyllaries, and the presence or absence of papillae on the achenes. These characters were not used directly in the phenetic analysis. Character selection, and specimen selection and identification followed the methodology of Chmielewski (1993). Data were collected for 16 quantitative characters for each specimen following Chmielewski (1994a) with the exceptions that mea- surements were made on the middle cauline leaf as opposed to either the lower or upper-one, and that neither style length nor lobe length were included among the characters analyzed. The reasons for selecting these characters have already been stated elsewhere (Chmielewski and Chinnappa 1988a, 1991; Chmielewski et al. 1990a, 1990b; Chmielewski 1993). The SAS (SAS Institute Inc. 1989) DISCRIM pro- cedure (including the options list, simple, pool, slpool, crossvalidate, crosslist, and posterr) was used to perform classificatory discriminant analysis (fol- lowing Chmielewski 1994a) on the specimens of A. densifolia and A. pulvinata. Rates of correct identifi- cation and Geisser classification probabilities (Pimentel and Frey 1978; Pimentel 1979) were used as indicators of separation between the species. Tests for equality of group centroids were performed as part of the analysis. The same data set was used to both define and evaluate the classification criterion. Classification through crossvalidation (SAS cross- validate option) eliminated the problem of circularity in the analysis because each specimen was identified using a discriminant function that was computed from specimens exclusive of the specimen being classified. The classification criterion was subse- quently used to classify collections from the disjunct Montana population, as well as the small leaved individuals from British Columbia, to either A. den- sifolia or A. pulvinata. A posteriori classification of CHMIELEWSKI: DENSE-LEAVED PUSSY’ S-TOES 315 specimens through the use of the classification crite- rion assumes that specimens classified in this man- ner do in fact belong to either A. densifolia or A. pul- vinata. Results Evaluation of the discriminant function through the use of crossvalidation classification indicated that 98.2% (Antennaria densifolia, N=69, 100%; A. pulvinata, N=149, 97.4%) of the 222 specimens were assigned (with the highest probability) to the correct a priori species. Geisser assignment probabil- ities for the 218 correctly assigned specimens aver- aged (mean + standard deviation) 0.9809 + 0.0770 for A. densifolia and 0.9811 + 0.0807 for A. pulvina- ta. The Mahalanobis distance between species cen- troids and associated F-value indicated that they were significantly different (P < 0.0001). The low error count in conjunction with the high Geisser assignment probabilities indicated that a posteriori classification of the Montana and British Columbia collections through the use of the classification crite- rion would yield acceptable results. The five small-leaved individuals from British Columbia were consistently classified through the use of the classification criterion to A. densifolia. Geisser assignment probabilities averaged 0.9665 + 0.0523 for these specimens. Three of the four specimens from the disjunct Montana population of A. densifolia were classified using the classification criterion to A. pulvinata. Geisser assignment probabilities averaged 0.9638 + 0.0575 for these specimens. A single staminate spec- imen (Lackschewitz 4611, MONTU73034) was assigned to A. densifolia (0.9970). If however, pistil- late and staminate plants were analyzed separately and only those characters identified by stepwise dis- criminant analysis were subsequently used in the classificatory discriminant analysis, all five small- leaved individuals from British Columbia were assigned to A. densifolia (0.9812 + 0.0331), and all four Montana specimens from the disjunct popula- tion of A. densifolia were assigned to A. pulvinata (0.9255 + 0.1216). Discussion Results based on the a posteriori classification of individuals from the disjunct Montana population previously identified as Antennaria densifolia do not support Bayer’s (1989a) view that they belong to that species, but rather that they bear morphological affinity to A. pulvinata. This conclusion is no doubt partly due to the very different interpretation we have relative to morphological variation within A. pulvinata and A. aromatica (Bayer 1989a, 1989b, 1991; Chmielewski and Chinnappa 1988b; Chmielewski 1993, 1994b) especially with respect to glandularity. 316 Bayer (1989a) stated that A. aromatica occurred only east of the continental divide in predominantly unglaciated areas and hypothesized that it could be restricted to the Front Ranges east of the divide as a result of climatic factors, that is, these mountains receiving less rainfall than those to the west in west- ern Montana and Idaho. Although I agree with Bayer (1991) that two specimens cited by Chmielewski and Chinnappa (1988b) were misidentified at the time, the authors did also report a single British Columbia collection that occurs west of the continental divide in an area which was previously glaciated. Bayer (1991) did not verify the status of the latter collec- tion. The present study also includes several addi- tional collections of A. pulvinata from British Columbia as well as one from Idaho. The remoteness of the subalpine habitat in which the species occurs has likely contributed to the paucity of collections. Although I do not agree with Bayer (1989a) that the previously cited Montana collections represent disjunct A. densifolia, five specimens from British Columbia were classified through the use of the clas- sification criterion to A. densifolia. These specimens were all pistillate and may represent a polyploid apomictic form of A. densifolia. The occurrence of A. densifolia in southern British Columbia is not sur- prising, however, as Bayer (1989a) noted that a more widespread preglacial distribution was likely. As with A. pulvinata, the lack of previous reports may simply be a consequence of the remoteness of suit- able subalpine to alpine habitat. It is possible that in the future additional collections may be found which will close the distributional gap between these south- ern British Columbia populations and those in the southern Mackenzie Mountains. Additionally, it is equally likely that more southern populations of A. densifolia will be discovered in the future. Antennaria densifolia and A. pulvinata may be distinguished according to the following characteris- tics. Basal leaves in A. densifolia are typically less than 6.0 (average 5.4) mm long and the involucre less than 5.5 (average 4.9) mm long. Staminate plants are slightly smaller on average with involu- cres typically less than 4.5 (average 4.2) mm long. The upper cauline leaves in this species typically ter- minate in flat, scarious, linear-lanceolate tips. Basal leaves in A. pulvinata are typically greater than 6.0 (average 8.7) mm long and the involucres typically greater than 5.5 (average 6.2) mm long. The upper cauline leaves in this species typically lack the flat, scarious, linear-lanceolate tips. Staminate plants are slightly smaller on average with involucres typically greater than 4.5 (average 4.8) mm long. The foliage of A. pulvinata may be covered in stalked glands but this is never true of A. densifolia. British Columbia Specimens: Mount Apex, SW of Penticton, occasional in bare, gravelly-rocky areas at summit and on upper slopes, THE CANADIAN FIELD-NATURALIST Vol. 110 elevation 7000’ and up, J.A. Calder and D.B.O. Savile 10708, July 18, 1953, DAO443861. Paradise Mine about 15 mi W of Windermere, occasional on bare shale summit, elevation 8500’, J.A. Calder and D.B.O. Savile 11320, Aug 1, 1953, DAO683766. Baldy Mountain, approximately 7.5 mi ENE of Littlefort, 51°27’N and 120°03’W, occasional in rocky areas near summit, elevation 7228’, J.A. Calder, J.A. Parmelee, and R.L. Taylor 19866, Aug 1, 1953 DAO683760. Mount McLean at Lillooet, occasional on rocky-gravelly slope, elevation 7500’, J.A. Calder, D.B.O. Savile, and J.M. Ferguson 15576, Sept 6, 1954, DAOS576733. Cathedral Lakes District, Ashnola Range, between Red Mountain and Mount Bomford, 49°04’N and 120°12’W, common in grassy rocky area at 7850’, J.A. Calder, J.A. Parmelee, and R.L. Taylor 19660, Aug 2, 1956, DAOS576739. Acknowledgments The curators at ALTA, CAN, CAS, CM, COLO, DAO, F, GH, LEA, MO, MONTU, MOR, MT, NDG, OS, PH, RM, SLRO, UAC, UB, UC, and US are thanked for the loan of specimens. Literature Cited Bayer, R. J. 1988. Typification of western North American Antennaria Gaertner (Asteraceae: Inuleae); Sexual species of sections Alpinae, Dioicae, and Plantaginifoliae. Taxon 37: 292-298. Bayer, R. J. 1989a. A systematic and phytogeographic study of Antennaria aromatica and A. densifolia (Asteraceae: Inuleae) in the western North American Cordillera. Madrono 36: 248-259. Bayer, R. J. 1989b. A taxonomic revision of the Antennaria rosea (Asteraceae: Inuleae: Gnaphaliinae) polyploid complex. Brittonia 41: 53-60. Bayer, R. J. 1991. A note on the geographic range of Antennaria aromatica Evert (Asteraceae: Inuleae). SIDA 14: 505-506. Bayer, R. J. 1992. Some observations on morphometric analysis of Antennaria (Asteraceae: Inuleae): Reply. Canadian Journal of Botany 70: 2316-2317. Chmielewski, J. G. 1993. Antennaria pulvinata Greene: The legitimate name for A. aromatica Evert (Asteraceae: Inuleae). Rhodora 95: 261-276. Chmielewski, J. G. 1994a. The Antennaria frieseana (Asteraceae: Inuleae) polyploid complex: Morphological variation in sexual and agamospermous taxa. Canadian Journal of Botany 72: 1018-1026. Chmielewski, J. G. 1994b. Evaluation of the taxonomic status of Antennaria bayardi, Antennaria brunnescens, and Antennaria foggii (Asteraceae: Inuleae). Canadian Journal of Botany 72: 1775-1777. Chmielewski, J. G., and C. C. Chinnappa. 1988a. The genus Antennaria (Asteraceae: Inuleae) in North America: Multivariate analysis of variation patterns in Antennaria rosea sensu lato. Canadian Journal of Botany 66: 1583-1609. Chmielewski, J. G., and C. C. Chinnappa. 1988b. Range extension of Antennaria aromatica Evert (Asteraceae: Inuleae). SIDA 13: 256-258. 1996 Chmielewski, J. G., and C. C. Chinnappa. 1990. The genus Antennaria (Asteraceae: Inuleae). in arctic North America: Chromosome numbers and taxonomic notes. Rhodora 92: 264-276. Chmielewski, J. G., and C. C. Chinnappa. 1991. Gender dependent and independent morphological dimorphism in sexual and apomictic Antennaria monocephala sensu lato. Canadian Journal of Botany 69: 1433-1448. Chmielewski, J. G., and C. C. Chinnappa. 1992. Commentary on Bayer’s use of incomplete data matrices and weighted qualitative characters in phenetic studies of North American Antennaria: Commentary. Canadian Journal of Botany 70: 2313-2315. Chmielewski, J. G., C. C. Chinnappa, and J. C. Semple. 1990a. Patterns of intraspecific variation in Antennaria alborosea, A. corymbosa, A. marginata, A. microphylla, A. parvifolia, and A. umbrinella. Plant Systematics and Evolution 169: 123-150. Chmielewski, J. G., C. C. Chinnappa, and J. C. Semple. 1990b. The genus Antennaria Gaertner (Asteraceae: Inuleae) in western North America: Morphometric anal- ysis of A. alborosea, A. corymbosa, A. marginata, A. microphylla, A. parvifolia, A. rosea, and A. umbrinella. Plant Systematics and Evolution 169: 151-175. Douglas, G. W., G. W. Argus, H. L. Dickson, and D. F. Brunton. 1981. The rare vascular plants of the Yukon. Syllogeus Number 28, National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario. CHMIELEWSKI: DENSE-LEAVED PUSSY’ S-TOES Sly) Holmgren, P. K., N. H. Holmgren, and L. C. Barnett. 1990. Index Herbariorum, 8th edition. Part I. The herbaria of the world. The New York Botanical Garden, Bronx, New York. Pimentel, R. A. 1979. Morphometrics: The multivariate analysis of biological data. Kendall/Hunt Publishing Co., Dubuque, Iowa. Pimentel, R. A., and D. F. Frey. 1978. Multivariate analy- sis of variance and discriminant analysis. Pages 247-274 in Quantitative ethology. Edited by P. W. Colgan. John Wiley & Sons, New York, New York. Porsild, A. E. 1945. The alpine flora of the east slope of Mackenzie Mountains, Northwest Territories. Bulletin Number 101, Biological Series Number 30, National Museum of Canada, Ottawa, Ontario. Porsild, A. E. 1975. Materials for a flora of central Yukon Territory. National Museum of Natural Sciences Publications in Botany, Number 4, National Museums of Canada, Ottawa, Ontario. Porsild, A. E., and W. J. Cody. 1980. Vascular plants of Continental Northwest Territories, Canada. National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario. SAS Institute Inc. 1989. SAS/STAT™ user’s guide, ver- sion 6, Fourth Edition, Volume 1, SAS Institute Inc, Cary, North Carolina. Received 14 August 1995 Accepted 29 January 1996 Dispersal Characteristics of Two-year-old Beavers, Castor canadensis, in Western Montana TIMOTHY R. VAN DEELEN! and DANIEL H. PLETSCHER Wildlife Biology Program and Montana Cooperative Wildlife Research Unit, University of Montana, Missoula, Montana 59812 ‘Current address: Illinois Natural History Survey, 607 E. Peabody Drive, Champaign, Illinois 61821 Van Deelen, Timothy R., and Daniel H. Pletscher. 1996. Dispersal characteristics of two-year-old Beavers, Castor canadensis, in western Montana. Canadian Field-Naturalist 110(2): 318-321. The movements of 22 two-year-old Beavers (Castor canadensis) were monitored in montane habitat using radio-telemetry. Twelve did not disperse. Dispersal distance, dispersal dates, and settlement dates of 10 dispersers varied. Disperser survival rate was 0.70. Wide variation in dispersal dates, and settlement dates, suggests the existence of a summertime sub-popula- tion of transient Beavers, unattached to traditional colonies. Key Words: Beavers, Castor canadensis, Montana, dispersal. Two-year-old Beavers disperse from natal colonies to establish new colonies, and to replace breeders lost to trapping or natural mortality (Bradt 1938; Svendsen 1980). However, not all two-year- olds disperse from their natal colonies. Observations of colony composition indicate delayed dispersal (Brooks et al. 1980; Peterson and Payne 1986) that might be related to high population density (Bergerud and Miller 1977; Payne 1982; 1984; Busher et al. 1983). Dispersers generally leave natal colonies in late winter or early spring and live as a “floating” popu- lation of transients before settlement (Townsend 1953; Aleksiuk 1968; Svendsen 1980; Allred 1981). They typically move along waterways through occu- pied territory but are prevented from staying by resi- dent Beavers (Townsend 1953; Aleksiuk 1968; Allred 1981; Hodgdon and Lancia 1983). Nearly all of what is known about Beaver disper- sal comes from mark-recapture studies, a technique limited by recapture success. Mark-recapture studies also provide little information on the timing of dis- persal and settlement, or movement during the dis- persal-settlement interval. We used radio-telemetry to describe the timing and movement characteristics associated with the dispersal of two-year-old Beavers in western Montana. Study Area Study areas consisted of four secondary drainages of the Clark Fork River in west-central Montana (approximate latitude 47° 00’ N, approximate longi- tude 114° 30’ W); Upper Willow and Meadow Creeks in Granite County, Rattlesnake Creek in Missoula County, and Fish Creek in Mineral County. Dominant bank vegetation in all areas con- sisted of willows (Salix spp.), alders (Alnus spp.), and Red Osier Dogwood (Cornus stolonifera). Surrounding uplands were montane conifer forests (Jackson 1991). Mean annual precipitation was 40—80 cm/year and mean annual snowfall was 80—160 cm/year. Mean January temperature was -1.1—-17.8°C with 150-210 days/year when temperatures were below freezing (O°C, United States Department of the Interior Geological Survey 1970). In spite of the cold temperatures, these streams were free-flowing and relatively ice-free during the winters of 1989/90 and 1990/91. Springtime high water periods occurred between mid-April and late June (Van Deelen 1991). Methods Beavers were captured in Hancock live-traps (Hancock Trap Co., Custer, South Dakota) baited with fresh Cottonwood (Populus trichocarpa), aspen (Populus spp.), or willow twigs and a castor-based lure. In 1989 and 1990, trapping began the third week of March and continued through the first week of June. We immobilized live-trapped Beavers with 150-200 mg of ketamine hydrochloride and 2.5 mg of acepromazine maleate to facilitate morphological measurement and sex determination (Jackson 1991). We assigned age-classes based on morphological development (Patric and Webb 1960; Payne 1979; Jackson 1991). Two-year-old Beavers were intraperitoneally implanted with radio-transmitters (Model IMP400, Telonics Inc., Mesa, Arizona.) using methods described by Jackson (1991). The next day, Beavers were released at the capture site. Radio-transmitters were configured for a minimum battery life of 24 months. We located transmitter-equipped Beavers weekly in 1989 and every other day in 1990. The increased effort in 1990 was an attempt to refine the move- 318 1996 ment data. We recorded locations for each Beaver from the time of their release in the spring until con- struction of a food cache and continued use of the same daytime resting site suggested that the Beaver had selected a site to spend the winter. Dispersal date was defined as the date of the last location in the natal colony. Settlement date was defined as the date of the first location in the subsequent over-wintering site. Beavers alive at the end of the field season were located the following spring to determine over-win- ter mortality. We located Beavers during daylight hours using signal strength to “home in on” daytime resting sites (White and Garrott 1990), and used fixed-wing aircraft to find Beavers which had moved extensively. Nocturnal or daytime movements that did not require radio-equipped Beavers to establish new resting sites were not considered. For each record of telemetry locations, we used a plan measure (Alvin and Co., Inc., Windsor, Connecticut) to measure the shortest stream distance from release site to Beaver location (= daytime resting site). We recorded distances to the nearest 0.1 km. Dispersers were defined as those Beavers that left their capture site (assumed to be within the natal colony’s home range) without returning. In all cases, dispersals involved capture site to settlement site dis- tances >2.2 km, the stream distance home range of Beavers in Manitoba (Novakowski 1965). Interpretations of dispersal movements were neces- sarily subjective. Ours were based on a knowledge of colony locations in the study areas and the assumption that subadult Beavers trapped in a colony were pre-dispersal members of that colony. A two-tailed Wilcoxon Rank Sum test (Johnson and Bhattacharyya 1987) was used to test for sex- biased dispersal distance, and to test if the distribu- tions of median, maximum upstream, and maximum downstream movements differed between 1989 and 1990. Simple linear regression (Sokal and Rohlf 1995) was used to determine if dispersal dates and dispersal-settlement interval lengths were related to dispersal distances. A G-test (Sokal and Rohlf 1995) was used to test if dispersal proabability was indepen- dent of sex. We used the KAPLAN computer program (copyright 1988, Missouri Department of Conser- vation, Columbia, Missouri) for survival analysis. It is based on Pollock et al.’s (1989) modification of the Kaplan-Meier estimator for the staggered entry of radio-marked animals. Samples from 1989 and 1990 were pooled for survival analysis because of small sample size. Survival was estimated from the Beavers’ release in the spring until 6 April of the fol- lowing year. The survival estimate is roughly equiva- lent to the life table statistic p, (Caughley 1977). Results We radio-equipped eight Beavers in 1989 and 14 in 1990. There was no mortality due to the surgical procedures. The Beavers from 1989 were located VAN DEELEN AND PLETSCHER: DISPERSAL OF BEAVERS SY) 279 times and 46 movements > 2.5 km from the release site were detected. We obtained 1299 loca- tions for the Beavers from 1990 and detected 468 movements > 2.5 km from the release site. The mean maximum upstream movement was 3.0 km (SD = 3.5, n = 8, range = 0.2—5.5 km) in 1989 and 4.7 km (SD = 4.7, n = 14, range = 0.1 - 15.1 km) in 1990. The mean maximum downstream movement was 7.0 km (SD = 13.7, n = 8, range = 0.5 - 40.6 km) in 1989, and 4.7 km (SD = 5.0, n = 14, range = 0.1 - 15.7 km) in 1990. Average median movements were 2 km! (SDe=)2 1h 8, rane =; 028-9529) kim) ran 1989, and 1.2 km (SD = 1.0, n = 14, range = 0.1 - 3.7 km) in 1990. There were no differences in the distributions of median movement distances (Z = - 0.61, P = 0.54), maximum upstream distances (Z = 1.00, P = 0.32), or maximum downstream distances (Z = 0.55, P = 0.58) for Beavers between 1989 and 1990. We saw no evidence of over-land travel. Three of the Beavers from 1990 were probably captured outside of their natal colonies since they were captured and released at sites that were not near known active colonies. Two of these three had fresh puncture wounds on the hip and tail areas which are indicative of intra-specific fighting (Novak 1987). By contrast, none of the juvenile Beavers caught in active colonies had similar wounds. Movements for the Beavers caught outside of their natal colony were analyzed as if the capture/release site were the natal colony. One of these Beavers settled and was classi- fied as a non-disperser. The other two continued moving and were classified as dispersers. Four of the dispersing Beavers settled within 16 days. The remaining six dispersers settled between 35 and 181 days. Twelve Beavers (five in 1989, seven in 1990) did not disperse. Three of the non- dispersers and one of the dispersers made explorato- ry movements to resting sites > 2.5 km from their natal colonies and then returned. Mean dispersal date (17 May) was highly variable (n = 10, SD = 44 days, range: 7 April - 20 August), as was mean settlement date (24 July, n= 10, SD = 86 days, range: 9 April - 12 November), and the length of the dispersal-settlement interval (x = 68 days, n = 10, S.D. = 69, range: 2— 181 days). Dispersal distance was highly variable (x = 7.7, n = 10, SD = 5.9 km, range = 2.9— 22.2 km), and not cor- related with dispersal date (r = 0.29, P = 0.38), or the length of the dispersal-settlement interval (r = 0.16, P0576): The sex ratio of the 10 dispersing Beavers was 6 males : 3 females : 1 unknown. The probability of dispersal was independent of sex (G = 0.52, P = 0.47) as was dispersal distance (Z = 0.36, P = 0.64). Of the eight Beavers from 1989, two (one dispers- er, and one non-disperser) died during the summer of unknown causes and one (a disperser) died during the winter of unknown causes. One (a non-disperser) was censored (censorships indicate loss of radio sig- 320 nals despite extensive aerial searches). The sample from 1990 had four mortalities and one censorship. Two (non-dispersers) were removed by a commer- cial trapper during the trapping season in 1990-1991, one (a non-disperser) was probably killed by a Black Bear (Ursus americanus), and one died (a disperser) during the winter of unknown causes. The estimated annual survival rate for the pooled samples of two-year-old Beavers was 0.67 (n = 22, SE= 0.11). For dispersers it was 0.70 (n = 10, SE = 0.14) and for non-dispersers it was 0.64 (@=—25SE (0116): Discussion Dispersal by two-year-old Beavers generally coin- cides with the birth of the colony’s kits, spring runoff, and high water (Svendsen 1980; Hodgdon and Lancia 1983). Dispersal dates were variable in Montana Beavers, and generally coincided with the 10 April - 29 June high water period in 1990 (Van Deelen 1991). Idaho Beavers in montane habitat dispersed in both upstream and downstream directions (Leege 1968) as did Beavers in this study. Michigan Beavers dispersed over land (Bradt 1938). In the rugged terrain of the western Montana, Beavers are more restricted to travel along the stream courses (Allred 1981), although Leege (1968) reported that one Idaho Beaver crossed the divide between two adjacent drainages. Dispersal distances in this study varied, but were comparable to the mean airline distance of 5.6 km reported for Idaho Beavers (Leege 1968). At least three non-dispersers and one disperser exhibited large (>2.5 km) exploratory movements away from the natal colony. Nocturnal locations might have enabled us to identify more explorations. These might provide potential dispersers with a method of assessing local colony density by encountering scent mounds (Aleksiuk 1968) and other Beavers (Aleksiuk 1968; Bergerud and Miller 1977; Allred 1981). High density might cause potential dispersers to delay dispersal or return to the natal colony per- manently (Molini et al. 1980; Payne 1982). Twelve of the 22 radio-equipped two-year-olds in this study did not disperse. This is consistent with frequent reports of non-breeding adults in Beaver colonies, and might indicate a high population densi- ty relative to the habitat’s carrying capacity (Novakowski 1965; Gunson 1970; Payne 1982; Busher et al. 1983; Hodgdon and Lancia 1983). Alternatively, Svendsen (1980) reported 100% dis- persal for two-year-old Beavers in a high density population that had no predators and was protected from trapping by being in a state park. We believed that Beaver densities in the study areas were relative- ly high during the course of the study because fur prices were low and trapping was light. THE CANADIAN FIELD-NATURALIST Late summer settlement and variation in settle- ment times among dispersers (this study) results in a summer sub-population of transient juvenile Beavers (Townsend 1953; Aleksiuk 1968; Molini et al. 1980) and is probably linked to pair formation. Svendson (1988) found that pair formation among dispersing Ohio Beavers occurred throughout year but peaked during September, October, and November as tran- sients either paired with other transients that recently Vol. 110: | 4 | a settled at suitable sites, or were incorporated into existing family groups where the same-sex adult was | missing. Acknowledgments This study was funded by the Montana | Department of Fish Wildlife and Parks through the Montana Cooperative Wildlife Research Unit (MCWRU), the Lolo and Deerlodge National Forests, and the Montana Chapter of The Wildlife Society. We thank H. T. Harris field assistance, H. S. Hash of the Montana Department of Fish Wildlife and Parks for safe flying and logistical support, and the MCWRU staff. Literature Cited Aleksiuk, M. 1968. Scent-mound communication, territo- riality, and population regulation in the Beaver (Castor canadensis Kuhl). Journal of Mammalogy 49: 759-762. Allred, M. 1981. The potential use of Beaver population behavior in Beaver resource management. Journal of the Idaho Academy of Sciences 17: 14—24. Bergerud, A. T., and D. R. Miller. 1977. Population dynamics of Newfoundland Beaver. Canadian Journal of Zoology 55: 1480-1492. Bradt, G. W. 1938. A study of Beaver colonies in Michigan. Journal of Mammalogy 19: 139-162. Brooks, R. P., M. W. Fleming, and J. J. Kennelly. 1980. Beaver colony response to fertility control: evaluating a concept. The Journal of Wildlife Management 44: 568-575. Busher, P. E., R. J. Warner, and S. H. Jenkins. 1983. Population density, colony composition, and local move- ments in two Sierra Nevadan Beaver populations. Journal of Mammalogy 64: 314-318. Caughley, G. 1977. Analysis of vertebrate populations. John Wiley, New York. 234 pages. Gunson, J. R. 1970. Dynamics of the Beaver of Saskatchewan’s northern forest. M.S. thesis. University of Alberta, Edmonton. 122 pages. Hodgdon, H. E., and R. A. Lancia. 1983. Behavior of the North American Beaver Castor canadensis. Acta Zoologica Fennica 174: 99-103. Jackson, M.D. 1991. Beaver dispersal in Western Montana. M.S. thesis. University of Montana, Missoula. 77 pages. Johnson, R., and G. Bhattacharyya. 1987. Statistics principles and methods. John Wiley, New York. 578 pages. Leege, T. A. 1968. Natural movements of Beaver in southeastern Idaho. The Journal of Wildlife Management 32: 973-976. Molini, J. J., R. A. Lancia, J. Bisher, H. E. Hodgdon. 1980. A stochastic model of Beaver population growth. 1996 Pages 1215-1245 in Worldwide furbearer conference proceedings. Edited by J. A. Chapman, and D. Pursey. Frostburg, Maryland. Novak, M. 1987. Beaver. Pages 283-312 in Wild fur- bearer management and conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. Ontario Ministry of Natural Resources, | Toronto, Ontario. | Novakowski, N.S. 1965. Population dynamics of a Beaver population in northern latitudes. Ph.D. thesis, __ University of Saskatchewan, Saskatoon. 155 pages. Patric, E. F., and W. L. Webb. 1960. An evaluation of three age determination criteria in live Beavers. The Journal of Wildlife Management 24: 37-44. Payne, N. F. 1979. Relationship of pelt size, weight, and age for Beaver. The Journal of Wildlife Management 43: 804-806. _ Payne, N. F. 1982. Colony size, age, and sex structure of _ Newfoundland Beaver. The Journal of Wildlife Management 46: 655-661. | Payne, N.F. 1984. Mortality rates of Beaver in Newfoundland. The Journal of Wildlife Management 48: 117-126. | Peterson, R. P., and N. F. Payne. 1986. Productivity, size, age, and sex structure of nuisance Beaver colonies in Wisconsin. The Journal of Wildlife Management 50: 265-268. VAN DEELEN AND PLETSCHER: DISPERSAL OF BEAVERS 321 Pollock, K. H., S. R. Winterstien, C. M. Buunk, and P. D. Curtis. 1989. Survival analysis in telemetry stud- ies: the staggered entry design. The Journal of Wildlife Management 53: 7-15. Sokal, R. R., and F. J. Rohlf. 1995. Biometry. Third edi- tion. W. H. Freeman and Co.. New York. 887 pages. Svendsen, G. E. 1980. Population parameters and colony composition of Beaver (Castor canadensis) in southeast Ohio. American Midland Naturalist 104: 47-56. Svendsen, G. E. 1989. Pair formation, duration of pair bonds, and mate replacement in a population of Beavers. Canadian Journal of Zoology 67: 336-340. Townsend, J. E. 1953. Beaver ecology in western Montana with special reference to movements. Journal of Mammalogy 34: 459-479. United States Department of the Interior Geological Survey. 1970. The national atlas of the United States of America. Washington, D. C. 417 pages. Van Deelen, T. R. 1991. Dispersal patterns of juvenile Beavers in western Montana. M. S. thesis, University of Montana, Missoula. 91 pages. White, G. C., and R. A. Garrott. 1990. Analysis of wildlife radio-tracking data. Academic Press Inc., San Diego, California. 383 pages. Received 14 August 1995 Accepted 19 December 1995 Microhabitats of Two Peromyscus (Deer and White-footed Mice) Species in Old Fields and Prairies of Wisconsin GAIL E. KANTAK Department of Biology, Saginaw Valley State University, University Center, Michigan 48710 Kantak, Gail E. 1996. Microhabitats of two Peromyscus (Deer and White-footed mice) species in old fields and prairies Ov Wisconsin. Canadian Field-Naturalist 110(2): 322-325. Peromyscus spp. were live-trapped in old fields and prairie remnants of southern Wisconsin. Microhabitat data were al lected at each capture site and at random sites. The Prairie Deer Mouse, P. maniculatus bairdi, was the more abundan’ species, but the Northern White-footed Mouse, P. leucopus noveboracensis, also maintained resident, though smaller, pop: ulations in these open field habitats. Compared to random trap stations, trap stations capturing Peromyscus spp. had more bare substrate, less dead vegetation, taller vegetation, lower forb diversity, and were closer to cultivated fields. Comparec, to P. maniculatus, P. leucopus selected microhabitats which had more small-scale three-dimensional structure and whict | were closer to isolated trees. ! Key Words: Prairie Deer Mouse, Peromyscus maniculatus bairdi, Northern White-footed Mouse, Peromyscus leucopus noveboracensis, microhabitat, old fields, prairies, Wisconsin. The Prairie Deer Mouse, Peromyscus maniculatus bairdi, is characteristic of old field and prairie habi- tats, whereas the Northern White-footed Mouse Peromyscus leucopus noveboracensis is typically a woodland inhabitant (Blair 1940; Harris 1952; Madison 1977). Although densities of P. leucopus noveboracensis are highest in wooded habitat, this subspecies regularly occurs in a variety of old field successional stages (Beckwith 1954; Getz 1961; Hirth 1959; Jackson 1961; Lackey 1978; Root and Pearson 1964; Verts 1957). Both of these mice main- tain resident populations in old fields and prairies of southern Wisconsin (Kantak 1981; Stromberg 1979a) and have comparable diets (Cogshall 1928; Drickamer 1970; Whitaker 1966). This poses a ques- tion as to how these ecologically similar rodents are able to coexist in these habitats. In species that share habitats and diets, differences in microhabitats may be ecologically significant (Kantak 1983). Therefore, the objective of this study was to examine structural and vegetational characteristics of the microhabitats of P. maniculatus bairdi and P. leucopus novebora- censis in old fields and prairies in southern Wisconsin to determine if differences at this spatial scale could account for the coexistence of these species. Study Area and Methods This study was conducted on 10 sites in south- central Wisconsin. This is the “sand prairie” region of the Wisconsin River Valley, a term which acknowledges both the sandy nature of the substrate and the dry prairie vegetation found there (Curtis 1959). Ten study sites at four geographic localities were chosen to include a range of successional stages from recently disturbed fields to undisturbed a22 prairie. Geographic coordinates of these locales are: Peetz Prairie, Sauk County, 1.5 miles west of Prairie Du Sac at 43°17'16” N, 89°47'23” W; Schluckebier: Prairie, Sauk County, 1.5 miles west of Prairie Du Sac at 43°17'30" N, 89°47'30" W; Spring Green) Preserve, Sauk County, | mile north of Spring Green at 43°12'06” N, 90°02'57” W; Arena area, Iowa County, 0.5 mile north and 0.5 mile west of Arena at. 43°10'38" N, 89°54'46" W Prairie sites were dominated by Big Bluestem (Andropogon gerardi), Little Bluestem (A. scoparius) and Indian Grass (Sorghastrum nutans). The most recently disturbed sites (1 — 3 years) were dominated by Sandbur (Cenchrus longispinus), Foxtail (Setaria glauca) and Quackgrass (Agropyron’ repens). Older disturbed sites (5 — 15 years) had increasing representation of prairie species and dom-, inant species such as Junegrass (Koeleria cristata), Fall Witchgrass (Leptoloma cognatum) and Canada Bluegrass (Poa compressa). On each site, 50 Sherman live-traps (5 by 6.4 byt 16.5 cm) were set in a5 X 10 grid pattern with! 10-m intervals. In each of the four localities, live-| traps were also placed along a 100-m transect with) 10-m trap intervals in a windbreak (row of trees) or! woodland adjacent to the grassy habitat containing: the grids. All traps were baited with mixed bird seed. ) Grids were trapped at 7-10 day intervals during: August-October 1978 and May-October 1979; tran-/ sects were trapped during June-October 1979. Trap) stations were not permanently located and_varied somewhat from one trapping session to the next to } maintain independence of captures. Peromyscus were identified by using a field dis- | criminant function developed by Stromberg (1979b) | for mice collected from the study area. At each trap, y 1996 station where Peromyscus was caught the percent cover of the following variables was measured in 1m? quadrats: bare area, dead vegetation, lichens or moss, Selaginella spp., corn stalk remnants, sedges, Opuntia sp., shrubs, grasses, and forbs. Cover esti- mates of individual grass and forb species were used to calculate grass species diversity and forb species diversity (Shannon index, Magurran 1988). Also measured at each trap station were maximum height of vegetation, distance to the nearest tree within 200- m, distance to the nearest windbreak or woods with- in 200-m, and distance to the nearest cultivated field within 200-m. These microhabitat variables were also measured once a month from June-September 1979 at 20 random trap stations at each grid. Recapture rates of Peromyscus were compared with a G-test (Sokal and Rohlf 1981) on the number of individuals and captures of each Peromyscus species in the grids. Discriminant analysis (SPSS 1988) was used to distinguish microhabitats among the three groups- P. maniculatus, P. leucopus, and random trap stations. The direct entry method of variable selection was used because of statistical advantages over stepwise procedures (Habbema and Hermans 1977; James and McCulloch 1990). Rank- transformed discriminant analysis was applied to the data because it permitted use of the F- test and other parametric statistical tests when assumptions of nor- mality and homogeneous covariance matrices were not met by the raw data (Conover and Iman 1980, 1981; Potvin and Roff 1993). Results A sampling effort of 10100 trapnights within field and prairie sites yielded 164 captures of P. maniculatus, 44 captures of P. leucopus and 20 cap- tures of juvenile Peromyscus spp. which could not be identified to species and were not included in the analysis. Recapture rates of P. leucopus and P. man- iculatus did not differ (G = 0.554, p > 0.10). Sampling in the transects in adjacent windbreaks or woody habitats resulted in three captures of P. man- iculatus and 70 captures of P. leucopus in 1260 trap- nights of sampling effort. Two significant functions of microhabitat data were identified by discriminant analysis (Table 1). The first function explained 87% of the variance and separated stations of both species of Peromyscus from random stations. Significant variables separat- ing scores for Peromyscus stations from random sta- tions included bare area, low ground cover (lichens and moss), vegetation height and distance to wind- breaks. Variables contributing heavily to the dis- criminant scores of random stations included dead vegetation cover and forb diversity. The second dis- criminant function explained 13% of the variance and separated P. maniculatus stations from P. leuco- pus stations. Peromyscus maniculatus scores indicat- ed that bare area, low or dead vegetation cover and KANTAK: Two PEROMYSCUS IN OLD FIELDS AND PRAIRIES 323 TABLE 1. Summary of discriminant function analysis com- paring P. maniculatus bairdi, P. leucopus noveboracensis, and random microhabitats: significance, coefficients, and classification. Discriminant Functions #1 #2 Eigenvalue 0.29826 0.04461 % Variance 86.99 13.01 Chi-square Statistic 303.6 43.495 Degrees of Freedom 32 15 Significance p< 0.0000 p< 0.0001 Discriminant Analysis Results Standardized DF Coefficient Variable #1 #2 Bare Area Cover 0.54940 0.65866 Dead Vegetation Cover -0.43315 0.42621 Lichens and Moss Cover 0.37455 0.21999 Opuntia sp. Cover -0.05137 -0.41628 Shrub Cover -0.03533 -0.30405 Forb Diversity -0.32580 -0.17023 Vegetation Height 0.46152 -0.24062 Tree Distance -0.27748 0.32626 Windbreak Distance 0.31172 -0.08488 Cultivated Field Distance -0.05656 -0.43759 Group Means of Discriminant Functions Function 1 Function 2 P. maniculatus 1.05065 0.25204 P. leucopus 1.13431 -0.88378 random stations -0.27812 -0.00306 Classification Matrix Predicted Group Membership (%) Actual Group n Pm.b. Pin. random P. maniculatus 164 56.7 22.0 21.3 P. leucopus 44 295 59.1 11.4 random stations 799 18.9 12.0 69.1 Total cases correctly classified: 66.6% distance to trees were important contributors, where- as variables contributing most to P. leucopus scores included amount of three-dimensional cover such as of shrubs and Opuntia sp., vegetation height, and distance to cultivated fields. Discussion Despite large heterogeneity in microhabitat fea- tures across the range of successional stages studied, discriminant analysis identified several common fea- tures which classified Peromyscus capture stations. The two species of Peromyscus were not intermin- gled randomly across these habitats but each selected microhabitats associated with key habitat features involving amount of cover, three-dimensional struc- ture, and distance to trees or cultivated fields. However, the variables incorporated into these dis- criminant functions might not be those to which Peromyscus are actually responding but could sim- ply be correlated to unmeasured parameters. 324 Therefore, the ecological importance of significant variables must be considered. The first discriminant function determined that most of the variability in the data set occurred between random stations and stations capturing either species of Peromyscus. The standardized dis- criminant function coefficients show that Peromyscus trap stations had taller living vegetation and less dead vegetation than random trap stations, perhaps related to food supply or quality of cover with respect to risk of predation. Peromyscus sta- tions also had more bare substrate and low growth like lichens and moss, which might be related to for- aging efficiency for seeds on the ground. Lesser forb diversity relative to random stations may reflect selection by Peromyscus spp. for microhabitats with more monotypic cover of preferred plant species. The second discriminant function showed that field- caught P. maniculatus and P. leucopus were not co- existing in the same microhabitats. Peromyscus leu- copus selected microhabitats with features character- istic of their more typical woodland habitat, that is, with more three-dimensional structure (shrub and Opuntia cover). Also suggestive of wooded habitat were the taller vegetation and closer proximity of trees. Peromyscus maniculatus appeared to be more of a grassland generalist whose capture sites were best characterized by what they lacked- heavy vege- tative cover and trees. Peromyscus leucopus individuals in my study sites were not all transients or migrants between wooded windbreaks and fields because recapture rates were as high as those of the resident Prairie Deer Mouse. Also, no P. leucopus individual was caught in the windbreaks once it had already been captured in the fields. However, the converse did occur- three P. leucopus individuals captured in the windbreaks early in the summer were later recaptured in the fields. This suggests that emigration from wind- breaks to fields may occur. Because of higher cap- ture rates in windbreaks compared to fields, one might hypothesize that population pressure underlies this movement to grassy habitats. In my study sites, P. leucopus is a microhabitat specialist, concentrat- ing its activities in the most structurally diverse areas. It may be able to appropriate these microhabi- tat patches from resident P. maniculatus because of its larger body size (Jackson 1961; Stromberg 1979a) and dominant and aggressive behavior (Fitch 1963; Kantak 1983). This is supported by field experiments in which the persistence of P. manicula- tus populations was negatively correlated with the density of P. leucopus, and removal of one species was followed by an increase in the other (Master 1977). Given its generalist diet and competitive advantage over P. maniculatus, why P. leucopus is not more successful in grassy habitats is unclear. Differential predation was suggested but could not THE CANADIAN FIELD-NATURALIST Vol. 110 be demonstrated in field experiments by Stromberg (1979a). Microhabitats should be selected not only to reduce competition and predation but also to provide appro- priate shelter and microclimate, nest sites, and food. Hence a single factor explanation is not likely to total- ly explain the complex phenomenon we call micro- habitat selection. Nevertheless, if the question is how sympatric species with similar diets coexist, examina- tion of the microhabitat should be the first step. Differences in microhabitats reflect the collective effects of a number of selective forces, and further research must sort out their relative contributions. Acknowledgments I thank the following people for assistance during the planning phase and field work of this study: E.W. Beals, T.C. Moermond, D. Rogers, M. Stromberg and G. Stephenson. The Nature Conservancy and O. Peetz allowed use of study sites. Field work was supported by the University of Wisconsin, and manuscript preparation was support- ed by Saginaw Valley State University, Michigan. 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The effect on interspecific competi- tion on habitat utilization by two species of Peromyscus. Ph.D. thesis, University of Michigan, Ann Arbor. 179 pages. KANTAK: Two PEROMYSCUS IN OLD FIELDS AND PRAIRIES 325) Potvin, C., and D. A. Roff. 1993. Distribution-free and robust statistical methods: viable alternatives to paramet- ric statistics? Ecology 74: 1617-1628. Root, P. G., and P. G. Pearson. 1964. Small mammals in the early stages of old field succession on the New Jersey piedmont. Bulletin of the New Jersey Academic Society 9: 21-26. Sokal, R. R., and F. J. Rohlf. 1981. Biometry. W. H. Freeman and Company, New York. SPSS, Inc. 1988. SPSS-X User’s Guide, 3rd edition. SPSS Inc., Chicago, Illinois. Stromberg, M. R. 1979a. Experimental analysis of habi- tat performance and direct observation of deermice (Peromyscus) in southern Wisconsin. Ph.D. thesis, University of Wisconsin, Madison. 224 pages. Stromberg, M. R. 1979b. Field identification of Pero- myscus leucopus and P. maniculatus with discriminant analysis. Transactions of the Wisconsin Academy of Science, Arts and Letters 67: 159-164. Verts, B. J. 1957. The population and distribution of two species of Peromyscus on some Illinois strip-mined land. Journal of Mammalogy 38: 53-59. Whitaker, J. O., Jr. 1966. Food of Mus musculus, Pero- myscus maniculatus bairdi and Peromyscus leucopus in Vigo County, Indiana. Journal of Mammalogy 47: 473-486. Received 29 August 1995 Accepted 9 January 1996 Peatlands: A New Habitat for the Upland Sandpiper, Bartramia longicauda, in Eastern Canada SOPHIE CALME and STEPHANIE HADDAD Centre de recherche en biologie foresti¢re, Université Laval, Sainte-Foy, Québec G1K 7P4 Calmé, Sophie, and Stéphanie Haddad. 1996. Peatlands: A new habitat for the Upland Sandpiper, Bartramia longicauda, | in eastern Canada. Canadian Field-Naturalist 110(2): 326-330. Although there have been reports of Upland Sandpipers breeding in peatlands of northwestern Canada, our study is the first to demonstrate widespread use of peatlands in other parts of the species’ range. In the summer of 1995, while conducting a study on the breeding birds of Quebec’s peatlands, we recorded a total of 11 pairs of Upland Sandpipers, in five peatlands | in a small region on the south shore of the St. Lawrence River. In the previous year, four pairs had been observed in two of the same peatlands. These peatlands were the largest in the region (160 ha to 496 ha), and consisted mainly of open treeless habitat. The mean density of breeding pairs in 1995 was 0.59 per 100 ha. The availability of large open areas, a low preda- tion rate, and infrequent human disturbance indicate that peatlands could constitute a suitable habitat for the species. Bien que des mentions de nidification de maubéche des champs aient été rapportées dans les tourbiéres du nord-ouest cana- dien, la présente étude démontre pour la premiere fois l'utilisation de cet habitat par l’espéce dans |’est de son aire de dis- tribution. Au cours de |’été 1995, lors d’une étude sur les oiseaux nicheurs des tourbiéres du Québec, 11 couples de Maubéche des champs, ont été observés dans cing tourbiéres localisées dans une petite région de la rive sud du fleuve Saint-Laurent. L’année précédente, quatre couples avaient été observés dans deux des mémes tourbieres. Ces tourbiéres, les plus grandes de la région (160 ha a 496 ha), étaient principalement constituées d’ habitat ouvert dépourvu d’ arbres. La densité moyenne de couples nicheurs était de 0.59 pour 100 ha en 1995. La disponibilité de grandes surfaces ouvertes, un faible taux de prédation, ainsi qu’un niveau de perturbation humaine presque inexistant pourraient faire des tourbiéres un habitat convenable pour I’espéce. Key Words: Upland Sandpiper, Bartramia longicauda, peatland, grassland, open habitat, Québec. The Upland Sandpiper (Bartramia longicauda) is a species of the American grasslands, scarcely and locally distributed in much of its range (Peterson 1994). It usually inhabits grassland pastures, tall- grass prairies, blueberry barrens, airport runways, and recently burned or mowed areas (Askins 1992). In the Northwest, Upland Sandpipers are occasional- ly observed in clearings in spruce muskeg (Palmer 1967), on montane grasslands (Gabrielson and Lincoln 1959), and even in peatlands (Godfrey 1986). Like many North American game species, Upland Sandpipers experienced near-extinction at the turn of the century. This decline was halted around 1920, with the help of the Migratory Bird Convention, (Osborne and Peterson 1984), as no open season has since been declared for this species. In spite of a period of recovery marked by the apparent westerly expansion of the species’ range, the number of Upland Sandpipers began to decline again in almost every northeastern and central region. For instance, its numbers in Illinois decreased by approximately 95% between 1956-1958 and 1978-1979 (Ano- nymous, cited in Askins 1992). In the New England states, the species became uncommon to rare, and it *See Documents Cited section before Literature Cited. is still a species of special concern in many of these states (Vickery 1992). Moreover, the Upland Sandpiper was on the Blue List from 1975 to 1986 (Tates 1986), and is one of the species classified as threatened on its breeding grounds (Thompson et al. 1992). However, the results from the Breeding Bird Survey (BBS) show a positive trend of the species population in its entire area of distribution between 1966 and 1994 with a mean percent of annual change of 2.0% (p < 0.001). Most of this increase seems to have taken place before 1980, as the recent BBS trend (1980-1994) indicated a rate of annual change of 0.2% (non significant). In Québec, for the same period, this mean percent of annual change reached -3.0%, though this value was non-significant (Sauer et al. 1995*). A recent summary on shore- birds based on documented data and surveys indi- cates a possible general decline of the Upland Sandpiper throughout Canada with the current popu- lation estimated at 2000 individuals (Morrison et al. 1994). Therefore, while some studies show encour- aging signs for the species, the overall status of the Upland Sandpiper warrants further attention. In this paper we report on recent observations of Upland Sandpipers in peatlands. We discuss these findings in relation to previous knowledge of the species, while emphasizing their importance in regards to Upland Sandpiper’s present status. 326 1996 Methods Study sites This study was conducted in 1994 and 1995, between 5 June and 14 July. In 1994, we visited 137 peatlands, roughly distributed along the Saint Lawrence River lowlands, between the Québec/Vermont border and Havre-Saint-Pierre on the North Shore. In 1995, 72 peatlands were sampled south and west of Quebec City (Figure 1). In 1995, we also counted birds in open habitats, 1.e., mostly hayfields (41) and pastures (10), abandoned farm- lands (7), recent clearcuts (4), grain crops (6), fields of clover (3), food crops (2), cranberry farms (1), and young plantations (1). Sampling methods In 1994 the fixed-radius point-count was used to measure species diversity. One 10-min observation period was carried out in each of the 137 peatlands, where all birds seen or heard within a radius of 100 m were recorded. When peatlands were large enough, a second point count was conducted at a dis- tance of at least | km from the first point count. Plots were located at least 150 m from the peatland’s CALME AND HADDAD: HABITAT FOR THE UPLAND SANDPIPER Sil edge. Vegetation was sampled in each circular plot by visually estimating the percentage of the different vegetation types, as well as the percentage of open water. Vegetation types consisted of the following strata: forbs, ericaceous shrubs, trees <2 m, trees between 2 and 5 m, and trees > 5 m. In 1995 the point-count technique was again used. We placed one count in each distinct habitat type in the peatland; thus their number depended on the peatland’s heterogeneity. In the surrounding open habitats we used single point-counts. In the peatlands, birds were also sampled along 200-m wide transect strips. We recorded the positions of all birds seen or heard in the strips. All transect lines were situated at least 200 m from the peatland edge and 300 m from each other. The total area sampled in each peatland represented a minimum of 30% and a maximum of 67% of its surface. Each peatland was surveyed once during the breeding season. Like many other shorebirds, Upland Sandpipers are mostly silent during incubation. However, we considered that failure to detect breeding Upland Sandpipers was unlikely since our surveys started around the first week of June. By Quebec City and vicinity FiGuRE 1. Study area in 1995. The peatlands with Upland Sandpipers are indicated by solid triangles (4), while solid cir- cles (@) indicate other peatlands sampled, and the empty triangle (A) represents the field with three observed pairs. Numbers correspond to the locatiosn on Table 1. On the inserted map, the northern limit of distribution of Upland Sandpiper in Québec is indicated by the dotted line (...), and the study area is represented by the small rectangle. 328 THE CANADIAN FIE this date, the main incubation period is completed and chicks begin to leave the nest in southern Québec (Yank and Breton 1995). To determine whether the area of the peatland had a possible influence on the presence of Upland Sandpipers, the peatlands sampled in 1995 were divided into two groups, with and without Upland Sandpipers (n = 5 and n = 27, respectively). We assume that the peatlands without Upland Sandpipers used for the analysis, had similar open habitat, based on the presence of the Savannah Sparrow (Passerculus sandwichensis), a species characteristic of open habitats, the habitat structure, and the interpretation of aerial photographs. We digitized 1: 15 000 aerial photographs of the peat- lands to measure peatland area. We used a two- tailed t-test to compare the two groups of peatlands. Results In 1994, four pairs were observed in two separate but neighboring peatlands (Table 1 and Figure 1). In 1995, 11 pairs were recorded in five peatlands (Table 1 and Figure 1), and three others in a hayfield (Figure 1). The peatlands with Upland Sandpipers were situated in a small region characterized by a mixture of lands dedicated to agriculture and forestry. Typically, these peatlands were the largest in the surrounding region, and they consisted mainly of open habitat. The vegetation was dominated either by ericaceous shrubs such as Ledum groenlandicum and Kalmia angustifolia or forbs such as Carex sp. Shrubs were scarce and usually found on peatland edges known as fen lags. Trees, mainly Larch (Larix laricina) and/or Black Spruce (Picea mariana), were rather rare and scattered: the overall tree cover rarely exceeded 15% (Table 1). The results from the Student t-test show that the two groups of peatlands were significantly different (t = 4.999, p < 0.0001). The mean areas of peatlands used by the Upland Sandpipers was 375 + 134 ha (n = 5) compared to 122 + 98 ha (n = 27) for unused peatlands in the same region. The birds were usually heard whistling before they became aware of us; as we approached their ter- ritories the sandpipers were visibly disturbed by our LD-NATURALIST Vol. 110 FiGure 2. An Upland Sandpiper perched on a spruce, its wings semi-erect in an unstable position. presence (Figure 2). They flew around us, calling repeatedly. These alarm calls often attracted other individuals, up to five. In peatland 4 (Figure 1), on 14 June, we also observed a chick, while an adult was performing a distraction display. Discussion In 1995, the mean population density in the peat- lands was 0.59 pairs/100 ha. This latter value could be higher if we considered only the type of habitat used by the Upland Sandpipers, thus eliminating “islands” of denser vegetation. To our knowledge, no other data are available concerning population densities in Québec. Despite the small sample size, the population density in our study was lower than the 1.1 nest/100 ha found for Upland Sandpiper in a twenty-year study conducted in Illinois (Buhner- kempe and Westemeier 1988). According to TABLE |. Sites of observations of Upland Sandpipers in peatlands in Quebec, and number of territories found in 1994 and 1995. Area Location (ha) 46°04'N;72°06’W 160 46°19'N;72°11'W = 338 46°19'N;72°49'W ss 4418 46°23'N;72°53’W 496 46°22'N;72°35’W = 463 Pa, 2 Locality Saint-Valére Sainte-Marie-de-Blandford Notre-Dame-de-Lourdes Villeroy Sainte-Anastasie ABWN Vegetation cover in observation sites Number of (%) territories eric. forb >5m 2-5m <2m 1994 1995 90 2 0 3 2 - 1 35 65 5 1 1 - 70 15 5 5 1 2 4 15 40 3 10 8 2 4 7 80 0 0 1 - 1 1996 Osborne and Peterson (1984), an ideal habitat could support up to eight pairs/100 ha. That value was sup- ported by a study in the Wisconsin prairies in which two Upland Sandpiper pairs had territories of 8.1 and 12.1 ha (DeGraaf and Rudis 1987). The results show that Upland Sandpipers pre- ferred large peatlands. The smallest peatland in which an individual was observed (160 ha) was of intermediate size, but the four other peatlands were the largest ones in the region studied in 1995. Area- sensitivity by Upland Sandpiper is not limited to peatlands: in Missouri and Illinois, Upland Sandpipers were not found in grassland habitat of less than 10 ha and 30 ha, respectively (Samson 1980; Herkert 1991). Closer to eastern Canada, Vickery et al. (1993) found a relationship between the presence of Upland Sandpipers and the area of grasslands in eastern Maine. The species was very rare in patches of habitat smaller than 50 ha, and reached an incidence of 50% in grasslands of more than 200 ha. As with many other grassland specialists (Askins 1992), the continued loss and fragmentation of habi- tat may be the major cause of the Upland Sandpiper decline. On breeding grounds, urban and surburban development, afforestation on abandoned farmlands (see Askins 1992), as well as changes in agricultural practices, contribute to the loss of suitable nesting habitat. In Québec, the disappearance of traditional family-owned farms, and the extensive cultivation of corn have both been related to the decline of the species (Yank and Breton 1995). Threats to the species are not restricted to its breeding grounds; the Upland Sandpiper also faces an important reduction of its main wintering habitat, the South American pampas. Like the North American Great Plains, the Argentinian and Urugayan pampas have been replaced by extensive crops of corn and wheat. The region’s natural ecosystem has become so scarce that the outlook for its avifauna seems even more pressing than the situa- tion of tropical forest species (Terborgh 1989). The presence of a breeding bird may be a good indication of the suitability of a site. However, breeding success is a guarantee of its quality (Van Horne 1983) and the most important obstacle to breeding success is often nest predation (Martin 1988). Here, we experimentally measured nest pre- dation with artificial nests in six large and relative- ly open peatlands of the region previously described, including Villeroy, Sainte-Anastasie and Sainte-Marie-de-Blandford. The apparent predation rate on nests averaged 12% (n = 114), with no obvious effect of distance from edge (unpublished data). This value is similar to results found by Burger at al. (1994). In their study, the rate of pre- dation on artificial nests averaged 16.2% (n = 216) for six prairie fragments larger than 130 ha in CALME AND HADDAD: HABITAT FOR THE UPLAND SANDPIPER 329 Missouri. Our results thus suggest that peatland habitat is as suitable as grassland habitat for breed- ing Upland Sandpipers in Québec. Two hypotheses may explain the lack of previous observations of Upland Sandpipers in peatlands. First, until recently, peatlands have been relatively overlooked by ornithologists and this could explain the absence of documented observations of the species in peatlands. Nevertheless, the Upland Sandpiper call and behavior make it readily detectable even by non-ornithologists. A second hypothesis is that the Upland Sandpiper has recently found new sites, along with a new suitable habitat. This plasticity would be highly valuable for the species, given the scarcity of its “traditional” habitat in northeastern America. In peatlands, disturbance by humans or predators seems rare, and there is no risk for the brood to be destroyed by farm machin- ery, as is often the case in fields (Ehrlich et al. 1988). These observations, based on only two years of data, give a limited picture of the situation of the Upland Sandpiper in eastern peatlands, even though geographic coverage was wide. Consequently, they should be considered as a starting point for more investigations on the potential of peatlands for sus- taining Upland Sandpiper populations, and possibly those of other grassland species. Acknowledgments We thank Jacques Ibarzabal, Jean-Francois Rousseau, and Nicolas Wampach for their assistance in the field, and especially J.-F. Rousseau for his original illustration in Figure 2. We also thank André Desrochers, Bruno Drolet, Cheri Gratto- Trevor and Anthony Erskine for their helpful com- ments on this manuscript. S. Calmé was supported by a National Science and Engineering Research Council of Canada doctoral fellowship, and S. Haddad by a Wildlife Habitat Canada Fund. Documents Cited Sauer, J. R., B. G. Peterjohn, S. Orsillo, and J. E. Hines. 1995. The North American Breeding Bird Survey Homepage. Version 95.1 Internet Homepage (Address: http://www.im.nbs.gov/bbs.htpnl). Literature Cited Askins, R. A. 1992. Population trends in grassland, shrubland, and forest birds in Eastern North America. Pages 1-34 in Current Ornithology, Volume 11. Edited by D. M. Power. Plenum Press, New York. Buhnerkempe, J. E., and R. L. Westemeier. 1988. Breeding biology and habitat of Upland Sandpiper on prairie-chicken sanctuaries in Illinois. Transactions, Illinois State Academy of Science 81: 153-162. Burger, L. D., L. W. Burger, and J. Faaborg. 1994. Effects of prairie fragmentation on predation on artificial nests. Journal of Wildlife Management 58: 249-254. DeGraaf, R. M., and D. D. Rudis. 1987. New England wildlife: habitat, natural history, and distribution. United 330 States Department of Agriculture, Forest Service, Northeastern Forest Experiment Station, Broomall, General Technical Report NE-108. Ehrlich, P. R., D. S. Dubkin, and N. D. Wheye. 1988. The Birders Handbook: A Field Guide to the Natural History of North American Birds. Simon and Schuster, Fireside Books. Gabrielson, I. N., and F. C. Lincoln. 1959. The Birds of Alaska. Stockpole Company and Wildlife Management Institute, Harrisburg. Godfrey, W. E. 1986. The Birds of Canada. Revised edi- tion. National Museums of Canada, Ottawa. Herkert, J. R. 1991. An Ecological Study of the Breeding Birds of Grassland Habitat within Illinois, Ph.D. disser- tation, University of Illinois in Urbana-Champaign. Martin, T. E. 1988. Habitat and area effects on forest bird assemblages: is nest predation an influence? Ecology 69: 74-84. Morrison, R. I. G., A. Bourget, R. Butler, H. L. Dixon, C. Gratto-Trevor, P. Hicklin, C. Hyslop, and R. K. Ross. 1994. A preliminary assessment of the status of shorebirds populations in Canada. Canadian Wildlife Service Progress Notes Number 208. Osborne, D. R., and A. T. Peterson. 1984. Decline of the Upland Sandpiper (Bartramia longicauda) in Ohio: an endangered species. Ohio Journal of Science 84: 8-10. Palmer, R. S. 1967. The Shorebirds of North America. Viking Press, New York. Peterson, R. T. 1994. Les oiseaux de l’est de l’ Amérique du Nord. Les guides Peterson, Editions Marcel Broquet, La Prairie. Samson, F. B. 1980. Island biogeography and the conser- vation of prairie birds. Proceedings, North American Prairie Conference 7: 293-305. THE CANADIAN FIELD-NATURALIST Vol. 110 Tate, J., Jr. 1986. The Blue List of 1986. American Birds 40: 227-236. Terborgh, J. 1989. Where Have all the Birds Gone? Princeton University Press, Princeton. Thompson, F. R., S. J. Lewis, J. Green, and D. Ewert. 1992. Status of neotropical migrant landbirds in the Midwest: Identifying species of management concern. Pages 145-158 in Status and Management of Neotropical Migratory Birds. Edited by D. M. Finch and P. W. Strangel. United States Department of Agriculture, Forest Service, Rocky Mountains Forest and Range Experiment Station, Fort Collins. General Technical Report RM-229. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47: 893-901. Vickery, P. D. 1992. A regional analysis of endangered, threatened, and special concern birds in the northeastern United States. Transactions Northeast Section Wildlife Society 48: 1-10. Vickery, P. D., M. L. Hunter, Jr., and S. M. Melvin. 1993. Effects of habitat area on the distribution of grassland birds in Maine. Conservation Biology 8: 1087-1097. Yank, R., and L. Breton. 1995. Maubéche des champs. Pages 482-483 in Les oiseaux nicheurs du Québec : Atlas des oiseaux nicheurs du Québec méridonial. Edited by J. Gauthier and Y. Aubry. Association québé- coise des groupes d’ornithologues, Société québécoise de protection des oiseaux, Service canadien de la faune, Environnement Canada, région du Québec, Montréal. Received 27 September 1995 Accepted 15 December 1995 Notes Northern Pocket Gophers, Thomomys talpoides, with White Pelage from Alberta GILBERT PROULX!, LorI LOUNSBURY2 and HAROLD N. BRYANT? ‘Alpha Wildlife Research & Management Ltd., 9 Garnet Crescent, Sherwood Park, Alberta T8A 2R7 ?County of Red Deer, 4758 - 32 Street, Red Deer, Alberta T4N OM8 3Provincial Museum of Alberta, 12845 - 102 Avenue, Edmonton, Alberta TSN 0M6 Proulx, Gilbert, Lori Lounsbury, and Harold N. Bryant. 1996. Northern Pocket Gophers, Thomomys talpoides, with white pelage from Alberta. Canadian Field-Naturalist 110(2): 331. We report the capture of two Northern Pocket Gophers (Thomomys talpoides) with a complete lack of integumentary pig- ments. Key Words: Alberta, Northern Pocket Gopher, Thomomys talpoides, pelage. The color of the pelage of pocket gophers (Geomyidae) is highly variable (Chase et al. 1982). In Thomomys, it varies from almost black through gray and brown to almost white; the ventral pelage is only slightly paler than the dorsal pelage (Nowak and Paradiso 1983). In Alberta, Northern Pocket Gophers (Thomomys talpoides) typically have a steel-gray dorsal pelage (Smith 1993); individual hairs have a brownish band toward the tip. Reports of true albinism (the complete lack of melanin pig- mentation resulting in white pelage and pink eyes, Searle 1968) in Thomomys are rare. Burnett (1925) described a Northern Pocket Gopher (Thomomys talpoides) from Colorado, with a deep cream pelage, a large white throat-patch, a light tail, and almost white feet and legs. No information was provided about the eye color. Storer and Gregory (1934) reported complete albinism in three pink-eyed indi- viduals of Botta’s Pocket Gopher (Thomomys bottae) from California. In May 1994, farmer D. Fear from Cremona (51° 33’ N, 114° 32’ W), Alberta, captured two adult Northern Pocket Gophers that were completely white from nose to tail. One was a female (Provincial Museum of Alberta accession number 95.2.1; weight 169.8 g; total length 230 mm; tail 60 mm; hindfoot 30 mm; ear 7 mm) and the other was a male (PMA 95.2.2; weight 158.1 g; total length 235 mm; tail 60 mm; hindfoot 30 mm; ear 7 mm). The animals were received frozen but the eye color was not recorded. For this reason, it is uncertain whether these animals were true albinos. Other instances of white gophers have been verbally reported to us by farmers from central Alberta. To our knowledge, these specimens are the first con- firmed cases of Northern Pocket Gophers with a complete lack of integumentary pigments. Acknowledgments We thank Daniel Fear of Cremona, Alberta, for bringing these white pocket gophers to our attention and for providing the specimens. Bill Weimann pre- pared the specimens for the Museum collection. Literature Cited Burnett, W. L. 1925. Dichromatism and albinism in Thomomys talpoides clusius. Journal of Mammalogy 6: 129. Chase, J. D., W. E. Howard, and J. T. Roseberry. 1982. Pocket gophers. Pages 239-255 in Wild mammals of North America. Edited by J. A. Chapman and G. A. Feldhamer. The Johns Hopkins University Press, Baltimore, Maryland. Nowak, R. M., and J. L. Paradiso. 1983. Walker’s mam- mals of the world. Fourth edition. The Johns Hopkins University Press, Baltimore, Maryland. 1362 pages. Searle, A. G. 1968. Comparative genetics of coat colour in mammals. Logos Press, London, United Kingdom. 308 pages. Smith, H. C. 1993. Alberta mammals. An atlas and guide. Provincial Museum of Alberta, Edmonton, Alberta. 238 pages. Storer, T. I., and P. W. Gregory. 1934. Color aberrations in the pocket gopher and their probable genetic explana- tion. Journal of Mammalogy 15: 300-312. Received 16 June 1995 Accepted 17 October 1995 ajo) 332 THE CANADIAN FIELD-NATURALIST Vol. 110 First Record of a Chum Salmon, Oncorhynchus keta, from the Thompson River: Adams River Spawning Grounds, British Columbia D. W. WEtcu! and J. N. TILL” 'Department of Fisheries and Oceans, Pacific Biological Station, Nanaimo, British Columbia V9R 5K6 *Department of Fisheries and Oceans, 555 W. Hastings St., Vancouver, British Columbia V6B 5G3 Welch, D. W., and J. N. Till. 1996. First record of a Chum Salmon, Oncorhynchus keta, from the [hempson River: Adams River spawning grounds, British Columbia. Canadian Field-Naturalist 110(2): 332-334. The first record of a Chum Salmon (Oncorhynchus keta) from the upper reaches of the Fraser River, British Columbia, Canada is based on a carcass of a partly spent mature male found on the lower Adams River spawning grounds, a tributary of the South Thompson River. The record is of interest because it is a major range extension for Chum Salmon within the Fraser River drainage. The migration timing of this fish may indicate that genetic selection has occurred for a run timing significantly earlier than the norm for Fraser River chum, and may suggest that a small population of Chum Salmon is present in the Adams River area. Key Words: Chum Salmon, Oncorhynchus keta, spawning grounds, first record, Fraser River, Thompson River, Adams — River, British Columbia. The Fraser River is a major producer of British Columbia Chum Salmon (Oncorhynchus keta), and the world’s largest single producer of Pacific salmon. Although populations of all the other North American species of Pacific salmon are widely dis- tributed throughout the Fraser watershed, popula- tions of Chum Salmon are almost exclusively restricted to the lower reaches of the river. Only limited records of Chum Salmon upriver of Hope, British Columbia, exist (Figure 1; Lee et al. 1980; Zallen and Farwell 1985; Farwell et al. 1987; FHIP 1990 and 1992); all are scattered observations of small numbers of Chum Salmon in creeks close to the Fraser River mainstem. Only a few hundred Chum Salmon have been recorded upstream of Hope (which lies 263 km from the river mouth at an elevation of 61.m) between 1953 and 1989 (FHIP 1992), with all but three observations lying within 60 km of Hope. The farthest upstream obser- vation is of a total of six Chum Salmon observed in THOMPSON RIVER HELLS GATE HOPE FicurE |. Map of the Fraser River watershed, southwestern British Columbia. 1981-1982 in the Seton River, near Lillooet, British Columbia (Farwell et al. 1987; DFO 1990, 1992; FHIP 1992), about 140 km upstream of Hope at an elevation of 202 m (Palmer 1972). The second far- thest upstream observation is of 23 Chum Salmon in Texas Creek (southeast of Lillooet), 125 km upstream of Hope, at an elevation of 213 m (Farwell et al. 1987). We report here the first evidence for the presence of Chum Salmon in the Thompson River, on the lower Adams River Sockeye Salmon (Oncorhynchus nerka) spawning grounds, approximately 483 km from the river mouth (elevation 350 m). Field Observations We found a male Chum Salmon in an advanced state of decomposition on a gravel bar in the lower reaches of the Adams River spawning grounds on 29 October 1994, approximately 50 m from river entry into Shuswap Lake. Field staff from the Department of Fisheries and Oceans had first noticed the salmon several (perhaps 4 to 5) days earlier at the same location. When initially observed it had been mistaken for an unusually large male Sockeye Salmon. The large size (73.9 cm FL, 57.5 cm postorbital- hypural length), lack of red pigmentation, and pro- nounced kype all suggest that the salmon was not a Sockeye (Figure 2). A scale sample confirmed the tentative identification as a Chum Salmon (Figure 3). Examination of the gonads showed that although partly decomposed, they appeared to be partially spent, and significant abrasion of the lower lobe of the caudal fin was also evident (Figure 2). Discussion Although the possibility of a hoax cannot be com- pletely discounted, it is unlikely that this chum had 333 Sockeye Salmon (below). Note the difference in size, the pronounced kype, dentition, and wear on the lower caudal fin of the Chum Salmon. Not evident in this photograph is the bright red colouration of the Sockeye Salmon; the Chum Salmon is greyish-black with slight dorsal mottling. been carried to the spawning grounds after capture somewhere else. Several patches of white fungus were evident on the skin, a common freshwater infection in spawning salmon near death. If the fish had been caught in salt water, it is unlikely that the colouration would have been as dark as was observed, or that the fungus infection would have been as extensive. The resorption evident on the scale samples also indicates the chum had spent some time in freshwater prior to death. The apparent extension to the distribution of Chum Salmon within the Fraser River is unusual as the Adams River spawning grounds are located up a major tributary approximately 270 km past Hope, and 483 km past the mouth of the Fraser. Chum Salmon undertake extensive (> 2000 km) river migrations in a number of large Asian and North American rivers (see review by Salo 1991); but rivers where extensive migration occur generally have relatively shallow gradients compared with the Fraser River. Immediately upstream of Hope the flow of the Fraser River is very swift because of the narrow canyon that the river must flow through. The chum appeared to have died on the spawning grounds around 25 October (or earlier). The median survival time of Fraser River chum tagged on the spawning grounds is 10 days (Palmer 1972). Assuming (because spawning was only partially complete) that the chum arrived on the Adams River spawning grounds about one week prior to death, and that upriver migration took about 18 days (the value for Adams River Sockeye Salmon; Gilhousen 1980), freshwater entry occurred no later than the end of September. This timing is very early compared with most Fraser River chum, which spawn in the lower Fraser from November to January (Beacham and Starr 1982; Salo 1991), and suggests that there may have been some genetic selection for an earlier than nor- mal run timing. Indeed, if the chum was a stray born from some part of the known spawning distribution of Fraser chum, it is very unlikely that the upriver migration rate could be as rapid as that of the direct- ed migration of Adams Sockeye to their spawning grounds, since more time would have to be spent in searching for suitable spawning areas. Combined with the extreme range extension relative to the known distribution of Chum Salmon within the Fraser River, it seems reasonable to speculate that there may be a small population of early-run Chum ferred area of the Chum Salmon. Note the lack of a freshwater growth zone, the globular reticula- tions (A) characteristic of Chum Salmon (Bilton et al. 1964), and the three annuli. In addition, the overall appearance of the scale is characteristic of Chum Salmon, with a bolder, more open appear- ance, and broader shoulders than is evident on the scales of other species (Mosher 1969). The indi- vidual is age 0.4. Salmon spawning on the Adams River, rather than a single male both unusually early in its run timing and far from its birthplace. Acknowledgments Partial funding was received under Grant Number 061-42-2317, from the Canadian Department of Foreign Affairs, Industry, and Trade. THE CANADIAN FIELD-NATURALIST Vol. 110 Literature Cited Beacham, T. D., and P. Starr. 1982. Population biology of Chum Salmon, Oncorhynchus keta, from the Fraser River, British Columbia. Fisheries Bulletin 80: 813-825. Bilton, H. T., D. W. Jenkinson, and M. P. Shepard. 1964. A key to five species of Pacific Salmon (Genus Oncorhynchus) based on scale characters. Journal of the Fisheries Research Board of Canada, 21: 1267-1288. Farwell, M. K., N. D. Schubert, K. H. Wilson, and C. R. Harrison. 1987. Salmon escapements to streams entering statistical areas 28 and 29, 1951 to 1985. Canadian Data Report Fisheries and Aquatic Sciences, Number 601. FHIP. 1990. Fish Habitat Inventory and Information Program. Stream Summary Catalogue. Subdistrict 29E, Chilliwack. Department of Fisheries and Oceans, Vancouver, British Columbia. Unpaginated. FHIP. 1992. Fish Habitat Inventory and Information Program. Stream Summary Catalogue. Subdistrict 29F, Lillooet. Department of Fisheries and Oceans, Vancouver, British Columbia. Unpaginated. Gilhousen, P. 1980. Energy sources and expenditures in Fraser River sockeye salmon during their spawning migration. International Pacific Salmon Fisheries Commission Bulletin 22. 51 pages. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina Biological Survey, Raleigh, North Carolina. Publication Number 1980-12. 854 pages. Mosher, K. H. 1969. Identification of Pacific Salmon and Steelhead Trout by scale characteristics. U.S. Fish and Wildlife Service Circular 317: 1-17. Palmer, R. N. 1972. Fraser River Chum Salmon. Fisheries Service, Canadian Department of the Environment. Technical Report 1972-1. 284 pages. Salo, E. O. 1991. Life history of Chum Salmon (Oncorhy- nchus keta). Pages 231-309 in Pacific Salmon Life Histories. Edited by C. Groot and L. Margolis. University of British Columbia, Vancouver, British Columbia. Zallen, M., and M. Farwell. 1985. Description and evalua- tion of the 1984 survey of Chum Salmon spawning in the Fraser River mainstem. ESL Environmental Sciences Ltd. Vancouver, British Columbia. 21 pages. [Report prepared for the Department of Fisheries and Oceans, Fraser River, Northern British Columbia, and Yukon Division, 610 Derwent Way, New Westminster, British Columbia V3M 5P8.] Received 8 June 1995 Accepted 25 September 1995 1996 NOTES 335 A Fisher, Martes pennanti, with Multiple Amputations GILBERT PROULX and PAMELA J. COLE Alpha Wildlife Research & Management Ltd., 9 Garnet Crescent, Sherwood Park, Alberta T8A 2R7 Proulx, Gilbert, and Pamela J. Cole. 1996. A Fisher, Martes pennanti, with multiple amputations. Canadian Field- Naturalist 110(2): 335. An adult male Fisher (Martes pennanti) captured twice, and possibly three times, in a leghold trap suffered amputations of the left hind leg at the knee joint and most of the digits of the right and left front feet. Key Words: Fisher, Martes pennanti, furbearer, leghold trap, trapping, amputations, Alberta. Fishers (Martes pennanti) caught in leghold traps sustain severe injuries such as broken bones and amputations (Quick 1953; Coulter 1960). From 1990 to 1993, we recorded self-mutilations and amputa- tions in 118 (15%) out of 762 Fishers harvested in Alberta (Cole and Proulx 1994). During this study, we collected a three-legged male adult Fisher from the northeastern region. The left hind leg had been amputated at the knee level. The lesion was well healed and fibrosed over. On the right front foot, digits 1, 2 and 3 had also been amputated at the metacarpal joint and the terminal phalange of the fourth digit was missing. The injuries had healed over. The animal was captured in a leghold trap dur- ing the 1992-1993 trapping season. During this cap- ture, it had chewed off the digit 3 terminal and mid- dle phalanges and the digit 4 terminal phalange of the left front foot. The amputations observed on the right front foot were similar to the fresh self-mutilations observed on the left front foot. However, because they were healed over, they likely occurred during an earlier capture. We are uncertain about the exact cause of the hind leg amputation. It could be the consequence of a bad fall or fight with another animal. On the other hand, in the northeastern region of Alberta, Number 3 leghold traps are set for Lynx (Felis lynx) and Coyote (Canis latrans), and Fishers are acciden- tally captured in these traps (Cole and Proulx 1994). According to Coulter (1960), these traps often break Fisher leg bones and the animals may lose part or most of a leg. This Fisher may have suffered a hind leg amputation during such a capture. On the basis of the age of the lesions, i.e., only the right front foot and the left hind leg were covered with scar tissue, and the unlikelihood that the animal could have been caught by both these limbs at the same time, we con- clude that this Fisher could have been captured at three different instances in a leghold trap. The present observation demonstrates the repeti- tive use by an animal of extreme measures to escape a trap in at least two instances, and the recuperative potential of a Fisher subjected to consecutive serious trauma. Acknowledgments We thank Floyd Kunnas, Alberta Fish and Wildlife Division, for providing the specimen and veterinary pathologist Detlef Onderka, Alberta Agriculture, for conducting the postmortem exami- nation which confirmed our observations. Literature Cited Cole, P. J., and G. Proulx. 1994. Leghold trapping: a cause of serious injuries to Fishers. Martes Working Group Newsletter 2(1): 14-15. Coulter, M. W. 1960. The status and distribution of Fisher in Maine. Journal of Mammalogy 41: 1-9. Quick, H. F. 1953. Wolverine, Fisher and Marten studies in a wilderness region. Transactions of the North American Wildlife Conference 18: 512-533. Received 24 May 1995 Accepted 11 October 1995 336 THE CANADIAN FIELD-NATURALIST Vol. 110 Second Record and Possible Breeding of the Eurasian Wigeon, Anas penelope, in the District of Mackenzie, Northwest Territories MICHAEL A. FOURNIER and JAMES E. HINES Canadian Wildlife Service, P.O. Box 637, Yellowknife, Northwest Territories X1A 2N5 Fournier, Michael A., and James E. Hines. 1996. Second record and possible breeding of the Eurasian Wigeon, Anas penelope, in the District of Mackenzie, Northwest Territories. Canadian Field-Naturalist 110(2): 336-337. Eurasian Wigeons, Anas penelope, were observed in the Yellowknife area on seven occasions between 1990 and 1992. These included the second record of this species in the District of Mackenzie. The temporal distribution and close proximi- ty of six of these observations suggested that breeding of Eurasian Wigeon may have occurred in the Yellowknife area dur- ing this period. Key Words: Eurasian Wigeon, Anas penelope, Northwest Territories, possible breeding. In North America, the Eurasian Wigeon, Anas penelope, has been recorded in virtually every state, province, and territory north of Mexico but there is no proven evidence of breeding (Edgell 1984). It is considered rarest in the westerly interior (Palmer 1976) although there are a number of records from interior Alaska (Edgell 1984). The Canadian status of this species has been defined as scarce autumn and spring transient and local winter resident (Godfrey 1986). In northwestern Canada, Campbell et al. (1990) classified it as casual in summer and autumn in interior British Columbia; in Yukon Territory it is considered rare (Anonymous 1992); and, in Alberta, Pinel et al. (1991) listed thirteen records in 1971-1980 which represent an increase in abundance over previous decades. The first report of A. penelope in the District of Mackenzie was apparently erroneous. Russell (1898) supposedly collected a specimen in 1892 at the Stagg River near Yellowknife. However, Preble (1908) later noted that Ridgway had identified Russell’s specimen as an American Wigeon, Anas americana. The first valid and apparently only record of A. pene- lope in the District of Mackenzie to date was a single male, observed by Salter et al. (1974) on 14 May 1973 on the Mackenzie River at Fort Simpson. We observed a pair of wigeons, the male of which was positively identified as A. penelope, on a road- side pond about 45 km northwest of Yellowknife (62°27'N, 114°22’W) on 6 June 1990. The female was not observed well enough to detect the subtle differences separating females of A. penelope and A. americana. The latter is a common breeding species in the Yellowknife area, and mixed pairs must occur because hybrids are occasionally recorded (Watson 1970; Edgell 1984; Merrifield 1993). Regardless, this observation constitutes the second record of A. penelope for the District of Mackenzie. Subsequent observations of A. penelope occurred on 23 and 31 May 1991. On both occasions a single male was seen on a roadside pond 32 km northwest of Yellowknife. The dates of these observations (coincident with the egg-laying period of A. ameri- cana in the Yellowknife area), as well as the behaviour of this male (remaining on or near the same pond for at least a nine-day period) suggested that he might have had a mate nesting in the vicinity. Another observation of a single male occurred on 1 June 1991 on Great Slave Lake approximately 5 km from the city of Yellowknife. Further observations of A. penelope occurred in a third consecutive year. A single male was observed on 27 May 1992 on the same pond where the male was seen twice in May 1991. Another single male (presumably the same individual) was observed on 28 May and 3 June 1992, on a pond less than 2 km away. No observations of A. penelope were reported in the Yellowknife area prior to 1990 or since 1992, although waterfowl studies have continued in the area at the same level of intensity from 1985 to the present. All of the observations we report here, except the bird on Great Slave Lake, likely represent only one or two individuals. The remaining six observations occurred within a distance of 13 km, five of these within a distance of 2 km. In particular, it seemed highly unlikely that three observations on the same pond and two less than 2 km away, during two con- secutive years, were coincidental. Rather, they sug- gested migrational homing by a male, or a male(s) following a philopatric female, and may constitute circumstantial evidence of breeding. Migrational homing of waterfowl is a well-recog- nized phenomenon (Anderson et al. 1992). As sum- marized by Blohm and Mackenzie (1994), migra- tional homing of female ducks to natal and breeding areas has received considerable attention in the liter- ature, whereas homing of unmated males and pairs, although known to occur, are less well documented. Anderson et al. (1992) presented data on homing rates of a variety of waterfowl. Although data for A. penelope were not included, data for A. americana indicated an approximate homing rate of 33% to 1996 natal areas and 44% to breeding areas for females, and 8% to breeding areas for males (no data avail- able for natal homing in males). Based on these data, it seems most probable that our observations were of a male(s) following a homing female, although we cannot eliminate the possibility of homing in an unpaired male. There has been much conjecture over the years regarding the breeding of A. penelope in North America. Richardson (cited in Baird et al. 1884), suggested that it bred in northern interior Canada to 68° north. This idea was reiterated by Phillips (1923) and the evidence in its support was summarized by Hasbrouck (1944). However, Hasbrouck’s argument was largely based on the supposed record of Russell (1898). Hasbrouck apparently never verified the specimen himself, nor was he aware that Preble (1908) had proven the record false. Forbush (1925, cited in Watson 1970) also believed the species was breeding in North America. In more recent literature, many authors discount (e.g. Bellrose 1976) or appear indifferent to (e.g., Johnsgard 1975; Palmer 1976) the possibility of a breeding population in North America. However, Edgell (1984), following a thor- ough analysis of North American records of A. pene- lope, stated; “Whether or not the wigeon is a breed- ing member of the American avifauna is still open to question. The possibility remains high, ..., and inter- mittent breeding almost to be expected from the fre- quency and timing of records and the regular occur- rence of wintering and summering pairs.” Other Eurasian species (e.g., Little Gull, Larus minutus, and Common Black-headed Gull, Larus ridibundus) have eastablished small, local breeding populations in North America in recent decades (Godfrey 1986). Although we do not subscribe to Richardson’s notion of a traditional breeding ground for this species located in subarctic Canada, we believe it likely that A. penelope or mixed pairs of A. penelope and A. americana occasionally breed in North America. Further, we believe breeding of A. pene- lope may have occurred in the Yellowknife area dur- ing the period 1990-1992. Acknowledgments M. Kay, D. Kuhn, and J. Sirois provided field observations. A. J. Erskine, P. Latour, and an anony- mous referee provided valuable comments on the draft manuscript. Literature Cited Anderson, M. G., J. M. Rhymer, and F. C. Rohwer. 1992. Philopatry, dispersal, and the genetic structure of waterfowl populations. Chapter 11 in Ecology and man- agement of breeding waterfowl. Edited by B. D. J. Batt, A. D. Afton, M. G. Anderson, C. D. Ankney, D. H. NOTES Sioi/ Johnson, J. A. Kadlec, and G. L. Krapu. University of Minnesota Press, Minneapolis. 635 pages. Anonymous. 1992. Birds of the Yukon. Field checklist. Yukon Department of Renewable Resources, Whitehorse. 3 pages. Baird, S. F., T. M. Brewer, and R. Ridgway. 1884. The water birds of North America. Reprint Edition 1974. Arno Press Inc. 1089 pages. Bellrose, F. C. 1976. Ducks, geese and swans of North - America. Stackpole Books, Harrisburg. 540 pages. Blohm, R. J., and K. A. Mackenzie. 1994. Additional evidence of migrational homing by a pair of mallards. Journal of Field Ornithology 65: 476-478. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, and M. C. E. McNall. 1990. The birds of British Columbia. Volume 1. Nonpasserines. Royal British Columbia Museum, Victoria. 514 pages. Edgell, M. C. 1984. Trans-hemispheric movements of Holarctic Anatidae: the Eurasian wigeon (Anas penelope) in North America. Journal of Biogeography 11: 27-39. Godfrey, W. E. 1986. The birds of Canada. Revised edi- tion. National Museum of Natural Sciences, Ottawa. 595 pages. Hasbrouck, E. M. 1944. Apparent status of the European widgeon in North America. Auk 61: 93-104. Johnsgard, P. A. 1975. Waterfowl of North America. Indiana University Press, Bloomington. 575 pages. Merrifield, K. 1993. Eurasian X American wigeons in western Oregon. Western Birds 24: 105-107. Palmer, R. S. 1976. Handbook of North American birds. Volume 2, Waterfowl (Part 1). Yale University Press, New Haven. 521 pages. ; Phillips, J. C. 1923. A natural history of the ducks. Volume II. The genus Anas. Reprint Edition 1986, Dover Publications Inc., New York. 409 pages. Pinel, H. W., W. W. Smith, and C. R. Wershler. 1991. Alberta birds, 1971-1980. Volume 1. Non-passerines. Provincial Museum of Alberta, Natural History Occasional Paper Number 13, Edmonton. 243 pages. Preble, E. A. 1908. Biological investigation of the Athabasca-Mackenzie region. U.S. Department of Agri- culture, Bureau of Biological Survey, Washington. 557 pages. Russell, F. 1898. Explorations in the far north, being the report of an expedition under the auspices of the University of Iowa during the years 1892, 93, and ‘94. University of Iowa, Iowa City. 290 pages. Salter, R., W. J. Richardson, and C. Holdsworth. 1974. Spring migration of birds through the Mackenzie Valley, N.W.T. April-May, 1973. Chapter I in Ornithological studies in the Mackenzie Valley, 1973. Edited by W. W. H. Gunn, W. J. Richardson, R. E. Schweinsburg, and T. D. Wright. Arctic Gas Biological Report Series, Volume 28. LGL Ltd., Environmental Research Associates. 168 pages. Watson, G. E. 1970. A presumed wild hybrid Baldpate X Eurasian wigeon. Auk 87: 353-357. Received 21 June 1995 Accepted 13 October 1995 338 THE CANADIAN FIELD-NATURALIST Vol. 110 Red Fox, Vulpes vulpes, kills a European Beaver, Castor fiber, Kit NILS B. KILE, PETTER J. NAKKEN, FRANK ROSELL, and SIGURD ESPELAND Telemark College, Department of Economics, Environment and Sport Studies, N-3800 Bg, Norway Kile, Nils B., Petter J. Nakken, Frank Rosell, and Sigurd Espeland. 1996. Red Fox, Vulpes vulpes, kills a European Beaver, Castor fiber, kit. Canadian Field-Naturalist 110(2): 338-339. We observed an adult Red Fox (Vulpes vulpes) attack, kill and partially consume a 2-month-old female kit European Beaver (Castor fiber) near its lodge in Norway. The inner organs were consumed first. One adult beaver apparently attempted to frighten the fox away by tail-slapping. Key Words: Red Fox, Vulpes vulpes, European Beaver, Castor fiber, predation, southeast Norway. On 31 July 1994, during a beaver census in Southeast Norway (58°39'N, 7°59’E), a Red Fox (Vulpes vulpes) was observed killing a European Beaver (Castor fiber) kit (see Kile and Nakken 1995). The observer was seated with binoculars about 100 meters from the lodge. At 1930 hours a kit emerged from the lodge and started to eat Beaked Sedge (Carex rostrata) 15 meters from the lodge, on shore about 2 meters from the water’s edge. One hour later, first a single adult, and then, several min- utes later, another adult with kit emerged from the lodge and swam away. One hour and 50 minutes after the adults left an adult Red Fox approached the lodge (Figure 1). A large stone was situated between the fox and the foraging kit, so neither saw the other. The fox walked to the top of the lodge and suddenly stiffened, apparently having smelled the kit or heard it eating (the kit was still not visible). The observer could not hear the kit eating. The distance between the two animals was then about 15 meters. The fox crouched into attack position, and after a short pause attacked and killed the kit while still on shore. The struggle lasted a few seconds. The kit did not attempt to flee or fight back, apparently having been caught completely unaware by the fox. No sounds were heard from either kit or fox. The fox dragged the kit approximately 10 meters to a flat stone under a large spruce (Picea abies) and pro- ceeded to feed on the carcass. Five minutes later an adult beaver appeared and swam to the vicinity of the kill site, circling 20-30 meters from the water’s edge before gliding towards the shore where the fox was feeding. The fox stood up and walked a few steps towards the adult. The beaver then tail-slapped, appar- ently in an attempt to frighten it, or to alert other beavers (Wilsson 1971). The fox ignored the tail-slap and returned to its prey. The adult beaver swam near the the lodge for the next 10 minutes, about 40 meters from the fox, before finally diving and disappearing. Directly after this the observer frightened the fox away and collected the remains of the kit. The remains of the carcass weighed about 2 kg (estimated live weight at least 2.5 kg). Based on the mean parturition date for this region (Syvertsen 1976; Morner 1990) we estimated the age at 2 months, the sex was female (see Wilsson 1971). The breast had been opened from the ventral side. The sternum, heart, lungs, stomach, cecum and both intestines had been eaten, as well as most of the liver and ribs, and part of the muscle from the right fore- arm. Two cervical vertebrae and 11 thoracic verte- brae were missing and apparently eaten. Inspection of the skin revealed two puncture marks, with subdermal hemorrhaging, at the upper edge of each scapula, from the fox’s lower and upper canines. The fox had bitten over the spinal column between the shoulders. Judging from the short dura- tion of the struggle, it seems likely that the spinal cord had been severed, though direct evidence of this was not available due to the advanced stage of car- cass consumption. This is apparently the first reported observation of a Red Fox killing a beaver. Payne and Finlay (1975) reported a Red Fox attacking a beaver, but they did not observe the attack itself, and the beaver was still alive. Novak’s (1987) review of the beaver mentions ° Beyond observer's < vision = = a Beyond observer's vision 1 Scale (meters) 50 FiGuRE |. The relative positions of the observer, Fox, Beaver kits and Beaver adults in numeric order: 1) observer; 2) kit kill site; 3) single adult appears and swims away; 4) another adult and kit appears and swims away; 5) attack route of fox; 6) stone between fox and kit: 7) fox eats beaver; 8) return route of adult beaver and 9) site of tail-slapping. 1996 the Red Fox as a predator of little importance for beaver populations. The Red Fox was not considered as a potential predator in Hill’s (1982) review. Tyurnin (1984) did not record any attacks by Red Fox on beaver. Red Fox predation on beaver seems to be unusual. Only young beaver (1-2 months) are probably susceptible to fox attack while feeding on land and before having fully developed escape behavior (Wilsson 1971). Male kits are reportedly more cautious than females (Shadle 1956). Acknowledgments We thank Dr. H. Parker for reviewing several drafts of the manuscript. Literature Cited Hill, E. P. 1982. Beaver (Castor canadensis). Pages 256-281 in Wild mammals of North America Biology, Management, and Economics. Edited by J. A. Chapman and G. A. Feldhamer. John Hopkins University Press, Baltimore and London. Kile, N. B., and P. J. Nakken. 1995. Koloni- og kullstgr- relse hos bever (Castor fiber) i Gautestad-omradet, Evje og Hornes kommune, Aust Agder. Hovedoppgave ved 3-arig studium i natur og miljévernfag. Hggskolen i Telemark. Norway. 32 pages. NOTES B39) Morner, T. 1990. Fodseltid hos svenska biivrar (Castor fiber). Statens Veterinarmedicinska Anstalt, Uppsala, Sweden. 13 pages. Novak, M. 1987. Beaver. Pages 283-312 in Wild Furbearer Management and Conservation in North America, Ontario. Edited by M. Novak, J. A. Baker, M. E. Obbard and B. Malloch. Ontario Ministry of Natural Resources. Payne, N. F., and C. Finlay. 1975. Red Fox attack on beaver. Canadian Field-Naturalist 89: 450-451. Schadle, A. R. 1956. The American Beaver. Animal Kingdom. [Pages 59, 60, 61]. Syvertsen, K. 1976. Undersgkelse av bever (Castor fiber) i Agder-fylkene-Sgr-Norge. Populasjonstetthet, alders- bestemmelse, reproduksjon og aktivitet. Hovedfags- oppgave ved Universitetet i Bergen 1976. 180 pages. Tyurnin, B. N. 1984. Factors determining numbers of the river beavers (Castor fiber) in the European North. Soviet Journal of Ecology [in English] 14: 337-344 [Translated from Ekologiya number 6: 43-51]. Wilsson, L. 1971. Observations and experiments on the ethology of the European beaver (Castor fiber L). Viltrevy 8: 115-266. Received 29 June 1995 Accepted 29 September 1995 Polar Bear, Ursus maritimus, Depredation of Canada Goose, Branta canadensis, Nests ARTHUR E. SMITH! and MICHAEL R. J. HILL? ‘Department of Wildlife Ecology, 1630 Linden Drive, Room 226, University of Wisconsin, Madison, Wisconsin 53706 "Department of Zoology, The University of Western Ontario, London, Ontario N6A 5B7 Smith, Arthur E., and Michael R. J. Hill. 1996. Polar Bear, Ursus maritimus, depredation of Canada Goose, Branta canadensis, nests. Canadian Field-Naturalist 110(2): 339-340. Evidence for depredation of four marked Canada Goose nests by Polar Bear in May 1995 on Akimiski Island, James Bay, Northwest Territories, is described. Thirty-two additional marked nests were untouched. Key Words: Canada Goose, Branta canadensis, Polar Bear, Ursus maritimus, nest depredation. Polar Bears (Ursus maritimus) have been reported to consume the eggs of Lesser Snow Geese (Chen caerulescens) (F. Cooch in Abraham et al. 1977), Light-bellied Brent Geese (Branta bernicla hrota) (Madsen et al. 1989), and eggs of unspecified or unknown origin (R6mer and Schaudinn 1900; Loughrey 1956; Harrington 1965; Russell 1975), but we found no published reports documenting Polar Bears depredating Canada Goose eggs or nests. While conducting Canada Goose (Branta canadensis interior) nest searching on 28 May 1995, we found four Canada Goose nests on Akimiski Island, Northwest Territories, which had apparently been depredated by a Polar Bear. All four nests were with- in 200 m of one another along the willow (Salix spp.) transition zone paralleling the coastline, approximately 8 km east of Houston Point (53°12'N, 81°08’ W). Each depredated nest had been marked on 11 May 1995 with a 10 x 12 cm orange plastic flag attached to a one m wire, placed 20 m north of each nest site. At that time, the nests had clutches of two, three, four, and four intact eggs, but egg-laying was proba- bly not complete as indicated by floating (Westerskov 1950). Additional Canada Goose nests (n = 32), within eight km in either direction along the coastline of the four destroyed nests, were also marked on 11 May but were not destroyed by Polar Bears. On 28 May, each destroyed nest had nest material and egg fragments scattered within a two m 340 radius around the nest bowl. Two of the nests had one egg and one nest had two eggs which were com- pletely flattened, but were still held together by the egg membrane. These four eggs were also found within two m of the nest; the fourth nest did not have flattened eggs associated with it. Contents from the flattened eggs were not dried to the shells suggesting that they had not been broken while in or near the nest bowl. All of the nest flags showed evidence of being chewed, and the flag wires had been bent in several directions, sometimes up to 45°, and three of four wires had been pulled out of the ground. Polar Bear tracks were clearly visible in the mud leading to and from the nests and nest flags, as well as in the general direction of the next closest destroyed nest. No other predator tracks were observed. The hind tracks were measured at approxi- mately 18 cm wide; adult Black Bear (Ursus ameri- canus) hind tracks are approximately nine cm wide (Murie 1974). It is likely that the bear visited the nests sometime after 23 May as a rain storm occurred the afternoon of 22 May and continued until mid-day on 23 May, and neither the bear tracks or egg shell fragments seemed affected by rain drops or rain-caused mud splashings. Human activities were minimized at each nest during marking and the depredation event occurred well after the last visit to the nest, therefore there was probably little, if any, human scent left at the nest sites at the time of nest depredation. There were no sightings of Polar Bears during either the 11 May or 28 May visits. The most southern boundary of Polar Bear range occurs in James Bay near a polynya that forms south of Akimiski Island in winter (Stirling 1988: 64-65). Here the ice pack is usually completely melted by mid-July (Russell 1975), the earliest area to do so adjacent to Canada Goose nesting grounds. Upon melt, the bears move onto the mainland and islands, forming potential Polar Bear predation on geese. However, the Canada Goose nesting period in this area typically extends from early May to mid-June, making it unlikely that Polar Bears encounter many goose nests. However, fresh bear tracks have been - observed in May on Akimiski Island in previous years, and a Polar Bear was encountered on the north shore of the island 24 May 1993 (J. O. Leafloor, per- THE CANADIAN FIELD-NATURALIST Vol. 110 sonal communication). On 30 May 1995, Polar Bear tracks were found in close proximity to approximate- ly 30 destroyed Lesser Snow Goose nests on the coast 25 km west (81°30'W longitude) of the depre- dated Canada Goose nests (S. R. McWilliams, per- sonal communication). This observation indicates that the depredation of the Canada Goose nests in 1995 was not an isolated incident, and that when Polar Bears are present in goose nesting areas they may consume goose eggs and should be regarded as potential nest site predators. However, at this time, we do not believe that Polar Bears are major preda- tors of goose nests on Akimiski Island. Acknowledgments Funding and logistical support were provided by the Ontario Ministry of Natural Resources, Moosonee office. We thank Jim Leafloor, Don Rusch, Scott McWilliams, Jonathan Thompson, A. J. Erskine and two anonymous reviewers for comments on earlier drafts. Literature Cited Abraham, K. F., P. Mineau, and F. Cooke. 1977. Unusual predators of Snow Goose eggs. Canadian Field- Naturalist 91(3): 317-318. Harrington, C. R. 1965. The life and status of the polar bear. Oryx 8(3): 169-176. Loughrey, A. G. 1956. The polar bear and its protection. Oryx 3(5): 233-239. Madsen, J., T. Bregnballe, and F. Mehlum. 1989. Study of the breeding ecology and behaviour of the Svalbard population of light-bellied brent goose Branta bernicla hrota. Polar Research 7: 1-21. Murie, O. J. 1974. A field guide to animal tracks. Second edition. Houghton Mifflin Company, Boston. 375 pages. Romer, F. and F. Schaudinn. 1900. Fauna Arctica I. Gustav Fischer, Jena. Russell, R. H. 1975. The food habits of polar bears of James Bay and southwest Hudson Bay in summer and autumn. Arctic 28(2): 117-129. Stirling, I. 1988. Polar Bears. University of Michigan Press, Ann Arbor. 220 pages. Westerkov, K. 1950. Methods for determining the age of game bird eggs. Journal of Wildlife Management 14: 56-67. Received 26 July 1995 Accepted 14 December 1995 1996 NOTES 341 Wood Turtles, Clemmys insculpta, in the Fresh-tidal Hudson River ERIK KIviAT and JAMES G. BARBOUR Hudsonia Limited, Bard College Field Station, Annandale, New York 12504 Kiviat, Erik, and James G. Barbour. 1996. Wood Turtles, Clemmys insculpta, in the fresh-tidal Hudson River. Canadian Field-Naturalist 110(2): 341-343. Twelve records of the Wood Turtle, including one viable egg clutch, 1973-1995, were associated with freshwater tidal swamps on the Hudson River in New York. This is the first report of a Wood Turtle population in tidal habitat. Key Words: Wood Turtle, Clemmys insculpta, estuary, fresh-tidal swamp, Hudson River, New York, wetland. Little herpetological field work has focused on temperate estuaries which continue to be altered for industry, housing, transportation, and recreation. Reptile and amphibian use of estuaries (including freshwater tidal environments) may be limited by tidal fluctuation, salinity, water depth and breadth, currents, ice scouring, temperature, storms, preda- tion, or pollution (Bleakney 1958: 44; Neill 1958; Dunson 1986; Kiviat 1989; Lillywhite and Ellis 1994). The Wood Turtle (Clemmys insculpta) is widespread in association with nontidal streams and ponds of the Hudson Valley of eastern New York. Here we report 12 records from four sites on the tidal Hudson River (Table 1). Mean tide range is ca 1.2 m and maximum salinity < 0.1 ppt in the reaches of the Hudson River where Wood Turtles have been found. The four sites (Table 1) contain extensive complexes of intertidal and supratidal tree or shrub-dominated swamp, intertidal marsh, tidal creeks in swamp and marsh, tidal mouths of tributary streams, and upland habitats (woods and old fields). Irregularly, tidally flooded (supratidal) pools are also present at all sites except Vanderburg Cove. The supratidal hardwood swamps resemble nontidal floodplain habitats with which the Wood Turtle is associated elsewhere in the region. Beaver (Castor canadensis) food caches on the bot- toms of tidal creeks, and the burrows of Beaver and Muskrat (Ondatra zibethicus) in the creek banks are possible overwintering microhabitats. It is not clear whether the Hudson River Wood Turtles use the tidal swamp creeks as primary habi- tat, or if they are resident in nearby nontidal streams and move into the tidal wetlands to forage; e.g., dur- ing the unusually cold spring of 1994. Although the Hudson River Wood Turtles could have been flood- washed or human-transported, the consistent associ- ation with tidal swamp (Table 1), a habitat of restricted extent on the Hudson, suggests a resident population. A probable exception is the hatchling found in May 1973 in the tidal mouth of a tributary to the Tivoli Bays, 1100 m from the nearest tidal swamp. During the preceding two days 56 mm of rain fell, suggesting this individual was washed downstream to the estuary. Although the geographic distribution of the Wood Turtle approaches or adjoins estuaries in 12 other states and provinces from Nova Scotia to Virginia (see range map, Ernst et al. 1994), we know of only one record of a Wood Turtle in an estuary other than the Hudson River (Parker River, Newbury, Essex County, Massachusetts, 1975, captured and released by David Taylor; Tom French, Massachusetts Natural Heritage Program, personal communica- tion). Inclusion of Wood Turtle with the fresh-tidal marsh biota in Odum et al. (1984) was based on Kiviat (1978) (the 1974 Tivoli record, Table 1) and McCormick (1970); the latter may refer to diked (nontidal) marsh as noted by McCormick (1970) and Harding (1986: 56). The Wood Turtle is rare on the Coastal Plain compared to regions farther inland (Reed 1956; Latham 1971; Hardy 1972; Gilhen 1984; Norden and Zyla 1989; Ernst and McBreen 1991; Klemens 1993; Miller 1993). The Hudson River is the only estuary that transects the Appalachian Mountains, and the ca 130 km long fresh-tidal reach does not touch the Coastal Plain. Acknowledgments The sites are administered by the Northern Catskills Audubon Society, Hudson River National Estuarine Research Reserve, and Southlands Farm. Scenic Hudson, Inc., the New York Natural Heritage Program, and the Town of Rhinebeck supported our field work. Michael W. Klemens, C. Lavett Smith, and Gretchen Stevens commented on drafts of this paper, Bard College Field Station — Hudsonia Contribution 64. Literature Cited Bleakney, J. S. 1958. A zoogeographical study of the amphibians and reptiles of eastern Canada. National Museum of Canada Bulletin 155. 119 pages. Dunson, W. A. 1986. Estuarine populations of the snap- ping turtle (Chelydra) as a model for the evolution of marine adaptations in reptiles. Copeia (3): 741-756. Ernst, C. H., R. W. Barbour, and J. E. Lovich. 1994. Turtles of the United States and Canada. Smithsonian Institution Press, Washington, D.C. 578 pages. Ernst, C. H., and J. F. McBreen. 1991. Wood turtle. Pages 455-457 in Virginia’s Endangered Species; THE CANADIAN FIELD-NATURALIST Vol. 110 342 (EOI UONRIS Plely eSa][0D preg) perp Surpyorey | ‘payoiey sssa g ‘osng Aq poyeqnouy, “spueyjoM [epl) pue siayem [epyy Aq papunodins st purysi ey p] Wy) ‘pepooyy Ay[epy Ayrepn3aut st [ood ay Le “CLI uoNtIg platy edel[0D preg, ‘qyeurxoiddy ‘O19Y}] PUNOJ 319M S[LNPIAIPUT ddIY} SBI] IV P2PIPUI YIeID WOH suey wo suaurIoads ay} pur sydessojoyd uo suraned yoroyg fe-nseyd “(@[1M C66 ey) Jo ydessojoyd & pourwexe eIATY pue opin} WOH SWeY 066] 24) Jo ydessojoyd ev pourwexe noqieg) poydesso.0yd, ‘(P66 Sulids ul parp) sureuras AreyuoUIseIy, (42019 sinqs|[e4) Areinqin 199} OA} SUTSSTU Jo yNoU! [epy wor aTeul 9861 Ayunog sseyoing ZeIATY ANA Wi ¢ Play pyo purydy Wpy =: 190190 OE I Gal uO SSo€L uSS,CSolV ‘AOD Ysinqiapue A (TIE Meg) Areingin [etuuaisad Jo you ‘au0z KAI Aqunos sseyoing WATS] YA [epHisqut Jo adpe saddp) sulyoeH RA IT 07 6S1 iV, SSoEL 100, 100CP ‘skeg TOALL E861 Ayunog sseyoing SAQP AA WIRTTTEAN cPurjs] JesniD ‘yred UG apy AP 9RD OT 091 ubb SSo€L iS, 100th ‘skeg TOALL prewod Porapsog ({[IF) peorey Jo yoNID oun OT OC O9T uVVSSo€L ul V,COoCY ‘sheg TOALL duwiems 1661 Ayunoy sseyoing JOPeIN (OD WeITTTA sod) [eph Ul prod WI yOpV sunt 17 OC O9T ut Ci SSotL uSV,COoCP ‘sheg TOALL durems [epn ul ,o[eul S661 AUNOD ausd0ID Auog yeroqod yeord Jofeu ‘peop puno{ NPV tudy 0¢ O0C 181 OV TSotL ev0$, 11 oCV ‘Yoo WOH suey yaolo [epn Jofew Woy Wi py “ysreul pue dwems qniys jepn oyeul +661 AJUNOD UddID elnoqieg *H soulef Aq pasepsog peor Iq WNpVv AR PI 00C [8 w0S,1So€L u00, CL oCV “YeorD WOH suey dures [ep ul C661 Ajunog udaeIH Auog yesoqed PeOl UIP 1B Joos ole] WNpVy AW TI 00C I8T w0S,TSo€L u00, CL oCV ‘eer WOH suey zpoy seurer dwiems 39] JUO1J paulojep 0661 Ayunod usetH ‘Auag ye1oqod sel} [ep Ur yuRG Yoor) ({aTeuey) INPV sunt 07 00C 181 ev00,CSo€L euv00, C1 oCP “YeorD WOH suey dure ms oyeul +661 Aqunod aueaID 7 eA Ma 9o7) [epy ul yueq Yoo) NPY tudy 17 O00C [8 ue LSotL ul€,CloCt “Year WOH suey dures Ajunog eiquinjod 201) [ep Aq posap10q a[PUlay +661 *yoorg yodyo01¢$ Inoqieg “DH soulef (ITF) proses uo ;pesq WNpVy eune €T 6C col wl, OVotL uC 81oCV Jo WNo|| JOAIasqQ yeqiqey SsP[D arg ey uyy opnmiisuoTy A opnine’y N Aq[B0'T ‘dures [ept) paleurtuop-qniys Jo 901) Jo soreyoey ul vare ajeutxoldde = By (ANID YIOX MON) purS] ULWeYULP] JO pus UJAYyINOS ay) aAogK SIAISWO[PY JOATY = WIPY “YOR MON “JOATY UOspny [epl) JowMYSIJ 9Y} WOIJ SpIOSEI BIN], POOM “[ ATAVL 1996 Proceedings of a Symposium. Edited by K. Terwilliger. McDonald and Woodward Publishing Company, Blacksburg, Virginia. 672 pages. Gilhen, J. 1984. Amphibians and reptiles of Nova Scotia. Nova Scotia Museum, Halifax. 162 pages. Harding, J. J. 1986. Marsh, meadow, mountain; Natural places of the Delaware Valley. Temple University Press, Philadelphia, Pennsylvania. 267 pages. Hardy, J. D., Jr. 1972. Reptiles of the Chesapeake Bay region. Chesapeake Science 13(Supplement): $128-S 134. Kiviat, E. 1978. Hudson River east bank natural areas, Clermont to Norrie. Nature Conservancy, Boston, Massachusetts. 115 pages. Kiviat, E. 1989. The role of wildlife in estuarine ecosys- tems. Pages 438-476 in Estuarine Ecology. By J. W. Day, Jr., C. A. S. Hall, W. M. Kemp, and A. Ydafiez- Arancibia. John Wiley and Sons, New York. 558 pages. Klemens, M. W. 1993. Amphibians and reptiles of Connecticut and adjacent regions. State Geological and Natural History Survey of Connecticut Bulletin 112. 318 pages. Latham, R. 1971. Dead wood turtles found on eastern Long Island. Engelhardtia 4(4): 32. Lillywhite, H. B., and T. M. Ellis. 1994. Ecophysio- NOTES 343 logical aspects of the coastal-estuarine distribution of acrochordid snakes. Estuaries 17(1A): 53-61. McCormick, J. 1970. The natural features of Tinicum Marsh, with particular emphasis on the vegetation. Pages 1-104 in Two Studies of Tinicum Marsh; Delaware and Philadelphia Counties, Pennsylvania. Conservation Foundation, Washington, DC. 123 pages + folded map. Miller, R. W. 1993. Comments on the distribution of Clemmys insculpta on the Coastal Plain of Maryland. Herpetological Review 24(3): 90-93. Neill, W. T. 1958. The occurrence of amphibians and rep- tiles in saltwater areas, and a bibliography. Bulletin of Marine Science of the Gulf and Caribbean 8(1): 1-97. Norden, A., and J. Zyla. 1989. The wood turtle, Clemmys insculpta, on the Maryland Coastal Plain. Maryland Naturalist 33(1-2): 37-41. Odum, W. T., T. J. Smith, HI, and C. MclIvor. 1984. The ecology of tidal freshwater marshes of the United States East Coast: A community profile. U.S. Fish and Wildlife Service, FWS/O17. 177 pages. Reed, C. F. 1956. Distribution of the wood turtle, Clemmys insculpta, in Maryland. Herpetologica 12(1): 80. Received 1 August 1995 Accepted 21 December 1995 Denning Behaviour of Non-gravid Wolves, Canis lupus L. DAvip MEcH!, MICHAEL K. PHILLIPS?, DOUGLAS W. SMITH?2, and TERRY J. KREEGER? 1U. S. National Biological Service, Patuxent Environmental Science Center, 11510 American Holly Drive, Laurel, Maryland 2708 Mailing address: North Central Forest Experiment Station, 1992 Folwell Avenue, St. Paul, Minnesota 55108 2Yellowstone Center for Resources, P. O. Box 168, Yellowstone National Park, Wyoming 82190 3Wyoming Game & Fish Department, Bosler Route Box 8101, Wheatland, Wyoming 82201 Mech, L. David, Michael K. Phillips, Douglas W. Smith, and Terry J. Kreeger. 1996. Denning behavior of non-gravid Wolves, Canis lupus. Canadian Field-Naturalist 110(2): 343-345. Wild Wolves (Canis lupus) that had produced pups in earlier years but were not currently pregnant, and ovariectomized captive Wolves, dug dens during and after the whelping season even though they produced no pups. These observations suggest that den digging is not a function of pregnancy or of ovarian estrogen or progesterone. We hypothesize that increasing prolactin in spring elicits or mediates den-digging behavior. Key Words: Wolf, Canis lupus, den, behavior, estrogen, progesterone, prolactin, pups, pseudopregnancy, reproduction. Because Wolves, Canis lupus, dig dens primarily around parturition time and use the dens almost exclusively for raising pups, den digging might be a function of reproductive hormones such as proges- terone or estrogen. Apparently all mature female Wolves undergo the same hormonal changes during the luteal phase of the reproductive cycle whether gravid or not (Seal et al. 1979). The non-gravid luteal phase is known as pseudopregnancy. Presumably pseudopregnant Wolves behave similar- ly to gravid Wolves in seeking out and digging a suitable den, but no information about this subject has been recorded. This article reports on denning behavior of pseudopregnant Wolves as well as of ovariectomized Wolves. Data for this study were available from two study areas, Ellesmere Island, Northwest Territories, Canada (80°N, 86°W) (Mech 1988, 1995), and Yellowstone National Park, Wyoming (45°N 110°W) (Cook 1993); and from a pack of four 2- year-old captive Wolves, including three females, at the International Wolf Center in Ely, Minnesota (48° N, 92° W). In the first area, an identifiable Wolf, “Whitey,” born in 1987 and habituated to the senior author was observed each summer 1987-1995 from > 1m 344 (Mech 1988, 1995). Whitey produced pups in early June 1990, 1991, 1992, and 1994 in or near her natal den (Mech 1988, 1993; Mech and Packard 1990) and consistently remained with her pups throughout June (Mech 1995). However, in 1993 and 1995 she did not produce pups, as evidenced by close observation by the senior author. There was a lack of nipple development and she failed to localize consistently at a den. In 1993, none of Whitey’s known dens showed sign of digging or use on 2 July, and she was observed intermittently from 5 to 30 July 1993 trav- eling with her mate and 1992 offspring (Mech in press). However, Whitey visited her natal den area with her pack from 2200 hours on 13 July 1993 to 1220 hours on 14 July. Her mate even brought an Arctic Hare (Lepus arcticus) leveret to the den as if to give it to Whitey. However, Whitey was tem- porarily away from the den, and after 45 minutes, a yearling stole the Hare. Whitey lay near the den at 0002 hours, and from 0329 to 0439 hours, she dug out one entrance and entered the den six times for periods of 7-19 minutes each. She continued to sleep or rest until 1220 hours 14 July when the pack left. In 1995, Whitey was observed away from the den on 5 July and was not observed at her natal den dur- ing three aerial checks for the senior author by Department of Defense personnel from 28 May to 29 June. None of her dens showed signs of use on 29 June, although her natal den was not checked closely on that date. On 5 July, however, Whitey was observed 8 km from her natal den and had no nipple development, contrary to her appearances during her pup- bearing years. Whitey’s natal den was freshly dug out on 4 July 1995 although no pups were present. Whitey’s mate visited the den area on that date. On 5 July, Whitey visited the den at 1621 hours. She entered the den several times, inspected its various entrances, and dug it out, all for 18 minutes. Then she slept near the den from 1709 on 5 July to 2 0513 hours on 6 July. At 1800 hours on 5 July, Whitey’s 3-year-old male offspring arrived and entered and inspected the den for about a minute. Whitey also walked into the den for a few seconds at 1829 hours on 5 July. Then the male slept near Whitey through = 0513 hours on 6 July. Both Wolves were there when we left on 0513 hours but were gone at 1650 hours on 6 July and 1800 hours on 7 July. The Yellowstone Wolf, OO5F, was captured along with her mate (004M) and four young males in Alberta, Canada on 10 January, 1995. Her teats indi- cated that she had bred in the past (Mech et al. 1993). She and her packmates were placed in a 1- acre enclosure in Yellowstone and held for 10 weeks before being released in late March. Phillips and Smith monitored the behavior of Wolf OO5F by direct observation while she was in captivity and by THE CANADIAN FIELD-NATURALIST Vol. 110 radio-tracking and direct observation after her release with her pack from 31 March to 31 July 1995. During confinement, OOSF and 004M exhibit- ed breeding behaviors, and a bloody discharge from OOSF’s vulva was apparent. Wolf OO5F first showed signs of denning behavior on 29 April. Meanwhile two other females captured in the same area at the same time as Wolf OOSF and similarly transported, held, and released in the same region, produced pups about 26 April. Wolf OO5F explored Coyote (Canis latrans) dens several times from 29 April through 12 May. For about 2 hours on 29 April, OOSF and her packmates were observed near an unoccupied Coyote den (hole 1). Her mate and one yearling spent a few minutes excavating the hole. On 3 May, the pack bedded near a second unoccupied Coyote den (hole 2), about 400 m west of hole 1. From 0615 to 0821, OOSF slept at the den entrance, and from 0821 to 0830 she entered the den three times, each time disappearing for about a minute. At 0843, she entered hole 1, leaving at. 0845. At 1225 hours, OOSF, her mate, and a yearling bedded at a third unoccupied Coyote den (hole 3) about 4.8 km east of holes | and 2. Again, through- out 7 May, the entire pack was observed near holes | and 2, and on I1 May, OOS5F entered hole | for 8 minutes. On 12 May, between 1750 and 1844 hours, OO5F dug intensively at a fourth hole about 400 m west of hole 3. Her mate also dug for about three minutes. The hole was large, and O005F sometimes disap- peared. Clouds of dust indicated that she continued to dig while out of sight. The only time OO5F stopped digging is when displaced by a small herd of Bison (Bison bison) that approached the hole for 3—5 minutes. Wolf OOSF then returned and resumed dig- ging. At 1844 hours, OO5F ceased digging and rejoined the pack which had traveled about 2.0 km east. During the next few weeks, OO5F and pack remained within a few km of holes 1-4, but no more digging was seen. By early July, the pack began fre- quenting Pelican Valley about 32 km south of the holes. Three female captive Wolves in Minnesota were ovariectomized by Kreeger on 6 May 1994, when about 1-year old. During spring 1995, only one of the three showed any vaginal bleeding (7-10 days), but it was light. No other courtship or reproductive behavior was apparent. On 20 and 21 July 1995, at least two of the female Wolves, not including the one that showed minor bleeding, dug a large den. (Wild Wolves use dens in this area from about 27 April through June [Mech unpublished]). These two wolves probably were completely ovariectomized since they showed no bleeding. The latter observa- tion tends to support the report of a wild Wolf that had never ovulated but still paired with a male, held a territory, and restricted her movements for several 1996 weeks during the denning season, although it was unknown whether she actually attended a den (Mech and Seal 1987). The above observations suggest that den digging is not a function of pregnancy or even of ovarian estrogen or progesterone. Conceivably, the den dig- ging could have been in response to heat, as is typi- cal of sled dogs. However, this would not explain the Ellesmere or Yellowstone observations because heat was not a factor in those cases. A possible endocrine-based explanation for such denning behavior may lie with rising prolactin con- centrations. Prolactin, a hormone secreted from ante- rior pituitary lactotrophs, induces both maternal and paternal behavior in a variety of species (Zarrow et al. 1971; Dixson and George 1982; Bridges et al. 1985; Numan 1988; Gubernick and Nelson 1989). Both intact or neutered, male and female Wolves, show a strong circannual prolactin rhythm (Kreeger et al. 1991). Prolactin peaks just prior to summer sol- stice, coincident with whelping. All Wolves in a pack, including unrelated mem- bers, begin feeding other pack members even before whelping (Fentress and Ryon 1982). Thus, pups are not required for this behavior. Based on these obser- vations, it was hypothesized that increasing prolactin in spring elicits parental behavior in Wolves (Kreeger et al. 1991). Our observations of den digging by males and by females, in the absence of pregnancy or elevated progesterone or estrogen, support this hypothesis and lead us to further hypothesize that prolaction elicits or mediates den digging in Wolves. These observations also indicate that merely see- ing a Wolf at a den, or finding evidence of digging during the denning season, should not lead to the conclusion that pups were produced or will be during the current parturition season. Acknowledgments These projects were supported by the U.S. Fish and Wildlife Service, U.S. National Biological Service, the U.S.D.A. North Central Forest Experiment Station, the U. S. National Park Service, and the International Wolf Center. For the Ellesmere work, the logistical support of the Polar Continental Shelf Project, Atmospheric Environment Services, and High Arctic International are also greatly appre- ciated. The help of several field assistants and of International Wolf Center observers is also gratefully acknowledged. C. S. Asa and T. J. Meier suggested improvements to the manuscript. NOTES 345 Literature Cited Bridges, R.S., R. DiBiase, D. D. Loundes, and P. C. Doherty. 1985. Prolactin stimulation of maternal behavior in female rats. Science 227: 782-784. Cook, R.S. 1993. Ecological issues on reintroducing wolves into Yellowstone National Park. Scientific Monograph NPS/NRYELL/NRSM-93/22. U.S. Department of the Interior, National Park Service. 328 pages. Dixson, A. F., and L. George. 1982. Prolactin and parental behaviour in a male New World primate. Nature 299: 551-553. Fentress, J. C., and J. Ryon. 1982. A long-term study of distributed pup feeding in captive wolves. Pages 238-260 in Wolves of the World. Edited by F. H. Harrington, and P. C. Paquet. Noyes Publications, Park Ridge, New Jersey. Gubernick, D. J., and R. J. Nelson. 1989. Prolactin and paternal behavior in the biparental California mouse, Peromyscus californicus. Hormonal Behavior 23: 203-210. Kreeger, T. J., U.S. Seal, Y. Cohen, E. D. Plotka, and C.S. Asa. 1991. Characterization of prolactin secretion in gray wolves (Canis lupus). Canadian Journal of Zoology 69: 1366-1374. Mech, L. D. 1988. The Arctic Wolf: Living with the pack. Voyageur Press. Stillwater, Minnesota. 128 pages. Mech, L. D. 1993. Resistance of young wolf pups to inclement weather. Journal of Mammalogy 74: 485-486. Mech, L. D. 1995. A ten-year history of the demography and productivity of an arctic Wolf pack. Arctic 48: 329-332. Mech, L. D. in press. Summer movements and behavior of a pup-free arctic wolf, Canis lupus, pack. Canadian Field-Naturalist. Mech, L. D., and U. S. Seal. 1987. Premature reproduc- tive activity in wild wolves. Journal of Mammalogy 68: 871-873. Mech, L. D., and J. M. Packard. 1990. Possible use of Wolf, Canis lupus, den over several centuries. Canadian Field-Naturalist 104: 484-485. Mech, L. D., T. J. Meier, and U. S. Seal. 1993. Wolf nip- ple measurements as indices of age and breeding season. American Midland Naturalist 129: 266-271. Numan, M. 1988. Maternal behavior. Pages 1569-1645 in The physiology of reproduction. Edited by E. Knobil, and J. Neill. Raven Press, Ltd., New York. Seal, U. S., E. D. Plotka, J. M. Packard, and L. D. Mech. 1979. Endocrine correlates of reproduction in the wolf. Biology of Reproduction 21: 1057-1066. Zarrow, M. X., R. Gandelman, and V. H. Denenberg. 1971. Prolactin: is it essential hormone for maternal behavior in the mammal? Hormonal Behavior 2: 343-354. Received 17 August 1995 Accepted 4 December 1995 346 THE CANADIAN FIELD-NATURALIST Vol. 110 Evidence of Successful Chinook Salmon, Oncorhynchus tshawytscha, Spawning in the St. Lawrence River, near Cornwall, Ontario SANDRA C. RIBEY and FRANCOIS CHAPLEAU Ottawa-Carleton Institute of Biology, Department of Biology, University of Ottawa, P.O. Box 450, Station A, Ottawa, Ontario KIN 6N5 Ribey, Sandra C., and Fran¢gois Chapleau. 1996. Evidence of successful Chinook Salmon, Oncorhynchus tshawytscha, spawning in the St. Lawrence River, near Cornwall, Ontario. Canadian Field-Naturalist 110(2): 346-347. On 7 and 20 June 1994, two young (36.8 and 42 mm) Chinook Salmon (Oncorhynchus tshawytscha) were captured in the St. Lawrence River, just down-stream from Cornwall, Ontario. The specimens are attributed to natural reproduction. This is the first record of Chinook Salmon successful spawning in the St. Lawrence River and the easternmost documentation of natural reproduction of this species in the Great Lakes-St. Lawrence System. Key Words: Chinook Salmon, Oncorhynchus tshawytscha, natural reproduction, St. Lawrence River, Cornwall. The Chinook Salmon (Oncorhynchus tshawytscha), a species native to the Pacific Ocean, was first introduced into the Great Lakes basin in 1873 (Mills et al. 1993). A spectacular sport fishery now exists for this species in Lake Ontario (Smith 1985). Although some natural propagation is sus- pected, self-perpetuation is minimal and intense stocking programs have been in place since 1967 to maintain the fishery (Mills et al. 1993). Some natural reproduction has been recorded in the tributaries of the Salmon River in New York State (Smith 1985). In eastern Canada, voucher specimens (Royal Ontario Museum) of young chinook suspected to be the result of natural reproduction have been taken from several streams and rivers along the north shore of Lake Ontario. The easternmost record of chinook spawning in Canada is in Shelter Valley Creek (near Cobourg, Ontario). No record of successful spawn- ing exists for the St. Lawrence River. On 7 and 20 June 1994, two young Chinook Salmon (36.8 and 42 mm, respectively) were seined (minnow seine, 10 m) on the north shore of the St. Lawrence River near Cornwall, Ontario (Figure 1). Fish were identified using the key to young salmonids in McPhail and Lindsey (1970). In addition to basic salmonid features, both fish had 19 anal-fin rays (typical of Pacific salmons), a series of parr marks (8 and 10, respectively) taller than the vertical eye diameter and an adipose fin with a clear unpigmented area. The combination of the latter two characters is typical for young Chinook Salmon. The first fish was caught in the morning of 7 June on the eastern shore of Flanigan’s Point in Glen Walter, approximately 5 km east of Cornwall (45°02'00"N; 74°38'03”W). It was captured on a sandy substrate sprinkled with a few large rocks and macrophytes. Seven other species were captured in the same haul: Fundulus diaphanus (2 specimens), Etheostoma exile (3), Notemigonus crysoleucas (50), Pimephales notatus (59), Notropis heterodon (1), Notropis volucellus (2) and Notropis heterolepis (1). The second specimen was captured in the after- noon of 20 June on the north shore of the St. Lawrence River, just east of the city of Cornwall, approximately 1.6 km upstream of the mouth of Grays Creek (45°01'30"N; 74°40'54”W). The habi- tat consisted of a rubble and gravel substrate with dense algae cover on the rocks. The following fish were captured in the same seine haul: Fundulus diaphanus (2), Etheostoma exile (4), and Pimephales notatus (28). Smith (1985) indicates that spawning occurs in August and September in New York State. Scott and Crossman (1973) indicate that freshwater popula- tions may spawn in rivers flowing into lakes or on gravel shoals in the lake. Eggs (diameter: 6-7 mm; 8 000-13 600) hatch the following spring into alevins, stay two or three weeks in the nest and then become free-swimming (Scott and Crossman 1973). Adult Chinook Salmon have been caught in the Cornwall area. A large gravid female (89 cm; 9.3 kg; 5 years old) caught in November 1978 near Cornwall (45°02'N; 74°44’W) is preserved in the fish collec- tion of the Canadian Museum of Nature (NMC 78- 0372). No stocking of Chinook Salmon has been done in the St. Lawrence River and the sampling sites are situated at approximately 150 km from the east end of Lake Ontario. The small size of our spec- imens and the date of capture makes it impossible to infer an origin from fish that have spawned in the tributaries of Lake Ontario. Consequently, it is con- cluded that both specimens are direct evidence of successful natural reproduction which occurred in the fall of 1993. The location of the spawning ground is unknown but it seems most parsimonious to infer that it has occurred somewhere along the north shore of the St. Lawrence River near Cornwall. Dumont et al. (1988) indicated that the furthest downstream report of a Chinook Salmon in the St. Lawrence River was at Pointe-au-Pére, near Rimouski, in 1986. Spawning of Chinook Salmon 1996 NOTES 347 FiGuRE 1. Young Chinook Salmon (Oncorhynchus tshawytscha) captured in the St. Lawrence River, near Cornwall (NMC 95-002; 36.8 mm). was observed in the St. Lawrence River, 30 km upstream from Montreal (Dumont et al. 1988) but no evidence of successful reproduction (survival of young) was reported. This is, therefore, the first doc- umented record of Chinook Salmon natural repro- duction in the St. Lawrence River and the eastern- most of successful natural reproduction for this species in the Great Lakes-St. Lawrence System. Acknowledgments These specimens were captured during a fish sur- vey done in the St. Lawrence River, near Cornwall, as part of a large multidisciplinary project funded by a tri-council grant dealing with Ecosystem Recovery on the St. Lawrence, obtained and administered by the Institute for Research on Environment and Economy at the University of Ottawa. Both speci- mens were deposited in the fish collection of the Canadian Museum of Nature (NMC 95-002 (36.8 mm specimen), NMC 95-003 (42 mm speci- men). Lara L. Ridgway, Lee Willard, Martin D. Lemay and David Bajurny participated in the field- work. Courtaulds Fibres Canada allowed us to live in one of their cottages while working in Cornwall. The St. Lawrence River Institute of Environmental Sciences (Cornwall) provided us with logistical help during the field season. Erling Holm (Royal Ontario Museum) confirmed the identification of one (NMC 95-002) specimen. Our most sincere thanks to all the above-mentioned people and institutions for their help. Literature Cited Dumont, P., J. F. Bergeron, P. Dulude, Y. Mailhot, A. Rouleau, G. Ouellet, and J. P. Lebel. 1988. Introduced salmonids: Where are they going in Quebec watersheds of the Saint-Laurent River? Fisheries 13: 9-17. McPhail, J. D., and C. C. Lindsey. 1970. Freshwater fish- es of northwestern Canada and Alaska. Fisheries Research Board of Canada Bulletin‘ Number 173. 381 pages. Mills, E. L., J. H. Leach, J. T. Carlton, and C. L. Secor. 1993. Exotic species in the Great Lakes: A history of biotic crisis and anthropogenic introductions. Journal of Great Lakes Research 19: 1-54. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin Number 184. Ottawa, Ontario. 966 pages. Smith, C. L. 1985. The inland fishes of New York State. The New York State Department of Environmental Conservation. Albany, New York. 522 pages. Received 3 August 1995 Accepted 11 October 1995 348 THE CANADIAN FIELD-NATURALIST Vol. 110 Apparent Longevity Records for Red Foxes, Vulpes vulpes, in Labrador Tony E. CHUBBS!, and FRANK R. PHILLIPS2 'Department of National Defence, 5 Wing Goose Bay, Postal Station A, Happy Valley - Goose Bay, Labrador, Newfoundland AOP 1S0 2Newfoundland-Labrador Tourism Division, Government of Newfoundland and Labrador, Box 3027, Station B, Goose Bay, Labrador, Newfoundland AOP 1E0 Chubbs, Tony E., and Frank R. Phillips. 1996. Apparent longevity records for Red Foxes, Vulpes vulpes, in Labrador. Canadian Field-Naturalist 110(2): 348-349. Two Red Foxes (Vulpes vulpes), trapped in November 1994 near Goose Bay, Labrador, were aged at 8 years 7 months and 10 years 8 months. These two ages exceed the oldest record in the literature by | month to 2 years 3 months, respectively. Key Words: Red Fox, Vulpes vulpes, age, annuli, Labrador. Red Foxes, Vulpes vulpes, are important to wildlife managers because of their ecological role as predators, their aesthetic, recreational and fur value, and their role in disease transmission (Allen and Sargeant 1993). Although researchers have concen- trated on feeding habits (Sklepkovych 1994), repro- ductive patterns (Smits and Slough 1993), and dis- persal (Allen and Sargeant 1993), most studies have recorded very low frequencies (< 3 percent) of foxes in older (> 5 years) age classes. Red Foxes have been commonly aged at 5 to 7 years (Allen 1984; Allen and Sargeant 1993) and some have exceeded 7 years. Storm et al. (1976) found in their study of a midwest fox population only two (1M, 1F) foxes to survive 6 years and only 3 percent of 1987 Red Foxes were trapped 3 to 6 years after tagging. Allen and Sargeant (1993) reported Red Foxes being recovered up to 8.6 years after being tagged as pups, of which four foxes (3F,1M) were recovered >7 years after being tagged. It appears that the 8.5 year old female fox (Matsons 1991*) recovered by Tullar (1983), 7 years after tagging, is the maximum reported age of a live captured Red Fox. Female foxes surviving > 2 years may be especially important to populations because of age-related increases in reproductive performance (Allen 1984; Allen and Sargeant 1993). On 12 November 1994, a cross colour phase, female fox was snared by Phillips near Canadian Forces Base, Goose Bay, in south-central Labrador (53°22.44'N, 60°27.27'W). Seventeen days later on 29 November 1994, Phillips snared a silver colour phase, male fox approximately 15 m from the first capture location. Both foxes, especially the male, exhibited excessive tooth wear. The skulls of both specimens were retained. Lower canines from both foxes were extracted and aged using cementum annuli (Matson’s, Milltown, Montana). Ages as of *See Documents Cited section before Literature Cited. their last birthdays were reliably determined at 8 years for the female and 10 years for the male (Matson 1991). Both age determinations were given a “A” rating (90% accuracy) or + | year for the 8 to 15 year age category. If we arbitrarily assign an average birth date for fox cubs in this area of 1 April, the ages of the female and male at death were 8 years 7 months and 10 years 8 months, respectively. Mounted slides displaying longitudinal tooth sec- tions of each canine (Catalogue Number NFM MA- 80.1 [female cross]) and NFM MA-81.1 [male sil- ver]) and the skulls (Catalogue Number NFM MA- 80.2 [female cross] and NFM MA-81.2 [male sil- ver]) have been deposited in the Newfoundland Museum of Natural History. Acknowledgments Publication costs were provided by the Department of National Defence through the Low- Level Flying, Wildlife Monitoring and Mitigation Program. The Department of Natural Resources, Newfoundland-Labrador Wildlife Division provided the necessary funding for laboratory analysis. We give special thanks to J. Maunder of the Newfoundland Museum of Natural History for his continued support and advice. Document Cited (marked * in text) Matson, G. 1991. Age determination by tooth cementum analysis - Histological preparations. Unpublished Progress Report Number 12. 5 pages. Literature Cited Allen, S. H. 1984. Some aspects of reproductive perfor- mance in female Red Fox in North Dakota. Journal of Mammalogy 65: 246-255. Allen, S. H., and A. B. Sargeant. 1975. A rural mail car- rier index of North Dakota Red Foxes. Wildlife Society Bulletin 3: 74-77. Allen, S. H., and A. B. Sargeant. 1993. Dispersal patterns of Red Foxes relative to population density. Journal of Wildlife Management 57: 526-533. 1996 Andrews, R. D., G. L. Storm, R. L. Phillips, and R. A. Bishop. 1973. Survival and movements of transplanted and adopted red fox pups. Journal of Wildlife Management 37: 69-72. Sargeant, A. B. 1972. Red fox spatial characteristics in relation to waterfowl predation. Journal of Wildlife Management 36: 225-236. Sargeant, A. B., S. H. Allen, and D. H. Johnson. 1981. Determination of age and whelping dates of live red fox pups. Journal of Wildlife Management 45: 760-765. Sklepkovych, B. 1994. Arboreal foraging by Red Foxes, Vulpes vulpes, during winter food shortage. Canadian Field-Naturalist 108(4): 479-481. Smits, C. M. M., and B. G. Slough. 1993. Abundance NOTES 349 and summer occupancy of Arctic Fox, Alopex lagopus, and Red Fox Vulpes vulpes, dens in the northern Yukon Territory, 1984-1990. Canadian Field-Naturalist 107(1): 13-18. Storm, G. L., R. D. Andrews, R.L. Phillips, R. A. Bishop, D. B. Siniff, and J. R. Tester. 1976. Morphology, reproduction dispersal and mortality of midwestern red fox populations. Wildlife Monograph 49. 82 pages. Tullar, B. F. 1983. An unusually long-lived red fox. New York Fish and Game Journal 30: 227. Received 26 September 1995 Accepted 16 January 1996 Observation of Repeated Use of a Wolverine, Gulo gulo, Den on the Tundra of the Northwest Territories JOHN LEE! and ALLEN NIPTANATIAK2 1Department of Renewable Resources, Government of Northwest Territories, 600, 5102-50 Avenue, Yellowknife, Northwest Territories X1A 3S8 2Coppermine, Northwest Territories XOE OEO Lee, John, and Allen Niptanatiak. 1996. Observation of repeated use of a Wolverine, Gulo gulo, den on the tundra of the Northwest Territories. Canadian Field-Naturalist 110(2): 349-350. The use of the same small area for denning by one or more Wolverines over three consecutive years is reported. Key Words: Wolverine, Gulo gulo, denning, Northwest Territories. Although several studies have occurred in recent years (Hornocker and Hash 1981; Magoun 1985; Gardner 1985; Whitman et al 1986; Banci 1987) there are many gaps in our understanding of Wolverine ecology. For instance, little is known of Wolverine denning requirements or of the distribu- tion of dens on the landscape (Banci 1994). Wolverine dens can vary from simple rest beds to complex natal dens with extensive tunnel networks (Pulliainen 1968; Magoun 1985). Natal dens are found in a variety of habitats, and in tundra and alpine situations are frequently associated with rocky scree slopes and large snowdrifts (Magoun 1985; Bevanger 1992). Although Inuit hunters have report- ed active Wolverine dens in the same location over several years (A. Niptanatiak, unpublished data), we are unaware of published reports describing annual den or denning area fidelity. We report on the repeat- ed use of a Wolverine den site over three years. We located a Wolverine den in April 1993 on the tundra of the Northwest Territories, approximately 160 km southeast of Coppermine (latitude 66°30'N, longitude 113°00’W). The den was in a large hard- packed snowdrift on the lee side of a south easterly facing rock outcrop. The snowdrift extended from the top of the rock face to a take shore, a vertical dis- tance of approximately 25 m and a horizontal dis- tance of 100 m. There was an opening at the very top of the drift and one lower down the slope approxi- mately 75 m away, almost at the lake shore. Several sets of fresh Wolverine tracks came and went from both openings. Many sets of older tracks were evi- dent in the vicinity. We visited the site briefly every few days over the next three weeks and always encountered fresh tracks. In March 1994 we revisited the area and found the den again active. The two den openings were in the same positions as in 1993 and Wolverine tracks were evident entering and leaving both openings. Tracks of various ages were present in the area. We fol- lowed a set of fresh tracks leading from the den and captured an adult female Wolverine. As the Wolverine was pursued, she headed directly toward the den and was captured less than 1 km from it. She did not appear to be lactating or pregnant. There was no way of knowing if this Wolverine was the same animal which used the den in 1993. In August 1994, we again visited the area. With snow absent, the ground beneath the rock face was a jumble of angular boulders varying in diameter from about 40 cm to several metres. Under some of the overhangs created by the boulders was evidence of rudimentary beds pressed in the lichen and grass. Scattered between the positions where the upper and 350 lower openings had been were several places where scats and food remains were concentrated. The remains varied in age from recent to very old, suggest- ing that this site may have been used over many years. In March 1995, we again visited the site. A den was located approximately 300 m north of the origi- nally described den on the same rock face. No Wolverines were observed but fresh tracks and numerous older tracks were observed. Only one opening was evident. The numerous tracks in the vicinity, the repeated use of the same area for denning over three years of observation, the apparent extent of the den, and the presence of an adult female would suggest that this may have been a natal den site. Denning habitat on the Canadian tundra should not be limited because rocky outcrops and extensive snowdrifts abound. Although we were unable to con- firm the den was used by the same Wolverine in all three years, this den site was used each spring. On the tundra, Arctic Foxes (Alopex lagopus) (Macpherson 1969) and Wolves (Canis lupus) (Mech and Packard 1990) often reuse dens in successive years. In northwestern Alaska, Magoun (1985) located several Wolverine natal dens, four of which were used by the same female over four years. Although these four dens were not in the exact location, they were within | to 2 km of each other, or all within approximately 4 km? (Magoun, personal communi- cation 1995). In one year no kits were produced and the den was abandoned. In the years when kits were present, this female’s home range during the den- ning/summer period was calculated at 56 and 99 km? (Magoun 1985). The area chosen for denning, conse- quently, represented about 4% to 7% of her home range. Considering that the home ranges of adult females with young may be exclusive (Magoun 1985) and that the number of adult males are related to the number of females (Magoun 1985; Banci 1987), the number and distribution of natal dens could have potential to provide an indication of the density of resident Wolverines on the tundra. In view of increased mineral exploration occurring on the tundra of the Northwest Territories, questions of Wolverine den site fidelity are pertinent. Answers THE CANADIAN FIELD-NATURALIST Vol. 110 to such questions are just a small part of Wolverine ecology that should be understood to allow assess- ment and mitigation of impacts as development pro- ceeds. Acknowledgments I thank J. Nishi for his contributions toward logistics and funding and K. Poole for his helpful comments on the manuscript. Literature Cited Banci, V. A. 1987. Ecology and behaviour of wolverines in Yukon. Upublished M.Sc. Thesis, University British Columbia, Vancouver. 178 pages. Banci, V. A. 1994. Wolverine. Pages 99-127 in The Scientific Basis for Conserving Forest Carnivores; American Marten, Fisher, Lynx, and Wolverine in the Western United States. USDA Forest Service General Technical Report RM-254. Bevanger, K. 1992. Report on the Norwegian wolverine (Gulo gulo). \UCN Small Carnivore Conservation 6: 8-10. Gardner, C. L. 1985. The ecology of wolverines in southcentral Alaska. Upublished M.Sc. Thesis, University Alaska, Fairbanks. 82 pages. Hornocker, M. G., and H.S. Hash. 1981. Ecology of the wolverine in northwestern Montana. Canadian Journal of Zoology 50: 1286-1301. Magoun, A. J. 1985 Population characteristics, ecology, and management of wolverines in northwestern Alaska. Unpublished Ph.D. thesis, University of Alaska, Fairbanks. 197 pages. Macpherson, A. H. 1969. The dynamics of Canadian arc- tic fox populations. Canadian Wildlife Service Report Series number 8. 52 pages. Mech, L. D., and J. M. Packard. 1990. Possible use of wolf (Canis lupus) den over several centuries. Canadian Field-Naturalist 104: 484-485. Pulliainen, E. 1968. Breeding biology of the wolverine (Gulo gulo) in Finland. Annales Zoologica Fennici 5: 338-344. Whitman, J.S., W. B. Ballard, and C. L. Gardner. 1986. Home range and habitat use by wolverines in southcentral Alaska. Journal of Wildlife Management 50: 460-463. Received 6 October 1995 Accepted 23 January 1996 ———S—————————— 1996 NOTES 351 Home Range Size of Bushy-tailed Woodrats, Neotoma cinerea, in Southwestern Alberta MICHAEL G. TOPPING and JOHN S. MILLAR Ecology and Evolution Group, Department of Zoology, University of Western Ontario, London, Ontario N6A 5B7 Topping, Michael G., and John S. Millar. 1996. Home range size of Bushy-tailed Woodrats, Neotoma cinerea, in south- western Alberta. Canadian Field-Naturalist 110(2): 351-353. The home range sizes of Bushy-tailed Woodrats (Neotoma cinerea) in the Canadian Rockies, were measured over a three year period (1992-1994) by means of radio telemetry. Males had significantly larger mean home ranges than females (6.12 ha and 3.56 ha respectively), but there were no differences within each sex between years. Home range sizes from this study were >20 times larger than home ranges reported for other species of Neotoma. The proportionately larger home ranges of Bushy-tailed Woodrats may be due to their reliance on fissures within rocky outcrops for den sites. Key Words: Bushy-tailed Woodrat, Neotoma cinerea, den site, home range, radio telemetry, Alberta. Woodrats (Neotoma spp.) occupy habitats ranging from boreal woodland to desert and thornscrub (Vaughan 1990). Two habitat requirements common to all members of the genus are succulent plant food (at least 50% water by weight) and shelter, provided by rocks, shrubs or trees (Finley 1958; Vaughan 1990). Energy conserving adaptations such as small litter size (Vaughan and Czaplewski 1985), late matu- ration (Egoscue 1962) and storage of food (Atsatt and Ingram 1983) are common in this genus. Reliance on shelter provides a further method of conserving ener- gy in woodrats. The majority of Neotoma species con- struct nests from sticks, twigs, leaves and bones (Finley 1958) which provides protection from both predators and temperature extremes (Brown 1968; Vaughan 1990). This reliance on a nest and protected den site has resulted in the majority of Neotoma species exhibiting a den-centered ecology, with small home ranges and central place foraging (Raun 1966; Finley 1990). However, the majority of studies on for- aging ecology of Neotoma have been performed on species inhabiting the central or southern areas of North America (Fitch 1947; Murphy 1952; Hamilton 1953; Finley 1958; Findley et al. 1975). As such, little is known about the home range size and foraging ecology of the more northern species of woodrat, Neotoma cinerea. The Bushy-tailed Woodrat (Neotoma cinerea) is the largest and most northerly living of the genus, reaching a weight of up to 500g and inhabiting the mountainous regions of Western North America, as far north as the southern Yukon, and as far south as northern New Mexico (Finley 1958). They utilize fis- sures in rocky outcrops for den sites (Finley 1958; Escherich 1981; Hickling 1987). We measured the home range sizes of Bushy-tailed Woodrats over three breeding seasons in the Kananaskis Valley, southwestern Alberta (51° N, 115° W). Woodrats from four adjacent rocky outcrops (elevation: 1670 m.a.s.l.) were monitored by weekly live trapping and radiotelemetry between late April and August of 1992-1994. Outcrops were surrounded by mature mixed coniferous (White Spruce, Picea glauca, Douglas Fir, Pseudotsuga menziesii, and Lodgepole Pine, Pinus contorta) forest with a mixed shrub understory (Bearberry, Arctostaphylos uva-ursi, Common Juniper, Juniperus communis; Buffalo- berry, Sheperdia canadensis; and Ribes spp.). Positional data were gathered from live trapping and radio telemetry. Tomahawk live traps were set at fixed locations along the base of each outcrop every week. Depending on the length of the outcrop (170 — 300 m) 20-40 traps were set each night. Live trap- ping provided demographic data, and allowed radio- collars (Model No. PD-2C, Holohil Systems Ltd., Ontario) to be fitted to reproductively active individ- uals. Radio telemetry was carried out once per week on each outcrop, from 0000 —.0400 hrs. We scanned the range of radiocollar frequencies at 10 minute intervals, and recorded data on the signal strength, and direction from a telemetry station of known and fixed position. The receiver was calibrated for dis- tance, allowing the position of the animal to be plot- ted on a scale map of the study area. By plotting an observation area curve, we deter- mined that a minimum of 14 independent locations were required to accurately represent the home range size. Therefore, we only used data from individuals that had 14 or more locations. We determined home range size for 5 males and 9 females in 1992, 8 males and 9 females in 1993, and 10 males and 8 females in 1994 using the Minimum Convex Polygon method, from the RANGES IV program (Kenward 1990). In all years, males had significant- ly larger home ranges than females (F=9.31; df=1,43; p=0.004), but home ranges within each sex did not differ between years (F=2.14; df=2,43; p=0.130) (Table 1). In comparison to home range sizes from other species of Neotoma, which range from 0.1-1.08 ha 352 TABLE |. Mean (+SE) home range sizes of male and female Bushy-tailed Woodrats during 1992-1994. Sample sizes are in parentheses; minimum and maximum are shown below mean values. Year Male Home Female Home Range size (ha) Range size (ha) 1992 5.49 + 1.48 (5) 3.13 + 0.65 (9) (1.59 - 9.18) (1.65 - 6.90) 1993 5.10 + 0.84 (8) 3.15 + 0.82 (9) (1.93 - 10.06) (0.13 - 7.43) 1994 7.26 + 0.70 (10) 4.44 + 1.33 (8) (3.84 - 11.19) (0.91 - 10.44) (Johnson 1952; Goertz 1970; Cranford 1977; Vaughan and Schwartz 1980; Kelly 1989; Lynch et al. 1994), home range sizes from this study were much larger (Table 2). This variation in home range sizes within Neotoma species may be a reflection of the productivity of the habitat, since larger home ranges are often indicative of lower productivity (Harestad and Bunnell 1979). However, no informa- tion is available on the relative degree of productivi- ty in the habitats occupied by members of the genus Neotoma, so we were unable to determine whether variation in productivity between habitats could be one cause of the variation in home range sizes. The larger home range sizes may also be a direct consequence of N. cinerea requiring existing fis- sures in rocky outcrops for den sites. Neotoma species that are able to manipulate or alter the envi- ronment in order to create den sites are not depen- dent on existing geographic structures for den sites, and may establish their nests in close proximity to other resources, resulting in small home ranges (Table 2). A suitable den site for N. cinerea may be located at considerable distances from other neces- sary resources. Therefore, N. cinerea may require large home ranges in order to have access to all requirements for survival. The benefits of occupy- ing an appropriate den site (which may be located at some distance from other resources) may out- weigh the costs of maintaining a large home range. THE CANADIAN FIELD-NATURALIST Vol. 110 Acknowledgments J. Goddard, J-P. Bechtold and N. Swerhun provid- ed valuable assistance in the field. Thanks also to T. Teferi for advice on radio telemetry at the start of this study. The University of Calgary Kananaskis Field Station provided facilities which allowed the undertaking of this study. M. Chase provided critical review and discussion on earlier drafts of this manuscript. This study was supported by a Commonwealth Scholarship to M. Topping and a Natural Sciences and Engineering Research Council of Canada grant to J. Millar. Literature Cited Atsatt, P. R., and T. Ingram. 1983. Adaptation to oak and other fibrous, phenolic-rich foliage by a small mam- mal, Neotoma fuscipes. Oecologia 60: 135-142. Brown, J. H. 1968. Adaptation to environmental tempera- ture in two species of woodrats, Neotoma cinerea and N. albigula. Miscellaneous Publications. Museum of Zoology of the University of Michigan 135: 148. Cranford, J. A. 1977. Home range and habitat utilization by Neotoma fuscipes as determined by radiotelemetry. Journal of Mammalogy 58: 165-172. Egoscue, H. J. 1962. The bushy-tailed woodrat: a labora- tory colony. Journal of Mammalogy 43: 328-337. Escherich, P. C. 1981. Social biology of the bushy-tailed woodrat, Neotoma cinerea. University of California Publications in Zoology 110: 1-132. Findley, J. S., A. H. Harris, D. E. Wilson, and C. Jones. 1975. Mammals of New Mexico. University of New Mexico Press, Albuquerque. 360 pages. Finley, R. B. 1958. The woodrats of Colorado: distribu- tion and ecology. University of Kansas Museum Natural History Publications 10: 213-552. Finley, R. B. 1990. Woodrat Ecology and Behavior and the interpretation of Paleomiddens. Pages 28-42 in Packrat Middens: the last 40,000 years of biotic change. Edited by J. L. Betancourt, T. R. Van Devender and P. S. Martin. University of Arizona Press, The Arizona Board of Regents, Arizona. Fitch, H.S. 1947. Predation by owls in the Sierran foothills of California. The Condor 49: 137-151. Goertz, J. W. 1970. An ecological study of Neotoma floridana in Oklahoma. Journal of Mammalogy 51: 94-104. TABLE 2. Mean Home Range size (ha) of males and females for five species of Neotoma. The method by which data was collected is also shown. Species Male Female Home Home Range Range N. floridana 0.26 ha 0.17 ha N. fuscipes 0.23 ha 0.19 ha 1.08 ha 0.05 ha 0.44 ha 0.36 ha N. lepida 0.34 ha 0.11-ha N. micropus 0.10 ha 0.13 ha 6.12 ha 3.56 ha N. cinerea Method Source of data collection Live Trapping Goertz (1970) Telemetry Cranford (1977) Telemetry Kelly (1989) Telemetry Lynch et al. (1994) Nocturnal tracking | Vaughan and Schwartz (1980) Live Trapping Johnson (1952) Trap/Telemetry this study 1996 Hamilton, W. J. 1953. Reproduction and young of the Florida wood rat, Neotoma f. floridana. Journal of Mammalogy 34: 180-189. Harestad, A.S., and F. L. Bunnell. 1979. Home range and body weight — a reevaluation. Ecology 60: 389-402. Hickling, G. J. 1987. Seasonal reproduction and group dynamics of bushy-tailed woodrats. Ph. D. dissertation, Zoology Deptartment, University of Western Ontario, London, Ontario. 163 pages. Johnson, C. W. 1952. The ecological life history of the packrat, Neotoma micropus, in the brushlands of Southwest Texas. M.S. thesis. University of Texas, Austin, Texas. 115 pages. Kelly, P. A. 1989. Population ecology and social organi- zation of Dusky-footed woodrats, Neotoma fuscipes. Ph. D. dissertation, Zoology Department, University of California at Berkeley, California. 191 pages. Kenward, R. 1990. RANGES IV. Software for analyzing animal location data. Institute of Terrestrial Ecology, Wareham, Dorset, U.K. Lynch, M.F., A. L. Fesnock, and D. V. Vuren. 1994. Home range and social structure of the Dusky-footed woodrat. Northwestern Naturalist 75: 73-75. NOTES 353) Murphy, M.F. 1952. Ecology and helminths of the Osage woodrat, Neotoma floridana osagensis, including description of Longistriata neotoma (Trichostrongylidae). American Midland Naturalist 48: 204-218. Raun, G. G. 1966. A population of wood rats, Neotoma micropus, in southern Texas. Bulletin of Texas Memorial Museum 11: 1-62. Vaughan, T. A. 1990. Ecology of Living Packrats. Pages 14-27 in Packrat Middens: the last 40,000 years of biot- ic change. Edited by J. L. Betancourt, T. R. Van Devender and P. S. Martin. University of Arizona Press, The Arizona Board of Regents, Arizona. Vaughan, T.A., and N.J. Czaplewski. 1985. Reproduction in Stephens’ woodrat: The wages of folivory. Journal of Mammalogy 66: 429-443. Vaughan, T. A., and S. T. Schwartz. 1980. Behavioral ecology of an insular woodrat. Journal of Mammalogy 61: 205-218. Received 22 August 1995 Accepted 18 December 1995 News and Comment Errata: The Canadian Field-Naturalist 109(3) Pringle, James S. 1995 [1996]. The history of the explo- ration of the vascular flora of Canada 109(3): 291-356 On page 333, right column, lines 49 and 50. “Live” should deleted and replaced by “Luc”. “Saucier” should be deleted and replaced by “Goulet”. On page 352, right column, line 40, “Saucier” should be deleted and replaced by “Goulet” Pringle, James S. 1995 [1996]. The history of the explo- ration of the vascular flora of Greenland. Canadian Field-Naturalist 109(3): 362-375. On page 368, two lines of text are missing from the bottom of the right column, add: “ing the 1930s have been cited by Adams and Norwell (1936) and Kleppa (1973).” Pringle, James S., Compiler. 1995 [1996]. Combined index to personal names. Canadian Field-Naturalist 109(3): 378-382. On page 379, right column, after line 42, insert Goulet, 1:333;: On page 381, right column, line 11, delete “Saucier, I. 333” Thanks to Jacques Cayouette, Agriculture and Food Canada, Ottawa. FRANCIS R. COOK Rana-Saura: Amphibian Follow-up Project: Atlas of Amphibians and Reptiles of Quebec Volume 2, number 2, December 1995, of the newsletter of the Atlas of Amphibians and Reptiles of Quebec project contains a summary of the third ses- sion (1995) results. Compared to 1994 there was a 44% increase in participation to 56 people. Also included is an analysis of road surveys in 1993 and 1994 and “End of Season” comments for 1995 with a full list of participants. The recently published Atlas of Amphibians and Reptiles of Quebec (reviewed in The Canadian Field-Naturalist 109(4): 493, 1995) is described. The newsletter is produced by the St. Lawrence Valley Natural History Society, with the assistance of the Ministére de l’Environnement et de la Faune, Quebec. Copies of the Atlas are available from the Society at $12 per copy plus $3 for postage and handling (in United States funds for U.S. orders). SYLVIE MATTE St. Lawrence Valley Natural History Society, 21 125 Che- min Ste.-Marie, Ste-Anne-de-Bellevue, Quebec H9X 3L2 *Phone: 514-457-9449. Sea Wind: Bulletin of Ocean Voice International Sea Wind, volume 9, number 3, pages 1 to 36, July-September 1995 has a feature article on “Offshore and deep-sea marine reserves and Parks: A major gap in protected area networks” by D. E. MacAllister who also contributed “Advisory body to Biodiversity Convention” covering the Subsidiary Body for Scientific, Technical and Technological Advice (SBSTTA) created to provide advice to the Conference of Parties of the Biodiversity Convention. T. E. Reimchen of the University of Victoria has written “Estuaries, energy flow, and biomass extraction in Gwaii Haanas” [South Moresby] National Park Reserve in British Columbia. Sea Wind, volume 9, number 4, October- December 1995 is a special issue on the Status of the World Ocean and its Biodiversity by D. E. McAllister covering I. Status of Biodiversity, Biological Resources and Conservation Initiatives, II. Strategies and Action. There is a seven page ref- erence section to round out this largely point-form comprehensive discussion. OCEAN VOICE INTERNATIONAL P.O. Box 37926, 3332 McCarthy Road, Ottawa, Canada K1V OWO *Phone 613-264-986 *Fax 613-264-9204 *E-mail ah 194 @freenet.carleton.ca 354 1996 NEWS AND COMMENT 355 FROGLOG: IUCN/SSC Declining Amphibian Populations Task Force The December 1995 issue (number 15) of the newsletter of the Declining Amphibians Task Force (DAPTF) sponsored by the World Conservation Union (formerly called the “International Union for the Conservation of Nature” - IUCN), Species Survival Commission (SSC), contains an update on the North American Monitoring Program, Abiotic Disturbances in the Lesser Antilles, Biological Control of the Cane Toad Bufo marinus in Australia, Observations from Long-term Population Studies in Switzerland, US Regional Group Meetings, Publications of Interest, and a list of New Working Group Chairs and some address changes. There is also a plea for donations to continue the work of the Decline in Amphibian Populations Task Force. The editor is John Baker, Department of Biology, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom. The February 1996 issue (Number 16) is entirely devoted to an overview of some of the work being conducted in Canada on amphibian populations, including a summary of the 1995 Decline in Amphibian Populations in Canada (DAPCAN) fifth annual meeting. This issue has been guest edited by Christine Bishop of the Canadian Wildlife Service who has been a major force in the success of DAP- CAN since its inception at a workshop held in Burlington, Ontario, in the fall of 1991. The May 1966 issue (Number 17) features News from South Asia, Aliens Exterminate Amphibians, DAPCAW VI and The Boreal Dip Net, US Great Lakes Declining Amphibians Conference, South- western US Working Group/NAAMP Documents Web Meeting, Earthwatch Funding, Endangered Species Bulletin, More Donations to Task Force, Meetings, Publications of Interest, and New PAPTF Working Group chair. Further information on the Declining Amphibians Task Force and copies of its newsletter FROGLOG are available from: STEPHEN CORN Task Force Chair, Midcontinent Ecological Science Center 4512 McMurry Avenue, Fort Collins, Colorado, USA 80525-3400 FROGLOG can also be found on the World Wide Web at the following URL: http://arcs-info.open.ac.uk/info/ news- letters/FROGLOG.html Bullfrog Management in Ontario: Workshop Proceedings The Southern Region Science and Technology Transfer Unit of the Ontario Ministry of Natural Resources has printed Bullfrog Management in Ontario: Workshop Proceedings. This is the result of session held 3 October 1994 at which invited presen- tations were given by Francis Cook (Bullfrog life history), Bob Johnson (Global perspective on amphibians), Michael Berrill (Ontario Bullfrog per- spective), Ron Brooks (Amphibian monitoring and Bullfrog research), Hans von Rosen (Eastern Ontario Bullfrog management perspective) and Ross Cholmondeley (Bullfrog management program sum- mary - 1994). Some highlights from the original ver- bal presentations are summarized in point form and followed by excerpts from three discussion groups on Monitoring, Research and Experimental Management Needs, and Bullfrog Harvest. The focus of the session was an examination of reported Bullfrog declines in Ontario and the possible causes and solutions. The most important resulting action in the short term was the suspension of commercial Bullfrog harvest in Ontario in 1995. The most useful inclusion is the 1994 Program Summary which makes available the results of sur- veys and questionnaires in eastern Ontario in tabular and graph form (by Ross Cholmondeley, co-editor with K. Coleman). Texts were apparently transcribed from oral presentations by uncredited Ministry staff and not proof-read by presenters, hence a few slips in delivery or transcription that otherwise might have been detected before publication have been included, such as the lapsus credited to me that there are 4000 species of amphibians in North America (a total which actually applies to the whole world). [Future printings will be corrected]. Another discor- dant note is the cover picture of a stylized frog with dorsolateral folds, a feature which is uniquely absent in Bullfrogs among Canadian ranid frogs. Copies are available from the Ontario Ministry of Natural Resources, Southern Region Science & Technology . Transfer Unit, Oxford Avenue, Box 605, Brockville, Ontario K6V 5Y8. FRANCIS R. COOK 356 Frogwatch 96 In Nova Scotia, the Frogwatch program is to be continued for a third consecutive year. In 1994 a pilot project was instigated and in 1995 this was expanded to be province-wide. Both seasoned “Peeper seekers” and newcomers in Nova Scotia are invited to participate in the 1996 efforts. These began in mid-March with the listening for, recording and reporting of Spring Peeper (Hyla crucifer) calls. This will run through to mid-June. As well, wet places where amphibians live will be studied though the summer. For more information, participation forms and kits, please contact: Frogwatch 96 c/o The Nova Scotia Museum of Natural History at address, phone, fax or e-mail given below. THE CANADIAN FIELD-NATURALIST Vol. 110 Frogwatch 96 is a partnership of the Nova Scotia Museum of Natural History, The Clean Nova Scotia Foundation, Envirosphere Consultants Ltd, and is also supported by Wildlife Habitat Canada, Canada Trust Friends of the Environment Foundation, Environment Canada, The Nova Scotia Department of Education, and Seagull Pewter. Nova ScCoTIA MUSEUM of NATURAL HISTORY 1747 Summer Street, Halifax, Nova Scotia, Canada B3H 3A6 *Phone 902-424-3563 * Fax 902-424-0560 * e-mail: Frogwatch @nsm.ednet.ns.ca The Second Annual International Symposium on Biology and Conservation of Owls of the Northern Hemisphere The Second Annual International Symposium: Biology & Conservation of Owls of the Northern Hemisphere is scheduled for 5-9 February 1997 at the Delta Winnipeg Hotel, Winnipeg, Manitoba under the auspices of the Manitoba Department of Natural Resources. International enquiries have indicated that a second symposium (the first was also sponsored by the Manitoba Department of Natural Resources in February 1987) 10 years later would provide a welcome forum for discussion of progress in biology and management of owls and their habitats and presentation of the accumulated research results. Included will be the 62 species (in 19 genera) of owls recognized as occurring in the Northern Hemisphere in K. H. Voous 1988 (Owls of the Northern Hemisphere. William Collins & Co. Ltd., London. 320 pages). Features of the meeting Second International Conference on Raptors The Raptor Research Foundation’s Second International Conference on Raptors will be held at the University of Urbino, in Urbino, Italy, 2-5 October 1996. It will cover all aspects of the study and conservation of birds of prey throughout the world. Also associated with this event is an important international symposium entitled “Raptors and Energy Development’, that will take place during the conference on 3-4 October 1996. This sympo- sium will address the interaction of birds of prey and wildlife in general with energy-related infrastructure and activities, as well as their impact, management and regulation. For additional information on this symposium contact one of its co-chairs, Thomas E. Krueger (address below) or Dr. Patricia L. Kennedy, Colorado State University, Department of Fish and will include Scientific Paper Sessions, Poster Sessions, Workshops, Field Trips, Exhibits and Owl Art Display, Films and Videos, Banquet and Social Gatherings. Blocks of rooms have been reserved at a special rate at the Delta Winnipeg Hotel. Details on program availability and preparation and submis- sion of abstracts are available form the addresses below. Dr. JAMES R. DUNCAN Chair, Program Committee Dr. ROBERT W. NERO Manitoba Conservation Data Centre, Department of Natural Resources, 200 Saulteaux Crescent, Winnipeg, Manitoba, Canada R3J 3W3 * Phone 204-945-7465 (Duncan) * 204-945-6817 (Nero) * Fax 204-945-3077 (both); e-mail owl @envirolink.org Wildlife Biology, Colorado Springs, Colorado (970- 491-6597; Fax 970-491-5091). The conference will also include a symposium on the status and conservation of Italian raptors, round-tables on various timely topics, moderated film and video sessions, local field trips, and the possibility for attendees to participate in longer cul- tural and nature tours, as well as tours of Urbino itself. Tentatively planned are the following scientific sessions along with the General Session: Applied Behavioral Ecology, Asian Raptor Studies, Conser- vation and Management Techniques, Genetics, Migration and Dispersal, Patterns of Raptor Diversity in South America, Population Ecology, and Toxicology. Papers and poster will be presented either in English and Italian. 1996 The deadline for receipt of abstracts in | August 1996. Contact the Conference Committee Chair- person Massimo Pandolfi, Instituto de Sciencze Morfologische, Via Oddi 21, 61029 Urbino, Italy *Phone +39-722-328033 *Fax +39-722-329655) for more information or to submit abstracts. NEWS AND COMMENT 35/7) THOMAS E. KRUEGER, JR. Scientific Director, The Italian Raptor Association, Via Eulambio, 14/A, 34072 Gradisca d’Isonzo (Gorizia), Italy *Phone: +39-481-960918 *Fax: +39-481-99876 Recovery of Nationally Endangered Wildlife in Canada (RENEW) The latest recovery plan for an endangered Canadian species was mailed in February 1996. Earlier notices for this series appear in The Canadian Field-Naturalist 109(1): 124 (numbers 1 to 11) and 109(2): 266 (numbers 12 and 13). The new report is: National Recovery Plan for the Newfoundland Martin. RENEW Report Number 14. 29 pages. dated “August 1995”. RENEW: Fifth Annual Report 1994/1995 The Fifth Annual Report 1994/95 of the Recovery of National Endangered Wildlife (RENEW) [Canada] is now available from the Canadian Wildlife Service (34 pages). It contains an explanation of what RENEW is, a From the Chair introduction by David R. Bracket; RENEW at the Crossroads; Cross-Canada check-up: species with and species without approved recovery plans; Finance: funding by species, list of donors; Profiles: Dale Hjertaas; Theresa Aniskowicz, Cormick Gates; The recovery teams; For more infor- mation (where and how to obtain, by province); RENEW members (federal and provincial depart- Available from: Recovery of National Endangered Wildlife, Ottawa, Ontario K1A OH3. CANADIAN WILDLIFE FEDERATION 2740 Queensview, Ottawa, Ontario, Canada K2B 1A2 ments and societies). An insert is titled “1994 List of Species at Risk Designated by the Committee on the Status of Endangered Wildlife in Canada”, and con- tains all species designated Extinct, Extirpated, Endangered, Threatened, and Vulnerable up to, and including, the annual meeting of COSEWIC annual meeting of two years ago. SIMON NADEAU RENEW Secretariat, c/o Canadian Wildlife Service, Environment Canada, Ottawa, Ontario, Canada K1A 0H3 Canadian Wildlife Service LRTAP Biomonitoring Program Part 2, Food chains monitoring in Ontario lakes: _ Taxonomic codes and collections by D. K. McNicol, M.-L. Mallory, and B. E. Bendell. 1996. Canadian Wildlife Service Technical Report Series number 246. 32 pages. This is the second in a series describing ongoing research and monitoring activities of the Canadian Wildlife Service Long Range Transport of Air Pollutants (LRTAP) Biomonitoring Program from 62 small lakes and wetlands in the Algoma, Muskoka, and Sudbury regions of Ontario sampled on a regular basis for various prey of resident water- fowl. Many of these prey are acid-sensitive and absent from lakes degraded by acid precipitation. ~ This report contains information on locations, meth- ods, and timing of collections and summarizes the macroinvertebrate, fish, and amphibian taxa taken between 1987 and 1994. There is a valuable biogra- phy in which 25 of 45 items included are earlier aspects of this large study coauthored by one or more of the present authors. Other reports in this series include: The Canadian Wildlife Service LRTAP Biomonitoring Program Part 1. A strategy to monitor the biological recovery of aquatic ecosystems in Canada from the effects of acid rain. Canadian Wildlife Service Technical Report Series number 245. 28 pages. The Canadian Wildlife Service LRTAP Biomonitoring 358 Program Part 3. Site locations, physical, chemical and biological characteristics. Canadian Wildlife Service Technical Report Series number 248. For more information on the CWS LRTAP Biomonitoring Program or to obtain copies of any of the reports in the series please contact: THE CANADIAN FIELD-NATURALIST Vol. 110 DONALD K. McCNICOL Environment Canada, Environmental Conservation Branch, Canadian Wildlife Service, Ontario Region, 49 Camelot Drive, Nepean, Ontario, Canada K1A 0H3 Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) One of the outstandingly successful and continu- ing initiatives in advancing the description and anal- ysis of Canada’s fauna is the Biological Survey of Canada (Terrestrial Arthropods) which coordinates research projects and effectively serves as communi- cation centre for those conducting original work on arthropods in Canada. Its latest newsletter (volume 15, number 1, spring 1996) includes a summary of a meeting of the Scientific Committee for the Biological Survey of Canada (Terrestrial Arthropods, October 1995; a project update: arctic The Boreal Dip Net Volume 1, Number 1, Winter-Spring 1996: the new Newsletter of the Working Group on Amphibian and Reptile Conservation in Canada was mailed out in March. The “Working Group” is an expansion of the IUCN/SSC Task Force on Declining Amphibian Populations in Canada (DAP- CAN) in order to include the study of reptiles as well as amphibians. DAPCAN becomes a subgroup of the new “Working Group”. Previously the Canadian Association of Herpetologists (CAH) has included DAPCAN activities in their newsletter which became, informally, the publication of DAPCAN. The new newsletter is an independent venture. The first meeting of the new Working Group on Amphibian and Reptile Conservation in Canada will be held on 5-7 October 1996 in conjunction with the IUCN/SSC Task Force on Declining Amphibians in Canada at the University of Calgary, Alberta. The program will include a symposium on the The Ontario Chorus Volume 1, Number 2, February 1996, the second issue of a newsletter for participants in volunteer amphibian monitoring programs in Ontario, has been distributed by Environment Canada. The intent of this publication is to report highlights of the various monitoring programs and focus on issues of concern to Ontario herpetologists, environmentalists and pro- gram volunteers. This issue features articles titled “Volunteer Amphibian Monitoring Programs Gain invertebrate zoology; a list of selected acronyms; a list of requests for material or information required for studies of the Canadian fauna 1996, and many other news and general interest items. Copies of the newsletter can be obtained from: H. V. DANKS Head, Biological Survey of Canada (Terrestrial Arthro- pods), Canadian Museum of Nature, P.O. Box.3443, Station D, Ottawa, Ontario, Canada K1IP 6P4 Conservation of Reptile Populations in Canada and a workshop on Herpetological Atlasing. For more information contact: STAN A. ORCHARD Chairman and DAPCAN National Co-ordinator, 1745 Bank Street, Victoria, British Columbia, Canada V8R 4V7 *Phone/Fax 604-595-7556 * E-mail: sorchard @islandnet.com CAROLYN SEBURN Vice-chairman and DAPCAN Co-ordinator for Eastern Canada, 930 River Road, RR#4, Kemptville, Ontario, Canada KOG 1JO * Phone/Fax 613-258-6142 * E-mail: aw964 @freenet.carleton.ca ANDREW DIDIUK Vice-chairman and DAPCAN Co-ordinator for Western Canada, Saskatchewan Monitoring Project and Saskatchewan Herpetology Atlas Project, 314 Egbert Avenue, Saskatoon, Saskatchewan, Canada S7N 1X1 * Phone 306-975-4087 * Fax 306-975-4089 * E-mail: did- iuka @ desoto.wxe.sk.doe.ca Momentum”, “Volunteering in Great Lakes Marshes”, “Ontario Hosts Amphibian Population Conferences” and “The Appreciation Corner”, the last listing participants in 1995 of two programs: “Road Call Counts” and “Backyard Surveys”. CANADIAN WILDLIFE SERVICE Canada Centre for Inland Waters, P.O. Box 5050, 867 Lakeshore Road, Burlington, Ontario, Canada L7R 4A6 1996 Missouri Botanical Garden 1996 Update The 1996 AABGA Annual Conference “Roots in the Past, Routes to the Future” will be hosted 30 May - | June. A major question to be addressed is the readiness for the Information Highway, the Internet and Virtual Gardening. Ground will be broken this spring for the Gardens’s $19.4 million research center, to be con- structed with special “base isolator” technology to prevent earthquake damage. This building will house the herbarium and library, provide adequate working space for researchers, and serve as a major interna- tional resource of scientific information for corpo- rate, government and university researchers working in biotechnology and related fields. A new Applied Research Department will be head- ed by Dr. James Miller, formerly assistant head of the Madagascar Department. The new department’s activities will include “bioprospecting” projects for the U.S. National Cancer Institute and Monsanto Corporation, providing both organizations with plant samples for testing for medical, anti-fungal, and anti- insect properties. Another program is the creation of a bank of plant samples for DNA extraction. An index will be available on the Internet to encourage use of the bank by molecular researchers. The Flora of Venezuelan Guayana, the initial two volumes of a projected 11, was co-published with the Timber Press, Inc. in late 1995. The Flora of North America, will be supported by an continuing grant awarded by the National Science Foundation of nearly $1 million over the next 3 years. When complete it will include 14 printed vol- umes, a CD-ROM, a new regional database, and a Global Biodiversity Volume 5, Number 4, Spring 1996, contains 48 pages and includes papers on “Cop-1: The NGO per- spective” by Elizabeth May; “Deforestation - out of control in Venezuela” by Julio Cesar Centeno; “The re-emergence of fibre hemp in Canada by Sean Twomey; “Linking gastronomic sin and environ- mental virtue: Growing bananas and chocolate organically in Costa Rica” by William O. McLarney; “Spanish national inventory on road mortality of ver- tebrates” by Javier Caletrio, Jose Maria Fernandez, Javier Lopez, and Federico Roviralta. “A portrait of biodiversty” pictures a wasp of the genus Evania with a text of fact on the relevance and diversity of its group. A Views section features “Evangelicals urge [U.S.] Congress to strengthen the Endangered Species Act”. A Biotech Corner contains “Indigenous person from Papua New Guinea claimed in U.S. government patent” by Pat Mooney, NEWS AND COMMENT 359 variety of information on Internet, including a full- feature WWW site and easy access to the Flora database. The Flora of Chile was published in August 1995 by the University of Conception Press in Conception, Chile, in collaboration with internation- al coordinators which included the Missouri Botanical Garden. The Garden library collections have been comput- erized and are available through Internet. Included are the general collection of 110 000 volumes of monographs and journals, and specialized collections of rare books, manuscripts, historical documents, and personal papers. Instruction for accessing the catalog is available by E-mail to molib@mobot.org. The Garden Herbarium now includes more than 4.5 million specimens of mosses, ferns, gymno- sperms, and flowering plants. Each specimen is being bar coded for ease of tracking loans, etc. The Garden’s database, TROPICOS, now con- tains more than 740 434 of the million published plant names, and information on more than 180 830 type specimens, more than 673 770 distribution records, nearly 347 187 synonyms, literature reports on chromosome counts, and ways in which plants have been used by, or had an effect on, humans. The Missouri Botanical Garden has a WWW Home Page at http://www.mobot.org. A special “news” section is going up soon. DELLE WILLETT Public Relations, Missouri Botanical Garden, P.O. Box 299, St. Louis, Missouri 63166-0299 *Phone 314-777-5100 and “Bioprospecting or biopiracy?” by Linda C. Nowlan. Other sections cover News (Biodiversity news, Cyberdiversity, and Biodiversity meetings), Reviews (both books and periodicals) and The Last Word, a message corner from the President of the Canadian Museum of Nature* “Sustainable yield: Is it possible?” This is the only feature to include a pic- ture of the author. Price of copies and subscriptions may be obtained from: Dawn Arnold, Business Manager (613-993-5908), or Susan Swan, Subscription Manager (623-990-6671), CANADIAN MusEuM of NATURE, P. O. Box 3443, Ottawa, Ontario K1P 6P4 *FAX 613-990-0318 *e-mail *Subsequently retired. The Ottawa Citizen 8 June 1996, p. A4, “Museum head leaves post.” 360 Canadian Species at Risk 3 April 1996 The Committee on the Status of Endangered Wildlife in Canada has released Canadian Species at Risk: April 1996 a 20-page listing of animals and plants designated by the Committee up to and including its most recent Annnal Meeting in April 1996. In is divided into three parts (1) contains species in the Extinct, Extripated, Endangered, Threatened, and Vulnerable categories: (2) contains species examined and designated in the Not At Risk catego- Amphipacifica: Journal of Systematic Zoology Volume IH, Number 2 is dated 10 April 1966 and contains an Editorial and two feature articles “The Amphipod Superfamily Hadzioidea on the Pacific Coast of North America: Family Melitidae. Part I. The Melita Group: “Systematics and Dis- tributional Ecology” by Norma E. Jarrett and E. L. Bousfield, pages 3-74; and “The Amphipod Family Oedicerotidae on the Pacific Coast of North America. THE CANADIAN FIELD-NATURALIST Vol. 110 ry and (3) Species examined and designated in the Indeterminate category because of insufficient scien- tific information. Also included is a record of status reexaminations. Copies can be obtained from: SYLVIA NORMAND Committee on the Status of Endangered Widlife in Canada, Ottawa, Ontario K1A OH3 * Phone: (819) 997-4991 Part I. The Monoculodes and Synchelidium Generic Complexes: “Systematics and Distributional Ecology” by E. L. Bousfield and Andrée Chevrier, pages 75-148. E.L. BOUSFIELD 611-548 Dallas Road, Victoria, British Columbia Canada V8V 1B3 Editor’s Report for The Canadian Field-Naturalist Volume 109 (1995) Ninety-five research, observation, synthesis, or tribute manuscripts were submitted to The Canadian Field-Naturalist in 1995. By the calender year’s end, evaluations by associate editors and additional reviewers were complete for all but 24. Of those returned to authors, 44 were subsequently accepted after requested revisions had been made; 24 of these were published in volume 108, the remaining 20 were with the printer for 110(1) and (2). The journal continues to have a healthy and broad circulation. The totals for The Canadian Field- Naturalist individual and institutional subscribers for 1995 are given in Table 1. A geographic tabulation of members of The Ottawa Field-Naturalists’ Club in 1995 is given in the Membership Committee sec- tion of the Report to Council, page 368 in this issue. The total circulation in 1995 (subscribers and mem- berships combined) was 1811 copies per issue. With its generalized content in research and observation articles and notes, very strong Book Review section, frequent tributes, news and comment items, and occasional historical and survey contributions, The Canadian Field-Naturalist contributes to the knowl- edge of the natural history of northern North America and relevant adjacent areas supported by a moderate annual membership in The Ottawa Field- Naturalists’ Club or non-member subscription to The Canadian Field-Naturalist. In addition, authors and their institutions generally share in the cost of publi- cation through page, table, figure and reprint charges. (In general, membership [about 40% of dues] contribution and subscriptions [the full amount] account for approximately half the costs of publication, charges make up the balance). Issue mailing dates for volume 109 were: (1) 21 July, (2) 12 October 1995, (3) 9 January 1996, and (4) 22 February 1996. The time from receipt to return for revision, and from acceptance to publica- tion was reduced during the year. (Receipt and acceptance dates are given at the end of each article and note in each issue), and the number of submis- sions rose slightly. The volume totalled 542 pages, the largest single issue (4) was 152 pages. Major contributions in this volume were a catalogue of freshwater algae in Quebec by Michael Poulin, Paul B. Hamilton, and Marc Proulx in 109(1): 27-110, for which the Canadian Museum of Nature contributed substantially to page costs. Additionally, 109(3) was devoted to the three-section history of exploration of the vascular flora of Canada, St. Pierre et Miquelon, and Greenland by James S. Pringle, with publication made possible by the cosponsorship of the Missouri Botanical Garden, St. Louis, Missouri. The number of articles and notes is summarized in Table 2 by topic, the totals for Book Reviews and New Titles in Table 3, and the distribution of pub- 1996 TABLE 1. Subscriber totals to The Canadian Field- Naturalist 1995 Canada USA Foreign Total Individuals 170 61 8 239 Institutions 206 266 46 518 Totals 376 327 54 757 Note: See Annual Report, Membership, for 1995 totals for membership in The Ottawa Field-Naturalists’ Club, page 368, which are additional to the subscriber totals given here. lished pages among issues in Table 4. Looking ahead to volume 110, number | will be a Special Issue: “A Life with Birds: Percy A. Taverner, Canadian Ornithologist, 1875” a biography by John L. Cranmer-Byng, Professor Emeritus, Department of History, University of Toronto. Taverner was the author of Birds of Eastern Canada (1919), Birds of Western Canada (1926), and Birds of Canada (1934) all published by the National Museum of Canada and all standard references of their time. I am particularly grateful to Henri Ouellet, W. Earl Godfrey, Ross James, and Daniel Brunton for their discussions and recommendations on this manu- script, to Ross for the initial suggestion that we con- sider publishing it, and to John and Margaret Cranmer-Byng for coming to Ottawa for two delightfully productive editorial conferences on the manuscript and their subsequent response to all edi- torial questions and suggestions. Arch Stewart and Mireille Boudreau, Library, Canadian Museum of Nature, provided bibliographic help and Bonnie Livingstone, Publications section, Canadian Museum of Nature, arranged approvals for the Museum photo in 109(3) and the majority in the Taverner issue to be published in 1996. Thanks are also due the journals Amphipacifica and Global Biodiversity for running notices on The Ottawa Field-Naturalist’s Club’s contributions to the biodi- TABLE 2. Number of articles and notes published in The Canadian Field-Naturalist Volume 109 (1995) by major field of study. Subject Articles Notes Total Mammals 8 12 20 Birds 8 10 18 Amphibians and reptiles 2 3 5 Fish 1 1 D Invertebrates 1 1 2 Plants 9 4 13 Other ik 0 1 Totals 30° 31 61° “a Tribute to Henry Mousley in News and Comment in 109(2). NEWS AND COMMENT 361 TABLE 3. Number of reviews and new titles published in Book Review section of The Canadian Field-Naturalist Volume 109 by topic. Reviews New Titles Zoology 32 63 Botany 8 25 Environment 22 52 Miscellaneous 9 8 Young Naturalists 0 38 Totals wal 186 versity publication through The Canadian Field- Naturalist and its predecessors since 1880 (Transactions to 1887, replaced by The Ottawa Naturalist to 1919). M.O.M. Printers, Ottawa, set and printed the jour- nal and special thanks are due Emile Holst and Eddie Finnigan and their staff. Wanda J. Cook proof-read the galleys for the volume. Mickey Narraway remained on standby call for any additional assis- tance to the Editor. Regretfully, she has elected to step down from this role in 1996. Bill Cody contin- ued as Business Manager, assisted by Lois Cody. Lois will be relinquishing her position as assistant to the Business Manager in 1996, after decades of for- mal and informal contributions in this role, and her contribution and dedication will be hard to duplicate. Bill oversaw the compilation, and proof-read and edited, the Index for volume 109 which was diligent- ly prepared by Leslie Cody, who on her marriage this year elected to change her name to Durocher. Wilson Eedy continued his fine contribution as Book Review Editor including the compilation of New Titles despite increasing world travel on other duties during the year. Other strengths in the journal are the Associate Editors who review manuscripts in their particular TABLE 4. Number of pages published in The Canadian Field-Naturalist Volume 109 (1995) by section (number of manuscripts in parenthesis). Issue number: -l- -2- -3- -4- Total Articles 110 (6)108(12) 87 (3) 50 .(8)355 (29) Notes 12 (6) 15 (8) O(O) 39(17) 66 (31) News and Comment 9(12) 16 (8) 4() 2 (4) 31(7) Book Reviews’ 11(17) 8 (7) 4(2) 39(45) 62 (71) Index -— -— SiC) AO e(Gb) Daye (C2) Advice to Contributors On O) ae CL) MO} (Opal CL) hits 2a (2) Next issue notice — Gael) Total pages: 142 148 100 152 542 “Total pages for Book Review section include both reviews and new titles listings but parenthesis figures include only the number of reviews. 362 field and often provide recommendations of addi- tional reviewers. Robert Anderson (Research Division, Canadian Museum of Nature, Ottawa), Warren Ballard (Wildlife Research Unit, Faculty of Forestry Research, University of New Brunswick, Fredericton, New Brunswick), Charles D. Bird (Erskine, Alberta), Robert R. Campbell (Woodlawn, Ontario), Brian W. Coad (Research Division, Canadian Museum of Nature, Ottawa), Anthony J. Erskine (Canadian Wildlife Service, Sackville, New Brunswick), W. Earl Godfrey (Curator Emeritus, Canadian Museum of Nature), Diana Laubitz (Researcher Emeritus, Canadian Museum of Nature), and William O. Pruitt, Jr. (Department of Zoology, University of Manitoba, Winnipeg) continued to serve as Associate Editors in 1995. George La Roi (University of Alberta, Edmonton, Alberta) contin- ued as Coordinator of the Biological Flora of Canada series. At the end of the year, Diana Laubitz was determined to relinquish her contribution as an Associate Editor to coincide with winding up her research projects at the Museum. Her evaluations and encouragement will be greatly missed. I am indebted to her both for thoughtful and incisive advice and counselling over the six years she has served as an Associate Editor for invertebrates (exclusive of insects), and for the many more years she has been my colleague both as an active member of the Ottawa Field-Naturalists’ Club and as a staff member at the National Museum of Canada/ Canadian Museum of Nature. Supplementing the views of Associate Editors, the following kindly returned reviews on request for one or more manuscripts in 1995: Stewart Alexander (Whitehorse, Yukon), R. C. Anderson (University of Guelph, Guelph, Ontario), Brad Andres (U. S. Fish and Wildlife Service, Anchorage, Alaska), C. Davison Ankney (University of Western Ontario, London, Ontario), George W. Argus (Canadian Museum of Nature, Ottawa, Ontario), J. Roger Bider ° (MacDonald Campus of McGill University, Ste Anne de Bellevue, Quebec), David M. Bird (MacDonald Campus of McGill University, Ste. Anne de Bellevue, Quebec), David A. Boag (Brentwood Bay, British Columbia), Stan Boutin (University of Alberta, Edmonton, Alberta), Ronald J. Brooks (University of Guelph, Guelph, Ontario), Luc Brouillet (Institut de recherche en biologie végé- tale, Montréal, Québec), Daniel F. Brunton (Ottawa, Ontario), Robert Elner (Canadian Wildlife Service, New Westminster, British Columbia), Mike Cadman (Canadian Wildlife Service, Guelph, Ontario), Ludwig Carbyn (Canadian Wildlife Service, Edmonton, Alberta), Paul M. Catling (Agriculture Canada, Ottawa, Ontario), Jacques Cayouette (Agriculture Canada, Ottawa, Ontario), Francois Chapleau (University of Ottawa, Ottawa, Ontario), Jerry G. Chmielewski (Slippery Rock University, Slippery Rock, Pennsylvania), William J. Crins THE CANADIAN FIELD-NATURALIST Vol. 110 (Ontario Ministry of Natural Resources, Huntsville, Ontario), Peter Ross Croskery (Grimsby, Ontario), E. J. Crossman (Royal Ontario Museum, Toronto, Ontario), Ron Cumberland (Department of Natural Resources and Energy, Fish and Wildlife Branch, Fredericton, New Brunswick), Stephen Darbyshire (Agriculture Canada, Ottawa, Ontario), Terry A. Dick (University of Manitoba, Winnipeg, Manitoba), Kathyrn Dickson (Canadian Wildlife Service, Hull, Quebec), T. G. Dilworth (University of New Brunswick, Fredericton, New Brunswick), Michael G. Dolinski (Alberta Agriculture, Food and Rural Development, Edmonton, Alberta), George W. Douglas (British Columbia Conservation Data Centre, Victoria, British Columbia), David Duncan (Saskatchewan Wetland Conservation Corporation, Regina, Saskatchewan), Patricia A. Duncan and James R. Duncan (Carven, Saskatchewan), Erica Dunn (Canadian Wildlife Service, Ottawa, Ontario), Leonard J. Easton (Nova Scotia Agriculture College, Truro, Nova Scotia), Carl H. Ernst (George Mason University, Fairfax, Virginia), M. Brock Fenton (York University, North York, Ontario), Graham Forbes (University of New Brunswick, Fredericton, New Brunswick), Bill Freedman (Dalhousie University, Halifax, Nova Scotia), John Fryxell (University of Guelph, Guelph, Ontario), Daniel Gagnon (Université du Québec a Montréal et Biodome de Montréal, Montréal, Québec), Cheri Gratto-Trevor (Canadian Wildlife Service, Saskatoon, Saskatchewan), Patrick T. Gregory (University of Victoria, Victoria, British Columbia), Erich Haber (National Botanical Services, Ottawa, Ontario), Fred Harrington (Mount St. Vincent University, Halifax, Nova Scotia), Vernon L. Harms (University of Saskatchewan, Saskatoon, Saskatchewan), Ken Harris (Ontario Ministry of Natural Resources, Carleton Place, Ontario), Stuart Hay (Université de Montréal, Montréal, Québec), David Henry (Canadian Heritage Directorate - General Prairie and Northern Region, Winnipeg, Manitoba), Thomas Herman (Acadia University, Wolfville, Nova Scotia), Erling Holm (Royal Ontario Museum, Toronto, Ontario), Eli Holmes (University of Washington, Seattle, Washington), Stuart Houston (Saskatoon, Saskatchewan), Ross James (Royal Ontario Museum, Toronto, Ontario), Stephen R. Johnson (LGL Limited Environmental Research Associates, Sidney, British Columbia), Daniel M. Keppie (University of New Brunswick, Fredericton, New Brunswick), Brina Kessel (University of Alaska-Fairbanks Museum, Fairbanks, Alaska), Michael Kingsley (Institut Maurice La Montagne, Mont-Joli, Québec), Gordon L. Kirkland, Jr. (Shippenburg University, Shippenburg, Pennsylvania), Richard W. Knapton (Long Point Bird Observatory, Port Rowan, Ontario), M. Steve Lapan (Bureau of Fisheries, Waterton, New York), J. Donald Lafontaine 1996 (Agriculture Canada, Ottawa, Ontario), Raymond LeClair (Université de Trois Riviéres, Trois Riviéres, Quebec), Richard D. Leonard (Winnipeg, Manitoba), Frederick G. Lindzey (Wyoming Cooperative Fish and Wildlife Research Unit, Laramie, Wyoming), Ross D. MacCulloch, (Royal Ontario Museum, Toronto, Ontario), W. Bruce McGillivray (Provincial Museum of Alberta, Edmonton, Alberta), B. MacKinnon, (University of New Brunswick, Fredericton, New Brunswick), Douglas B. McNair (Buckingham, North Carolina), Frank F. Mallory (Laurentian University, Sudbury, Ontario), Martin K. McNicholl (Burnaby, British Columbia), Mary Meagher (National Biological Survey, Yellowstone National Park, Wyoming), L. David Mech (US Fish & Wildlife Service, St. Paul, Minnesota), J. K. Morton (University of Waterloo, Waterloo, Ontario), David Nagorsen (Royal British Columbia Museum, Victoria, British Columbia), Robert W. Nero (Manitoba Natural Resources, Winnipeg, Manitoba), Thomas W. Nudds (University of Guelph, Guelph, Ontario), Henri Ouellet (Canadian Museum of Nature, Ottawa, Ontario), Gerry Parker (Canadian Wildlife Service, Sackville, New Brunswick), Alex Peden (Royal British Columbia Museum, Victoria, British Columbia), Michel Poulin (Canadian Museum of Nature, Ottawa, Ontario), Gilbert Proulx (Alpha Wildlife and Management Limited, Sherwood Park, Alberta), Scott Redhead (Agriculture Canada, Ottawa, Ontario), Austin Reed (Canadian Wildlife Service, Ste. Foy, Quebec), Randall Reeves (Okapi Wildlife Associates, Hudson, Quebec), James D. Rising (University of Toronto, Toronto, Ontario), Frederick W. Schueler (Bishops Mills, Ontario), J. C. Semple (University of Waterloo, Waterloo, Ontario), Doug Skinner (D. A. Westworth and NEWS AND COMMENT 363 Associates Ltd., Edmonton, Alberta), Kenneth W. Stewart (University of Manitoba, Winnipeg, Manitoba), Rudolph F. Stocek (Maritime Forest Ranger School, Fredericton, New Brunswick), Mike Sullivan (Department of Natural Resources and Energy, Fish and Wildlife Branch, Fredericton, New Brunswick), Tom Sullivan (University of British Columbia, Vancouver, British Columbia), Edmund S. Telfer (Canadian Wildlife Service, Edmonton, - Alberta), Shaun Thompson (Ontario Ministry of Natural Resources, Kemptville, Ontario), Michelle Wheatley (Kilpisjarven biologinen asema, Finland), Robert E. Wrigley (Winnipeg, Manitoba), Heather Whitlaw (University of New Brunswick, Fredericton, New Brunswick), Stan Van Zyll de Jong (North Augusta, Ontario), Dennis Voigt (Ontario Ministry of Natural Resources, Maple, Ontario), D. E. Yarborough (University of Maine, Orono, Maine). [In the list of reviewers in the Editors Report for volume 108 (1994) in The Canadian Field-Naturalist 109(2): 269-271, Graham Forbes, University of New Brunswick, Fredericton, New Brunswick, was omitted and Heather Whitlow’s surname was misspelled “Whitler”. My apologies to both] I am also indebted to Frank Pope, President of the Ottawa Field-Naturalists’ Club, the Club Council, Chairman Ron Bedford and the Publications Committee of the OFNC for their support, to The Canadian Museum of Nature for allowing me to con- tinue to use some space and facilities at the Macoun Nature Centre at Aylmer, and to Joyce for encour- agement at home, where most of the editing is done, throughout the year. FRANCIS R. COOK Editor Minutes of the 117th Annual Business Meeting of The Ottawa Field-Naturalists’ Club, 9 January 1996 Place and Time: Community Gallery, Canadian Museum of Nature, Metcalfe and McLeod Streets, Ottawa, 19:30 h Chairperson: Attendance: Frank Pope, President Frank Pope opened the meeting by asking mem- bers to spend the first half hour reviewing copies of the minutes of the previous meeting, a proposed amendment to Article 17 of the Constitution, the Treasurer’s Report, and the reports of Committees. 1. Minutes of the Previous Meeting No errors or omissions were identified. It was moved by Dave Moore (2nd Fenja Brodo) that the minutes be accepted. (Motion Carried) 2. Business Arising from the Minutes There was no business arising from the minutes. 3. Proposal to Amend Article 17 of the Constitution Frank Pope introduced a proposal to amend Article 17 of the Constitution which had been pub- lished previously in Volume 108, Number 3, of THE CANADIAN FIELD-NATURALIST. When the present wording of Article 17 was approved at the 114th Annual Business Meeting it was criticized for per- mitting a major revision to the Constitution in one block. The intent of the proposed amendment was to resolve that problem by permitting more than one change at a time only when those changes are “edito- rial and do not alter the intent of any article”. Under any other circumstances, changes to the Constitution must be introduced and approved individually. The meeting approved the proposal. (Motiori Carried) 4. ‘Treasurer’s Report Gillian Marston opened her remarks by noting the favourable report from the Club’s Auditor. In reviewing details of the financial statements, she noted an increase in Member’s Equity from $216 097 to $233 483 and an increase in the de Kiriline-Lawrence Fund from $10 478 to $12 749. There was a significant increase in the cost of pub- lishing THE CANADIAN FIELD-NATURALIST, as a result of an increase in the cost of paper. It was moved by Gillian Marston (2nd Ken Young) that the Treasurer’s report be accepted. (Motion Carried) Thirty-one persons attended the meeting. 5. Committee Reports Frank Pope introduced each of the Committee reports and asked for comments and questions. Several members asked for clarification of some points. In response to a question regarding the report of the Executive Committee, Frank said that an ad hoc committee had been established to review the use of the Member’s Equity in meeting the objectives of the Club, and it had made three recommendations to the Council; a) set aside money for the special publica- tions recommended by the Publications Committee, b) support land acquisition by organizations such as the Nature Conservancy of Canada, c) assist environmen- tal educators in the Ottawa region. The Council is still considering these recommendations. It was moved by Fenja Brodo (2nd Gillian Marston) that the Committee reports be accepted. (Motion Carried) 6. Nomination of the Auditor It was moved by Ken Young (2nd Gillian Marston) that Janet Gehr continue as Auditor for another year. (Motion Carried) 7. Report of the Nominating Committee On behalf of the Nominating Committee, Barbara Campbell presented the following slate of candidates for the 1996 Council. New members are indicated with an asterisk: Dave Moore Michael Murphy Dave Smythe Eileen Evans Gillian Marston Ron Bedford Jeff Harrison Fenja Brodo Cendrine Huemer Lee Cairnie Ann MacKenzie Bill Cody Patricia Narraway Francis Cook Frank Pope President Vice-President Recording Secretary Corresponding Secretary Treasurer Other Council Members 364 -1996 Ellaine Dickson Tom Reeve* Colin Gaskell Jane Topping Alan German* Chris Traynor* Christine Hanrahan Ken Young One member of the 1995 Council, Carol German, chose not to stand for re-election, and one member, Trix Geary, resigned during the term of the council. It was moved by Barbara Campbell (2nd Peter Hall) that the proposed slate be accepted. (Motion Carried) 8. New Business Frank Pope thanked Lois Cody for her long and valuable contribution to the Club as the Treasurer’s Assistant, and in particular her willingness to contin- ue temporarily after her official retirement in September, to ensure a smooth transition to a com- puter based accounting system. Frank reviewed the long and fruitful association between the Club and the Canadian Museum of Nature, and the awkward situation in which the Club found itself when it criticized the decision by the Museum to build on a wetland site in Aylmer. In the subsequent discussion of the Museum’s action and the Club’s response, several members expressed strong disagreement with the Museum’s action. It was moved by Ian Whyte (2nd Fenja Brodo) MINUTES OF 117TH ANNUAL BUSINESS MEETING 365 that the minutes record the fact that the meeting expressed strong disagreement with the Museum’s choice of building site in Aylmer. (Motion Carried) In his remarks as outgoing President, Frank said that he was proud to have played a part in the history the Club. He thanked The Canadian Museum of Nature, and Agriculture and Food Canada for the use of their facilities, and members of Council and of the Club for their support and encouragement during his four years as President. 9. Presentation by the Fletcher Wildlife Garden Committee Jeff Harrison and Peter Hall gave a presentation on the history and development of the Fletcher Wildlife Garden with before and after slides to show the changes that have taken place. Thousands of hours of volunteer effort are now producing rewards in the form of habitat growth, visitors to the site, and an Interpretive Centre that will enable many more activities at the Garden. For the future, the Garden will remain a volunteer organization and more effort will be made to raise operating and capital funds to ensure that it is self-sufficient. 10. Adjournment At 22:05 h, it was moved by Lee Cairnie (2nd Ian Whyte) that the meeting be adjourned. (Motion Carried) DAVE SMYTHE Recording Secretary Committee Reports for 1995 to The Ottawa Field-Naturalists’ Club Awards Committee The Following awards were presented at the annu- al Soiree held on April 28, 1995: 1994 — MEMBER OF THE YEAR AWARD: Tony Beck for his enthusiastic involvement in the OFNC see- dathon, Bird Records Sub-committee, Chairmanship of the Birds Committee, and for his continuing ser- vice on bird walks and coordinated field trips. 1994 — GEORGE MCGEE SERVICE AWARD: Ron Bedford for his 14 years of service as Chairman of the Publications Committee, for his longtime record of service on OFNC Council, and his participation on field trips. 1994 — CONSERVATION AWARD (OFNC MEMBER): Jan Huggett a second Conservation Award, for his continuous commitment to conservation in the National Capital Region and for his leadership in mobilizing of the citizens of Aylmer Quebec, to pro- tect the Wychwood pine forest. 1994 — CONSERVATION AWARD (NON-MEMBERS): The Goulbourn Environmental Advisory Committee for the conversion of the abandoned Richmond sewage lagoons into a wetland conservation area, thereby establishing new freshwater marsh habitats for resi- dent water birds and providing educational opportu- nities for local residents and naturalists. Because the Club has maintained its full comple- ment of twenty-five Honorary Members, there were no Honorary Memberships confirmed this year. Also, there was no recommendation for the Anne Hanes Natural History Award. BILL ARTHURS 366 Birds Committee A plan of action for the reactivation of the Bird Records Sub-Committee in early 1996 was developed. The Committee oversaw the Late Fall Count. The turnout was much improved over 1994 perhaps due to the change back to a single day event. The Committee also oversaw the Ottawa-Hull Christmas Bird Count, and officially accepted responsibility for the Dunrobin-Breckenridge Christmas Bird Count from Bruce Di Labio who will continue to oversee it. The Committee tended the Bird Status Line. A new permanent number (860-9000) was installed in downtown Ottawa to minimize long distance charges for birders in outlying areas. The Rare Bird Alert was updated to reflect the changing composition of the birding community. The archiving of materials from the Committee’s activities is nearly complete, including nearly 200 slides of rare birds. The Committee took care of the Club feeders and raised over $1000 from the Seedathon for their maintenance. Arrangements were made to have the American Birding Association include the OFNC checklists in its sales catalogue. TONY BECK Computer Management Committee The Computer Management Committee ensures the efficient and controlled use of the computer assets of the Club. It maintains and improves exist- ing systems and increases awareness of how comput- er facilities may be of benefit to other committees. This year, members of the Committee assisted in the selection of a computer system for the Treasurer’s Assistant and the replacement of the computer used by the Membership Committee. They also provided various support services to the users of the computer system used to publish Trail & Landscape. MICHAEL MURPHY Conservation Committee The Committee liaised with local Conservation Authorities and Provincial Ministries concerning various programs including the Landowner Resources Centre, the Forest Diversity Project and watershed studies of the Jock River. Members of the Committee participated in the Marlborough Forest Advisory Committee of the Regional Municipality of Ottawa-Carleton (RMOC) and reviewed environmental policies in the RMOC official plan. The Wetlands Working Group, of which OFNC is a member, submitted recommenda- tions to the RMOC Council in an attempt to recon- cile naturalist and landowner conflicts with regard to the Ontario Wetlands Policy (1992) as it is to be applied in Ottawa-Carleton. THE CANADIAN FIELD-NATURALIST Vol. 110 The Committee made various submissions to the RMOC, the National Capital Commission (NCC), and the Ontario Municipal Board to advocate the protection and enhancement of natural areas in Ottawa-Hull. These included submissions on a pro- posed subdivision in the Leitrim Wetland (Gloucester), the “outsourcing” of park management by the NCC (Gatineau Park), the Canadian Museum of Nature Consolidation Project (Aylmer), the Kanata North Expansion proposal (Kanata), the Transportation Master Plan (RMOC) and the Natural Environmental Study (RMOC). : MICHAEL MURPHY Education & Publicity Committee With the help of approximately 45 volunteers, the Committee set up the Club’s display at four different locations. The Committee responded to nine requests for speakers and seven requests for walk leaders from various Church groups, Seniors homes, scout- ing groups and others. Posters were designed and posted to advertise monthly meetings of the Club. Judges and prizes were arranged for the Ottawa Science Fair. New slide presentations were provided during the year, and work continues on a new set of slides. The FreeNet connection was refined and a plan for expansion was started. Work continues on a tele- phone list of environmental and other agencies for the use of members. DAVE Moore Excursions & Lectures Committee During the past year, a varied program of 46 same-day outdoor activities was scheduled for Club members, four of which required the use of a char- tered bus. An indoor workshop on bird identification by sight and sound was an additional event. Approximately 80% of all outings were conducted within the Ottawa district. A weekend outing to Algonquin Park and a four-day excursion to Point Pelee were also arranged. Traditional OFNC social functions such as the Annual Soiree, Members’ Slide Night, and the Annual Picnic once again proved to be successful ventures. The Committee supported a motion to Council by the Membership Committee to discontin- ue the New Members Night in response to dwindling attendance over the past two years. A series of eight evening lectures featuring a stim- ulating mix of speakers, was well attended. COLIN GASKELL Executive Committee On February Ist, the Committee met to discuss the issues anticipated in the coming year. It decided to continue the current format of the Annual Business Meeting. It decided to continue holding the monthly 1996 meetings at the Canadian Museum of Nature notwith- standing the Museum’s new policy of charging for the use of rooms, and concern was expressed about the site in Aylmer chosen by the Museum for its new facility. The Club’s relationship with the Federation of Ontario Naturalists and the Canadian Nature Federation was discussed. The healthy financial reserves of the Club, led to a decision to set up an ad hoc committee to investigate possible uses of some of these funds to further the objectives of the Club. On June 29th, the Committee met to discuss trans- ferring the Club’s financial accounts from a manual system to a computer system and to fill a future vacancy in the position of Treasurer’s Assistant. It was decided to establish a committee to implement the transition. FRANK POPE Finance Committee The Committee considered the recruitment of an assistant for the Treasurer to replace Lois Cody who retired at the end of the fiscal year, and the replace- ment of the current ledger based accounting system with one based on a software package. KEN YOUNG Fletcher Wildlife Garden Committee For the past two years the project has been slowed awaiting transfer of responsibility for Building 138 (the interpretive centre) from Agriculture Canada. With the project in jeopardy due to lack of a facility, it was decided to “occupy” the building with Agriculture Canada’s unofficial approval. Between February and April approximately 50 volunteers in many working bees, cleaned and paint- ed the Centre. A workshop for tour leaders was con- ducted. The public opening on Earth Day April 22nd attracted 200 people and subsequent open houses and tours during the Summer and Fall attracted an additional 300 people. Tables and chairs were purchased for the Centre and after April, all Garden Committee meetings were held there. The Council met there in June and September. A donation box designed to look like a Great Blue Heron was built and placed in the Centre attracting donations of about $100. Twenty four t-shirts with the Garden logo were made and given to key volunteers, stakeholders, and members of the media. Two spon- sors donated $4500 without solicitation. In September and October, painting was complet- ed, a septic line was installed, interior plumbing replaced, and storm windows were fixed and installed, at a total cost of about $10,000. In the model backyard garden, some perennials were added, and a cedar hedge was planted. A troop from Scouts Canada adopted the new woods and planted 500 trees in May. In the old woodlot, more woodland plants were brought in. MINUTES OF 117TH ANNUAL BUSINESS MEETING 367 The self-guided trail brochure was reprinted including a french edition. A dispensing box was affixed to the door so that visitors can take a brochure and tour the site when the Centre is closed. Over the year, volunteers contributed approximately 1000 hours of time. JEFF HARRISON Macoun Field Club Committee The Committee met six times during the year to plan the weekly schedule for the children and young people of the Club. Field trips and camping trips were balanced with indoor presentations by mem- bers of the local naturalist and scientific community. A gift from the Excursions & Lectures Committee made possible an all-day bus trip to Murphy’s Point Provincial Park for the whole Club. A determined effort was made to welcome a stu- dent with a serious hearing impairment into the Club. The willing acquisition of sign language by both other children and the leaders encourages us to think it will be possible to accommodate deaf mem- bers in the future. The Club published the 49th issue (in the Club’s 47th year) of the Little Bear, in June, with an impor- tant section on members’ Study Area projects. ROBERT LEE Membership Committee The total membership paid up for 1995, was 1054; an increase of 21 from 1994. There were 138 new members, including 80 Individual, 57 Family, and one Sustaining. A detailed breakdown of these num- bers with the 1994 numbers in brackets, follows: Four complimentary one-year memberships were presented to outstanding young participants in the Annual Science Fair. Mr. Claude E. Garton was added to “Golden Anniversary” membership list. Claude joined the Club in 1945 and was awarded an Honorary mem- bership in 1985. PATRICIA NARRAWAY Publications Committee The publications Committee oversees and advises on all aspects of the Club’s publications. It met three times in 1995. With the appearance of four issues of The Canadian Field-Naturalist (Volume 108, Numbers 3, 4 and Volume 109, Numbers 1, 2) in 1995, and two more ready to go to press, the publication of the Journal is essentially back on schedule. These four issues contained 566 pages, 36 articles, 34 notes, one COSEWIC article, 63 book reviews, 221 new titles, one commemorative tribute, and 18 pages of news and comments. All of the Associate Editors accepted reappointment for 1995. The number of subscrip- tions to The Canadian Field-Naturalist (CEFN) remains stable. 368 Volume 29 of Trail & Landscape was published in four issues that contained 172 pages, with again a large emphasis on bird-related articles. Three issues were accompanied by a one-page insert of the Green Line. During 1995 some special requests were consid- ered. The OFNC applied for, but was unsuccessful in obtaining, a grant from Science Culture Canada towards the publishing of a book for young natural- ists by Joe Shepstone. The Club approved a grant of $2000 towards the publication of the Butterflies of THE CANADIAN FIELD-NATURALIST CANADA Vol. 110 FOREIGN Canada by P. Hall, D. Lafontaine, and R. Layberry. The Canadian Field-Naturalist also agreed to pub- lish, with the cooperation of the Missouri Botanical Garden, three articles on the history of botanical exploration in Canada, Greenland, and Saint-Pierre and Miquelon. The Canadian Field-Naturalist has also arranged to publish a biography of P. A. Taverner, in a future issue. RONALD BEDFORD The Ottawa Field-Naturalists’ Club Financial Statements: Year ended September 30, 1995 Auditor’s Report 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, 1995, and the statements of operations and members’ equi- ty. 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 accepted auditing standards. Those standards require that I plan and perform an audit to obtain reasonable assurance whether the financial statements are free of material misstatement. An audit includes examining evidence supporting the amounts and disclosures in the financial statements. An audit also includes assessing the accounting principles used and signifi- cant estimates made by management, as well as eval- uating the overall financial statement presentation. In common with many non-profit organizations, The Ottawa Field-Naturalists’ Club derives some of its revenue from memberships, donations, and fund rais- ing activities. These revenues are not readily suscep- tible to complete audit verification, and accordingly, my verification was limited to accounting for the amounts reflected in the records of the organization. In my opinion, except for the effect of the adjust- ments, if any, which I might have determined to be necessary had I been able to satisfy myself 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 of September 30, 1995, and the results of its operations for the year then ended in accordance with generally accepted accounting principles. JANET M. GEHR Chartered Accountant North Gower, Ontario 6 January 1996 ee OO ee 1996 MINUTES OF 117TH ANNUAL BUSINESS MEETING The Ottawa Field-Naturalists’ Club Balance Sheet September 30, 1995 1995 ASSETS .- CURRENT ASSETS (Cashier eS 270,404 Accounts Receivable ...... 9,808 Interest Receivable ......... 4,299 Prepaid Expenses............ 1,608 286,119 RIXEDI (NOES) rete — LAND - Alfred Bog........ 3,348 289,467 1994 3,348 275,752 LIABILITIES, FUNDS AND MEMBERS’ EQUITY CURRENT LIABILITIES Accounts Payable............. 22,800 Deferred Income ...........:.: 12,068 34,868 FUNDS (Note 4)..............555 14,616 LIFE MEMBERSHIPS............ 6,500 CLUB RESERVES ............0622 100,000 GENERAL EQUITY .............. 133,483 289,467 The Ottawa Field-Naturalists’ Club Statement of Members’ Equity September 30, 1995 1995 Excess INCOME (EXPENDITURES) The Ottawa Field- Naturalists’ Club.......... 1,476 Canadian Field-Naturalist 15,126 16,602 OTHER INCOME (ALLOCATIONS) Donations - Misc. upon membership renewal .... 3,503 Allocation to Kiriline-Lawrence Fund (2,719) 784 MOTAMINCOME Rss eee. 17,386 MEMBERS’ EQUITY, Beginning of Year........ 216,097 MEMBERS’ EQUITY, Endo year sn eee 233,483 30,327 9,871 40,198 12,957 6,500 100,000 116,097 275,752 The Ottawa Field-Naturalists’ Club Statement of Operations - OFNC Year Ended September 30, 1995 1995 INCOME Memberships:see-s-sesee 14,406 T&L Subscriptions and Backslssucsse eee 503 IN tEKeS tee eee es 33212 OthemSaleseee ee 2,885 Special Publications......... 139 Motaliincomes seers 21,145 EXPENSES OPERATIONS EXPENSES Atfiliationitees}es ns 2,134 @omputerensec serene 3,397 Depreciationee.s.e ee - Membershipe iene 1,702 Office assistant................. 748 Operations scree 3,088 OFNC GST Rebate .......... (499) Total Operations Expenses 10,570 CLuB ACTIVITY EXPENSES (Net) SOIC ee See 209 itd Sieestee secrete ee — Gonservationeeeen ee 139 Education and Publicity ... 157 Excursions and Lectures .. (713) Fletcher Wildlife Garden . 1,200 Macoun'@lubeesse 613 Trail & Landscape............ 7,494 Total Club Activity Expenses 9,099 INCOME OVER EXPENSES... 1,476 369 1994 14,099 645 1,864 1,311 149 18,068 295 33955 1255 2,564 750 2,450 (591) 10,678 370 The Ottawa Field-Naturalists’ Club Statement of Operations - CFN Year Ended September 30, 1995 1995 INCOME Membershipsis:..2:0 0st 9,699 Subseriptions:..... we PES TIT! Sub=otalies Aes 35,470 Reprints: 22 se See 6,542 Publication charges .......... 31,026 Back numbers: 2/228: 316 Interest and exchange....... 16,485 Total Income......2.:....:. 89,839 EXPENSES Publishing ei Gecse ccs 49,557 IREPUIntSrarceecseeseshaneetaae 6,495 @irculationss22 3 6,743 Citi Gree ates ae vacsereeess 2,475 Office:assistante eek 4,997 Officeysuppliesi...3--2.22.° 2,344 AGVertising 2.1 ates. 145 ION OLanlaye sees ea 4,500 CEN GST Rebate.............. (2,543) 74,713 INCOME OVER EXPENSES...... 15,126 THE CANADIAN FIELD-NATURALIST 1994 9,317 19,029 28,346 8,848 34,737 225 10,339 82,495 37,443 5,956 5,369 3,707 4,675 3,736 134 3,000 (3,154) 60,866 21,629 Vol. 110 The Ottawa Field-Naturalists’ Club Notes To The Financial Statements September 30, 1995 1. Authority and Activities The Ottawa Field-Naturalists’ Club is a non-profit organization incorporated under the laws of Ontario (1884). The Ottawa Field-Naturalists’ Club pro- motes the appreciation, preservation and conserva- tion of Canada’s natural heritage; encourages inves- tigation and publishes the results of research in all fields of natural history and diffuses information on these fields as widely as possible. It also supports and cooperates with organizations engaged in pre- serving, maintaining or restoring environments of high quality living things. 2. Significant Accounting Policies Membership, subscriptions and donations are recorded as received. All other revenues and expen- ditures except for inventory are accounted for on the accrual basis. Memberships are allocated to The Canadian Field-Naturalist publication on a pre-deter- mined percentage. Supplies, records, tapes and other items held for resale are expensed when purchased. Fixed assets acquired after 1989 are expensed. Fixed assets acquired prior to 1990 were recorded at cost and depreciated on a straight line basis. 3. Fixed Assets 1995 1994 COSt ieee age 16,746 16,746 Accumulated Depreciation..... 16,746 16,746 NetBook! Waluess) Que iaie - - 4. Funds 1995 1994 Baldwin Memorial Fund........ - 358 SECM Ath OMe. heer scctscsete ee 585 888 Anne Hanes Memorial Fund... 870 815 de Kiriline-Lawrence Fund.... 12,749 10,478 AlfrediBoouitss Meunier 412 418 14,616 129571 Book Reviews ZOOLOGY Manual of Ornithology: Avian Structure and Function By Noble S. Proctor and Patrick J. Lynch. 1993. Yale University Press, New Haven and London. xi + 340 pp., illus. U.S. $40. This book includes topics expected from this type of work but it is not an introduction to ornithology. Although systematics, topography, and field tech- niques are discussed and occupy separate chapters, this work is primarily a guide to bird anatomy and function for undergraduate students or graduate stu- dents who may not have been acquainted earlier with the structure of birds. Feathers, skeleton, muscula- ture, digestive system, circulatory system, respirato- ry system, urogenital and endocrine system, and ner- vous system are the topics of the other chapters. An appendix on bird classification, a bibliography, and an index are also part of the book. The text is clearly written and abundantly illustrat- ed. The illustrations are accurate and of excellent quality even if some of them are presented in an unusual perspective, at least one that is not found in classical presentations of anatomical topics. Each is accompanied by an extensive caption detailing the contents of the plate and containing a great deal of information about the illustrated topics and instruc- tions on dissection where applicable. Chapter 3, enti- tled “Topography”, will be useful to anyone interest- ed in birds, not only students or ornithologists but also to bird watchers and biologists who need to use ornithological references, because it contains, defines, and illustrates most of the terminology of modern works particularly on external anatomy. In general the text is complete and up-to-date, the authors having incorporated the most recent informa- tion available at publication time. The writing is clear and easy to follow. I found it easy to retrieve information because the layout is clear and well organized, key words being highlighted in bold face characters. The definitions of ornithological terms are precise, succinct, and carefully worded. The Downy Waterfowl of North America By Colleen Helgeson Nelson. 1993. Delta Station Press, Portage la Prairie, Manitoba. xx + 302 pp., illus. $49.95 + $5.00 shipping and handling. After many years in preparation, this exhaustive work on the downy plumages of North American ducks, geese, and swans has appeared and is fulfill- Each chapter contains a two-page worksheet of questions where one can verify how much has been retained from reading the previous pages. The work- sheets are not unlike those found in Pettingill’s Ornithology in laboratory and field (1970) but are more detailed and elaborate. There is also a one-page list of references pertaining to the chapter. These ref- erences appear also in the main bibliography at the end of the book and although this subject-oriented list of references may appear to be repetitious it helps the reader to find references on a given topic more easily. Chapter 12, “Field techniques”, seems at first to be somewhat out of place in this book but I found its contents useful in presenting various techniques nec- essary for the study of birds. It should be compulsory reading to all students who plan to undertake studies on birds. Amateurs and bird watchers also will find a wealth of valuable information. The “Classification of Birds” given in appendix is based on the Sibley and Monroe system which has been clearly explained in Chapter 2 (Systematics). This book will not replace the recent ornithology manuals because of its scope. However, it should be consulted by all students enrolled in an ornithology course. I recommend its reading also to amateurs and bird watchers who will find clear explanations about topics often difficult to access elsewhere in the ornithological literature. This book in spite of its limitations constitutes a complementary reference to other standard ornithology manuals. The quality of the text, illustrations, presentation, and overall pro- duction, makes it an essential element in a balanced ornithological library. HENRI OUELLET 175 avenue de la Citadelle, Hull, Québec J8Z 3L9 ing the expectations of all those who were aware of the author’s lifetime devotion to this enormous task. The book is attractively produced on high quality paper with an easy to read typography, plain design, and sturdy binding. In the introduction, the author outlines the proce- Sil BIZ dure she has followed in the preparation of the book and defines the parameters used in her descriptions of young birds as well as problems related to the use of live birds versus museum specimens when describing colours of plumage or soft parts. This is an extensive part where many aspects of young waterfowl biology are discussed succinctly but ade- quately for the non-initiated reader to obtain a full understanding of the text related to each species. Colours and their nomenclature are treated with par- ticular care as they are used extensively in the rest of the book and because the identification of several species in down plumage depends on an accurate colour evaluation. Fifty-three species are treated in the book and the author has followed her own taxonomic arrangement to present the information. Each tribe is introduced by a section on its distribution and taxonomy followed by another on the general appearance of the birds and their behaviour. Within each tribe, the following information is given for every species, English, French, and Spanish names, scientific name, author, appearance, variation, source of specimens, and other references; a short section called “discussion” is fre- quently inserted after the section on variation. The sections on appearance and variation of each species are particularly useful as the author describes the downy plumage in great detail and gives a com- plete summary of the known variation, including that of the soft parts coloration (bill, legs, eyes). These descriptions when read carefully will provide the necessary information for the identification of any downy young either in the field or in the laboratory. The author is particularly careful in her descriptions and in the use of colour terminology so that the text is clear and easy to understand. The discussion fol- lowing many species contains information on a vari- ety of topics such as behaviour, anatomy, measure- ments, colorations, and field identification in relation to the findings of other authors as reported in the lit- erature. A list of the specimens used by the author appears in the section entitled “Source of specimens” and a shorter section, “Other references’, lists refer- ences peculiar to the species where it is included. The part dealing with “References” contains twen- ty-five pages and appears to be complete at least up to the publication of the book. It is followed by the three appendices and a table of contents. The first appendix gives a list of measurements in metric units for all the species treated at some or all the standard stages of development from 8 to 96 hours, and includes mass (weight), exposed cul- THE CANADIAN FIELD-NATURALIST Vol. 110 men, two tarsal lengths, and middle toe with claw. Appendix B is entitled “Color Descriptions” and deals with a detailed depiction of the colours of the down plumage as well as that of the soft parts called here “unfeathered parts” of all the species dealt with in the book. The colour descriptions are meticulous and based on the Munsell colour notation which pro- vides an indispensable and bias free element of com- parison. “Appendix C” entitled “Identification Keys” constitutes an important part of the book. It starts with a key to genera and is followed by keys to species within each genus. This part can be consid- ered as one of the highlights of the work. In addition to the succinct but precise text, the outstanding black-and-white line drawings of each species com- plement the text and provide the necessary informa- tion for accurate identifications of downy young. The illustrations are of an unusually high quality and can suffice in many instances for identifying an unknown duckling without having to refer to the text. They are not only accurate but also delightful. This is where the author has contributed an outstand- ing accomplishment to waterfowl studies along with the ten color plates and life sketches scattered through the main text. The colour plates are excep- tional in quality and reproduction to the extent that they could easily be considered as works of art. Form a scientific point of view, the postures of the birds are meticulous and the colours of the plumage and soft parts are accurate. The taxonomic sequences and classification adopted by the author may not be in agreement with that of many recent proposals but this should not be an important issue at this time because the taxonomy of many groups, including waterfowl, is currently under review as a result of new techniques of analy- sis and current taxonomic studies. Results of future studies may support the taxonomic proposals of the author and may contribute to a better understanding of the evolution and relationships of these birds. All those interested in waterfowl biology, particu- larly those who have to identify downy waterfowl, will find this book an essential reference in their study or work. The price is more than reasonable for a book of this quality. It is therefore with great plea- sure that I recommend it to anyone working with waterfowl and also to anyone interested in birds in general. HENRI OQUELLET 175 avenue de la Citadelle, Hull, Québec J8Z 3L9 : ! | 1996 Last of the Curlews By Fred Bodsworth. 1995. Counterpoint, Washington. 192 pp., illus. U.S. $15. The vast number of Eskimo Curlews of the past century were decimated by market hunting, not an unfamiliar story in North America. What is riveting about this species is the tantalizing hints of its linger- ing as a few scattered survivors still occasionally reported along the traditional spring and fall migra- tion routes, mixed with flocks of other shorebirds, some similar enough in appearance to leave a ele- ment of doubt to sight identifications. This slim volume is a reprint of the 40-year-old- classic, originally published in 1955 by Dodd, Mead & Company, in which Bodsworth imagines a year in the life of a surviving Eskimo Curlew, and in doing so graphically re-creates its behaviour and ecology centred on its nine-thousand-mile migration route from the far north of North America to the far south of South America. Particularly poingant is a fleeting companionship with a potential mate which does not survive to the nesting grounds. Although this is biol- Bats: A Community Perspective By James S. Findley. 1993. Cambridge University Press, Cambridge. xi + 167 pp., illus. U.S. $19.95. This is a good book for professional community ecologists working on bats or interested in compar- ing their study species with bats. The first third of the book is somewhat introductory: chapter | justi- fies the study of bat communities. Much is made of the fact that bats are exceptional, being small yet K- selected mammals who live in stable habitats. Chapter 2 provides brief descriptions (3—6 sen- tences) of each bat family and their habits. The non- _chiroptologist will do well to read this section care- fully, as family or genus names regularly appear in the rest of the book. Chapter 3 gives an overview of the methods used in the field to catch bats and to determine their movements, interactions, and diet. This chapter ends on a presentation of ecomorpholo- gy, an analytical field which uses multivariate tech- niques to group morphologically similar species in the hope that ecological similarities will be reflected in such groups. The remaining two-thirds of the book is where the beef is for the community ecologist. Chapter 4 describes the bat communities of the following five geographical regions: temperate North America, Europe and Russia, tropical Africa, tropical Asia and Australia, Central and South America. Exhaustive tables list the species found at various sites within these zones, sometimes with abundance indexes. Chapter 5 looks at the influence of food, foraging areas, roosting sites, heat, and water, and concludes BOOK REVIEWS 513 ogy from a personal birds-view of events, instinct is stressed. An effective text contrast is provided by stark verbatim excerpts from the scientific literature which introduce many chapters. This edition is illustrated by Abigail Rorer who has redrawn from the originals by the late Terry M. Shortt. It begins with a forward by W. S. Merwin out- lining his personal discovery of the book and subse- quent championing of its re-printing. Fred Bodsworth has added an Epilogue documenting the scattered evi- dence of Eskimo Curlew survival in the past four decades, and Murray Gell-Mann an Afterward touch- ing on other North American abundant species deci- mated by over-exploitation and why we should care, ending starkly: “The human race must get used to the simple idea that the Earth is really finite. The sooner this happens, the happier the outcome will be.” FRANCIS R. COOK RR 3, North Augusta, Ontario KOG 1RO that although some of these factors may be limiting to bat populations, competition for them does not appear to be important. Chapters 6 and 7 deal with the global pattern of bat distribution in terms of species number, biomass, and trophic types. The first of these two chapters is more descriptive, with a lot of ecomorphograms, whereas the second one is more analytical, concluding that bat diversity seems to depend on habitat area and on the number of ancient or modern refuges (partitions in habitat). Chapter 8, a 5-page summary where parallels are established with plants, birds, rodents, and freshwa- ter fishes, ends by restating that bat species seem to have arisen in refuges and managed to coexist there- after, without much evidence for the role of competi- tion and resource limitation. Proponents of these two concepts as important factors shaping communities can take solace in several studies mentioned in the book where competition was apparent, and in the author’s admission that, as a whole, studies in the field of bat communities are still relatively few, owing to the difficulty of studying a nocturnal ani- mal for which species are sometimes hard to identify in the hand. Notwithstanding this last comment, the author did a good job of reviewing the literature, giv- ing 206 references. The book is well produced, although I did catch six typos and found graph lines to be too thin. The writ- ing gets technical in places (particularly where eco- morphology is concerned) but it is otherwise clear, and the summaries at the end of each chapter, if not at 374 the end of chapter sections, are well done. The book is part of the Cambridge Studies in Ecology series; it is too specialized for the field naturalist, but commu- nity ecologists should find it useful. THE CANADIAN FIELD-NATURALIST Vol. 110 STEPHAN REEBS Département de biologie, Université de Moncton, Moncton, New Brunswick E1A 3E9 No Room for Bears: A Wilderness Writer’s Experiences with a Threatened Breed By Frank Dufresne. 1991. Alaska Northwest Books, Bothell, Washington. 252 pp., illus. Reprinted from 1965 edition. U.S. $12.95; $15.95 in Canada. Dufresne draws upon years of personal experience with people and bears to craft first-hand accounts of human behaviour in bear country and the response of bears to the people they encounter. While some chapters address black bears and polar bears, most of this book focuses on the brown bears of Alaska. There are many interesting, and sometimes chilling, accounts of human-bear encounters at the fire side, in tent camps, around fishing holes, at photographer blinds, and at remote cabins. Dufresne rounds the book out with information about bear biology, evolution, and folklore; he makes anecdotal reference to bear species inhabit- ing other continents as well. In addition to bear Bird Life of Woodland and Forest By Robert J. Fuller. 1995. Cambridge University Press, New York. xiii + 244 pp., illus. U.S. $64.95. This is an excellent and well-written account of woodland birds in Britain. Overall, the book is read- able, informative, and provides interesting insights and questions about bird life in forested habitats. It is unfor- tunate that the book deals largely with British birds which might limit its interest to Canadian naturalists. The book is well organised and the initial chapters provide a historical perspective on the forests of Britain and Europe followed by a discussion of how birds use woodland habitats. A chapter on the abun- dance and distribution of woodland birds is well thought out and discusses factors which determine the diversity of birds. Comparisons with avifaunas in mainland Europe and North America are made where appropriate and the book is well referenced through- out. The book has chapters on broad-leaved forest, coniferous forest, scrub forest, and upland woods, and includes sections on human managed “woodland” systems such as coppice, wood-pasture, and heath. One aspect of the book that I found refreshing was the insights into bird distribution and behaviour which reflect the dynamic interactions between many species of birds and their habitats. The long history of human intervention in forested habitats in Britain, well over 2000 years, has left no old growth forest and very little mature forest; the majority being variously managed types of forests. Associated mortality through direct encounters with humans, Dufresne addresses threats to the Alaskan brown bear resulting from habitat alienation and destruc- tion. In his plea for the preservation of North American bears, Dufresne provides a series of rec- ommendations for people living and working in bear country. Because this book was written in 1965, some of the information on bear biology and evolution is outdated. But for those interested in mid-century human-bear interactions, this book will make a worthwhile addition to your library. PAUL A. GRAY Department of Natural Resources, P.O. Box 8070, Cause- way, Harare, Zimbabwe with changes in management of forests and their species composition have come changes in the distri- bution of birds. Dr. Fuller’s reference to the general change in the habitat of the mistle thrush over a hun- dred years or so, from coniferous forests, through deciduous forest to largely suburban habitats indi- cates the adaptability of some species to changing environments. Likewise, the variation in habitat use in different parts of a bird’s range, indicate the diffi- culty in finding appropriate indicators for measuring the health of forests. In central and eastern Europe, coal tit, goldcrest, bullfinch, and mistle thrush use coniferous habitats while in Britain they may be found in a range of deciduous habitats. Major concepts of bird distribution such as edge effects, patchiness of habitat, and stand structure are discussed. Dr. Fuller draws on a wide personal back- ground of research as well as that of the British Trust for Ornithology and other researchers and naturalists to make this a valuable book. The inclusion of a chapter on woodland in a changing countryside pro- vide much food for thought and provide an under- standing of how man’s use and management of forests through millennia have affected the distribu- tion and abundance of birds. MARK E. TAYLOR Geomatics International Inc., 3370 South Service Road, Burlington, Ontario L7N 3M6 1996 BOOK REVIEWS S75) A Supplement to Distribution and Taxonomy of Birds of the World By Charles G. Sibley and Burt L. Monroe, Jr. 1993. Yale University Press, New Haven, Connecticut. vi + 108 pp. US $25.00. The publication in 1990 of Distribution and Taxonomy of Birds of the World by Charles G. Sibley and Burt L. Monroe became a landmark in ornithological literature (see review in The Canadian Field-Naturalist 107: 377-378, 1993) in spite of the fact that the classification proposed in it continues to be challenged. The authors did not consider their work as definitive and asked for suggestions or cor- rections. Readers responded to their request and “extensive” comments and suggestions were received. These and the changes identified by the authors form the basis for the current supplement. The supplement is divided into two parts and con- sists of a detailed account and compilation of the changes and corrections from the original work. The first section is short (13 pages) and deals with changes in classification, including sequences of species, and adjustments in scientific and English names. The second part (pages 14—108) is entitled “complete update” and comprises all the modifica- tions that the authors have brought their initial text. A World Checklist of Birds By Burt L. Monroe, Jr. and Charles G. Sibley. 1993. Yale University Press, New Haven and London. xiv, 393 pp. U.S. $50.00. This typical checklist contains the 9702 species and 2063 genera recognized by Sibley, Ahiquist, and Monroe in their earlier works, including a recent supplement. The classification adopted in this book is identical to that of their earlier books and has become “a taxonomic listing in the Sibley-Ahlquist- Monroe (SAM) classification”. A section on how to use the checklist appears in the short introduction and is followed by a list of the abbreviations and symbols which are abundantly used in the text. The book includes all the taxonomic categories found in the SAM classification, as well as an updated num- ber of genera and species. Each species occupies a single line and the scien- tific name is preceded by a box allowing one to record the sighting of that species. The English name follows with a brief outline of the geographic distri- bution and status of the species depicted by abbrevi- ations and symbols given in parentheses. Initially, the abbreviations may be confusing to follow but one becomes familiar with them after having They have revised the species counts for all cate- gories where such counts appeared in the earlier book. The changes deal mainly with the results of recent taxonomic work at the species level resulting in the recognition of species splits which resulted in different species counts at the higher taxonomic cat- egories. On the other hand, the overall classification at the higher taxonomic levels, based on earlier work on DNA-DNA hybridization by Sibley and associ- ates, remains as in the original book. Corrections and amendments to the original publication affect all sections including the world numbers, maps, gazetteer, references, and index. The present work is a soft-cover book that will be indispensible to all who need to consult the original volume or use its information but the high price of both works, particularly of the supplement, may dis- suade many potential buyers. It is hoped that all libraries where the original work is available will acquire the supplement. HENRI OQUELLET 175, avenue de la Citadelle, Hull, Québec, Canada J8Z 3L9 referred to the introduction a few times. The species line ends with a space permitting the user to register brief comments. Because the SAM classification is different from those used by the majority of ornithologists and bird watchers, the index of genera will be a useful tool to locate species in the list for those familiar with sci- entific names. Likewise, the extensive index of English names will be appreciated by those search- ing for the position of species in the list and who are not familiar with scientific names or the SAM classi- fication. This checklist is a high quality production with good typography, clear design, compact format, excellent binding, and strong paper. This may explain its high price. It should be useful and appre- ciated by those who keep records of their sightings of birds anywhere in the world and by those who have to refer to a complete listing of birds of the world in their work or studies. HENRI OUELLET 175 avenue de la Citadelle, Hull, Québec, Canada J8Z 3L9 376 THE CANADIAN FIELD-NATURALIST Vol. 110 Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics Jonathan Roughgarden. 1995. Oxford University Press, New York. xvi + 200 pp. U.S. $44.50. The past 20 years or so have seen a revolution in evolutionary ecology. Much of this revolution is due to the development and application of mathematical models. Catalyzing this advancement is the advent of desktop computers. Virtually everyone can now follow the models, and understand the predictions derived from them. Roughgarden, a recognized lead- er in the field of theoretical ecology, has apparently taken the next logical step in education, that of lucid- ly providing the theoretical basis of ecological theo- ry, along with the software to examine the conse- quences and predictions derived from various forag- ing strategies. Unfortunately, the PC software pack- age designed to accompany the book must be pur- chased separately ($22.50), and was not provided for review by the publisher. (Reader beware: the soft- ware package “is intended for readers familiar with computer programing.”’) Roughgarden’s new book is an attempt to synthe- size ecological, evolutionary and biogeographic theo- ries. The central topics include the mathematical models and applications for foraging strategies, com- munity assemblage, and food webs. He provides a discussion on the evolution of body size, competi- tion, and coevolution. He has attempted to intertwine these data with geological development to formulate a new vicariance theory for the evolution of the insu- lar anoles. The goal of the book is very lofty, and, unfortunately some of these goals fall far short of being realized. For example, although Roughgarden’s expertise in theoretical ecology is clearly evident, his understanding of the application of phylogenetic methods to questions of evolution is wanting. A sound phylogenetic analysis is central to discussions of evolution, be it body size, colonization sequences, or ecology. Much of his “phylogenetic analysis” has little to do with phylogeny, but rather is aimed at a controversy over whether character displacement has occurred once (the most parsimonious, but not pre- ferred explanation) or several times. As evidence for his point of view, Roughgarden cites an unpublished phylogenetic study that used size characters evaluat- ed by generalized gap coding. Apart from the circu- latity of evaluating the evolution of size on a phy- logeny constructed from size, generalized gap coding has been shown to be a poor, if not outright invalid, method for coding data. The extent of the phyloge- netic methods problem is obvious from his primary “cladogram” (Figure 3.2). It has albumin immuno- logical distances (AID) as synapomorphies! Evidence for one node is cited as “Genetic distance between wattsi pogus and other wattsi populations is greater than the distance between w. schwartzi and w. wattsi.” Cladograms should be based on inheritable characteristics, and organisms do not inherit genetic distances. There are other problems with the text that reveal a dated approach to biogeography and systematics. For example, in 1989 the anole lizards became mem- bers of the family Polychridae, and not Iguanidae; the former Iguanidae now consists of eight families. Similarly, the lizard family Agamidae has been sub- sumed as a subfamily into the Chamaeleonidae. Apart from the theoretical ecology, the book has little to offer. Overall, it is a wealth of information on theoretical ecology as it applies to anole lizards and is exceptionally easy to read, especially if one is well-versed in math. However, the conclusions about the evolution of the group, and their biogeography, must await a sound application of phylogenetic methodology. ROBERT W. MURPHY Department of Ichthyology and Herpetology, Royal Ontario Museum, 100 Queen’s Park, Toronto M5S 2C6 1996 BOTANY BOOK REVIEWS S)1/T/ An Orchid Flora of Puerto Rico and the Virgin Islands By James D. Ackerman. 1995. New York Botanical Garden, Bronx, New York. 203 pp., illus. Cloth US $35.00 Floras are essential tools of biologists because they provide a basis for research and for the under- standing and management of biodiversity. We need good floras and we need a lot more of them. Any good flora is a very significant contribution. This orchid flora contains descriptions of 145 species as well as information on distribution, habitat, flower- ing time, pollination, and classification. There are 97 plates illustrating approximately that many species as well as keys to genera and species. The book con- cludes with an explanation of the excluded species, literature cited, checklist and classification, a valu- able glossary in both English and Spanish, and an index to scientific names. Ackerman’s book is a very significant contribution. It is a model flora. The keys are mostly well con- structed, and the illustrations range from good to very good quality. It is accurate, easy to use, current and essentially complete with respect to the information available and the species covered. The fact that the keys are in both Spanish and English reflects a very sensible decision to increase communication and utili- ty since Spanish is the major language of researchers in the surrounding region. The “taxonomic notes” are especially useful in explaining decisions of the author regarding classification and sometimes even provid- ing rather extensive taxonomic histories and new information on patterns of variation. This flora will be of use to anyone interested in the plant life of the region covered. It will also be of great value to specialists in orchid systematics and ecology, and researchers generally. Orchid growers and hobby- ists may also find it useful, but of limited scope. The only areas where this book falls a little short are in the four-page introduction and the alphabetical sequence of genera. Genus names change and when things are organized that way one cannot always find them easily. To identify plants and collect information about related groups it helps to have all related and similar groups together. That is the reason for a phylo- genetic classification system, for expending effort to develop a reliable system, and for using it. Users of floras however, often disagree on the relative merits of phylogenetic versus alphabetical organization. The introduction briefly describes geography, cli- mate, the six life zones, orchid collections providing the basis for the work, species richness, endemics, geographical associations, and text organization. Considering that all these subjects are covered on four pages the reader can imagine that “brief” is given new meaning. However, in these days of con- straints on funding, it is increasingly necessary to frame work within the needs and interests of the largest possible user group, and to provide justifica- tions and fit work into a bigger picture. That is why an introduction is an opportunity. A few pages on orchid conservation in Puerto Rico would have been interesting in view of the fact that the majority of the orchids in Puerto Rico evidently occupy an ecologi- cal zone (lower montane rainforest) that covers less than 0.1% of the island. This must certainly mean a challenge for conservation efforts. Details on pat- terns of occurrence, major contributions of orchid research on the island to orchid biology, and biology generally (there have been some), more details on how this work fits into the larger region, a brief dis- cussion of research directions for the Carribean region would all have been useful. Even a paragraph introducing the biggest and one of the most taxo- nomically complex (and irresistable) plant families in the world would have taken advantage of an opportunity. The 20 000-25 000 species indicated under the family description by Ackerman is a con- servative estimate. The introduction is lacking in illustrations and diagrams. Only one map is included (political subdi- visions on the endpapers). Consequently it is hard to get a quick handle on the place and the orchid flora. In summary we have here a model flora with a brief introduction and an alphabetical instead of nearest neighbour arrangement. It is reassuring to see a prominent scientist like Ackerman (at least 50 refereed papers of exceptional quality on ecology, pollination, and systematics) step away from the frontier for a while to provide a foundation for future research and help us cope with the biodiversity out- side the front door. PAuL M. CATLING 8 Scrivens Drive, R.R. 3, Metcalfe, Ontario KOA 2P0 378 ENVIRONMENT THE CANADIAN FIELD-NATURALIST Vol. 110 Radiation Hazards to Fish, Wildlife and Invertebrates: A Synoptic Review By Ronald Eisler. 1994. Biological Report 26, Contaminant Hazard Reviews Report 29, U.S. Department of the Interior, National Biological Services, Washington D.C. 20240. 124 pp., illus. No cost. - Ronald Eisler has compiled data from a large list of sources to provide a compendium of reported haz- ards to various types of wildlife. His introductory chapter gives a brief explanation of ionising radia- tion, radionuclides, and measurement terms. He then looks at both natural and human-generated sources of radiation in terms of their origin and potential magnitude. The next chapter, on radionuclides in the environment, is broken into sections on abiotic mate- rial, aquatic ecosystems, birds, and mammals. Finally he looks at the effects of radiation on several forms of living organisms. The information is drawn from 352 references listed at the back of the report, along with a short glossary. My immediate reaction to this work was that the author had produced a compact useful reference that I would use time and again. I did have some misgiv- ings about the inferences created by the form of the sentences. For example, he often lumps nuclear power and weapons together in the same phrase. The reader is left with the impression that these sources contribute equally to human dose. Later, when he discusses the data, you can see the measured differ- ence in contribution but it can be hard to shake those first impressions. Some paragraphs have a curious sequence of sentences. The section on the Great Lakes tells of “significant” amounts of radioactivity from reactors and mine waste but gives no data to establish what is meant. The second sentence talks of “low levels” of fallout nuclides and is followed by a table of results with a cumulative total of over 21 x 10° Bq/km’. Incidently it is unclear how this table was derived and if it accounts for decay over the 30 year collection period. As I dug deeper I became more concerned. I began to question the data itself. Table 5 is a typical annual whole body dose from all sources. Global weapons fallout is quoted as 0.05 mSv, a little on the high side (0.04 mSv is a more typical estimate) but not worth an argument. Table 7 shows a breakdown by nuclide of this fallout dose. The total from fallout is 4.45 mSv. The author notes this is 1.85 times natu- ral background but dées not question or comment on this unusually high figure. For comparison the aver- age annual dose to Canadian atomic workers in 1994 was 2.86 mSv. A potentially confusing point is the practice of quoting doses to animals in mSv units, which is the unit used for equivalent and effective doses in humans. The relationship between absorbed dose and equivalent and effective doses has not been established for non-human species. It would be preferable to state the dose in units of absorbed dose, i.e. the gray (Gy), and provide information on the type of radiation. This inconsistency may not stem from the author but those whom he is quoting. Nevertheless, he needs to explain the limitations on the data presented. Much of the time I noted that the questionable data came from a few specific references that I was not familiar with. I reviewed the reference list and realised that many Canadian authors were missing. There is no mention of Swanson and her colleagues at University of Saskatchewan, of Brunskill and coworkers at the Experimental Lakes Area or of Sheppard at AECL’S Pinewa research unit to name a few. I felt I was being parochial by thinking only of Canadians so I checked for European material and I was left wondering. There are only two IAEA docu- ments quoted out of the whole library of reports pro- duced by that agency. Finally when I came to use this document to find out information I had a hard time. I wanted to know the lethal dose to bacteria. The first problem I encountered is the lack of an index so you have to thumb through the document. I eventually found a value for protozoans, as close as I could come to bacteria, in a large table. It was some time later I noticed a small bar chart that included bacteria and was able to estimate that ten to over 10 000 Gy were required to achieve a 50% death rate. This is a useful reference report and I believe I will find that I will use it frequently. However, I will not use the data from it until I have verified its accu- racy. I would prefer that it be rewritten into a text I can both use and trust. The author needs to remem- ber that this is a U.S. Government publication and this automatically confers a level of credibility. The author is responsible for ensuring the information given is accurate and can be read in context. A good editorial stance is needed to produce statements that are understandable and cannot be misquoted. Roy JOHN 1-613 McPherson Avenue, Saskatoon, Saskatchewan S7N 0X7 1996 Prehistoric Alaska Edited by P. Rennick. 1994. Alaska Geographic Volume 21, Number 4. The Alaska Geographic Society, P.O. Box 93370, Anchorage, Alaska 99509-3370, U.S.A. 112 pp., illus. U.S. $19.95. Alaska Geographic, published since 1972, pro- duces quarterly issues, each thematic and focussed on a specific aspect of Alaska’s geography or natural history. This colourful and interesting issue concen- trates on “Prehistoric Alaska’, referring to all geo- logic time. It consists of eleven main chapters; most chapters are written by scientific experts, a couple are contributed by professional writers. The first four chapters deal with geologic and palaeoenvironmental themes; the final seven chapters focus on the region’s human history. The opening article (“Prehistoric Alaska: The Land” by F. H. Wilson and F. R. Weber) describes the geologic development of Alaska and is preceded by a timeline for orientation. This chapter concen- trates on the assembly of the various terranes that make up the region. It leaves an impression of a highly mobile system, albeit on a vast time-scale. Although forming the immediate background and context for the human history, less attention is paid to Quaternary glaciation or geomorphology. Though it includes a series of maps, I found this chapter heavy-going and hard to follow. “The Terrible Lizards” or dinosaurs are next, described by staff writer L. J. Campbell and accompanied by a dinosaur bestiary. Dinosaur fossils from Alaska are a relatively new discovery. Despite hints in the 1960s and 1970s, major finds were only made in the 1980s. Serious scientific attention and exploration for new sites have occupied barely a decade. Most fossil have been recovered from the Colville River valley, not far inland from the Arctic coast. Dinosaur remains in such high latitudes have important impli- cations for the interpretation of their ecology. Campbell emphasizes that Alaska is a huge area that will take many years to survey for suitable fossil localities and that there are few people and restricted funds for doing the work. The same comments could equally well apply to the archaeological resources discussed later. The dinosaur discussion places emphasis on a Late Cretaceous Land Bridge connec- tion between Asia and North America. This is not mentioned in the geology section and, indeed, the illustrations included in that chapter leave the impression that there has been no such land connec- tion until the Pleistocene. The next two chapters (“Alaska Vegetation: What the Fossil Record of the Past 20 Million Years Shows” by T. Ager and “Pleistocene Mammals” by P. E. Matheus) focus on biological themes arising from the botanical and faunal records from Alaska. Our modern image of Alaska as windswept tundra BOOK REVIEWS 379) and coniferous forest needs modification. Ager’s dis- cussion shows that Alaska has at times, such as the Early and Early-Middle Miocene, supported broadleaved thermophilous trees, such as lime (Tilia), hickory (Carya), and elm (Ulmus). These remains also suggest that the climate was considerably warmer than present, Ager estimates by up to 12° — 18°F mean annual temperature. Widespread conifer- ous vegetation has developed since the late Tertiary. These ancient forests and grasslands (“Mammoth Steppe’) were inhabited in the late Pleistocene by a varied assortment of now extinct and extant fauna, including the eponymous mammoth. Matheus pres- nets an interesting discussion of the mammals’ ecolo- gy, showing how they could co-exist by exploiting slightly different niches in the Beringian landscape. He tackles the fascinating question of the demise of the Pleistocene megafauna by offering two linked explanations. Changing environments may have made the terrain less favourable for some species, such as camels and horses, which are extirpated in Alaska although living elsewhere, and mammoths, which are extinct. Matheus suggests that other mam- mals, especially carnivores such as cheetahs and lions, were simply too specialized to be capable of exploiting the changed Beringian environment. Animals that had the most success were those such as wolves and brown bears that could exploit a variety of prey species and had more eclectic dietary habits. The human history section opens with a back- ground article by W. Workman (“The First of the Last: Pioneer Human Settlers on the Last Frontier’’) that provides a succinct chronologic survey of the main archaeological subdivisions in Alaska. The next five short chapters each concentrate on a differ- ent region and/or site in Alaska. Workman outlines several themes in Alaskan archaeology including the role the region played in the initial incursion of peo- ple from Asia to North America, whenever that might have occurred. From his account, the oldest generally-accepted archaeological sites found so far in Alaska date to the millennium 12 000 — 11 000 yr BP. Two of these ancient sites are highlighted in these chapters. Science writer Lee Dye describes the Mesa site, located on the north slope of the Brooks Range, which may date to 11 700 yr BP. The account of the furore surrounding the press confer- ence to announce the site called by the Bureau of Land Management makes for instructive reading: the public found it fascinating, some colleagues found it outrageous. The importance of serendipity in site discovery is clear from this account. D. R. Yesner and K. J. Crossen introduce the Broken Mammoth site in central Alaska where the earliest archaeologi- cal remains date from about 11 800 yr BP. Whether these and similarly-aged sites really record “the environment and lifeways of the first Alaskans” 380 (page 92), as Yesner and Crossen claim for the Broken Mammoth site, remains a matter of contro- versy. What is clear, however, is that by the early Holocene people were established throughout Alaska. Among the slightly later sites are Trail Creek, dating from 9500 yr BP, mentioned by Jeanne Schaaf in her discussion of “Seward Peninsula Prehistoric Lifeways”, and Ground Hog Bay in the Alaska Panhandle, also dating to 9500 yr BP, includ- ed in W.M. Olson’s survey of “A Prehistory of Southeast Alaska”. Workman comments that “the question of early man is only one of a number of interesting and important topics in Alaska archaeology” and makes the point that “recent prehistory is both more approachable ... and perhaps of greater relevance” to today’s Aboriginal people (page 75). On this theme, D. W. Veltre describes the problems of reconstruct- ing pre-Contact Aleut cultures from the fragmentary archaeological record and oral traditions. Some sites may have considerable antiquity; Veltre mentions that the location of the modern village of Nikolski may have been continuously occupied for the last 4000 years. Much of Olson’s discussion of southeast Alaska also concentrates on later prehistory, espe- cially the emergence of the varied coastal Aboriginal groups. A captivating narrative by Herbert Anungazuk, describing a prehistoric hunter’s experi- ence of whale-hunting in western Alaska, puts a human face to the archaeology. The authors in this section adopt a conservative position and espouse a late entry (post-Late Wisconsinan maximum) position for the first migra- tion of people into North America via Alaska. On this theme, the final chapter, “Molecular Evolutionary Genetics of Indigenous Northern Populations”, is an attempt by G. F. Shields to put a more objective spin on this debate. Shields reports on studies of mitochondrial DNA, which is transmit- ted through the female line and can be used as a “molecular clock”. These analyses show groups of different genetic diversity. Mapped data show clear differences between Aboriginal populations in north- ern North America (and Siberia) and those to the south. Shields points out some incongruities between the pattern and chronology produced by genetic and linguistic evidence, noting that genetic and linguistic groups do not coincide. It would be useful to com- pare these data with dental evidence analyzed by physical anthropologists such as Christy Turner. The main body of the text is followed by a list of museums displaying relevant Alaskan geological, archaeological and ethnographic material, a glossary, a bibliography, and an index. The glossary consists exclusively of geologic terms. I think it would have been useful to include some archaeological terms, such as “component”, “bifacial tools”, and “‘burin”’. The bibliography contains much useful material. However, general and highly technical publications THE CANADIAN FIELD-NATURALIST Vol. 110 are combined. I would prefer these readings arranged in the same subdivisions as the chapters, making it easier for a reader to follow-up on a specific topic, with the more general readings, such as Dale Guthrie’s excellent book on Blue Babe, highlighted in a “Further reading” section at the beginning. I was surprised that E. C. Pielou’s outstanding book (After the Ice Age: The Return of Life to Glaciated North America) is not listed; as an accessible introduction to the complex topic of palaeoecology it would be a useful addition. The text is profusely illustrated with maps, paint- ings, and especially colour photographs. Most of these images are excellent, particularly those illustrating the various archaeological sites. Having read much about Alaskan sites in recent years, I was especially interest- ed in the archaeological photographs. These include artifact pictures and excavation scenes as well as gen- eral scenery shots. The pictures of Blue Babe and, in particular, the ground squirrel carcass (page 57) require scales. The photomicrograph of the pollen slide (page 42) needs an indication of magnification or scale. A location map (page 8) includes the sites mentioned in the articles. Among the illustrations are two major full-page paintings by local artist Tom Stewart, one of dinosaurs, the other of Pleistocene mammals. They are certainly eyecatching, although I must admit that personally I did not like them. The dinosaur painting in particular suffers by juxtaposition with the truly excellent reconstructions by Jan Sovak and Vladimir Krb. Although human history is the focus of a third of the text, this painting is used as the cover illustration, Most writers have managed to keep the technical jargon to a minimum but there are a few places where it slips in. In this regard, the chapter by Olson stands out, including unexplained terms such as “detachment of platform tablets”, and “moiety sys- tem”. There are a few intrusive spelling and gram- matical errors, e.g., “forbs” is consistently given as “forbes”, and “gate” appears for “gait”. Most chap- ters are clear, absorbing and well-written, but I did note some awkward sentence structures, especially in Campbell’s contribution. Writing for the public is very difficult — I speak here from personal experience — and the selection of material from such a vast region and subject-area must have been hard. This volume is a worthy endeavour and clearly represents the outcome of much hard work by the authors and production team. Criticisms aside, I feel that this issue of Alaska Geographic gives a valuable introduction to some fascinating topics in a complex and diverse region. It deserves to be widely-read. ALWYNNE B. BEAUDOIN Provincial Museum of Alberta, 12845-102nd Avenue, Edmonton, Alberta TSN OM6 1996 Functions of Nature By Rudolf S. deGroot. 1992. Wolters-Noordhoff, Groningen, Netherlands. 345 pp. illus. Nfe 80. Twenty-first century historians will mine the stacks, CDs, computerized data bases, and many other yet to be invented electronic searching tools to examine humanity’s march through time. From their computers will spill detailed and varied accounts of the forces, factors, and events that shaped the evolu- tion of human culture, society, and economy. Historians will dissect, expose, highlight, analyze, and perhaps provide prophetic advice for their read- ers. And while human conflict will persist as a domi- nant theme of their “backgazing”, I anticipate that humanity’s struggle to find its place in the ecosphere will be a rich area of investigation as well. I expect that the 1990s will be described as a decade of global chaos, of continued and increasing ecospheric degradation, of economic upheaval, and continued human conflict. But too, historians just may be able to describe the 1990s as a “turn around” decade, a decade in which societies throughout the World accepted and embraced the need for signifi- cant and unprecedented change, change characterized by a commitment to ecological sustainability, change focused on finding balance between people who use the ecosphere and the capacity of the ecosphere to provide for human survival, security, a sense of belonging, self-esteem, and self-actualization. Among the thousands of books and articles avail- able to them, historians will look for key documents that helped shape the 21st Century. Functions of Nature may well be selected for mention in the foot- notes as one of the early attempts to comprehensive- ly grapple with techniques to describe and value the many forces and factors that drive the Earth’s ecosystems and the people who live and work in them. A primary purpose of the book is to contribute to the development of methods that translate envi- ronmental data into applied information for planning and decision-making in more objective and system- atic ways. In Chapters one and two, deGroot describes eco- logical evaluation and assessment methods, and defines 37 functions that he believes capture the spectrum of values required to plan for and make ecologically-based decisions. The functions are orga- nized according to four themes. “Regulation Functions” reflect the capacity of the ecosphere to provide and regulate ecological processes and life- support systems. They include the regulation of ener- gy balance, chemical composition of the atmosphere and the oceans, fixation of the solar energy and biomass production, and storage and recycling of organic matter, nutrients, and human waste. “Carrier ~ BOOK REVIEWS 381 Functions” characterize the capacity of ecosystems to provide space to meet the physical needs of humans, including habitation, cultivation, industry, transportation, recreation, and nature protection. The “Production Functions” define natural resources that humans use to survive and prosper, including air, water, food, fuel and energy, fertilizer, medicines, and raw materials for clothing. The “Information Functions” contribute to human psychological health by providing opportunities for educational, spiritual, and cultural development and experiences. Integrated planning and decision-making is a central theme throughout these two chapters. Chapter three describes many of the values ascribed to ecosystems, outlines various methods available to assess the monetary value of environ- mental functions, and provides examples. Values include: conservation value, existence value, human health value, option value, consumptive use value, and productive use value. Chapter four contains three case studies employing these functions and val- ues in a moist tropical forest ecosystem (Darien National Park in Panama), a temperate wetland ecosystem (Dutch Wadden Sea), and an oceanic island ecosystem (Galapagos National Park in Ecuador). In the concluding chapter, deGroot argues that the concept of “sustainability” must be a prima- ry element in decision-making, and describes the use of the “functions of nature” in planning, environ- mental impact assessment, and cost-benefit analyses. The text is punctuated with examples, illustra- tions, tables, and the parameters of the 37 functions are listed and defined in an Appendix. This is an important book for people working to understand ecological sustainability and the design and imple- mentation of an ecosystem approach to management. It contributes to current efforts aimed at defining and implementing an holistic and integrated approach to caring for the ecosphere’s natural assets, including the people who live and work in it. It is regrettable that deGroot did not write this book within the con- text of an ecologically meaningful spatial framework — a context in which people and organizations can use described and mapped ecosystems to understand and manage for the array of ecological, cultural, social, and economic conditions and forces at work in the World. Evaluating the ecosphere’s natural assets and determining the roles and responsibilities of humans who live and work in it is a complex sub- ject. deGroot has made a good start. PAUL A. GRAY Department of Natural Resources, PO Box 8070, Causeway, Harare, Zimbabwe 382 THE CANADIAN FIELD-NATURALIST Vol. 110 Biodiversity in Canada: A Science Assessment for Environment Canada By Biodiversity Science Assessment Team. 1994. Environment Canada, Ottawa. 245 pp., illus. + 24 pp. summary volume. Free. If you are looking for a book on the biodiversity of Canada, this is not it. Rather, this is a science assessement on biodiversity issues. What is a science assessement? The preface of the book claims that a science assessment “reviews the state of knowledge on a particular issue or topic in order to identify implications for policy and for further research.” Separate chapters address threats to biodiversity from a variety of land uses (e.g. forestry and agri- culture) and other environmental pressures such as the introduction of exotic species and pollution. The books summarizes a vast amount of research on bio- diversity issues, unfortunately the chapters are wild- ly uneven in their thoroughness. For example, the literature cited section alone for the chapter on agri- culture is over twice as long as the entire chapter on forestry! One of the strong points of the book is its focus on what should be done. Each chapter contains recom- mendations for additional research and policy direc- tions. While some are needlessly vague, they still provide some guidance on these issues. It remains to be seen if the Government of Canada is indeed com- mitted to preserving biodiversity. To decide for yourself if the government is heading in the right direction, you can obtain the complete book or a summary booklet (with the key recommendations) free of charge from Environment Canada in either English or French (819-997-1095). DAVID SEBURN Seburn Ecological Services, 930 River Road South, R.R.4, Kemptville, Ontario KOG 1JO Restorations of Endangered Species: Conceptual Issues, Planning and Implementation Edited by Martin L. Bowles and Christopher J. Whelan. 1994. Cambridge University Press, Cambridge. xiii + 394 pp., illus. U.S. $49.95. The reintroduction or restoration of endangered species 1s increasingly common as more and more species become threatened with extirpation or extinction. This handy volume compiles a number of papers, most of which were first presented at a sym- posium on the subject held in 1990. As the sub-title suggests, the papers tackle three main themes: 1) Conceptual Issues — from organizational guidelines for recovery teams to genetic considerations for viable populations; 2) Restoration Planning — case studies that outline factors that should be considered ahead of time such as the role of biological invasions on the management of rare plant species and the effects of metapopulation dynamics on the long-term Everglades: The Ecosysem and Its Restoration Edited by Steven M. Davis and John C. Ogden. 1994. St. Lucie Press, Delray Beach, Florida. xv + 826 pp., illus. + map. U.S. $110. As humankind continues to gain knowledge of the biosphere it lives in, the significance of previous undisturbed ecosystems comes to light. With this enlightenment comes the realization of the need to attempt restoration of what once was. Everglades: the Ecosystem and Its Restoration is a treatise which provides the details of such an enlightment describ- success of restorations; 3) Implemented Restorations — results of various restoration projects that have been undertaken, including the swift fox program in western Canada. The strength of this book is its integration of dif- ferent fields: theory and practice, botany and zoolo- gy. It’s rare to see an article on caribou next to one on a thistle. Nonetheless, the diversity of fauna cov- ered could have been broader, as all three of the ani- mal restoration papers are devoted to mammals. Still, this is a valuable reference tool for wildlife managers and could be an excellent textbook for a senior level course in restoration ecology. DAVID SEBURN Seburn Ecological Services, 930 River Road South, R.R. 4, Kemptville, Ontario KOG 1JO ing “details of the past and present Everglades,” with “its justification” only being “realized in the future Everglades.” The book is stated to be “an outgrowth of the 1989 Everglades Symposium in Key Largo, six adaptive environmental assessment workshops, intensive com- munication and interaction among contributors ... of topics through 1992.” Everglades has a contributing authorship of 57 authors with 27 institutional affilia- tions. There are 31 chapters, with an emphasis on 1996 hydrologic aspects, organized in five sections: (I) The Everglades Issues in a Broader Perspective, (II) Spatial and Temporal Characteristics of Ecosystem Driving Forces, (III) Vegetation Components and Processes, (IV) Faunal Components and Processes, and (V) Toward Ecosystem Restoration. Each chapter has an abstract. The text is organized in a logical and very readable manner. Davis and Ogden provide a detailed overview of a unique ecosystem and its problems. One lesson the book emphasizes is the enormous cost disruption of a key ecosystem and the resulting attempt at restora- tion can have. A restoration which, as the editors BOOK REVIEWS 383 state in the final chapter, will provide a new Everglades, not the Everglades of the past, and if successful will rekindle the wildness and richness of the former system. The book is not the final chapter in the restoration of the Everglades but an update and identification of possible future direction. Everglades: the Ecosystem and Its Restoration should be of interest and a good read to those with an interest in the Everglades as well as wetland restoration. M. P. SCHELLENBERG 434 4th Avenue SE, Swift Current, Saskatchewan S9H 3M1 Reinventing Nature? Responses to Postmodern Deconstruction Edited by Michael E. Soulé and Gary Lease. 1995. Island Press, Washington, D.C. xvii + 186 pp., illus. Cloth U.S. $34.95. Paper U.S. $17.95 The world of humanities often questions the valid- ity of the scientific or technical view of nature. In Reinventing Nature the editors furnish a number of essays collected from a conference in Santa Cruz which was-organized for an exchange of views between the worlds of the natural sciences and the humanities. At the conference and in the book the editors wanted to raise the question “whether per- ceptions and conceptions of nature ... differ enough between cultures to affect the way these cultures would wish to maintain or manage nature in the rem- nants of remaining habitat.” Soulé and Lease provide the reader a dialogue from leading thinkers from the fields of philosophy, history, literature, public policy, forestry, and others. The reader is first provided the essay abstracts and then a short glossary. Both items are useful and ide- ally located for help while navigating through the discussion. The discussion reveals “certain contemporary forms of intellectual and social relativism can be just as destructive to nature as bulldozers and chain saws.” For this reason alone this book should be a must read for individuals in conservation biology, restoration, and ecology work to name a few. The editors provide a work which will provide a feast for continued dialogue on “nature”. M. P. SCHELLENBERG 434 4th Avenue SE, Swift Current, Saskatchewan S9H 3M1 384 NEw TITLES Adaptation and natural selection in caves: the evolu- tion of Gammarus minus. 1995. D. C. Culver, T. C. Kane, and D. W. Fong. Harvard University Press, Cambridge, Massachusetts. 223 pp., illus. U.S. $ 35. Atlas des oiseaux richeurs du Québec méridionol. 1995. Par J. Gauthier et Y. Aubry. L’ Association québe- coise des groupes d’ornithologues, CP 1000 Succursale M, Montreal HIV 3RZ. xviii + 1295 pp., illus. $150. Available also in English. +Atlas saisonnier des oiseaux du Québec. 1995. Par A. Cyr et J. Larivée. Presses de I’ Université de Sherbrooke et Société de Loisir ornithologique de I’ Estrie, Sherbrooke. viii + 709 pp., + transparent, illus. $56.95 au Canada; U.S. $56.95 extérieurs. A chorus of frogs. 1995. By J. P. Hunt. Silver Burdett Press, Parsippany, New Jersey. 40 pp., illus. Cloth U.S. $14.95; paper U.S. $7.95. The company of wolves. 1995. By P. Steinhart. Knopf, New York. xviii + 374 pp., illus. U.S. $25. +A critical review of the aerial and ground surveys of breeding waterfowl in North America. 1995. By G. W. Smith. Biological Science Report 5. National Biological Service, Washington. ili + 252 pp., illus. Free. The grizzly bears of Yellowstone: their ecology in the Yellowstone ecosystem, 1959-1992. 1995. By J. J. Craighead, J. S. Sumner, and J. A. Mitchell. Island Press, Washington. 560 pp., illus. U.S. $100. Hawks, Owls, and other birds of prey. 1995. By. D. Fourie. Silver Burdett Press, Parsippany, New Jersey. 40 pp., illus. Cloth U.S. $14.95; paper U.S. $7.95. High country heritage: New England’s mountain flow- ers. 1996. By. J. Wallner and M. J. DiGregorio. Mountain Press, Missoula, Montana. 224 pp., illus. U.S.$16. Insects through the seasons. 1996. By G. Waldbauer. Harvard University Press, Cambridge, Massachusetts. 304 pp., illus. U.S.$24.95. In the country of gazelles. 1995. By F. R. Walther. Indiana University Press, Bloomington. 1x + 162 pp., illus. U.S. $24.95. *Masters of the ocean realm: whales, dolphins, porpoises. 1995. By J. E. Heyning. UBC Press, Vancouver. 112 pp., illus. $24.95. The origins of right and wrong in humans and other animals. 1996. By. F. deWaal. Harvard University Press, Cambridge, Massachusetts. 384 pp., illus. U.S. $24.95. The others, how animals made us human. 1996. By P. Shepard. Island Press, Washington. 390 pp., illus. U.S. $24.95. The penguins: Spheniscidae. 1995. By T. D. Williams. Oxford University Press, New York. xvi + 295 pp., illus. USS. $60. +Proceedings of the 6th International Grouse Sym- posium. 1995. Edited by D. Jenkins. Proceeding of a sym- posium Udine, Italy, 20-24 September 1993. World Pheasant Association, Reading, United Kingdom. iv + 175 pp.. ulus. £15. THE CANADIAN FIELD-NATURALIST Vol. 110 Seals and sea lions. 1995. By D. G. Gordon. Monterey Bay Aquarium, Monterey Bay California. 64 pp., illus. U.S. $9.95. *Shells of Atlantic and gulf coasts and the West Indies. 1995. By R. T. Abbott and P. A. Morris. Peterson Field Guide Series. Houghton Mifflin, Boston. 350 pp., illus. U.S. $26.95. Whales, dolphins, and porpoises: the visual guide to all the world’s cetaceans. 1995. By M. Carwardine. Dorling Kindersley, New York. 256 pp., illus. cloth U.S. $ 29.95; paper U.S. $17.95. *Where to watch birds in Africa. 1995. By N. Wheatley. Princeton University Press, Princeton. 432 pp., illus. U.S. $35. +The wind masters: the lives of North American birds of prey. 1995. By P. Dunne. Houghton Mifflin, New York. U.S. $22.95. Botany The book of rhododendrons. 1995. By M. Kneiler. Timber Press, Portland, Oregon. 160 pp., illus. U.S. $45. +Botanical reconnaissance of the Tuxedni Wilderness Area, Alaska. 1995. By S. S. Talbot, S. L. Talbot, and S. L. Welsh. Biological Science Rept. 6. National Bilogical Service, Washington, 41 pp., illus. Free. Canada’s vegetation: a world perspective. 1995. By G. A. J. Scott. McGill-Queen’s University Press, Montreal. Xvill + 361 pp., illus. Ericas of South Africa. 1995. By D. Schumann and G. Kirsten. Timber Press, Portland, Oregon. 274 pp., illus. WS.'$5995; Flora of the Venesuelan Guyana, Volume 1: introduc- tion and Volume 2: Pteridophytes, Spermatophytes (Acanthaceae-Araceae). 1995. Edited by J. A. Stéyer- mark, P. E. Berry, and B. K. Holst. Timber Press, Portland Oregon. c400 pp., illus. U.S. $49.95 and c650 pp., illus. US. $65. +Flore laurentienne. 1995. Par Frére Marie-Victorin. Troisiéme édition. Les Presses de 1’ Université de Montréal, Montréal. xv + 1083 pp., illus. $65. +Greenland lichens. 1995. By. E. S. Hansen. Rodos, Copenhagen. 124 pp., illus. DK. 150. *A history of the orchid. 1995. By. M. A. Reinikka. Timber Press, Portland, Oregon. 340 pp., illus. U.S. $29.95. Lecythidaceae of a central Amazonian moist forest. 1995. By S. A. Mori and N. Cunha. Memoirs vol. 75. New York Botanical Garden, Bronx. 55 pp., illus. U.S. $ 12.590. Manual of orchids. 1995. Edited by J. Stewart. Timber Press, Portland, Oregon. 403 pp., illus. U.S. $49.95.“ +Mountain plants of the pacific northwest: a field guide to Washington, western British Columbia, and South- eastern Alaska. 1995. By. R. J. Taylor and G. W. Douglas. Mountain Press, Missoula, Montana. vi + 437 pp., illus. U.S. $ 20. 1996 Nueva flora de Chile. 1995. Editado por C. Marticorena y R. A. Rodriguez. Volume 1. Proyecto Flora de Chile, Universidad de Concepcion, Concepcion, Chile. Ch$ 20,900 (U.S. $70). Plants and their names: a concise dictionary. 1995. By R. Hyamond and R. Pankhurst. Oxford University Press, New York. x + 545 pp. U.S. $29.95. The prairie keepers: secrets of the grassland. 1995. By. M. Houle. Addison-Wesley, New York. 266 pp., U.S. $20. +Plants of the maritimes: a bibliography for agricul- ture, resource management, landscape planning, and biological research. 1995. By P. M. Cattling, S. Porebski, and B. S. Brookes. CanaColl Foundation, 1010 Carling Avenue, Ottawa, Ontario K1A 0C6. 65 pp. $10. The rhododendron species, volume iv: azaleas. 1995. By H. H. Davidian. Timber Press, Portland, Oregon. 184 pp., illus. U.S. $54.95. *Sonoran desert plants: an ecological atlas. By R. M. Turner, J. E. Bowers, and T. L. Burgess. University Arizona Press, Tuscon. 500 pp., illus. U.S. $70. “La symbiose mycorhizienne: états des connaissances. 1995. Edité par J. A. Fortin, C. Charest, et Y. Piche. Orbis, Frelighsburg, Québec. viii + 195 pp., $24.95. Utah wildflowers: a field guide to northern and central mountains and valleys. 1995. By R. J. Shaw. Utah State University Press, Logan. v + 218 pp., illus. Environment *Biodiversity and conservation of neotropical montane forests. 1995. Edited by S.P. Churchill, H. Balslev, E. Forero, and J.L. Luteyn. Proceedings of a symposium, the New York Botanical Garden, Bronx, June, 1993. New York Botanical Garden, Bronx. 700 pp. U.S.$ 85. *Conservation biology in theory and practice. 1995. By G. Caughely and A. Gunn. xii + 459 pp., illus. U.S. $44.95. Ecology and management of tidal marshes: a model from the Gulf of Mexico. 1995. Edited by C. L. Coultas and Y. P. Hsieh. St. Lucie Press, Delray Beach, Florida. e325 pp., illus. U.S. $59.95. +Ecology of infectious diseases in natural populations. 1995. Edited by B. T. Grenfell and A. P. Dobson. Cambridge University Press, New York. xii + 521 pp., illus. U.S. $59.95. y+An ecosystem approach to living sustainably: a per- spective for the Ministry of Natural Resources. 1995. By P.A. Gray, L. Demal, D. Hogg, D. Greer, D. Euler, and D. DeYoe. Ontario Ministry of Natural Resources, Peterborough. vi + 77 pp., illus. Free. *Kvolutionary naturalism. 1995. By. M. Ruse. Toutledge, New York. x + 361 pp., illus. U.S. $49.95. {Expanding partnerships in conservation. 1995. Edited by J. A. McNeely. Island Press, Washington. xvi + 302 pp., U.S. $34.95. {This fragile land: a natural history of the Nebraska sandhills. 1995. By P. A. Johnsgard. University Nebraska Press, Lincoln. xv + 256 pp., illus. U.S. $35. BOOK REVIEWS 385 *Fundamentals of conservation biology. 1995. By M. L. Hunter, Jr. Blackwell Science, Cambridge, Massachu- setts. xiv + 482 pp., illus. U.S. $42.95. Guardians of the parks: a history of the National Parks and Conservation Association. 1995. By J. C. Miles. Taylor & Francis, Washington. xviii + 363 pp., illus. U.S. $29.95. Invasions of the land: the transitions of organisms from aquatic to terrestrial life. 1995. By M. S. Gordon and E. C. Olson. Columbia University Press, New York. xv + 312 pp., illus. U.S. $65. Marine conservation for the 21st century. 1995. By H. Viders. Best, Flagstaff, Arizona. xvi + 350 pp., illus. Cloth U.S. $29.50; paper U.S. $24.95. The national parks of Canada. 1994. By. K. McNamee. Key Porter Books, Toronto. 224 pp., illus. $45. The northern forest. 1995. By D. Dobbs and R. Ober. Chelsea Green, White River Junction, Vermont. xxvi + 356 pp., U.S. $23. Our national park system: caring for America’s great- est natural and historic treasures. 1995. By D. F. Rettie. University of Illinois Press, Urbana. xvi + 293 pp., illus. Partnership for the planet: an environmental agenda for the United Nations. 1995. By H. F. French. World- watch Institute, Washington, 71 pp., illus. U.S. $5. *A place to walk: a naturalist’s journal of the Lake Ontario Waterfront Trail. 1995. By A. Karstad. Natural Heritage/Natural History, Toronto. 120 pp., illus. $19.95. The value of life: biological diversity and human soci- ety. 1996. By S. R. Kellert. Island Press, Washington. 280 pp., illus. U.S. $24.95. +Wild ideas. 1995. Edited by D. Rothenberg. Univeristy Minnesota Press, Minneapolis. xxvii + 225 pp., Cloth U.S. $49.95; paper U.S. $19.95. +Wild Thailand. 1995. By B. Stewart-Cox. MIT Press, Cambridge, Massachusetts. 208 pp., illus. U.S. $40. The world of wilderness: essays on the power and pur- pose of wild country. 1995. Edited by T. H. Watkins and P. Byrnes. Roberts Rinehart, Boulder Colorado. Miscellaneous Always Rachel: the letters of Rachel Carson and Dorothy Freeman, 1952 - 1964. 1995. Edited by M. Freeman. Beacon Press, Boston. xxx + 567 pp., illus. U.S. $35. *+Minutes of meetings, 1932 - 1935, of the McIlwraith Ornithological Club, London, Ontario, Canada. 1996. By W. W. Judd. Phelps Publishing, London. 111 + 106 pp., $10. Darwinism evolving: systems dynamics and the geneal- ogy of natural selection. 1995. By D. J. Depew and B. H. Weber. MIT Press, Cambridge, Massachusetts. xii + 588 pp., illus. U.S. $49.95. Illustrated dictionary of science. 1995. Edited by M. Allaby. Facts on File, New York. 256 pp., illus. U.S. $29.95. 386 *The year is a circle: a celebration of Henry David Thoreau. 1995. By V. C. Friesen. Natural Heritage/Natural History, Toronto. xvii + 123 pp., illus. $24.95. Books for Young Naturalists African Elephants. 1995. By R. Smith. Lerner, Minneapolis. 48 pp., illus. U.S. $14.21. Alligators. 1995. By F. Staub. Lerner, Minneapolis, 48 pp., illus. U.S. $ 14.21. Animal behavior science projects. 1995. By N. W. Cain. Witey, New York. x + 162 pp., illus. U.S. $12.95. Bats, bugs, and biodiversity: adventures in the Amazonian rain forest. 1995. By S. E. Goodman. Atheneum, New York. 46 pp., illus. Birds. 1995. By B. Taylor. Dorling, Kindersley, New York. 160 pp., illus. U.S. $5.95. A bold carnivore: an alphabet of predators. 1995. By C. Powell. Roberts Rinehart, Boulder, Colorado. 28 pp., illus. U.S. $14.95. Of bugs and beasts: fact, folklore, and activities. 1995. By L. J. Livo, G. McGlathery, and N. J. Livo. xxi + 215 pp., illus. U.S. $23.50. Dinosaurs. 1995. By N. Clark and W. Lindsay. Dorling Kindersley, New York, 160 pp., illus. U.S. $5.95. Duck. 1995. By S. Savage. Thomson Learning, New York. 32 pp., illus. U.S. $13.95. Exploring the world of animals. 1995. By P. R. Durant. Watts, New York. 48 pp., illus. U.S. $19.30. Exploring the world of plants. 1995. By P. R. Durant. Watts, New York. 48 pp., illus. U.S. $19.30. How do we know animals can think? 1995. B. S. Parker. Raintree, Austin. Texas. 40 pp., illus. U.S. $15.95. How to be a nature detective. 1995. By M. E. Selsam. Harper Collins, New York. 32 pp., illus. Cloth U.S. $14.95; paper U.S. $4.95. Insects. 1995. By L. Mourd and S. Brooks. Dorling Kindersley, New York. 160 pp., illus. U.S. $5.95. Inside the amazing Amazon. 1995. By D. Lessem. Crown, New York. 38 pp., illus. U.S. $18. THE CANADIAN FIELD-NATURALIST Vol. 110 An introduction to northern California birds. 1995. By H. Clarke. Mountain Press, Missoula, Montana. ix + 189 pp., illus. U.S. $14. The manatee. 1995. By A., V., and R. Silverstein. Millbrook Press, Brookfield, Connecticut. 64 pp., illus. U.S. $15.90. ; My shell. 1995. By L. Patchett. Garth Stevens, Milwaukee. 32 pp., illus. U.S. $12.95. A nest full of eggs. 1995. By P. B. Jenkins. Harper Collins, New York. 32 pp., illus. Cloth U.S. $14.95; paper U.S. $4.95. Outside and inside snakes. 1995. By. S. Markle. MacMillan Books for Young Readers, New York. 40 pp., illus. U.S. $16. Owls: Whoo are they? 1996. By K. Jarvis and D. W. Hold. Mountain Press, Missoula, Montana. 64 pp., illus. Paper U.S. $10; cloth U.S. $16. Primates: lemurs, monkeys, and you. 1995. By I. Tattersall. Millbrook Press, Brookfield, Connecticut. 72 pp., illus. U.S. $16.90. Puffins climb, penguins rhyme. 1995. By B. McMillan. Gulliver Books, San Diego. 32 pp., illus. U.S. $14. Sea turtles. 1995. By. G. Gibbons. Holiday House, New York. 30 pp., illus. U.S. $15.95. The tree in the ancient forest. 1995. By C. Reed-Jones. Dawn, Nevada City, California. 30 pp., illus. U.S. $6.95. The whale: giant of the ocean. 1995. By V. Tracqui. Translated by T. Froggatt. Charlesbridge, Waterdown, Massachusetts. 27 pp., illus. U.S. $6.95. Whales. 1995. By E. D. Stoops, J. L. Martin, and D. L. Stone. Sterling, New York. 80 pp., illus. U.S. $14.95. What’s alive. 1995. By K. W. Zoehfeld. Harper Collins, New York. 32 pp., illus. U.S. $ 14.95. Who eats what: food chains and food webs. 1995. By P. Lauber. Harper Collins, New York. 32 pp., illus. Cloth U.S. $15; paper U.S. $4.95. Zipping, zapping, zooming bats. 1995. By A. Earle. Harper Collins, New York. 32 pp., illus. Cloth U.S. $14.95; paper U.S. $4.95. *assigned for review tavailable for review TABLE OF CONTENTS (concluded) Wood Turtles, Clemmys insculpta, in the fresh-tidal Hudson River ERIK KIVIAT and JAMES G. BARBOUR Denning behavior of non-gravid Wolves, Canis lupus L. DAviD MEcH, MICHAEL K. PHILLIPS, DOUGLAS W. SMITH, and TERRY J. KREEGER Evidence of successful Chinook Salmon, Oncorhynchus tshawytscha, spawning in the St. Lawrence River, near Cornwall, Ontario SANDRA R. RIBEY and FRANCOIS CHAPLEAU Apparent longevity records for Red Foxes, Vulpes vulpes, in Labrador Tony CHUBBS and FRANK R. PHILLIPS Observation of repeated use of a Wolverine, Gulo gulo, den on the tundra of the Northwest Territories JOHN LEE and ALLEN NIPTANATIAK Home range size of Bushy-tailed Woodrats, Neotoma cinerea, in a southwestern Alberta MICHAEL G. TOPPING and JOHN S. MILLAR News and Comment Errata: The Canadian Field-Naturalist 109(3) — Rana-Saura: Amphibian Follow-up Project: Atlas of Amphibians and Reptiles of Quebec — Sea Wind: Bulletin of Ocean Voice International — FroGioc: IUCN/SSC Declining Amphibian Populations Task Force — Bullfrog Management in Ontario: Workshop Proceedings — Frogwatch 96 — The Second Annual International Symposium on Biology and Conservation of Owls of the Northern Hemisphere — Second International Conference on Raptors — Recovery of Nationally Endangered Wildlife in Canada (RENEW) — RENEW: Fifth Annual Report 1994/1995 — Canadian Wildlife Service LRTAP Biomonitoring Program — Newsletter of the Biological Survey of Canada (Terrestrial Arthropods) — The Boreal Dipnet — The Ontario Chorus — Missouri Botanical Garden 1996 Update — Global Biodiversity — Canadian Species at Risk, April 1996 — Amphipacifica: Journal of Systematic Zoology — Editor’s Report for The Canadian Field-Naturalist Volume 109 (1995) Minutes of the 117th Annual Business Meeting of The Ottawa Field-Naturalists’ Club 9 January 1995 — Committee Reports for 1995 — Financial Statements Book Reviews Zoology: Manual of Ornithology: Avian Structure and Function — The Downy Waterfowl of North America — Last of the Curlews — Bats: A Community Perspective — No Room for Bears: A Wilderness Writer’s Experiences with a Threatened Breed — Bird Life of Woodland and Forest — A Supplement to Distribution and Taxonomy of Birds of the World — A World Checklist of Birds — Anolis Lizards of the Caribbean: Ecology, Evolution, and Plate Tectonics Botany: An Orchid Flora of Puerto Rico and the Virgin Islands Environment: Radiation Hazards to Fish, Wildlife, and Invertebrates: A Synoptic Review — Prehistoric Alaska —Functions of Nature — Biodiversity in Canada: A Science Assessment for Environment Canada — Restorations of Endangered Species: Conceptual Issues, Planning and Implementation — Everglades: The Ecosystem and Its Restoration — Reinventing Nature? Responses to Postmodern Deconstruction New Titles Mailing date of the previous issue 110(1): 30 April 1996 341 343 346 348 349 Soil 354 364 S71 Si 378 384 THE CANADIAN FIELD-NATURALIST Volume 110, Number 2 Articles Recent discoveries of southern vascular plants at their northern limits in the granite barrens area of Lennox and Addington County, Ontario VIVIAN R. BROWNELL, C. SEAN BLANEY, and PAUL M. CATLING Additions and range extensions to the vascular plant flora of the Northwest Territories, Canada WILLIAM J. Copy Ecological replacement of the Deer Mouse, Peromyscus maniculatus, by the White-footed Mouse, P. leucopus, in the Great Lakes region CHARLES A. LONG A contribution to the biology of the White-beaked Dolphin, Lagenorhynchus albirostris, in waters off Newfoundland DOonG JIN HAI, JON LIEN, DAWN NELSON, and KRISTINA CURREN Identification of Greater Scaup, Aythya marila, and Lesser Scaup, A. affinis, ducklings COLEEN H. NELSON Seed age-germination relationships in Plains Rough Fescue, Festuca altaica subspecies hallii J.T. Romo White-tailed Deer, Odocoileus virginianus, summer dispersion areas in Ontario Jim D. BROADFOOT, DENNIS R. VoIGT, and TIM J. BELLHOUSE Berry consumption by the American Robin, Turdus migratorius, and the subsequent effect on seed germination, plant vigour, and dispersal of the Lowbush Blueberry, Vaccinium angustifolium D. R. CROSSLAND and S. P. VANDER KLOET Group hunting forays of wintering Northern Harriers, Circus cyaneus: An adaption of juveniles? THOMAS BOSAKOWSKI and DWIGHT G. SMITH The Dense-leaved Pussy’s-toes, Antennaria densifolia (Asteraceae: Inuleae): An addition to the vascular flora of British Columbia JERRY G. CHMIELEWSKI Dispersal characteristics of two-year-old Beavers, Castor canadensis, in western Montana TIMOTHY R. VAN DEELEN and DANIEL H. PLETSCHER Microhabitats of two Peromyscus (Deer and Whitefooted mice) species in old fields and prairies of Wisconsin GAIL E. KANTAK Peatlands: A new habitat for the Upland Sandpiper, Bartramia longicauda, in eastern Canada SOPHIE CALME and STEPHANIE HADDAD Notes Northern Pocket Gophers, Thomomys talpoides, with white pelage from Alberta GILBERT PROULX, LORI LOUNSBURY, and HAROLD N. BRYANT First record of a Chum Salmon, Oncorhynchus keta, from the Thompson River: Adams River spawning grounds, British Columbia D. W. WELCH and J. N. TILL A Fisher, Martes pennanti, with multiple amputations GILBERT PROULX and PAMELA J. COLE Second record and possible breeding of the European Wigeon, Anas penelope, in the District of Mackenzie, Northwest Territories MICHAEL A. FOURNIER and JAMES E. HINES Red Fox, Vulpes vulpes, kills a European Beaver, Castor fiber, kit NILs B. KILE, PETTER J. NAKKEN, FRANK ROSELL, and SIGURD ESPELAND Polar Bear, Ursus maritimus, depredation of Canada Goose, Branta canadensis, nests ARTHUR E. SMITH and MICHAEL R. J. HILL ISSN 0008-3550 1996 278 28% 29¢ 298 30: 31 32( 33 38 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 110, Number 3 July-September 1996 The Ottawa Field-Naturalists’ Club | FOUNDED IN 1879 Patron His Excellency The Right Honourable Roméo LeBlanc, P.C., C.C., C.M.M., C.D., Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse infor- | mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- — ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Clarence Frankton Stewart D. MacDonald Loris S. Russell Irwin M. Brodo W. Earl Godfrey Verna Ross McGiffin Douglas B.O. Savile William J. Cody C. Stuart Houston Hue N. MacKenzie Pauline Snure Ellaine Dickson George F. Ledingham Eugene G. Munroe Mary E. Stuart R. Yorke Edwards Thomas H. Manning Robert W. Nero Sheila Thomson Anthony J. Erskine Don E. McAllister Hugh M. Raup | 1996 Council President: David W. Moore Ronald E. Bedford Jeff Harrison | Vice-Presidents: Michael Murphy Fenja Brodo Cendrine Huemer Lee Cairnie Ann MacKenzie | ; f William J. Cody Patricia Narraway | Recording Secretary: David Smythe Brancisik. Cook Frank Pope Corresponding Secretary: Eileen Evans Ellaine Dickson Tom Reeve : ay: : Colin Gaskell Jane Topping | Treasurer: Gillian Marston Fete See Ghiischaynar Christine Hanrahan Ken Young Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. For information on Club activities telephone (613) 722-3050. | The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed 1n this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211 Copy Editor: Wanda J. Cook Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1JO Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9 Associate Editors: Robert R. Anderson Robert R. Campbell W. Earl Godfrey Warren B. Ballard Brian W. Coad William O. Pruitt, Jr. Charles D. Bird Anthony J. Erskine Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor with the exception of book reviews which should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $23 per calendar year. Libraries and other institutions may subscribe at the rate of $38 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $23 includes a subscription to The Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $5.00 to cover postage. 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. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Date of this issue: July-September 1996 (November 1996). Back Numbers and Index ; Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, and Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa | Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager. Cover: A pod of Long-finned Pilot Whale, Globicephala melas, drawn by Dawn Nelson. See COSEWIC status report by Nelson and Lien, pages 511-524. The Canadian Field-Naturalist Volume 110, Number 3 ~ July-September 1996 Endemic Vascular Plants of British Columbia and Immediately Adjacent Regions GEORGE W. DOUGLAS British Columbia Conservation Data Centre, 780 Blanshard Street, Victoria, British Columbia V8V 1X4 Douglas, George W. 1996. Endemic vascular plants of British Columbia and immediately adjacent regions. Canadian Field-Naturalist 110(3): 387-391. Forty-six endemic vascular plant taxa are listed for British Columbia and immediately adjacent regions. These taxa occur within 12 endemic regions. These endemic taxa are of some importance thus resource managers should be aware of their conservation and preservation responsibilities. Key Words: Endemic vascular plants, endemic regions, British Columbia. Vascular plant floristics have recently become a topic of interest for many resource managers. These managers, often in response to public concern with conservation or preservation issues, are now attempt- ing to deal with such issues as endangered species, biodiversity, old growth forests and gap analysis when developing protected areas strategies. When examining these, and other issues dealing with natur- al landscape features, the vascular plant flora of a region often becomes one of the main focal points. Some of the aspects examined during these floristic studies include the numbers of endemic plants, rare plants, threatened and endangered plants and their current status with respect to their extant occurrences. This paper examines the endemic vascular plants of British Columbia and immediately adjacent regions. Forty-six plants, which have all or a large part of their range in British Columbia, are considered here. These plants were chosen for both geographical and political reasons. In the majority of cases the rare plants of a political area, such as British Columbia, are peripheral species which have more extensive populations outside the area considered here. Although all possible efforts should be expended to preserve these populations (which may contain unique genetic material and are part of a region’s distinct natural biodiversity) it may not always be catastrophic if these efforts sometimes do not suc- ceed. Nearby populations in other jurisdictions may not be much different due to the restricted range of the plant(s) in British Columbia. Of greater concern are those plants which have all or a large part of their range within British Columbia. These regional endemics are especially important if their present populations are not secure from a conservation ~ standpoint. Of the 46 regional endemics treated in this paper, 34 are currently being tracked by the British Columbia Conservation Data Centre, an agency which has the responsibility of tracking over 600 provincially and globally rare native vascular plants existing within the province. It is therefore the responsibility of provincial resource managers to ensure that these 34 endemics, and many of the oth- ers which may become vulnerable in the future, retain their viability through adequate habitat protec- tion. It is also the responsibility of the latter man- agers to convince managers in other political jurisdictions that they too should take part in the conservation strategy for each of these endemics. The 46 regional endemics treated in this paper occur within 12 endemic regions. Only one of these regions, that associated with the Queen Charlotte Islands, has been well documented with respect to endemics (Calder and Taylor 1968). Some of the taxa in other groups have been mapped in regional floras (e.g., Hulten 1968; Packer 1983) or other floristic treatments (Gillett 1963; Mulligan 1974; Ceska and Ceska 1988; Douglas 1982, 1995; Douglas et al. 1996; Straley et al. 1985) while most are known only from range descriptions in regional floras (e.g., Hitchcock et al. 1955-1969; Douglas et al. 1989-1991, 1994) or research papers (Douglas and Packer 1988; Henderson et al. 1990; Packer 1972, 1983; Soreng 1991). These endemic regions and their vascular plant components are as follows: 1. Queen Charlotte Islands-Northern Vancouver Island-Southeastern Alaska Region This region includes endemic taxa associated with glacial refugia located on the Queen Charlotte 387 388 THE CANADIAN FIELD-NATURALIST Vol. 110 FiGuRE |. Endemic elements of British Columbia and immmediately adjacent regions. | - Queen Charlotte Islands- Northern Vancouver Island-Southeastern Alaska, 2 - Southern Vancouver Island-Lower Fraser Valley, 3 - Coast- Cascade Mountains-Selkirk Mountains, 4 - Rocky Mountains (Southern Canada-Northern U.S.). Islands (Calder and Taylor 1968). These plants often have ranges which also extend to adjacent islands or the mainland (Figure 1). For instance, seven of the 10 plants in this region range south to northern Vancouver Island. These include Queen Charlotte Avens (Geum schofieldii Calder & Taylor*), Queen Charlotte Isopyrum U/sopyrum savilei Calder & Taylor*), Calder’s Lovage (Ligusticum calderi Mathias & Const.*), Alp Lily (Lloydia serotina [L.] Richenb. ssp. flava Calder & Taylor*), Taylor’s Saxifrage (Saxifraga taylori Calder & Savile*), Queen Charlotte Butterweed (Senecio moresbiensis [Calder & Taylor] G.W. Dougl. & G. Ruyle- Dougl.*) and Queen Charlotte Twinflower Violet (Viola biflora L. ssp. carlottae Calder & Taylor*). *Taxa marked with an asterisk are those currently being tracked by the British Columbia Conservation Data Centre. Ligusticum calderi, Senecio moresbiensis, and Glabrous Dwarf Willow (Salix reticulata L. ssp. glabellicarpa Argus*), also range north to southeast- ern Alaska. Cordate-leaved Saxifrage (Saxifraga nel- soniana D. Don ssp. carlottae [Calder & Savile] Hult.*) also occurs in the Coast Mountains on the | adjacent mainland. Newcombe’s Butterweed (Senecio newcombei Greene*) is the only species known to be restricted entirely to the Queen Charlotte Islands at this time. 2. Southern Vancouver Island-Lower Fraser River Valley Region This region includes the remaining two (of nine) taxa which have their ranges entirely within British Columbia (Figure 1). Both species, Vancouver Island Beggarsticks (Bidens amplissima Greene*) and Macoun’s Meadow-foam (Limnanthes macounit Trel.*), occur in recently glaciated areas thus indi- cating that at certain times during glaciation parts of the area must have been ice-free. Bidens amplissima — 1996 DOUGLAS: ENDEMIC VASCULAR PLANTS OF BRITISH COLUMBIA 389 FIGURE 2. Endemic elements of British Columbia and immmediately adjacent regions. 5 - Vancouver Island Ranges-North Cascades, 6 - North Pacific Coast, 7 - Northern Rocky Mountains, 8 - Cascade Mountains-Rocky Mountains. occurs on the southeast coast of Vancouver Island and in the lower Fraser River Valley. Limnanthes macounii ranges along the southeastern coast of Vancouver Island as well as on some of the adjacent islands. 3. Coast Mountains-North Cascade Mountains- Selkirk Mountains Region This region ranges from the Coast Mountains of British Columbia (S of 57°N) to the North Cascade Mountains of southern British Columbia and north- ern Washington, and has a small outlier in the Selkirk Mountains of southeastern British Columbia (Figure 1). Lance-fruited Draba (Draba lonchocarpa Rydb. var. thompsonii [C.L. Hitchc.] Rollins*) is the only taxon within this region. 4. Rocky Mountain (Canada-Northern U.S.) Region Plants in this region range south (from about 54°N) in the Rocky Mountains of Alberta and British Columbia to northern Idaho and/or northern Montana (Figure 1). Twelve taxa occur in this element. These species include Lake Louise Arnica (Arnica louiseana Farr*), Pink Agoseris (Agoseris lackschewitzii Henderson & Mosely*), Bourgeau’s Milk-vetch (Astragalus bourgovii A. Gray*), Hooker’s Thistle (Cirsium hookeriana Nutt.), Woolly Daisy (Erigeron lanatus Hook.*), Three- lobed Daisy (E. trifidus Hook.*), Sandberg’s Desert-parsley (Lomatium sandbergii (Coult. & Rose] Coult. & Rose*), Dwarf Poppy (Papaver alpinum L.*), Oval Penstemon (Penstemon ellipti- cus Coult. & Fisch.), Lyall’s Phacelia (Phacelia lyallii [A. Gray] Rydb.*), Banff Bluegrass (Poa laxa Haenke ssp. banffianna Soreng*) and High Alpine Butterweed (Senecio conterminus Greenm.*). 5. Vancouver Island Ranges-North Cascade Mountains Region This region contains a single alpine species, Salish Daisy (Erigeron salishii G.W. Dougl. & Packer*). It occurs both in the central Vancouver Island Ranges 390 THE CANADIAN FIELD-NATURALIST Vol. 110 —---- 2 FIGURE 3. Endemic elements of British Columbia and immmediately adjacent regions. 9 - Vancouver Island Ranges- Olympic Mountains, 10 - North Cascade Mountains, 11 - Shuswap Highlands-Okanagan Valley-Columbia River Basin, 12 - Northern Mountains and Plateaus. and the North Cascade Mountains of Washington (Figure 2). 6. North Pacific Coast Region This region extends along the coast from Vancouver Island north to southeastern Alaska (Figure 2). The seven taxa in this region are likely associated with coastal glacial refugia since most of their range was glaciated. Taxa in this group are Fern-leaved Goldthread (Coptis asplenifolia Salisb.), Lance-fruited Draba (Draba lonchocarpa var. vestita O.E. Schulz*), Swamp Gentian (Gentiana dou- glasiana Bong.), Bird’s-beak Lousewort (Pedicularis ornithorhyncha Benth.), Alaska Holly Fern (Polystichum setigerum [K. Presl] K. Presl.*), Cooley’s Buttercup (Ranunculus cooleyae Vasey & Rose), Menzies’ Burnet (Sanguisorba menziesii Rydb.*). 7. Northern Rocky Mountain Region The taxa in this region range north (from about 52°N) in the Rocky Mountains of southern Alberta and British Columbia to southeastern Yukon and southwestern Northwest Territories (Figure 2). The three taxa in this group are Boreal Paintbrush (Castilleja fulva Pennell*), Jordal’s Locoweed (Oxytropis jordalii Porsild ssp. davisii [Welsh] Elisens & Packer*) and Raup’s Willow (Salix raupii Argus*). Castilleja fulva also has a single disjunct station in the southwestern Yukon. 8. Cascade Mountains-Rocky Mountains Regions This region ranges from the Cascade Mountains of southern British Columbia and northern Washington to the Rocky Mountains of southern British Columbia, southern Alberta and northern Montana (Figure 2). Five taxa are included in this range: Timber Milk-vetch (Astragalus miser Dougl. vars. miser and serotinus |Gray ex Cooper] Barneby), Deer Paintbrush (Castilleja cervina Greenm.), Golden Fleabane (Erigeron aureus Greene) and Columbia River Locoweed (Oxytropis columbiana St. John.*). ——EE 1996 9. Vancouver Island Ranges-Olympic Mountains Region This endemic region occurs in the southern Vancouver Island Ranges and the Olympic Mountains of Washington (Figure 3). A single species, Olympic Mountain Aster (Aster paucicapi- tatus [B.L. Robins.] B.L. Robins.*), belongs to this element. 10. Southern Coast Mountains-North Cascade Mountains Region This endemic region contains species which occur in the southern Coast Mountains and the Cascade Mountains of southwestern British Columbia and northern Washington (Figure 3). This group includes two taxa, Elmer’s Paintbrush (Castilleja elmeri Fern.) and Elmer’s Butterweed (Senecio elmeri Piper). 11. Shuswap Highland-Okanagan Valley-Columbia River Basin Region This region ranges from the Shuswap Highlands of southcentral British Columbia through the Okanagan Valley to the northern Columbia River Basin of northern Washington (Figure 3). The single species in this region is Okanogan fameflower - (Talinum sediforme Poelln.*). | 12. Northern Mountains/Plateaus Region A single species, Mount Sheldon Butterweed (Senecio sheldonensis Porsild*) occurs in this region. It ranges through most of northern British Columbia, southern Yukon and southwestern | Northwest Territories (Figure 3). Acknowledgments I thank Denise Manning for technical assistance, Gail Harcombe for preparing the maps and Adolph Ceska for reviewing the manuscript. Literature Cited _ Calder, J. A. and R. L. Taylor. 1968. Flora of the Queen Charlotte Islands, Part 1. Canada Department of Agriculture, Research Branch. Monograph 4. 659 pages. Ceska, A., and O. Ceska. 1988. Status report on the Macoun’s meadowfoam Limnanthes macounii. COSEWIC Report, Ottawa. [unpublished]. DOUGLAS: ENDEMIC VASCULAR PLANTS OF BRITISH COLUMBIA SMI Douglas, G. W. 1982. The sunflower family (Asteraceae) of British Coiumbia. Volume 1. Senecioneae. British Columbia Provincial Museum Occasional Papers Series Number 23. 180 pages. Douglas, G. W. 1995. The sunflower family (Asteraceae) of British Columbia. Volume 2. Astereae, Anthemideae, Eupatorieae and Inuleae. Royal British Columbia Museum, Victoria. 393 pages. Douglas, G. W., and J. G. Packer. 1988. Erigeron salishii. G.W. Douglas and J.G. Packer, a new Erigeron (Asteraceae) from British Columbia and Washington. Canadian Journal of Botany 66: 414-416. Douglas, G. W., G. B. Straley, and D. Meidinger. 1989-1991, 1994. The vascular plants of British Columbia. Parts 1—4. British Columbia Ministry of Forests, Victoria. Douglas, G. W., G. B. Straley, and D. Meidinger. 1996. Rare vascular plants of British Columbia. British Columbia Ministry of Environment, Lands and Parks. Victoria. [in press]. Gillett, J. M. 1963. The gentians of Canada, Alaska and Greenland. Research Branch, Canada Department of Agriculture Publication 1180. 99 pages. Henderson, D. M., R. K. Moseley, and A. F. Cholewa. 1990. A new Agoseris (Asteraceae) from Idaho and © Montana. Systematic Botany 15: 462465. Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W. Thompson. 1955-1969. Vascular plants of the Pacific Northwest. Parts 1-5. University Washington Press, Seattle. Hulten, E. 1968. Flora of Alaska and neighbouring territo- ries. Stanford University Press, Stanford. 1008 pages. Mulligan, G. A. 1974. Cytotaxonomic studies of Draba nivalis and its close allies in Canada and Alaska. Canadian Journal of Botany 52: 1793-1801. Packer, J. G. 1972. A taxonomic and phytogeographical review of some arctic and alpine Senecio species. Canadian Journal of Botany 50: 507-518. Packer, J. G. 1983. Flora of Alberta: Sparganium angusti- folium and Erigeron trifidus. Canadian Journal of Botany 61: 359-366. Soreng, R. J. 1991. Notes on new infraspecific taxa and hybrids of North American Poa (Poaceae). Phytologia 71: 390-413. Straley, G. B., R. L. Taylor, and G. W. Douglas. 1985. The rare vascular plants of British Columbia. Syllogeus 28. National Museums of Canada, Ottawa, Ontario. 165 pages. Received 10 December 1993 Accepted 12 March 1996 Catatropis lagunae n. sp., Trematoda, Notocotylidae, parasite d’ oiseaux de mer CHRISTIANE BAYSSADE-DUFOUR!, JEAN-LOUIS ALBARET!, HELENE FERMET-QUINET!, ET KHEMAiS FARHAT? "Muséum National d’Histoire Naturelle, Laboratoire de Biologie parasitaire, Protistologie, Helminthologie, 61, rue Buffon F 75231 Paris cedex 05 *Institut Santé et Développement, Les Cordeliers, Laboratoire de Parasitologie expérimentale, 15-21, rue de I’Ecole de Médecine, F 75270 Paris cedex 06 Bayssade-Dufour, Christiane, Jean-Louis Albaret, Hélene Fermet-Quinet, et Khémais Farhati. 1996. Catatropis lagunae n. sp., Trematoda, Notocotylidae, parasite d’ oiseaux de mer. Canadian Field-Naturalist 110(3): 392-402. Des cercaires émises par des Mollusques Hydrobia ulvae des cétes atlantiques de France, s’enkystent sur des algues Ulva lactuca; la consommation d’algues porteuses de métacercaires contamine en laboratoire de jeunes Oiseaux Anatidae Anser |} anser, Anas platyrhynchos et Cairina moschata. Les Mammiferes Mesocricetus auratus et Mus musculus sont réfractaires. Les vers adultes obtenus dans les coecums digestifs des Oies et Canards ont les caractéristiques des Notocotylidae Liihe, 1909 et répondent a la définition du genre Catatropis Odhner, 1905. La poche du cirre trés longue occupant la moitié antérieure du corps, la vésicule séminale externe trés volumineuse et le cirre recouvert de tubercules caractérisent une espece nouvelle appelée Catatropis lagunae dont la morphologie est complétée par des observations au microscope élec- tronique a balayage. Les vers pondent des oeufs operculés a filaments polaires qui n’ éclosent pas dans l’eau sauméatre. L’hote vertébré naturel est inconnu mais pourrait étre |’ Anatidae migrateur Branta bernicla bernicla, présent en hiver sur le site et consommateur d’algues vertes marines ou saumatres. Les cercaires mires, monostomes, apharyngées, 42 nom- breuses cellules kystogeénes, a trois taches antérieures, a glandes adhésives postérieures, 4 pores excréteurs s’ouvrant dans le tiers antérieur de la queue, correspondent au type monostomi des Notocotylidae. Leur chétotaxie céphalique et caudale est décrite. L’intérét des papilles ventrales des vers adultes est souligné, elles ont une action pathogéne sur les hétes. Mots clés : Catatropis lagunae, Trematoda, Notocotylidae, morphologie, cycle biologique, chétotaxie, MEB, Anser anser, Anas platyrhynchos, Cairina moschata, Hydrobia ulvae. L’aire étudiée est une lagune saumiatre, pres de la cdte orientale de l’ile d’Oléron, a Boyardville (Charente-Maritime); elle est séparée de 1’Océan Atlantique par un cordon littoral de sable et terre. Elle appartient a la Réserve Naturelle de l’ile. Son fond est constitué de sable vaseux. Sa végétation comprend essentiellement des algues vertes Ulva lactuca, Ulothrix sp. et une algue bleue Microcoleus chthonoplastes. Elle est fréquentée, en automne et hiver par plusieurs milliers de Bernaches cravants Branta bernicla bernicla, de Tadornes de Belon Tadorna tadorna, Anatidae, d’ Huitriers-pies Haematopus ostralegus, Haematopodidae, par des Cormorans Phalacrocorax sp. Phalacrocoriacidae, des Hérons cendrés Ardea cinerea et des Aigrettes Egretta garzetta, Ardeidae, au total par plusieurs centaines de milliers d’oiseaux migrateurs, selon Mahéo (1991) et Walmsey (1991). Des Goélands argentés Larus argentatus et des Mouettes L. sp., Laridae, y sont également abondants toute |’ année. Les Bernaches cravants stationnent fréquemment au bord de la lagune; elles peuvent également tenir leurs quartiers d’hiver sur d’autres cétes frangaises, ainsi que sur les cétes néerlandaises et anglaises tan- dis qu’elles estivent en Sibérie Occidentale, dans la Péninsule de Taymyr, selon Cramp et al. (1977). La Bernache Branta bernicla hrota venant du Canada et du Groenland n’a pas été signalée a Oléron mais sur les cOtes de Normandie. Toutefois depuis la création récente de la Réserve, un nombre croissant d’especes migratrices est attiré dans l’ile. Les vertébrés aquatiques de la lagune sont de jeunes poissons, et les invertébrés, de trés nombreux Mollusques Prosobranches ainsi que des arthropodes et des isopodes. Le but de ce travail est de connaitre les Trématodes parasites des espéces animales protégées en partant ~ des stades larvaires: les cercaires, mises par des Mollusques naturellement infestés et en réalisant le cycle biologique, en laboratoire, sur des modeéles expérimentaux. Matériel et méthodes. Des Mollusques Hydrobiidae Hydrobia ulvae | (Pennant, 1777) naturellement infestés, sont récoltés, | dans la lagune, au mois de décembre; apres exposi- _ tion a la lumiére, ils émettent des cercaires dont | plusieurs sont dessinées vivantes a la chambre claire, d’autres étant imprégnées au nitrate d’argent selon la ~ méthode de Combes et al. (1976), pour la mise en — évidence de leurs- récepteurs sensoriels ou sensilles. — La majorité d’entre elles s’enkyste, moins d’une heure apres I’ émission, sur Ulva lactuca. Quelques 592 | | 1996 BAYSSADE-DUFOUR, ALBARET, FERMET-QUINET, ET FARHATI: CATATROPIS LAGUNAE N. SP. Métacercaire Ulva lactuca 393 Miracidium | / Sporocyste YA Rédie et Cercaire Hydrobia ulvae FiGureE 1. Cycle biologique de Catatropis lagunae n. sp. métacercaires sont dessinées vivantes; les autres, fixées sur des algues, sont données en nourriture a un Canard de Barbarie Cairina moschata 77 TM, deux Oies cendrées Anser anser 76 TM et 72 TM, un Canard colvert Anas platyrhynchos 78 TM, cha- cun agé de quelques jours et élevé a l’abri de conta- minations parasitaires, ainsi qu’a un Hamster Mesocricetus auratus et quatre Souris Mus musculus. C. moschata consomme a J zéro, 80 métacercaires, puis a J + 2, 30 métacercaires; il est autopsié 11 jours apres la premiére infestation. A. anser 76 TM consomme 4a J zéro, 100 métacercaires, puis a J +13, 200 métacercaires; son sacrifice a lieu 24 jours apres la premiére infestation. A. anser 72 TM ingére 100 métacercaires et est autopsié a J + 27. A. platyrhynchos recoit, a J zéro, 150 métacer- caires puis a J +1, 100 métacercaires; il est autopsié 15 jours aprés la premiére infestation. Chez les qua- tre Anatidae, des vers adultes sont recueillis. Quelques uns, mis dans du sérum physiologique ou dans de |’eau de mer, pondent. Les autres sont fixés a l’alcool, au formol 4 10%, ou a la glutaraldéhyde, 394 certains étant aplatis entre lame et lamelle. Une par- tie des vers de 77 TM, 76 TM et 72 TM est colorée au Carmin chlorhydrique alcoolique, dessinée et mesurée a la chambre claire; une partie des vers de 77 TM, 76 TM et 78 TM est déshydratée a l’alcool, passée au point critique au CO2, métallisée a 1’Or- Palladium et observée au microscope électronique a balayage JEOL 840 A. Le Hamster et les quatre Souris ayant ingéré des métacercaires sont sacrifiés, M. auratus a J +21, M. musculus, respectivement a J +5, 4+15,+21 et +21. Résultats Un total de 82 Trématodes adultes, appartenant manifestement a la méme espéce, est recueilli dans les coecums digestifs des Anatidae: 10 chez C. moschata, 45 chez A. anser 76 TM, 7 chez A. anser 72 TM et 20 chez A. platyrhynchos. Mauratus et M. musculus w hébergent aucun parasite. Cycle biologique du Trématode. Le cycle biologique (figure 1) admet deux hdtes successifs. Le premier h6te naturel est le Mollusque marin ou saumatre Hydrobia ulvae. Cette espéce, dont plusieurs milliers de représentants ont été récoltés a Oléron, a produit des cercaires pendant plus de deux mois; celles-ci, émises a la lumiére, s’enkystent essentiellement sur les algues Ulva lactuca; les métacercaires sont infestantes quelques heures a quelques jours apres |’enkystement. Rothschild (1938) et Deblock (1980) signalent la présence de cercaires, issues de rédies, de genre indéterminé et de morphologie identique a celle de notre cercaire chez les Hydrobia des cotes anglaises et frangaises de la Manche; le cycle biologique s’y réalise donc. Il est possible qu’il se déroule aussi dans d’autres pays et continents puisque, selon Scarlato ( 1987), H. ulvae est présent des cotes du Maroc a celles de la Mer Blanche (Sibérie Occidentale, Russie). H. ulvae n’existe pas sur les cotes atlantiques américaines, mais l’espéce proche, H. totteni Morrison, 1954, est présente, du Labrador au New Jersey, selon Abott (1974). Les hétes définitifs expérimentaux sont C. moschata, A. anser et A. platyrhynchos. Les vers deviennent adultes et ovigéres en 9 jours environ. Les hétes définitifs naturels du parasite sont incon- nus. Les seuls Oiseaux consommateurs d’algues vertes marines ou saumatres sont les Bernaches cra- vants, selon Cramp et al. (1977) ; elles peuvent donc s’infester comme nos modéles, par ingestion de métacercaires enkystées sur U. lactuca. Branta ber- nicla bernicla, présente a Oléron, pourrait étre 1’ hdte naturel du parasite. Les oeufs des vers sont éliminés avec les déjec- tions des Anatidae expérimentalement infestés; lais- sés seuls, dans l’eau de mer ou saumatre, ces oeufs THE CANADIAN FIELD-NATURALIST Vol. 110 n’évoluent pas mais y demeurent pendant plusieurs semaines; laissés avec des Hydrobia vivants, ils dis- paraissent, en quelques jours, probablement consom- més par les Mollusques. Les oeufs de la plupart des familles de Trématodes éclosent dans |’eau, apres maturation; parmi les exceptions, Wright et Bennet (1964) signalent ceux de Notocotylidae qui éclosent dans le tube digestif des Mollusques qui les consom- ment et deviennent ainsi infestants pour leurs pre- miers hétes. Ces oeufs, contrairement au cas trés général, contiennent un sporocyste et non un miracidium, selon Murrils et al. (1985). Description du Trématode Les vers adultes. Les dimensions du corps et celles des principaux organes de 19 spécimens sont données dans le tableau 1. Les vers 77 TM ont entre 9 et 11 jours, ils sont les plus jeunes; ils sont déja ovigéres. Les vers 72 TM ont 27 jours, ils sont les plus agés. Les vers 76 TM sont séparables en deux lots: les plus longs, vraisemblablement agés de 24 jours et les plus courts, 4gés de 11 jours. Les vers longs de 76 TM et tous ceux de 72 TM ( agés de 24 et 27 jours) ont des tailles significativement supérieures a celles des vers courts de 76 TM et a celles de 77 TM, agés de 9 a 11 jours. Ceci indique que les parasites continuent a grandir apres leur maturité sexuelle et qu'une premiére infestation n’ empéche pas la réussite d’une seconde. Le métraterme, la poche du cirre et espace occupé par les vitellogenes sont plus longs chez les vers agés que chez les jeunes; par contre, le nombre de papilles ventrales, la taille de l’ovaire, des testicules et de la ventouse orale ne semblent pas varier ou varient trés peu avec |’ age. Le corps des adultes (figure 2, 3) est allongé, a bords paralléles, 4 concavité ventrale, il est pourvu d’une cuticule a épines larges dans la région antérieure, fines dans la région postérieure et a mélange d’épines larges et fines dans une zone de transition. Le tégument porte, ventralement, dans la région médiane de la moitié postérieure du corps, une créte longitudinale, rouge clair sur le ver vivant, bordée a gauche et a droite de 6 a 9 papilles, approx- imativement symétriques. La ventouse orale est ter- minale et de diamétre relativement faible. L’ appareil digestif comporte une bouche, un oesophage court, des coecums modérément sinueux et trés longs, pas- sant entre les glandes vitellogénes et l’utérus, puis entre les testicules et l’ovaire et se terminant au voisinage du pore excréteur. L’appareil génital male comprend, dans la région postérieure du corps, deux testicules symétriques, latéraux, profondément lobés sur leur bord externe, deux canaux déférents conflu- ant rapidement en un canal médian unique, une volu- | mineuse vésicule séminale externe repliée plusieurs fois sur elle- méme, une poche du cirre trés longue | atteignant la moitié du corps et comprenant une | 1996 BayssADE-DUFOUR, ALBARET, FERMET-QUINET, ET FARHATI: CATATROPIS LAGUNAE N. SP. KS S aS Wena AN ( Fe SN SSS = a SS ZW WN = Ss SSS = Ss ow) SSS 395 a 200U™ FicurE 2. Catatropis lagunae n. sp. , A: créte médiane et papilles ventrales de l’adulte. B: morphologie générale de V’adulte. C: détail des épines au niveau du pore excréteur. D: détail de la poche du cirre et du métraterme. E: détail du carrefour génital. F: oeuf du métraterme. THE CANADIAN FIELD-NATURALIST 396 STOX LTO 010x610 070X610 910X810 910X810 €LO0XTI0 b1T0 X80 rl OXxLto 8TOXx1c0 60'0 X 910 STOX610 910X810 €l0x8st0 cl0X8T0 cTLOX810 TLOXLIO cL 0x60 Il'0 x10 OLO0Xx€l0 ITOXST0O cTLOXPIL0 ITOX+v10 OlOxel0 aTIkAO 60°0 < 760 60°0 x 160 60°0 X ZO'T 60°0 X 980 60'0 X 76'0 O10 X 880 60°0 x SL°0 80°0 x $80 010X060 L00X TL'0 60°0 x L9°0 60°0 X 8L'0 60'0 x 09°0 60°0 x €9°0 010x790 L0'0 x 190 60°0 x c9°0 L0°0 x 6$°0 90'0 x 09°0 90'0 x $9°0 60°0 x 6S°0 80'0 x SS'0 L0';0 x $S'0 UID ENA sTT vel vil scl 601 cO'T cot Lil LOT LO'T £60 £60 £60 08°0 88°0 cL 0 08°0 99°0 OL'0 cL'0 99°0 09°0 £9°0 7] sano np ayo0g 60°0 x 79°0 010x990 L0'0 x $90 O10x€s'0 L0'0 x $90 60'0 x 790 80°0 x 09°0 IT0xL9°0 60'0 x 89°0 90'0 x PS'0 90'0 X LS'0 90°0 x 09°0 LO0x1S°0 S0°0 x OF°0 S0'0 x 870 SO0XxrE0 90'0 X 6£°0 80°0 X F7'0 80°0 X £70 90'0 STO 80'0 X L770 80°0 xX 070 80'°0 X £70 ‘d sayfevtA. 60°0 5ekm~? (Whitlaw and Lankester 1994a). In a more detailed study for Ontario, Whitlaw and Lankester (1994b) came to the conclu- sion that Moose could co-exist with infected deer, in relatively stable numbers, over at least a decade and that Moose numbers were inversely related to the intensity of meningeal worm larvae passed by deer. Whitlaw and Lankester (1994b) concluded that inter- actions between the two cervids and the nematode were more subtle than previously believed and that further studies were required to evaluate the relative importance of the meningeal worm among limiting factors of Moose populations. For example, the like- lihood of transmission to Moose could depend on the age structure of co-habiting deer populations since Slomke et al. (1995) showed that the number of meningeal worm larvae passed by deer was inversely related to deer age. Along with the renewed interest in the role of the meningeal worm on population dynamics of Moose, Schmitz and Nudds (1994) modelled the Moose- deer-meningeal worm system with differential equa- tions and developed a sensitivity analysis. Model solutions were most sensitive to the competitive effect of Moose on deer, to parasite-induced mortali- ty rate of moose, and to the rate of increase of the intermediate host, three parameters for which there is little empirical information. The objectives of our study were to census and map the cases of Moose exhibiting signs of pare- laphostrongylosis in Québec and to evaluate the role of the meningeal worm in the population dynamics of Moose living in sympatry with White-tailed Deer. We considered two hypotheses concerning the trans- mission of the meningeal worm from deer to Moose: 1) there exist refugia in which Moose are not infect- ed; as Moose density increases in refugia, animals are progressively forced to disperse out and the pro- portion of infected individuals increases with Moose density; 2) the probability of Moose infection depends only on deer density. If the first hypothesis were true, then the meningeal worm might regulate Moose populations (sensu Messier 1991); the second scenario implies that the meningeal worm is a limit- ing factor (sensu Messier 1991), affecting population rate of increase and carrying capacity of the Moose range without any stabilizing effect. Study area and methods The study focused on Moose showing signs of infection caused by the meningeal worm throughout Québec although deer occurred only in the south (Figure 1). The extreme south of Québec is covered by deciduous forests with the exception of agricul- tural lands and urban areas; highest deer densities (up to 13°km?: F. Potvin and L. Breton, unpub- THE CANADIAN FIELD-NATURALIST Vol. 110 lished) occur in this region. Boreal forest, dominated by Black Spruce (Picea mariana), Jack Pine (Pinus banksiana) and Balsam Fir (Abies balsamea), covers northern Québec, beginning around 49°—50° latitude; deer are absent from this biome and Moose density is low (0.05—0.10 animalsekm~). In between, mixed forests dominate; deciduous tree species, namely maples (Acer saccharum, A. rubrum), birches (Betula alleghaniensis, B. papyrifera) and aspen (Populus tremuloides), are progressively replaced northward by conifers. Deer occupy the southern portion of the mixed forests (Figure 1b) at low densi- ty (< 3e¢km*: F. Potvin and L. Breton, unpublished) whereas Moose reach the highest density in this tran- sition zone; i.e., 0.20—0.40 individualsekm~ (Anonymous 1993). We collected information on symptomatic Moose using a questionnaire distributed to the personnel of the Ministere de l'Environnement et de la Faune throughout Québec, with the exception of the two northernmost regions, which encompassed Anticosti Island. The questionnaire included a summary sheet describing the life cycle of the meningeal worm and listed typical signs of infection in Moose. The first question pertained to the number of cases of sympto- matic Moose observed by the respondent during the last year while the second question covered previous years without time limits. For each case, the respon- dent completed a separate form pertaining to observed signs, location (hunting zone, locality, uni- versal transverse mercator grid locus), date, sex and age class (calf and older) of the animal, and respon- dent’s name and address. The questionnaire was dis- tributed to conservation officers, wildlife technicians and biologists. Most reports were precise in time and space because they were completed by conservation officers who are accustomed to note special events. It was possible to eliminate double reports in all cases. Questionnaires were distributed in June 1992 and returned before March 1993. We did not set a time limit on reported cases, but the analysis focused mostly on the last 10 years, 1983-1992. We assumed that symptomatic Moose suffered from pare- laphostrongylosis based on Gilbert (1974), who showed a close agreement between signs and detec- tion of worms in the nervous system of moose. Apparent infection rate was expressed as the aver- age annual number of cases per 10 000 Moose in win- ter, taking each hunting zone where symptomatic Moose were observed as a sampling unit (Figure 1a). Moose population sizes were measured from winter aerial census, corrected for visibility bias (Créte et al. 1986); we used the most recent estimates of the five- year census program for monitoring Moose in Québec as Moose densities have been relatively stable during the last decade (Anonymous 1993). Deer density was estimated from the average buck harvest for the study period; i.e., 1983-1992, during which deer populations 1996 DUMONT AND CRETE: MENINGEAL WORM IN SOUTHERN QUEBEC 415 eservoir lanicouagan sD AR HUNTING ZONES Sept-lles ILE ‘ D'ANTICOSTI SS Chicoutimi Chibougamau ® Témiscaming ONTARIO B 1983-1990 A 1991-1993 — White-tailed deer distribution Ficure 1. (A) Location of current hunting zones in southern Québec; (B) Northern limit of the White-tailed Deer range in winter, and location of reported Moose showing signs of infection by the meningeal worm during the 10-year peri- od 1983-1992. 416 exhibited strong and variable fluctuations (G. Lamontagne and C. Daigle, unpublished). Registration of hunter-harvested deer is compulsory in Québec. Buck harvests have been used for manage- ment purposes in Québec as an index to population size because they have generally paralleled fluctua- tions in size of deer wintering areas which have been mapped from the air since the mid 1970s. We used simple and multiple linear regression models for evaluating the strength of the relationships relating apparent infection rate to Moose and deer density (SAS Institute Inc. 1985). The proportion of calves and the sex ratio of symptomatic Moose were compared to the corresponding variables in winter populations with the x? test; for this comparison, we pooled composition counts of all hunting zones where at least 1 infected Moose was reported. Results We received 143 questionnaires that reported 89 different Moose with signs of meningeal worm infection. We deleted five reports coming from Saint-Félicien Zoo (hunting zone 18: Figurela) where Moose and deer have been kept together in a large forested enclosure. No other cases were report- ed in this region although most conservation officers there completed the questionnaire. Since hunting zoning changed in the early 1980s and deer density varied markedly between the 1970s and the 1980s, we kept only those reports from the last 10 years (1983-1992) for further analysis. All but two reports of symptomatic Moose originated from the White-tailed Deer range (Figure 1b). Cases were concentrated in the region to the north and northwest of Montréal, and around Rimouski in east- ern Québec. Most reports in the Rimouski area pre- ceded 1990, before the deer population crashed due to a series of harsh winters aggravated by coyote (Canis latrans) predation (Créte and Lemieux 1994). Moose of both sexes appeared equally susceptible to the meningeal worm since the sex ratio among symptomatic animals (403602; n = 72) did not dif- fer statistically (x? = 2.39, 1 d.f.; P > 0.10) from that observed among living Moose in winter (326 :68 2; n = 1095). In contrast, calves constituted only 3% of the reported cases (n = 79), which is significantly lower than their percentage in the population (28% (n = 1509); x? = 23.1, df. = 1; P< 0.005). The majority of cases (47/82) were concentrated in late winter and spring (March-June) whereas incidence was lowest between August and December (14 cases). Only two symptomatic calves were observed, both in January. The most frequent signs noted were lack of fear of humans, nonchalance, circling movements, and equi- librium loss. In most reports, many signs were observed in an individual animal simultaneously. The number of reports per 10 000 Moose varied by almost two orders of magnitude between hunting THE CANADIAN FIELD-NATURALIST / | Vol. 11058 | zone 13, where the infection rate was low, and zone 8 whet it averaged 37.5 cases — 10 000 moose! « | year! (Table 1). Apparent infection rate exhibited a i positive linear relationship with the index of deer density ( r? = 0.80; 8 d.f.; P = 0.0004); the addition | of Moose density as a second independent variable in a multiple linear regression did not improve the: model significantly (P = 0.34). We found no signifi-- cant relationship between apparent infection rate andy Moose density (r? = 0.25; P = 0.14). : Discussion One might question the validity of evaluating the role of the meningeal worm on Moose demography using a questionnaire because such an approach pro- vides semi-quantitive results that can be biased. In particular, animals showing little fear of humans could have been incorrectly recorded as infected moose. This bias, if it existed, must be very small, given the close agreement between reported cases. and deer distribution. In this regard, reports from the area of Saint-Félicien Zoo are meaningful: all the cases observed in the zoo where deer and Moose coexist, were declared, whereas the same observers reported no case elsewhere in the region, where only Moose are present. The two cases observed outside of the deer range west of Amos (Figure 1b) could be explained by the presence of deer across the border in Ontario (Whitlaw and Lankester 1994b). Moreover, the decrease in the number of sympto- matic Moose reported in southeastern Québec after the deer population crashed also suggests that our monitoring tool of infection was relatively sensitive. The apparent infection rate could also be biased by differences in road access among hunting zones, in observer:moose ratios, in hunting pressure and in wolf density; infected Moose are most likely vulner- able to hunting or predation. Road access in forested areas of southern Québec has not been measured, but it is relatively good everywhere in the deer range, most of which is private property (Anonymous 1993). North of the deer range, accessibility is gener- ally lower. Hunting pressure for Moose is high everywhere in southern Québec, but the bow is the only weapon allowed in zones 6, 7, 8, 9 and 10 (east- ern half)(Anonymous 1993). Wolves (Canis lupus) are mostly restricted to the north of the deer range? consequently there could be an underestimation of— apparent infection rate in zones 11, 13 and 15 due iol | predation and road access. The relationship that we obtained between relativel deer density and apparent infection rate of Moose by the meningeal worm suggests that deer abundance constitutes the major risk factor of Moose infection. Deer, most of which harbour meningeal worms in Québec ( Bindernagel and Anderson 1972; Beaulieu- Goudreault 1981; Claveau and Filion 1984), spreac larvae in the environment. As deer density increases 1996 DUMONT AND CRETE: MENINGEAL WORM IN SOUTHERN QUEBEC 417 TABLE 1. Average annual (1983-1992) apparent rate of Moose infection by the meningeal worm as estimated through a questionnaire, mean Moose density, average deer buck harvest (1983-1992) and actual deer density, for all hunting zones where at least one symptomatic Moose was observed. Hunting zone Infection rate Moose density Deer harvest Deer density* (cases*10 000 moose!) (animalsekm7”) (bucke10 km”) (deerekm~) 1 1.4 (5*) 0.18 0.35 0.02 2 3.6 (9) 0.22 - 0.47 0.27 3-4-6 4.9 (3) 0.04 1.74 N.A.4 7 1.5 (1) 0.17 0.90° N.A. 8 37.5 (3) 0.02 5.89 SIDE 9 13.4 (13) 0.22 0.89 P07, 10 TAC) 0.16 2.54 2.88 11 Dri) 0.11 73 1.89 13 0.4 (2) 0.23 0 N.A. 15 0.6 (3) 0.10 0 N.A. “number of reported cases > estimated through the size of deer wintering areas; only bow hunting allowed © estimated from aerial surveys between 1991 and 1995, according to Potvin et al. (1992) 4 not available © only the portion of the zone located to the south of Fleuve Saint-Laurent larvae and risk of infection for Moose also increase. Although our sample size is small, our results sug- gest that the risk of infection was independent of Moose density; our results provide no evidence of refugia for Moose living in sympatry with deer at the scale of Québec hunting zones; i.e., many thousands of square kilometres. Infection rates of Moose appeared very low. In hunting zone 8 where the rate was the highest, fewer than 1% of Moose were found with signs of pare- laphostrongylosis each year. Obviously, many cases passed undetected, and this value may be higher, by an unknown amount. Monitoring of radio-tagged Moose also suggests that paralephostrongylosis is a marginal limiting factor. We monitored, for other purposes, 149 Moose for a total of 73 968 days in two study areas of southern Québec (intersection of zone 10, 11 and 12: Courtois and Créte 1988; Rimouski area: Courtois et al. 1994) where deer den- sity was low. We detected only one case of mortality attributable to the meningeal worm: i.e., an annual mortality rate of 0.5% + 0.05 (S.E.) (Heisey and Fuller 1985). However, this estimate is also underes- timated because Wolves were present in one study _area and hunting was allowed in both. j As the infection rate by the meningeal worm was not positively density-dependent, it is unlikely that this parasite regulates Moose populations in southern : | Québec. When suitable habitat exists, Moose density will mostly depend on forage, predation and hunting _(Créte et al. 1981: Messier and Créte 1985; Créte 1989). But the meningeal worm constitutes a limiting _ factor that diminishes the rate of increase of Moose _ populations in proportion to deer density. At very high _ deer density, Moose may be incapable of colonizing _ deer range. We cannot determine this threshold densi- ty because we could not compute an absolute mortali- ty rate caused by the parasite, only an index to it. However, it seems that deer-induced infections cannot exclude Moose from some areas of Québec. Indeed, Moose have persisted on Anticosti Island even with deer densities as high as 15 individualsekm7 (Potvin et al. 1991), which exceeds densities observed else- where in southern Québec (G. Lamontagne and C. Daigle, unpublished). Symptomatic Moose have been observed on Anticosti Island (A. Gingras, personal communication). Besides the meningeal worm, other factors could determine the southern limit of the Moose range: quantity and quality of forage in sum- mer and winter (Saether 1985; Créte 1989), competi- tion with White-tailed Deer (Nudds 1990), or thermal stress (Renecker and Hudson 1986). Acknowledgments The Ministére de l'Environnement et de la Faune du Québec supported this study. We are grateful to the conservation officers, wildlife technicians, biolo- gists, park wardens and veterinarians who completed our questionnaire. G. Lamontagne and C. Daigle pro- vided unpublished information whilst J. Bouchard drew the figure. M.W. Lankester and F. Messier kindly reviewed an earlier version of this manuscript. Literature Cited Anonymous. 1993. Plan de gestion de l’orignal 1994- 1998. Objectifs de gestion et scénarios d’exploitation. Québec, Ministére du Loisir, de la Chasse et de la Péche, report ISBN 2-551-15697-1. Anderson, R. C. 1964. Neurologic disease in Moose infected experimentally with Pneumostrongylus tenuis from White-tailed Deer. Pathologia Veterinaria 1: 289-322. Anderson, R. C. 1965. An examination of wild Moose exhibiting neurologic signs in Ontario. Canadian Journal of Zoology 43: 635-639. 418 Anderson, R. C. 1972. The ecological relationships of meningeal worm (Pneumostrongylus tenuis Dougherty) and native cervids in North America. Journal of Wildlife Diseases 7: 304—309. Beaulieu-Goudreault, M. 1981. Etude des endoparasites et plus spécialement du ver des méninges (Parelaphostrongylus tenuis) du cerf de Virginie (Odocoileus virginianus) a Vile d’ Anticosti. M. Sc. the- sis, McGill University, Montréal. Bindernagel, J.A., and R.C. Anderson. 1972. Distribution of the meningeal worm in White-tailed Deer in Canada. Journal of Wildlife Diseases 36: 963-966. Claveau, R., and J.-P. Filion. 1984. Fréquence et distrib- ution du ver des méninges chez le cerf de Virginie de Vest du Québec. Naturaliste Canadien 111: 203-206. Courtois, R., and M. Créte. 1988. Déplacements quoti- diens et domaines vitaux des orignales du sud-ouest québécois. Alces 24: 78-89. Courtois, R., D. Sigouin, J.-P. Ouellet, A. Beaumont, and M. Créte. 1994. Mortalité naturelle et d’origine anthropique de l’orignal au Québec. Québec, Ministére de l’Environnement et de la Faune, Report Number 94- 2466-10. Créte, M. 1989. Approximation of K carrying capacity for Moose in eastern Québec. Canadian Journal of Zoology 67: 373-380. Créte, M., and R. Lemieux. 1994. Dynamique de popula- tion des coyotes colonisant la péninsule gaspésienne. Québec, Ministére de 1’7Environnement et de la Faune, Report No. 94-2429-07. Créte, M., R. J. Taylor, and P. A. Jordan. 1981. Optimization of Moose harvest in southwestern Québec. Journal of Wildlife Management 45: 598-611. Créte, M., L.-P. Rivest, H. Jolicoeur, J.-M. Brassard, and F. Messier. 1986. Predicting and correcting heli- copter counts of Moose with observations made from fixed-wing aircraft in southern Québec. Journal of Applied Ecology 23: 751-761. Gilbert, F. F. 1974. Parelaphostrongylus tenuis in Maine. II. Prevalence in moose. Journal of Wildlife Management 38: 614-616. Gilbert, F. F. 1992. Retroductive logic and the effects of meningeal worms: a comment. Journal of Wildlife Management 56: 614-616. Heisey, D. M., and T. K. Fuller. 1985. Evaluation of sur- vival and cause-specific mortality rates using telemetry data. Journal of Wildlife Management 49: 668-674. Karns, P. D. 1967. Pneumostrongylus tenuis in deer in Minnesota and implications for moose. Journal of Wildlife Management 37: 299-303. Kerney, S. R., and F. F. Gilbert. 1976. Habitat use by White-tailed Deer and Moose on sympatric range. Journal of Wildlife Management 40: 645-657. Kelsall, J. P., and W. H. Prescott. 1971. Moose and deer behaviour in snow. Canadian Wildlife Service, Report Series Number 15. THE CANADIAN FIELD-NATURALIST Vol. 110 Messier, F. 1991. The significance of limiting and regu- lating factors in the demography of Moose and White- tailed Deer. Journal of Animal Ecology 60: 377-393. Messier, F., and M. Créte. 1985. Moose-wolf dynamics and the natural regulation of Moose populations. Oecologia (Berlin) 65: 503-512. Nudds, T. D. 1990. Reproductive logic in retrospect: the ecological effects of meningeal worms. Journal of Wildlife Management 54: 396-402. Nudds, T. D. 1992. Reproductive logic and the effects of meningeal worms: a reply. Journal of Wildlife Management 56: 617-619. Potvin, F., L. Breton, and A. Gingras. 1991. La popula- tion de cerfs d’ Anticosti en 1988-1989. Québec, Ministére du Loisir, de la Chasse et de la Péche, report Number SP 1779. Potvin, F., L. Breton, L.-P. Rivest, and A. Gingras. 1992. Application of a double-count aerial survey tech- nique for White-tailed Deer, Odocoileus virginanus, on Anticosti Island, Québec. Canadian Field-Naturalist 106: 435-442. Prescott, W. H. 1974. Interrelationships of the Moose and deer of the genus Odocoileus. Le Naturaliste canadien 101: 493-504. Renecker, L. A., and R. Hudson. 1986. Seasonal energy expenditures and the thermoregulatory responses of moose. Canadian Journal of Zoology 64: 322-327. SAS Institute Inc. 1985. SAS user’s guide. 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The co- occurence of moose, White-tailed Deer and Parelaphostrongylus tenuis in Ontario. Canadian Journal of Zoology 72: 819-825. Received 11 September 1995 Accepted 31 January 1996 _ Dietary Flexibility of Shorebirds in the Western Hemisphere _ SUSAN K. SKAGEN and HEATHER D. OMAN Midcontinent Ecological Science Center, National Biological Service, 4512 McMurry Avenue, Fort Collins, Colorado 80525-3400 Skagen, Susan K., and Heather D. Oman. 1996. Dietary flexibility of shorebirds in the western hemisphere. Canadian Field-Naturalist 110(3): 419-444. _ Regional patterns of shorebird diets were examined by reviewing 75 papers reporting prey of 43 shorebird species through- - out the western hemisphere. Collectively, shorebirds consumed a wide variety of invertebrate taxa, including 12 phyla, 22 classes, 72 orders, 238 families, and 404 genera of invertebrates. The most widely represented invertebrate classes were Insecta, Malacostraca, Gastropoda, Polychaeta, and Bivalvia. The ten most widely studied shorebird species exhibited con- _ siderable dietary breadth, consuming an average of 36 (range 23-65) families of invertebrates. Fifteen invertebrate families were common to the diets of seven or more of these ten species. For all shorebird species evaluated, there was little dietary overlap in invertebrate taxa between geographic regions, especially between tidal and inland areas. Diet similarities of _ species and guilds of shorebirds within regions and of coexisting species within studies were high. The flexible nature of _ food choice in shorebirds influences management approaches toward providing vital food resources for shorebirds during all seasons. Management efforts should focus on maintaining hydrologic regimes and ecosystem processes that promote the growth and maintenance of invertebrate populations in general; specific taxa need not be targeted. Successful maintenance _ of wetlands will ensure that naturally-occurring populations of invertebrates occur and are accessible to shorebirds. Key Words: Shorebirds, Charadriiformes, prey, invertebrates, opportunistic foraging, dietary overlap. Shorebirds (Charadriiformes; 12 families) are an ecologically diverse group of migratory birds found throughout many regions of the world. Collectively, _ shorebirds are morphologically diverse, with body _ lengths ranging from 130 mm to 650 mm and bill lengths ranging from 13 mm to 219 mm (Hayman et al. 1986). Shorebirds occupy a broad array of habi- tats, including tidal flats, sandy beaches, grass - uplands and wet meadows, agricultural fields, and freshwater wetlands (Hayman et al. 1986; Colwell _ and Oring 1988). They capture a broad spectrum of - invertebrates, primarily by gleaning them from the surface of mud, water, and emergent vegetation, or by probing in moist substrates. Research on shorebird foraging has covered a broad array of topics, including feeding assemblages, niche dimensions, competition, partitioning of habi- _ tats, prey selection, and digestive performance (Recher 1966; Baker 1977; Baker and Baker 1973; Connors et al. 1979; Myers et al. 1979a,b; Quammen 1982; Baldassare and Fischer 1984; Gratto et al. 1984; Myers 1984; Colwell and Landrum 1993; _ Piersma et al. 1993). Information on seasonal and _ regional patterns of prey choice by shorebirds may _ lend insight toward the conservation and protection _ of wetlands that provide critical food resources. The maintenance of healthy viable invertebrate popula- tions is especially important in highly modified land- scapes with changing habitat conditions upon which many shorebirds rely, especially during migration (Skagen and Knopf 1993, 1994). In this paper, we review a sample of data on shorebird diets in the western hemisphere and pro- ~ vide information on the broad spectrum of inverte- brates utilized by shorebird species. We examine dietary differences and similarities of species among regions and dietary overlap among co-occurring species. The implications of our findings on wetland management for shorebirds are discussed. Methods A literature search was conducted to compile pub- lished papers and documents that report invertebrate prey items used by shorebirds in the western hemi- sphere. In these papers, prey items consumed by shorebirds were identified by examining stomach/ esophageal and gizzard contents, pellets, and fecal droppings, or by directly observing foraging. The studies were inconsistent in presentation of inverte- brates consumed, some reporting only orders or fami- lies whereas others reported genera or species. We organized invertebrates taxonomically according to Borror and White (1970), Kozloff (1987), Turgeon et al. (1988), Pennak (1989), and Brusca and Brusca (1990), with additional reference to Levi and Levi (1968) and George and George (1979). We recognize that many potential biases are inher- ent in a data set combining studies with varying objectives and collection methods. Therefore, no attempt was made to quantify the information beyond occurrence of prey taxa in diets, and dietary similarities were computed based on presence/ absence. Basic descriptions of the spectrum of inver- tebrates in shorebird diets are provided. The data were divided into eight distinct regions: Pacific (north and south), Interior (north and south), 419 420 Breeding (4, 5) Migration (3, 4) Pacific Region Migration (7, 22) Winter (9, 9) Nonbreeding (3, 6) Year-round (3, 3) THE CANADIAN FIELD-NATURALIST Breeding (9, 17) Migration (4, 13) Interior Region Breeding (4, 3) Migration (10, 28) Vol. 110 Breeding (1, 1) Migration (10, 7) < e Atlantic Region Migration (9, 11) Gulf Region Winter (3, 9) Central America South America Migration (2, 12) Winter (6, 10) FIGURE |. Geographic distribution of 75 scientific studies that reported shorebird diets in the western hemisphere. Parentheses contain the number of studies and the number of shorebird species represented in eight regions during breeding, migration, and winter- ing periods. Atlantic (north and south), Gulf of Mexico, and Central/South America (modified from Helmers 1992). North and south divisions occurred at the United States—Canada border (Figure 1). Location of study, time of year, and general habitat type (inter- tidal or inland wetlands) were also recorded. Shorebird species were grouped according to body size and foraging guild (Table 1; Helmers 1992). In studies reporting diets of several species in a local area, we calculated similarity indices for pairs of species/guilds based on presence/absence of invertebrate taxa. We also calculated similarity indices to determine dietary overlap in different regions for a given species and for different species within the same region. We used two similarity coef- ficients, Jaccard [C, = = ja + b - j)] and Sorenson [C, = 2j/(a + b)], where a = the number of invertebrate taxa consumed by species/guild A, b = the number of invertebrate taxa consumed by species/guild B, and j = the total number of invertebrate taxa com- t i mon to both species/guilds (Magurran 1988). Values | of both indices range from 0 to 1, with 0 indicating ~ dissimilarity and 1 complete similarity. Results | We reviewed 75 papers that reported prey items of | shorebirds in the western hemisphere. Of these stud- | ies, 53% identified prey items by examination of | stomach/esophageal and gizzard contents, 24% dur- | ing foraging observations, 7% by examination of | pellets and fecal droppings. Sixteen percent of the © papers cited results of other research, and one study | determined food items experimentally. We found reference to diets of 43 shorebird species. To further — examine patterns of dietary overlap on a larger geo- graphic scale, we. reviewed an additional 16 papers on diets of two species in northern Europe that have — been widely studied in both the western and eastern hemispheres, the Black-bellied Plover (Fluvial ) squatarola) and Dunlin (Calidris alpina). 1996 SKAGEN AND OMAN: DIETARY FLEXIBILITY OF SHOREBIRDS 421 TABLE 1. Composition of shorebird guilds based on body size and foraging method (modified after Helmers 1992). Regions of study (see Figure 1) are as follows: AtlIN and AtlS (Atlantic coast north and south of the US-Canada border), IntN and IntS (North American interior region north and south of the US-Canada border), PacN and PacS (Pacific coast north and south of the US-Canada border), Gulf (US side of the Gulf of Mexico), and SAm (Central and South America). References are given for studies that report diets. References for two species in NEur (Northern Europe) are given in parentheses. Guild Common and Scientific Names Plover Black-bellied Plover, Pluvialis squatarola gleaner Lesser Golden Plover, Pluvialis dominica Collared Plover, Charadrius collaris Snowy Plover, Charadrius alexandrinus Wilson’s Plover, Charadrius wilsonia Semipalmated Plover, Charadrius semipalmatus Piping Plover, Charadrius melodus Killdeer, Charadrius vociferus Mountain Plover, Charadrius montanus Oyster- American Black Oystercatcher, catcher Haematopus bachmani prober/prier Small Sanderling, Calidris alba gleaner/ prober Semipalmated Sandpiper, Calidris pusilla Western Sandpiper, Calidris mauri Regions AtIN, AtlS, IntN, IntS, PacS, SAm [NEur] AtIN, IntN, IntS SAm IntS, PacS SAm AtIN, AtlS, IntN, IntS, Gulf, PacS, SAm AtlN IntS IntS, PacS PacS AtlS, IntN, IntS, Gulf, PacS, SAm AtlN, AtlS, IntN, IntS, Gulf, PacN, PacS, SAm IntS, PacN, PacS, SAm References Recher 1966; Baker 1974; Schneider 1978,1983,1985; Hicklin and Smith 1979; Schneider and Harrington 1981; Wishart et al. 1981; Myers 1986; Michaud and Ferron 1990; Helmers 1991; Colwell and Landrum 1993 [Goss-Custard et al. 1977a,b; Evans 1979; Pienkowski 1981, 1983; Zwarts and Drent 1981; Pienkowski et al. 1984; Baird et al. 1985; Evans 1986; Durell and Kelley 1990]. Brooks 1967b; Baker 1977; Bengston et al. 1978; Wishart et al. 1981; Brown and Harris 1988: Helmers 1991. Strauch and Abele 1979. Reeder 1951; Purdue 1976. Strauch and Abele 1979; Schneider 1983. Reeder 1951; Recher 1966; Baker 1973,1977; Hicklin and Smith 1979; Strauch and Abele 1979; Lewis 1983; Schneider 1983; Myers 1986; Michaud and Ferron 1990; Helmers 1991. Shaffer and Laporte 1994. Baldwin 1971b; Rundle 1982; Baldassare and Fischer 1984; Helmers 1991; Fair et al. 1995. Baldwin 1971a. Hartwick and Blaylock 1979; Morrell et al. 1979. Reeder 1951; Couch 1966; Danks 1971; Schneider 1978; Gochfeld and Burger 1980; Myers et al. 1979a,1980; Connors et al. 1981; Duffy et al. 1981; Schneider and Harrington 1981; Lewis 1983; Myers 1984,1986,1988; Helmers 1991. Brooks 1967a,b; Holmes and Pitelka 1968; Baker 1973,1977; Harrington and Groves 1977; Ashkenazie and Safriel 1979; Hicklin and Smith 1979,1984; Schneider 1978; Duffy et al. 1981; Schneider and Harrington 1981; Lewis 1983; Gratto et al.1984; Myers 1986; Boates and Smith 1989; Michaud and Ferron 1990; Napolitano and Ackman 1990; Wilson 1990; Helmers 1991; Gratto-Trevor 1992; Colwell and Landrum 1993. Reeder 1951; Couch 1966; Recher 1966; Senner 1979; Duffy et al. 1981; Quammen 1982,1984; Schneider 1983; Baldassare and Fischer 1984; Senner et al. 1989; Helmers 1991; Colwell and Landrum 1993. (Continued) 422 THE CANADIAN FIELD-NATURALIST TABLE |. Continued. Guild Medium gleaner/ prober Common and Scientific Names Least Sandpiper, Calidris minutilla Small Baird’s Sandpiper, Calidris bairdii White-rumped Sandpiper, Calidris fuscicollis Purple Sandpiper, Calidris maritima Red Knot, Calidris cantus Pectoral Sandpiper, Calidris melanotos Dunlin, Calidris alpina Stilt Sandpiper, Calidris himantopus Short-billed Dowitcher, Limnodromus griseus Long-billed Dowitcher, Limnodromus scolopaceus Greater Yellowlegs, Tringa melanoleuca Lesser Yellowlegs, Tringa flavipes Solitary Sandpiper, Tringa solitaria Willet, Catoptrophorus semipalmatus Spotted Sandpiper, Actitis macularia Regions AtlN, AtlS, IntN, IntS, Gulf, PacS, SAm IntN, IntS, PacN AtlS, IntN, IntS, SAm IntN AtlS, IntN, IntS, PacS, Gulf IntN, IntS, PacN AtlS, IntN, IntS, Gulf, PacN, PacS, [NEur] IntN,IntS AtIN, AtlS, IntN, IntS, Gulf, PacS, SAm IntN, IntS, PacN, PacS IntS, PacS, SAm IntN, IntS, Gulf, SAm IntS AtlS, IntN, Gulf, PacS, SAm SAm Vol. 110 References Couch 1966; Recher 1966; Brooks 1967a,b; Baker 1973,1977; Hicklin and Smith 1979; Lewis 1983; Schneider 1983; Baldassare and Fischer 1984; Gratto et al. 1984; Myers _ 1986; Brown and Harris 1988; Helmers 1991; Colwell and Landrum 1993. Drury 1961; Holmes and Pitelka 1968; Lewis 1983; Baldassare and Fischer 1984; Helmers 1991. Drury 1961; Lewis 1983; Myers 1986; Helmers 1991; Parmelee 1992. Danks 1971. Sperry 1940; Recher 1966; Myers 1986; Helmers 1991; Colwell and Landrum 1993. Brooks 1967a,b; Pitelka 1959; Holmes and Pitelka 1968; Rundle 1982; Lewis 1983; Helmers 1991. Couch 1966; Holmes 1966; Recher 1966; Brooks 1967a,b; Holmes and Pitelka 1968; Baker 1973,1977; Senner 1979; Quammen 1982,1984; Buchanan et al. 1985; Myers 1986; Senner et al. 1989; Brennan et al. 1990; Helmers 1991; Colwell and Landrum 1993 [Bengtson and Svensson 1968; Davidson 1971; Bryant 1979; Goss-Custard et al. 1977a,b; Rands and Barkham 1981; Lifjeld 1984; Pienkowski et al. 1984; Baird et al. 1985; Kelsey and Hassall 1989; Durell and Kelly 1990]. Brooks 1967a,b; Baker 1977; Lewis 1983; Baldassare and Fischer 1984; Helmers 1991. Sperry 1940; Reeder 1951; Baker 1973,1977; Schneider 1978,1983,1985; Hicklin and Smith 1979; Mallory and Schneider 1979; Schneider and Harrington 1981; Quammen 1982; Myers 1984,1986; Helmers 1991. Sperry 1940; Spawn 1941; Recher 1966; Quammen 1982,1984; Lewis 1983; Baldassare and Fischer 1984; Helmers 1991. Spawn 1941; Reeder 1951; Brooks 1967a,b; Schneider 1983; Robert and McNeil 1989; Helmers 1991 . Spawn 1941; Brooks 1967a,b; Baker 1973, 1977; Lewis 1983; Baldassare and Fischer 1984; Rundle 1982; Robert and McNeil 1989; Michaud and Ferron 1990; Helmers 1991. Helmers 1991. Recher 1966; Stenzel et al. 1976; Gochfeld and Burger 1980; Schneider 1983,1985; Myers 1986; Helmers 1991. Schneider 1983. (Continued) 1996 TABLE |. Concluded. Guild Common and Scientific Names American Woodcock, Scolopax minor Common Snipe, Gallinago gallinago Buff-breasted Sandpiper, Tryngites subruficollis Large Hudsonian Godwit, Limosa haemastica gleaner/ prober Marbled Godwit, Limosa fedoa Whimbrel, Numenius phaeopus Long-billed Curlew, Numenius americanus Turnstone Ruddy Turnstone, Arenaria interpres gleaner/ Black Turnstone, Arenaria melanocephala prober Surfbird, Aphriza virgata Avocet/ Stilt American Avocet, Recurvirostra americana gleaner/ Black-necked Stilt, Himantopus mexicanus sweeper Phalarope Wilson’s Phalarope, Phalaropus tricolor pelagic Northern Phalarope, Phalaropus lobatus gleaner Red Phalarope, Phalaropus fulicara The distribution of studies and the numbers of shorebird species represented within regions by sea- son are delineated in Figure 1. Many studies reported results that encompassed several shorebird species or more than one region. The Pacific (north and south), Interior (north and south), and Atlantic (north and south) regions were well represented, cited in 38%, 30%, and 26% of the studies, respectively. Eight studies (11%) were in Central/South America, and only three studies (4%) were conducted in the Gulf of Mexico region. Studies that were conducted dur- ing the breeding season were predominately in the Northern Interior region; the majority of studies in the Southern Interior region and in the Atlantic regions were conducted during migration. Diets of wintering birds were recorded primarily in three regions, the Southern Pacific, the Gulf of Mexico, and Central / South America regions. Invertebrates consumed by western hemisphere shorebirds were taxonomically diverse, representing 12 phyla, 22 classes, 72 order’, 238 families, and 404 SKAGEN AND OMAN: DIETARY FLEXIBILITY OF SHOREBIRDS 423 Regions References IntN, IntS Sperry 1940. IntS, PacS Sperry 1940; Spawn 1941; White and Harris 1966; Brooks 1967a,b; Fritzell et al. 1979; Rundle 1982. IntS Helmers 1991. IntN, IntS Baker 1977; Helmers 1991. IntS, PacS Reeder 1951; Recher 1966; Helmers 1991. SAm Schneider 1983,1985; Velasquez and Navarro 1993. PacS Stenzel et al. 1976. AtlS, IntN, Danks 1971; Gochfeld and Burger 1980; Gulf, SAm Wishart et al. 1981; Schneider 1983,1985; Myers 1986. PacN Norton et al. 1990. PacN Norton et al. 1990. IntS, PacS Recher 1966; Hamilton 1975; Quammen 1982,1984; Baldassare and Fischer 1984; Mahoney and Jehl 1985. IntS, SAm Hamilton 1975; Robert and McNeil 1989. IntS Brooks 1967b; Baldassare and Fischer 1984; Mahoney and Jehl 1985; Brown and Harris 1988; Jehl 1988. IntN, IntS, Baker 1977; Mercier and Gaskin 1985; PacS Jehl 1986; Brown and Harris 1988. IntN Danks 1971; Dodson and Egger 1980. genera (Appendix 1). The classes with the most reported taxa were Insecta (phylum Arthropoda, sub- phylum Uniramia; 117 families and 242 genera), Malacostraca (phylum Arthropoda, subphylum Crustacea; 33 families and 47 genera), Gastropoda (phylum Mollusca; 29 families and 40 genera), Bivalvia (phylum Mollusca; 14 families and 21 gen- era), and Polychaeta (phylum Annelida; 11 families and 15 genera). The orders containing the most reported taxa were Coleoptera (class Insecta, 33 fami- lies and 142 genera), Diptera (class Insecta; 28 fami- lies and 28 genera), Hemiptera (class Insecta; 18 fam- ilies and 31 genera), Decapoda (class Malacostraca; 14 families and 19 genera), Hymenoptera (class Insecta; !1 families and 22 genera), and Amphipoda (class Malacostraca, 9 families and 16 genera). Seeds were occasionally reported as minor items in shorebird diets, usually comprising less than 5% but occasionally about 10% of prey items (Couch 1966; White and Harris 1966; Baker 1977; Senner 1979: Rundle 1982; Mercier and Gaskin 1985; Vol. 110 THE CANADIAN FIELD-NATURALIST 424 panuiuod 8I Ol I I G I € al Ol vioidiwisy ¢ € I I Tel € viajdowoy G I I el G eyeUuOpO I I I I eiajdorawieydq I I I R[OquIa|[OD BOOSU] 9 (6 I I ¢ ll, t pooeuind, oeprydo10g avplewury I¢ v € (6 8 v iL 8 Ol 6 € L 6 epodrydury iL I I I c I L € epodosy] v I C I A € BIORpPIvUrR |, 8T It c I iG € I - 9 G TE Il epodeoeq BOBISODR|LIN| epodoimry winyAyg el C I I I € I G C Dai (eposensQ) I I al I vorovsoy,L (4 I I I eploonordiepy ¢ I I G I Teal (epodado) epodoy[ixeypy C I I I voBNsSOyUO, I I I I eIZ.0prID € I I I I poe souy epodoryourig vpodoimiry wnyAyd 9 I I I I I I AE € eprxejojdey vyovysosl[Q v I I I I L I eprloqes I I L I epr[oqoloL, I I di I eproruny SeploION Of G I S I € ¢ C Cc v ¢ AE 4 eproopol[Ayd septfondeD I] I a I Z G Z I i € eprfoudeS Cc I I ale I eplyMeLiy 6 I I c Z I I ai I epruoids c I I aE I eprtulqiO vyoeyoA[od eprpouuy winyAyd I I epo}eulaN untAYyg € I Z Af, I BdLIOUIOU01N]9H pjdouy 0 voyIoulaN winyAyd I I I I (eydiowounoe.ia) aeisuodsowag viafliog winjAyg I I IE (eplIojiurUelo.) evurpoores umnyAyg zSoltuey §=ssouenns9Q “TTIM HANI OddS INNdG VSHT VSHS VSHM AUNVS TddS Tddd yeyiqeH = Sol ue] IOp1O SseID uowwo0y) jo Jaquinyy saroads priqaioys jo d}V.AQOLIOAUT dIVAQOIAUT [B10], JOquInyy ‘pajouap ore ‘(]) puelur pue (1) fepy ‘adAj yeyqvy [eloues OY, “Sassejoqns ase SasoyjUored UT SOUL UOXE) dRIQIIAU] “SISIP PALQIOYS Ul SIOPIO djB.IGOJIOAUT JO JOULIIOCUIT DALAL JO XOPUT ISIBOD ¥ SB SdAJOS S9dUdLINIDO Jo Jaquinu sy{, ‘arsydstwuey w9}saM Uy} Ur satoads pItqar1OYs Ud} JO SJoIP dy} UT (SINDDO ATIWUL] YTIYM UL SUOTSII JO JoqUINU >< SdTTIUIR} JO JAqUINU) sd9oUdLINI9O JO JOqUINU PUL So}PIGOOAUT JO SOT[IUL] JO JOQUINN' ‘7 ATAVL, 425 DIETARY FLEXIBILITY OF SHOREBIRDS SKAGEN AND OMAN 1996 ‘saloads p1lga1oys Ua} dy} JO SLOU 10 UIAVS JO SJOIP UT PoLINd9O Jey) Sore, “TOTEM - TTIM ‘S82]MOTI2X asso] - FAAT “JOYUMod pal[tg-40ys - OS ‘UNG - INA “Iedidpueg ysvoT - yy] ‘Jodidpurg pareujedruisg - WSas ‘Jedidpueg usaisaqy - WSAA\ ‘SUT[Iapues - CNVS ‘1eAo]d payeuyediuag - TqAS “IOAo[d PIf[9q-YRIG - TAA ‘SMO]JOJ sv are sorsads psrgaroys 10} sapoD, c9 EG OS 8C EG 8c Of 8E €C JOIp Ul SOT[IWUILJ BBIQVIOAUT JO IOQUINN] Ta el LI al CE Gl 91 TT a SaIpnys JO Jaquinyy v L 9 L 8 4 9 is 9 paquasaidar suorsai Jo Jaquinyy € I I sadyyoIascC, eyeployg wnyAyg I I if I aL, I eprpody voprlolimuyjo[oH vyeuLapouryoy wnyAyg G I I tall, RIeWINV Rye[NONIVUy epodoryoeig umnyAyg 9 Z I 4 tf I eploky IePLI9Ud A, seprlunlaL Gy I g ¢ € v 9 € ¢ ¢ L 9 Bplorous A eepynAW 8 I é I I I I ic I eplopnky I I L I epropnonN BIATCATY I I Al I vyeuowndosryo1y € I I I I I vroydojeuuoseg € C I ll l voov]japruresikg SUpriTesseN] I] I I I C I € I als P vpodenseso0an aepligoipAH 9 I (6 € (6 ¢ G v € L ¢ epodonsesosayy I I Tf I epodonsesooryo1y epodonsey vosnyfoyy wunpAyd £ I I I Iga WRoy 9 G if i I I I avourly L I I I I I I db I vinsoydry BILIBOTJOYO I I I viajdopida] v € I I (4 via doyoif, depl[owosAIYD oepiquie,) c9 61 G II 9 Ol € ¢ L Cc I 81 vraidoajoy sepipodoysrjoq oepupxydg oeprndiy, oeprurouoly) LL 81 ¢ el L Il 9 1 6 € Iba CT vioidiq I I I via}doyviC vi € 9 I I I G LI 9 viojdousw Ay Sole SeoustINnI9O) TIM AAAI OGdS INNG VSAI VSHS VSHM GNVS IWddS aWddd yeyqeH = Sol Turley JOpsJO S410) UOUIWIOD) jo fe isardads piiqor0yg ae d}.IQOVIOAUT o}B.IQOIOAUT panuyuoy “7 ATaV 1 426 Brown and Harris 1988; Brennan et al 1990; Fair et al. 1995). Seeds were prevalent, however, in the diets of some fall migrants in the interior region (Brooks 1967a; Baldassarre and Fischer 1984), com- prising 10-20% of the diets of Common Snipe (Gallinago gallinago), Pectoral Sandpipers (Calidris melanotus), Least Sandpipers (C. minutilla), Western Sandpipers (C. mauri), Long-billed Dowitchers (Limnodromus scolopaceus), American Avocets (Recurvirostra americana), and, in one study (Baldassare and Fischer 1984), nearly 40% of the diet of Stilt Sandpipers (C. himantopus). We summarized the diets of the ten most widely studied species, which represented three foraging guilds, plovers (gleaners), small gleaner/probers, and medium-sized gleaner/probers. We present a coarse index of the relative importance of each invertebrate order in the diet of each of these species; this index is the number of occurrences (the number of invertebrate families multiplied by the number of regions in which the family was reported: Table 2). All ten species exhibited considerable dietary breadth, consuming an average of 36 (range 23-65) families of invertebrates. The invertebrate orders most frequently cited in the diets of these ten species in inland areas were Diptera (primarily chironomid larvae, tipulids, and ephydrids) and Coleoptera (primarily carabid beetles). In inter- tidal areas, the orders Veneroida (primarily tellinid and venerid clams), Amphipoda (primarily gammarid and corophid amphipods), and Phyllodocida (primari- Dolichopodidae Muscidae Curculionidae (Arachnida) Vi [ ae Hydrophilidae | Physidae NS “ SOUTH ™ INTERIOR ~~ (intand) THE CANADIAN FIELD-NATURALIST Chironomidae Vol. 110 ly nereid polychaete worms) were the most often cited (Table 2). There was extensive dietary overlap between these ten species, with 15 families of inverte- brates common to the diets of seven or more of the ten species (Table 2). Nereid polychaetes, chironomid lar- vae, and venerid clams were in the diets of all ten of these shorebird species. Minimal dietary overlap for six of these ten shorebird species occurred between inland and tidal areas; only the insect families Dolichopodidae, Chironomidae, and Corixidae occurred in both inland and tidal areas. The diet of the Semipalmated Sandpiper (C. pusil- la) is represented by 22 studies across all eight regions and is the most extensively studied of all shorebird diets. Collectively, this species consumed a wide variety of invertebrates, spanning 25 orders, 37 families, and one additional subclass. Classes with the most families were Insecta (14 families and 2 additional orders), Malacostraca (7 families and one additional order), and Polychaeta (5 families). There was very little overlap in diets between inland and tidal areas in North America and little overlap in taxa consumed in the north Atlantic and south Atlantic regions (Figure 2, Table 3). Dunlin diets, referenced in 17 studies across six North American regions, spanned 19 orders and 50 families of invertebrates, with two additional sub- classes. Insecta (23 families) and Malacostraca (10 families were the most commonly reported food items. Bivalvia was represented by five families, and (Harpacticoida) (lsopoda) Corophiidae Podoceridae Nannastacidae Littorinidae Muricidae Nassaridae Nuculidae Chiridotidae A Spionidae AN Nereidae \ if Hydrobiidae \ \ Crangonidae \ / Veneridae SOUTH Orbiniidae ATLANTIC Maidanidae (tidal) Haustoridae if \ Gammaridae / Limulidae Ya \ Mytilidae es ‘\ a Telinidae 4 Sa map FiGuRE 2. Families of invertebrates reported in diets of Semipalmated Sandpipers in four geographic regions. Circles delineate diet overlap among regions. Taxon names in parentheses are orders or subclasses. Data on the North Interior region are from the breeding season and for other regions from migration. ee ee 1996 SKAGEN AND OMAN: DIETARY FLEXIBILITY OF SHOREBIRDS 427 TABLE 3. Similarity coefficients (CG = Jaccard and C, = Sorenson; Magurran 1988) for regional pairs for individual species. C; C, Semipalmated Plover North Atlantic tidal - Central American tidal 0.09 0.16 North Atlantic tidal - North Interior inland 0.04 0.08 North Interior inland - Central American tidal 0 0 South Pacific tidal - Central American tidal 0.17 0.29 Black-bellied Plover North Atlantic tidal - South Atlantic tidal 0.11 0.20 South Pacific tidal - South Atlantic tidal 0.27 0.42 South Pacific tidal - Central American tidal 0.30 0.46 Northern Europe tidal - North Atlantic tidal 0.10 0.18 Semipalmated Sandpiper North Interior inland - South Interior inland 0.22 0.36 Interior inland - Atlantic tidal 0.06 0.12 North Atlantic tidal - South Atlantic tidal 0.18 0.30 Dunlin Pacific tidal - Pacific inland 0.10 0.18 Pacific tidal - Interior inland 0.05 0.10 North Pacific tidal - South Pacific tidal 0.31 0.47 Pacific inland - Interior inland 0.25 0.40 Pacific tidal - Northern Europe tidal 0.23 0.38 Polychaeta and Gastropoda by four families each. Diet similarity and overlap Dietary overlap between regions and between tidal We evaluated diet similarity across regions for and inland areas again was minimal (Figure 3). four species by calculating similarity coefficients PACIFIC Arenicolidae (inland) (tidal) Gammandae Cardiidae Saldidae Ichneumonidae Chironomidae Sciaridae — netehenadiaes — Dolichopodidae | Mycetophilidae Tipulidae cE 5 Carabidae Muscidae NORTH 7 Ansiogammaridae Staphylinidae Chrysomelidae INTERIOR a Corophiidae Dytiscidae (intand) Hydrobiidae Linyphiidae 7A Mytilidae Wp, Tellinidae IN Veneridae / Myidee Chironomidae / Dytiscidae Physidae \ | Spionidae Tenthredinidae \ Capitellidae Braconidae | | Nereidae Tubificidae sueyeilidae Aegidae Talitridae ceayecyaae SOUTH \ Tanaidae Phoxocephalidae Haliptidae INTERIOR Hyalidee Hydrophilidae (inland) \ Aare Curculionidae Wf \ Nannastacidae dh NN re Nitidulidae Nassariidae S SS SOUTH x ae — PaciFic— (tidal) FIGURE 3. Families of invertebrates reported in Dunlin diets in five geographic regions in North America. Circles delineate diet overlap among regions. Taxon names in paren- theses are orders or subclasses. Data from the inland North Pacific and North Interior regions are from the breeding season, on the tidal North Pacific and South Interior regions from migration, and the South Pacific from migration and winter. 428 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 4. Similarity coefficients (C, = Jaccard and C, = Sorenson; Magurran 1988) of pairs of species and foraging guilds. Species codes are as follows. SEPL - Semipalmated Plover, BBPL - Black-bellied Plover, SESA - Semipalmated Sandpiper, DUNL - Dunlin. C; ee Within-region species pairs, all studies in region SEPL - SESA (North Interior, inland) 0.42 0.59 SESA - DUNL (North and South Interior, inland) 0.35 0.51 BBPL - DUNL (Northern Europe, tidal) 0.32 0.48 Within-study species pairs, for comparison SEPL - SESA (North Interior, inland; n = 2) 0.64 0.75 SESA - DUNL (North/South Interior, inland; n= 4) 0.67 0.77 BBPL - DUNL (Northern Europe, tidal; n = 4) 0.59 0.74 Means for all within-study pairs Species pairs within guilds (n = 29) 0.51 0.62 Guild pairs (n = 28) 0.55 0.67 (a) SEMIPALMATED SEMIPALMATED PLOVER ~ SANDPIPER ~~ SS 7 ‘ if \ / rellnte Anostraca) \ / Hymenoptera) Conchostraca) \ ; 4 Ceratopogonidae Copepoda) Stratiomyidae | Chironomidae (Collembola) \ Haplidae Tipulidae Corixidae | Dytiscidae | Ephydridae Psychodidae (Araneae) Dolichopodidae Carabidae | Hydrobiidae \ Muscidae Staphylinidae \ Hydrophilidae Dytiscidae / Chrysomelidae Physidae / Curculionidae / Z vA Fa rae a _———_ — SEMIPALMATED pe DUNLIN _ SANDPIPER nN N N N\ Anostraca) : nostraca leceeroet aca} / \ Collembola) / Tipulidae \ Gopecedel Dolichopodidae \ Ostracoda Muscidae Cicadellidae Corixidae Chrysomelidae Stratiomyidae | (Hymenoptera) | Curculionidae Haplidae Ceratopogonidae Dytiscidae Nitidulidae | Ephydridae \ Hydrophilidae Psychodidae \ Physidae i Carabidae / Staphylinidae / / 7 7 as - FIGURE 4. (a) Invertebrate families reported in diets of Semipalmated Plovers and Semipalmated Sandpipers in the north Interior inland region. (b) Invertebrate families reported in diets of Semipalmated Sandpipers and Dunlin in north and south Interior inland regions. Circles delineate diet overlap between species. Taxon names in paren- theses are orders or subclasses. _ 1996 _ based on families (or orders) for pairs of regions. _ Again, we include dietary information for two of _ these widely-studied species for northern Europe for _ additional perspective. There was relatively little overlap in diets between regions (Table 3); C. values averaged 0.16 and ranged from 0 to 0.31. C, values were higher, averaging 0.26 and ranging from 0 to 0.47. Similarity coefficients for pairs of the same major habitat type (inland or tidal) were greater than for pairs with contrasting habitat types (average C. = 0.20 for same and 0.05 for different habitat types). We also evaluated diet similarities of three pairs of species with sufficient records within the same region (Table 4). There was substantial overlap in diet between Semipalmated Plovers and Semipalmated Sandpipers in the north Interior inland region, and between Semipalmated Sandpipers and Dunlin in the Interior inland regions (Figure 4a,b) and between Black-bellied Plovers and Dunlin in northern Europe. Without exception, similarity indices for the species pairs exceeded the values for species across region pairs, as presented in Table 3. When species occurred within the same study, the diet similarities for the same three species pairs were even greater (Table 4). In 29 studies including more than one member of a foraging guild, there was substantial diet overlap between species within guilds (Table 4), as indicated by the similarity coefficients, average C. = 0.51 (+ 0.29 sd, n = 29) and average C, = 0.62 (+ 0.26 sd, n = 29). In 28 studies with more than one foraging guild represented, diet overlap between guilds was similarly extensive (Table 4), represented by average C = 0.55 (+ 0.26 sd, n = 28) and average C, = 0.67 (+ 0.22 sd, n = 28). Discussion Collectively, shorebird diets span a tremendous breadth of invertebrate taxa. Many shorebird species are euryphagic or cosmopolitan in food choice, and local studies often do not reveal the potential breadth of diet choice for-a species across its geographic range. Dietary flexibility allows for the exploitation of variable resources and is highly advantageous to many shorebirds that migrate across vast landscapes and inhabit a variety of wetland types, both inland and tidal, during their yearly cycles. Patterns of dietary similarity/overlap in shorebirds are also consistent with a flexible foraging strategy. Our analyses showed the greatest dietary similarities between pairs of shorebird species or guilds within a given study; species pairs within a given region were intermediate in diet similarity, and the same species in different regions had the lowest similarity. These findings probably reflect the similarities in food resources within studies and regions and the differ- ences between seasons and regions. The observed patterns of dietary similarities would be expected if shorebirds choose the predorninant food items avail- able within each local area. SKAGEN AND OMAN: DIETARY FLEXIBILITY OF SHOREBIRDS 429 Several authors have noted the opportunism in shorebirds, especially species in the small and medi- um sandpiper guilds (Couch 1966; Recher 1966; Holmes and Pitelka 1968; Davidson 1971; Thomas and Dartnell 1971; Gochfeld and Burger 1980; Lewis 1983; Fair et al. 1995). In these studies, opportunism meant taking prey in accordance with its availability (Couch 1966; Recher 1966; Thomas and Dartnall 1971); altering food choice to take advantage of inver- tebrates available (Davidson 1971), or concentrating where prey is abundant (Holmes and Pitelka 1968). Lewis (1983) and Skagen and Knopf (1993, 1994) emphasized the advantages of flexible, opportunistic strategies for shorebirds that depend on highly unpre- dictable resources. Several examples demonstrate that shorebirds combine opportunistic foraging with specific prey selection. Even though many shorebird species are euryphagous, selection for prey type and size has been documented through comparisons of diets of coexisting species (Strauch and Abele 1979; Lifjeld 1984; Senner et al. 1989; Michaud and Ferron 1990) and through comparisons of the abundances of prey items in sediments and in shorebird diets (Holmes 1966; Brooks 1967a; Baker 1977; Bengston et al. 1978; Gratto et al. 1984; Velasquez and Navarro 1993; Fair et al. 1995). Redshanks (Tringa totanus) selected for polychaete size only when the preferred size was abundant (Goss-Custard 1977a). In another study, Redshanks selected amphipods above a threshold size relative to their proportion in the sub- strate (Goss-Custard 1969). Morrell et al. (1979) proposed that diets of Black Oystercatchers (Haematopus bachmani) are determined primarily by prey availability and only secondarily by distinct preferences for prey type. Brooks (1967a) and Holmes (1966) found birds selecting a few preferred items and taking other foods relative to their avail- ability in the substrate, and Schneider (1978) report- ed the selective removal of numerically dominant invertebrate species. Species and foraging guilds substantially overlap in diet within a geographic region and within a local study area. Even in light of reported dietary differ- ences, many authors commented on the high degree of dietary overlap among coexisting species (Couch 1966; Recher 1966; Holmes and Pitelka 1968; Baker 1977; Hicklin and Smith 1979; Senner et al. 1989; Baldassarre and Fischer 1984; Lifjeld 1984; Brown and Harris 1988; Michaud and Ferron 1990). Extensive dietary overlap may occur if coexisting species forage opportunistically on a limited number of invertebrate taxa in a small area (Holmes and Pitelka 1968). Dietary overlap based on invertebrate taxonomy at the family level, as discussed here, does not neces- sarily reflect a lack of ecological segregation. Niche separation may be achieved as a result of behavioral differences or use of different substrates or micro- 430 habitats (Drury 1961; Recher 1966; Bengtson and Svensson 1968; Thomas and Dartnall 1971; Baker and Baker 1973; Baker 1977; Connors et al. 1979; Senner et al. 1989; Strauch and Abele 1979; Quammen 1982; Lewis 1983; Baldassare and Fischer 1984; Grant 1984; Helmers 1991). For example, habitat partitioning based on tide cycles, substrate, microhabitat features, and water depth is well documented (Recher 1966; Brooks 1967a; Burger et al. 1977; Colwell and Oring 1988; Helmers 1991; Colwell and Landrum 1993). The flexible nature of food choice in shorebirds has important implications for the management and restoration of wetland habitats. Because shorebirds utilize a broad range of invertebrate taxa, efforts to maintain vital food resources for shorebirds during all seasons should focus on maintaining hydrologic regimes and ecosystem processes that promote the growth and maintenance of invertebrate populations in general. Particular invertebrate taxa need not be targeted. Wetland management, restoration, and cre- ation that enhances naturally-occurring populations of invertebrates will undoubtedly be successful if the invertebrates are accessible to shorebirds. Acknowledgments We are grateful to Boris Kondratieff and Paul Opler for reviewing the taxonomic arrangements of the invertebrates. Scott Kahan and Erin Muths assist- ed in the literature review and data compilation. We thank Patricia L. Kennedy, Murray Laubhan, and Erin Muths for their valuable comments on earlier versions of the manuscript. 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Received 27 September 1995 Accepted 8 February 1996 433 DIETARY FLEXIBILITY OF SHOREBIRDS SKAGEN AND OMAN 1996 panuljuo,) s[reus DIUOKA J, sjreus pigopaH] suasyuo. vpoorulue pnut DIONUULY aepriqoipAH epodonsesosayy syoduity EIS) sepl[oed saqliou DUIJIAIN, aePNLION sjodunty DIJOT asepimoy evpodonsesoavyoiy eIyouRIqosolg DJ MOgAN I, deprapruresAg voov]jopruuesAg soyqqnq-[a1eg [ouueyo pUuld0ajoy oeplipueydess Dyjakvos DSNJOY sopqqnq-[areq DIMOISOPE depisnjoy vopidseyeydap eryourigoyisidG epodonsey vOSNTOJ suvozociq paiouny deprT[ayeuiny eplyjayeuny eyeUlae[o}oe[AY vozoAIg syfoys dure] DINSUIT _— eyeuUlONny pye[Nonseuy vepodoryorig soyoady pyjapqodiayy depr{[apqodiay voulpniiy eplulpniiy DAOpK]Od oidsojqa.ls oepruoids epluoids SNLMIULM Id SULIOMBRI SUI SIAAAN sayjuvan JEPIOION sdyjydan oeprAjydan DAIIKID) aepladk[H eploopoT[Ayd SULIOM puRs sojdojo2s oepUIgICE eplUlqiO aeproydo epueydo SIMAULAQUINT depLoULIquin | eprorung DIWAOfMAAND depl[NyeLD eplyMeLuiD SULIOM OOqUIeG pyjauawuda]y oeprurpley SNJSDULOJON SNISDUOAIIA LY pyjandvy seprfaudes SWIOMSN] DIONUAIDGAY deprljoorusry eprjaudes pyovyodog ; S2PlOAPOUUUny seproyiqn | psoydoqojoy]y WIOMY}IeS SNILAQUNT deprloliquin’] eprxejo[dey vyavysosI[O vpljouuy vipsy vaploipAH vozoIpAH eleplud SULIOMpUNOI epoleuaN| SUIIOM stosogoid SNINIDAGIAID oeprloury BOLIOWIIU0IN}OH rjdouy RoLIOUION sosuods Jayemysaly oepryisuods epliaposo[dey vydiowounseia) aeisuodsowsqd vIdJLOg SUBIOJTUTLUR IO} PPLOJIUIUeIO. esOTNONeJO;NULIDH vulpoors vioydosyseuoosies aIWVN NOWNOD SNNdD ATINVA Ric(an(@) ssvToans ssv1D. WNTAHdEN§ WOTAHd ‘o1oydsttuay UI9]SOM JY} UI SJOIP PALqoIOYsS JO SaIpnys G/L UT poyodar suo} Adid 93v.1Ga}IOAUT JO JXQJUOS SIWIOUOXB], “| XIGNAddV THE CANADIAN FIELD-NATURALIST Vol. 110 434 panuyuoy SapPIO9 wnipsvzouny oeplupies) a1e\se IADISY oeplieisy PPlOIgUd A Ja}sho DasISEQ aeplons¢C sdoyjeos uajaad aeplunoad eploanseC, Jassnur as1oY SN]OIpO| sjessnul sna oeplnA eplopnA piwojdty S][2ysJos Di oeplAjy ePLOA| s][ous ye DADpDUuYy oeploly eplooly BIyoULIqI [Owe T] wueyoynu DININN oeprynonyy eprloynonNny vIyouRrlqojolg RIA[RALG uloy sues diy yoru} SIQAOUD] UIOY SURI ISP OMI DULOSI]AET o1AS Yse SHINDAK aepriqiouryd sjreus yonod vskyd aeprsAyd DIONUSDIS sjodurl] Jaye MyYsory pavuucT BIIOSSOJ UOP[OS DIADSSOJ oeplovuuA] vloydoeuwiose g. snduvjay oepriqol[q DUAY sndwieyau snduvjay depipoduryayy eyeuowndoaryory pyeuowyng DIasuDy oeprluin y, MEO) seplAllO s]Ieus UOOUI sanuyog oeplonen Bsseu SNLUDSSDN sjreus pnut DSSDUDATT oeplliessen, syjayMsop DIJA@INN aeprlounyyy SePI[[OUISIL IA] [leus dAop Jeun] DIAL [ [reus aaop Apaais sTyoADUy oepryfaquinjos yyoum WNUIIING — aeprurong vpodo.ses0oN BIVATBA ISPLI oI} DIDAIDA aepneale A voudys sisdoauays aeprisdoausys pynsuly LPIOSSTY s]foys wr0y paplyjlad depIprIUILjog s]JoysuooUl oeprloneNn sopyurmtiad DULLOINIT oeprluLIonrTy] vunde] DUNIVT oeplunoe’] aS sepisdormpiiad DULOISDIG wniygg deprIyIad vpodonsesosajy BIYIULIQOSOIg epodonsey BOSNT[OW aNVN NOWNWOD SANay ATINVA Ric(et(@) ssv10ans SSVI) WOTAHdENS WOTAHd ‘panuyuoy “| XIGNaddy 435 DIETARY FLEXIBILITY OF SHOREBIRDS SKAGEN AND OMAN 1996 panuluoy snu1oipo0ad¢ snadouvd&ing aepipodAsQ snyjDUsipa—d aepIpoy’y sqeio pues DILAIU ovepiddiyy sqvid o10Ys snsdvasKyovd sqevio pnur snsdpssiwuapy ovpisdein duis uOsUDAD oepluosuvl) uowan]vg aepueg SnulgIDIZ sqeid ony AdIUDD aepligours dumtys jsoys DSSDUDILIDI oeprisseuely[eg sqeio Jo[pply aepipodkog YSIJMvIO SnNADQUDD aeplor|sy epodroaq voelsoov[eung BORI]SOOL|LIA] duwitiys pases ‘spoov.so studkr0j KJ aepipuidks epoornsO SNUDIAO J, pDAOMa Kn oeplouls |, snupjpz0pnas sa8vdoajuag oepisodonuay DIADIV spodadoos prouryes snuojdvig oepruoj}deiq vplourye) sprodojoAo oepidoaka eplodojoka spodadoo eprloonoediey epodados sajoeureq DIAN aeprfadeas volovioy,L, eIpodtui epodoy[ixey duiys wieyjo voBsSOyouoD snsopay) suvlad0peyo SNIAIINANT piuydvg Seal IOV DU1OW aepruydeg vIQIOPR[D dumtiys outig Arey DIWUAJAY oepliiwoyiy vovlsouy epodoryourig voov}snig epodoimsy wed poTfoysprey snua, suey DIJQUISUDA T, DIDYJOJOd , Weld Was UMOIG IJADISVAD sued Wos puiuay deploud A, DUNT au: suIe[o DUOIDIN oeplury [aL uryo vod WNnIpISi_ depriioeyds wryo Jans JIeMp pIUulyny aepLoR| sT[oys os8pam xpuoqg oeprlovuod pnbijis Sepr[ey nD BPlOIOUs A, BY SUCIG HOU a BIATRATY vosn{ [OW aINVN NOWNOD saNay ATINVA Bic(ah(@) ssvTOgns SsvTD WNTAHdEN§ WOTAHd ‘panuljuoyd “| XIGNAddV THE CANADIAN FIELD-NATURALIST Vol. 110 436 panuyuo) SJOYOLIO [AUIS snpiydoyinag aepIplioey [AID E stoddoyssvis oepIpLoy eiajdoyuig sorpAeu pyaeq Syavqyyv oepnoeg eiaydorawaydyq R108AINg sjreysutids R[OqUIdT[OD eloskiady pIOaSU] PIUIeIIU) dumtiys poas snjoouvdg ovpreur |, pyaysojdaT ovpreuryeied vooepreur y, puiosavydsoxy oepruoriaeyds snulouy deplajop] ssnq Mos o1yenbe DUDIOMIIXA deplurlollD Sn]]aSY oeply[asv pAanyjvKy oepunyUy pjaulv0y aepisoy epodos] S1JKJSOANAXE) punoojdaT sidspjxduva deprlor}seuueN uOmNaT oeptiuoona’] eoovuing, DUDIYSDA I. svayy yoraq vaplojsayItE svayy yoraq DISAYIAC svapy yoraq pyajpay oeplyye |, piyoyng aeplias0pog snxoydvivg aepreydasoxoyg SajsayIsO]] V oepleAH snxoydoyoi SNLANDISNDYOJOA SMLLOISNDYOY SNIAOJSNDYOYIUDIV oeplsioisney SNADUUUDS OUIYIT SNADUMUDL oeplieulUey wniydo10d aepriydoi0g SnADUUUDSOIsSUy —- deplvWIWURSOIsUY stoddoypues aoyjiduy oeproyjduy epodiydury ePLVollod snadouvg py diay oepmyjuex duntys pnw pigasody ovpiiqesoda, sqeio onyq sajoaulyva aeprlunyiog snapuad oeplovuad SnANSD SAISIANSVG aepunsedg sqeio I9]pply DIN aeprpodAoQ epodesaq ROBISOITRIA| voovjysnia epodommuy aNVN NOWNOD SANAD ATINVA c(ens(@) SSVTOENS ssv19 WO TAHdENS§ WOTAHd ‘panuljuo) “| XIGNaddY 437 DIETARY FLEXIBILITY OF SHOREBIRDS SKAGEN AND OMAN 1996 panuljuo,) sajaauayy SNISTYISN SNYJaJSOUsD]y polyroso[Y) pspupg ssnq yuns snoyaody aeprluroyeyuag SIOUIUIIMS yor Dald aepyoouo}ON s8nq JayeM Suldsa10 SILOIO]Ag oepliooneN s8nq jasuep SIQDN oepiqeNn ssnq jueyd oepliyy SPII[2AOSOUI DYaAosayy IEPIIAOSI|Y SNYJAANAIG SnISAN s14090aH) s8nq paos snucy oeplavsAT sIoplys 19]eM STAID) depLayH pydns]pH s8nq SUIMO.LING snapsuvd ovplupAd DXIMODOYINA DXILOIDU]D DXLUOD udtROg J3}eM DXIAOIOJIAY depIxLiog ssnq o1s90u aeplusvoULOD S3Nq PaI}00J-Jeo] SNZ1MOD S3Nq Po}00J-Jeo] sniafny depla1oD ssnq JayeM jurIs pwuojsojag depHeUO}sojog ssnq JaMo[y deplLoooyUYy viaduiay sd} viajdoueshy Djja11aas8aq oepliaidoypiyg ddI] Palq wnjpydaso0djoD aepruodousy varydol|ey SOT[Jouo}s viajydooa}g SUMAYID] ; 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Loggerhead Shrike, Lanius ludovicianus, in Southeastern Alberta RONALD R. BJORGE and DAvID R. C. PRESCOTT 1Alberta Environmental Protection, Natural Resources Service, Suite 404, 4911-5Ist Street, Red Deer, Alberta T4N 6V4 2Alberta NAWMP Centre, 5th Floor, 10011-109th Street, Edmonton, Alberta T5J 3S8 Bjorge, Ronald R., and David R. C. Prescott. 1996. Population estimate and habitat associations of the Loggerhead Shrike, Lanius ludovicianus, in southeastern Alberta. Canadian Field-Naturalist 110(3): 445-449. We used 26 randomly-selected 41.5 km? blocks to census Loggerhead Shrike populations in core breeding areas of south- eastern Alberta in 1993. We estimated the breeding population in the 23 600 km? study area (approximately one-third of the species’ breeding range in Alberta) to be 2477 pairs (95% confidence interval of 1588 to 3365 pairs), which is higher than total population estimates made previously for the province as a whole. The number of breeding pairs in a block was highest where shrubs were prevalent, and shrikes were found more often than expected in areas with farmyards/shelterbelts and road allowances, and where a variety of land uses were present. Almost half of all shrikes were observed >200 m from a road, and would probably be undetectable by roadside surveys. Furthermore, only 63% of birds near roads were detected from a moving vehicle. We therefore suggest that only 32.9% of birds on the study area would be detected by a roadside survey. Key Words: Loggerhead Shrike, Lanius ludovicianus, habitat selection, population size, censuses, Alberta. Loggerhead Shrike (Lanius ludovicianus) popula- tions are considered to be in jeopardy throughout much of their range. In the United States and Canada, populations have declined significantly since the 1960s (Cadman 1985; Morrison 1981; Robbins et al. 1986), although numbers have been relatively stable over the past decade (Price et al. 1995). The Loggerhead Shrike is classified as endan- gered in eastern Canada and threatened in western Canada (Johns et al. 1994). In the United States, the Loggerhead Shrike has been included in the Audubon Blue List for declining species (Tate 1986). Several states in the Great Lakes region have listed the species as being endangered (Hands et al. 1989). The management of sensitive species requires measurements of populations size and trends. This is especially true when recovery plans tie management goals to specific population sizes (Johns et al. 1994). Population size and trends of the Loggerhead Shrike have been estimated by a variety of methods. These include Christmas Bird Counts (Morrison 1981), Breeding Bird Surveys (Robbins et al. 1986), road- side surveys (Telfer et. al. 1989; Robert and Laporte 1991; Smith and Kruse 1992), and complete counts in the case of small or localized populations (Cadman 1985; Wershler 1989*). In Alberta, the range of the Loggerhead Shrike appears to have contracted substantially during *Unpublished, see Document Cited Section which follows Acknowledgments. ~ recent decades (Telfer et al. 1989; Collister 1994), but the current population size of the species is unknown. Wershler (1989*) estimated a provincial population of 350 pairs based on complete counts of sites which were known to support shrikes, and extrapolations to other areas of known habitat. An assumption of that approach was that few shrikes were located elsewhere in the province, and that such populations were at very low densities. Extensive roadside surveys conducted by Telfer et al. (1989) indicated “fewer than 1000 pairs” in the province. However, information from the Alberta breeding bird atlas project (Semenchuk 1992), and casual observa- tion by wildlife professionals, suggested that Loggerhead Shrikes may be more abundant. Our primary objective was to estimate the popula- tion size of Loggerhead Shrikes in the core breeding areas of Alberta using a random sampling technique. This approach has been used to estimate widely-dis- persed populations of other endangered or relatively rare species (Leighton et al. 1979; Schmutz 1984). Random sampling allows investigators to determine distribution over large areas and to estimate popula- tions with a measure of precision. Secondary objec- tives were to describe habitat features of areas occu- pied by shrikes, and to assess the effectiveness of random block censuses versus roadside surveys that have previously been used to estimate populations of the Loggerhead Shrike in Alberta (Telfer et al. 1989). Study Area and Methods The study area encompassed 23 600 km? in the mixed-grass ecoregion (Strong and Leggatt 1992) of east-central Alberta. Native grasslands in this area 445 446 are dominated by Needle-and-Thread (Stipa comata), Blue Grama (Bouteloua gracilis), and Northern (Agropyron dasystachyum) and Western (A. smithii) wheatgrass. Willows (Salix spp.) and Trembling Aspen (Populus tremuloides) are the dominant woody vegetation, although Thorny Buffaloberry (Sheperdia argentea) is locally common as single trees or clumps where sufficient moisture is present. Overall, the region receives 156 mm (median) of summer precipitation, which usually accumulates during fewer than 10 days. Cattle grazing and cereal crop production are the dominant land uses. The study area enclosed about 35% of the current range of the Loggerhead Shrike in the province, and contained the majority of confirmed breeding records found during the bird atlas project (Semenchuk 1992). Population estimates were derived from censuses conducted in 26, 6.4 x 6.4 km (41.5 km’) blocks that enclosed about 4.7% of the total study area (Figure 1). Most blocks were originally selected by Schmutz (1984, 1989) for estimating populations of Ferruginous Hawks (Buteo regalis). We opted to reuse established study blocks for several reasons. First, we had no prior knowledge of shrike popula- tions in these blocks or within the study area as a whole, and therefore had no reason to stratify our sampling effort based on local shrike densities. Second, the original selection criteria for these >. Coronation Ss Hanna Drumheller Ficure |. Study area and location of 41.5 km? blocks used to survey Loggerhead Shrike populations in south- eastern Alberta in 1993. THE CANADIAN FIELD-NATURALIST Vol. 110 blocks was not biased towards particular habitat fea- tures that might influence shrike distribution and abundance. Finally, there would be a higher proba- bility of sufficient time and money to repeat the study in future years if more than one species could be inventoried at a time. In areas where the study | area extended beyond the hawk study boundary, new survey blocks were randomly selected. All study plots were mapped on a 1:500 000 Alberta base map to facilitate locating the block on the ground. A 1:50 000 NTS map was used for more detailed infor- | mation while conducting the inventory. Following field trials of the census protocol in 1992, intensive sampling of the survey blocks occurred between 15 June and 7 July 1993. Observers usually worked independently, and spent ! between 2 and 30 (average 12) party hours in each block using a combination of truck, all-terrain vehi- cle and foot travel. To ensure that weather did not | influence detectability of birds, we avoided periods with rain, temperatures >25°C, or winds >25 km/h. All areas containing woody vegetation within the block were surveyed. Due to time constraints, we focused on locating adults, and did not actively search for nests. When detected, the position of birds was recorded on standardized data sheets and 1:50 000 maps. Shrikes observed near the edge of blocks were included only if they were within the block when first encountered. Independent observa- tions of shrikes within 400 m of another observation were considered to be either the same pair or the same individual, unless simultaneous observation of birds indicated otherwise. This distance would include the majority of the area utilized by a breed- ing pair of shrikes in this area (Collister 1994). Observation of a single shrike was regarded as an indicated pair. This was a reasonable assumption because repeat visits to nine territories (six in 1992, three in 1993) where single observations of shrikes were originally made, later revealed two adults and a nest with eggs or young in all cases. Furthermore, Collister (personal communication) concluded that occurrence of unpaired adult shrikes on his intensive study area in south-eastern Alberta was essentially | non-existent. Population size in the study area was estimated by calculating the mean number of shrike pairs per block, and extrapolating this value, with 95% confidence limits, to the study area as a whole. We determined habitat associations of Loggerhead | Shrikes within the study area on two spatial scales. First, we determined the relative proportions of pre- dominant land use in each of the census blocks by plotting different habitats on field data sheets of each | block. Specific land uses considered were the num- | ber of farmyards/shelterbelts, and the percentage of annual cultivation, native pasture and tame grass | (pasture or hayland). We used Spearman rank corre- | lations (Conover 1980) to test for associations | 1996 between the relative abundance of each habitat type in a block with the number of indicated breeding pairs of shrikes in that block. We also counted the number of clusters of shrubs and trees in a block, and divided woody plant density into LOW (< 50 clusters), MEDIUM (50 to 100), and HIGH (> 100). A cluster ranged from a few trees or shrubs to many trees covering up to 2 ha. The effect of shrub density on the number of breeding pairs was analyzed using Kruskall-Wallis tests and simultaneous test proce- dures (Conover 1980). Second, we tabulated the fre- quency of occurrence of five different land uses (annual cultivation, native pasture, tame grass, road or railway rights-of-way, and farmyards/shelterbelts) within 400 m of points where shrikes were first encountered. This radius was chosen because it is the maximum distance that shrikes routinely travel from their nest sites while foraging (Collister 1994). We then randomly selected an equal number of 400-m _ radius circles from the study blocks using a random number generator. The frequency of occurrence of each habitat type, as recorded on field sheets, was then compared between observed and random sites using chi-square tests. We also determined the total number of these habitat types occurring in each 400 m circles, and compared values between observed and random circles using Mann-Whitney U-tests. To determine the relative efficiency of roadside versus block surveys in censusing Loggerhead Shrikes, we classified the positions of shrike encoun- ters in relation to the distance from the nearest road as follows: NEAR, 0-200 m; MEDIUM, 201-400 m; and FAR, > 400 m. Chi-square tests were used to test for differences in frequency of occurrence of shrikes across distance. For calculating expected val- ues, we estimated that 17.8% of the study area occurred in the NEAR category, 17.6% occurred in the MEDIUM class, and 64.6% occurred > 400 m from roads. Results Loggerhead Shrikes were encountered in 21 of 26 (80.8 %) study blocks during 1993. A total of 113 breeding pairs was observed during the census, for a mean density of 4.35 + 0.75 (SE) pairs per 41.5 km? block (range 0 to 14). There were 569.3 blocks such in the 23 600 km* study area, for an overall popula- tion of 2477 pairs and a 95% confidence interval of 1588 to 3365 pairs. Native pasture was the dominant land use in the study blocks (49.6 + 6.3% of area), followed by annual cultivation (43.4 + 6.6%) and tame grass (5.0 + 6.3%). The number of farmyards/shelterbelts in a block ranged from 0 to 14 (mean = 6.08 + 0.83). There were no correlations between the percent of any of the above major land uses with the number of breeding shrike pairs observed in a block (all rt, < 0.23, all p > 0.27). However, there was a rela- BJORGE AND PRESCOTT: LOGGERHEAD SHRIKE IN SOUTHEASTERN ALBERTA 447 TABLE |. Comparison of major land-uses within a 400-m radius of Loggerhead Shrike observations (“occupied”, n= 113), and in 113 randomly-selected circles within study blocks. Frequency of Occurrence (%) Land Use Occupied Random x? Tame Grass Sy) LS 0.9 Native Pasture 61.9 63.7 0.1 Cultivation Wilce/ 60.2 353 Rights-of-Way 63.7 SD Diaries Farmyards/Shelterbelts 45.1 12.4 29567 *P < 0.0001 tionship between the relative density of woody vege- tation and the number of breeding pairs (T = 7.0, df= 2, p< 0.04), with significant differences in breeding pairs occurring between the LOW (2.33 + 0.66 pairs, n= 12) and HIGH (6.6 + 1.6 pairs, n = 7) shrub den- sities (p < 0.05). The number of shrike pairs observed in blocks with MEDIUM shrub density (5.6 + 1.6 pairs, n = 7) was not significantly different from either the LOW or HIGH classes (p > 0.05). Land-use characteristics of occupied and random 400-m plots are given in Table 1. The overall distrib- ution of habitats in the two classes was different (x?= 19.3, df = 4, p < 0.001), with the frequency of occurrence of rights-of way and farmyards/shelter- belts being significantly higher in occupied sites (both x? >21.7, df = 2, p < 0.0001). Shrikes also pre- ferred sites that contained several habitat types with- in a 400-m radius. Occupied sites contained 2.6 + 0.1 habitat types, whereas random sites contained only 1.8 + 0.1 types (Mann-Whitney U-test, T = 6.2, p < 0.0001). Only 13 of 113 observed pairs (11.5%) occurred in areas that contained a single major habi- tat type, compared to 38.9 % of random locations. The frequency of occurrence of shrikes was dependent on the distance from roads (x? = 30.0, df = 2, p < 0.0001), with the majority of pairs (52.2%) being detected within 200 m. An additional 6.2% were found between 200 and 400 m, whereas 41.6% were found > 400 m from roads, and unlikely to be routinely detected by roadside surveys. Discussion Our major objective was to estimate the popula- tion size of Loggerhead Shrikes within the core breeding area of Alberta using a random sampling technique. Our results show that shrike populations are substantially higher than previously estimated (350-1000 pairs; Telfer et al. 1989; Wershler 1989*). In our study area alone, we estimate with 95% confi- dence that between 1588 and 3365 pairs of shrikes (mean of 2477 pairs) occurred during the study peri- od. This might be a conservative estimate, because we assumed that birds observed within 400 m of each other were members of the same pair. However, 448 adjacent nests of Loggerhead Shrikes in Alberta have occasionally been observed less than 80 m apart (Collister 1994). We did not actively search for nests during our survey, so the frequency of nests closer than 400 m to each other, and the resulting impact on estimated pair densities in a survey block is unknown. Although the study area we sampled likely contains the highest densities of shrikes in Alberta, it encompassed only about 35% of the species’ provincial breeding range. Thus, the true provincial population might reasonably be estimated to be double the size of the population that we calcu- lated for our study area (i.e., close to 5000 pairs). Previous population studies of the Loggerhead Shrike in Alberta may have under-estimated popula- tion sizes for several reasons. First, complete counts of known breeding areas (e.g., Wershler 1989*) cover a fairly small geographic range where shrikes are most likely to be encountered, and do not sample vast areas of the province where shrikes occur in rel- atively low densities. Roadside surveys (e.g., Telfer et al. 1989) cover extensive geographical areas, and tend not to be biased towards where shrikes are expected to occur. However, our study indicated that only 52.2% of shrikes located in randomly-selected blocks were found within detection distance of a road. Furthermore, birds in our study area were typi- cally associated with shelterbelts and clumps of shrubs. Even though shrikes often perch conspicu- ously on vegetation, adequate censusing of these habitats was achieved only by walking around or through the potential nesting cover. Vehicle surveys conducted at 50-70 km/hr (Telfer et al. 1989), and on only one side of shrub or tree patches, would over- look many resident shrikes that we recorded via intensive ground searches. In fact, incidental obser- vations conducted during previous roadside surveys on the prairies revealed many sample blocks where shrikes were overlooked by observers in vehicles (Telfer et al. 1989). We made no systematic effort to determine the percentage of birds known to occur near roads that were detected from a moving vehicle. However, of the 27 pairs of shrikes observed within 200 m of roads by the senior author, 17 (63%) were initially detected while driving. This value, coupled with the proportion of shrikes that were found near roads, suggests that only 32.9% of the population would be detected from an extensive roadside survey in our study area. Despite this, roadside surveys may be the most efficient means of surveying broad geo- graphical areas, but only if appropriate corrections for detectibility are applied. Such correction factors likely vary among habitat types, and would need to be cal- culated for each geographical area being surveyed. Loggerhead Shrikes are heterogeneously distrib- uted over the study area, with the number of indicat- ed breeding pairs in a 41.5 km? block ranging from 0 to 14. Pair numbers were highest in blocks with rela- THE CANADIAN FIELD-NATURALIST Vol. 110 tively high concentrations of native trees and shrubs, and in areas where farmyards/shelterbelts and rights- of-way were more common than expected by chance. The association with woody vegetation is not surprising, given the nesting requirements of shrikes, and the relative scarcity of trees and shrubs in many parts of the species’ range in Alberta. Accordingly, we concur with Telfer (1992) who rec- ommended planting appropriate shrubs for nesting and perching as a means of increasing local shrike populations. Our data indicate that shrubs would best be planted in areas with considerable vegetation diversity, such as on the interface of tame and native pastures. Road allowances, which typically separate different land uses, and provide power lines and fences for perching (Bohall-Wood 1987) and tall grasses for foraging (Prescott and Collister 1993; Chavez-Ramirez et al. 1994) would also be suitable for shrub and tree plantings. Given the tendency of shrikes to inhabit farmyards, we also suggest that programs designed to preserve existing tree growth in abandoned farmyards be an important component of management plans for Loggerhead Shrikes on the Canadian prairies. Acknowledgments We thank J. Allen, C. Calihoo, D. Collister, K. Froggatt, E. Hofman, S. Johnston, H. Kiliaan, L. Lester, Gy Niemanse Is\\Pocock; "Ji, Potter..k Podlubney, C. Tremblay and S. Visser for capable field assistance. D. Collister provided heipful input concerning various aspects of the study protocol. L. Beattie drafted the figure. J. Schmutz kindly provid- ed locations of randomly-selected census blocks. Land-owners are thanked for providing access to their properties, and for their interest. K. DeSmet, A. Erskine, E. Ewaschuk and two anonymous reviewers commented on earlier drafts of this paper. Document Cited (marked in text with *) Wershler, R. M. 1989. Nesting habitat and abundance of the Loggerhead Shrike (Lanius ludovicianus) in two key areas of southern Alberta. Unpublished report to the | World Wildlife Fund Canada and Alberta Fish and Wildlife. 27 pages. Literature Cited Bohall-Wood, P. 1987. Abundance, habitat use, and perch use of Loggerhead Shrikes in north-central Florida. Wilson Bulletin 99: 82-86. Cadman, M. D. 1985. Status report on the Loggerhead | Shrike (Lanius ludovicianus) in Canada. Commitee on | the Status of Endangered Wildlife in Canada, Ottawa. 86 » pages. Chavez-Ramirez, F., D..E. Gawlik, F. G. Prieto, and R. D. Slack. 1994. Effects of habitat structure on patch use | by Loggerhead Shrikes wintering in a natural grassland. | Condor 96: 228-231. Collister, D. M. 1994. Breeding ecology and habitat preservation of the Loggerhead Shrike (Lanius Iudovi- 1996 cianus) in southeastern Alberta. Unpublished M.Sc the- sis, University of Calgary, Calgary, Alberta. 161 pages. Conover, W. J. 1980. Practical nonparametric statistics. John Wiley and Sons, New York. 493 pages. Hands, H. M., R. D. Drobney, and M. R. Ryan. 1989. Status of the Loggerhead Shrike in the northcentral United States. Missouri Cooperative Fish and Wildlife Research Unit, Columbia, Missouri. 15 pages. Johns, B., E. Telfer, M. Cadman, D. Bird, R. Bjorge, K. DeSmet, W. Harris, D. Hjertas, P. Laporte, and R. Pittiway. 1994. National recovery plan for the Loggerhead Shrike. Recovery of Nationally Endangered Wildlife Committee Report number 7, Ottawa, Ontario. 32 pages. Leighton, F. A., J. M. Gerrard, D. W. A. Whitfield, and W. J. Maher. 1979. An aerial census of Bald Eagles in Saskatchewan. Journal of Wildlife Management 43: 61-69. Morrison, M. L. 1981. Population trends of the Loggerhead Shrike in the United States. American Birds 35: 754-757. Prescott, D. R. C., and D. M. Collister. 1993. Characteristics of occupied and unoccupied Loggerhead Shrike territories in southeastern Alberta. Journal of Wildlife Management 57: 346-352. Price, J., S. Droege, and A. Price. 1995. The summer atlas of North American birds. Academic Press, New York. 364 pages. Robbins, C. S., D. Bystrak, and P. H. Geissler. 1986. The breeding bird survey: its first fifteen years, 1965-1979. United States Fish and Wildlife Service Resource Publication number 157. 196 pages. BJORGE AND PRESCOTT: LOGGERHEAD SHRIKE IN SOUTHEASTERN ALBERTA 449 Robert, M., and P. Laporte. 1991. History and current sta- tus of the Loggerhead Shrike in Quebec. Canadian Wildlife Service Progress Notes number 196. 6 pages. Schmutz, J. K. 1984. Ferruginous and Swainson’s Hawk abundance and distribution in relation to land use in southeastern Alberta. Journal of Wildlife Management 48: 1180-1187. Schmutz, J. K. 1989. Hawk occupancy of disturbed grass- lands in relation to models of habitat selection. Condor 91: 362-371. Semenchuk, G. P. Editor. 1992 . The atlas of breeding birds of Alberta. Federation of Alberta Naturalists, Edmonton, Alberta. 390 pages. Smith, E. L., and K. C. Kruse. 1992. The relationship between land use and the distribution and abundance of Loggerhead Shrikes in south-central Illinois. Journal of Field Ornithology 63: 420-427. Strong, W. L., and K. R. Leggatt. 1992. Ecoregions of Alberta. Alberta Forestry, Lands and Wildlife, Edmonton. 59 pages. Tate, J. R., Jr. 1986. The Blue List for 1986. American Birds 40: 227-236. Telfer, E. S., C. Adam, K. DeSmet, and R. Wershler. 1989. Status and distribution of the Loggerhead Shrike in western Canada. Canadian Wildlife Service Progress Notes number 184. 4 pages. Telfer, E. S. 1992. Habitat changes as a factor in the decline of the western Canadian Loggerhead Shrike (Lanius ludovicianus) population. Canadian Field- Naturalist 106: 321-326. Received 30 October 1995 Accepted 14 February 1996 Four Records of the Chestnut Lamprey, I[chthyomyzon castaneus, New to Ontario CLAUDE B. RENAuD!, SANDRA C. RIBEY? and FRANCOIS CHAPLEAU2 ‘Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 ?Ottawa-Carleton Institute of Biology, Department of Biology, University of Ottawa, P.O. Box 450, Station A, Ottawa, Ontario KIN 6N5 Renaud, Claude B., Sandra C. Ribey, and Francois Chapleau. 1996. Four records of the Chestnut Lamprey, Jchthyomyzon castaneus, new to Ontario. Canadian Field-Naturalist 110(3): 450-453. We report on the widespread, although rare, presence of the parasitic Chestnut Lamprey (/chthyomyzon castaneus) in Ontario based on four collections and five specimens. This report increases the number of lamprey species for the province to five, three of which are parasitic. The Chestnut Lamprey is sympatric with three of the four other Ontario lamprey species. Key Words: Chestnut Lamprey, /chthyomyzon castaneus, Ontario, Petromyzontidae. We report on the widespread although rare occurrence of the parasitic Chestnut lamprey, Ichthyomyzon castaneus Girard, 1858, in Ontario (Figures | and 2). During the course of routine cura- torial work, three collections of lampreys in The Canadian Museum of Nature (CMN) were found to contain four transformed specimens of this species. A fourth collection of a single immature transformed specimen (149.0 mm TL) was made on the 3 August 1994 in the St. Lawrence River off the northeast side of eastern Colquhoun Island (45°01'N 74°38'W) near Cornwall, Glengarry County, and has since been deposited in the Canadian Museum of Nature (CMN) [formerly National Museum of Canada, NMC] (alphanumeric catalogue code NMC95-7). The specimen was collected in a trapnet by one of us (SCR) in swift, very clear water over a rocky and weedy substrate. The first collection (NMC 86-856) was made on the 18 May (year unrecorded) in the Mad River (44°24'N 79°54'W), Simcoe County, Georgian Bay watershed, Lake Huron basin. One transformed male 90°W 85°W 80°W FiGurE |. Geographic distribution of [chthyomyzon casta- neus in Ontario. Chestnut Lamprey in spawning condition (224.5 mm TL), two transformed Northern Brook Lampreys (Ichthyomyzon fossor), three transformed Silver Lampreys (J. unicuspis) and 15 transformed Sea Lampreys (Petromyzon marinus) were collected in a FIGURE 2. Side-view of two transformed Ichthyomyzon castaneus from Lake of the Woods, Ontario (NMC70- 223). The specimen above measures 164.5 mm in total length and the one below 144.0. 450 1996 RENAUD, RIBEY, AND CHAPLEAU: CHESTNUT LAMPREY 451 FiGurE 3. Drawings of the oral disc in the two species of Ichthyomyzon. Left side: I. castaneus, Sac River, Missouri, 20-23 May 1951 (closed disc measures 15.0 mm). Note the seven bicuspid endolaterals; Right side: [. unicuspis, Manitoba, no other collection data (closed disc measures 20.0 mm). Note the absence of bicuspid endolaterals. Modified from Vladykov and Kott (1980) with permission from the Canadian Journal of Fisheries and Aquatic Sciences. large trap by M. Milson. The Mad River, therefore, harbours three parasitic lamprey species. The second collection (NMC 83-918) was made on the 6 July 1966 in the Chippewa River (46°58'N 84°21'W), Algoma District, Batchawana Bay water- shed, Lake Superior basin. One spent transformed female Chestnut Lamprey (207.5 mm TL) and one transformed Sea Lamprey were collected by elec- trofishing by members of the Sea Lamprey Control Centre in Sault-Ste-Marie. The third collection (NMC 70-223) was made in the summer of 1970 in Lake of the Woods, south- west of Burton Island (48°59'N 94°27'W), Rainy River District. Two immature transformed Chestnut Lampreys (144.0-164.5 mm TL) and 64 transformed Silver Lampreys were collected by Pat O'Connor, a commercial fisherman. The specimens were collect- ed in still, clear, and brown-tinged water over a stony substrate, at a maximum depth of 20 m. Specimens of Ichthyomyzon were identified to species using Lanteigne (1981). Teeth nomenclature follows Vladykov and Follett (1967). According to Lanteigne (1981), the Chestnut Lamprey is distin- guished from the Silver Lamprey (Figure 3), another parasitic species with which it could be confused by the presence of 2-8 (usually 4-8) bicuspid endolater- als (n = 36; X = 5.9) as opposed to 0-2 (usually 0) bicuspid endolaterals (n = 43; X = 0.06). Four of the five transformed specimens in this study possessed either four or five bicuspid endolaterals. The fifth transformed specimen (NMC 83-918) possessed at least three bicuspid endolaterals; the condition of one of its endolaterals could not be ascertained because it was missing. On the other hand, the 67 transformed specimens of the Silver Lamprey in this study possessed only 0-2 bicuspid endolaterals 300 N Y/A \. unicuspis [_] |. castaneus 290 280 50 40 30 No. of specimens 20 3 4 5 Cetew into Oal© No. of Bicuspid Endolaterals FIGURE 4. Histogram of frequencies of the number of bicuspid endolaterals in Ichthyomyzon unicuspis and I. castaneus based on the data contained in Table 1 minus the two specimens for which the condition in one tooth could not be ascertained. 452 TABLE |. Comparison of various dentitional counts in Ichthyomyzon unicuspis versus I. castaneus. Number of Lateral Rows Number of Anterior Rows Number of Bicuspid Endolaterals Source foo) NN Species Hubbs and Trautman (1937) Present Study 6.61 6.58 8.54 6.10 219 234 40 8 0.03 0.30 245 I. unicuspis 54 Hubbs and Trautman (1937) Present Study 3 19 75 AN I. castaneus NN nN N *One endolateral tooth missing for which the number of cusps could not be ascertained. THE CANADIAN FIELD-NATURALIST Vol. 110 (Table 1). A histogram combining the data in Hubbs | and Trautman (1937) and this study with regards to the number of bicuspid endolaterals clearly shows | that /. unicuspis has a strong mode at 0 and I. casta- neus a strong mode at 6 (Figure 4). Examination of | the number of anterior and lateral rows, two charac- ters suggested by Hubbs and Trautman (1937) of | lesser diagnostic value, were not found useful in this | study (Table 1). These latter two counts included the » marginal teeth as in Hubbs and Trautman (1937). In Canada, the Chestnut Lamprey has been report- | ed previously only from Saskatchewan and Manitoba (Lanteigne 1992). It is not listed in Mandrak and Crossman (1992a), and is not even included in their list of potential fish invaders of Ontario. The distribution of the Chestnut Lamprey in | Ontario matches a pattern that Mandrak and) Crossman (1992b) broadly term “distribution limited | to the Great Lakes and Nelson River watersheds”. Its. present-day distribution (Lanteigne 1992) teads us to) suggest that its presence in Lake of the W: ie is the | result of a post-Wisconsinan reinvasion from the | Missourian and/or Mississippian Refugium while its | presence in the Chippewa, Mad and St. Lawrence | rivers is the result of a reinvasion of the species from | the Mississippian Refugium. Lanteigne (1992) ) reports that the Chestnut Lamprey presently occurs | in both the Upper and Lower Missouri as well as the | Upper and Lower Mississippi River basins. According to Lanteigne (1981), the Chestnut | Lamprey and the Silver Lamprey occur sympatrical- ly in the Hudson Bay, Upper Mississippi River and Ohio River basins. The basins of lakes Superior and | Huron and of the St. Lawrence River can now be added to this list. Ontario material of transformed individuals of J. unicuspis held at the Canadian Museum of Nature consist of 36 collections totalling 153 specimens. | Ontario material at the Royal Ontario Museum amounts to 48 collections and 84 specimens (Erling Holm, personal communication). On this basis, the number of Ontario specimens of /. castaneus report- ed here represents only 2.10% of the Ontario speci- mens of /. unicuspis and is an indication of the for- mer's relative rarity. The Committee on the Status of Endangered Wildlife in Canada designated the species I. castaneus as vulnerable in 1991 (Lanteigne 1992), and this designation is still appropriate, , despite its broader distribution recorded here, , because of its apparent rarity. | Acknowledgments The specimen caught in the St. Lawrence River was part of a broad fish survey of the river, near Cornwall, sponsored by a tri-council grant (Federal Green Plan, Government of Canada) obtained by the Institute for Research on Environment and Economy | of the University of Ottawa (UO). Lara L. Ridgway, | ; | | | ' 1996 Lee Willard, Martin D. Lemay and David Bajurny participated in the fieldwork. We thank Sylvie Laframboise, CMN, for bringing an odd Chippewa River lamprey to our attention; André Dumouchel for producing Figure 2; the Audiovisual and Reprography Services of UO for the layout of Figure 3; and Alison Murray, CMN, for producing Figure 4. We also thank Erling Holm for permission to use information on material from the Royal Ontario Museum and the four reviewers for forcing us to take a second look at our data. Literature Cited Hubbs, C. L., and M. B. Trautman. 1937. A revision of the lamprey genus [chthyomyzon. Miscellaneous Publications of the Museum of Zoology, University of Michigan (35): 7-109 + 2 plates. Lanteigne, J. 1981. The taxonomy and distribution of the North American lamprey genus I[chthyomyzon. unpub- lished M.Sc. thesis, University of Ottawa, Ottawa, ida. xi + 150 pages. RENAUD, RIBEY, AND CHAPLEAU: CHESTNUT LAMPREY 453 Lanteigne, J. 1992. Status of the Chestnut Lamprey, Ichthyomyzon castaneus, in Canada. Canadian Field- Naturalist 106(2): 14-18. Mandrak, N. E., and E. J. Crossman. 1992a. A checklist of Ontario freshwater fishes annotated with distribution maps. Royal Ontario Museum, Toronto. v + 176 pages. Mandrak, N. E., and E. J. Crossman. 1992b. Postglacial dispersal of freshwater fishes into Ontario. Canadian Journal of Zoology 70(11): 2247-2259. Vladykov, V. D., and W. I. Follett. 1967. The teeth of lampreys (Petromyzonidae): their terminology and use in a key to the Holarctic genera. Journal of the Fisheries Research Board of Canada 24(5): 1067-1075. Vladykov, V. D., and E. Kott. 1980. Description and key to metamorphosed specimens and ammocoetes of Petromyzonidae found in the Great Lakes region. Canadian Journal of Fisheries and Aquatic Sciences 37(11): 1616-1625. Received 17 October 1995 Accepted 9 April 1996 Rare and Endangered Fishes and Marine Mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: X ROBERT R. CAMPBELL 666 Bayview, RR1 Woodlawn, Ontario KOA 3M0 Campbell, Robert R. Editor. 1996. Rare and endangered fish and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: X. Canadian Field-Naturalist 110(3): 454461. Nine status reports representing the 1994 fish and marine mammal status assignments by the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) have been prepared for publication and two species previously assigned status reviewed. Committee (COSEWIC) and Subcommittee (Fish and Marine Mammals) activities are briefly discussed as are the implications of a British Columbia Court of Appeal ruling rending habitat sections of the Fisheries Act inoper- | able for non-commercial or non-game species. Tabular lists of fishes and marine mammal species assigned status to April 1994 and species currently under consideration are presented and new category definitions are outlined. Neuf rapports de statut relativement aux poissons et aux mammiferes marins auxquel ont été attribué un statut en 1994 ont | été préparés pour publication et on a mis a jour les rapports de deux espéces qui possédaient un statut. Les activitités du | Comité (CSEMDC) et du souscomité (des poissons et des mammifeéres marins) sont bri¢vement discutées, de méme que les implications d’une décision prise par la cour d'appel de la C.-B., qui rend inopérantes les sections de |’habitat de la Loi sur La Péche pour les especes non-commerciales ou non-sportives. Des listes sous forme tabulaire des espéces de poisson et de mammiferes marins qui ont recu un statut en date d’avril 1994, des especes toujours a l'étude et les nouvelles définitions de catégorie sont présentées. Key Words: Rare and Endangered species, fish, marine mammals, COSEWIC. Previous submissions (Campbell 1984 through 1993), have stated and repeated the intent of the Subcommittee on Fish and Marine Mammals to pub- lish the status reports (on those species of fish and marine mammals) which the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) has reviewed, approved and used as a basis of assigning status to species in jeopardy in Canada. The nine reports presented in this issue are the fish and marine mammal component of those species assigned status in 1994 (see Table 1). Progress COSEWIC has undertaken to make available to all Canadians supporting information on each species classified (see Cook and Muir 1984). The Fish and Marine Mammal Subcommittee used The Canadian Field-Naturalist as one step in achieving the goal. A series of reports have appeared from 1984 through 1993 [see The Canadian Field- Naturalist 98(1): 63-133; 99(3): 404-450; 101(2): 165-309; 102(1): 81-176 and 102(2): 270-398; 103(2): 147-220; 104(1): 1-145; 105(2): 151-250; 106(1): 1-72; 107(4): 395-546]. Commencing in April 1994 the Canadian Wildlife Federation agreed to support COSEWIC report preparation through a system of matching grants and some $40 000 ($10 000 contributed by the Canadian Wildlife Service) was available to support the vari- ous subcommittees in 1993 through 1994. As of April 1994, COSEWIC has reviewed the status of 85 fish species, one marine invertebrate, and 38 marine mammals (Table 1). Of the 124 species considered six were found to be indetermi- nate (five fish, one marine mammal), 52 (21 fish, 30 marine mammals, one marine invertebrate) were found not to require status designation and another 43 (38 fish, five marine mammals) were designated as vulnerable, mainly due to natural rarity, leaving 23 species of immediate concern which are of inter- est to the RENEW (Recovery of Nationally Endangered Wildlife) organization which was estab- lished in 1990 to oversee the development of recov- ery teams and plans for such species listed by COSEWIC (CWS 1993). There are currently 26 status reports on fish and 10 on marine mammal species under review or in preparation (Table 2).* As well, some 54 additional species of fish, two of marine mammals and 12 | marine invertebrates have been identified as being worthy of consideration (see Campbell 1993: Table 3). Many may be found to not require status desig- nation, but the process serves to bring together the information necessary to make a formal determina- *Editor’s note: Some of these were subsequently presented | at the 1996 Annual Meeting of COSEWIC. 454 1996 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS TABLE |. Fish and Marine Mammal Species with Assigned COSEWIC Status to 14 April 1994. Species Fish Lake Sturgeon Bloater Blueback Herring Striped Shiner Redfin Shiner Hornyhead Chub River Chub Ghost Shiner Blackchin Shiner Cutlips Minnow Leopard Dace Mountain Sucker Golden Redhorse Least Darter Tesselated Darter River Darter Green Sunfish Longear Sunfish Spoonhead Sculpin Brook Silverside Y-Prickleback Darktail Lamprey Bering Cisco Flathead Catfish Northern Madtom Pixie Poacher Lake Lamprey‘ Chestnut Lamprey Northern Brook Lamprey Shortnose Sturgeon Green Sturgeon White Sturgeon Spotted Gar Spring Cisco* Squanga Whitefish‘ Kiyi Pacific Sardine Redside Dace Silver Chub Pugnose Shiner Bigmouth Shiner Silver Shiner Rosyface Shiner Pugnose Minnow Speckled Dace Umatilla Dace Central Stoneroller Banded Killifish — Newfoundland Blackstripe Topminnow Lake Chubsucker Bigmouth Buffalo Black Buffalo Spotted Sucker River Redhorse Greenside Darter Brindled Madtom Redbreast Sunfish Orangespotted Sunfish Warmouth Fourhorn Sculpin — Arctic Islands Scientific Name Acipenser fulvescens Coregonus hoyi Alosa aestivalis Luxilus chrysocephalus Lythrurus umbratilis Nocomis biguttatus Nocomis micropogon Notropis buchanani Notropis heterodon Exoglossum maxillingua Rhinichthys falcatus Catostomus platyrhynchus Moxostoma erythrurum Etheostoma microperca Etheostoma olmstedi Percina shumardi Lepomis cyanellus Lepomis megalotis Cottus ricei Labidesthes sicculus Allolumpenus hypochromus Lethenteron alaskense Coregonus laurettae Pylodictis olivaris Noturus stigmosus Occella impi Lampetra macrostoma Ichthyomyzon castaneus Ichthyomyzon fossor Acipenser brevirostrum Acipenser medirostris Acipenser transmontanus Lepisosteus oculatus Coregonus sp. Coregonus sp. Coregonus kiyi Sardinops sagax Clinostomus elongatus Macrhybopsis storeriana Notropis anogenus Notropis dorsalis Notropis photogenis Notropis rubellus Opsopoeodus emiliae Rhinichthys osculus Rhinichthys umatilla Campostoma anomalum Fundulus diaphanus Fundulus notatus Erimyzon sucetta Ictiobus cyprinellus Ictiobus niger Minytrema melanops Moxostoma carinatum Etheostoma blennioides Notorus miurus Lepomis auritus Lepomis humilis Lepomis gulosus Myoxocephalus quadricornis Status RANSDR? RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RAISIFSD> RAISIFSD RAISIFSD RAISIFSD RAISIFSD Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable Date Assigned April 1986 April 1988 April 1980 April 1993 April 1988 April 1988 April 1988 April 1993 April 1994 April 1994 April 1990 April 1991 April 1989 April 1989 April 1993 April 1989 April 1987 April 1987 April 1989 April 1989 April 1991 April 1990 April 1990 April 1993 April 1993 April 1991 April 1986 April 1991 April 1991 April 1980 April 1987 April 1990 April 1983™ April 1992 April 1988 April 1987 April 1987 April 1987 April 1985 April 1985 April 1985 April 1983° April 1994 April 1985 April 1980! April 1988 April 1985 April 1989 April 1985 April 1994 April 1989 April 1989 April 1983™ April 1983° April 1990 April 1985 April 1989 April 1989 April 1994 April 1989 (Continued) 455 456 TABLE |. Continued. Species Giant Stickleback‘ Unarmoured Stickleback* Blackline Prickleback Bering Wolffish Lake Simcoe Whitefish* Blackfin Cisco Shortnose Cisco Shortjaw Cisco Black Redhorse Copper Redhorse* Eastern Sand Darter Channel Darter Margined Madtom Enos Lake Stickleback‘ Shorthead Sculpin Deepwater Sculpin — Great Lakes Acadian Whitefish® Aurora Trout® Salish Sucker Paddlefish Gravel Chub Longjaw Cisco Deepwater Cisco Banff Longnose Dace* Blue Walleye Marine Molluscs Northern Abalone Marine Mammals Sea Otter Sea Mink Hooded Seal Bearded Seal Steller Sea Lion Northern Elephant Seal Ringed Seal California Sea Lion Atlantic Walrus Eastern Arctic Northwest Atlantic Baird's Beaked Whale Beluga Beaufort Sea THE CANADIAN FIELD-NATURALIST Scientific Name Gasterosteus sp. Gasterosteus sp. Acantholumpenus mackayi Anarhichas orientalis Coregonus clupeaformis ssp. Coregonus nigripinnis Coregonus reighardi Coregonus zenithicus Moxostoma duquesnei Moxostoma hubbsi Ammocrypta pellucida Percina copelandi Noturus insignis Gasterosteus sp. Cottus confusus Myoxocephalus thompsoni Coregonus huntsmani Salvelinus fontinalis timagamiensis Catostomus sp. Polyodon spathula Erimystax x-punctata Coregonus alpenae Coregonus johannae Rhinichthys cataractae smithi Stizostedion vitreum glaucum Haliotis kamtschatkana Enhydra lutris Mustela macrodon Cystophora cristata Erignathus barbatus Eumetopias jubatus Mirounga angustirostris Phoca hispida Zalophus californianus Odobenus rosmarus rosmarus Berardius bairdi Delphinapterus leucas Western and Southern Hudson Bay High Arctic Eastern Hudson Bay St. Lawrence River S.E. Baffin Island Ungava Bay Common Dolphin Grey Whale Northeast Pacific Northwest Atlantic Risso's Dolphin Short-finned Pilot Whale Longfinned Pilot Whale Northern Bottlenose Whale Pygmy Sperm Whale Atlantic White-sided Dolphin Pacific White-sided Dolphin Delphinus delphis Eschrichtius robustus Grampus griseus Globicephala macrohynchus Globicephela malaena Hyperoodon ampullatus Kogia breviceps Lagenorhynchus acutus Lagenorhynchus obliquidens Status Vulnerable Vulnerable Vulnerable Vulnerable Threatened Threatened Threatened Threatened Threatened Threatened Threatened Threatened Threatened Threatened Threatened Threatened Endangered Endangered Endangered Extirpated Extirpated Extinct Extinct Extinct Extinct N/A® Endangered Extinct RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR Extirpated RANSDR RANSDR RANSDR Vulnerable Threatened Endangered Endangered Endangered RANSDR RANSDR Extirpated RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR RANSDR Vol. 110 Date Assigned April 1980 April 1983 April 1989 April 1989 April 1987 April 1988 April 1987 April 1987 April 1988 April 1987 April 1994 April 1993 April 1989 April 1988 November 1983 April 1987 April 1983 April 1987 April 1986 April 1987 April 1987 April 1988 April 1988 April 1987 April 1985 April 1988 May 1978! April 1985 April 1986 April 1994 April 1987 April 1986 April 1989 April 1987 April 1987 April 1987 April 1992 April 1986 April 1993 April 1992 April 1988 April 1983 April 1990 April 1988 April 1991 April 1987 April 1987 April 1990 April 1993 April 1994 April 1993 April 1994 April 1991 April 1990 | (Continued) | 1996 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS 457 TABLE 1. Concluded. Species Scientific Name Status Date Assigned Northern Right Whale Dolphin Lissodelphis borealis RANSDR April 1990 Hubbs' Beaked Whale Mesoplodon carlhubbsi RANSDR April 1989 Blainville's Beaked Whale Mesoplodon densirostris RANSDR April 1989 True's Beaked Whale Mesoplodon mirus RANSDR April 1989 Stejneger's Beaked Whale Mesoplodon stejnegeri RANSDR April 1989 Narwhal Monodon monoceros RANSDR April 1986° Dall's Porpoise Phocoenoides dalli RANSDR April 1989 False Killer Whale Pseudorca crassidens RANSDR April 1990 Striped Dolphin Stenella coeruleoalba RANSDR April 1993 Bottlenose Dolphin Tursiops truncatus RANSDR April 1993 Cuvier's Beaked Whale Ziphius cavirostris RANSDR April 1990 Blue Whale Balaenoptera musculus Vulnerable April 1983! Fin Whale Balaenoptera physalus Vulnerable April 1987 Sowerby's Beaked Whale Mesoplodon bidens Vulnerable April 1989 Harbour Porpoise Phocoena phonoeca Northwest Pacific RAISIFSD April 1991 Northwest Atlantic Threatened April 1990 Humpback Whale Megaptera novaeangliae Northwest Atlantic Vulnerable April 1985 Northeast Pacific Threatened April 1982! Bowhead Whale Balaena mysticetus Endangered April 1980! Right Whale Eubalaena glacialis Endangered April 1980* * RANSDR -— Use of NIAC (Not in Any Category) dropped in 1988 and subsequently converted. RANSDR is not a category = Report Accepted No Status Designation Required. > RAISIFSD — the use of a new list “Report Accepted Insufficient Scientific Information For Status Designation” was approved at the 1990 General Meeting. ° Endemic to Canada 4 Vulnerable “Rare” category changed to “Vulnerable” in 1988. Dates Assigned of 1988 or earlier indicate date of original Rare status assignment. These were subsequently converted to Vulnerable at the 1990 General Meeting based on the advice of the Fish and Marine Mammal Subcommittee. * Updated April 1987 — no status change. Updated April 1984 — no status change. & Updated April 1987 — previous status of “Endangered” assigned April 1985. » N/A —Status Not Assigned. COSEWIC has no mandate for invertebrates. Report accepted and recommended RANSDR Nn tatus agreed to, but not assigned. pdated April 1986 — no status change. J K Updated April 1985 and April 1990 — no status change. ™ Updated April 1994 — no status change. tion that they are not presently at risk. The Subcommittee will, as opportunity allows, attempt to document the status of all of these species and assign designations. In addition to soliciting further status reports on species of concern, the Subcommittee continues to obtain updates on the status of selected species as new information becomes available, or in the 10- year review process initiated in 1993 (see Campbell 1993: Table 3) for those species not updated since their initial status assignment. In that regard, updated reports on the Spotted Gar and Spotted Sucker were considered at the April 1994 General Meeting and the original (Table 1) vulnerable status reconfirmed for each. These reports are not included in the following series as the only differences from the original reports U Updated April 1985 — North Atlantic stock downlisted to “Vulnerable”. U (Spotted Gar —Parker et al. 1980; Parker and McKee 1984a: Spotted Sucker — Parker et al. 1980; Parker and McKee 1984b) were the inclusion of new records. The only Spotted Gar record since 1975 that could be considered bona fide is the 1986 collection of two specimens from Rondeau Bay (ROM 51555) [Campbell 1994a]. The only new information on the Spotted Sucker in Canada since that reported by Parker et al. (1980) and Parker and McKee (1984b) are the 10 additional specimens obtained (see Campbell 1994b: Table 1) from Lake Erie and Lake St. Clair tributaries. Protection An important 1983 court ruling which impacts on endangered fishes in Canada and COSEWIC reports has been previously overlooked. In the past the 458 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 2. Fish and Marine Mammal Species for which Status Reports are in preparation, or under review — to 14 April 1994, Species Scientific Name Fish Atlantic Sturgeon Lake Sturgeon? Red (Arctic) Char Bull Trout Mira Whitefish! Opeongo Whitefish! Lake Cisco Lake Whitefish Pygmy Whitefish Round Whitefish Pygmy Smelt Redfin Pickerel Grass Pickerel Chain Pickerel Chiselmouth Western Silvery Minnow Eastern Silvery Minnow Weed Shiner Bluntnose Minnow Jasper Longnose Sucker! Spinynose Sculpin Cultus Pygmy Coastrange Sculpin! Mottled Sculpin Shorthead Sculpin? Texada Stickleback! Bluefin Tuna Marine Mammals Minke Whale Sei Whale Bowhead Whale? Blue Whale? Dwarf Sperm Whale Whitebeaked Dolphin Killer Whale Sperm Whale Northern Fur Seal Harp Seal 'Endemic to Canada *Updated Status Report Acipenser oxyrhynchus Acipenser fulvescens Salvelinus alpinus ssp. Salvelinus confluentus Coregonus sp. Coregonus sp. Coregonus artedi Coregonus clupeaformis Prosopium coulteri Prosopium cylindraceum Osmerus spectrum Esox americanus americanus Esox americanus vermiculatus Esox niger Acrocheilus alutaceus Hybognathus argyritis Hybognathus nuchalis regius Notropis texanus Pimphales notatus Castostomus castostomus lacustris Asemichthys taylori Cottus aleuticus Cottus bairdi Cottus confusus Gasterosteus sp. Thunnus thynnus Balaenoptera acutorostrata Balaenoptera borealis Balaena mysticetus Balaenoptera musculus Kogia simus Lagenorhynchus albirostris Orcinus orca Physeter catadon Callorhinus ursinus Phoca groenlandica Proposed Status i ? ? Vulnerable Vulnerable Threatened Endangered Threatened — Lakes Erie and Ontario 9 Vulnerable Vulnerable Vulnerable — Quebec Vulnerable Vulnerable ? Vulnerable - British Columbia 9 Vulnerable Vulnerable Vulnerable Vulnerable Vulnerable — British Columbia Threatened — British Columbia Vulnerable — British Columbia, Alberta Vulnerable Vulnerable ? ? ? Endangered Vulnerable Vulnerable 9 RS) oS) eased “Protection” sections of these reports usually con- tained a general phrase to the effect that no specific legislation was in place for the protection of fish species, but that general protection, if required, was available under habitat sections of The Fisheries Act of 1867 (this would also apply for marine mammals, but specific Marine Mammal Regulations are also promulgated under the Act). However, in 1994, it was brought to our attention that in 1983 the British Columbia Court of Appeal, in a decision to become known as the “Tsitika Judgement”, (resulting from a case between Regina and MacMillan Bloedel involving degradation and destruction of fish habitat in a section of the Tsitika River in British Columbia) ruled that the Fish Habitat Sections 31 and 33 of the Fisheries Act do not apply to those habitats that support fish which have no commercial value or sport value. In other words, there is no gen- eral vehicle to provide protection for minnows, for- age species etc., only game fish and commercial species (J. Ptolemy, Fisheries Branch, British Columbia Ministry of Environment Lands and Parks, Victoria, B.C.; personal communication). The Supreme Court of Canada refused the Federal Government appeal of the decision and apparently the Department of Fisheries and Oceans has. no intent to address the situation through amendments to the Act or by Regulation under the Act (D. Good, Department of Fisheries and Oceans, Ottawa, Ontario; personal communica- 1996 CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS 459 TABLE 3. COSEWIC definitions of conservation status. Category (Abbreviation) Species Report Accepted: Insufficient Scientific Information For Status Designation (RAISIFSD) Inderminate (I) Report Accepted: No Status Designation Required (RANSDR) Not At Risk (NAR) Vulnerable Threatened (T) Endangered (E) Extirpated (XT) Extinct (X) Definition 1988? — 1994 any species, subspecies or geographically separate population not a category, its use as a list was first approved in 1990 not a category, its use as a list was first approved in 1988 to replace NIAC! any indigenous species of fauna or flora that is particularly at risk because of low or declining numbers, occurrence at the fringe of its range or in restricted areas, or for some other reason, but is not a threatened species any indigenous species of fauna or flora that is likely to become endangered in Canada if the factors affecting its vulnerability do not become reversed any indigenous species of fauna or flora that is threatened with imminent extirpation or extinction throughout all, or a significant portion, of its Canadian range, owing to human action any indigenous species of fauna or flora no longer existing in the wild in Canada but occurring elsewhere any species of fauna or flora formerly indigenous to Canada, but no longer existing anywhere New any indigenous species, subspecies, variety or geographically defined population of wild fauna and flora a species for which there is insufficient scientific information to support status designation a species that has been evaluated and found to be not at risk a species of special concern because of characteristics that make it particularly sensitive to human activities or natural events a species that is likely to become endangered if limiting factors are not reversed A species facing imminent extirpation or extinction a species no longer existing in the wild in Canada, but occurring elsewhere a species that no longer exists 'For definitions prior to 1988 see Cook and Muir (1984). *NIAC = not in any category, used as list until 1988. tion). It has been left to provincial jurisdictions to deal with this through their legislative processes. To date we are not aware of any action taken by the provinces and any such attempt to deal with it at the provincial level would be problematic since the Constitution Act of 1867 gives the Federal Government clear jurisdiction over all Canadian fisheries waters, including the territorial seas and all inland waters. It is now clear that wording referring to general protection under habitat sections of the Act, in the “Protection” section of status reports accepted and approved prior to 1994 is in error, except for the 11 commercial and/or game species (see Table 1) listed. Definitions In its early days, one of the first problems that the Committee had to deal with was the formulation of a set of definitions of categories to be used. This was a formidable task and it is evident that the inau- gural workers gave considerable thought to the process. An overview of each year’s status assign- ments, as exemplified by this subcommittee’s reports (see Table 1 in Campbell 1984 through 1994) indicates that the original definitions (see 460 Cook and Muir 1984) have, in the main, provided an sound basis for status designations. As the Committee and its work evolved, it became evident that some definitions, or the lack thereof, gave cause for concern. Subsequently in 1988 the category “Rare” became “Vulnerable” with a change in definition and the use of NIAC (Not In Any Category) dropped in favour of an official list (not a category) to be known as “Report Accepted, No Status Designation Required” [RANSDR] (see Campbell 1989, 1990). In 1990, an additional list was created for species for which there was “Insufficient Scientific Information For Status Designation (see Campbell 1991) which became known by is acronym, “RAISIFSD”. However, the use of RAISIFSD, as a list, did not adequately signify the purpose of having such a list...that appropriate jurisdictions attempt to obtain the lacking information in order to facilitate status designation. To that end, the Committee decided to formalize the list as a category at its 1994 meeting. - At the same time, some changes in wording to cate- gory definitions were made to remove redundancy and make more clear the meaning. Also, in order to once and for all end the confusion regarding RANS- DR, a new category was established — “Not At Risk” (NAR) and the use of the RANSDR list dropped. The new definitions, in comparison to the preceding are presented in Table 3. As these changes were made at the 1994 meeting, deliberations at that meeting were based on the existing definitions as reflected in Table 1. Status assignments in future years will reflect the new definitions. Subcommittees have been directed to re-evaluate all species current- ly listed under RANSDR for recommendations to the next meeting as to the correct listing subject to the new definitions. Concluding Remarks The nine reports included in the following series are reports on the status of the respective species in Canada. Status was assigned by consensus of the COSEWIC Committee based on these reports which are published under the name(s) of the original author(s). The reports have undergone minor editing to provide a brief introduction and some degree of consistency in format and presentation. Acknowledgments The members of COSEWIC and the Fish and Marine Mammal Subcommittee would like to extend their thanks to the various authors who have so gen- erously contributed their time and talents in support of COSEWIC. The Committee also wishes to acknowledge the members of the Subcommittee for their unstinting efforts in reviewing the reports and their helpful comments. THE CANADIAN FIELD-NATURALIST Vol. 110 COSEWIC is grateful to the Canadian Wildlife Federation, World Wildlife Fund Canada, the Canadian Wildlife Service, the Canadian Museum of Nature and the Royal Ontario Museum for assis- tance provided in cash and kind. A special mention to Francis Cook and The Canadian Field-Naturalist for assistance in publication and editing and to all members of COSEWIC for their dedication and interest in the future of Canada's flora and fauna. We gratefully acknowledge the financial support provided by Environment Canada, the Province of British Columbia, and the Canadian Nature Federation which permitted the contracting several new reports. Literature Cited Campbell, R. R. 1984. Rare and endangered fishes of Canada: The Committee on the Status of Endangered Wildlife in Canada (COSEWIC) Fish and Marine Mammal Subcommittee. Canadian Field-Naturalist 98(1): 71-74. Campbell, R. R. 1985. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammals Subcommittee Status Reports: II. Canadian Field-Naturalist 99(3): 404-408. Campbell, R. R. 1987. Rare and endangered fish and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: III. Canadian Field-Naturalist 101(2): 165-170. Campbell, R. R. 1988. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: IV. Canadian Field-Naturalist 102(1): 81-86. Campbell, R. R. 1989. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: V. Canadian Field-Naturalist 103(2): 147-157. Campbell, R. R. 1990. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: VI. Canadian Field-Naturalist 104(1): 1-6. Campbell, R. R. 1991. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: VII. Canadian Field-Naturalist 105(2): 151-156. Campbell, R. R. 1992. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: VIII. Canadian Field-Naturalist 106(1): 1-6. Campbell, R. R. 1993. Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: IX. Canadian Field-Naturalist 107(4): 395—401. Campbell, R. R. 1994a. An updated report on the status of the Spotted Gar, Lepisosteus oculatus, in Canada. Report to the Committee on the Status of Endangered Wildlife in Canada (COSEWIC), Canadian Wildlife Service, Ottawa, Ontario. 7 pages. Campbell, R. R. 1994b. An updated report on the status of the Spotted Sucker, Minytrema melanops, in Canada. Report to the Committee on the Status of Endangered Wildlife in Canada (COSEWIC), Canadian Wildlife Service, Ottawa, Ontario. 6 pages. 1996 Cook, F. R., and D. Muir. 1984. The Committee on the Status of Endangered Wildlife in Canada (COSEWIC): History and progress. Canadian Field-Naturalist 98(1): 63-70. CWS. 1993. RENEW: Recovery of nationally endangered wildlife. Report Number 3. Canadian Wildlife Service, Ottawa, Ontario CW70-2/1993: 1-55. Parker, B., P. McKee, B. Hindley, M. Brinkman, M. J. Fenton, K. Rowan, J. Tanner, and C. Goddard. 1980. Rare, threatened and endangered fish species of southern Ontario: Status reports. Beak Consultants report to Department of Supply and Services, Department of Fisheries and Oceans and the National Museum of Natural Sciences, Ottawa, Ontario. 238 pages. CAMPBELL: FISHES AND MARINE MAMMALS OF CANADA STATUS REPORTS 461 Parker, B., and P. McKee. 1984a. Status of the Spotted Gar, Lepisosteus oculatus, in Canada. Canadian Field- Naturalist 98(1): 80-85. Parker, B., and P. McKee. 1984b. Status of the Spotted Sucker, Minytrema melanops, in Canada. Canadian Field-Naturalist 98(1): 104-109. Accepted 13 March 1996 Editor’s note: A complete list of designations of the Committee on the Status of Endangered Wildlife in Canada (COSEWIC) to April 1996 for all groups is avail- able from Sylvia Normand, Committee on the Status of Endangered Wildife in Canada, Ottawa, Ontario K1A 0H3 * Phone: (819) 997-4991. The Status of the Eastern Sand Darter, Ammocrypta pellucida, in Canada* + ERLING HOLM and NICHOLAS E. MANDRAK! Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario MSS 2C6 Present address: Department of Biological Sciences, Fort Hays State University, Hays, Kansas 67661 Holm, Erling, and Nicholas E. Mandrak. 1996. The status of the Eastern Sand Darter, Ammocrypta pellucida, in Canada. Canadian Field-Naturalist 110(3): 462-469. The Eastern Sand Darter, Ammocrypta pellucida, is a small slender member of the family Percidae. It is widely and dis- junctly distributed in north central North America. The degradation of its preferred sand-bottomed habitat has led to the decline of populations in the United States where it is classified by the American Fisheries Society as Threatened. Many populations have also declined or been extirpated in Canada. Recent collections from lakes Erie and St. Clair, and the Grand, Sydenham, and Thames rivers indicate that some populations still exist in Ontario. It has not been captured in recent sampling in Québec. It is recommended that the Eastern Sand Darter be classified as Threatened in Canada. Le dard de sable de l’est, Ammocrypta pellucida, est un membre, petit et élancé, de la famille des Percidae. Il est trés répandu et dispersé dans le centre nord de |’ Amérique du Nord. La dégradation des habitats aux fonds sablonneux, qu’il préfére, a provoqué un déclin de populations aux Etats-Unis, ot I’ American Fisheries Society l’a classifié “menacé”. Beaucoup de populations ont aussi décliné ou sont disparues au Canada. De récents échantillonnages des lacs Erie et St- Clair et des riviéres Grand, Sydenham et Thames indiquent qu’il en existe encore des populations en Ontario. Au Québec, il y a absence de données récentes. On recommandent, pour le dard de sable de lest, le statut de “menacé” au Canada. Key Words: Eastern Sand Darter, Dard de sable, Ammocrypta pellucida, Percidae, threatened, Ontario, Québec. The Eastern. Sand Darter (Dard de Sable), Ammocrypta pellucida (Putnam, 1863), belongs to a genus in the family Percidae which includes species commonly known as sand darters because its mem- bers are found in large creeks, rivers, and lakes with sandy bottoms. They are small pellucid fishes which are known to bury themselves completely or with only their eyes and snout showing. Scott and Crossman (1973) noted that Ammocrypta pellucida, an uncommon species in Canadian waters, has prob- ably declined in abundance from former levels. This report summarizes our current knowledge of the dis- tribution and status of the species in Canada. Description Species in the genus Ammocrypta are generally dis- tinguished from other darters (tribe Etheostomatini) by translucent and slender elongate bodies which are usually incompletely scaled. The Eastern Sand Darter (Figure |) differs from the other six species of the genus in the following characteristics. It is pale white, yellowish or silvery coloured with a series of 10 to 14 lateral dark spots usually located entirely below the lateral line scale row. These spots are slightly smaller than the pupil, and are frequently rounded anteriorly and oblong posteriorly. The median fins are not pig- mented. Ammocrypta pellucida is one of the most elongate species of Ammocrypta, with body depth entering into standard length usually 8 to 9 times. There are usually 10 to 12 transverse scale rows on each side, 4 to 7 of these below the lateral line, and 9 to 11 (usually 10) preoperculomandibular canal pores. The pelvic rays of adult males are darkly pigmented and have small tubercles. Simon et al. (1992) described larval characteristics of five sand darter species, including Ammocrypta pellucida. Average adult size ranges from 46 — 71 mm total length (TL), and maximum recorded size is 81 mm TL (from Scott and Crossman 1973; Williams 1975; Trautman 1981). Although the genus Ammocrypta is recognized in the widely accepted Common and Scientific Names of Fishes from the United States and Canada (Robins et al. 1991), Simons (1991, 1992) proposes that Ammocrypta be downgraded to the subgenus level and that six species within the subgenus, including Ammocrypta pellucida, be placed in the genus Etheostoma. His study indicates that the genus Ammocrypta is not monophyletic, and when reduced to a monophyletic group (by removing one species), Ammocrypta exhibits a similar amount of character variation as the other Etheostoma subgenera Boleosoma and Ioa. However, there has been insuffi- *Reviewed and Approved by COSEWIC 14 April 1994, status assigned — Threatened. +Contribution number 58 of the Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario. 462 1996 HOLM AND MANDRAK: STATUS OF THE EASTERN SAND DARTER 463 FiGurRE |. The Eastern Sand Darter, Ammocrypta pellucida (Putnam), 59 mm TL, Sydenham River, Lambton Co. (ROM 56997). Drawing by Anker Odum. cient time for this proposed name change to be scru- tinized and accepted by the scientific community and the well-established scientific name Ammocrypta pellucida is retained here. Distribution In North America, the Eastern Sand Darter has been found in the Ohio River basin in Ohio, Indiana, Illinois, Kentucky, W. Virginia, and Pennsylvania. It has been recorded from the Lake Huron, Lake St. Clair and Lake Erie drainages in Michigan, Ohio, New York, and Ontario. It has not been recorded in the Lake Ontario drainage, but occurs farther east in Saint- Laurent (St. Lawrence River) and Lac Champlain drainages of Québec, Vermont, and New York (see inset, Figure 2). Recent reductions in distribution have been documented in Kentucky (Kuehne and Barbour 1983), Illinois (Smith 1971), Ohio (Trautman 1981), Michigan (Smith et al. 1981), and Pennsylvania (Cooper 1983). Since 1970, Ammocrypta pellucida has been recorded at new localities in New York (Smith 1985), Ontario, and Québec. In Ontario, Ammocrypta pellucida was collected prior to 1970 from the Ausable River, the Sydenham River (Kent, Lambton, and Middlesex counties), and the Thames River, Catfish Creek, Big Otter Creek, and Big Creek. It was first collected in the Canadian waters of Lake Erie in 1953 at Pelee Island, and in both eastern and western basins of Lake Erie in 1957. Since 1970, specimens have been collected at new locations in Lake Erie at Rondeau Bay and Long Point Bay, Lake St. Clair, and in the Grand River [see Figure 2 (Appendices of all records are on file and available on request from COSEWIC, Canadian Wildlife Service, Ottawa, Ontario K1A 0H3)]. In Québec, Ammocrypta pellucida was collected prior to 1970 in Lac des Deux-Montagnes near Montréal, Saint-Laurent near Sorel, and in some larger tributaries of Saint-Laurent: Riviere Chateauguay, Riviere L’ Assomption, Riviére Yamaska, Riviére Saint-Francois, Riviere Yamachiche, and Riviere Gentilly. Since 1970, spec- imens have been collected in Riviére Richelieu, Chenal aux Ours (a channel between the Berthier Islands at the western end of Lac St-Pierre), Riviere Becancour, Riviere aux Orignaux, and Petite Riviere du Chéne (see Figure 2). On the map of Hocutt (1980), one distribution point appears to be on the Canadian side in the St. Lawrence River around Cornwall, Ontario. Since we could not find any capture records in the vicinity of Cornwall, we conclude that this point probably rep- resents the record for the Little Salmon River near Fort Covington, New York (Smith 1985). We were unable to verify the authenticity of a distribution point about 15 km upstream from the only record we have located in Riviere L’ Assomption, Québec. According to C. H. Hocutt, D. E. McAllister (editors of the Atlas of North American Freshwater Fishes (Lee et al. 1980); and J. D. Williams (U.S. Fish and Wildlife Service Exotic Fish Lab, Gainsville Florida) Canadian distribution records on the map of Hocutt (1980) are based on Mongeau et al. (1974, 1979), Williams (1975), and Mongeau (1979a). The upper L’Assomption River record could not be found in any of these publications. Protection Ammocrypta pellucida receives no special protec- tion in Canada (but see Habitat section). It is listed as threatened in the United States by the American Fisheries Society (Williams et al. 1989). It is clas- sified as Endangered in Ohio, Michigan, Pennsylvania, and New York and listed as of Special Concern in Indiana and Kentucky (Johnson 1987). Population Sizes and Trends Ammocrypta pellucida has declined in, or has been extirpated from, many areas of its North American range. Formerly widespread and abundant prior to 1900, the Eastern Sand Darter displayed a steady decline in abundance between 1925 and 1950 in Ohio. Despite thorough investigations, few speci- mens were captured after 1955 (Trautman 1981). It has been decimated in the upper Wabash River, a major portion of its former range in Hlinois (Smith 1971). Cooper (1983) suggested Ammocrypta pellu- cida has disappeared from the Monongahela drainage in southwestern Pennsylvania. Few popula- tions still survive in New York (Smith 1985). 464 Kuehne and Barbour (1983) reported declines in Kentucky in the upper Kentucky and Licking rivers. In Canada, populations have declined or been extir- pated from several areas. However, recent collec- tions of specimens indicate that Lake Erie, Lake St. Clair, and several rivers in southwestern Ontario and Québec continue to support populations. Studies have not been specifically conducted to estimate population sizes of Ammocrypta pellucida in Canada. However, changes in population size may be inferred from sampling data. Between 1922 and 1958, the Eastern Sand Darter was collected at 13 sites in six rivers (Ausable, Sydenham, Thames, Catfish, Big Otter, Big) in southwestern Ontario. An intensive sampling program was conducted between 1970 and the mid 1980s by the Ontario Ministry of Natural Resources (OMNR), Royal Ontario Museum (ROM), and the Canadian Museum of Nature (Mandrak and Crossman 1992). During this time, the Eastern Sand Darter was collected at seven new sites in three river systems (Thames, Sydenham, Grand). At least six of the 13 sites sampled prior to 1959 in five streams (Ausable, Thames, Catfish, Big Otter, Big) were sampled during this program, but the species was captured at none. Between 1989 and 1991, ROM surveys specifically targeted suitable sand-bottomed habitats. All 13 sites where Ammocrypta pellucida was known prior to 1959 were sampled. It was captured at only three of those sites. Seven sites in three river systems (Thames, Sydenham, Grand), including five of the seven sites where the species was captured between 1970 to 1987, and two new sites were also sampled. The Eastern Sand Darter was found at all of these sites. A single specimen was collected in a Lake Huron tributary, the Ausable River, in 1928. Subsequent sampling at this site in 1936 and in 1982, and in 1974 at five sites within 5 km of the capture site, failed to collect specimens. Therefore, it probably no longer occurs in the Ausable River. Ammocrypta pellucida has been collected in two Lake St. Clair tributaries. Forty-eight specimens were collected in the Thames River “at Muncey” in 1923 (Hubbs and Brown 1929). Although it was not captured “near Muncey” in a 1941 sampling, it was recorded during sampling downstream from Muncey (the community of Muncey in Gazetteer of Canada, Ontario: 42°49’N, 81°29’ W) in the Thames between Wardsville and the Moravian Indian Reserve in 1958 (ROM Accession 482). The Eastern Sand Darter has been collected at four other sites both upstream and downstream of Muncey in the 1970s. The 1989 to 1991 ROM surveys found Ammocrypta pellucida at most locations in the Thames River where they had been captured in the past including the upstream and downstream extremes. Therefore, it can be conclud- ed that the range of the Eastern Sand Darter has not been reduced in the Thames River. However, the THE CANADIAN FIELD-NATURALIST Vol. 110 number of specimens captured in one sampling in 1923 is approximately equivalent to the total number of specimens captured in at least 22 subsequent sam- pling attempts indicating that its abundance has declined in the Thames River. Ammocrypta pelluci- da was collected in the Sydenham River at Strathroy in 1927, Alvinston in 1929, and downstream at the mouth of one of its tributaries, Fansher Creek, in 1972. It was also captured in the “East Sydenham River” in 1983 (Wilfrid Laurier University 8123). However, no additional locality data for this record were available. Eight collections, made by the ROM at seven sites with sandy bottoms between Strathroy and Alvinston in 1991, failed to capture the Eastern Sand Darter. It was collected in the Sydenham River | at and 700 metres below the mouth of Fansher Creek during fish surveys conducted in 1989 and 1991. In | 1991, it was captured farther downstream at another location in the Sydenham River. The Eastern Sand | Darter still exists in the Sydenham River but because there are no historical data for the Sydenham River downstream of Alvinston, it cannot be determined whether the range of Ammocrypta pellucida has been reduced or has shifted downstream. The Eastern Sand Darter was collected in three cen- tral Lake Erie tributaries prior to 1970 and one tribu- tary in 1987. Specimens were collected in Big Creek and Big Otter Creek in 1923 and 1955, and in Catfish Creek in 1922 and 1941. Between 1973 and 1990, four sampling attempts in both Big Creek and Catfish Creek, and nine sampling attempts in Big Otter Creek failed to capture specimens. Therefore, it is probable that Ammocrypta pellucida no longer occurs in Big Creek, Big Otter Creek, and Catfish Creek. In 1987, the Eastern Sand Darter was first captured in the Grand River, at Brantford. The Grand River has been sampled in the vicinity of Brantford between 1966 and 1976 (W. Yerex, Grand River Conservation Authority, Brantford, Ontario; personal communica- tion). However, the site of the 1987 capture of Ammocrypta pellucida was not sampled during this | period and there is no evidence in records available that it was sampled prior to 1966. This population is 60 km from the nearest known native population in Big Creek. It may be the result of an unrecorded intro- duction, or a remnant of a formerly wider range frag- mented by environmental or cultural impacts. | Additional specimens caught in 1991 confirm that this population is established and self-reproducing. Ammocrypta pellucida was first collected in the Canadian waters of Lake Erie at Pelee Island in 1953 (University of Florida 9911). Additional specimens were collected in both eastern and western basins | during trawls conducted in 1957 (Scott and Crossman 1973), in Rondeau Bay in 1975, and in the | western basin in 1984 and 1985. OMNR index net- ting trawls in Long Point Bay conducted since 1972, captured specimens every year between 1979 and 1996 HOLM AND MANDRAK: STATUS OF THE EASTERN SAND DARTER 465 Ammocrypta pellucida o captured before 1970 e captured 1970 - 1991 0 25 50 etre Kilometres FiGurE 2. Canadian distribution of Ammocrypta pellucida, with inset of North American dis- tribution modified from Hocutt (1980). 1987, except 1983. Neither sampling techniques nor personnel had changed in the OMNR index netting program in Long Point Bay (S. Nepszy, Lake Erie Fisheries Research Station, OMNR, Wheatley, Ontario; personal communication). Therefore, the appearance of specimens in Long Point Bay only after 1978 was not the result of changes in expertise of detection or sampling techniques and may repre- sent the establishment of a new population. The Eastern Sand Darter was first collected in the Canadian waters of Lake St. Clair in Mitchell’s Bay by an OMNR small trawl study conducted between 1983 and 1985. The results of the study show an overall decline in numbers of captured specimens (1983 97 specimens, 0.6 specimens/tow; 1984: 66, 0.4; 1985: 26, 0.2). This does not necessarily indicate a decline of Ammocrypta pellucida in the lake, as this trend may be the result of other factors such as normal fluctuations in year-class numbers or variations in sam- pling techniques (Nepszy, personal communication). Due to the limited sampling of suitable habitats in Lake Erie and Lake St. Clair, it is difficult to deter- mine the present status of Eastern Sand Darter popu- lations at specific locations within these lakes. However, the population in Long Point Bay appeared to be stable during the nine-year period from 1979 to 1987. It is possible that populations of Ammocrypta pellucida exist in areas of sandy habitat in lower Lake Huron. These areas have not been sampled by the methods used in Lake Erie and Lake St. Clair (B. Payne, Lake Huron Fisheries Assessment Unit, OMNR, Owen Sound, Ontario; personal communication). Before 1970, Ammocrypta pellucida was collected in southwestern Québec at two sites on Lac des deux Montagnes, one site on Saint-Laurent, and 11 sites on six Saint-Laurent tributaries. Since 1970, it has been collected at one site in Chenal aux Ours, and at 37 sites on seven Saint-Laurent tributaries. Ammocrypta pellucida was collected in Lac des Deux Montagnes in 1941 and 1946. Mongeau and Massé (1976) and Mongeau et al. (1980) did not report its capture in their studies of the waters around Montréal between 1964 and 1977. A sampling attempt in 1990 by ROM on a shallow, sandy beach at the 1941 site, Anse a L’Orme, failed to capture any Ammocrypta. The only recorded capture of the Eastern Sand Darter in Saint-Laurent (not including lake expansions) was reported by Cuerrier et al. (1946). It was not recorded in a 1973 survey of 325 466 sampling stations in Saint-Laurent between Montréal and Sorel (Massé and Mongeau 1976). Cuerrier et al. (1946) reported that the Eastern Sand Darter was par- ticularly abundant in Riviere Saint-Frangois in the Lac Saint-Pierre region. However, it has not been captured in the Riviere Saint-Francois since 1944 despite sampling by Service de |’ Aménagement de la Faune, Ministére du Loisir, de la Chasse et de la Péche (MLCP)-Montréal between 1965 and 1975 (Mongeau and Legendre 1976) and in 1991 (Audet and St-Onge 1992). Fourteen specimens were collect- ed at a single site on Riviere L’ Assomption in 1969. No specimens were captured at or near this site (Station 4) nor at any of the other 15 sites sampled on this river during electrofishing surveys in 1990. At station 4, the water was described as very turbid and the substrate consisted of 100% clay (St-Onge 1992). The Eastern Sand Darter probably no longer occurs in Lac des Deux Montagnes, Saint-Laurent, Riviere L’ Assomption, and Riviére Saint-Francois. Ammocrypta pellucida was collected at 12 sites on Riviere Chateauguay between 1941 and 1976. Vladykov (1942) reported the capture of three speci- mens in June 1941 from Riviere Chateauguay near Ste. Philomeéne village (now Mercier). Cuerrier et al. (1946) later documented the capture in August 1943 of about 180 specimens at a site near the city of Chateauguay. A fish survey of Riviere Chateauguay, where the main course of the river was sampled at approximately 0.32 km intervals from the mouth to the headwaters, was conducted by MLCP-Montréal during 1975 and 1976 (Mongeau et al. 1979). The Eastern Sand Darter was collected at 10 of 287 sam- pling stations distributed along approximately 55 km of the main course of the river and at one site in one of its tributaries, Riviere Trout. It ranked 31 out of 53 total species in frequency of occurrence in the collections. However, it was not recorded from the city of Chateauguay where it had been previously reported as abundant (Cuerrier et al. 1946). In Riviere Yamaska, Ammocrypta pellucida was cap- tured between 1963 and 1971 at four of 120 sam- pling stations within a 5 km stretch of the river. It ranked 37 out of 59 species in frequency of occur- rence in the collections (Mongeau 1979a). No speci- mens were caught during sampling in the Riviere Chateauguay conducted in 1993 (Nathalie La Violette, University of Toronto, Toronto, Ontario; personal communication). It has been recorded from three sites in Riviere Yamachiche near the mouth in 1944 and 1972. As the result of lack of recent sam- pling, it is not possible to determine the current sta- tus of the Eastern Sand Darter in Riviére Yamachiche. The Eastern Sand Darter was collected at one site in 1941 and at two sites in 1982 in Riviére Gentilly where it likely still exists. Since 1970, the Eastern Sand Darter has been col- lected in four additional tributaries of Saint-Laurent, THE CANADIAN FIELD-NATURALIST Vol. 110 and in Chenal aux Ours (in 1974). It has been cap- tured once at single sites in Riviére Becancour (1981; ROM 42091), Riviere aux Originaux (1982; MacFarlane et Durocher 1984), and Petite Riviére du Chéne (1982; MacFarlane et Durocher 1984). In 1970, Ammocrypta pellucida was collected at 19 of 159 sampling stations in a 60 km stretch of the main channel of Riviere Richelieu from McMasterville to its mouth. It ranked 30 of 60 total species in frequen- cy of occurrence in the collections (Mongeau 1979b). In 1974, four specimens were collected at one site less than a kilometre from the mouth of Riviere Richelieu, but at none of the other 481 sites sampled in the Lac St. Pierre region (Massé and Mongeau 1974). Because there has been no sampling since 1982, it is not possible to determine the current status of Ammocrypta pellucida in the rivers where it has been captured since 1970. It is apparent that populations of Ammocrypta pel- lucida in tributaries to the north central shore of Lake Erie, the upper Sydenham River, and the Ausable River have been extirpated. Analysis of the results of sampling after 1958 by ROM, OMNR, and NMC indicate that the decline of populations in Ontario streams occurred prior to 1970. Results of | sampling between 1990 and 1993 in Québec suggest that populations in the Chateauguay, Yamaska, l’Assomption and St-Frangois rivers have declined or been extirpated. However, capture results may be affected by differences in sampling gear (the 1990s sampling was conducted using boat and back-pack electrofishers, whereas seine nets were used in earli- er sampling). The current status of other populations, is unknown as the result of the lack of recent sam- pling. Based on recent sampling, populations are considered extant in Lake Erie, Lake St. Clair and its tributaries, the Grand River, and Riviere Gentilly. Habitat The preferred habitat of Ammocrypta pellucida is sand-bottomed areas in streams and rivers, and sandy shoals in lakes (Scott and Crossman 1973). It has also been collected over a bottom of “limestone ter- races covered with a thin layer of mud” in Riviere Chateauguay (Vladykov 1942), in riffles over rubble and gravel, and on silted sand bottoms in the Sydenham River (ROM 56997). It has been found in waters that are clear, tea-coloured and highly turbid (secchi depth =15 cm). In these waters, aquatic vege- tation ranged from absent to some submerged macrophytes present, and current ranged from still to swift (unpublished data, ROM). Siltation associated with agricultural practices has led to the significant reduction of the preferred sand- bottomed habitat of the Eastern Sand Darter. For example, a significant increase in turbidity, indica- tive of an increased potential for siltation, has been documented in Big Creek as the result of an increase 1996 in cropped area between 1931 and 1961 associated with the introduction of tobacco. Tobacco farming is characterised by a high percentage of exposed ground, which on the Norfolk Sand Plain results in considerable erosion and siltation (Whillans 1977). This impact is not limited to Big Creek, but has like- ly occurred in all central Lake Erie watersheds where tobacco farming was practised. Provincial legislation exists that nominally pro- tects the habitat of Ammocrypta pellucida. The Ontario Lakes and Streams Improvement Act pro- hibits the impoundment or diversion of watercourses which leads to silt accumulation. The Land Stewardship II program of the Ontario Ministry of Agriculture and Food (OMAF), designed to reduce the erosion of agricultural lands, has the potential to slow the degradation of remaining critical habitat by reducing siltation. However, this program is volun- tary and can only be implemented with the coopera- tion of landowners. In Québec, habitat is generally protected by the Environmental Quality Act, and may be protected by the Ecological Reserves Act if a species is “threatened with disappearance or extinc- tion.” Specific protection can be provided through the Endangered Species Act and conservation laws. General Biology Reproductive Capability Based on a study in the Scioto River drainage in southern Ohio (Spreitzer 1979), Ammocrypta pellu- cida females are ready to spawn at age 1+ if they have reached a standard length of 36 mm. Fecundity is low but comparable to many Etheostoma species. Total number of eggs for ova-bearing females ranged from 22 to 829 (mean = 343.1) and the num- ber of mature ova in fecund females ranged from 30 to 170 (mean = 71.0). Larger females produced more eggs. Based on female fecundity and the great dis- parity in sizes of individuals of the same year class, the 1974 spawning season was protracted, ranging from May to mid-August. Sex ratio was determined to be 1:1 during the entire year, including the spawn- ing season. Water temperature during spawning sea- son ranged from 14.4°C to 24.4°C In captivity, the Eastern Sand Darter has been observed to spawn at water temperatures between 20.5°C and 23°C. During spawning the male mounts the female, and eggs are deposited when the pair have vibrated and buried their tails and caudal peduncles in the substrate. “Sneaker males” often joined mating pairs (Johnston 1989). A well-oxy- genated substrate such as unsilted sand is likely required for high egg survivorship. Spreitzer (1979) suggested that the spawning season was synchro- nized with low silt levels in the habitat. Examination of the gonads of 17 specimens in the ROM collec- tion indicated that Ammocrypta pellucida probably spawns between late June and late July in Ontario. HOLM AND MANDRAK: STATUS OF THE EASTERN SAND DARTER 467 Species Movement The movements of the Eastern Sand Darter are virtually unknown. Most darters are sedentary, and migrations are rare (Page 1983). However, Johnston (1989) suggested male Ammocrypta pellucida may have congregated in an area sampled in the Tippecanoe River, Indiana, in July 1987. Spreitzer (1979) gave evidence that some individuals may migrate to feed when local chironomid population levels are low. Behaviour/Adaptability The diet of Ammocrypta pellucida has been report- ed to be limited to midge larvae, blackfly larvae and possibly entomostracans by its small mouth size and restricted habitat (Scott and Crossman 1973; Smith 1979; Cooper 1983). In southern Ohio, chironomid larvae comprised an average of 94.4% of the diet of the Eastern Sand Darter. Oligochaetes and cladocer- ans comprised significant, but smaller, proportions in June and November, respectively (Spreitzer 1979). Fossorial behaviour is well-developed in Ammocrypta. Daniels (1989) provided evidence indi- cating that burying is an adaptation to maintain posi- tion on the relatively homogenous sand beds, partic- ularly during periods of extremely high or low flow. His experiments suggested that Ammocrypta does not bury itself to avoid predators or to ambush prey. Low oxygen levels in silted substrate may discour- age complete burial, or reduce the length of burial time. This may have a negative survival effect by increasing the amount of energy expended to main- tain position in its habitat. Limiting Factors Siltation of critical habitat, impoundments, and deterioration of water quality from effects such as chemical pollution and acid mine drainage, are factors attributed to the decline of the Eastern Sand Darter in Ohio, Illinois, and Kentucky (Smith 1971; Barnes 1979; Trautman 1981; Burr and Warren 1986). Poor water quality near urban areas such as Montréal and Chateauguay, Québec, may have caused its decline or extirpation in those areas (Scott and Crossman 1973). Siltation seems to be the leading cause of significant loss of critical habitat in Canada. Silt reduces the available substrate oxygen, necessary for fossorial behaviour and egg survivorship. It has caused the decline and extirpation of Ammocrypta pel- lucida in some rivers where it was formerly abundant. Special Significance of the Species Ammocrypta pellucida is classified as Threatened in the United States and Threatened or Endangered in most U.S. states where it is present. Some Ammocrypta (Ammocrypta beani, Ammocrypta bifascia, and Ammocrypta meridiana) are considered common by Page and Burr (1991). Other species in the genus such as Ammocrypta clara and 468 Ammocrypta vivax have been extirpated from parts of their range (see Becker 1983; Robison and Buchanan 1988). Therefore, it can be concluded that the genetic diversity, expressed in behaviour, ecolo- gy, and morphology represented in the genus Ammocrypta 1s in jeopardy. Evaluation Population declines will continue in areas where the Eastern Sand Darter is still present, unless the sil- tation of critical habitat and continued degradation of water quality is prevented. If further siltation and water quality degradation is prevented, populations surviving in areas of remaining critical habitat should stabilize. Extirpated populations may be re- established only if silt is removed from preferred habitat and dispersal from an extant donor popula- tion is possible. Due to physical barriers and limited dispersal capabilities, it is unlikely that populations would become re-established naturally in waterbod- ies without extant populations. Populations of Ammocrypta pellucida in the Grand River and Lake Erie appear to be stable. Many other populations have declined or been extir- pated in Canada. The population trend of the remain- ing populations cannot be assessed due to lack of adequate sampling in the last 10 years. It is likely that all populations will decline or become extirpated if further loss of critical habitat is not prevented. Therefore, until sampling of these populations is undertaken to determine their stability, it is recom- mended that the Eastern Sand Darter be classified as Threatened in Canada. Acknowledgments Financial support was provided by the Royal Ontario Museum, Environmental Youth Corps grants to E. J. Crossman and EH and a World Wildlife Fund grant to E. J. Crossman. Frederique Arnaud, David Boehm, Lisa Dorman, Kira Dunham, Colleen Fennesey, William Ramshaw, Marty Rouse, and Franklin Wong assisted with field work. William Ramshaw, Frederique Arnaud, and Kira Dunham assisted with verification and summary of distribu- tion records. Distribution maps were drawn by William Ramshaw. We are grateful to the following people who pro- vided information and distribution records: Pierre Dumont and Gilles Roy (MLCP, Montréal), Yves Mailhot (MLCP, Trois Rivieres), Steve Nepszy (Lake Erie Fisheries Research Station, OMNR), Nathalie La Violette (Québec Ministére de l'Environnement et de la Faune), Larry Witzel (Lake Erie Fisheries Assessment Unit, OMNR), D. E. McAllister (NMC), Douglas Nelson (University of Michigan Museum of Zoology), Daryl Coulson (Cambridge, OMNR), Warren Yerex (Grand River THE CANADIAN FIELD-NATURALIST Vol. 110 Conservation Authority). E. J. Crossman, J. S. Nelson, and D. E. McAllister provided useful com- ments on the manuscript. Literature Cited Audet, R. and J. St-Onge. 1992. Recueil de données brutes sur la faune piscicole récoltée dans la Riviére Saint-Frangois (aoait et septembre 1991). Direction de la qualité des cours d’eau, Ministere de L’ Environnement du Québec, ENVIRODOC No EN 920301, Rapport interne QE-92-14. Barnes, M. D. 1979. Eastern sand darter, Ammocrypta pellucida and other fishes from the streams of the Wayne National Forest. Ohio Journal of Science 79(2): 92-94. Becker, G. C. 1983. Fishes of Wisconsin. The University of Wisconsin Press. Madison Wisconsin. Burr, B. M., and M. L. Warren, Jr. 1986. A distribu- tional atlas of Kentucky fishes. Kentucky Nature Preserves Commission, Scientific and Technical Series Number 4. Cooper, E. L. 1983. Fishes of Pennsylvania and the north- eastern United States. The Pennsylvania State University Press. Cuerrier, J.-P., F. E. J. Fry, and G. Fontaine. 1946. Liste préliminaire des poissons de la région de Montréal et du Lac Saint-Pierre. Le Naturaliste canadien 73: 17-32. Daniels, R. A. 1989. Significance of burying in Ammo- crypta pellucida. Copeia 1989(1): 29-34. Hocutt, C. H. 1980. Ammocrypta pellucida (Agassiz), Eastern Sand Darter. Page 620 in Atlas of North American Freshwater Fishes. Edited by D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer. North Carolina State Museum Natural History, Publication Number 1980-12. Hubbs, C. L., and D. E. Brown. 1929. Materials for a dis- tributional study of Ontario fishes. Transactions Royal Canadian Institute 17(1): 1-56. Johnson, J. E. 1987. Protected fishes of United States and Canada. American Fisheries Society, Bethseda, Maryland. Johnston, C. E. 1989. Spawning in the eastern sand darter, Ammocrypta pellucida (Pisces: Percidae), with comments on the phylogeny of Ammocrypta and related taxa. Transactions of the Illinois Academy of Science 82(3 and 4): 163-168. Kuehne, R. A., and R. W. Barbour. 1983. The American darters. The University Press of Kentucky, Lexington, Kentucky. Lee D.S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer. Editors. 1980. Atlas of North American Freshwater Fishes. North Carolina State Museum Natural History, Publication Number 1980-12. MacFarlane, A. et L. Durocher. 1984. Inventaire ichthy- ologique de plusieurs tributaires de la rive sud du lac Saint-Pierre et du fleuve Saint-Laurent (Région de Gentilly). Ministére du Loisir, de la Chasse et de la Péche, Direction régionale des Trois-Riviéres, S. A. E. F., Coordination de la faune aquatique. Mandrak, N. E., and E. J. Crossman. 1992. A checklist of Ontario freshwater fishes annotated with distribution maps. Royal Ontario Museum Life Sciences Publication. f 1996 Massé, G., and J.-R. Mongeau. 1974. Répartition géo- graphique des poissons, leur abondance relative et bathymétrie de la région du lac Saint-Pierre. Service de 1’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Massé, G., and J.-R. Mongeau. 1976. Influence de la nav- igation maritime sur la répartition géographique et l’abondance relative des poissons du fleuve Saint- Laurent entre Longueuil et Sorel. Service de 1’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Rapport Technique. Mongeau, J.-R. 1979a. Les poissons du bassin de drainage de la rivigre Yamaska, 1963 a 1975. Service de 1’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Mongeau, J.-R. 1979b. Dossiers des poissons du bassin versant de la Baie Missisquoi et de la Riviere Richelieu, 1954 a 1977. Service de |’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Mongeau, J. R., A. Courtemanche, G. Massé, and B. Vincent. 1974. Cartes de répartition geographique des espéces de poissons du sud du Québec, d’aprés les inventaires ichthyologiques effectués de 1963 a 1972. Serice de | Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1979. Les Poissons du bassin de drainage de la riviére Chateauguay, leur milieu naturel, leur répartition géo- graphique et leur abondance relative. Service de l’Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1980. La répartition géographique des poissons, les ensemence- ments, la péche sportive et commerciale, les frayéres et la bathymétrie du fleuve Saint-Laurent dans le Bassin de La Prairie et les Rapides de Lachine. Service de l’Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Rapport Technique No. 06-29. Mongeau, J.-R., and V. Legendre. 1976. Les resources fauniques du bassin inférieur de la Riviére Saint- Francois: évolution des populations en dix ans 1965-1974. Service de 1’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Rapport Technique. Mongeau, J.-R., and G. Massé. 1976. Les poissons de la région de Montréal, la péche sportive et commerciale, les ensemencements, les frayéres, la contamination par la mercure et les PCB. Service de 1’ Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Robins, C. R., R. M. Bailey, C. E. Bond, J. R. Brooker, E. A. Lachner, R. N. Lea, and W. B. Scott. 1991. Common and scientific names of fishes from the United States and Canada. American Fisheries Society Special Publication 20, Bethseda, Maryland. Robison, H. W. and T. M. Buchanan. 1988. Fishes of Arkansas. University of Arkansas Press, Fayetteville, Arkansas. HOLM AND MANDRAK: STATUS OF THE EASTERN SAND DARTER 469 Page, L. M. 1983. Handbook of Darters. TFH Publi- cations, Inc. Ltd. Neptune City, New Jersey. Page, L. M., and B. M. Burr. 1991. A field guide to freshwater fishes. Houghton Mifflin Company, Boston. Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- es of Canada. Fisheries Research Board of Canada Bulletin 184. Simon, T. P., E. J. Tyberghein, K, J. Scheidegger, and C. E. Johnston. 1992. Descriptions of protolarvae of the sand darters (Percidae: Ammocrypta and Crystallaria) with comments on systematic relationships. Ichthyo- logical Exploration of Freshwaters 3(4): 347-358. Simons, A. M. 1991. Phylogenetic relationships of the crystal darter, Crystallaria asprella (Teleostei: Percidae). Copeia 1991(4): 927-936. Simons, A. M. 1992. Phylogenetic relationships of the Boleosoma species group (Percidae: Etheostoma. Pages 268-292 in Systematics, historical ecology and North American freshwater fishes. Edited by R. L. Mayden. Stanford University Press, Stanford, California. Smith, C.-L. 1985. The inland fishes of New York State. New York State Department of Environmental Conservation. Albany, New York. Smith, G. R., J. N. Taylor, and T. W. Grimshaw. 1981. Ecological survey of the fishes in the Raisin River drainage, Michigan. Papers of the Michigan Academy of Science, Arts, and Letters 13(3): 275-305. Smith, P. W. 1971. Illinois streams: a classification based on their fishes and an analysis of factors responsible for disappearance of native species. Illinois Natural History Survey. Biological Notes Number 76. Smith, P. W. 1979. The fishes of Illinois. University of Illinois Press. Urbana, Illinois. Spreitzer, A. E. 1979. The life history, external morphol- ogy, and osteology of the eastern sand darter, Ammocrypta pellucida (Putnam, 1863), an endangered Ohio species (Pisces: Percidae). M. Sc. thesis (unpub- lished). The Ohio State University. Columbus, Ohio. St-Onge, J. 1992. Recueil de données brutes sur la faune piscicole récoltée dans la Riviere L’ Assomption (aout et septembre 1990). Direction de la qualité des cours d’eau, Ministére de L’Environnement du Québec, ENVRO- DOC No EN 920117, Rapport interne QE-92-10. Trautman, M. B. 1981. The fishes of Ohio. Ohio State University Press. Columbus, Ohio. Vladykov, V. D. 1942. Two fresh-water fishes new for Québec. Copeia 1942(3): 193-194. Whillans, T. 1977. Fish community transformation in three bays within the lower Great Lakes. M. Sc. thesis, University of Toronto. Toronto, Ontario. Williams, J. D. 1975. Systematics of the percid fishes of the subgenus, Ammocrypta with descriptions of two new species. Bulletin of the Alabama Museum of Natural History, Number 1. Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. Contreras-Balderas, J. D. Williams, M. Navarro- Mendoza, D. E. McAllister, and J. E. Deacon. 1989. Fishes of North America Endangered, Threatened, or of Special Concern: 1989. Fisheries 14(6): 2—20. Accepted 13 March 1996 The Status of the Cutlips Minnow, Exoglossum maxillingua, in Canada**+ E. J. CROSSMAN and ERLING HOLM Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario MSS 2C6 Crossman, E. J., and Erling Holm. 1996. The Status of the Cutlips Minnow, Exoglossum maxillingua, in Canada. Canadian Field-Naturalist 110(3): 470-477. The Cutlips Minnow, Exoglossum maxillingua, is a stout bodied minnow distinguished from all other members of the fami- ly Cyprinidae in North America by its unique trilobed lower jaw. It is common in many upland regions of the Atlantic coast drainage of northeastern North America. It is rare, and there is evidence that populations have declined from levels in the 1930s in Ontario. It is more widespread in Québec where it has been found in numerous river systems from 1935 to 1989. Since 1977, there has been a significant decrease in the number of surveys in the limited area occupied by this species in Canada. Surveys might have provided more information on present status. There is, however, evidence of low and/or declining numbers in some river systems. Le bec-de-lievre, Exoglossum maxillingua, est un méné au corps trapu qui se distingue de tous les autres membres de la famille Cyprinidae en Amérique du Nord par sa machoire inférieure trilobée. Il est répandu dans de nombreuses régions hautes du bassin versant de ]'Atlantique dans le nord-est de l'Amérique du Nord. On le rencontre rarement en Ontario, ou sa population enregistre une baisse par rapport aux chiffres des années 1930. Il est plus répandu au Québec ow I'on a relevé sa présence dans nombre de réseaux fluviaux de 1935 a 1989. Depuis 1977, le nombre d'études effectuées dans la région restreinte occupée par cette espéce au Canada a diminué de fagon marquée. Si tel n'avait pas été le cas, nous disposerions probablement de plus de données sur la situation actuelle de l'espéce. I] semblerait toutefois que sa population est faible ou a la baisse dans certains réseaux. Key Words: Cutlips Minnow, Exoglossum maxillingua, Bec-de-Liévre, cutlips, eye-picker, Cyprinidae, minnows. The Cutlips Minnow, Exoglossum maxillingua, (Lesueur, 1817), is a stout bodied minnow (Figure 1) which can reach a total length of about 150 mm (Pappantoniou et al. 1984a). It can be distinguished from all other North American minnows by its unique trilobed lower jaw consisting of a central bony tongue-like lobe, two lateral fleshy lobes, and no maxillary barbels. The Tonguetied Minnow, Exoglossum laurae, the only other species in the genus and known only from the United States, has a lower jaw which is not as obviously trilobed and fre- quently possesses a maxillary barbel. Male and female Cutlips Minnows are approxi- mately equal in size and outside the reproductive season there are no obvious external differences between the sexes. During the reproductive season, mature males develop tubercles on the paired fins (Pappantoniou 1983). Larval development of the Cutlips Minnow has been described by Fuiman and Loos (1978) and Buynak and Mohr (1980). The lat- ter reference provides a key to six species of cyprinids, four of which are frequently found in association with Exoglossum maxillingua in Canada. Exoglossum maxillingua is currently considered to be derived from Exoglossum laurae (Gilbert and Lee 1980). Phylogenetic analysis indicates that the genus Exoglossum is most closely related to Phenacobius, a genus of minnows restricted to the Mississippi and Gulf of Mexico drainages of the United States (Coburn and Cavender 1992). Geographic variation in four morphometric and seven meristic characters has been investigated using 1247 specimens, includ- ing five individuals from the St. Lawrence drainage. Ten characters displayed significant geographic vari- ation between populations, but this variation was not correlated with latitude (Pappantoniou 1983). Distribution The Cutlips Minnow is found in eastern North America (see inset map, Figure 2) in the Atlantic drainage from the St. Lawrence and lower Ottawa river systems in Québec and Ontario south to North Carolina (Gilbert and Lee 1980). It is closely associ- ated with upland areas such as the Allegheny, Catskill and Adirondack mountains and is not found in lowland coastal areas such as most of New Jersey and the Delaware Peninsula. *Reviewed and approved by COSEWIC 14 April 1994, report accepted, no status designation required. *Contribution number 59 of the Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario. 470 1996 CROSSMAN AND HOLM: STATUS OF THE CUTLIPS MINNOW 471 FiGureE |. Cutlips Minnow, Exoglossum maxillingua, 84 mm TL, ROM 9213. St. Lawrence River, Leeds Co., Ontario. Drawn by Anker Odum, from Scott and Crossman (1973), by permission. Canadian distribution records were obtained from the Service de l'Aménagement de la Faune, Ministére du Loisir, de la Chasse et de la Péche, Québec (MLCP) [now part of Ministére de l’Environment et de la Faune] in Montréal (240 records), the Canadian Museum of Nature, formerly National Museum of Canada (NMC) (19), the Royal Ontario Museum (ROM) (18),eMLCP in Trois Riviéres (14), literature (4), a database obtained from the Ontario Ministry of Natural Resources (OMNR; edited by Nick Mandrak, Department of Zoology, University of Toronto and Department of Ichthyology and Herpetology, ROM) (2), and the database of the University of Michigan Museum of Zoology (1). These records were checked for corre- spondence of locality description and coordinates, entered into a database, and plotted on 1:50 000 maps (all records in the database are listed in an unpublished Appendix on file with COSEWIC and available on request). Records were plotted on a 1:1 000 000 rough base map from which Figure 2 was prepared. In Canada, Exoglossum maxillingua is found in the St. Lawrence River and its tributaries, from a tributary of the Riviere St-Denis near Saint-Pascal, Québec (the northern most record known) upstream in the St. Lawrence River to Ivy Lea, Ontario. It is known from the lower Ottawa River system as far upstream as Riviere du Diable in the Riviere Rouge system (see Table 1, Figure 2). Nash (1908) stated that the species occurred in Lake Ontario. There are no voucher specimens to substantiate its presence in the Ontario portion of that lake but it has been reported from New York tributaries (Crossman and Van Meter 1979). Protection Exoglossum maxillingua is not legally protected in North America but it is listed as of Special Concern in North Carclina (Johnson 1987). In Canada, no specific legislation exists for the protection of this species buf laws which incidently protect its habitat include: the Ontario Lakes and Streams Improvement Act which prohibits the impoundment or diversion of watercourses which leads to siltation; the voluntary Land Stewardship I program of the Ontario Ministry of Agriculture and Food which is designed to reduce the erosion of agri- cultural lands and thus reduce siltation of habitat. In Québec, habitat is generally protected by the Environmental Quality Act, and may be protected by the Ecological Reserves Act if a species is “threat- ened with disappearance or extinction.” The species has been given little, or no attention, but could be given specific protection under provincial legislation (Endangered Species Act and law on faunic habitats) if required. Population Size and Trends The Cutlips Minnow is reported fairly common in its United States range (Gilbert and Lee 1980). It is particularly abundant in Pennsylvania (Cooper 1983) and New York (Smith 1985). In Ontario, the Cutlips Minnow has been collected on 14 occasions at 12 different sites (see Table 1). No voucher specimens are known before 1936. Exoglossum maxillingua was captured at six sites between 1936 and 1938 from the Delisle River, Lake St. Francis, the St. Lawrence River, and two small tributaries of the St. Lawrence River, Hoasic Creek and an unnamed creek. In 1943, bait dealers consid- ered it to be common in the St. Lawrence around Ivy Lea and in the Delisle River below Alexandria (Toner 1943). The Cutlips Minnow has not been captured recent- ly at any of the six sites in Ontario where it was taken in the 1930s. Attempts to capture it in Hoasic Creek (ROM Accessions 1276 and 5501) in 1967 and 1989 were unsuccessful. Surveys by the OMNR in the Ontario portion of the Delisle River system in 1973 and 1978 (ROM Accessions 2364 and 3765) also failed to capture the species. Attempts to cap- ture the species in the St. Lawrence River at Ivy Lea in 1967 (ROM Accession 1276) were also unsuc- cessful. Relatively intensive sampling was conducted by OMNR, ROM and NMC from the late 1960s to 472 THE CANADIAN FIELD-NATURALIST Exoglossum maxillingua o captured before 1950 e captured 1950 - 1989 0 25 50 ————— Kilometres FiGureE 2. Distribution of the Cutlips Minnow in Canada, Exoglossum maxillingua. A point may represent more than one capture at different sites within the area of the circle. Inset: North American distribution modified from Gilbert and Lee (1980). the mid 1980s (Mandrak and Crossman 1992). Despite the much larger scale of sampling, compared to that in the 1930s, the surveys captured the species at only six additional sites in the Raisin River sys- tem, Little Rideau Creek, and at one site in the St. Lawrence River. The unsuccessful capture attempts and the scarcity of records after 1970 suggest that populations in Ontario have declined in abundance from former levels. The species is more widely distributed in Québec where 284 collections have been made at 274 sites. The species was first recorded in 1935 from the upper section of the Ulverton River (Saint-Frangois river system) and in a few streams of the Nicolet system. At that time, Exoglossum maxillingua was consid- ered to be one of the rarest minnows in the Eastern Townships (Richardson 1935). In 1941, the species was also recorded from below the Rapides du Rocher Fendu, Lac Saint-Louis, and the Chateauguay, Chaudieére, and Saint-Denis river systems. Surveys conducted by MLCP in several major St. Lawrence River systems from 1960 to 1982 captured the species in many of the rivers where it was for- merly known and found it at numerous other sites in the St. Lawrence River of Québec (see Table 1). Distribution records indicate that Exoglossum max- illingua occurs at the greatest number of locations in the Riviere Chateauguay system (82 records, 1941 to 1989) and in the St. Lawrence River below Rapides de Lachine in the Montréal region (36 records, 1967 to 1989). In the Chateauguay it is particularly com- mon in the streams of the upper half of the drainage. It ranked 22nd out of 53 different species in relative frequency of occurrence in the collections in a 1975 to 1976 survey of the entire river system (Mongeau et al. 1979). It was captured in 1973 at 20 of 108 seining stations in a 25 km stretch of the St. Lawrence River below the Jacques-Cartier bridge at Montréal. At a few of these sites it was captured in considerable numbers (Massé and Mongeau 1976). Further upstream, immediately below the Rapides de Lachine, it was captured in 8 of 114 seining stations in 1977 (Mongeau et al. 1980). However, it is not currently considered common in the Montréal region (Dumont and Roy, personal communication, 1989). It is moderately abundant in the drainage of the Riviére Chaudiére (24 records, 1949-1977), Riviere du Chéne (20 records, 1971), and Riviére Nicolet (13 records, 1935-1977). In other river systems it is not as common. For example, it was captured at only RS EA RES A CD ce nD 1996 CROSSMAN AND HOLM: STATUS OF THE CUTLIPS MINNOW 473 TABLE |. Canadian rivers and river systems in which Exoglossum maxillingua has been captured 1935-1989. Year(s) of capture are followed by number of distribution records for each year in brackets. Names according to Répertoire Toponymique du Québec (1978. Editeur Officiel du Québec), and Gazetteer of Canada, Ontario (1988, Energy, Mines and Resources Canada. Ottawa). (DR = Division de Rencensement; NL = not listed in gazetteer, underlined records are from Ontario; the rest are from Québec) River system Year (Number of records) River system Year (Number of records) Riviere Saint-Denis system Ruisseau Saint-Andre 1971(1) Creek (NL) 1941(1) Creek (NL) 1971(1) Riviere Ouelle ; 1964(3),1968(4) Riviere Bayonne system Bras Saint-Nicholas 1975(2) Riviere Bayonne 1971(5) Riviere du Sud 1964(7) Ruisseau Bibeau 1971(1) Riviere Etchemin 1962(1) Riviere la Chaloupe 1971(1) Riviere Boyer system Channels above Lac Saint-Pierre Riviere Boyer 1971(5) Chenal aux Ours 1971(2) Riviere Boyer Sud 1971(2) Chenal du Nord 1971(1) Riviere Boyer Nord 1971(3) Riviére Richelieu 1965(1),1970(4) Riviere Chaudiere system Riviere L'Assomption system Riviere Chaudiére 1949(1),1976(3) Riviere de l'Achigan 1968(1) Riviere Saint- Victor 1965(8),1971(1) Lac Saint-Louis 1941(2),1942(1) Riviere Beaurivage 1964(9) Riviere Chateauguay system Riviere du Cing 1977(1) Riviere Chateauguay 1941(7),1942(2), Ruisseau Tring 1977(1) 1946(2),1960(3), 1961(1), Riviere du Chéne system (DR: Lotbiniere) 1963(1),1976(21) Riviére du Chéne 1971(8) Ruisseau Dewitt 1970(1),1976(2) Riviere Huron 1971(1) Creek (NL) 1977(1) Riviere Henri 1971(2) Riviere aux Anglais 1976(5) Bras d'Edmond 1971(1) Ruisseau Robson 1976(1) Riviere du Bois Claire 1971(2) Ruisseau Allen 1976(4) Riviere aux Chevreuils 1971(5) Riviére aux Outardes 1976(2) Creek (NL) 1971(1) Ruisseau Mitchel 1963(3),1976(3) Petite Riviere du Chéne system (DR: Lotbiniere) Creek (NL) 1976(4) Petite Riviere du Chéne 1 1982(2) Riviere Hinchinbrook 1961(1),1963(2),1976(4),1989(2) Riviere du Creux 1982(2) Riviere Trout 1941(1),1976(7) Ruisseau l'Espérance 1982(1) Ruisseau Oak 1967(1),1976(1) Creek (NL) 1982(1) Riviere du Chéne system (DR: Deux Montagnes) Riviere Sainte-Anne system Petite Riviere du Chéne 1970(1) Riviere Sainte-Anne unknown (between 1979-1980)(1) Lac des Deux Montagnes tributaries Riviere Charest unknown (between 1979-1980)(1) Riviere a la Raquette 1964(1) Riviere aux Orginaux system Riviere Rigaud 1964(1),1965(1),1966(1),1972(1) Riviere aux Orginaux 1982(3) Riviere du Nord system Ruisseau Santorio 1982(1) Lac Saint-Denis 1966(1) Riviere Gentilly system Lac Gémont 1967(1) Riviere Gentilly 1982(3) Lac La Riviere 1967(1) Riviere Gentilly Sud-Ouest 1982(1) Riviere Dalesville 1976(1) Riviere Bécancour 1964(1) Riviere de l'Ouest 1975(2),1976(2) Riviere Yamachiche 1972(6),1973(1) Riviere Rouge system Riviere Nicolet system Riviere du Diable 1968(1) Riviere Nicolet 1977(3) Little Rideau Creek 1978(1),1989(1) Riviere Nicolet Centre 1977(1) Riviere Delisle 1936(1),1938(1),1946(1),1970(1) Riviere Nicolet Nord-Est 1977(1) Riviere Baudette 1970(1) Riviere Nicolet Sud-Ouest 1935(1),1977(2) Lake St. Francis 1938(1) Riviere des Rosiers 1935(1) Raisin River system Ruisseau Francoeur 1935(1) Raisin River 1973(1) Riviére Bulstrode 1977(2) North Raisin River 1973(1),1989(1) Creek (NL) 1977(1) St. Lawrence creek (NL) 1938(1) Riviere Saint-Frangois system Hoasic Creek (Nash Creek) 1938(1) Riviere Saint-Franc¢ois St. Lawrence River (from a point downstream of unknown (between 1964 and 1974)(1) Riviere Richelieu upstream to Ivy Lea) Riviere Ulverton 1935(1) downstream of Riviere Richelieu 1971(1) Riviere Maskinongé 1967(1) Montréal-below Rapides de Lachine Riviere Yamaska system '41(2),'67(3),'72(1),'73(20),'77(8),'83(1),'89(1) Riviere Saint-David 1970(4) below Rapides du Rocher Fendu 1942(1),1980(5) Riviere du Chicot system z at Cardinal 1981(1) Riviere du Chicot 1971(5) at Ivy Lea 1936(2),1937(1) 474 four of 159 fishing stations in the Riviere Richelieu in 1970 (Mongeau 1979b). It is known from only two sites in the Saint-Francois river system (Mongeau and Legendre 1976; Richardson 1935) and from only four sites sampled between 1963 and 1975 in a tributary of the Riviere Yamaska (Mongeau 1979a). It was taken in 1980 from below the Rapides du Rocher Fendu but has not been cap- tured again in Lac Saint-Louis despite attempts in 1965 and 1968 (Mongeau and Massé 1976). Knowledge of the distribution of the Cutlips Minnow in Quebec increased dramatically. However, little sampling has been carried out since 1977 (P. Dumont and G. Roy, Ministére de l'Environnement et de la Faune, Montreal, Quebec; personal communication). Therefore, because most sites have been sampled only once the current status of most populations in Quebec, is undetermined. Table 1 summarizes our knowledge of where, in what year, and how frequently Exoglossum max- illingua has been captured. It summarizes 298 records representing 286 different sites in 38 river systems, 82 rivers and creeks in those systems, and three lakes. Two sites were sampled on three sepa- rate occasions and eight sites were sampled on two separate occasions. Information on sampling effort is not shown but investigators which have this informa- tion can use Table | to better assess the status of the species in each river system. Habitat In the United States, the Cutlips Minnow is usual- ly found in small to moderately sized clear streams, 4.6 to 15 m in width. It prefers quiet pools or chan- nels with gentle to moderately swift current. It has been found in depths of 0.15 to 1.2 m, in water with temperatures ranging from 0 to 26°C, and over firm bottoms of rubble, gravel, boulders, and cobbles. Instream cover such as large rocks, logs, vegetation, or overhanging banks is an important component of the habitat of this species (Hankinson 1922; Van Duzer 1939; Haase and Haase 1975; Cooper 1983; Pappantoniou 1983; Smith 1985). In Connecticut, total alkalinity ranged from 7 to 137 and hardness 22 to 184 both mg/l equivalent CaCo,. Bottom type (percent occurrence) consisted of stone or rubble (35%), gravel (30%), silt (13%), rock (9%), muck (9%), and sand (4%) (Whitworth et al. 1968). In Canada, Exoglossum maxillingua is found pri- marily in clear or tea-coloured rivers or creeks, on firm rocky bottoms, frequently mixed with one or more combinations of gravel, sand, and mud. In Québec, they are frequently found on hard clay and shale bottoms (Dumont and Roy, personal communi- cation). Aquatic vegetation is frequently present and water current varies from still to fast, but is most fre- quently described as slow. It has been found in water up to 26°C in June and July. Streams are usually THE CANADIAN FIELD-NATURALIST Vol. 110 small with a width of 1-20 m, but populations have also been found where the St. Lawrence River and its lake-like expansions are several kilometres wide. In the St. Lawrence, it was most commonly found at the lower end of rapids. The Cutlips Minnow is also known from lakes at elevations of up to 380 m in the Laurentians in Québec. General Biology There is no information on the biology of | Canadian populations, but several studies have been | carried out in New York and Pennsylvania. Age at | spawning is unknown but sizes are reported. In cen- tral New York, nest building by a 76 mm (three- inch) male was reported. However, spawning males usually averaged 102 to 140 mm (4 to 51% inches). Females were usually not over 76 mm (3 inches). Spawning season in the Susquehannah river system of central New York lasted approximately seven | weeks in one year (19307). It began around 25 May and lasted to the middle of July. Spawning occurred | in the daytime, peaking at mid-day and late after- noon at temperatures of 17 to 21.5°C. Length of spawning period varied from one to eight days | depending on the period during the spawning season (Van Duzer 1939). Spawning may occur later in Québec. Richardson (1935) indicated that specimens captured in the Eastern Townships in the latter half | of August and early September had well-developed ovaries and testes. In New York and Pennsylvania, females outnum- bered males in collections during July 1979 and in monthly collections from the winter of 1979/1980 to the winter of 1980/1981. The ratio of males to females ranged from 1:1.1 to 1:1.8. The lower num- ber of males in the collections was attributed to high- er male mortality caused by nest building and defense activities (Pappantoniou 1984a,b). In suitable areas, nests are often built very close to each other. Observations by Van Duzer (1939) indi- cated that the nest is built by a lone male. After spawning, it may be driven off by a larger male which may or may not continue nest building prior to spawning. The smaller male may attempt to con- tinue to add stones to the nest or spawn in the absence of the larger male. During spawning from one to 12 females may congregate on one nest, but only one pair spawns at a time. Age composition of populations in New York and Pennsylvania has been shown to differ considerably. Predominant age classes vary from I+ in eastern Pennsylvania (Pappantoniou 1984a) to ILI+ in south- eastern New York (Pappantoniou 1984b). The fol- lowing sizes at amnulus formation were determined for age classes in the Waccabuc River in eastern Pennsylvania: I, 37 to 52 mm; II, 63 to 81; Ill, 88 to 108; IV, 110 to 126 (Pappantoniou 1984b). Previous studies indicated that overlap in sizes occur between . 1996 year classes (Breder and Crawford 1922; Haase and Haase 1975). Maximum age is usually IV+, but specimens have been found to be V+ in a fertile stream in Pennsylvania (Haase and Haase 1975). Fecundity varied between 345 to 1177 eggs/female (X = 792 + 2 standard deviations of 281.3) in Waccabuc Creek in southeastern New York. Fecundity was considerably lower in the Titicus River, New York (x = 371.9 + 182.6). The fecundity of the female is apparently not directly correlated with size (Pappantoniou 1983). Some migration into deeper water may occur in extremely cold or wet winters (Miller 1964). Haase and Haase (1975) found that the numbers of Cutlips Minnows declined in fall collections. During the spawning season, Exoglossum maxillingua moves to suitable areas. Exoglossum maxillingua is a relatively specialized bottom feeder, but is apparently able to shift to other food resources when its preferred food is unavail- able. Several studies have been conducted on its diet in New York and Pennsylvania (Breder and Crawford 1922; Haase and Haase 1975: Johnson 1981; Johnson and Johnson 1982; Pappantoniou 1983; Pappantoniou et al. 1984a,b). The studies indi- cated that Exoglossum maxillingua consumes a vari- ety of aquatic invertebrates, but chironomids, tri- chopteran larvae, and oligochaetes are the most important items. Younger individuals consume a much larger proportion of chironomids, whereas older individuals favour larger food items such as tri- chopterans, oligochaetes and plecopterans. Breder and Crawford (1922) found, in addition to unidenti- fied insect remains (34%) and a large proportion of oligochaetes and polychaetes (30%), the gut con- tained diatoms and plant remains (15%) which they believed were being digested. Seasonal variation of the benthos was reflected in the diet in the Delaware River in Pennsylvania. When chironomid and tri- chopteran populations were low in September, they fed more on molluscs (Haase and Haase 1975). Successful reproduction of the Cutlips Minnow depends on availability of a specific type of habitat. Spawning habitat in the Susquehannah River system in central New York consisted of a firm rubble bottom overlain by an abundance of gravel. Depending on its size, the male selects flat stones with angular margins or thin edges that are 6 to 24 cm wide. Large flat rocks and submerged logs will offer protection during nest-building, spawning, and defence of eggs and fry (Van Duzer 1939). The Cutlips Minnow avoids the stronger current sought by other mound-building cyprinids such as the Creek Chub, Semotilus atromac- ulatus, and the River Chub, Nocomis micropogon (Miller 1964). Apparently, current must be sufficient- ly strong to insure a constant change of water and pre- vent excessive siltation, but gentle enough to prevent the removal of stones as small as 6 cm. CROSSMAN AND HOLM: STATUS OF THE CUTLIPS MINNOW 475 The Cutlips Minnow is sometimes called the eye- picker because it is known to deliberately attack the eyes of other species. Pappantoniou (1983) suggested that the incidence of eye-picking behaviour in Cutlips Minnows apparently increases with intra-specific den- sity. He suggested that in crowded conditions such as those which occur in pools in the summer, the Cutlips Minnow can increase access to limited resources by attacking the eyes of other species. Limiting Factors The presence of the Common Shiner, Luxilus cornu- tus, may adversely affect the reproduction of the Cutlips Minnow. The Common Shiner has been known to breed on the nest of the Cutlips Minnow while the Cutlips Minnow attempted to spawn. The presence and nervous motion of the shiners on the nest always lessened, and sometimes stopped, the spawning of the Cutlips Minnow. Attempts by the male Cutlips Minnow, occasionally assisted by the female, to drive the shiners off the nest were seldom successful (Van Duzer 1939). Miller (1964) noted, however, that Exoglossum maxillingua selects quiet channels not usually frequented by breeding shiners and chubs and spawned in late May, whereas the Common Shiner spawned in the first half of May. However, he also noted that the Common Shiner preferred the nest of the Cutlips Minnow over the nests of the chub, Nocomis, and Fallfish, Semotilus corporalis. Nothing is known about the susceptibility of the Cutlips Minnow to predation. Its sluggish nature may make it more vulnerable than other small fishes, but its habit of hiding under rocks and logs and in beds of aquatic vegetation would protect it. Temperature is probably an important limiting factor. It was noted (Pappantoniou et al. 1984b) that, in general, Cutlips Minnows in New York State were more long-lived and robust than Pennsylvania coun- terparts. This was attributed to the generally milder climatic conditions in southeastern New York State. Harsher climatic conditions, probably also adversely affects the life-span of Canadian populations. Scott and Crossman (1973) stated that the Cutlips Minnow prefers warm streams. This preference may limit its northward dispersal. The Cutlips Minnow is probably intolerant of tur- bidity and excessive siltation, both consequences of agricultural and urbanization activities (Scott and Crossman 1973). Flooding may increase mortality of eggs and fry if they are carried downstream beyond the nest during spawning and early develop- ment of the species. Flooding may have caused the reduced 1972 year-class of Cutlips Minnows in the Delaware River in eastern Pennsylvania. High water increases turbidity and scours the benthos which adversely affects food availability (Haase and Haase 1975). 476 Special Significance of the Species The Cutlips Minnow possesses some unique mor- phological and behaviourial characteristics. Its lips are unlike any other North American minnow. It is known to attack and consume the eyes of other species of fishes, a behaviour useful in experiments on the effec- tiveness of eye camouflage (Pappanto-niou 1983). This species is one of few minnows which demon- strate post-hatching care of fry (Smith 1991). Evaluation Canadian populations are at the northerly fringe of the range of the species. It has been found in 82 Trivers or creeks and three lakes in Ontario and Québec from Ivy Lea, Ontario to Saint-Pascal, Québec (Table 1). Although it has been captured at numerous sites in Québec from 1960 to 1980, most of these waters have not been surveyed in the last 15 years and there is little information on their present status. In Ontario and some areas of Québec, evi- dence indicates that it has declined. Although the Cutlips Minnow has been found in numerous river systems, it has shown decline in some. Possible rea- sons for this decline may include both natural and cultural factors such as species competition, preda- tion, flooding, turbidity, siltation, cold temperatures, and over-harvest by bait fisherman. The species, never abundant in Canada, is rarer in Ontario than Québec. Although surveys in Québec have been limited since 1977 there is no evidence to indicate the species is in decline there (M. Huot, Ministére de l'Environnement et de la Faune, Québec, Québec; personal communication). Although surveys have been limited since 1977 there is evidence to suggest the species is in decline in Ontario mainly due to habitat degradation and compe- tition with increasing populations of Common Shiners. Acknowledgments Financial support was provided by the Royal Ontario Museum, the World Wildlife Fund and the Environmental Youth Corps grant programme. Frederique Arnaud, David Boehm, Lisa Dorman, Kira Dunham, Colleen Fennesey, William Ramshaw, and Marty Rouse, assisted with field work. Frederique Arnaud, Kira Dunham, and William Ramshaw assisted with verification and summary of distribution records. Distribution maps were drawn by William Ramshaw. Pierre Dumont and Gilles Roy (MLCP, Montréal) and Yves Mailhot (MLCP, Trois Riviéres) provided information and distribu- tion records for Exoglossum maxillingua in Québec. Literature Cited Breder, C. M., Jr., and D. R. Crawford. 1922. The food of certain minnows. A study of of the seasonal dietary cycle of six cyprinoids with especial reference to fish culture. Zoologica 2: 287-327. Buynak, G. L., and H. W. Mohr, Jr. 1980. Larval devel- opment of Stoneroller, Cutlips Minnow, and River Chub THE CANADIAN FIELD-NATURALIST Vol. 110 with diagnostic keys, including four additional cyprinids. The Progressive Fish-Culturalist 42(3): 127-135. Coburn, M. M., and T. M. Cavender. 1992. Interrela- tionships of North American cyprinid fishes. Pages 328-373 in Systematics, historical ecology, & North American freshwater fishes. Edited by R. L. Mayden. Stanford University Press, California. Crossman, E. J., and H. D. Van Meter. 1979. Annotated list of the fishes of the Lake Ontario Watershed. Great Lakes Fishery Commission Technical Report. Number 36. Cooper, E. L. 1983. Fishes of Pennsylvania and the north- eastern United States. The Pennsylvania State University Press. Fuiman, L. A., and J. J. Loos. 1978. Morphological changes during the larval development of the Cutlips Minnow, Exoglossum maxillingua. Transactions of the American Fisheries Society 107(4): 605-612. Gilbert, C. R., and D. S. Lee. 1980. Exoglossum max- illingua (Lesueur), Cutlips minnow. Page 58 in Atlas of North American Freshwater fishes. Edited by D. S. Lee, C.R. Gilbert, C. H. Hocutt, R. A. Jenkins.-D. E: McAllister, and J. R. Stauffer Jr. North Carolina Biological History Survey, Publication Number 1980-12. Haase, R., and B. L. Haase. 1975. Feeding ecology of the cutlips minnow, Exoglossum maxillingua, in the Delaware River at Bushkill, Pennsylvania. Proceedings of the Pennsylvania Academy of Sciences 49: 67—72. Hankinson, T. L. 1922. Nest of the cut-lips minnow, Exoglossum maxillingua (LeSueur). Copeia 102: 1-3. Johnson, J. E. 1987. Protected fishes of the United States and Canada. American Fisheries Society, Bethesda, Maryland. Johnson, J. H. 1981. The summer diet of the cutlips min- now, Exoglossum maxillingua, in a central New York stream. Copeia 1981: 484-487. Johnson, J. H., and E. Z. Johnson. 1982. Observations on the eye-picking behaviour of the cutlips minnow, Exoglossum maxillingua. Copeia 1982(3): 711-712. Mandrak, N. E., and E. J. Crossman. 1992. A checklist of Ontario freshwater fishes annotated with distribution maps. Royal Ontario Museum Life Sciences Publication. Massé, G., and J.-R. Mongeau. 1976. Influence de la navi- gation maritime sur la répartition géographique et l'abon- dance relative des poissons du fleuve Saint-Laurent entre Longueuil et Sorel. Rapport Technique. Service de l'Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Miller, R. J. 1964. Behavior and ecology of some North American cyprinid fishes. American Midland Naturalist 72: 313-357. Mongeau, J.-R. 1979a. Les poissons du bassin de drainage de la Riviére Yamaska, 1963 a 1975. Rapport Technique. Service de l'Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche, Québec. Mongeau, J.-R. 1979b. Dossiers des poissons du bassin versant de la Baie Missisquoi et de la Riviére Richelieu, 1954 4 1977. Service de l'Aménagement de la Faune, Ministére du Tourisme. de la Chasse et de la Péche, Québec. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1979. Les Poissons du bassin de drainage de: la riviére Chateauguay, leur milieu naturel, leur répartition géo- 1996 graphique et leur abondance relative. Rapport Technique. Service de l'Aménagement de la Faune. Ministére du Tourisme. de la Chasse et de la Péche, Québec. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1980. La répartition géographique des poissons. les ensemence- ments, la péche sportive et commerciale, les frayéres et la bathymétrie du fleuve Saint-Laurent dans le Bassin de La Prairie et les Rapides de Lachine. Rapport Technique No. 06-29. Service de l'Aménagement de la Faune, Ministére du Tourisme, de la Chasse et de la Péche. Québec. Mongeau, J.-R., and V. Legendre. 1976. Les ressources fauniques du bassin inférieur de la Riviére Saint- Francois: Evolution des populations en dix ans, 1965- 1974. Rapport Technique. Service de l'Aménagement de la Faune. Ministére du Tourisme. de la Chasse et de la Péche, Québec. Mongeau, J.-R., and G. Massé. 1976. Les poissons de la région de Montréal. la péche sportive et commerciale. les ensemencements. les frayéres. la contamination par le mercure et les PCB. Service de l'Aménagement de la Faune, Ministére du Tourisme. de la Chasse et de la Péche. Québec. Nash, C. W. 1908. Manual of vertebrates of Ontario. Legislative Asssembly of Ontario, Toronto, Ontario. Pappantoniou, A. 1983. Aspects of the life history of the cut- lips minnow Exoglossum maxillingua (Pisces: Cyprinidae). Ph.D. thesis. Fordham University, New York. Pappantoniou, A., G. Dale, and R. E. Schmidt. 1984a. Aspects of the Life History of the Cutlips Minnow, CROSSMAN AND HOLM: STATUS OF THE CUTLIPS MINNOW A477 Exoglossum maxillingua, from two eastern Pennsylvania streams. Journal of Freshwater Ecology 2(5): 449-457. Pappantoniou, A., R. E. Schmidt, and G. Dale. 19845. Ecology and life history of the Cutlips Minnow, Exoglossum maxillingua, from a southeastem New York stream. Northeastern Environmental Science 3(3): 75-79. Richardson, L. R. 1935. The fresh-water fishes of south- eastern Quebec. PhD thesis. Department of Zoology. McGill University. Montreal. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. Ottawa, Ontario. Smith, C. L. 1985. The inland fishes of New York State. N. Y. State Department of Environmental Conservation. Smith, R. J. F. 1991. Chapter 18. Social behaviour, hom- ing and migration. Pages 507-529 in Cyprinid fishes. Systematics. biology and exploitation. Edited by I. J. Winfield and J. S. Nelson. Chapman and Hall, London. Toner, G. C. 1943. Ecological and geographical distribu- tion of fishes in eastern Ontario. M. A. thesis. University of Toronto. Toronto, Ontario. Van Duzer, E. M. 1939. Observations on the breeding habits of the cutlips minnow. Exoglossum maxillingua. Copeia 1939: 63-75. Whitworth, W. R., P. L. Berrien, and W. T. Keller. 1968. Freshwater fishes of Connecticut. State Geological and Natural History Survey of Connecticut, Bulletin 101. Accepted 13 March 1996 The Status of the Lake Chubsucker, Erimyzon sucetta, in Canada*t NICHOLAS E. MANDRAK' and E. J. CROSSMAN Center for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 'Present address: Department of Biological Sciences, Fort Hays State University, Hays, Kansas 67661 Mandrak, Nicolas E., and E. J. Crossman. 1996. The status of the Lake Chubsucker, Erimyzon sucetta, in Canada. Canadian Field-Naturalist 1 10(3): 478-482. The Lake Chubsucker, Erimyzon sucetta, 1s a robust, moderately deep-bodied member of the sucker family with a wide head, blunt snout and small, slightly inferior, protrusible mouth. It has a disjunct North American distribution, and in Canada, is found only in southwestern Ontario. This species is declining in many parts of its range. The most significant threats to the Lake Chubsucker in Canada are the drainage or siltation of its critical habitat. In Canada, it prefers a habitat susceptible to human perturbation, has only been collected in low numbers at few localities, and is at its northeastern range limit. It has been classified as Vulnerable in Canada. Le sucet de lac, Erimyzon sucetta, est un membre de la famille des meuniers, robuste et relativement trapu, avec une téte large, un nez émoussé et une petite bouche protractile, l}égerement sous la téte (inférieure). Sa distribution est éparse en Amérique du Nord et, au Canada, on ne le retrouve que dans le sud-ouest de |’Ontario. L’espéce est en déclin dans la majeure partie de son aire de répartition. Au Canada, les pires menaces pour le sucet de lac sont le drainage et la sédimen- tation dans son habitat principal. Au Canada, il préfere les habitats vulnérables ot il y a risque de perturbations humaines; on n’en a récolté que de petits nombres dans quelques localités et il se trouve a la limite nord-est de son aire de répartition. Par conséquent, jusqu’a ce qu’un échantillonnage adéquat soit entrepris pour évaluer la situation des populations canadi- ennes, on recommande que le sucet de lac ait le statut de “menacé” au Canada. Key Words: Lake Chubsucker, sucet de lac, Erimyzon sucetta, Catostomidae, vulnerable, Ontario. Erimyzon sucetta (Lacepede, 1803), the Lake Chubsucker, belongs to a genus of suckers (family Catostomidae) which includes only three species. It is the only representative of this genus to occur in Canada, and is found only in southwestern Ontario. This report summarizes our current knowledge of the distribution and status of the species in Canada. The Lake Chubsucker is a robust, slightly com- pressed fish with a moderately deep-arched back, thick caudal peduncle and wide head with a blunt snout (Figure 1). It has a small, slightly inferior, suc- torial, protrusible mouth (Scott and Crossman 1973). The dorsal surface of its body 1s deep olive to green- ish-bronze, the ventral surface is green-yellow to yellow-white. Scales above lateral line are dark- edged giving a cross-hatched appearance. A lateral band, if present, has been documented as continuous in adults (Pflieger 1975; Trautman 1981; Rutherford et al. 1985; Robison and Buchanan 1988), or broken into blotches or transerve bands (Anonymous 1962; Scott and Crossman 1973; Douglas 1974; Page and Burr 1991). Preserved adult specimens from Ontario exhibit both continuous and blotched lateral bands when present. Adult size may reach a maximum of 410 mm total length (TL; Page and Burr 1991), although Ontario specimens seldom exceed 254 mm TL (Scott and Crossman 1973). A dorsal fin with a short base, fewer than 20 rays and without a rounded or pointed anterior lobe differentiates the genus Erimyzon from the genera Carpiodes, Cycleptus and Ictiobus. Erimyzon differs from other genera of Catostomidae by the presence of an oblique mouth and absence of a lateral line. In Canada, the creek chubsucker, Erimyzon oblongus, has been reported only in New Brunswick, but not since 1873 (Cox 1896). Cox (1896) erroneously listed this record as E. sucetta based on Adams (1873) who listed it as Moxostomus oblongus (=Erimyzon oblongus). Scott and Crossman (1959) concluded that “it seems high- ly unlikely that it [Erimyzon oblongus] ever occurred in New Brunswick”. It is present in the American drainage of the Great Lakes in the tribu- taries of southwestern Lake Erie and southeastern Lake Ontario; therefore, all Erimyzon specimens collected in Ontario should be examined closely for Erimyzon oblongus. There are differences in the lit- erature in the description of the distinguishing char- acteristics between the largely sympatric Erimyzon sucetta and Erimyzon oblongus (Cook 1959; Scott and Crossman 1973; Douglas 1974; Smith 1979; Trautman 1981; Rutherford et al. 1985; Robison and Buchanan 1988; Page and Burr 1991), probably as “Reviewed and Approved by COSEWIC 14 April 1994, status assigned — Vulnerable yContribution number 57 of the Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario 478 1996 MANDRAK AND CROSSMAN: STATUS OF THE LAKE CHUBSUCKER 479 Ficure 1. The Lake Chubsucker, Erimyzon sucetta. Reprinted with permission from Scott and Crossman 1973. the result of the natural variation of these characters over the geographic ranges of these species. The eye is generally larger in Erimyzon sucetta (eye diame- ter ranges from 3.4 to 6.0 times into head length (HL); 3.2 to 5.2 with 3.0 and 7.2 extremes in the 55 preserved Ontario specimens examined) than in Erimyzon oblongus (5.5 to 6.2 times into HL). Erimyzon sucetta generally has a lower lateral line scale count (32 to 40; 32 to 40 in Ontario specimens examined) and higher dorsal ray count (10 to 13; 10 to 13 in Ontario specimens examined) than Erimyzon oblongus (37 to 47; 8 to 11). Erimyzon oblongus is more slender (body depth 3.2 to 4.2 times into standard length (SL)) than Erimyzon sucetta (2.4 to 3.2 into SL; 3.0 to 4.0 in Ontario specimens examined). Prior to spawning, male Lake Chubsuckers develop a falcate anal fin and 3 tuber- cules (a reduced fourth tubercule is occassionally present; Douglas 1974) on each side of the snout (Robison and Buchanan 1988). Distribution Erimyzon sucetta exhibits a disjunct North American distribution (see inset map, Figure 2). A southern element is centred around the Gulf States and extends northward from the Arkansas River through the Mississippi Valley to southern Illinois; east of the Mississippi River to the Atlantic Seaboard northward to southern Virginia; and, west of the Mississippi River to eastern Texas. A northern ele- ment encompasses a southern Great Lakes drainage. Distribution is fragmented between the two main areas of distribution. Trautman (1981) hypothesized that this fragmentation was the result of northeastern range expansion during the warm Hypsithermal Period (ca. 7000 to 5000 years before present), and subsequent range contraction and fragmentation dur- ing the wane of this Period. In recent times, the dis- tribution of Erimyzon sucetta appears to be decreas- ing in many states, and it is now considered extirpat- ed in Iowa and New York (Becker 1983; Smith 1985). However, it was first recorded in Oklahoma in 1982 (Rutherford et al. 1985). In Canada, Erimyzon sucetta has been collected only in the drainages of the Niagara River, and lakes Erie, St. Clair and Huron in southwestern Ontario (Figure 2, all records are listed in an unpublished Appendix to the Status Report on file with COSEWIC Canadian Wildlife Service, Ottawa, in Ontario). Records by Small (1883) for “Hartwell’s locks”, Ottawa, and by Halkett (1913) for the St. Lawrence River and tributaries are probably erro- neous. Hubbs and Brown (1929) felt that the Lake Chubsucker was probably present in Ontario, although none had been collected. Scott (1952) reported that this species was first captured in Ontario in 1949, and suggested that its presence was the result of recent natural migration northward. Mandrak (1990) stated that Erimyzon sucetta dis- persed through glacial waterbodies into the Lower Peninsula of Michigan and along the south shore of Lake Ontario during the late Pleistocene. Through these, and adjacent glacial waterbodies, the Lake Chubsucker would have had the opportunity to dis- perse into the lower Great Lakes and subsequently into Ontario. Mandrak (1990) suggested that it was not collected prior to 1949 due to low population numbers and the difficulty of sampling its preferred habitat; therefore, he concluded that the species should be considered native to Ontario. Despite more recent sampling [Appendices containing all records for the species are on file with COSEWIC and avail- able on request (Canadian Wildlife Service, Ottawa, Ontario KOA 3MO)], the Lake Chubsucker was col- lected only prior to 1970 in Tee Creek, a Niagara River tributary, in Jeanettes Creek, a Thames River tributary, and at the mouth of Big Creek. Despite previous sampling, it has been collected only since 1970 in three tributaries of Big Creek, and the Old Ausable Channel, a Lake Huron tributary. The species has been collected before and after 1970 in Lake Erie at Long Point, Rondeau Harbour, Point 480 Pelee and in Lake St. Clair. No specimens have been collected since 1983. Protection Erimyzon sucetta receives no special protection in Canada (but see “Habitat” section). It is legally pro- tected in New York and Ohio, and is of “Special Concern” in Indiana, Kentucky, Missouri and Wisconsin (Johnson 1987). Population Size and Trend No attempt has been made to determine the popu- lation size of Erimyzon sucetta in Ontario. Only 30 collections of the Lake Chubsucker have been made in Ontario, and most of these collections yielded fewer than five specimens. Therefore, it is difficult to assess population sizes and trends. The sampling data suggest that populations are stable at Point Pelee where the Lake Chubsucker had been collected in 1949, 1968, 1972 and 1983, and in Rondeau Bay where it had been collected in 1953, 1954, 1955, 1963, 1975 and 1983. Trends in the remaining popu- lations in Ontario are not known. Habitat The preferred habitat of Erimyzon sucetta is clear, still, well-vegetated waters, such as those provided by backwaters, bayous, drainage ditches, floodplain lakes, marshes, oxbows, sloughs and wetlands, with substrates of gravel, sand and silt mixed with organic debris (Douglas 1974; Pflieger 1975; Smith 1979; Trautman 1981; Burr and Warren 1986; Robison and Buchanan 1988). In Ontario, the Lake Chubsucker has been captured primarily in heavily vegetated, stagnant bays, channels, ponds and swamps with low turbidity and substrates of clay, silt, sand and organic debris. In 1974, a single specimen was collected in Lyndecock Creek, a Big Creek tributary, in a habitat described as being moderately flowing with abundant floating vegetation over a clay and silt substrate. It is likely that the number and quality of areas containing the critical habitat of the Lake Chubsucker are decreasing, as the result of the draining of wetlands and increases in siltation associated with agricultural practises in southwestern Ontario. Federal and provincial legislation exists that pro- tects the habitat of Erimyzon sucetta. The National Parks Act protects habitat in Point Pelee National Park, and the Ontario Provincal Parks Act protects habitat in Long Point, Pinery and Rondeau provin- cial parks. The flora, fauna and its habitat in the lat- ter two parks are protected by the Ontario Wilderness Areas Act. The Ontario Lakes and Streams Improvement Act prohibits the impound- ment or diversion of watercourses which leads to sil- tation. The voluntary Land Stewardship II program of the Ontario Ministry of Agriculture and Food is designed to reduce the erosion of agricultural lands. THE CANADIAN FIELD-NATURALIST Vol. 110 This program has the potential to slow the degrada- tion of critical habitat by reducing siltation. General Biology i In North America, the annual spawning season of | Erimyzon sucetta varies within the period March and | July (Cooper 1983). Examination of the gonads of | several preserved specimens from Ontario indicated | that Erimyzon sucetta likely spawns between late | April and June in Ontario. At spawning time, the | Lake Chubsucker moves to marshes to spawn (Loftus and Kushlan 1987). Depending on the size of | the female, between 3000 and 20 000 eggs (Bennett and Childers 1966) are broadcast over submerged vegetation and hatch at water temperatures between 22°C and 29°C (Cooper 1983). The Lake Chubsucker is omnivorous and its diet consists of plankton, small crustaceans and molluscs, aquatic insects, and filamentous algae and other | plant matter which sometimes comprise over 70% of | its diet (Cooper 1983; Robison and Buchanan 1988). Erimyzon sucetta 1s tolerant of low O, levels (Odum and Coldwell 1955; Cooper 1983) and intolerant of siltation, turbidity and high stream gradients | (Trautman 1981). Limiting Factors Siltation, increased turbidity and loss of critical | habitat are factors attributed to the decline of the | Lake Chubsucker throughout its distribution (Lee et. al 1980; Trautman 1981; Burr and Warren 1986). | Draining of wetlands and siltation appear to be the | leading causes of significant loss of critical habitat in Canada. Special Significance of the Species Erimyzon sucetta is declining throughout most of its North American range and is the only representa- tive of its genus presently known in Canada. It can © be concluded that the behavioural, ecological and | genetic diversity represented by the genus Erimyzon | is in jeopardy in Canada. In Canada, the drainage or siltation of critical habi- tat appear to be the most significant threats to | Erimyzon sucetta. Population declines will occur in | areas where the Lake Chubsucker is still present |) ) unless further drainage or siltation of critical habitat is | prevented. If further degradation of critical habitat is | | prevented, extant populations should become stable. | | Evaluation Populations of Erimyzon sucetta at Point Pelee | and Rondeau Harbour on Lake Erie appear to be sta- | ble. Trends in the remaining populations cannot be | assessed due to lack of adequate resampling. In | Ontario, it has only been collected in low numbers at |} few locations, it prefers a habitat highly susceptible | to human perturbation, and the populations represent | 1996 MANDRAK AND CROSSMAN: STATUS OF THE LAKE CHUBSUCKER 481 Erimyzon sucetta o captured before 1970 ecaptured 1970 - 1983 te) 25 50 Kilometres FIGURE 2. Canadian distribution of Erimyzon sucetta. Inset map: North American distribution of the Lake Chubsucker (modified from Lee et al. 1980). Note: this species has not been recorded in Canada since 1983. ' the northeastern range limit of the species. Therefore, until adequate sampling of these popula- tions is undertaken to determine their stability, it is _ recommended that the Lake Chubsucker be classi- | fied as Vulnerable in Canada. _ The choice of status is effected by differences in population trends in protected habitats (e.g., Point © Pelee and Rondeau parks) and unprotected habitats _ (e.g., riverine habitats). It is obvious that the popula- tions in the protected habitats are, by definition, Vulnerable. The unprotected habitats, for which there is poor information, account for a significant | portion of the Canadian range of the Lake _ Chubsucker. As the result of human actions, popula- | tions in unprotected habitats may be threatened with imminent extirpation which suggests a status of Endangered. Additional information may lead to a reevaluation of the status assigned. Acknowledgments E. Holm, D. E. McAllister, J. S. Nelson and A. _ Peden provided useful comments on the manuscript. Financial support for this project was provided by an Environmental Youth Corps grant to EJC, the Royal Ontario Museum, and a World Wildlife Fund grant to EJC. F. Arnaud, K. Dunham, E. Holm and W. Ramshaw assisted with the resampling of some cap- ture sites, and the verification and summary of distri- bution records. Distribution maps were drawn by W. Ramshaw. Additional distributional data were pro- vided by the Canadian Museum of Nature Ottawa, and the University of Michigan Museum of Zoology, Ann Arbor, Michigan. Literature Cited Adams, A. L. 1873. Field and forest rambles, with notes and observations on the natural history of eastern Canada. Henry S. King and Company, London. 333 pages. Anonymous. 1962. Some North Carolina freshwater fishes. North Carolina Widlife Resources Commisssion, Raleigh. Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison. 1052 pages. Bennett, G. W., and W. F. Childers. 1966. The lake chub- sucker as a forage species. Progressive Fish-Culturist 28: 89-92. Burr, B. M., and M. L. Warren, Jr. 1986. Distributional atlas of Kentucky fishes. Kentucky Nature Preserves Commission, Scientific and Technical Series Number 4. 398 pages. 482 Cook, F. A. 1959. Freshwater fishes in Mississippi. Mississippi Game and Fish Commission, Jackson. 239 pages. Cooper, E. L. 1983. Fishes of Pennsylvania and the north- eastern United States. The Pennsylvania University Press, University Park. 243 pages. Cox, P. 1896. Catalogue of the marine and freshwater fish- es of New Brunswick. Bulletin of the Natural History Society 13: 62-75. Douglas, N. H. 1974. Freshwater fishes of Louisiana. Louisiana Wildlife and Fisheries Commission, Baton Rouge. 443 pages. Halkett, A. 1913. Checklist of the fishes of the Dominion of Canada and Newfoundland. King’s Printer, Ottawa. 138 pages. Johnson, J. E. 1987. Protected fishes of United States and Canada. American Fisheries Society, Bethesda. 42 pages. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. 1980. Atlas of North American freshwater fishes. North Carolina Biological Survey Publication 1980-12. North Carolina State Museum of Natural History, Chapel Hill. 867 pages. Loftus, W. F., and J. A. Kushlan. 1987. Freshwater fishes of southern Florida. Bulletin of the Florida State Museum of Biological Sciences 31: 147-344. Mandrak, N. E. 1990. The zoogeography of Ontario fresh- water fishes. MSc. thesis. University of Toronto, Toronto. 190 pages. Odum, H. T., and D. K. Coldwell. 1955. Fish respiration in the natural oxygen gradient of an anaerobic spring in Florida. Copeia 1955: 104-106. THE CANADIAN FIELD-NATURALIST Vol. 110 Page, L. M., and B. M. Burr. 1991. A field guide to fresh- water fishes: North America, north of Mexico. Houghton Mifflin Company, Boston. 432 pages. Pflieger, W. L. 1975. The fishes of Missouri. Missouri Department of Conservation, Jefferson City. 343 pages. Robison, H. W., and T. M. Buchanan. 1988. Fishes of Arkansas. University of Arkansas Press. 536 pages. Rutherford, D. A., A. A. Echelle, and O. E. Maughan. 1985. An addition to the fish fauna of Oklahoma: Erimyzon sucetta (Catostomidae). Southwestern Naturalist 30: 305-306. Scott, W. B. 1952. Records of the western lake chubsucker, Erimyzon sucetta kennerleyi, from Ontario. Copeia 1952: 203. Scott, W. B., and E. J. Crossman. 1959. The freshwater fishes of New Brunswick: a checklist with distributional notes. Contributions of the Royal Ontario Museum Division of Zoology and Paleontology 51. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Bulletin of the Fisheries Research Board of Canada 184. 966 pages. Small, H. B. 1883. Fishes of the Ottawa district. Transactions of the Ottawa Field Naturalists Club 4, 1882-1885: 31-47. Smith, C. L. 1985. The inland fishes of New York state. New York Department of Environmental Conservation. 522 pages. Smith, P. W. 1979. The fishes of Illinois. University of Illinois Press, Urbana. 314 pages. Trautman, M. B. 1981. The fishes of Ohio. Ohio State University Press, Columbus. 683 pages. Accepted 13 March 1996 The Status of the Blackchin Shiner, Notropis heterodon, in Canada* J. HOUSTON 374 Fireside Drive, R.R. 1 Woodlawn, Ontario KOA 3M0O Houston, J. 1996. The Status of the Blackchin Shiner, Notropis heterodon, in Canada. Canadian Field-Naturalist 110(3): 483-488. The native range of the Blackchin Shiner, Notropis heterodon, in Canada extends westward from southwestern Quebec to southwestern Manitoba. Little information is available on the biology and ecology of the species and the present status in Quebec is uncertain as recent survey data are unavailable. The species is apparently secure in Ontario, but populations in southwestern Ontario should be monitored as streams in this area are subject to disturbances which could alter water quali- ty as the species appears to be particularly sensitive to turbidity and siltation. Although only recently added to the fauna of Manitoba, the species appears to be native to the province and more widespread and abundant than previously thought. Au Canada, I’aire de répartition du menton noir, Notropis heterodon, s’étend du sud-ouest du Québec jusqu’au sud-ouest du Manitoba. On dispose de peu d’information sur la biologie et l’écologie de l’espéce, et la situation actuelle au Québec est mal connue vu I|’absence de données récentes de relevé. L’espéce semble étre en sécurité en Ontario, mais les popula- tions du sud-ouest de la province devraient étre surveillées a cause des perturbations qui pourraient affecter la qualité de Veau dans certains couloirs de cette région; ce méné semble particuliérement vulnérable a la turbidité et a l’envasement. On a longtemps cru que cette espéce était une addition a la faune du Manitoba mais, on la considére maintenant, plutot comme indigéne de cette province ou elle est plus répandue et abondante qu'on croyait. Key Words: Blackchin Shiner, Notropis heterodon, menton noir, Cyprinidae, minnows, Ontario, Quebec, Manitoba. The Blackchin Shiner, Notropis heterodon (Cope, 1865), is a small cyprinid closely resembling the _ Pugnose Shiner, Notropis anogenus. The Canadian _ distribution was formerly thought to be restricted to Ontario and Quebec (Scott and Crossman 1973), but is now known to include southern Manitoba (Stewart _ et al. 1985), where it has been listed as being of spe- cial concern (Johnson 1987). _ Description Notropis heterodon (Figure 1) is a small minnow averaging about 64 mm in length. The body is slen- _ der and terete straw coloured above and white to sil- very below. The scales on the back have darkened edges and there is a prominent black lateral line from _ the tip of the snout to the caudal fin. The black pig- ment extends into the chin, accounting for the ver- nacular name and the black midlateral stripe often has a zig-zag appearance (E. J. Crossman, and E. Holm, Royal Ontario Museum, Toronto, Ontario; personal communication). Spawning fish may devel- op a pale yellow tinge to the ventral surface (Scott - and Crossman 1973) and breeding males develop tiny tubercles on the dorsal surface of the head and the upper surface of the pectoral rays (Trautman 1981). The fins are translucent, the dorsal is sharply _ pointed and situated above the insertion of the pelvics, the caudal is distinctly forked (see Scott and Crossman 1973; Smith 1979; Trautman 1981; Smith 1985) for definitive descriptions). The species closely resembles the Pugnose Shiner from which it can be distinguished by the silvery peritoneum and the smaller, upturned mouth of Notropis anogenus (Scott and Crossman 1973; Smith 1979). Blackchin Shiners also superficially resemble the Weed Shiner (Notropis texanus), differing in the pointed dorsal fin and larger eye, 2 pharygeal teeth on the lesser row on each side and an included lower jaw; the Mimic Shiner (Notropis volucellus), where the lateral band is very pale, and does not extend for- ward to the tip of the snout (Mimic Shiners also have dark pigment at the base of the anal fin), and a pha- ryngeal tooth count of 0,4-4,0; the Blacknose Shiner (Notropis heterolepis), where the lateral band is not as distinct and the pigmentation does not extend to the lower jaw and has 0,4-4,0 pharyngeal teeth and no mid-dorsal black band (Smith 1979). The species may also be confused with the Bridle Shiner (Notropis bifrenatus) to which it may be closely related (Gilbert 1980), but the ranges overlap only in Quebec and eastern Ontario. Distribution The range of the Blackchin Shiner is limited to the Great Lakes Basin and the extreme upper Mississippi basin of the northern United States and southern Canada (Figure 2). In the U.S. the species ranges from eastern North Dakota east to New York, probably no further south than 40°N (Scott and Crossman 1973; Gilbert 1980). *Reviewed and approved by COSEWIC 14 April 1994, report accepted, no status designation required. 483 484 THE CANADIAN FIELD-NATURALIST Vol. 110 FiGure |. Blackchin Shiner, Notropis heterodon, [(Gravid female, 50.5 mm (C22 NMC68- 0206, 21 July 1968) drawing by Sally J. Gadd, courtesy Don E. McAllister, Canadian Museum of Nature]. In Canada, the species is most widely distributed and abundant in Ontario, although the range more or less extends from the Upper St. Lawrence near Trois-Rivieres, Quebec, west to southern Manitoba (Figure 2; Appendices of all records are on file and -available on request from COSEWIC). The species is limited to the extreme southwest- ern region of Quebec (Figure 2) where it is known from tributaries of the Ottawa River in Hull, Pontiac, Papineau and Gatineau counties, and of the St. Lawrence River as far downstream as Trois- Rivieres (Mongeau et al. 1979; Bergeron and Brousseau 1983). Blackchin Shiners have also been collected from the south shore of the St. Lawrence from the Richelieu and Chateauguay watersheds (Mongeau et al. 1974). No Ottawa River collections north of Pontiac County are known, although the species has been recorded from streams on the Ontario side of the river north to at least Temagami (Scott and Crossman 1973; ROM 34900). This per- ceived absence may be a sampling artifact, since suitable habitat does exist; surveys in the area should be on the alert for this species in any collec- tions made from local streams. Current Ontario records (Scott and Crossman 1973; Appendices of all records are on file and available on request from COSEWIC) suggest a dis- junct distribution (Figure 2). The species occurs in the Ottawa River watershed north to Pembroke and throughout southwestern Ontario in lakes Ontario, Erie and Huron and their tributary streams, north to Sault Ste Marie. Blackchin Shiners have only recently been recorded from Canadian Lake Superior tributaries, Crossman and Holm (personal communication) indicate a collection from the Goulais River (46°45'N, 84°16"W) in 1990 (OMNR 5842). The species has been recorded from several streams in Michigan tributary to Lake Michigan (Scott and Crossman 1973). It is also known in Ontario from the Lake of the Woods watershed, the Rainy River watershed in Quetico Park and the English and Wobigoon river watersheJs of the Kenora District of northwestern Ontario (so. nan and McAllister 1986; Stewart 1988). Th: records from the Hudson Bay, Moose Kiver drainage (Ontario Ministry of Natural Resources E2203 E36): Present knowledge of the Blackchin Shiner also suggests a disjunct distribution in Manitoba (Figure 2), where it was first collected in 1973 (ROM 29840) from Oak Creek, near the junction of the Souris and Assiniboine rivers (Stewart et al. 1985). Further records followed in 1982 from the Assiniboine watershed in Spruce Woods Provincial Park (see Stewart et al. 1985) and from Dauphin Lake (Babaluk and Harbicht 1984). Notropis het- erodon also occurs in the Winnipeg River in Manitoba and in the Lake of the Woods Watershed, in Falcon Lake Manitoba (Stewart 1988). Protection Blackchin Shiners are not subject to any specific protection in Canada. The species has been consid- ered to be of “special concern” in Manitoba (Johnson 1987), but has not as yet been considered for protec- tion under the provincial Endangered Species Act. In Quebec the species has been given little, or no atten- tion, but could be given specific protection under provincial legislation (Endangered Species Act and. laws on faunic habitats). In the U.S., Blackchin Shiners have been consid- ered to be of “special concern” only in New York State (Johnson 1987), even though it has been elimi- nated from Ohio waters since 1950 (Trautman 1981), and Iowa (Scott and Crossman 1973). The range is also apparently decreasing in Minnesota and Illinois in concert with habitat loss due to siltation (Eddy and Underhill 1974; Smith 1979). Population Sizes and Trends There is no recent information from Quebec on this species where it is largely known only from A ae ae j { | 4 4 j 4 | i | 4 | i 5 4 ‘ t 1996 HOUSTON: STATUS OF THE BLACKCHIN SHINER 485 FiGureE 2. North American range of the Blackchin Shiner (Notropis heterodon). [Based on NMC, OMNR and ROM collection records and Gilbert (1980)]. presence and absence data (P. Houde, Ministere du Loisir, de la Chasse, et de la Péche, Direction de la Gestion de la Faune, Hull, Quebec; personal commu- nication). Mongeau et al. (1979) did not find this fish to be common in the Chateauguay system, although it appears to be fairly common in the Ottawa River tributaries of Hull, Pontiac and Gatineau counties (D. E. McAllister, Canada Museum of Nature, Ottawa, Ontario; personal communication). In short, there is insufficient information for an assessment of population sizes and trends of this species in Quebec where no general surveys have been conducted since the 1970s (Houde, personal communication). The Blackchin Shiner is generally common where suitable habitat (clean, clear, weedy waters) exists (McAllister, personal communication) and is widely distributed in southern Ontario where it appears in some areas in sufficient numbers to be utilized as a bait fish (Scott and Crossman 1973). Although many streams have undergone habitat degredation similar to that responsible for the demise of the species in Iowa and Ohio, there is no direct evidence of a" decline in the species in southern Ontario. The species was not known west of Sault Ste Marie until 1968 when a number of specimens were collected in Quetico Provincial Park (ROM 26052). Since that time, the Blackchin Shiner has been recorded at a number of sites in the Rainy River — Winnipeg River system in the Lake of the Woods area and west into Manitoba (Falcon Lake, Manitoba 49°41'N; 95°19'W) and is apparently abundant where found (Stewart et al. 1985; Stewart 1988). On the other hand, there has been no program of systematic surveys in the province so neither is there evidence to suggest the species is secure. As in northwestern Ontario, the species was not recorded from Manitoba until 1973 (Stewart et al. 1985), but based on the size of specimens examined and the abundance where found, it has probably been there for some time. Stewart et al. (1985) and Stewart 486 (1988) discussed the disjunct and scattered popula- tions of this species in Manitoba and concluded that the species was not introduced and entered Manitoba naturally, surviving in areas of suitable habitat. The Dauphin Lake drainage has no direct connection with the Assiniboine River drainage and the species could not have moved naturally between the two in a rela- tively short time (Stewart et al. 1985). Similar argu- ments could be made for the dispersal of these fish from Mississippi headwaters to Rainy River headwa- ters in Minnesota and subsequent downstream disper- sal in the Rainy River — Winnipeg River systems in northwestern Ontario and southeastern Manitoba (see Crossman and McAllister 1986). Stewart (1988) fur- ther concluded that the absence of the species from the Red River plain in Manitoba suggests westward dispersal through the Manitoba Great Lakes. As elsewhere, population size and trend informa- tion is lacking, but the Blackchin Shiner is apparently abundant where found in Manitoba (Stewart et al. 1985). The fact that it was not recorded in Manitoba and northwestern Ontario until the latter half of this century is probably related to lack of collecting effort and/or confusion with similar species such as the Blacknose Shiner, Notropis heterolepis. So far, con- tinued sampling effort has not produced additional records from the Winnipeg/Rainy River system or evidence of the species from the Interlake area of Manitoba, but has revealed the presence of the Weed Shiner in the Interlake region (A. J. Derksen, Fisheries Specialist, Manitoba Department of Natural Resources, Winnipeg, Manitoba; personal communi- cation). As in the U.S.A., the species is sensitive to siltation and may not be able to coexist with the Weed Shiner, since the two species have not been captured together in Manitoba (Derksen, personal communication). Habitat No specific details are available on habitat re- quirements other than that clear, clean, cool waters with plenty of submerged aquatic vegetation ap- pear to be essential for the success of the species (Trautman 1981; Scott and Crossman 1973). They prefer quiet pools in creeks and rivers and weedy inshore areas of lakes (Scott and Crossman 1973) where the fish are usually found over bottoms of clean sand and/or gravel and quickly disappear when waters become turbid, bottoms silty and aquatic veg- etation vanishes (Trautman 1981; Eddy and Underhill 1974; Smith 1979). There is no information on preferred water tem- peratures, but the range is restricted to glacial pot- hole lakes and bays and tributary streams (Trautman 1981; Gilbert 1980). The range does not extend south of the Wisconsonian glacial maxima (Gilbert 1980) suggesting that water temperature may be an important to the success of the species. THE CANADIAN FIELD-NATURALIST Vol. 110 Aquatic vegetation probably plays some role in | reducing the risk of predation and is probably also a_| factor influencing the availability of appropriate food | items. The diet consists of small crustaceans (clado- | cerans, copepods and other entomostracans) and | small insects (mainly Diptera) taken at the surface (Scott and Crossman 1973; Cooper 1983). General Biology The biology and ecology of this species have apparently not been well studied; there is very little information available on Canadian populations. The species is said to spawn in May to June in Illinois (Forbes and Richardson 1920); however, Smith (1979) indicates that Forbes and Richardson were | probably referring to the Weed Shiner, Notropis tex- | anus, rather than the Blackchin Shiner. In | Wisconsin, spawning occurs from June to August. | Gravid females were collected in many samples | taken through 9 August, and inspection of ovaries | indicated a wide range in egg development. This | suggests that spawning may extend over a period of | several weeks (Becker 1983). Blackchin Shiners are short-lived, most in Ohio, | not living past two years of age (Trautman 1981). In Ohio young-of-the year fish ranged from 18 to 36 mm by October, immatures 25 to 51 mm and adults 41-71 total length (Carlander 1969). Scott and | Crossman (1973) reported that adults from Canadian | waters averaged 51 to 61 mm; Stewart et al. (1985) recorded and average total length of 47.1 mm for | specimens taken in Spruce Woods Provincial Park, | Manitoba in 1982. The species is considered to be a specialized feed- |} er, due to the terminal position of the mouth | (Willsman 1979 in Smith 1985), eating mostly | Cladocera and flying insects (small Diptera) taken at the surface (Keast 1965; Smith 1985). There is a pro- | gression of foods from young to adult; the young fish feeding on algae and zooplankton while adults prefer larger zooplanters, water fleas and surface }} midges (Cooper 1983; Keast 1985). In some areas plants, including the algae Oedogonium and | Spirogyra, may constitute a large part of the diet | (Rimsky-Korsakoff 1930). No information on behaviour in the wild is avail- | able, although Abrahams and Colgan (1985) have uti- |}, lized the species in conducting some laboratory exper- |}, iments related to hydrodynamic benefits from school- | ing. Parasites of this fish (mostly trematodes and | nematodes) have been documented by Bangham and Hunter (1939), Bangham (1955) and Hoffman (1967). Limiting Factors As discussed previously, the species appears to have a narrow range of habitat requirements and responds quickly to changes in habitat and water quality. Trautman (1981) indicated that Blackchin |) 1996 Shiners disappeared almost immediately from Ohio waters at locations where human or other distur- _ bances resulted in increases in turbidity and siltation or decreased aquatic vegetation. Similar results have been noted in Illinois (Smith 1979), Iowa (Scott and Crossman 1973), and Minnesota (Eddy and Underhill 1974). Ambient water temperature could also be a limit- ing factor as the species appears to favour cooler waters of glacial lakes and the headwaters of tribu- tary streams; however, no information exists on the _ temperature requirements of the species. Fluctuations in water level may also be limiting. Eddy and Underhill (1974) have observed that Blackchin Shiners are abundant in Lake Itasca (Minnesota) for several years, and then rare for sev- eral years, before becoming abundant again. These fluctuations in population seem to correlate with ris- ing and falling water levels due to differences in annual rainfall, high and stable water levels are fol- lowed by population increases (Becker 1983) Special Significance of the Species The Blackchin Shiner is probably an important forage species where abundant (Scott and Crossman 1973; Cooper 1983). It is apparently easily main- tained in aquaria and has been used as a laboratory species (Abrahams and Colgan 1985). The disjunct distribution and habitat requirements _are of interest to science in relation to the zoogeo- graphic history and distribution of species subse- quent to the Wisconsin Period of glaciation. Its criti- cal habitat requirements could also make the species a useful indicator of changing water quality if the | previous occurrence at a specific site were known. Evaluation The Blackchin Shiner has a relatively wide distri- bution in Canada from southwestern Quebec to west- -ern Manitoba. There is insufficient information on which to determine the status of the species in Quebec, but it may be secure, at least in streams trib- utary to the Ottawa River which have undergone lit- tle or no disturbance since the last surveys undertak- -en in early 1970s. However, directed surveys are _Tequired to verify both this status and that no chemi- cal or physical changes have occurred within the aquatic habitat. There is no evidence that Ontario populations are in decline, although those of southwestern Ontario should be surveyed on a regular basis. Streams in this region are subject to a variety of human distur- bances which could affect critical habitat resulting in demise of the resident stocks. The species has been designated as being of spe- cial concern in Manitoba (Johnson 1987), based on earlier (prior to 1985) inforthation indicating the presence of the species at only one location in the HOUSTON: STATUS OF THE BLACKCHIN SHINER 487 province, i.e., from Oak Creek, near Treesbank. During the 1980s, the species was found to occur in abundance at several other locations (Babaluk and Harbicht 1984; Stewart et al. 1985; Stewart 1988) and might exist in other suitable habitat (continued sampling has not as yet provided additional records) as well. Given that its occurrence in Manitoba is of long standing and that it previously escaped detec- tion for a variety of reasons, there appears to be no reason why the Blackchin Shiner should not be con- sidered to be a naturally occurring species there. The Blackchin Shiner is of special concern in Manitoba because its distribution is mostly in agri- cultural areas of the Province; e.g., the small oxbow lakes along the Assiniboine River and the tributaries of Lake Dauphin. These areas are vulnerable to activities such as drainage works, channelization, damming, etc. At the present time, there is no indication that the overall status of the species in Canada is at risk or a COSEWIC listing would be appropriate. However Manitoba populations are of special concern and more information is required on the status of the species in Quebec. Acknowledgments Funding for production of the report was made available through World Wildlife Fund (Canada) and the Department of Fisheries and Oceans. The author is grateful for the assistance of A. Derksen, Manitoba Department of Renewable Resources; Don E. McAllister, Canadian Museum of Nature; E. J. Crossman, Royal Ontario Museum; G. Gale, Ontario Ministry of Natural Resources and P. Hood, Quebec Ministry of Recreation, Hunting and Fishing in pro- vision of collection records and advice, and to COSEWIC for the opportunity to present the report. Literature Cited Abrahams, M. V., and P. W. Colgan. 1985. Risk of preda- tion, hydrodynamic efficiency and their influence on school structure. Environmental Biology of Fishes 13(3): 195-202. Babaluk, J. A., and S. Harbicht. 1984. Range extension of the blackchin shiner, Notropis heterodon, to Dauphin Lake, Manitoba. Canadian Field-Naturalist 98(1): 58. Bangham, R. V., and G. W. Hunter, III. 1939. Studies on fish parasites of Lake Erie. Distribution studies. Zoologica 24(4); 27: 385-448. Bangham, R. V. 1955. Studies of fish parasites of Lake Huron and Manitoulin Island. American Midland Naturalist 53(1): 184-194. Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison, Wisconsin. 1052 pages. Bergeron, J. F., and J. Brousseau. 1983. Guide des pois- sons d'eau douce du Québec. Gouvernement du Québec, Ministére du Loisir, de la Chasse et de la Péche. Québec, Québec. Carlander, K. D. 1969. Handbook of freshwater fishery biology. Volume 1. Iowa State University Press, Ames Iowa. 488 Cooper, E. L. 1983. Blackchin shiner, Notropis heterodon (Cope). Page 106 in Fishes of Pennsylvania. Pennsylvania State University Press, University Park, Pennsylvania. Crossman, E. J., and D. E. McAllister. 1986. Zoo- geography of freshwater fishes of the Hudson Bay drainage, Ungava Bay and Arctic Archipelago. Pages- 53-104 in The zoogeography of North American fresh- water fishes. Edited by C. H. Hocutt and E. O. Wiley. John Wiley and Sons, New York, New York. Eddy, S., and J. C. Underhill. 1974. Northern fishes. University of Minnesota Press, Minneapolis, Minnesota. Forbes, S. A., and R. E. Richardson. 1920. The fishes of Illinois. State of Illinois Natural History Survey Division, Urbana Illinois. Gilbert, C. R. 1980. Notropis heterodon (Cope), Blackchin shiner. Page 271 in Atlas of freshwater fishes of North America. Edited by D. S Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer, Jr. North Carolina State Museum of Natural History, Biological Survey Publication Number 1980-12. Hoffman, G. L. 1967. Parasites of North American fresh- water fishes. University of California Press, Los Angeles, California. Johnson, J. E. 1987. Protected fishes of the United States and Canada. American Fisheries Society, Bethesda, Maryland. Keast, A. 1965. Resource subdivision amongst cohabiting fish species in a bay, Lake Opinicon, Ontario. Publications of the Great Lakes Research Division, University of Michigan 13: 106-132. Keast, A. 1985. Planktivory in a littoral-dwelling lake fish association: prey selection and seasonality. Journal of Fisheries and Aquatic Sciences 42: 1114-1126. Mandrak, N. E., and E. J. Crossman. 1992. A checklist of freshwater fishes of Ontario. Royal Ontario Museum, Toronto, Ontario. Mongeau, J.-R., A., G. Courtemanche, G. Massé, and B. Vincent. 1974. Cartes de répartition géographique des THE CANADIAN FIELD-NATURALIST Vol. 110 espéces de poissons du sud du Québec d'aprés les inven- | taires ichthyologiques effectués de 1963 a 1972. Québec _ Ministere du Tourisme, de la Chasse et de la Péche, | Service de I'Aménagement de la Faune, Rapport spécial — numero 4. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1979. Les poissons du bassin de drainage de la riviére Chateauguay, leur milieu naturel, leur répartition géo- graphique et leur abondance relative. Gouvernement du | Québec, Ministére du Tourisme, de la Chasse et de la | Péche. Québec, Quebec. H Rimsky-Korsakoff, V. N. 1930. The food of certain fishes _ of the Lake Champlain watershed. Pages 88-104 in A biological survey of teh Champlain watershed. Supplement, 19th Annual Report New York © Conservation Department 1929. i Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes | of Canada. Fisheries Research Board of Canada, Bulletin 184. Smith, C.- L. 1985. The inland fishes of New York State. New York State Department of Environmental © Consevation, Albany, New York. Smith, P. W. 1979. The fishes of Illinois. University of Illinois Press, Urbana Illinois. Stewart, K. W., I. M. Suthers, and K. Leavelsey. 1985. New fish distribution records in Manitoba and the role of a man-made interconnection between two drainages as an avenue of dispersal. Canadian Field-Naturalist 99(3): | 317-326. Stewart, K. W. 1988. First collections of the Weed Shiner, | Notropis texanus, 11 Canada. Canadian Field-Naturalist 102(4): 657-660. Trautman, M. B. 1981. The fishes of Ohio. Ohio State University Press, Columbus, Ohio. Willsman, G. A. 1979 Resource partitioning and mecha- | nisms of coexistence of blackchin and blacknose shiners (Notropis: Cyprinidae). Ph.D. dissertaliton, Michigan Stae University, Ann Arbor, Michigan. 130 pages. Accepted 13 March 1996 The Status of the Rosyface Shiner, Notropis rubellus, in Canada* J. HOUSTON 374 Fireside Drive, R.R. 1 Woodlawn, Ontario KOA 3MO * Houston, J. 1996. The Status of the.Rosyface Shiner, Notropis rubellus, in Canada. Canadian Field-Naturalist 110(3): 489494. The Rosyface Shiner, Notropis rubellus, is a small cyprinid with a Canadian distribution from the upper St Lawrence drainage of southwestern Quebec west through southern Ontario to southeastern Lake Superior. Other populations have been reported from the Whitemouth-Birch River system in southern Manitoba. The biology and ecology of the species has not been studied in Canada and abundance information is limited to records of occurrence, but outside of the Manitoba population, which is of concern. the species is not at risk. _ Le téte rose, Notropis rubellus, est un petit cyprinidé dont l’aire de répartition au Canada s’étend depuis l’amont du bassin versant du Saint-Laurent, dans le secteur sud-ouest du Québec, jusqu’a l’extrémité sud-est du lac Supérieur et englobe le sud de l’Ontario. D’autres populations ont été signalées dans le systéme des rivieres Birch et Whitemouth, au sud du ors Manitoba. Au Canada, ses caractéristiques biologiques et écologiques n’ont pas été étudiés et l’abondance de l’espéce est limitée aux nombres de mentions, mais l’espéce n’est pas en péril, sauf la population du Manitoba, qui a l’égard partic- uliére. Key Words: Rosyface Shiner, téte rose, Noi» The Rosyface Shiner, Notropis rubellus (Agassiz 1850), is a small cyprinid found in Canada from the upper St Lawrence river in the east, to extreme southeastern Lake Superior with a northwest exten- sion of distribution reaching southern Manitoba in the west (Scott and Crossman 1973). Information is lacking on relative abundance of the species in Canada, but it is widely distributed in the central part of the range. Since it has a limited distribution in Manitoba (Hinks 1943; Fedoruk 1969), which is sep- arated from the nearest location in Ontario by some 900 to 1000 km, Derksen (A. J. Derksen, Fisheries Specialist, Manitoba Ministry of Natural Resources, Winnipeg, Manitoba; personnel communication to Chairman, COSEWIC Fish and Marine Mammal ' Subcommittee) suggested that it should be consid- ered for COSEWIC review. Description This fish has a typically slender, elongate body (Figure 1), adults do not usually grow larger than 75 mm in standard length (Gilbert and Burgess 1980) or 89 mm in total length (Trautman 1981), averaging 51 to 76 mm in total length (Scott and Crossman 1973). The species is closely related to, and resem- bles the Emerald Shiner, Notropis atherinoides, and the Silver Shiner, Notropis photogenis. Clay (1975) indicated that the similarity of the Rosyface Shiner to the Emerald Shiner has led to erroneous identifi- cations and some old records may be misidentifica- tions. Adults of the species can be separated from the _ Emerald Shiner as the body of the latter is deeper and more compressed the origin of the dorsal fin is sy rubellus, Cyprinidae, southwestern Ontario. slightly posterior to the pelvic insertion, and the snout is blunt and shorter (Clay 1975; Smith 1985). The body is usually deeper and more compressed and there is no red colouration on breeding adults; the dorsal surface is usually not as darkly pigmented. It can be separated from the Silver Shiner by its lat- ter’s nine pelvic rays, prominent middorsal stripe, more anterior position of the dorsal fin, and the dark crescents between the nostrils (Smith 1985). In the Rosyface Shiner the anal fin typically has 10 or 11 rays, the dorsal fin is usually located about midway between the base of the caudal fin and the preopercle. The caudal fin is moderately forked with rounded lobes, the origin of the anal fin 1s located in line with the base of the last dorsal ray, and the pelvic origin and insertion are considerably anterior to the dorsal origin. The pectoral fins are low on the body and well forward, the length and width is greater in males than in females (Becker 1983). The lateral line is complete and somewhat decurved towards the anterior. The mouth is terminal, large and slightly oblique. Proportional measurements and counts are summarized in Scott and Crossman (1973) and Smith (1985). The normal colouration of the fish is overall sil- very with slight olivaceous colouration dorsally and silvery white ventrally, the fins are transparent. Breeding males develop 100 or more tubercles on the upper half of the head from snout to occiput (Becker 1983) and the anterior rays of the pectorals; a few may occur on the opercles and the scales along the lateral line. Tubercles may also be found on the upper surfaces of the pelvic, dorsal and anal fins *Reviewed and approved by COSEWIC 14 April, 1994, status assigned — Vulnerable, Manitoba. 489 490 THE CANADIAN FIELD-NATURALIST Vol. 110 FIGURE 1. Rosyface Shiner, Notropis rubellus, NMC 64 0203, approximately 80 mm. Drawing by Sally J. Gadd, courtesy D. E. McAllister, Canadian Museum of Nature. (Scott and Crossman 1973). Breeding males display orange to brick red colouration of the head (hence the name, Roseface Shiner) to the nape of the neck with the rest of the body being light orange to orange-yellow (Becker 1983). The opercle, sides and ventral surface to the anal fin are a lighter red and the pectoral, pelvic, dorsal and anal fins may be dif- fused with red. Females may also develop this red colouration, but it is generally paler than on the males and may be absent (Becker 1983); they may also develop tubercles on the head (Pfeiffer 1955; Scott and Crossman 1973; Smith 1985). Distribution Rosyface Shiners are found in central and eastern North America (Figure 2). In the east, the limit of the range is the upper St Lawrence River drainage of southern Quebec, south through New York and Vermont to the James River drainage of Virginia, but excluding the Delaware. The range extends west through most of the Great Lakes drainage (excluding most tributaries to the north shore of Lake Superior) to the Red River of the North drainage in Minnesota and north into Manitoba. West of the Appalachians it occurs in the upper Ohio and Mississippi River basins from Pennsylvania, possibly to North Dakota (one location record from the Sheyenne River), Minnesota and Iowa. East of the Mississippi it is found south to the Tennessee River drainage of North Carolina, Tennesse and Alabama and west of the Mississippi in the Ozark uplands of Arizona, Oklahoma and Kansas to southern tributaries of the Missouri River in Montana (Scott and Crossman 1973; Gilbert and Burgess 1980). In Canada, the species is most widely distributed in Ontario, although the range more or less extends from the upper St Lawrence near Quebec City, west to southcentral western Manitoba (Figure 2), with a hiatus on the north shore system of Lake Superior. The Manitoba records look unusual only if we look at the Canadian populations in exclusion of the distribu- tion in lowa, Wisconsin, Minnesota and the Dakotas. The species is limited to the extreme southern region of Quebec (Figure 2) where it is known from tributaries of the Ottawa River north and west at least to the Black River (Bergeron and Brousseau 1983), and of the St Lawrence River as far down- stream as the Nicolet River at the eastern end of Lake St Pierre, just upstream of Quebec City (Mongeau et al. 1974; Mongeau et al. 1979; Bergeron and Brousseau 1981). Rosyface Shiners have also been collected from the south shore of the St Lawrence from the Richelieu and Chateauguay rivers and other St Lawrence River tributaries east to Leclercville on the River Grande (ROM 42159, 46°34’°25”N, 71°59’41”W; Mongeau et al. 1974). The Quebec distribution appears to be restricted to the St Lawrence and Ottawa River drainages south of a line at approximately 46 N. In Ontario, the species occurs in southwestern Ontario (Figure 2) in streams draining into lakes Ontario, Erie, Huron and the eastern end of lake Superior and the Ottawa River watershed north to about the Mattawa River (Scott and Crossman 1973), as in Quebec at a line approximately at 46° N [1.e., the Great Lakes watershed and not found in the Hudson, James or Ungava Bay (Arctic watersheds)]. Rosyface Shiners have not been recorded from north shore Lake Superior tributaries, although they have been recorded from several streams in Michigan tributary to that lake (Scott and Crossman 1973; Smith 1979). Hinks (1943) referred to the species as occurring in the Lake of the Woods watershed, but there are no known records to support the statement. The species should be looked for there as suitable habitat exists and other cyprinids with similar habitat requirements and distri- butions, such as Notropis heterodon, previously not known west of Sault St Marie have recently been recorded from the Rainy River watershed in Quetico Park and the Wabigoon River watershed of the Kenora District of northwestern Ontario (Crossman and McAllister 1986; Crossman 1986; Stewart 1988). Literature records of the presence of the species in Manitoba where it is known (A. J. Derksen, Fisheries Specialist, Manitoba Department of Natural Resources, Winnipeg, Manitoba; personal communi- cation) from the Whitemouth-Birch river systems (tributary to the Winnipeg River which it joins above | an historically impassable falls) go back at least to Hinks (1943). However, this record is uncertain as he | | 1996 | TC OOOO EE Vela Sak HOUSTON: STATUS OF THE ROSYFACE SHINER cs} 2 Or oe 49] FicurE 2. North American range of the Rosyface Shiner, Notropis rubellus, (from sources cited in the text). bases the species presence on unsupported evidence that it occurred in the Lake of the Woods system. Fedoruk (1969) also lists the species among the fauna of Manitoba, but he did not give the distribution of any species. Scott and Crossman (1973) included its distribution in Manitoba as in the Red River in south- ern Manitoba, perhaps based on a 1955 collection by J. J. Keleher from the Whitemouth River at Whitemouth, Manitoba, catalogued at the Royal Ontario Museum (ROM 17539), but more likely because of the reports of the species from the Red River Basin in Minnesota (Derksen, personal com- munication). The species does not occur along the axis of the Red River in Manitoba (K. Stewart, Department of Zoology, University of Manitoba, Winnipeg, Manitoba; personal communication). No other reliable literature records are known, although the species has been collected several times since from the Whitemouth and as recently as 1984 (ROM 45731; NMC 84-0010, NMC-85-0002). A previously unreported 1976 collection from Lake Manitoba, at Delta Marsh (50°12’00”N, 98° 20°00” W), catalogued in the Canadian Museum of Nature Collection (NMC 76-0421), has been re-examined and identified as Notropis atherinoides, not Notropis rubellus (B. Coad, Canadian Museum of Nature, Ottawa, Ontario; personal communication). Protection Rosyface Shiners are not subject to any specific protection in Canada, but general protection is afforded, if required, in Manitoba through the provincial Endangered Species Act. In Quebec, the species has been given little, or no attention, but could be given specific protection under provincial legislation (Endangered Species Act and law on fau- nic habitats) if required. In the U.S., Rosyface Shiners are considered to be of “special concern” only in the states of Mississippi, North Carolina, North Dakota and South Dakota and is given protected status in Tennessee (Johnson 1987). It has been extirpated from most Ohio waters 492 since 1950 (Trautman 1981) and is in decline in Minnesota and Illinois in concert with habitat loss due to siltation (Eddy and Underhill 1974; Smith 1979). However, forms of the species have devel- oped in the Pecatonica and Sugar river basins of Wisconsin which are capable of withstanding con- siderable turbidity (Becker 1983). Population Sizes and Trends There is no recent information from Quebec on this species where it is largely known only from presence and absence data (P. Houde, Ministére du Loisir, de la Chasse, et de la Péche, Direction de la Gestion de la Faune, Hull, Québec; personal communication), although a few incidental collections in the 1980s did note the presence of the species at some locations where it had been recorded earlier (Appendices of all records are on file and available on request from COSEWIC). Mongeau et al. (1979) found this fish to be abundant where found and Mongeau et al. (1979) found it to be common in the Chateauguay system, and it appears to be fairly abundant in the Ottawa River tributaries of Hull, Pontiac and Gatineau coun- ties (McAllister and Coad 1974). The species is wide- ly distributed in southern Ontario, collection records of the Royal Ontario Museum (ROM, Canada Museum of Nature (NMC) and the Ontario Ministry of Natural Resources (OMNR) indicate that the species is usually numerous where found. There is no direct evidence of a decline in the species in southern Ontario where it has recently been recorded from the same streams where found earlier this century. Many of these streams, particularly in southwestern Ontario, have undergone habitat degradation similar to that responsible for the demise of the species in Minnesota, Illinois and Ohio. On the other hand, there is no evidence to suggest the species is secure, although the species is generally common where suit- able habitat (clean, clear, weedy waters) exists (McAllister, personal communication). The species is not known in Ontario west of Sault Ste Marie, but it should be looked for in Quetico Park [not found there in the ROM surveys (Crossman 1976)] and the Rainy River — Winnipeg River system in the Lake of the Woods area, and west into Manitoba where similar species such as the Emerald Shiner, have recently been recorded (Scott and Crossman 1973). Although listed among the fishes of Manitoba by Hinks (1943), the first verified record of the species appears to be the 1955 collection by Keleher in the Whitemouth River (ROM 17539). The species is apparently abundant in this system (Derksen, person- al communication). Stewart et al. (1985) and Stewart (1988) discussed the disjunct and scattered populations of the Blackchin Shiner, Notropis heterodon, in Manitoba and concluded that the species has been there for some time, but had gone unreported through lack of collecting effort and/or confusion with other species. Similar arguments could be made for the dispersal of THE CANADIAN FIELD-NATURALIST Vol. 110 Notropis rubellus. This species like several others (Crossman and McAllister 1986) occurs mainly in the most southerly regions of the Hudson Bay sys- | tem and gets into Canada only as far as the Whitemouth River and are non-existent north of this. These fishes are warm water adapted and have only recently (in geological time) arrived at their norther- ly limit or are probably still limited in their north- ward extensions by temperature preferences. Some | species like the Rosyface Shiner have extended their range north only while others like the Blackchin have extended east to Lake of the Woods and Quetico (Crossman and McAllister 1986). The fact that the Rosyface Shiner was not record- ed in Manitoba until the latter half of this century | might be related to lack of collecting effort and/or confusion with similar species such as the Emerald Shiner [the Silver Shiner, is known in Canada only from southwestern Ontario (Baldwin 1988)] which is common in southern Manitoba (Scott and Crossman 1973). Continued sampling effort may produce addi- tional records from the Winnipeg/Rainy River sys- tem and could provide evidence of the species from the Interlake area of Manitoba. However, Ken Stewart of the University of Manitoba has conducted extensive collections in and around the Winnipeg River and the Interlake area. So far these efforts have failed to turn up Rosyface Shiners in any of these areas, or in streams adjacent to the Whitemouth River (Derksen, personal communication). Habitat The typical habitat of the species is clear, fast flowing larger streams and small rivers over sub- strates of clean gravel (Gilbert and Burgess 1980). Generally said to avoid small and\or sluggish | streams (Clay 1975), it is often found in schools in riffles and clear pools in the lower portion of streams near the confluence with larger streams or rivers (Scott and Crossman 1973; Smith 1979) where aquatic insects are plentiful. It is intolerant of turbid- ity and siltation (Trautman 1981). Water temperature tolerances are not known, but Crossman and McAllister (1986) list this as a fish of warm waters, and water temperature may be limiting | to its Canadian range which is the northern limit for the species. However, Smith (1979) suggests it is | intolerant of high summer water temperatures. Clear, | silt free pools and water temperature of at leat 21°C | seem to be critical for spawning activity in New | York (Pfeiffer 1955). General Biology Apparently the biology and ecology of Canadian | populations of this species have received little or no | attention. The species has received some attention in | the U.S. where it is more widely distributed. In New York State, Rosyface Shiners spawn in late June when water temperatures are between 21 to 25°C | (Pfeiffer 1955). In Illinois, Pennsylvania and Ohio | 1996 spawning occurs in late May and early June (Reed 1957a; Smith 1979; Trautman 1981) and may contin- ue into late June (Reed 1957b). Reed (1957a) noted that spawning in Pennsylvania occurred at water tem- peratures of 20 to 22°C. Spawning occurs in shallow water over gravel in riffles (Pfeiffer 1955; Reed 1957a; Trautman 1981). Becker (1983) mentions that spawning occurred during bright sunny days in shal- low water area containing nests of minnows. Pfeiffer (1955) and Becker (1983) provide detailed descrip- tions of spawning behaviour where schools of eight to 12 fish rush into the spawning area, the males collid- ing with females. During these brief (5 to 6 second) sessions the fish vibrate over a depression in the grav- el and the eggs are released and fertilized. Reed (1957b) noted that later spawnings involved smaller groups of fish with a preponderance of males. Rosyface Shiners apparently spawn over the nests of Nocomis spp. (Reed 1957a) and Luxilus cornutus (Becker 1983). Hybrids involving Notropis rubellus and Luxilus cornutus are well documented in the U.S., and six collections of Luxilus cornutus X Notropis rubellus are catalogued in the ROM collection (E. J. Crossman, Royal Ontario Museum, Toronto, Ontario; personal communication). According to Smith (1979) this fish spawns over the nests of sunfish, but hybrids with sunfish are unknown and unlikely. There is a record between Notropis rubellus and Notropis volu- cellus (Bailey and Gilbert 1960). Rosyface and Silver shiners are often associates and spawn on the same riffles (Trautman 1981); however, the Silver Shiner spawns later (late June to early July in Ohio) and hybridization is unlikely. The development of young fish has not been well documented; however, Reed (1958) found that hatch- ing occurred in about 60 hours at 20°C and Pfeiffer (1955) found 5 mm fry eight days after spawning. The fish are apparently mature at one year of age (males and females); one year old females were found to have 600 eggs on average (egg diameter being 1.1 to 1.5 mm) and three-year-old females averaged 1175 (Pfeiffer 1955), the eggs are demersal and sticky (Becker 1983). The males appear to grow faster than the females in the first year while the females grow faster in the second and third years. The maximum age 1s apparently three years (Pfeiffer 1955). Rosyface Shiners are omnivorous; aquatic insects making up about 72% of the diet (Reed 1957b); algae, diatoms and inorganic material making up the remain- der. Smith (1985) reported that aquatic and terrestrial invertebrates account for about 92% of the food. Parasites of this species are listed by Hoffman (1967). There is no information on utilization by other fishes for forage. Limiting Factors The species appears to have a narrow range of habitat requirements and responds quickly to changes in habitat and water quality. Trautman (1981) indicated that since 1938 it has decreased in HOUSTON: STATUS OF THE ROSYFACE SHINER 493 numbers, and even been extirpated from some areas in Ohio due to increased turbidity and siltation. In Illinois, the fish is disappearing from streams modi- fied by impoundments and excessive siltation (Smith 1979). Clay (1975) also remarked on its absence from impounded waters in Kentucky. Similar reac- tions have also been noted in Minnesota (Eddy and Underhill 1974). Ambient water temperature could also be a limit- ing factor as the species appears to favour cooler waters of glacial lakes and the headwaters of tribu- tary streams, however no information exists on the temperature requirements of the species. Smith (1979) thought it to be intolerant of high summer water temperatures. Special Significance of the Species The Rosyface Shiner, like other small cyprinids, may be an important forage and bait species where abundant. Scott and Crossman (1973) indicated its potential in studies of water quality because of its sensitivity to turbidity and siltation. Cherry et al. (1977) did utilize the species in a study to determine avoidance reactions of fish to free residual chlorine. The disjunct distribution and habitat requirements are of interest to science in relation to the zoogeo- graphical history and distribution of species subse- quent to the Wisconsin glaciation. Evaluation The Rosyface Shiner has a relatively wide distrib- ution in Canada from southwestern Quebec to south central Manitoba. There is no indication that the species is in decline in Quebec and it is probably secure in that province at present, at least in streams tributary to the Ottawa River which have undergone little or no disturbance since the last surveys under- taken in early 1970s. There is no evidence that Ontario populations are in decline, although those of southwestern Ontario should be surveyed on a regular basis. Scott and Crossman (1973) felt that it may be less common in some parts of the range than it was two or three decades ago, but had no evidence to support the sup- position. Streams in southwestern Ontario are sub- ject to a variety of human disturbances which could affect critical habitat resulting in demise of the resi- dent stocks. The species was brought to the attention of COSEWIC for consideration because of its apparent rarity in Manitoba, where it was supposedly known from only one location in the province, the Whitemouth River. Its presence in Manitoba is undoubtedly of long standing, and it has likely previ- ously escaped detection for a variety of reasons. The species may be of particular concern in Manitoba because of its restricted distribution and depen- danceon clear, fast moving waters which are being threatened by agricultural activity (Stewart, personal communication). 494 There appears to be no reason why the Rosyface Shiner should be considered to be of special concern in Ontario or Quebec, but should be considered as vulnerable in Manitoba. Acknowledgments Funding for production of the report was made available through World Wildlife Fund (Canada) and the Department of Fisheries and Oceans. The author is grateful for the assistance of A. Derksen, Manitoba Department of Renewable Resources; D. E. McAllister, Canadian Museum of Nature; E. J. Crossman, Royal Ontario Museum; G. Gale, Ontario Ministry of Natural Resources and P. Hood, Quebec Ministry of Recreation, Hunting and Fishing in pro- vision of collection records and advice, and to COSEWIC for the opportunity to present the report. Literature Cited Bailey, R. M., and C. R. Gilbert. 1960. The American cyprinid fish Notropis kanawha identified as an interspe- cific hybrid. Copeia 1960(4): 354-357. Baldwin, M. E. 1988. Updated status of the Silver Shiner, Notropis photogensis, in Canada. Canadian Field- Naturalist 102(1): 147-157. Becker, G. C. 1983. Fishes of Wisconsin. University of Wisconsin Press, Madison, Wisconsin. 1052 pages. Bergeron, J. F., and J. Brousseau. 1983. Guide des pois- sons d’eau douce du Québec. Gouvernement du Québec, Ministére du Loisir, de la Chasse et de la Péche. Québec, Québec. Cherry, D. S., S. R. Larrick, K. L. Dickson, R. C. Hoehn, and J. Cairns Jr. 1977. Significance of hypochlorous acid in free residual chlorine to the avoid- ance response of spotted bass (Micropertus punctatus) and rosyface shiner (Notropis rubellus). Journal of the Fisheries Research Board of Canada 34: 1365-1372. Clay, W. M. 1975. Rosyface shiner, Notropis rubellus (Agassiz). Pages 169-170 in The Fishes of Kentucky. Kentucky Department of Fish and Wildlife Resources, Frankfort, Kentucky. Crossman, E. J. 1976. Quetico fishes. Life Sciences Miscellaneous Publications. Toronto, Royal Ontario Museum. Crossman, E. J., and D. E. McAllister. 1986. Zoogeog- raphy of freshwater fishes of the Hudson Bay drainage, Ungava Bay and Arctic Archipelago. Pages 53-104 in The zoogeography of North American freshwater fishes. Edited by C. H. Hocutt and E. O. Wiley. John Wiley and sons, New York, New York. Eddy, S., and J. C. Underhill. 1974. Northern fishes. University of Minnesota Press, Minneapolis, Minnesota. Fedoruk, A. N. 1969. Check list of and key to the fresh- water fishes of Manitoba. Manitoba Department of Mines and Natural Resources, Winnipeg, Manitoba. Canada Land Inventory Project Report 6: 1-98. Gilbert, C. R., and G. H. Burgess. 1980. Notropis rubel- lus (Agassiz), Rosyface shiner. Page 302 in Atlas of North American freshwater fishes. Edited by D. S. Lee, GR. Gilbert, ©. Ho Hocutt, RovA. Jenkins; Dw E: McAllister, and J. R. Stauffer Jr. North Carolina State Museum of Natural History, North Carolina Biological Survey Number 1980-12. THE CANADIAN FIELD-NATURALIST Vol. 110 Hinks, D. 1943, The fishes of Manitoba. Manitoba Department of Mines and Natural Resources, Winnipeg, Manitoba. Hoffman, G. L. 1967. Parasites of North American fresh- water fishes. University of California Press, Los Angeles, California. 486 pages. Johnson, J. E. 1987. Protected fishes of the United States and Canada. American Fisheries Society, Bethesda, Maryland. Mandrak, N. E., and E. J. Crossman. 1992. A checklist | of Ontario freshwater fishes annotated with distribution maps. Royal Ontario Museum Publications in Science, Toronto. McAllister, D. E., and B. W. Coad. 1974. Fishes of Canada’s national capital region. Fisheries and Marine Services Miscellaneous Publication 24. Mongeau, J.-R., A. Courtemanche, G. Massé, and B. Vincent. 1974. Cartes de répartition géographique des espéces de poissons du sud du Québec d’aprés les inven- taires ichthyologiques effectués de 1963 a 1972. Québec Ministére du Tourisme, de la Chasse et de la Péche, Service de l’Aménagement de la Faune, Rapport spécial numero 4. Mongeau, J.-R., J. Leclerc, and J. Brisebois. 1979. Les poissons du bassin de drainage de la riviére Chateauguay, leur milieu naturel, leur répartition géo- graphique et leur abondance relative. Gouvernement du Québec, Ministére du Tourisme, de la Chasse et de la Péche. Québec, Quebec. Pfeiffer, Reverand R. A. 1955. Studies on the life history of the rosyface shiner, Notropis rubellus. Copeia 1955(2): 95-104. Reed, R. J. 1957a. The prolonged spawning of the rosy- face shiner, Notropis rubellus (Agassiz), in southwestern Pennsylvania. Copeia 1957(3): 250. Reed, R. J. 1957b. Phases of the life history of the rosy- face shiner, Notropis rubellus, in southwestern Pennsylvania. Copeia 1957(4): 286-290. Reed, R. J. 1958. The early life history of two cyprinids, Notropis rubellus, and Campostoma anomalum pullum. Copeia 1958(4): 325-327. Scott, W. B., and E. J. Crossman. 1973. Freshwater fish- es of Canada. Fisheries Research Board of Canada Bulletin 184. 966 pages. Smith, C. L. 1985. Rosyface shiner Notropis rubellus (Agassiz 1850). Pages 173-175 in The inland fishes of New York State. New York State Department of Environmental Conservation, Ithica, New York. Smith, P. W. 1979. Rosyface shiner, Notropis rubellus (Agassiz). Pages 115-1116 in The fishes of Illinois. University of Illinois Press, Urbana, Illinois. Stewart, K. W., I. M. Suthers, and K. Leavelsey. 1985. New fish distribution records in Manitoba and the role of a man-made interconnection between two drainages as an avenue of dispersal. Canadian Field-Naturalist 99(3): 317-326. Stewart, K. W. 1988. First collections of the Weed Shiner, Notropis texanus, in Canada. Canadian Field- Naturalist 102(4): 657-660. Trautman, M. B. 1981. Rosyface Shiner, Notropis rubel- lus (Agassiz). Pages 327-329 in The fishes of Ohio, Revised Edition. Ohio State University Press, Columbus, Ohio. Accepted 13 March 1996 The Status of the Warmouth, Chaenobryttus gulosus, in Canada*t E. J. CRossMAN!, J. Houston? and ROBERT R. CAMPBELL? 'Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 *1374 Fireside Drive, R.R. 1, Woodlawn, Ontario KOA 3MO 5666 Bayview, R.R. 1 Woodlawn, Ontario KOA 3MO Crossman, E. J., J. Houston, and Robert R. Campbell. 1996. The Status of the Warmouth, Chaenobryttus gulosus, in Canada. Canadian Field-Naturalist 110(3): 495—500. The Warmouth, Chaenobryttus gulosus, is a recent addition to the southern Ontario freshwater fauna where it has been known since only 1966. Little is known of this species in its limited Canadian range where it is undoubtedly a pioneering species which may have gone undetected for several years prior to 1966. A breeding population is present at Point Pelee. Le crapet a bouche guerriére est une espéce nouvelle a la faune des eaux douces du sud de |’Ontario, ot elle est une migrante récente. On la connait ici depuis 1966, mais on connait trés peu sur cette espéce au Canada. Sans doute, elle est une espéce colonisatrice, et elle y existait probablement quelques années avant de sa découvert en 1966. Une population reproductive est présente a la Pointe Pelée. Key Words: Warmouth, Chaenobryttus gulosus, crapet a bouche guerriére, Centrarchidae, Ontario, status. The Warmouth, Chaenobryttus gulosus (Cuvier, 1829) [see Wainwright and Lauder 1992] has been reported from Canada (Crossman and Simpson 1984) where it is known from only two locations in southwestern Ontario. Although most likely a recent natural migrant, it might have been present and undetected for several years before the initial discoy- ery in 1966. A successful breeding population is pre- sent at Point Pelee (Crossman and Simpson 1984). The Warmouth (Figure 1) is a small, robust cen- trarchid (sunfish), seldom exceeding 310 mm in length or 234 mg in weight. The species superficially resembles the Rock Bass, Amblopites rupestris and could be mistaken for it and, to a lesser extent, the Green Sunfish, Lepomis cyanellus, another small centrarchid which may be found in similar locations. The Warmouth has fewer anal spines than the Rock Bass (three versus six). The presence of many teeth on the tongue and the strong dark lines radiating out from the eye distinquish the Warmouth from the Green Sunfish. In Chaenobryttus gulosus there are no black spots at the bases of the dorsal rays. There are, however, pale spots near the base of most soft rays of the dor- sal and anal fins. In contrast, adults of Lepomis cyanellus have one large black spot at the base of each of the posterior soft dorsal and anal rays, and the head is less densely pigmented. In the Warmouth the maxillary reaches to, or beyond, the pupil so the mouth is larger than that in the Pumpkinseed and Bluegill. The body of the Warmouth is ovoid, compressed, and somewhat broader than most sunfishes. The mouth 1s large and oblique with a projecting lower jaw. The pectoral fin is rounded and the caudal slightly forked. The colour of these fish varies from olive-yellow to brassy or muddy brown dorsally, shading to yellow or white below, a small red or orange spot may appear at the base of the vermicu- lated soft dorsal in breeding males. There are numer- ous darker markings on the sides which may be sug- gestive of vertical bands. The five or so dark lines radiating from the snout and eye across the cheeks and opercle help to distinguish this species (Clay 1962; Smith 1979). Young Warmouth have heavy stippling on the head, and six or seven dark bands that extend from the mid-dorsal line to the ventral surface (Smith 1979; Crossman and Simpson 1984). The young of many other centrarchids have dark vertical bands on the sides so the separation of young Warmouth requires very careful examination. Distribution The species is widely distributed in ponds, lakes and, occasionally, streams throughout the eastern United States (Cooper 1983). The northern limit appears to be along a line from southeastern Minnesota, central Wisconsin to Maryland (see Lee 1980: Figure 1), including western Pennsylvania only. The range extends south to Florida and the Gulf coast to Texas and the Rio Grande and west to *Reviewed and approved by COSEWIC 14 April 1994, status assigned — Vulnerable. +Contribution number 60 of the Centre for Biodiversity and Conservation Biology, Royal Ontario Museum, Toronto, Ontario. 495 496 THE CANADIAN FIELD-NATURALIST Vol. 110 Ficure |. Warmouth, Chaenobryttus gulosus. (Illustration by Duane Raver, U.S. Fish and Wildlife Service, courtesy Don E. McAllister, Canadian Museum of Nature, Ottawa, Ontario) New Mexico and Kansas. Successful transplants have been made west of the Rockies and in locations on the Atlantic slope (Lee in Lee et al. 1980). It has been introduced elsewhere in the U.S. (see McMahon et al. 1984). The closest populations had been one in the upper Huron River in Michigan, and several in the mid and lower reaches of Lake Erie tributaries in eastern Ohio from the Grand River to the Cuyahoga River and possibly one in a tributary (Toussaint River?) east of Toledo (Trautman 1981). In Canada, the species has been recorded only from southwestern Ontario (Figure 2) where its pres- ence was first discovered in 1966 (Crossman and Simpson 1984). Chaenobryttus gulosus has been taken from Lake Erie near Rondeau Provincial Park, about 35 km southeast of Chatham, Ontario (Table 1) and in two ponds at Point Pelee National Park 60 km further west of Rondeau. Protection The species is widespread and common in the US. and is not listed as of any concern in any part of its range there (Williams et al. 1989). Although given no specific protection in Canada, general protection is available, if required, through laws which protect the habitat of the species which include: the Ontario Lakes and Streams Improve- ment Act which prohibits the impoundment or diver- sion of watercourses which leads to siltation; the voluntary Land Stewardship II program of the Ontario Ministry of Agriculture and Food which is designed to reduce the erosion of agricultural lands and thus reduce siltation of habitat. The Point Pelee collection sites (see Table 1) are protected within the confines of the Point Pelee National Park. Similarly, one of the two Rondeau sites is within Rondeau Provincial Park. This is a naturalized species in Canada [according to the recent definition of the American Fisheries Society (Kohler and Courtenay (1986)]. As a result of the growing concern for the potential negative impact of introduced species on native species, and on biodiversity (see Billington and Hebert 1991), some would argue that this species warrants no pro- tection in Canada. Habitat Extensive information on various habitat require- ments for this species was provided by McMahon et al. (1984). The recorded captures of the Warmouth in Ontario have been from ecological conditions similar to those for other sunfishes (see Scott and Crossman 1973). Other warm water species such as Yellow Perch, Perca flavescens; Bowfin, Amia calva, and Brown Bullhead, Ameiurus nebulosus, were taken at the same sites (Crossman and Simpson 1984). Similar assemblages have been noted in the United States (Lewis and English 1949; Larimore 1957; Germann et al. 1974; Guillory 1979). Typically, the Warmouth occurs in the deeper, open water of swamps, sluggish (low gradient) streams, ponds, or lakes with mud, silt or organic detritus substrates (Guillory 1979; Smith 1979; Cooper 1983). In Ohio, this species was said to occur (Trautman 1981) in silt-free water with an abundance of vegetation and a mucky bottom. In Kentucky they are usually found in clear areas with rooted aquatic vegetation (Clay 1962), and in Illinois streams, are associated with pool areas with beds of vegetation or roots and deadheads (Smith 1979). In Missouri, 1996 CROSSMAN, HOUSTON, AND CAMPBELL: STATUS OF THE WARMOUTH 497 GREAT LAKES BASIN LAKE L NIPISSING ST. LAWRENCE — RIVER \ HURON \ FIGURE 2. Canadian (southern Ontario) distribution of the Warmouth, Chaenobryttus gulosus. Pflieger (1971) indicates that the species may occa- sionally be found in turbid waters, but prefers clearer waters. Cooper (1983) suggests they may tolerate brackish water to some degree. Trautman (1981) indicated that (in Ohio) the Warmouth preferred lakes, ponds, marshes and streams of low gradient with an abundance of aquatic vegetation and a mucky bottom covered with organic debris. Undoubtedly, this is a warm water species, but there is no information on temperature tolerances. This could be a factor in limiting distribution even though habitat otherwise suitable should be abundant throughout much of southern Ontario. In Ontario, the Species was captured at depths of 20 to 95 cm (ROM collection records), similar to those reported in the U.S. (Guillory 1979). Population Sizes and Trends Eight specimens were taken from Rondeau Park sites (Table 1) between 1966 and 1968. Although no further specimens have been collected at Rondeau, 53 specimens have been taken (all but nine were released alive) at Point Pelee (Table 1) between 1983 and 1989 (Crossman and Simpson 1984). Of these, 28 were adults and 18 young-of-the-year (Crossman and Simpson 1984) indicating a breeding population. Fisheries agencies along the Ontario side of Lakes Erie, St. Clair, and the Detroit and St. Clair rivers have been advised to be on the lookout for the species but, to 1992, no additional specimens had been noted. This is not surprising since no directed surveys for the species have been undertaken and it can be a difficult species to sample. Contact with personnel at Rondeau and Point Pelee parks in 1992 indicated that no recent records for the species had been noted. However, no assessment work that might have indicated presence or absence was car- ried out in 1991. The two individual populations of Chaenobryttus gulosus in Ontario would appear to comprise one or more pioneering colonies resulting from recent nat- ural dispersion, probably moving along the shore from Ohio to Lakes Erie and St. Clair rather than across the lake from Ohio (Crossman and Simpson 1984). Although first recorded in Canada in 1966 they probably arrived several years prior to that date. Pioneer populations often go undetected for some time prior to documentation (Crossman and 498 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Collection Records of Chaenobryttus gulosus in Canada. Number of Location Date Specimens Source Kent Co., Rondeau Provincial Park, 5 June 1966 1 RPM? F103-66 Lake Erie, 42°17'N, 81°51'W Kent Co., Rondeau Provincial Park, 1967 2 Crossman and Simpson (1984) - Lake Erie, 42°17'N, 81°51'W Kent Co., Rondeau Provincial Park, 1968 3 Crossman and Simpson (1984) Lake Erie, 42°17'N, 81°51'W Kent Co., Rondeau Provincial Park, June 1968 1 NMC? 88-0030 Possibly Lake Erie, 42°17'N, 81°51"'W one of the above specimens Kent Co., McLean Farm, outside 8 August 1968 ] ROMS 34267 Possibly one Rondeau Provincial Park, Lake Erie, of the above specimens 42°19'N, 81°51'W Essex Co., Point Pelee National Park, 31 March 1983 1 ROM 42752 Lake Pond, 41°57'44"N, 82°30'21"W Essex Co., Point Pelee National Park, 3 June - 18 October 1983 46 Crossman and Simpson (1984) Redhead Pond, 41°57'N, 82°30'W Essex Co., Point Pelee National Park, 3 June 1983 pl ROM 43022 Possibly part Redhead Pond, 41°57'N, 82°30'W of the above specimens Essex Co., Point Pelee National Park, 11 August 1989 1 ROM 56953 Lake Pond, 41°58'00"N, 82°30'20"W Essex Co., Point Pelee National Park, 11 August 1989 4 ROM 57157 near SE corner of Redhead Pond, 41°57'06"N, 82°30'20"W Essex Co., Point Pelee National Park, 11 August 1989 1 ROM 57164 NE corner of Redhead Pond 41°57'24"N, 82°30'20"W Rondeau Park Museum >National Museums of Canada — Canada Museum of Nature “Royal Ontario Museum Simpson 1984). In this case, they are a difficult species to sample and can be confused with the Rock Bass and Green Sunfish which are also known from the area. If we consider native species as those which arrived here between the time aquatic habitats became available with the gradual “retreat”, or fluc- tuations of the Wisconsinan ice, and the arrival of European settlers, this is not a native (indigenous) species. If the established population in Point Pelee National Park survives, the Warmouth would have to be considered a naturalized species in Ontario. The range may still be expanding (Mandrak 1989) where conditions are suitable and one might expect to find it at other sites along the shores of Lakes Erie and St. Clair, as suggested by Crossman and Simpson (1984). Biology Nothing is known of the biology of the Warmouth in Canada but, in general, its life history may be sim- ilar to that of the Green Sunfish (Smith 1979) and the Rock Bass (see Scott and Crossman 1973 for details). Information on age, growth and food for U.S. populations were provided by McMahon et al. (1984). Detailed life histories on the Warmouth have been reported from studies in lowa (Lewis and English 1949), Illinois (Larimore 1957) and Georgia (Germann et al. 1974). Other studies related to growth, length-weight relationships and population structure have been summarized by Carlander (1977), Panek and Cofield (1978), and (Guillory 1980) among others. Spawning occurs in spring and early summer over rubble or loose vegetation or debris. Spawning behav- iour is that typical of sunfishes (Cooper 1983). The spawning season in Florida is protracted, and females spawn several times in one season (Guillory 1980). In Ontario the length of season may limit this. Fecundity estimates range from 800 to 34 000 eggs per female, larger females producing more eggs. Maturity seems to be related more to size than age; depending on growth rate, Warmouth may spawn at age I or II. Size at maturity varies with latitude and food availability, ranging from 51 mm in Illinois to 102 mm in Georgia (Larimore 1957; Germann et al. 1974; Guillory 1980). As a result, one would expect individuals in Ontario to mature at lengths of 50 mm or less. The species has hybridized successfully elsewhere with many other sunfishes, including Rock Bass (Schwartz 1972, 1981; Smith 1979; Parker et al. 1985). Growth depends on habitat conditions (Panek and Cofield 1978; Guillory 1980). Individuals in 1996 some populations have been noted to grow to 152 mm in 13 months and in others only to 137 mm in six years (Cooper 1983). Several authors (e.g., Panek and Cofield 1978; Guillory 1980) have devel- oped length-weight relationships and condition fac- tors for the species. Individuals live at least eight or nine years and seldom exceed 310 mm in length or 340 mg in weight, although a O.9 kg individual was reported by the U.S. Fish and Wildlife Service (Don E. McAllister, Canadian Museum of Nature, Ottawa, Ontario; personal communication). The Warmouth is probably more piscivorous than other sunfishes; small fishes, crayfish and aquatic insects comprise the bulk of their diet (Smith 1979; Cooper 1983). Smaller individuals feed on zooplank- ton, midges and caddisfly larvae with a shift to fish and crayfish as the fish become larger (Guillory 1980; Cooper 1983). Little is known of parasites and diseases. Benz and Pohley (1980) have recorded the presence of the nematode Philometra sp. from the occulo-orbit of centrarchids from Georgia, including Chaenobryttus gulosus. Limiting Factors Basically the Warmouth is a warm-water species and its range extension may be limited by seasonal water temperature. Further study is required, but under present conditions, the species probably would not be found much further north in Canada than at the present latitude (see Mandrak 1989 for potential range expansion under climate warming). Except when spawning, the Warmouth is solitary, not forming schools (Cross 1967), and does not dominate or eradicate other populations of syntopic centrarchids such as the Green Sunfish or Bluegill, Lepomis macrochirus (Smith 1979). Guillory (1980) found it was usually one of the least abundant of the centrarchids. Availability of suitable forage species and predation by larger fishes such as the Largemouth Bass, Micropterus salmoides, occupy- ing the same habitat would limit populations. Smith (1979) indicated that siltation, drainage of natural marshes and lakes, and other disruptions leading to destruction of aquatic vegetation has led to a decline of the species in Illinois. The nematode Philometra sp., if present in Canada, would not be limiting. Special Significance of the Species The species is too rare in Canada to be of impor- tance either economically or as a forage species. However, its presence here is of zoogeographic interest as it presents another example of the recent northward dispersal of freshwater fishes. Although small and solitary in nature, in the United States the Warmouth-is common throughout most of its range and is popular in some areas as a CROSSMAN, HOUSTON, AND CAMPBELL: STATUS OF THE WARMOUTH 499 sport fish (Clay 1962), but less so than the Bluegill or crappies. It is a naturalized species with the poten- tial of adversely impacting native species and bio- diversity. Evaluation If the populations in Ontario survive, the species would have to be considered a naturalized part of the aquatic fauna of Ontario. Although the population in Point Pelee National Park is probably secure, the species should be considered rare in Canada where it is at the northeastern fringe of its range. Populations in Ohio are close to those of Point Pelee and the gen- eral distribution in that State have remained relativley unchanged in the latter half of this century, but the species is uncommon where found (Trautman 1981). Until the nature of the impact of this species is determined, these populations enjoy the protection of being within Federal or Provincial Parks, are subject to few threats and are vulnerable only to natural factors. Acknowledgments The World Wildlife Fund (Canada) and the Department of Fisheries and Oceans supported the production of this manuscript. Thanks are also due to the Royal Ontario Museum, the Canadian Museum of Nature, and the Ontario Ministry of Natural Resources for provision of collection records. The final version of the manuscript was improved by information provided by N. E. Mandrak and E. Holm, and by suggestions by reviewers. Literature Cited Billington, N., and P. D. N. Hebert. Editors. 1991. International Symposium on “The ecological and genetic implications of fish introductions (FIN).” Canadian Journal of Fisheries and Aquatic Sciences 48 (Supplement 1). Benz, G. W., and W. J. Pohley. 1980. A conspicuous Philometra sp. (Nematoda: Philometridae) from the oculo-orbits of centrarchid fishes. Proceedings of The Helminithological Society of Washington 47(2): 264-265. Carlander, K. 1977. Handbook of freshwater fishery biol- ogy. Volume II. Iowa State University Press, Ames, Iowa. Clay, W. M. 1962. A field manual of Kentucky fishes. Kentucky Department of Fish and Wildlife Resources, Frankfurt, Kentucky. Cooper, E. L. 1983. Fishes of Pennsylvania and the north- eastern United States. Pennsylvania State University Press, University Park, Pennsylvania. Cross, F. B. 1967. Handbook of fishes of Kansas. University of Kansas Museum of Natural History. Miscellaneous Publications 45. Crossman, E. J., and R. C. Simpson. 1984. Warmouth, Lepomis gulosus, a freshwater fish new to Canada. Canadian Field-Naturalist 98(4): 496-498. Germann, J., L. McSwain, D. Holder, and C. Swanson. 1974. Life history of the Warmouth in the Swannee 500 River and Okefenokee Swamp, Georgia. Proceedings of the Annual Conference of the Southeast Association of Game and Fish Commissioners 28: 259-278. Guillory, V. 1979. Species asemblages of fish in Lake Conway, Florida. Science 42: 158-162. Guillory, V. 1980. Life history of warmouth in Lake Conway, Florida. Proceedings of the Annual Conference of the South East Association of Fish and Wildlife Agencies 32: 490-501. Kohler, C. C., and W. R. Courtenay, Jr. 1986. American Fisheries Society position on introductions of aquatic species: a review of past initiatives. Fisheries 11(2): 39-42. Larimore, R. 1957. Ecological life history of the war- mouth (Centrarchidae). Bulletin of the Illinois Natural History Survey 27: 1-83. Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E. McAllister, and J. R. Stauffer Jr., Editors. 1980. Atlas of North American freshwater fishes. North Carolina Biological Survey Publication Number 1980-12. Lewis, W., and T. English. 1949. The warmouth, Chaenobryttus coronarius (Bartram), in Red Haw Hill Reservoir, lowa. Iowa State College Journal of Science 23: 317-322. Mandrak, N. E. 1989. Potential invasion of the Great Lakes by fish species associated with climatic warming. Journal of Great Lakes Research 15: 306-316. McMahon, T. E., G. Gebhart, O. E. Maughan, and P. C. Nelson. 1984. Habitat suitability index models and instream flow suitability curves: warmouth. U.S. Fish and Wildlife Service, Fort Collins, Colorado. PWS/OBS-82/10.67. 21 pages. Panek, F. M., and C. R. Cofield. 1978. Fecundity of bluegill and warmouth from a South Carolina blackwater lake. The Progressive Fish-Culturist 40(2): 67-68. THE CANADIAN FIELD-NATURALIST Vol. 110 Parker, H. R., D. P. Philipp, and G. S. Whitt. 1985. Relative developmental success of interspecific Lepomis hybrids as an estimate of gene regulatory divergence between species. Journal of Experimental Zoology 233: 451-466. Pflieger, W. L. 1971. A distributional study of Missouri fishes. University of Kansas Museum of Natural History Publications 20(3): 225-570. Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 183. [Third Printing 1979] Schwartz, F. J. 1972. World literature to fish hybrids with an analysis by family, species, and hybrid. Publications of the Gulf coast Research laboratory Museum. Number 3. Schwartz, F. J. 1981. World literature to fish hybrids with an analysis by family, species, and hybrid. Supplement 1. National Ocean and Atmospheric Administration Technical Report NMFS SSRF-750. Smith, P. W. 1979. The fishes of Illinois. University of Illinois Press, Urbana, Illinois. Trautman, M. B. 1981. The fishes of Ohio. Ohio State University Press, Columbus, Ohio. Revised Edition. Wainwright, P. C., and G. V. Lauder. 1992. The evolu- tion of feeding biology in sunfishes (Centrarchidae). Pages 472-491 in Systematics, historical ecology, and North American freshwater fishes. Edited by R. L. Mayden. Stanford University Press, Stanford, California. Williams, J. E., J. E. Johnson, D. A. Hendrickson, S. Contreras-Balderas, J. D. Williams, M. Novarro- Mendoza, D. E. McAllister, and J. E. Deacon. 1989. Fishes of North America, endangered, threatened, or of special concern. 1989. Fisheries 14(6): 2-21. Accepted 13 March 1996 | The Status of the Bearded Seal, Erignathus barbatus, in Canada* Hoty J. CLEATOR Department of Fisheries and Oceans, Freshwater Institute, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Cleator, Holly, J. 1996. The Status of the Bearded Seal, Erignathus barbatus, in Canada. Canadian Field-Naturalist 110(3): 501-510. Bearded Seals, Erignathus barbatus, reside year-round in Canadian arctic and sub-arctic waters. They are patchily distrib- uted at relatively low densities throughout their range, typically inhabiting areas of broken ice and open water with depths of less than 200 m. Estimates of population size are not available, although summing different indices obtained over the past 35 years for several different regions suggest that a minimum of 190 000 Bearded Seals may inhabit Canadian waters. Hunt sta- tistics collected between the late 1950s and mid-1980s for 46 arctic communities indicated an annual harvest of approxi- mately 3000 to 5000 animals. Current rates of harvest are available only for a few arctic communities, but hunters appear to be taking fewer seals now than they did several decades ago. Information on current sightings and catch-per-unit-effort obtained from hunters across the Arctic suggest that Bearded Seal stocks are stable in Canada. The collection of harvest information should be resumed and continued on a routine basis in all northern communities. Better estimates of age-specific rates of natural mortality and pregnancy are also needed, as well as current population size and growth rates. Les phoques barbus, Erignathus barbatus, vivent pendant toute l’année dans les eaux arctiques du Canada. Ils sont répartis par petits groupes épars dans toute leur aire, en général, sur des banquises disloquées et dans les eaux libres de glace a des profondeurs de moins de 200 m. On ne dispose pas a l’heure actuelle d’ estimations précises de la taille de la population et du taux de capture dans les eaux canadiennes. Toutefois, des données de relevés et des statistiques de chasse recueillies entre la fin des années 1950 et le début des années 1980 ont indiqué que le troupeau comptait au moins 190 000 bétes et que les cap- tures annuelles atteignaient environ 3 000 a 5 000 phoques. D’aprés des observations sur le terrain des phoques barbus et des informations sur les captures par unité d’ effort fournies par les utilisateurs de la ressource, les phoques barbus ne seraient pas en danger au Canada. La population canadienne n’est pas en danger immeédiat; pourtant, il faudrait reprendre la collecte de statistiques sur les captures. I] faut aussi de meilleures estimations du taux de mortalité par age et du taux de grossesse pour l Arctique canadien, ainsi que du taux de croissance actuel des populations et de leur taille. Key Words: Erignathus barbatus, Bearded Seal, phoque barbue, arctic marine mammals., Phocidae, pinnipeds, seals, status. southern end of the Okhotsk Sea (44°N) in the Pacific Ocean and northeastern Newfoundland (50°N) in the Northwest Atlantic Ocean (Allen 1880; Bearded Seals, Erignathus barbatus Erxleben 1777, are large phocids that inhabit arctic and sub- arctic waters. Adults (Figure 1) are typically grey in colour and unpatterned, although some animals have a brownish or reddish colouration, especially on the face and foreflippers. The external characteristics that distinguish this species from other northern pho- cids are square-shaped foreflippers, a disproportion- ately small head, numerous long mystacial vibrissae, and four teats. Adults average 2.1 to 2.4 m in length and attain maximum weights of 350 to 360 kg (Chapskii 1938; McLaren 1958a; Johnson et al. 1966; Burns 1967, 1981; Benjaminsen 1973). Some studies have reported that females are slightly longer than males (McLaren 1958a; Johnson et al. 1966; Burns 1967; Benjaminsen 1973), but there is no evi- dence to verify that these differences are statistically significant. Distribution The distribution of Bearded Seals is circumpolar and boreoarctic and extends as far south as the Ognev 1935; King 1983). A few extralimital occur- rences, of juveniles mostly, have been reported in the waters off Spain and Portugal, 40 to 44°N (Ray et al. 1982). In North American waters, strays have been found as far south as Cape Cod (Mansfield 1967) and recently a yearling male was found in poor con- dition on the southern shore of the St. Lawrence Estuary (Gosselin and Boily 1994). Within Canada, Bearded Seals reside year-round in arctic waters as well as in Hudson Bay, Hudson Strait, Labrador Sea and probably in James Bay (Mansfield 1967; Mansfield et al. 1975; Smith and Taylor 1977; Boles et al. 1980; Ray et al. 1982) (Figure 2). A small resident population in the waters off southern Labrador and southern and eastern Newfoundland may have been recently extirpated (Ray et al. 1982). Two subspecies have been suggested: Erignathus barbatus barbatus is distributed from the central * Reviewed and approved by COSEWIC 14 April, 1994, report accepted no status designation required. ~ 501 502 Canadian Arctic east to the Laptev Sea (north of Siberia) and Erignathus barbatus nauticus 1s distrib- uted from the central Canadian Arctic west to the Laptev Sea (Manning 1974). The exact boundary between the two subspecies is not clearly defined. There is disagreement on whether the differences in body size and growth rates of cranial bones between specimens taken from the two regions are sufficient to warrant subspecies distinction (Kosygin and Potelov 1971; Manning 1974; Smith 1981). Protection International This species has not been assigned a designation under the Convention on International Trade in Endangered Species of Wild Fauna and Flora (CITES). National Canada The Marine Mammal Regulations of the Fisheries Act of Canada (Canada Department of Fisheries and Oceans 1993) provides for the management and con- trol of the harvest of Bearded Seals in Canadian waters. The regulations permit residents to kill seals for food purposes. Native hunters may also take seals for social or ceremonial purposes. There are no restrictions on the sale or trade of pelts resulting from the harvest, although a marine mammal trans- portation licence (export permit) is required to trans- port marine mammal material out of the Northwest Territories. No sport hunting of Bearded Seals is per- mitted. Over the past decade, two coastal land claim agreements have come into effect: the Inuvialuit Final Agreement, which covers the Yukon North Slope and the northwestern portion of the Northwest Territories and the Nunavut Land Claim Agreement, which covers the central and eastern portions of the Northwest Territories. In 1976, the James Bay and Northern Quebec Agreement was signed. It covers the territory within western and northern Quebec, including marine mammals that occur there in estu- ary areas. The Makivik Offshore Claim, which is currently under negotiation, will cover the coastal areas around western and northern Quebec. Under the terms of the three signed agreements, the respon- sibility for managing arctic marine mammal popula- tions in Canadian waters, except off Labrador, lies with public wildlife management boards. Land claim beneficiaries and representatives from the federal, territorial and/or provincial governments sit on these boards. The Minister of Fisheries and Oceans is obliged to implement management decisions made by the public wildlife management boards except those that conflict with “disallowance provisions” (e.g., decisions that would conflict with the princi- ples of conservation) in the agreement. THE CANADIAN FIELD-NATURALIST Vol. 110 United States The United States Marine Mammal Protection Act of 1972 permits coastal Alaskan natives to harvest Bearded Seals for food or the production of cultural- ly authentic handicrafts and clothing. Commercial harvesting is prohibited. Population Size and Trend No recent population estimates are available for the entire Canadian Arctic, however earlier estimates are available for some areas. In 1958, McLaren (1958b) estimated that the waters of Foxe Basin, Hudson Bay, Hudson Strait/Ungava Bay, and off eastern and northern Baffin Island contained 186 000 Bearded Seals. This crude estimate was based on catch statistics (excluding animals struck and lost), the relative proportion of Bearded Seals to Ringed Seals, Phoca hispida, sighted during shipboard sur- veys, and the amount of suitable habitat within 16 km of shore and water depths of less than 200 m. In the past twenty years, population indices for Bearded Seals have been reported for parts of the Western Arctic and High Arctic based on aerial sur- veys. These population estimates have not been cor- rected to include animals that were below the surface of the water at the time of the survey because no cor- rection factors exist. Indices for the eastern Beaufort Sea in 1974 to 1979 range between 1200 and 3100 animals (Stirling et al. 1977, 1982; Kingsley and Lunn 1983). Aerial surveys flown in the High Arctic between the mid-1970s and early 1980s revealed low numbers of Bearded Seals (maximum densities: 0.05 to 0.24 seal/sq km) in areas of preferred habitat (Finley 1976; Koski and Davis 1979; Koski 1980a,b; Koski and Davis 1980; Kingsley et al. 1985); no population estimates were calculated for these sur- veys. Surveys conducted in the offshore waters of western Baffin Bay and eastern Lancaster Sound in late June-early July 1979 produced uncorrected esti- mates of 7400 to 9500 seals (Koski and Davis 1980). Few reliable harvest statistics are available for Bearded Seals because they are taken for domestic purposes, and in relatively small numbers. However, some harvest records were kept for 46 communities between the late 1950s and mid-1980s (Beaubier et al. 1970; Smith and Taylor 1977; Native Harvesting Research Committee 1976, 1979, 1982a,b; Smith 1981; Donaldson 1988; Pattimore 1985, 1986; Gamble 1984, 1987a,b). Roughly 2400 Bearded Seals were reported taken on an annual basis in the Canadian Arctic, excluding animals that were struck- and-lost. Losses due to sinking have not been ade- quately quantified, however in open-water hunts it is common for more than 50% of the animals struck to sink before they can be retrieved (Burns 1967). During the spring hunt, when seals are usually shot while they are hauled out on the ice to bask, struck- and-loss rates drop to approximately 25% (Burns and 1996 CLEATOR: STATUS OF THE BEARDED SEAL 503 FIGURE |. Bearded Seal hauled out on ice (courtesy Wayne Lynch®). Frost 1979). Adjusting the reported take to account for sinking losses yields a total annual kill that was probably closer to 3000 to 5000 during that period. Although only sparse harvest data exist, the num- bers of Bearded Seals taken seem to have declined in recent years. For example, between 1962 and 1980, an average of 66 Bearded Seals were reported har- vested on an annual basis in the Western Arctic (50 from Sachs Harbour, n = seven years; 16 from Holman, n = one year) (Smith and Taylor 1977; Smith 1981), whereas between 1987 and 1993, an average of 22 seals were reported taken annually by five Western Arctic communities (nine from Sachs Harbour, n = seven yrs; eight from Holman, n = seven years) (Fabijan, 1991a,b,c, 1995a,b,c). In Grise Fiord during 12 years between 1962 to 1984, an average of 25 Bearded Seals were reported har- vested each year (Smith and Taylor 1977; Pattimore 1986; Stewart et al. 1986; Donaldson 1988) com- pared with a reported annual average catch of 13 seals from 1992 to 1993 (C. Craig, Department of Fisheries and Oceans, Winnipeg, Manitoba; personal communication). In Lake Harbour during nine years between 1962 and 1984, an average of 111 seals were reported harvested each year (Smith and Taylor 1977; Pattimore 1986; Stewart et al. 1986; Donaldson 1988) compared with an annual average of 67 seals in 1992 to 1993 (Craig, personal communication). Changes in data collection techniques and envi- ronmental conditions over the past 20-30 years may account for some differences in reported harvests. The recent data were collected in each community by personal interviews, while older data were often collected by community officials (e.g., Royal Canadian Mounted Police officers) and from fur records. The trend toward declining harvests proba- bly also reflects an overall shift away from the tradi- tional Inuit lifestyle (e.g., reduced hunting effort, increased use of southern foods and equipment, use of snowmachines instead of dog teams). A few com- munities have maintained a more traditional way of life and their take of Bearded Seals has probably remained relatively constant or possibly increased slightly. For example, Igloolik hunters still harvest numbers of Bearded Seals for food and making rope and kamiks (B. Parker, Department of Renewable Resources, Government of Northwest Territories, Igloolik, NWT; personal communication). It is unlikely that the current harvest of Bearded Seals in the Canadian Arctic exceeds 4000 animals per year. Current estimates of maximum sustainable yield (MSY) are not available for the Canadian Arctic but two older estimates have been published. Based on data from the eastern Canadian Arctic, McLaren (1958b) suggested an MSY of “about 5% of the mid- year population” as being “wholly safe” and, based 504 THE CANADIAN FIELD-NATURALIST Vol. 110 FiGuRE 2. Distribution of Bearded Seals in Canadian waters. on kill data collected from the Sea of Okhotsk, Fedoseev (1973) also reported an MSY of 5%. An examination of the Western Arctic data, for which conservative population indices and current kill data are available, shows that the current rate of exploita- tion probably removes no more than 3.3% of the population each year. While this calculation relies on a number of assumptions and historic data for one small region of the Canadian Arctic, it suggests that the Bearded Seal populations that inhabit Canadian waters may be relatively stable. This perspective is supported by hunters who have not reported declines in sightings or catch-per-unit-effort (C. Furgal, Department of Environmental Studies, University of Waterloo, Waterloo, Ontario; L. Harwood, Department of Fish- eries and Oceans, Iqualuit, Northwest Territories; Parker; K. Seto, Department of Fisheries and Oceans, Iqaluit, NWT; personal communications). Habitat Bearded Seals prefer areas of moving ice and open water in depths of less than 150 to 200 m, and tend to avoid areas of thick shorefast ice (McLaren 1958b; Burns 1967; Mansfield 1967; Davis et al. 1975; Burns and Frost 1979; Stirling et al. 1982; Kingsley et al. 1985; Cleator and Stirling 1990). In a few areas, Bearded Seals maintain breathing holes in landfast ice throughout the winter (Mansfield 1967; Stirling and Smith 1977). This is probably because the ice in those areas freezes late and breaks up early (Stirling et al. 1983), and also because benthic pro- duction there may be high (Smith 1981). Bearded Seals have been observed in waters deeper than 500 m during March and April (Finley and Renaud 1980). They regularly use pack ice for resting, pup- ping and moulting (Fay 1974). General Biology Reproductive Capability Most males reach sexual maturity at 6 or 7 years of age (McLaren 1958a; Tikhomirov 1966; Burns 1967; Burns and Frost 1979; Smith 1981). In Canadian arctic waters, most breeding likely occurs between mid-April and late May, although males are in breeding condition from about mid-March to late June (McLaren 1958a). The mating system of the Bearded Seal is unknown although they are probably polygnyous (Budelsky 1992). Females may begin to ovulate at three to four years of age and most are pregnant for the first time at five or six years of age (McLaren 1958a; 1996 Tikhomirov 1966; Burns 1967; Potelov 1975; Burns and Frost 1979; Burns 1981; Smith 1981). Reported estimates of the ovulation rate range between 0.46 and 1.00 (Stirling et al. 1977; Smith 1981) while pregnancy rates range between 0.33 and 0.85 (Burns 1967; Stirling et al. 1977; Burns and Frost 1979; Smith 1981). Typical pregnancy rates in adult females (six years and older) are 0.80 to 0.85. Most sexually mature females probably reproduce annual- ly (Tikhomirov 1966; Burns 1967; Fedoseev 1973; Burns and Frost 1979; Smith 1981); however, fertili- ty rates may decline when resources become limited in an area (Burns 1981; Smith 1981). No evidence of reproductive senility has been reported. Females with pups usually ovulate and mate after their pups are fully weaned (Burns and Frost 1979). Implantation is delayed and occurs between mid-July and early August, approximately two months after breeding (Chapskii 1938; Burns and Frost 1979; Smith 1981). The single neonates are first observed between mid-April and early May (Chapskii 1938; McLaren 1958a; Johnson et al. 1966; Potelov 1975; Burns 1981). Lactation is estimated to last only 12 to 18 days (Burns 1967). At birth, the sex ratio is close to unity, while in older age classes females are often more numerous (Sleptsov 1943; Fedoseev 1973; Burns and Frost 1979; Smith 1981). Age-specific mortality rates have been calculated from life tables based on fitted age distributions derived from Bering-Chukchi seas har- vest data (Burns and Frost 1979). The higher mortal- ity rates reported for males may account for why adult females are slightly more numerous than adult males (Burns and Frost 1979). However, the prepon- derance of females in the catch statistics may reflect the increased vulnerability of females at specific times of year or differential segregation of the sexes (Smith 1981). Harvest data suggests that the maxi- mum life span in the wild ranges between 23 and 31 years of age (Benjaminsen 1973; Burns and Frost 1979; Smith 1981). Species Movement In the Bering and Chukchi seas, many Bearded Seals make well-defined seasonal movements in order to maintain contact with ice year-round (Johnson et al. 1966; Burns 1967). Aerial survey data suggest that Bearded Seals in Baffin Bay may also move between coastal and offshore areas to stay with the ice (Koski 1980a). Over much of their range, however, Bearded Seals appear to be relative- ly sedentary, undertaking more local movements in response to ice conditions (Vibe 1950; McLaren 1962; Fedoseev 1973). Throughout their range, Bearded Seals appear to be patchily distributed at relatively low densities (Burns 1967; Burns and Frost 1979; Stirling et al. 1982), although during the winter months prior to break-up and during early summer when the availability of ice pans for haulout CLEATOR: STATUS OF THE BEARDED SEAL 505 are limited, they are likely more aggregated than at other times of the year. Behaviour/Adaptability Bearded Seals typically occur alone or in small groups. Even when several individuals are hauled out on a single large floe, they position themselves evenly around the perimeter of the pan, facing in dif- ferent directions (Burns 1981). Their response to human disturbance is varied. While hauled out on the ice during warm calm days in late spring and summer, they are relatively undisturbed by the close presence of men, boats or low-flying aircraft. During winter, however, human activity on the ice will cause a basking seal to quickly enter the water or a swimming seal to surface several hundred metres from the sound source (Burns 1981). The reasons for these differences in behaviour are unknown although in late spring and summer, hauled-out seals have more access to escape routes. Like basking Ringed Seals, Bearded Seals may also be more reluctant to enter the water once dry and warm. Bearded Seals produce loud trill-like calls, that can last up to about 30 seconds in duration and can be heard for distances of 25 km or more underwater, between March and June (Stirling et al. 1983; Cleator et al. 1989). Limited evidence suggests that only males call and that they may use these calls to advertise breeding condition, territoriality, or both (Dubrovskii 1937; Chapskii 1938; Ray et al. 1969; Burns 1981; Stirling et al. 1983; Cleator et al. 1989). Typically, calling seals are distributed in a random arrangement, one to three km apart, and remain fair- ly stationary; this species may use lek-display (Budelsky 1992). There is evidence of geographical variation in the call repertoire, that appears to be rel- atively stable from one year to the next and may be characteristic of discrete breeding stocks (Cleator et al. 1989). Bearded Seals will consume a wide variety of food items including pelagic fishes (Burns and Frost 1979; Lowry et al. 1980; Finley and Evans 1983; Antonelis 1993); however, they are primarily benthic feeders and usually the bulk of their diet is made up of only a few species. Prey selection appeais to be based on the availability of prey, therefore the diet may vary within a relatively small area and over a short period (Antonelis et al. 1993). Limiting Factors Predation Polar Bears (Ursus maritimus) are the chief preda- tors of Bearded Seals. Limited data suggest that at least half of the Bearded Seals killed by bears are pups and subadults (Stirling and Archibald 1977). Occasionally, parts of young Bearded Seals have been found in the stomachs of walruses. Whether walruses actively prey on Bearded Seals or merely eat them as carrion has not yet been determined, 506 however indirect evidence of predation has been reported (Lowry and Fay 1984). Bearded Seals are often infested with large numbers of internal para- sites (see review by Burns 1981); however, it is not known to what extent parasitism contributes to nat- ural mortality. It is likely that Bearded Seal popula- tions near communities will continue to be hunted and hence locally depressed in numbers. However, the importance of this species to Inuit has diminished in many communities in recent years and some pop- ulations may be located so far from communities that they are not subject to human hunting pressure. If this situation persists, it is unlikely hunting will be a significant limiting factor on Bearded Seal popula- tions across the Canadian Arctic. Environmental Contamination and Human Disturbance DDT and related residues, PCBs, and heavy met- als (i.e., mercury, selenium, nickel, copper, zinc, and cadmium) are present in the tissues of Bearded Seals (see reviews by Burns 1981 and Kelly 1988). Only a few studies have been conducted in Canadian waters to date. Analysis of samples collected from animals in eastern Amundsen Gulf and eastern Hudson Bay has shown that Bearded Seals accumulate large amounts of mercury and selenium in the liver and that there may be regional differences in the level of contamination (Smith and Armstrong 1975, 1978). Positive correlations have been detected between mercury level and age, as well as between selenium level and age (Freeman and Horne 1973; Smith and Armstrong 1975, 1978). Concentration levels of PCB’s and other organochlorine contaminants have been examined in small samples of Bearded Seals and Ringed Seals caught in the waters off northern Quebec and Baffin Island (Thomas 1990; Muir and Rosenberg 1990). In general, concentration levels of organochlorine cont- aminants in the blubber are considerably lower in Bearded Seals. This difference may reflect the lower trophic level at which Bearded Seals typically feed compared to Ringed Seals (i.e., benthic organisms versus fish). Little is known about the effects of pes- ticides and heavy metals on marine mammals, and it is not known if they contribute to mortality in this species. Aside from local reductions in population num- bers as a result of hunting, the most serious threat to this species is industrial activity. Bearded Seals may be vulnerable to even relatively minor environmental disturbances because they are distributed in small, localized populations. Dredging, dumping, and acci- dental spills that affect benthic productivity, for example, are a concern in areas where Bearded Seals occupy shallow waters (e.g., southeastern Beaufort Sea) and rely solely on benthic species for food (see review by Kelly 1988). In deeper areas (e.g., Lancaster Sound) where Bearded Seals feed on a THE CANADIAN FIELD-NATURALIST Vol. 110 wider variety of prey, including pelagic fish, they are probably at less risk. Oil spills also pose a direct risk to the health of Bearded Seals that inhabit waters near a spill. Although direct exposure to oil has not been investi- gated in Bearded Seals, it would likely cause stress, eye irritations or damage, and accumulate in body tissues and fluids thereby causing tissue damage, as has been reported for other northern pinnipeds (Smith and Geraci 1975; Engelhardt et al. 1977; Geraci and Smith 1977). Mortality due to oiling is likely to be most significant in pups and older ani- mals, and during the moult when seals are undergo- ing physiological stress (Smith and Geraci 1975; Davis and Anderson 1976; Geraci and Smith 1977). Bearded Seals may have enzymatic mechanisms for metabolizing and excreting components of crude oil similar to those found in Harp Seals (Lyle Lockhart, Department of Fisheries and Oceans, Winnipeg, Manitoba, unpublished data). The effects of chronic exposure to weathered oil are unknown. Bearded Seals prefer areas of moving pack ice and open water, which also offer the best navigation routes for ships travelling through ice-choked arctic waters. High noise levels produced by vessels and other industrial activities can cause hearing damage in pinnipeds (see review by Myrberg 1990); howev- er, the Bearded Seals’ preference for pack ice with cracks and leads allows them to move away from most disturbance (Mansfield 1983). In spring, in areas where seal movement is restricted by ice and prolonged exposure to underwater noise from indus- trial operations (e.g., drill ships) could mask auditory communication, disruption of breeding activities and social organization may occur (Cleator and Smith 1994). Species Competition While there doesn’t appear to be an immediate interaction between this species and any commercial fisheries, there is currently a commercial fishery for Pandalid shrimp in eastern Davis Strait and a number of exploratory fisheries for whelks (Buccinum sp.), shrimp (Pandalus borealis and P. montagui), and Iceland Scallops (Chlamys islandica) off the eastern coast of Baffin Island and in Ungava Bay and Hudson Strait (D. Chiperzak, Department of Fisheries and Oceans, Inuvik, NWT; personal com- munication). As Bearded Seals appear to vary their diet according to the availability of different prey species in an area, the introduction of a commercial fishery may have little effect on them as long as abundant alternate prey are available. Bearded Seals appear to avoid areas heavily used by Walruses (Odobenus rosmarus). Inter-specific competition for benthic prey or predation by walrus- es may explain the low numbers of Bearded Seals in areas frequented by Walruses (Cleator and Stirling 1990). | 1996 Special Significance of the Species Historically, Bearded Seals were a valuable resource to the Inuit, who used their meat for food, their blubber for fuel, and their skin for dog-team traces, harpoon lines, footgear, and any other use that required strength and durability. However, with increased availability of substitutes in the past 20 years, its importance to many local economies has declined. Bearded Seals continue to be important prey for Polar Bears (Stirling and Archibald 1977), especially in areas of active ice (Smith 1980). Canada is under international obligation to protect the Polar Bear populations that reside within its boundaries so it is essential that Bearded Seal popu- lations are also protected. Evaluation There are no recent or accurate survey data avail- able to determine current population levels in Canadian waters or to permit the evaluation of popu- lation trends. However in the Western Arctic, current harvest data and abundance indices collected twenty years ago, and the fact that hunters have not raised concerns there about declining numbers of Bearded Seals, suggest that the current populations in that region may be relatively stable. Hunters in other regions of the Canadian Arctic also have not report- ed a decline in sightings or in catch-per-unit-effort. In northern Foxe Basin, for example, hunters report that Bearded Seals are abundant and that they still take considerable numbers of them (Parker, personal communication). Anecdotal comments like these suggest that most, if not all, Canadian Bearded Seal populations may be relatively stable in spite of hunt- ing pressure near communities. Nevertheless, given the patchy distribution and paucity of data on this species, each population should be treated as a dis- crete stock and managed on the basis of accurate and current estimates of population size and growth and data on rates of harvest and hunting loss. Only in this way, can we hope to ensure the long-term sustain- ability of this species in northern Canadian waters. Acknowledgments The preparation of this status report was funded by the Department of Fisheries and Oceans. I thank Susan Cosens, Lois Harwood, Stuart Innes, Pierre Richard, Barb Stewart, Rob Stewart, and three anonymous reviewers for constructive comments on the manuscript. Wayne Lynch generously provided the photograph. Literature Cited Allen, J. A. 1880. History of North American pinnipeds. U.S. Geological and Geographical Survey, Washington. Miscellaneous Publication 12. 785 pages. Antonelis, G. A., S. R. Melin, and Y. A. Bukhtiyarov. 1993. 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Interrelationships of THE CANADIAN FIELD-NATURALIST Vol. 110 arctic ocean mammals in the sea ice habitat. Pages 129-136 in Proceedings of the Circumpolar Conference on Northern Ecology. Volume II. National Research Council of Canada, Ottawa. Stirling, I., R. Archibald, and D. DeMaster. 1977. Distribution and abundance of seals in the eastern Beaufort Sea. Journal of the Fisheries Research Board of Canada 34: 976-988. Stirling, I, M. Kingsley, and W. Calvert. 1982. The dis- tribution and abundance of seals in the eastern Beaufort Sea, 1974-79. Canadian Wildlife Service Occasional Paper 47. 25 pages. Stirling, I., W. Calvert, and H. Cleator. 1983. Underwater vocalizations as a tool for studying the dis- tribution and relative abundance of wintering pinnipeds in the High Arctic. Arctic 36: 262—274. Thomas, D. J. 1990. Organochlorine residues in biota of the Canadian arctic. Seakem Group, Sidney, B.C. 91 pages. Tikhomirov, E. A. 1966. Reproduction of seals belong- ing to the family Phocidae in the North Pacific. Zoologicheskii Zhurnal 45: 275-281. (Fisheries Research Board of Canada Translation Series 1889). Vibe, C. 1950. The marine mammals and marine fauna in the Thule District (northwest Greenland) with observa- tions on ice conditions in 1939-41. Meddelelser om Grgnland 140: 1-117. Accepted 13 March 1996 Canada* _ Dawn NELSON! and Jon LIEN? | The Status of the Long-finned Pilot Whale, Globicephala melas, in | 'Biopsychology Programme, Memorial University of Newfoundland, St. John’s, Newfoundland A1B 3X9 Ocean Sciences Centre and Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland A1C5S7 Nelson, Dawn, and Jon Lien. 1996. The status of the Long-finned Pilot Whale, Globicephala melas, in Canada. Canadian | Field-Naturalist 110(3): 511-524. The Long-finned Pilot Whale, Globicephala melas, is a mainly pelagic species widely distributed in the cold temperate waters of the North Atlantic and the Southern Hemisphere. It regularly migrates in summer to Canadian inshore waters fol- | lowing spawning squid. Drive fisheries from 1947 to 1971 seriously depleted numbers of Globicephala melas off Newfoundland. Mass strandings represent a major known source of natural mortality for this species. The effects of inci- dental entrapments, pollutants, and fisheries for prey species remain relatively unknown, but these factors have the poten- tial for limiting this species. Globicephala melas has recently been used in satellite tracking and DNA fingerprinting stud- ies, and is a common subject in the study of mass strandings. There are few reliable recent population estimates for Globicephala melas, but even optimistic recovery forecasts based on drive fisheries in Newfoundland would produce a pre- sent population substantially lower than pre-whaling numbers. Given that there are no immediate threats to the population a COSEWIC status designation would not seem to be warranted at this time. Le globicéphale noir, Globicephala melas, est une espéce plutdt pélagique retrouvée un peu partout dans les eaux froides | tempérées de |’ Atlantique nord et de |’hémisphére austral. En été, il migre réguli¢rement vers les eaux cOtiéres canadiennes ala recherche de calmars. La chasse par rabattage effectuée de 1947 a 1971 en a décimé les effectifs dans les eaux cdtiéres de Terre-Neuve. Les echouages en masse sont une importante cause connue de mortalité, par contre, les incidences de cap- tures accidentelles, les élements polluants et l’exploitation des espéce-proies demeurent relativement mal connues. Elles ont toutefois un potentiel comme facteurs limitatifs de ’espece. Globicephala melas a été 1 objet d’études de repérage par | satellite et d’identification a ! ADN, ainsi que des échouages en masse. Aucune prévision démographique récente fiable nest disponible en ce qui concerne Globicephala melas, mais méme des prévisions optimistes quant au rétablissement des | effectifs décimés par la chasse par rabbatage effectuée dans les eaux de Terre-Neuve ne donneraient qu’un effectif nette- | ment inférieur a l’abondance pré-exploitation. Puisque elles ne font pas face a des menaces immédiates, une désignation de | statut par le CSEMDC n’est pas justifiée 4 ce moment-ci. Key Words: Long-finned Pilot Whale, globicéphale noir, Globicephala melas (= Giobieepuain melaena), toothed whales, | Odontoceti, Cetacea, marine mammals, status. The Long-finned Pilot Whale, Globicephala melas (Traill 1809), is variously known as the Northern |Pilot Whale, Atlantic Pilot Whale, Pothead, | Blackfish, Calling Whale, and Caa’ing Whale. It has | often been cited as Globicephala melaena, although | Globicephala melas is the more taxonomically cor- rect form of the name (Rice 1989). The species is characterized by a bulbous fore- | head, a falcate dorsal fin with a long base located far jeorward on the body, a slight beak, and sharply pointed pectoral flippers that may reach one-fifth of | the body length (Figure 1). It is slate-grey to black in ‘colour, although pale or albino individuals are some- times observed (Sergeant and Fisher 1957; Hain and Leatherwood 1982; Bloch 1994). An anchor-shaped | patch of greyish-white on the throat extends into a | grey stripe along the underside which expands fiscrae the navel and surrounds the genital area (Sergeant and Fisher 1957). Other markings may include a variable grey saddle behind the dorsal fin and a grey or white streak behind the eye, both of which may be more common in Southern Hemisphere populations (Scott 1942; Sergeant and Fisher 1957; Davies 1960; Aloncle 1972; Aguayo 1975). Young animals are lighter in colour overall, and the saddle and eye blaze, if present, apparently become evident after three to five years of age (Starrett and Starrett 1955; Sergeant and Fisher 1957; Bloch et al. 1994). Globicephala melas can be distinguished from the closely-related Short-finned Pilot Whale, Globicephala macrorhynchus, by differences in range, size, external markings, length of pectoral flippers, size of tail flukes, tooth count, and skull characteristics (Fraser 1950; Yonekura et al. 1980; Leatherwood and Reeves 1983; Bloch et al. 1994). Most differences are not entirely reliable, however, as there is some overlap in range and physical char- _*Reviewed and approved by COSEWIC 14 April 1994, report accepted; no status designation required. Sil 512 acteristics (Leatherwood et al. 1976; Van Bree et al. 1978; Yonekura et al. 1980; Casinos 1981; Powers et al. 1982; Nores and Perez 1988; Bloch et al. 1994). The teeth may provide the best distingushing charac- teristic: those of Globicephala macrorhynchus are generally fewer (seven to nine instead of 10 to 12 in both jaws) and larger (Sergeant 1959). Globicephala melas shows marked sexual dimor- phism. Males can reach lengths of approximately 620 cm and weights of three tons, while females are approximately 18 to 25% smaller (Sergeant 1962; Martin et al. 1987; Kasuya et al. 1988b; Desportes 1990; Bloch 1994). Males also have longer flippers and longer and wider flukes than females, and more often have the eye-streak marking (Bloch et al. 1994). Distribution Globicephala melas is widely distributed in cold temperate waters of the North Atlantic and southern oceans (Figure 2). The northern and southern forms, which are widely separated geographically and may vary in coloration, are sometimes regarded as sub- species: Globicephala melas melas in the north, and Globicephala melas edwardi in the south (Davies 1960; Aguayo 1975; Mitchell 1975a; Van Bree et al. 1978). The northern population ranges from Greenland, Iceland, the Barents Sea, and possibly the Baltic sea in the north, to Cape Hatteras in the west, and north- west Africa (including the Mediterranean) in the east (Sergeant and Fisher 1957; Mitchell 1975a; Leatherwood and Dahlheim 1978; Evans 1980; Nores and Perez 1988). In the eastern Atlantic, Globicephala melas appears to be rare in Italian waters, on the coasts of the Netherlands and Belgium, and the east coasts of Britain and Ireland (Evans 1980). It may or may not be present in the North Sea (Leatherwood and Dahlheim 1978; Evans 1980). In the western Atlantic, it is numerous in the region of Georges Bank, Scotian Shelf, outer Laurentian Channel, and Grand Bank from July - December, but is absent from inshore Labrador waters during summer (Sergeant 1979). Globi- cephala melas is found throughout the Gulf of St. Lawrence, although it seems to be more abundant in the southern portion of the Gulf and along the west coast of Newfoundland (Sergeant and Fisher 1957; Sergeant et al. 1970; Dunbar et al. 1977; Sears et al. 1981; Sears 1982). Globicephala melas is common off the east coast of the U.S., although abundance varies greatly. They are found along the shelf break from Cape Hatteras to the eastern tip of George’s Bank (Hain et al. 1981). In summer, they move from the shelf edge onto George’s Bank and into the Gulf of Maine (Hain et al. 1981; Powers et al. 1982), although they are present on George’s Bank throughout the year. THE CANADIAN FIELD-NATURALIST Vol. 110 Sightings are most common in the southern New | England mid-shelf and shelf-break in fall and winter | (Powers et al. 1982). | In the Southern Hemisphere, Globicephala melas | occurs mainly north of the Antarctic Convergence in | the cold currents (Humbolt, Falkland, and Benguela) | associated with the West Wind Drift (Mitchell | 1975a; Leatherwood and Dahlheim 1978; Van Bree | et al. 1978; Guiler et al. 1987; Miyazaki and Kato | 1988). Long-finned Pilot Whales inhabited the Sea of Japan until the 12th century, although there is no recent evidence of them in the Bering sea or North Pacific (Kasuya 1975; Kasuya et al. 1988a). | Sometimes, individuals of Globicephala | macrorhynchus taken off the coast of Japan are mis- | takenly identified as Globicephala melas (Ohsumi | 1975; Kasuya et al. 1988a). Globicephala melas appears to be distributed con- tinuously across the North Atlantic (Brown 1961; | Leatherwood and Dahlheim 1978). Separate stocks | have yet to be conclusively distinguished, although there is some evidence that the western and eastern } North Atlantic populations are distinct. Eastern ani- | mals appear to be slightly larger, and although the } population around Newfoundland was depleted by a | drive fishery in the 1960s, the eastern population | showed no corresponding decline in numbers | (Sergeant 1962; Mercer 1975; Mitchell 1975a,b; Moore et al. 1979; Martin et al. 1987; Bloch 1994). | Declines were also not observed in areas near | Newfoundland, such as Nova Scotia (Sergeant 1982). This may indicate the presence of subpopula- | tions within the western North Atlantic. | Although there is incomplete census data, Andersen (1988) hypothesized a parapatric distribu- tion between the east and west Atlantic, resulting in a limited exchange of genes between the two areas. Further, differences in the types of parasites carried by pilot whales from the Faroe Islands, the western ) Mediterranean, eastern North Atlantic waters near | France, and Newfoundland, suggest that individual whales may not routinely move between any of these regions (I.W.C. 1990a). However, recent increases in mass strandings reported from both sides of the: Atlantic suggest the possibility of a continuous dis- - tribution across the North Atlantic (D. E. Sergeant, | McGill University, Main Road, Hudson, Quebec; ; personal communication). In the eastern Atlantic, enzyme variation within and between schools of whales caught in the Faroes may imply some degree of reproductive isolation among schools (Andersen 1988). Further, differing concentrations of the elements mercury, cadmium, | and selenium found among Faroese whales suggests the existence of at least two sub-populations (Jean-Caurant 1987; Julshamn et al. 1987). Faroese | pilot whales have been historically classified into |) | | | 1996 > Ti Unum Tht ae NELSON AND LIEN: STATUS OF THE LONG-FINNED PILOT WHALE Sls) FicureE |. Long-finned Pilot Whale, Globicephala melas, (Drawing by D. Nelson, approximately 1/50 life size). one of two types depending upon the shape of the dorsal fin, but these differences are now attributed to yearly variations in blubber thickness (Andersen 1988; Bloch et al. 1994). In waters of the British Isles, there is some evi- dence of at least two separate populations; a norther- iy one which may be part of the stock centered on _ the Faroes, and a more southerly one (Evans 1980). | Protection | International Pilot whales are listed on Appendix I of the | Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES). _ National Canada: All whaling, except aboriginal, is prohib- ited in Canadian waters under the Marine Mammal ) Regulations the Fisheries Act of 1867 (as amended ' to date). Hunting can only be undertaken under | license. United States: Protection is afforded under the Marine Mammal Protection Act of 1972 as well as by the Packwood-Magnuson Amendment of the | Fisheries and Conservation Act, and the Pelly - Amendment of the Fisherman’s Protective Act. Population Size and Trends There are inadequate estimations of abundance for the entire northern or southern population of _Long-finned Pilot Whales. Western Atlantic A drive fishery existed at Cape Cod from the / mid-1700s to the 1920s, with the mean annual catch ' in the 1800s on the order of 2000 to 3000 animals | (Mitchell 1975b; Mead 1979). Although Globicephala melas was exploited to _ some extent in Newfoundland even before the 1900s, it became the target species of organized drive fish- _eries from 1947 to 1971, which took approximately 54 000 animals within this time span (Mercer 1975; | Sergeant 1982). Most whales were captured in Trinity Bay and Bonavista Bay, with fewer in Conception Bay and Notre Dame Bay (Sergeant 1962; Mercer 1975). The population in eastern Newfoundland waters is thought to have numbered 50 000 to 60 000 at the onset of the fishery, and was apparently decimated by the hunt. A peak kill of 10 000 whales in 1956 gave way to a steady decline, until annual catches numbered in the low hundreds (Mitchell 1974; Mercer 1975; Sergeant 1982). The whales were usu- ally herded by boats into shallow bays and killed with lances, although a few hundred were harpooned annually by crews on whaling vessels (Sergeant 1962). In 1971, the company which processed the animals reported to the Department of Fisheries that “Potheads” were virtually commercially extinct in Newfoundland. The drive fishery ended when com- mercial whaling on the Canadian Atlantic coast was banned by the Government of Canada on December 22, 1972 (Mercer 1975). There is no evidence of depletion in any other stocks of Globicephala melas. Some authors feel that the Newfoundland popula- tion may be recovering (Mitchell 1975a; Sergeant 1982), however, there is little new information on the status of the Newfoundland population. The most recent estimates for northeastern populations yield 4000 to 12 000, and sightings are relatively infre- quent (Hay 1982; Lynch 1987; I.W.C. 1990b; Lien 1980; Lien et al. 1980; Lien, unpublished data). It is unlikely that recovery is complete. If one assumed a Newfoundland population of 4000 animals in 1972, present numbers estimated from an optimistic 6% net recruitment rate would only be around 12 000 animals. Eastern Atlantic Statistics on the non-commercial pilot whale drive in the Faroe Islands have been found dating as early as 1584, and an unbroken record exists from 1709 to the present (Bloch et al. 1990a). The annual catch between 1709 and 1989 varied from zero to 4325 whales, with an average of 988 whales caught per year (Bloch et al. 1990a). In 1987, an estimate of 100 000 whales was made for waters east of Greenland, with the most animals occurring south- west of the Faroe Islands (Desportes 1990). There is no evidence of depletion, although uncertainties in abundance numbers and reproductive rates make 514 assessments tentative (I.W.C. 1990a). Hunting tech- niques in the Faroe Islands are similar to those that were utilized in the Newfoundland drive fishery (Bloch et al. 1990b). Small numbers of animals have been taken in Norway, West Greenland, Iceland, Scotland, and Ireland (Christensen 1975; Kapel 1975; Mitchell 1975a,b; O’Riordan 1975; Sergeant 1979; I.W.C. 1990b). Population estimates for eastern Atlantic waters yield approximately 100 000 whales, although there is uncertainty in this number (I.W.C. 1990b). Habitat Globicephala melas is a pelagic species which inhabits deep water throughout most of the year, although at times it moves inshore in pursuit of prey (Leatherwood and Dahlheim 1978; Sergeant 1979; Sergeant 1982; Dawson et al. 1985; Martin et al. 1987). Knowledge of habitat use by Globicephala melas is fragmentary for most seasons and for off- shore areas. Satellite tracking data indicate an affini- ty for shelf edges (B. Mate, Marine Sciences Center, Oregon State University, Newport Oregon; personal communication). General Biology Reproductive Capability The breeding season of Long-finned Pilot Whales around Newfoundland lasts from May to November, with the maximum number of births occurring in mid-August (Sergeant 1962). A summer breeding season is also found for pilot whales in the Mediterranean. The whales begin to congregate in July and calve by late September. In the Faroes, the average conception date is around June (Evans 1980; Amos and Dover 1990; Desportes 1990). Some ani- mals breed sucessfully year-round, however (Desportes 1990). A single calf, approximately 180 cm in length and weighing 100 kg is born after a gestation period of about 16 months (Frazer and Huggett 1959, 1973; Sergeant 1962; Martin et al. 1987; Bloch 1994). Twins appear rarely (Bloch 1994). Lactation can last 2.5 years or longer, although weaning and tooth eruption begin at around six months of age (Sergeant 1962; Martin et al. 1987; I.W.C. 1988; Desportes 1990; Bloch 1994). Some whales taken in the Faroes were found to have milk in their stomachs, even though they were over four years old (Desportes 1990). Females can be both lactating and pregnant (I.W.C. 1988; Bloch 1994). Male pilot whales show a rapid rate of growth until sexual maturity is reached at about 12 years of age, and four to five meters in length (Sergeant 1962; Martin et al. 1987; Kasuya et al. 1988b; Bloch 1994). Growth may be most rapid in the first two to three years of life (Martin et al. 1987; Kasuya et al. 1988b). After sexual maturity is reached, the growth THE CANADIAN FIELD-NATURALIST rate for males slows, and may cease at about 20 to 25 years of age (Sergeant 1962; Kasuya et al. 1988b; Bloch 1994). Females show a growth pattern similar to males, although they generally grow a bit slower (Sergeant 1962; Kasuya et al. 1988b; Desportes 1990; Bloch 1994). Females mature at six to 13 years and a length of three to four m (Sergeant 1962; I.W.C. 1988; Kasuya et al. 1988b; Bloch 1994), Males live up to 50 years, while females can live longer than 60 years (Kasuya et al. 1988b; Desportes 1990; Bloch 1994). One calf is produced about every three or four years (Sergeant 1962; Desportes 1990). At least three ovulations can occur in females during one breeding cycle (Sergeant 1962). Annual pregnancy rate is approximately 30%, (Perrin and Reilley 1984; Bloch 1994), and annual calf production is estimated at 10 to 13% (Sergeant 1962; Harrison 1969; Martin et al. 1987; Desportes 1990). A high reproductive capacity is retained at all ages, with some females surviving to a post-reproductive phase (Sergeant 1962; Martin et al. 1987; Kasuya et al. 1988b; Desportes 1990). This phase can actually begin quite early - one female examined in the Faroes was sexu- ally senile at 28 years — possibly as a result of ovar- ian exhaustion (Kasuya et al. 1988b; Desportes 1990). The annual mortality rate of males from age one to eight is about 5.8%, while for females it is 4.5% (Sergeant 1962). Kasuya et al. (1988b), examining the same data as (Sergeant 1962), estimated the mor- tality rate of males under 25 years of age to be about seven percent, while for females it is much lower at two percent. In young calves the sex ratio 1s approxi- mately equal, but the difference in mortality rates results in a declining percentage of males as cohorts age; the overall sex ratio at maturity being one male to two or three females (Sergeant 1962; Martin et al. 1987; Kasuya et al. 1988b; Desportes 1990; Bloch 1994). A higher percentage of females than males within a pod is reported from many sources (Sergeant et al. 1970; McLeod 1981; Crespo et al. 1985; Dawson et al. 1985; Martin et al. 1987; Amos and Dover 1990; Bloch 1994), although occasional- ly, mostly-male or all-male pods are found (Sergeant 1962; Geraci and St. Aubin 1977; Amos and Dover 1990; Desportes 1990). The higher rate of male mor- tality may result through intraspecific fighting among males (Bloch 1994). Species Movement Canadian Waters: Around Newfoundland, pilot whales show a marked seasonal variation in distribu- tion. They generally arrive on the Grand Banks in June and remain until late autumn (Sergeant 1962; Mercer 1975; Lien et al. 1980; Lynch 1987) Maximum abundance in summer appears to be along the southeast coast of Newfoundland: they are not as commonly seen in the inshore waters of the Vol. 110 | Fee, Rn een eee er 1996 NELSON AND LIEN: STATUS OF THE LONG-FINNED PILOT WHALE ILS) FIGURE 2. Range of the Long-finned Pilot Whale, Globicephala melas. Maritime provinces of Canada or southwards, although they are present (Mercer 1975; Leatherwood and Dahlheim 1978; Hain et al. 1981; Powers et al. 1982; Gaskin 1983). Summer inshore movements of pilot whales around Newfoundland are coincident with the arrival of the Short-finned Squid (Illex illecebrosus), which prefer water tem- peratures in the range of five to 15°C. (Frost and : Thompson 1933; Sergeant and Fisher 1957; Lien and Aldrich 1982). Pilot whales are often sighted in June, and again in October — November, along the continental slope from the entrance of the Gulf of Maine northward to _ the Laurentian Channel, the southwest edge of the ' Grand Bank, and the channel between the eastern edge of this bank and Flemish Cap (Sergeant 1962; Lynch 1987). They appear to be absent from this zone in winter (Sergeant 1962). There have been other widespread summer sight- ings offshore from New England, Nova Scotia, Iceland, in the Labrador Sea, and over oceanic depths ' east of the Grand Bank (Sergeant and Fisher 1957; Brown 1961; Mercer 1973, 1975; Boles 1980; Haycock and Mercer 1985; Sigurjonsson and Gunnlaugsson 1989). They may be found off the coast of Cape Cod as early as May (Starrett and _ Starrett 1955). Apparently, Long-finned Pilot Whales occur only north of 55°N latitude in the summer months, on both sides of the Atlantic (Brown 1961). Pilot whales migrate outside the Continental Shelf in winter, and are then known to inhabit areas on and east of the Grand Bank in North Atlantic Current waters (Sergeant and Fisher 1957; Sergeant 1962). There have also been winter sightings off New England and Greenland (Sergeant and Fisher 1957; Brown 1961; Mercer 1967, 1975; Parsons 1981). Other Areas: In West Greenland and Iceland, pilot whales generally stay offshore (Sergeant and Fisher 1957; I.W.C. 1990a). In European seas, this species occurs numerously between Iceland, the Norwegian coast and Great Britain in the summer, with its maximum concentration around the Faroe Islands (Sergeant and Fisher 1957; Leatherwood and Dahlheim 1978; I.W.C. 1990a). They are known to pursue squid into Icelandic fjords, and their presence in Faroese waters appears to be related to prey distri- bution and changes in water temperature (Grimpe 1933; Saemundsson 1939; Bloch et al. 1990a). There is little evidence to support any strong north-south migrational patterns in the Northern Hemisphere, although a seasonal movement of Southern Hemisphere whales into Antarctic waters has been postulated (Guiler et al. 1987; Martin et al. 1987). Behaviour Feeding: In Newfoundland coastal waters, Globicephala melas subsists mainly on Short-finned Squid, although Northern Cod (Gadus morhua) are common prey when squid are less plentiful (Sergeant 1962; Mercer 1975). Other organisms known to be taken in the western Atlantic include the amphipod Gammarus locusta and the squids Loligo pealii and Gonatus fabricii (Sergeant and Fisher 1957; Mercer 1967). When squid is abundant, it has been estimated that food intake may be 3 to 6% of body weight, or as much as 41 kg per day (Sergeant 1962). Although Illex illecebrosus is considered to be the main food item for Long-finned Pilot Whales in the western North Atlantic, individuals of consumable size are only available in the summer months (Mercer 1975). Prey species for the remainder of the year are not known, although oceanic squids (includ- ing other species of Ommastrephids) and oceanic fishes are probably candidates (Mercer 1975). Pilot whales have been known to consume Turbot, Reinhardtius hippoglossoides, in the winter (Mercer 1967). 516 Saemundsson (1939) considered Ommastrephes sagittatus to be the most commonly ingested squid in northern European waters, although pilot whales feed mainly on the squid Todarodes sagittatus off the Faroe Islands (Moore et al. 1979; Desportes 1990; Bloch 1994). Generally, prey species taken in Faroese waters are shoaling, oceanic, mid-water organisms (Desportes 1990). Other squids taken in eastern waters include Gonatus fabricii, Eledone sp., Teuthowenia sp., Taonius sp., Architeuthes sp., and Histioteuthis sp. (Evans 1980; Desportes 1990; Bloch 1994). Fish prey include Blue Whiting (Micromesistius poutassou), Northern Cod, Horse Mackerel (Caranx trachurus), flounder (Pleuro- nectidae), and Turbot (Sergeant 1962; Evans 1980; Bloch 1994). Fish and other organisms become more prevalent in the diet of pilot whales when squid are scarce (Desportes 1990; Bloch 1994). Schools of pilot whales are known to associate strongly with mackerel shoals off southwestern England (Evans 1980). There is also a correlation between herring numbers and pilot whale catches for the Faroes, which may result from squid con- centrating to feed on the herring (Evans 1980). Pilot whales apparently feed in a group, as the degree of digestion of squid in stomachs is always the same in groups driven ashore (Sergeant 1962). Globicephala melas has been observed feeding in groups and forming circle patterns which may indi- cate “herding” of squid (Weilgart 1985). Social Behaviour: Pelagic groups. of Globicephala melas generally consist of about 20 animals, although they may concentrate inshore in much larger numbers sometimes exceeding 200 animals (Sergeant and Fisher 1957; Clarke 1962; Sergeant 1962; Evans 1980; Hay 1982; Weilgart 1985; Amos and Dover 1990). In Newfoundland, groups that are driven ashore or mass strand are generally larger than pelagic assemblages; the mean being 85 animals (Sergeant 1962, 1982). It is spec- ulated that changes in group size may reflect vary- ing behaviours such as feeding, migration, or repro- duction, although pod sizes may also be underesti- mated when observed at sea (Sergeant and Fisher 1957; Sergeant 1962; Evans 1980; Weilgart 1985; Martin et al. 1987). Schools generally contain animals of various sizes and both sexes (Sergeant 1962; Amos and Dover 1990; Bloch 1994). The social structure of Globicephala melas has yet to be conclusively determined, although there is evidence of a matri- lineal pod organization (Amos and Dover 1990; I.W.C. 1990a; Desportes 1990). Further, because there is usually more than one mature male in a pod, a multi-male, polygynous mating system seems likely (Sergeant 1962; Martin et al. 1987). Males probably move frequently between pods; spending only a few months in any particular group THE CANADIAN FIELD-NATURALIST Vol. 110 (Amos and Dover 1990; Desportes 1990). This movement may begin when the males reach sexual maturity, during which time segregation into sepa- rate pods is possible (Kasuya et al. 1988b). All-male or mostly-male herds are rarely observed however, and there is no evidence that this species is generally segregated (Sergeant 1962; Martin et al. 1987; Amos and Dover 1990; Bloch 1994). Globicephala melas is a highly social species, and strong social bonds are often cited to explain mass stranding (Kritzler 1952; Geraci and St. Aubin 1977; Norris and Dohl 1980). Its social nature is also reflect- ed in a rich vocal repertoire, which includes a variety of whistles ranging from 0.5 to 5.0 Khz in frequency, double clicks, and the ability to produce two totally different signals simultaneously (Schevill 1964; Busnel and Dziedzic 1966; Taruski 1979; Herman and Tavolga 1980). Signature whistles and dialects may also exist (Taruski 1979; McLeod 1982). Sound production is known to vary with behavioural and environmental context (Taruski 1979; Weilgart and Whitehead 1990). Simplest sounds are emitted during periods of minimal activity such as resting behaviour, while more complex sounds occur when behaviour is vigorous and energetic, and involves more complex coordination within the group (Taruski 1979; Weilgart and Whitehead 1990). Differences in calling rate between large and small schools have also been observed (Taruski 1979). In the summer of 1987, an immature pilot whale was tracked for 95 days in the western North Atlantic through the use of a satellite-monitored radio tag (Mate, personal communication). Information obtained indicated that virtually all deep dives occurred either just before sunset or at night, and coincided with the nocturnal rise of prey items. Since few deep dives were recorded during the day, it was thought that the whale was probably feeding on surface shoaling fish at these times, if it fed at all. The highest swimming speeds were also found at night, suggesting fast prey-chasing or searching. Daily movements of up to 234 km were observed, with a mean of 80 km per day. The average number of dives in a 12 hour period varied from 636 to 1433, reflecting changes in the animal’s activity patterns. Swimming speeds averaged 3.3 km/h over the entire | period, while speeds above 16 km/h could be main- tained for periods exceeding three hours. Surface resting activity, sometimes of up to 15 minutes, was _ most common during the first three hours of sunrise | (Mate, personal communication). This species has been observed to form mixed — groups with Bottlenose Dolphins (Tursiops | truncatus), White-beaked Dolphins (Lagenor- hynchus albirostris), Atlantic white-sided Dolphins (Lagenorhynchus acutus), Killer Whales (Orcinus orca), and Tuna, Thunnus thynnus (Sergeant and Fisher 1957; Aloncle 1972; Leatherwood et al. 1976; Leatherwood and Dahlheim 1978; Evans 1980; Sears 1996 1982; Bloch 1994). On occasion, single stray Narwhals, Monodon monoceros, have been found associating with groups of Globicephala melas (Lien and Barney 1991). Behaviours such as spyhopping and lobtailing have been observed in pilot whales, while breaching may occur when groups are feeding (Weilgart 1985; Leatherwood et al. 1976). They are not known to ride bow waves (Leatherwood et al. 1976). Limiting Factors Parasites Although not a serious limiting factor, parasites are common in this species. External parasites include the Whale Louse, /socyamus delphini, which infests crevices in the skin and old wounds, and Conchoderma auritum, found on the teeth and gums, usually of older animals (Sergeant 1962; Martin et al. 1987; Balbuena and Raga 1991). The Whale Louse apparently infests mature males more heavily than other classes, possibly due to differences in behaviour of the males (Balbuena and Raga 1991). Internal parasites include the nematode Anisakis simplex which may originate in the stomach from ingested squid, in the nasal passages and middle ear sinuses, and Crassicauda carbonelli infesting the penis; also the trematodes Leucasiella delamurei and Odhneriella subtila from the small intestine, and Phyllobothium delphini embedded in the blubber of older animals (Sergeant 1962; Raga and Balbuena 1988, 1990; Balbuena et al. 1989). Some parasites, such as the trematode Pholeter gastrophilus and the nematode Anasakis simplex show a decline in older animals, which could suggest that heavy infestation could be a cause of mortality (Desportes 1990). Infections The influenza A virus, originating from avian sources, is known to be transferrable to pilot whales (Hinshaw et al. 1986; Chambers et al. 1989). Streptococcus equi, an equine pathogen rarely isolat- ed from other animal species, has been found in Globicephala melas (Higgins et al. 1980). Pollutants Organochlorines and heavy metals have been doc- umented in the tissues of Globicephala melas (Taruski 1975; Wagemann and Muir 1984; Muir et al. 1988). Martin et al. (1987) reported higher levels of PCBs in pilot whales stranded along British coasts than have been found in any other populations of this species. Mercury levels in consumable parts of Faroe pilot whales are known to approach or exceed safe limits (Andersen et al. 1987). Contaminants may be ' transfered from females to their young during preg- nancy and lactation (Desportes 1990). The effects of these pollutants have yet to be determined. However, high concentrations of heavy metals have been suggested as a possible cause for NELSON AND LIEN: STATUS OF THE LONG-FINNED PILOT WHALE ili stranding, and high levels of polychlorinated biphenyls have been associated with reproductive failure in other marine mammals (Muir et al. 1988). Predation Pilot whales often bear scars, especially on trailing edges of flippers, dorsal fin, and flukes (Sergeant 1962; Martin et al. 1987). Tooth scars, common on adult males, may attest to play or fighting for pod dominance rather than by attempted predation (Martin et al. 1987; Bloch 1994). Sergeant (1962) found no evidence of predation by Killer Whales or sharks for pilot whales around Newfoundland, although Killer Whales are known to prey on them in other areas (Katona et al. 1988). Sucker marks have also been observed, and are probably made while the whale is feeding on squid (Sergeant 1962; Martin et al. 1987; Bloch 1994). Mass Strandings Globicephala is the genus that most frequently mass strands. Such strandings sometimes involve entire pods (Geraci and St. Aubin 1977; Evans 1980; Sergeant 1982). Mass strandings of Long-finned Pilot Whales have been recorded from Ireland, Britain, France, Spain, New Zealand, South America, New South Wales, the Netherlands, Greenland, Newfoundland, Nova Scotia, Cape Breton Island, Sable Island, Prince Edward Island, Magdalen Islands, Miquelon Island, and Cape Cod (Starrett and Starrett 1955; Dawbin 1964; Sergeant et al. 1970; O’Riordan 1975; Husson and Van Bree 1976; Mead 1979; Sergeant 1979; Wood 1979; Nores and Perez 1982; Sergeant 1982; Crespo et al. 1985; Dawson et al. 1985; Sheldrick 1989; Kingsley, unpublished data). Table 1 lists reported mass strandings for eastern Canadian waters between 1957 and 1990. A rough estimate of annual mortality due to mass strandings in Newfoundland from 1975 to 1980 is about one percent of the population, and mass strandings seem to be increasing (Sergeant 1982). Recently, pilot whales have begun to strand almost yearly around Cape Cod (Sergeant, personal commu- nication). Sheldrick (1979) and Brown (1975) report an increase of pilot whale strandings on the British coast beginning in 1947, although their data included single-stranded animals. A more recent study sug- gests that British mass strandings have increased since about 1982 (Sheldrick 1989). Further informa- tion is needed to estimate the effect of mass strand- ings on pilot whale populations. It is possible that mass strandings represent the main source of coastal mortality for this species (Sergeant 1982). Incidental Catches Incidental catches of Globicephala melas in fish- ing gear have been frequently observed (Lien and Aldrich 1982; I.W.C. 1990a; Donovan and Perrin 1990; Kraus et al. 1990; Northridge 1990; Stenson 518 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Mass strandings of Globicephala melas for eastern Canadian Waters between 1957 and 1990. All reported strandings of more than one animal are listed (Kingsley, unpublished data). Year Date No. Locality 1957 4 August 12+ Cow Head, N 1959 2 October 18 Sable Island, N.S. 1960 1 August 58 Port Maitland, N.S. 1967 24 December 15 Glace Bay, Cape Breton Island, N.S. 1975 28 September 300 Charleston, Bonavista Bay, NF 1976 26 December 130+ Sable Island, N.S. 1978 17 September 63 Miquelon Island 1978 19 September 99 Pte aux Allouettes, Miquelon Island 1978 29 September ca.70 Musgrave Harbour, Bonavista Bay, NF 1979 14 July 135 Point aux Gaul, NF 1980 18-19 October 75 Point Leamington, NF 1980 25 October 18 Grand Beach, NF 1980 9-18 November 52 Bedeque Bay, PEI 1981 31 August 39 Little Burnt Bay, Notre Dame Bay, NF 198] 4 September 2 Chance Cove Trinity Bay, NF 1981 4 September 5 Branch St. Mary’s Bay, NF 1981 6 Sepember 27 George’s Bay, NS 1981 8 September 70 Port Hood, George’s Bay, N.S. 1981 13 October 13 Magdalen Islands 1981 27 July 23 Grand Bank, NF 1982 18 August 3 Bonavista, NF 1982 13 August 14 Pinkney’s Pt., Yarmouth N.S. 1990 9 August 2 Cheticamp, N.S. 1990 15 September 2 Sturgeon Bay, PEI 1990 ca. 30 September 2 S. Lakevale, N.S. Remarks Source Fall died Sergeant et al. 1970 refloated on later tide Sergeant et al. 1970 all died Sergeant et al. 1970 all died Sergeant et al. 1970 200 whales towed out, Mitchell 1977 many smaller whales restranded, approx 125 died 116 examined scattered over | mile of sandy beach dead in two tight groups Geraci and St. Aubin 1977 Mitchell 1980 Mitchell 1980 fishermen towed many Mitchell 1980 into deep water, approx. 54 died all died Lien 1980 all died Lien 1980 all died Lien 1980 all died Mitchell 1982 all died Lien and Aldrich 1982 both died Lien and Aldrich 1982 all died Lien and Aldrich 1982 13 died, remainder Loch 1983 released 25-30 died Loch 1983 all died Loch 1983 12 unbeached themselves, __ Lien et al. 1982 11 died 2 unbeached, Lien et al. 1982 1 died 12 released, 2 died Goodman 1984 ca. 40 whales attempted to strand but were successfully repelled, 2 died female and young moved to deeper water, female restranded on 16th, euthanized males, dead when found Tom Kiely, in litt., 10 Aug 1990 R.P. Johnston, DFO Charlottown, in litt., 19 Sep 1990 Kingsley, personal communication and Reddin 1990). Catches often involve younger and smaller animals which cause little damage to the fishing gear. Because of this, there is a tendency to under-report catches (Lien and Aldrich 1982; Lien et al. 1994). Further, in some jurisdictions where there are regulatory problems stemming from incidental catches, fishermen may try to avoid reporting animals that have been caught (Prescott et al. 1980). Lien and Aldrich (1982) investigated incidental catches of Globicephala melas in Newfoundland during 1983, a year of high squid abundance. The whales were commonly found in traps set for squid, although they also became caught in groundfish and herring gillnets. Of 43 animals reported entrapped, 87% died as a result of the entrapment. It was sug- gested that entrapments were probably more likely to be reported by fishermen in 1983 because they 1996 occurred in a small area and were considered a major problem in the fishery. Since 1983, inshore abun- dance of Globicephala melas has been quite low, and it is probable that isolated entrapments would not be reported (Lynch 1987; Lien et al. 1994). Other The exploitation of prey species by various fish- eries may have an impact on populations of Globicephala melas, although such effects have yet to be documented for this species (Lowry and Frost 1985). Special Significance of the Species The Faroese Fishery The non-commercial pilot whale hunt, or “grind” is an integral part of the traditions and culture of the Faroe Islands (Olafsson 1990). In recent years a high level of controversy has surfaced which involves ethical, ecological, and animal-welfare issues (Gibson-Lonsdale 1990; Joensen 1990; Olafsson 1990). In addition, high concentrations of mercury found to be present in consumable tissues has raised significant health questions (Andersen et al. 1987; Olafsson 1990). The pilot whale fishery of the Faroes does offer an excellent opportunity for studying the biology and social structure of these animals, as statistics have been kept as far back as 1584, and represent infor- mation about entire pods of whales (Desportes 1990; Bloch 1994). A newly-developed technique which may prove invaluable for answering questions about population biology is that of DNA fingerprinting, which can accurately assess paternity of individual animals (Amos and Dover 1990). DNA fingerprint- ing is a more accurate method for examining popula- tion structure than the use of coded tags or naturally occurring markings, because these offer only indirect evidence about breeding systems (Kaufman et al. 1987; Amos and Dover 1990). At the present time, data are being gathered with this technique from the Faroese fishery which will eventually provide infor- mation concerning individual movement between _ pods, age at first breeding, length of time spent with- in pods, and the relatedness of individuals within pods (Amos and Dover 1990). The Phenomenon of Mass Stranding As Globicephala is the genus that most frequently | mass strands, Globicephala melas is a common sub- _ ject for the study of this phenomenon (Geraci and St. _ Aubin 1977; Evans 1980; Sergeant 1982; Kirschvink 1990). No conclusive evidence yet exists as to the causes of mass strandings. Parasitism has been sug- _ gested as a possible factor (Dailey et al. 1979; Wood 1979; Morimitsu et al. 1986), yet it appears that _ mass-stranded animals are generally not diseased or | highly parasitized (Geraci 1979; Hall and Schimpff 1979; Ridgway 1979; Sergeant 1979; Wood 1979; NELSON AND LIEN: STATUS OF THE LONG-FINNED PILOT WHALE Sg Sergeant 1982; Morimitsu et al. 1986). Other specu- lations include confusion due to lack of familiarity with inshore waters, disturbances to the echolocation system because of an area’s underwater topography, problems in attention, anomalies in geomagnetism, or general stress from any number of possible sources (Geraci and St. Aubin 1979; Wood 1979; Dawson et al. 1985; Kirschvink 1990). As mass strandings of Odontocete species appar- ently occur where the animals are abundant, a better understanding of the phenomenon could provide evi- dence on the state of various populations (Sergeant 1979; Sergeant 1982). Satellite Tracking The development of satellite-monitored radio tracking represents another important breakthrough in the study of whale behaviour. A feasible monitor- ing system was first used in 1987 to record the behaviour of a pilot whale for 95 days in the western North Atlantic (Mate, personal communication). The pilot whale is a useful model for tracking studies because it is found in a variety of environments, its trophic niche is typical of many odontocetes, and it is easily captured (Mate, unpublished data). At the present time, satellite systems are still under devel- opment; they may eventually provide information including stock separation, energetics, acoustics, and the identification of critical habitats (Mate 1989). Evaluation Globicephala melas is widespread, and although numbers are considerably lower than prior to com- mercial whaling, they are still relatively abundant in the North Atlantic. Although there may be several sub-populations in Canadian waters which are somewhat distinct, exact relationships are unclear. It is known that Globicephala melas off Newfound- land was severely depleted by drive fisheries which ended in 1971. There have not been systematic efforts to assess populations and their recovery since that time. Even optimistic estimates would indicate that numbers are still low and well below pre-whaling population size. Mass strandings repre- sent a major known source of natural mortality for this species. The effects of incidental entrapments, pollutants, and fisheries for prey species remain rel- atively unknown, but these factors have the poten- tial for limiting this species. Although this sub-pop- ulation would meet the criteria for a COSEWIC des- ignation of vulnerable, if considered without regard to other North Atlantic populations, there is no evi- dence that the population is distinct and discrete. 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U.S. Department of Commerce, National Technical Information Service, Number PB 293890. Yonekura, M., S. Matsui, and T. Kasuya. 1980. On the external characteristics of Globicephala macrorhynchus off Taiji, Pacific coast of Japan. Scientific Reports of the Whales Research Institute 32: 67-95. Accepted 13 March 1996 The Status of the Pygmy Sperm Whale, Kogia breviceps, in Canada* ROBIN W. BAIRD!?, DAWN NELSON?, JON LIEN* and DAVID W. NAGORSEN? "Marine Mammal Research Group, Box 6244, Victoria, British Columbia V8P 5L5 *Biology Department, Dalhousie University, Halifax, Nova Scotia B3H 4J1 3Whale Research Group, 230 Mt. Scio Road, Memorial University of Newfoundland, St. John’s, Newfoundland AlC 5S7 Current address: 16 Maples Road, Raymond, Maine 04071 4Ocean Sciences Centre and Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland A1C 5S7 Royal British Columbia Museum, 675 Belleville Street, Victoria, British Columbia V8V 1X4 Robin W. Baird, Dawn Nelson, Jon Lien, and David W. Nagorsen. 1996. The status of the Pygmy Sperm Whale, Kogia breviceps, in Canada. Canadian Field-Naturalist 110(3): 525-532. The general biology, world-wide status and management of the Pygmy Sperm Whale, Kogia breviceps, with special refer- ence to its status in Canadian waters, is reviewed. Pygmy Sperm Whales appear to be uncommon in Canadian waters; there are several unconfirmed sighting records off British Columbia and only four stranding records off the Canadian east coast. Little is known about its biology or world-wide status, and although it is taken in small numbers both directly and inciden- tally in fisheries, no serious threats to its status are apparent. No COSEWIC designation is required regarding its status in Canadian waters. Le présent rapport résume la biologie générale, le statut international et la gestion du Cachalot Pygmée, Kogia breviceps, et s’attache plus particuliérement a son statut dans les eaux canadiennes. Le Cachalot Pygmée semble étre une espéce inhab- ituelle dans les eaux canadiennes; il existe plusieurs mentions visuelles de cachalots au large de la Colombie-Britannique, mais aucune n’a été confirmée. En outre, on ne dispose que de quatre mentions de specimens échoués le long de la céte est du Canada. On ne connait que peu de détails au sujet de sa biologie ou de son statut international. Bien que le Cachalot Pygmée soit capturé en petit nombre, que ce soit directement ou comme prise fortuite par les echeurs, il ne semble y avoir aucune menace grave a son statut. Son statut dans les eaux canadiennes ne nécessite donc aucune désignation par le CSEMDC. Key Words: Pygmy Sperm Whale, Cachalot Pygmée, Kogia breviceps, Canada, North Atlantic, British Columbia, status, Cetacea. Little is known about the Pygmy Sperm Whale, Kogia breviceps (de Blainville, 1838) [Figure 1]. We summarize here the current state of knowledge of the species, with special reference to its status and man- agement in Canadian waters. There are two species within the genus Kogia: Kogia breviceps, and the gated and laterally compressed, and the flukes are notched and concave along the rear margin. Stranded specimens of both species are occasionally confused with sharks, due to the shark-like mouth and the presence of a lightly pigmented bracket-shaped mark on each side of the head between the eye and flipper, _ Dwarf Sperm Whale, Kogia simus. Before 1966, however, most authors recognized one species, _ Kogia breviceps, within the genus (Yamada 1954; sometimes referred to as “false gills” (Figure 1). The two species can be discriminated by a variety of external and cranial characters (see Table 1). ————————— Se Handley 1966), resulting in considerable confusion as to which species is actually referred to in early publications. This taxonomic uncertainty can be attributed to the similarity of the two species and the _ scarcity of specimens. Both taxa are small (less than _ 3.8 m), have a small, underslung shark-like mouth which is set well back from the tip of the snout. They | are dark bluish-gray dorsally and shade from a lighter gray laterally to a dull white or pink on the belly (Leatherwood and Reeves 1983; Nagorsen _ 1985). Both have a bulbous head and, like the other member of the Family Physeteridae, the Sperm _ Whale (Physeter macrocephalus), they have a sper- maceti organ (Handley 1966). The tail stock is elon- Many authors note that they can be distinguished by the position of the dorsal fin on the back, set well posterior to the midpoint of the back for the Pygmy Sperm Whale and with the anterior insertion of the fin near the midpoint of the back for the Dwarf Sperm Whale, but Ross (1979) cautioned against using this character in isolation, as there is consider- able individual variation in both species. Pygmy Sperm Whales reach a greater length than Dwarf Sperm Whales, but the maximum length attained by Kogia breviceps is unclear. Caldwell et al. (1971) reported an individual of 4.25 m, but Ross (1979) suggested that the length may have been estimated, rather than measured. According to Ross (1979) the *Reviewed and Approved by COSEWIC 14 April 1994, report accepted no status designation required. 525 526 next-largest recorded specimen was 3.5 m in length. Leatherwood et al. (1988) noted that Pygmy Sperm Whales may grow to a maximum length of at least 3.7 m, and Eliason and Houck (1986) reported an animal 3.82 m in length from the records of the Smithsonian Institution. The taxonomic position of the genus Kogia within the Odontoceti is unclear (e.g., see Rice and Wolman 1990). Distribution The Pygmy Sperm Whale is found virtually world-wide in tropical and warm-temperate seas. In the western Pacific it has been reported from Japan in the north to New Zealand and Australia in the south (Omura and Takahashi 1981; Baker 1983; Brabyn 1991), and in the eastern Pacific from Washington State in the north to Peru and Chile in the south (Allen 1941; Scheffer and Slipp 1948; Hubbs 1951; Brownell 1969; Waerebeek et al. 1987). Pike and Giovando (1963) stated that Pygmy Sperm Whales were known to occur in the offshore waters of British Columbia, but we could find no documented records from the Canadian west coast. Four unconfirmed sighting reports are listed as Kogia breviceps from the British Columbia coast (Baird unpublished data), but positive confirmation awaits photographs or a specimen. Three records are confirmed from the adjacent waters of Washington State (Scheffer and Slipp 1948; Everitt et al. 1979; Osborne et al. 1988) and it is likely that this species will eventually be documented in B.C. Kogia simus has been recorded in British Columbia (Nagorsen and Stewart 1983). In the western Atlantic, Pygmy Sperm Whales have been recorded from Canada southward along the United States east coast, throughout the Gulf of Mexico, and as far.south as Brazil, Uruguay and Argentina (Piers 1923; Allen 1941; Gunter et al. 1955; Carvalho 1967; Hysmith et al. 1976; Geise and Borobia 1987). Four stranding records have been reported on the Canadian east coast (Figure 2; Table 2). Three of these were from Canadian waters, and one from the French Island of Miquelon (Piers 1923; Sergeant et al. 1970; Nelson et al. 1991). These are the most northern records in the western Atlantic. In the eastern Atlantic this species has been recorded THE CANADIAN FIELD-NATURALIST Vol. 110 from Ireland, the Netherlands and France south to South Africa (Allen 1941; Fraser 1974; Maul and Sergeant 1977; Teixeira 1979; Ross 1984). Pygmy Sperm Whales are found in the Indian Ocean (Leatherwood and Reeves 1989; Chantrapornsyl et al. 1991) but do not appear to have been recorded: in the Mediterranean Sea (Baccetti et al. 1991). Protection International The Pygmy Sperm Whale is listed under Appendix II of the Convention on International Trade in Endangered Species of Wild Fauna and Flora 1973 (CITES) (see Birnie 1982). Such a listing allows for the regulation of international trade between members and non-members of the conven- tion by requiring export permits from the country of origin. However, there appears to be no current inter- national trade in Pygmy Sperm Whale products. The International Whaling Commission (IWC) regulates the taking of whales in accordance with the current Schedule provisions, but whether this Commission’s mandate covers the Pygmy Sperm Whale is unclear, as members of the Commission are divided as to whether “whale” refers to all cetaceans, or to only some species (Klinowska 1987, 1991). National Canada: Until they were repealed in 1993, the Cetacean Protection Regulations of the Fisheries Act of Canada of 1867 protected all cetacean species from “hunting”. “Hunting” was defined as “to chase, shoot at, harpoon, take, kill, attempt to take or kill, or to harass cetaceans in any manner’, and could only be undertaken under licence. Aboriginal “hunting” however could be undertaken without licences. The Cetacean Protection Regulations were replaced with the Marine Mammal Regulations of the Fisheries Act in early 1993. These regulations appear to provide no more or no less protection, by stating only that “no person should disturb a marine mammal except when ... under the authority of these Regulations”. No pro- visions exist for regulation of incidental catches in fishing operations. Canada is not currently a member of the International Whaling Commission, having withdrawn in 1982 (IWC 1982). TABLE |. Distinguishable characteristics of Kogia breviceps and Kogia simus (Ross 1979; Leatherwood et al. 1988). Characters Kogia breviceps Maximum length! 3.8m Throat creases absent Dorsal fin height Dorsal fin position! Relative snout length longer Teeth in upper jaw absent Teeth in lower jaw 10-16 pairs 'See text for further details. usually less than 5% total body length posterior to mid point of back Kogia simus 2.7m usually present usually greater than 5% total body length near midpoint of back shorter up to 3 pairs 8-13 pairs _ Date 1996 BAIRD, NELSON, LIEN, AND NAGORSEN: STATUS OF THE PYGMY SPERM WHALE 527 FiGuRE 1. Pygmy Sperm Whale. Illustration by Dawn Nelson. United States: All cetaceans are protected through the Marine Mammal Protection Act of 1972, as well as through the Packwood-Magnuson Amendment of the Fisheries and Conservation Act and the Pelly Amendment of the Fisherman’s Protective Act. Population Size(s) and Trends In the recent IUCN Cetacean Red Data Book, Klinowska (1991) noted that there is insufficient information to accurately classify the world status of this species. No information is available on popula- tion sizes or trends, nor on stock identity. In fact, very few sightings of either species of Kogia have ever been reported, and knowledge of both species comes largely from stranded animals. In some areas, Pygmy Sperm Whales are among the most frequent- ly recorded stranded species. For example, Odell (1991) noted that they are the second most frequent to strand in the southeastern United States; off the Hawaiian Islands the Pygmy Sperm Whale is the fourth most frequently stranded species (Nitta 1991). In his analysis of strandings in New Zealand, Brabyn (1991) noted that although they do not represent the largest number of individuals (as. mass strandings are infrequent), Pygmy Sperm Whales are the most fre- quently recorded species. Many authors have inter- preted these frequent strandings as evidence that this species is fairly common. Recent numerous aerial sightings of Kogia (not discriminated to species) in the northern Gulf of Mexico seem to support this supposition (Jefferson et al. 1992). Only four strandings have been reported off the Canadian east coast and it appears that Pygmy Sperm Whales become more common in the south, with a greater number of records reported off the northeast U.S. coast (Early and McKenzie 1991). Habitat Pygmy Sperm Whales generally inhabit offshore waters in warm temperate and tropical areas. Brabyn (1991) suggested that the Mahia Peninsula area of the north island of New Zealand is a calving area for this species, based on a high proportion of mother/calf strandings. However, information on the habitat type is not presented. Klages et al. (1989) noted that both species in the genus feed on prey typ- ical of the continental slope, although Kogia simus feeds more inshore than Kogia breviceps. General Biology Reproduction Very little is known about the reproductive biolo- gy of this species. Most estimates of reproductive parameters are based on the examination of a small number of stranded animals, and should be consid- ered preliminary. Ross (1979) suggested that mating and calving occur from autumn through spring, and noted that while there was insufficient data to accu- rately estimate gestation, two alternative methods suggest a duration between seven and 11 months. Individuals have been recorded as being simultane- ously pregnant and accompanied by a calf, indicating that some females may breed annually (Ross 1979; Price et al. 1984; Eliason and Houck 1986). Ross (1979) reported that length at birth averages about 1.2 m, sexual maturity is attained at about 2.7 to 2.8 TABLE 2. Records of Kogia breviceps from the Canadian east coast. All records are of single animals, found dead. No con- firmed records from the Canadian west coast have been reported. Location Source Piers 1923! Halifax Harbour, Nova Scotia 17 January 1920 + 29 January 1969 } 8 December 1992 8 October 1990 Isle de Miquelon Sable Island, Nova Scotia Saint John, New Brunswick Sergeant et al. 1970 Nelson et al. 1991 McAlpine and Murison, in preparation ‘Katona et al. (1983) report a dead animal found under the ice in Halifax Harbour, Nova Scotia, in the winter of 1970, but _ this appears to be a misprint, referring to the animal found under similar circumstances in 1920. 528 m for females and 2.7 to 3.0 m for males, and physi- cal maturity is reached at lengths of 3.0 to 3.3 m for both sexes. The sex ratio of stranded animals varies. Brabyn (1991) found that of 79 Pygmy Sperm Whales that stranded in New Zealand, there were more than twice as many females as males. Off South Africa, the sex ratio of 16 adult and sub-adult animals was 1:1, while the sex ratio for 15 foetuses and calves was heavily biased towards males (12 males: three females) [Ross 1979]. Such disparate figures likely arise from the small sample sizes, and clearly more information is needed. Ross (1979, 1984) and Eliason and Houck (1986) discussed sectioning teeth for age determination, but no information is available to calibrate the deposi- tion rate of layers. Ross (1984) noted that one sexu- ally mature female which stranded with a calf had only 3.5 growth layers in the dentine. Assuming that one layer is deposited annually, this would imply the female was only about two years old when she first conceived (Ross 1984). No information on longevity is available. Movements Strandings off South Africa and the U.S. southeast coast occur throughout the year, possibly suggest- ing a lack of seasonal movements (Ross 1979; Leatherwood and Reeves 1983). Off South Australia however, Kemper and Ling’s (1991) analysis of stranding records indicates the presence of Pygmy Sperm Whales only during April through October. In the eastern North Pacific the majority of records are concentrated during the fall and winter (Eliason and Houck 1986), and along the coast of Europe, Fraser (1974) suggested that this species follows the North Atlantic current extension of the Gulf Stream in search of food. There is also evidence for a possible seasonal movement of Kogia breviceps near the west coast of New Caledonia, as strandings there occur mainly between June and December (Sylvestre 1988). Behaviour Behavioural observations are based on a few sightings at sea and on instances where individuals have been kept in captivity after stranding alive (e.g., see Sylvestre 1983). Pygmy Sperm Whales are seen singly or in groups of up to about six individuals (Allen 1941; Vidal et al. 1987; Leatherwood et al. 1988), and do not appear to regularly associate with other species of cetaceans or with seabirds (Au and Pitman 1988). Surface behaviour is typically undemonstrative; individuals typically rise slowly to the surface, produce an inconspicuous blow, and dive without showing the flukes. They are occasion- ally observed lying still at the water’s surface with the top of the head and back exposed. Allen (1941) noted that animals at the surface appeared to be very easy to approach closely, even after being har- THE CANADIAN FIELD-NATURALIST Vol. 110 pooned. However, as with several other species of cetaceans, Pygmy Sperm Whales may release a cloud of reddish brown feces into the water when startled (Leatherwood et al. 1988). The function of this behaviour is unclear, although Scott and Cordaro (1987) observed a Dwarf Sperm Whale mother and calf pair exhibiting this behaviour while trapped inside a tuna purse-seine net, and then apparently hiding within the cloud when approached by dol- phins which were also trapped. Pygmy Sperm Whales appear to feed primarily on cephalopods, as well as crustaceans and fish. Cephalopods recorded from stomach contents of this species include Abralia sp., Abraliopsis sp., Ancistrocheirus sp., Galiteuthis sp., Histioteuthis sp., Loligo vulgaris, Lycoteuthis diadema, Moroteuthis sp., Octopoteuthis cyeletron, Ommastrephes sp., Onychoteuthis boreali-japonicus, Phasmatopsis sp., Pygropsis sp., Pyroteuthis sp., Sepioteuthis australis, Taningia sp., Taonius pavo, Todarodes sp., Tuethowenia pellucida, and Vampyroteuthis sp. (Hale 1947; Raun et al. 1970; Ross 1979; Eliason and Houck 1986; Klages et al. 1989). Crustaceans recovered include Aristaeomorpha foliacea, Carcinides maenas, Gnathophausia ingens, Goneplax angulata, Hymenodora sp., Pandalopsis sp., Pandalus sp., Pasiphaea pacifica, and Penaeus californiensis (Allen 1941; Hale 1947; Scheffer and Slipp 1948; Raun et al. 1970; Ross 1979; Vidal et al. 1987). Fish recorded include Lampanyctus sp., Maurolicus muelleri, Photichthys argenteus, Pyrosoma sp., Rexea solandri, Scopelopsis multi- punctatus, and Symbolophorus sp. (Ross 1979, 1984). Raun et al. (1970) also noted the seaweed - Sargassum from the stomach of a stranded animal, but suggest that this may have been ingested acci- dentally. An isotope analysis of muscle tissue from a stranded animal off Miquelon suggested that the ani- mal had been feeding at the same trophic level as Sperm Whales; i.e., it had probably been feeding on squid in offshore waters (Nelson et al. 1991). This species likely uses echolocation to find prey, as echolocation-type clicks have been recorded from live-stranded animals (Caldwell and Caldwell 1991). Limiting Factors Data on natural mortality are scarce. There are no reported observations of Killer Whales (Orcinus orca) attacking Pygmy Sperm Whales (Jefferson et al. 1991), but the species has been recorded from the stomach contents of Killer Whales from both the Caribbean and the Indian Ocean (Perrin 1982). Long (1991) noted an apparent attack by a White Shark (Carcharodon carcharias) on a Pygmy Sperm Whale off California. Information on other causes of natural mortality is sparse. Virtually all recorded strandings of this species are of single animals or of cow/calf pairs. In New Zealand, Robson (1984) 1996 BAIRD, NELSON, LIEN, AND NAGORSEN: STATUS OF THE PYGMY SPERM WHALE 929 New Brunswick Nova Scotia (0) 250 km Jk ay SR 92 Py SIV A we Oey aps ro * Sais Aa x £ a ¥ es Atlantic Ocean i) FIGURE 2. Records of Kogia breviceps off the Canadian east coast. See Table 2 for details of records. reported one case of three dead individuals found together on a beach, in association with seven stranded False Killer Whales (Pseudorca crassi- dens), and Brabyn (1991) described a stranding of four individuals. Robson (1984) attributed many of the single strandings of this species to heavy infesta- tions of parasites in the head and inner ear. Parasites recorded from this species include the cestode Phyllobothrium delphini, and the nematodes Anisakis physeteris, Anisakis typica, Anisakis sim- plex, Phocanema kogiae, Stenurus sp., Terranova sp., and possibly Crassicauda sp. (Zam et al. 1971; Ross 1979; Vidal et al. 1987). Buck (1984) noted the presence of the bacteria Enterobacter agglomerans, Enterobacter cloacae, Pseudomonas cepacia, Pseudolmonas maltophila, Bacillus, Flavobacterium, and the yeasts Rhodotorula pallida, Rhodotorula rubra, Torulopsis, and Aureobasidium, but the role of such pathogens in natural mortality is unknown. Severe vascular disease, including thrombus forma- tion, has been noted in one animal (Roberts et al. 1964, cited in Sweeney and Ridgway 1975). There appear to be no known major threats (past or present) to this species (Klinowska 1991). Small numbers of Pygmy Sperm Whales have been taken in fisheries, both directly and incidentally. In previ- ous years, Kogia have been taken by shore-based whaling operations off Japan, but it is unclear if both species were included (Yamada 1954). Edmondson (1948) noted one animal which was speared, and another accidentally hooked and landed while fish- ing with a hand line off Hawaii. Van Waerebeek et al. (1987) found a specimen in a dump in Peru where the remains of fish and dolphins taken deliberately for human consumption were discarded. One animal was illegally killed in Australia in 1989 (Australia 1991). Whalers in the Timor Sea, Indonesia, have hunted Kogia (Weber 1923), although there is no evidence that they have been taken in recent years (Barnes 1991). Kogia, probably including Kogia bre- viceps, have been and probably still are taken occa- sionally in the Lesser Antilles (Caldwell and Caldwell 1975; Reeves 1988). Vidal et al. (1990) noted that Pygmy Sperm Whales were recorded caught in gillnet fisheries in Mexican or Central American waters, but presented no details on exact locations or numbers. This species is taken in direct- ed fisheries and incidentally in gillnet and seine fish- eries in the Philippines (Aragones et al. 1991; Dolar et al. 1991). Animals have also been caught inciden- tally in gillnets in the central North Pacific and off Sri Lanka (Omura et al. 1984; Leatherwood and 530 Reeves 1989). Ingestion of foreign objects (Jones 1984; Ross 1979, 1984) and boat collisions (Sylvestre 1988) may occasionally result in mortali- ty. Little has been reported on levels of environmen- tal contaminants in this species (Cockcroft et al. 1991). Evaluation Based on existing information the Pygmy Sperm Whale appears to be only a rare visitor to Canadian waters. No serious threats to its status in Canadian waters are apparent. As such, no COSEWIC status designation is required. Acknowledgments We thank the World Wildlife Fund (Canada) for financial assistance for the preparation of this report, Robert Campbell and COSEWIC for provid- ing assistance and support, Sherry Smrstik of the National Marine Mammal Laboratory for providing references, and Tamara Guenther and two anony- mous reviewers (Randall Reeves and David Gaskin) for helpful comments on the manuscript. We would also like to thank Donald McAlpine and Laurie Murison for allowing us to cite their manuscript in preparation. Literature Cited Allen, G. M. 1941. Pygmy sperm whale in the Atlantic. Field Museum of Natural History Zoology Series 27: 17-36. Aragones, L. 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Marine Mammal Science 3(4): 354-356. THE CANADIAN FIELD-NATURALIST Vol. 110 Vidal, O., K. van Waerebeek, L. T. Findley, and G. Alvarez-Manilla. 1990. Abstract. Cetaceans and gill- net fisheries in Mexico, Central America and the Caribbean: a preliminary review. Page 51 in Abstracts of the Symposium on Mortality of Cetaceans in Passive Fishing Nets and Traps, October 20-21, 1990, La Jolla, CA. i Waerebeek, K. van, J. C. Reyes, and B. A. Luscombe. 1987. A second record of the pygmy sperm whale Kogia breviceps (de Blainville, 1838) (Cetacea, Physeteridae) from the Peruvian coast. Zeitschrift fiir Saugetierkunde 52: 192-194. Weber, M. 1923. Die cetaceen der Siboga-Expedition. Siboga-Expeditie Monograph 58: 1-38. Yamada, M. 1954. Some remarks on the pygmy sperm whale, Kogia. Scientific Reports of the Whales Research Institute 9: 37-58. Zam, S. G., D. K. Caldwell, and M. C. Caldwell. 1971. Some endoparasites from small odontocete cetaceans collected in Florida and Georgia. Cetology 2: 1-11. Accepted 13 March 1996 Notes Tree-climbing by Arctic Ground Squirrels, Spermophilus parryii, in the Southwestern Yukon Territory ANNE H. Husss!, TIM KARELS, and ANDREA ByROM? Division of Life Sciences, Scarborough Campus, University of Toronto, Scarborough, Ontario MIC 1A4 'Present address: Department of Zoology, University of Western Ontario, London, Ontario N6A 5B7 Present address: Department of Zoology, University of British Columbia, Vancouver, British Columbia V6T 1W5 Hubbs, Anne H., Tim Karels, and Andrea Byrom. 1996. Tree-climbing by Arctic Ground Squirrels, Spermophilus parryii, in the southwestern Yukon Territory. Canadian Field-Naturalist 110(3): 533-534. Arctic Ground Squirrels (Spermophilus parryii) are commonly considered obligate ground-dwelling. However, seventy- eight Arctic Ground Squirrels were seen climbing or sitting in spruce trees and on deadfall and stumps 164 times in the the southwestern Yukon from 1991-1994. Trees, deadfall, and stumps, seemed to serve primarily as temporary refuges and/or predator surveillance sites, although some were occasionally used as nest sites. Arctic Ground Squirrels are not as obligate ground-dwelling as previously thought. Key Words: Arctic Ground Squirrel, Spermophilus parryii, climbing, nesting, boreal forest, Yukon. The Arctic Ground Squirrel (Spermophilus parryi) lives in a variety of habitats in North America ranging from Arctic and alpine tundra to boreal forest (Banfield 1974). This species has, how- ever, only been previously studied in tundra habitats where it is considered to be obligate ground- dwelling (Carl 1971; Green 1977; Woods 1980; McLean 1981; Lacey 1991). In the boreal forest, Arctic Ground Squirrels must not only contend with relatively high levels of predation (Hubbs and Boonstra, unpublished), but compensate for an obscured view of approaching predators. Here, we report on climbing of trees, deadfall, and stumps by Arctic Ground Squirrels within the Boreal Forest of the southwestern Yukon Territory (61°N, 138°W). Squirrels were observed from mid-April, when they emerged from hibernation, to late August, when they entered hibernation, on five 10 ha grids between 1991-1994 and on two additional 10 ha grids between 1992-1994. All grids were dominated by White Spruce (Picea glauca) with an understory of Bog Birch (Betula glandulosa), Grey Willow (Salix glauca), and grasses (Festuca altacia and Calamagrostis lapponica). Observations of squirrels climbing or sitting in live or dead trees, or on dead- fall and stumps greater than 1 m high were made during live-trapping and telemetry sessions. Seventy-eight of a possible 400 Arctic Ground Squirrels captured on the grids from 1991-1994 (19.5%) were observed in trees or on deadfall and stumps 164 times. These included 49 females (25 adults, 24 young-of-the-year), 16 males (7 adults, 9 young-of-the-year) and 13 young-of-the-year of unknown sex. Squirrels were seen on average, 2.5 times + 0.3 SE (median 1; maximum 13) per indi- vidual, at aboveground heights of 1.8 m + 0.2 SE (range 0.5 m - 6.0 m). Squirrels were most frequent- ly seen on deadfall (112 sightings), followed by trees (48 sightings with 21 in live trees and 27 in dead trees), and stumps (4). The majority of climbing events (156 or 95% of the total sightings) were observed when squirrels were approached by an observer, usually when squir- rels were away from their burrows or when their bur- row entrances were not readily accessible upon release from traps. Alarm calling frequently occurred once squirrels were in trees or atop deadfall and stumps. This suggests that squirrels used these struc- tures as temporary refuges against potential preda- tors, as has been reported for Woodchucks (Afarmota monax — Bowdish 1922; Cleveland 1922; Swihart 1982) and small mammals (Jedrzejewski et al. 1992), or as surveillance sites from which to monitor predator movements with the least visual obscurity. Columbian Ground Squirrels (S. columbianus) and Beldings Ground Squirrels (S. beldingi) have also been reported to use aboveground structures (rocks, burrow mounds, and fence posts) for vigilance (Tyser 1980; Sherman 1985; MacHutchon and Harestad 1990). The remaining 5% of climbing events involved six female squirrels (five adults, one young-of-the-year) D)518) 534 which appeared to be nesting within 1.5—2.0 m high woodpecker or Red Squirrel (Tamiasciurus hudsoni- cus) holes in dead spruce trees. Nesting was suggest- ed by a squirrel’s repeated presence in trees prior to morning emergence as determined from radio- telemetery and by the lactating status of four of the five adults. This behaviour was observed on only one grid with very high numbers (55-216 squirrels versus 7-119 on the other grids from 1991-1994; Hubbs and Boonstra, unpublished; A. Byrom, unpublished; T. Karels, unpublished) where burrow sites likely were limiting. Other than sites for nesting, refuge, or surveil- lance, squirrels may climb trees to obtain food, as observed for Woodchucks (D. M. Jedlicka, unpub- lished). Arctic Ground Squirrels will climb shrubs to feed on the buds, leaves, and flowers as will Columbian Ground Squirrels (Manville 1959) and Richardson Ground Squirrels (S. richardsonii: G. Michener, personal communication). Arctic Ground Squirrels have been observed climbing trees for food, but only when artificially supplemented foods have been provided (K. Stuart-Smith, personal com- munication). Squirrels will feed on spruce buds and arboreal lichens, but only when other food sources are scarce (Lincoln 1972; Batzli and Sobaski 1980). It therefore seems unlikely that the primary reason for Arctic Ground Squirrels climbing trees in the Boreal Forest of the southwestern Yukon was to obtain food. Our findings indicate that Arctic Ground Squirrels in the Boreal Forest will climb trees, deadfall, and stumps, and thus are not obligate ground-dwelling as previously suggested. The primary reason for climb- ing appears to be to gain access to temporary refuges and/or surveillance sites. This conclusion is based strictly on observational data, rather than experimen- tal analysis. To more accurately determine if this is the prime reason(s) for climbing, predators should be simulated and the frequency of tree-climbing deter- mined in simulated and unsimulated areas. To test the utility of these structures further, they could be added to unforested areas and their usage monitored (MacHutchon and Harestad 1990). Acknowledgments We thank R. Boonstra, B. Harrison, M. Havelka, D. Innes, N. McDonald, J. S. Millar, J. Murie, S. Teichert, M. Topping, S. Woolfenden for their help- ful criticism, and the Arctic Institute of North America. This research was funded by a Collaborative Special Project grant, an individual operating grant from the Natural Sciences and THE CANADIAN FIELD-NATURALIST Vol. 110 Engineering Research Council of Canada, and by a Canadian Commonwealth Scholarship to A. Byrom. This is contribution number 78 of the Kluane Boreal Forest Ecosystem Project. Literature Cited Banfield, A. W. F. 1974. The Mammals of Canada. University of Toronto Press, Toronto. 438 pages. Batzli, G. O., and S. T. Sobaski. 1980. Distribution, abundance, and foraging patterns of ground squirrels near Atkasook, Alaska. Arctic and Alpine Research 12: 501-510. Bowdish, B. S. 1922. Tree-climbing woodchucks. Journal of Mammalogy 3: 259. Carl, E. A. 1971. Population control in Arctic Ground Squirrels. Ecology 52: 395-413. Cleveland, P. 1922. Woodchucks climb trees. Journal of Mammalogy 3: 260-261. Green, J. E. 1971. Population regulation and annual cycles in the Arctic Ground Squirrel. M. Sc. thesis, University of British Columbia, Vancouver, British Columbia. Jedrzejewski, W., B. Jedrzejewska, and E. McNeish. 1992. Hunting success of the weasel Mustela nivalis and escape tactics of forest rodents in Bialowieza National Park. Acta Theriologica 37: 319-328. Lacey, E. A. 1991. Reproductive and dispersal strategies of male Arctic Ground Squirrels (Spermophilus parryii plesius). Ph.D. thesis, University of Michigan, Ann Arbour, Michigan. Lincoln, B. J. 1972. Behavior study of the Arctic Ground Squirrel. I.R.R.P. Science Results 3: 245-253. MacHutchon, A. G., and A. S. Harestad. 1990. Vigilance behaviour and use of rocks by Columbian ground squirrels. Canadian Journal of Zoology 68: 1428-1432. Manville, R. H. 1959. The Columbian ground squirrel in northwestern Montana. Journal of Mammalogy 40: 26-45. McLean, I. G. 1981. Social ecology of the Arctic Ground Squirrel Spermophilus parryii. Ph.D. thesis, University of Alberta, Edmonton, Alberta. Sherman, P. W. 1985. Alarm calls of Belding’s ground squirrels to aerial predators: nepotism or self-preserva- tion? Behavioural Ecology and Sociobiology 17: 313-323. Swihart, R. K. 1982. Scansorial behavior in woodchucks, Marmota monax. Canadian Field-Naturalist 96: 215-216. Tyser, R. W. 1980. Use of substrate for surveillance behaviors in a community of talus slope mammals. American Midland Naturalist 104: 33-38. Woods, S. E., Jr. 1980. The squirrels of Canada. National Museums of Canada, Ottawa. 199 pages. Received 13 October 1995 Accepted 13 February 1996 1996 NOTES 535 Do Female Sharp-tailed Grouse, Tympanuchus phasianellus, Copulate Only Once During a Breeding Season? LEONARD J. S. Tsust Department of Biology, York University, North York, Ontario M3J 1P3 Tsuji, Leonard J. S. 1996. Do female Sharp-tailed Grouse, Tympanuchus phasianellus, copulate only once during a breed- ing season? Canadian Field-Naturalist 110(3): 535-536. Fluid mixtures taken from the reproductive tracts of eight female Sharp-tailed Grouse (Tympanuchus phasianellus), as they entered the lek, did not contain sperm or sperm fragments. Although not definitive, this study adds further support to the assumption that female Sharp-tailed Grouse typically copulate only once during a breeding season. Key Words: Sharp-tailed Grouse, Tympanuchus phasianellus, oviduct, sperm, copulation. It has been assumed in classically lekking grouse species that females typically copulate only once during a breeding season (except for renesting attempts). This perception was based initially on two types of field observations: (1) mating peaks at a lek closely correlated with hatching peaks of the young, 37 to 44 days later (Dalke et al. 1963; Svedarsky 1979); and (2) marked females observed copulating one day were not seen during subsequent display periods, at the same lek (Lumsden 1968; Kermott 1982). In recent studies by Hoglund et al. (1990) and Alatalo et al. (1991), female Black Grouse (Tetrao tetrix) were radio-marked and their copulatory behavior recorded. In the study by Alatalo et al. (1991), 79% of females (n=23) were known to have copulated only once at a given lek, while Hoglund et al. (1990) reported that 83% of the females (n=23) were known to have copulated only once at a given lek. Hoglund et al. (1990) also reported that females that copulated more than once were often individuals associated with a disturbed mating. This trend to remate after a disturbed mating was also observed for female Sharp-tailed Grouse (Tympanuchus phasianellus) (Gratson 1989). Thus, it appears that among classically lekking grouse species, females typically copulate only once during a breeding sea- son, unless disturbed. However, if one acknowledges that females can visit more than one lek during a breeding season (e.g. Kruijt et al. 1972), it becomes difficult to rule out multiple copulations because it would be diffi- cult logistically, to observe all leks (during all morn- ing and evening display periods) in a specified region for the entire breeding season, even if the females were marked. Another test of the assumption would be to collect females during the breeding sea- son as they entered a lek, prior to copulation on that lek. Galliforms possess uterovaginal glands that store and constantly release sperm (Compton et al. 1978; Bakst 1981). Thus, female reproductive sys- tems could be examined for the presence of sperm to establish whether or not prior copulation had occurred. If females typically copulate only once, approximately 80% of the female reproductive tracts should be without sperm. Eight female Sharp-tailed Grouse were “harvest- ed” by native Canadians near Fort Albany, Ontario, 52°15'N; 81°35’'W, as the birds entered leks during the 1992-1994 breeding seasons. Reproductive tracts were processed using a technique modified from Bakst (1981). Oviducts were removed and flushed by syringe with 15% buffered formalin (after the distal end of the oviduct was ligated with 3-0 gut suture). Individual fluid mixtures (contain- ing 15% buffered formalin and the flushed oviduct contents) were then emptied into separate test tubes, centrifuged, and the sediment examined in wet mount (for 5 minutes) under a microscope (x 100, x 250 objectives) for either sperm or sperm frag- ments. No fluid mixture contained sperm or sperm fragments. Although the results do not provide definitive evi- dence, they add further support for the assumption that female Sharp-tailed Grouse mate only once dur- ing a breeding season. Acknowledgments I thank the native hunters who allowed the sal- vaging of grouse remains. Comments on the manuscript from A. J. Erskine and an anonymous reviewer were appreciated. James Bay General Hospital kindly provided space and use of their equipment. Partial funding was provided by a Northern Scientific Training Grant (Department of Indian and Northern Affairs). Literature Cited Alatalo, R. V., J. Hoglund, and A. Lundberg. 1991 Lekking in the Black Grouse - a test of male viability. Nature 352: 155-156. Bakst, M. R. 1981. Sperm recovery from turkeys at known intervals after insemination and oviposition. Journal of Reproduction and Fertility 62: 159-164. Compton, M. M., H. P. Van Krey, and P. B. Siegel. 1978. The filling and emptying of the uterovaginal sperm-host glands in the domestic hen. Poultry Science 57: 1696-1711. 536 Dalke, P. D., D. B. Pyrah, D. C. Stanton, J. E. Crawford, and E. P. Schlatterer. 1963. Ecology, productivity, and management of Sage Grouse in Idaho. Journal of Wildlife Management 27: 813-841. Gratson, M. W. 1989. Sexual selection on Sharp-tailed Grouse leks. Ph. D. dissertation. University of Victoria, Victoria, British Columbia. Hoglund, J., R. V. Alatalo, and A. Lundberg. 1990. Copying the mate choice of others? Observations on female Black Grouse. Behaviour 114: 221-231. Kermott, L. H. 1982. Breeding behavior in the Sharp- tailed Grouse. Ph. D. dissertation. University of Minnesota, St. Paul, Minnesota. THE CANADIAN FIELD-NATURALIST Vol. 110 Kruijt, J. P., G. J. DeVos, and I. Bossema. 1972. The arena system of the Black Grouse (Lyrurus tetrix tetrix L.). Proceedings of the International Ornithological Congress 15: 399-423. Lumsden, H. G. 1968. The displays of the Sage Grouse. Ontario Department of Lands and Forests, Research Branch, Research Report 83. Toronto, Ontario. : Svedarsky, W. D. 1979. Spring and summer ecology of female greater prairie chickens in northwestern Minnesota. Ph. D. dissertation. University of North Dakota, Grand Forks, North Dakota. Received 9 October 1995 Accepted 16 January 1996 Interactions of a White-winged Black Tern, Chlidonias leucopterus, with Arctic Terns, Sterna paradisaea, at Churchill, Manitoba JOSEPH R. JEHL, JR. Hubbs/Sea World Research Institute, 2595 Ingraham St., San Diego, California 92109 Jehl, Joseph R., Jr. 1996. Interactions of a White-winged Black Tern, Chlidonias lecopterus, with Arctic Terns, Sterna paradisaea, at Churchill, Manitoba. Canadian Field-Naturalist 110(3): 536-537. A White-winged Black Tern (Chlidonias leucopterus), a species of Eurasian provenance, appeared at Churchill, Manitoba, in June—July 1995, where it was persistently harassed by breeding Arctic Terns (Sterna paradisaea). Key Words: White-winged Black Tern, Chlidonias leucopterus, Arctic Tern, Sterna paradisaea, agonistic behaviour. On 24 June 1995, Arnet Sheppard of Ottawa, Ontario, videotaped an adult White-winged Black Tern (Chlidonias leucopterus) in alternate plumage at the mouth of the Churchill River, Manitoba. There are fewer than a dozen records for this temperate Eurasian species in Canada, and this was the first in central Canada and the subarctic. Sheppard’s video- tape and subsequent sightings and photographs con- firmed his identification (R. Koes, personal commu- nication). During Sheppard’s 5-minute observation period the tern’s behavior was unremarkable. It flew along the shoreline, landed for a few minutes among a small flock of Arctic Terns (Sterna paradisaea) and Bonaparte’s Gulls (Larus philadelphia), and then disappeared upriver toward the town of Churchill, after being attacked by a Parasitic Jaeger (Stercorarius parasiticus). On 27 June, Jan van Gils and Irene Tieleman, researchers from The Netherlands, told me of a black tern whose characteristics fitted C. leucepterus that they had seen near the Arctic Tern colony at West Twin Lake, 26 km south-east of Churchill. That evening, Scott Yaeger and I visited the area for 15 minutes at dusk and photographed a single White- winged Black Tern (Figure 1) near a small, marshy island where 16 pairs of Afctic Terns were just start- ing to lay. It seemed to be exploiting a major emer- gence of dragonflies and would fly | km from the island, either over the lake or to the edge of the for- est, catch a flying insect, then return to the colony, where it repeatedly attempted to land. On each approach the Arctic Terns immediately left their nests and aggressively chased it (Figure 2). Although the White-winged Black Tern managed to land twice, the harassment never ceased entirely and its time on the ground was momentary. By the next morning the tern — and the dragonflies — had dis- appeared. Although many observers visited the colony in the following days, the White-winged Black Tern never reappeared there, and it was not rediscovered until 13 July, when T. and B. Holcombe of Tonbridge, Kent, England, saw it back on the Churchill River near the townsite. During their 45 minutes of obser- vation it was “savagely pursued” by Arctic Terns, which on one occasion literally knocked it into the river. The aggression was confined to flight periods, and ceased when the tern landed among Bonaparte’s Gulls and Arctic Terns; interestingly, it was always the last bird in the flock to land. The following day, B. Chartier also saw Arctic Terns harassing the tern in the same region; again, chasing ceased when the tern landed. The Holcombes’ and Chartiers’ observa- | | 1996 copterus) at West Twin Lake, Churchill, Manitoba, 27 June 1995. tions were made within 0.5-1 km of an Arctic Tern colony of approximately 100 birds. The far northern and usually coastal breeding range of the Arctic Tern is so different from that of the temperate and marsh-nesting White-winged Tern that the two species can come together only by acci- dent. Thus, it will be difficult to learn whether the interactions described above represented a general response to intruders in the colony by Arctic Terns, which are generally considered to be very aggres- Sive, or a specific response to the bold black-and- white flight pattern of the White-winged Tern. That the former might have been the case is suggested by a further observation from Churchill (Y. Zharikov, 12 June 1996) of Arctic Terns chasing a Black Tern (Chlidonias niger); the latter is a dark-bodied, but less boldly-marked, North American species that is closely related to the White-winged Tern. Farther NOTES FiGurE 2. Two Arctic Terns (Sterna paradisaea, left) chas- ing a White-winged Black Tern (Chlidonias leu- copterus) as it attempted to land in their colony. south in the interior of North America, light- plumaged (e.g., Common [S. hirundo] and Forster’s [S. forsteri]) terns nest amicably with Black Terns (W. Scharf, personal communication). Additional observations of interactions between light- and dark- bodied terns in the breeding season would be of interest. Acknowledgments I thank Arnet Sheppard, Brenda and Tony Holcombe, Jan van Gils, Irene Tieleman, Scott Yaeger, Bonnie Chartier, Yuri Zharikov and Rudolf Koes for their help in the preparation of this account, and E. Dunn and A. J. Erskine for comments on the manuscript. Received 7 November 1995 Accepted 17 April 1996 538 THE CANADIAN FIELD-NATURALIST Vol. 110 Predation of an Eastern Chipmunk, Jamias striatus, by a Gray Squirrel, Sciurus carolinensis STEVEN D. FACCIO Vermont Institute of Natural Science, RR2 Box 532, Woodstock, Vermont 05091 Faccio, Steven D. 1996. Predation of an Eastern Chipmunk, Tamias striatus, by a Gray Squirrel, Sciurus carolinensis. Canadian Field-Naturalist 110(3): 538. I observed a Gray Squirrel (Sciurus carolinensis), attack and kill an Eastern Chipmunk (Tamias striatus) in Orange, Massachusetts. Published reports of Gray Squirrel predation upon other vertebrates consist only of bird eggs and nestlings. Key Words: Gray Squirrel, Sciurus carolinensis, Eastern Chipmunk, Tamias striatus, predation, Massachusetts. Animal foods are relatively uncommon in the diet of Gray Squirrels (Sciurus carolinensis) (Nichols 1958; Flyger and Gates 1990). Madsen (1964) indi- cated that squirrels frequently obtain calcium and other minerals from scavenged bones, shed deer antlers, and old turtle shells, while others report Gray Squirrels feeding on a variety of invertebrate prey (Hamilton 1943; Layne and Woolfenden 1958; DeGraaf and Rudis 1986). However, all published reports of Gray Squirrel predation on other verte- brates consisted of bird eggs and nestlings (Thoms 1922; Godin 1977). I report on a Gray Squirrel killing and apparently feeding on an Eastern Chipmunk (Tamias striatus). My observation was made on the shore of Lake Mattawa, in the town of Orange, Worcester County, Massachusetts (ca. 42° 35'N, 72° 20'W). This man- made lake is surrounded by both seasonal and per- manent homes within a mixed forest community dominated by White Pine (Pinus strobus), Eastern Hemlock (Tsuga canadensis), Red Oak (Quercus rubra), and Red Maple (Acer rubrum). The under- story along the lakeshore is dominated by low Blueberries (Vaccinium spp.) and Black Huckleberry (Gaylussacia baccata) along with vari- ous ferns and herbs. On 16 August 1993, attracted by a persistent screaming, I observed a Gray Squirrel attacking an Eastern Chipmunk. Clinging to the trunk of a White Pine approximately 1.5 m above the ground, the squirrel was grasping the screaming chipmunk by the back of the neck. The squirrel immediately car- ried the chipmunk up the tree to a leaf-nest approxi- mately 10 m high where visual contact was lost. The chipmunk continued vocalizing for 5-10 sec- onds until an object fell from the nest. The squirrel immediately descended the tree and retrieved what appeared to be the chipmunk's head, and returned to the nest. Since I did not witness the initial encounter I do not know if the squirrel actively pursued the chip- munk or preyed on it opportunistically. Other poten- tial food sources were abundant at the time of the observation including blueberries, huckleberries, mushrooms, and a compost pile well-stocked with table scraps, indicating that food shortage was not a likely explaination for this predation. Although no direct evidence exists, it seems likely that the squir- rel consumed the chipmunk based on the effort it expended to kill the animal, retrieve the head, and carry it to a nest. Literature Cited DeGraaf, R. M., and D. D. Rudis. 1986. New England Wildlife: Habitat, Natural History, and Distribution. General Technology Report NE-108. Broomall, Pennsylvania: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 491 pages. Flyger, V., and J. E. Gates. 1990. Fox and Gray Squirrels. Pages 209-229, In Wild Mammals of North America. Edited by J. A. Chapman and G. A. Feldhamer. Johns Hopkins University Press, Baltimore, Maryland. Godin, A. J. 1977. Wild Mammals of New England. The Johns Hopkins University Press, Baltimore, Maryland. 304 pages. Hamilton, W. J., Jr. 1943. Caterpillars as food of the gray squirrel. Journal of Mammalogy 24: 104. Layne, J. N., and G. E. Woolfenden. 1958. Gray squirrels feeding on insects in car radiators. Journal of Mammalogy 39: 595-596. Madsen, J. 1964. Gray and fox squirrels. Olin Mathieson Chemical Corporation, East Alton, Illinois. 112 pages. Nichols, J. T. 1958. Food habits and behavior of the gray squirrel. Journal of Mammalogy 39: 376-380. Thoms, C. S. 1922. Are squirrels bird-enemies? Bird Lore 24: 207. Received 31 October 1995 Accepted 30 January 1996 1996 NOTES 5a) Late Spring Arrival, Nesting, and Fall Departure by Common Nighthawks, Chordeiles minor, in Saskatchewan in 1995 Ray G. PouLin, PAUL A. BRADSHAW AND MARK D. GRAHAM Biology Department, University of Regina, Regina, Saskatchewan S4S 0A2 Poulin, Ray G., Paul A. Bradshaw, and Mark D. Graham. 1996. Late spring arrival, nesting, and fall departure by Common Nighthawks, Chordeiles minor, in Saskatchewan in 1995. Canadian Field-Naturalist 110(3): 539-540. In 1995, Common Nighthawks (Chordeiles minor) were late arriving from their spring migration, late laying eggs and late migrating from Saskatchewan. Common Nighthawks were observed near Buffalo Pound Provincial Park, Saskatchewan, on and up to 17 October and two and possibly three individuals were observed in Regina on 19 October. To our knowl- edge, this is almost 3 weeks later than has ever been recorded at this latitude. Although the ambient temperature was near 0°C, these nighthawks were not in torpor. Key Words: Common Nighthawk, Chordeiles minor, migration, Saskatchewan, torpor. Common Nighthawks (Chordeiles minor) are the most widely distributed of the North American caprimulgids. Breeding occurs from the Northwest Territories and southern Yukon in the north to Mexico and Central America in the south, and from the Atlantic to the Pacific ocean across most of the continent. Although little is known about the loca- tion of their wintering grounds, they are thought to reside throughout much of South America, as far south as Uruguay (Bent 1940; Poulin et al. in press). During spring migration, they arrive at the Canada/United States border (49°N, west of 99°W), during the last week of May and first week of June (Bent 1940; Rust 1947; Poulin et al. in press). In late summer, nighthawks gather in large flocks, some- times greater than a thousand birds, to migrate south. They begin their migration from southern Canada/northern United States (49°N) by the third week of August and are generally thought to have left these areas completely by the second or third week of September (Campbell et al. 1990). The timing of fall migration is likely dictated, at least partly, by the availability of food. Nighthawks feed on flying insects during crepuscular (dusk and dawn) hours. Cool temperatures, which prevail dur- ing spring and autumn, result in reduced numbers of flying insects for nighthawks to feed on and may influence the timing of migration. One physiological adaptation employed by several species of caprimulgids to cope with unfavourable conditions is torpor (Reinertsen 1983). Torpor allows these birds to reduce metabolism as a means of con- serving energy during unfavourable conditions. Inclement weather, including low ambient tempera- tures, extended periods of rain and/or strong winds are known to induce torpor in Common Poorwills (Phalaenoptilus nuttallii) (Brigham 1992; Kissner and Brigham 1993; Csada and Brigham 1994). Lasiewski and Dawson (1964) induced torpor in captive Common Nighthawks by starving them. However, only one of four birds survived a drop in body temperature below 25°C. Firman et al. (1993) found no evidence of torpor by Common Nighthawks in the Okanagan Valley, British Columbia, even though the summer of 1990 was much colder and wetter than normal. Brigham et al. (in press) provide limited evidence that Common Nighthawks are capable of entering tor- por. One nighthawk in British Columbia and another in Alberta were found in an apparent torpid state, at ambient temperatures of 4°C and 5°C respectively. Brigham et al. did not determine if the birds were able to successfully re-warm and fly away. At Coeur d'Alene, Idaho, the first observed south- ward flights of Common Nighthawks usually occur in August (66% of the time) but are often delayed until September (34% of the time)(Rust 1947). Between 1911 and 1946 the latest recorded sighting of a Common Nighthawk at Coeur d'Alene was 24 September (Rust 1947). Bent (1940) reports the following late sighting dates for comparable locations: Okanagan Landing, British Columbia, 15 September; Tacoma, Washington, 11 September; Weston, Oregon, 9 September; Banff, Alberta, 17 September; Meridian, Idaho, 15 September; East-end, Saskatchewan, 15 September; Columbia Falls, Montana, 28 September; Great Falls, Montana, 6 October; Yellowstone Park, Wyoming, 15 September; Winnipeg, Manitoba, 19 September; Charlson,, North Dakota, 22 September and a casual record of one bird at Allakaket, Alaska in either late September or early October. The Okanagan Valley of British Columbia (49° 18'N, 119° 31'W) is a semi-arid region composed of a series of lakes linked by the Okanagan River. This area typically attracts large numbers of nighthawks (Cannings et al. 1987) and one might expect that nighthawks would remain later into the summer there than any other place in Canada. The latest recorded date of a nighthawk in that area was 30 September and the mean departure date was 18 September (Cannings et al. 1987). 540 Migration compels fledgling nighthawks to achieve maturation and independence by approxi- mately the last week of August. Eggs are incubated for 17 to 20 days and young are able to fly well at 25 to 30 days of age but typically take 45 to 50 days to develop fully (Bent 1940). Therefore, in order for a chick to be capable of migration before the last day of August, eggs must be laid on or before 30 June, and must hatch on or before 16 July. In previous years (1990-1994), Common Night- hawks were observed by approximately the first week of June in Cypress Hills Provincial Park, Saskatchewan (49° 34'N, 109° 53'W) (R. M. Brigham, personal communication). During the sum- mer of 1995, the first sighting of a nighthawk in Cypress Hills was 28 May (R. G. Poulin), but not until 13 June was more than one nighthawk observed in an evening. This suggests that nighthawks were at least one week late arriving in 1995. On 22 July a nest was found with a female incu- bating a single egg, this egg hatched on 6 August. Although the fate of this chick is unknown, given the developmental time requirements, this chick would not have been ready to migrate until at least 21 September, three weeks later than nighthawks nor- mally leave Cypress Hills (R. M. Brigham, personal communication). On and up to 17 October 1995, at approximately 11:00 each day, Common Nighthawks were observed (M. D. Graham) flying near Buffalo Pound Provincial Park, Saskatchewan (50° 40'N, 105° 30'W). These were individual birds, not the large migratory flocks normally observed at the end of August/beginning of September. The night-time minimum temperature on 17 October was 2.4°C (Environment Canada). On 19 October, at least two, and possibly three Common Nighthawks were observed (P.A. Bradshaw) Regina, Saskatchewan (50° 25'N, 104° 39'W). At 07:00, one nighthawk was sitting alertly on a lamp post while one and possibly two other nighthawks flew near by. The overnight minimum on this date was -3.2°C and from this date onward overnight minimum temperatures did not rise above O°C until the end of the winter season (Environment Canada). The months of September and October were normal in terms of average temperatures and precipitation (Environment Canada). The observations of Common Nighthawks on 17 and 19 October are almost three weeks later than has ever been recorded at this latitude in western Canada, following the trend of the late spring migra- tion, and late breeding in 1995. An important impli- cation of the observations on the morning of 19 October is that these nighthawks were not in torpor during very cold temperatures (-3°C), where food THE CANADIAN FIELD-NATURALIST Vol. 110 availability was undoubtedly low. By comparison, Common Poorwills in the Okanagan, British Columbia used torpor consistently when night time temperatures fell below +1°C (Brigham 1992). Acknowledgments We thank R. Mark Brigham for his encourage- ment and advice, plus Danielle Todd and Matina C. Kalcounis for constructive reviews. Research on goatsuckers is funded by a Natural Sciences and Engineering Research Council (NSERC) Canada Research Grant to R. Mark Brigham and an Edgar A. Wahn Scholarship to R. G. Poulin. Literature Cited Bent, A. C. 1940. Life histories of North American cuck- oos, goatsuckers, hummingbirds, and their allies, part 1. United States National Museum Bulletin 176. Brigham, R. M. 1992. Daily torpor in a free-ranging goat- sucker, the Common Poorwill (Phalaenoptilus nuttallii). Physiological Zoology 65: 457-472. Brigham, R. M., K. H. Morgan, and P. C. james. In press. Evidence that free-ranging Common Nighthawks can enter torpor. Northwestern Naturalist 76. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowen, J. M. Cooper, G. W. Kaiser, and C. E. McNall. 1990. - | The Birds of British Columbia, Volume II. Royal British Columbia Museum, Victoria. Cannings, R. A., R. J. Cannings, and S. G. Cannings. 1987. Birds of the Okanagan Valley, British Columbia. Royal British Columbia Museum, Victoria. Csada, R. D., and R. M. Brigham. 1994. Reproduction constrains the use of daily torpor by free-ranging com- mon poorwills (Phalaenoptilus nuttallii). Journal of Zoology (London) 234: 209-216. Firman, M. C., R. M. Brigham, and R. M. R. Barclay. 1993. Do Free-Ranging Common Nighthawks Enter Torpor? Condor 95: 157-162. Kissner, K. J., and R. M. Brigham. 1993. Evidence for the use of torpor by incubating and brooding common poorwills Phalaenoptilus nuttallii. Ornis Scandinavica 24: 333-334. Lasiewski, R. C., and W. R. Dawson. 1964. Physiological responses to temperature in the common nighthawk. Condor 66: 477-490. Poulin, R. G., S. D. Grindal, and R. M. Brigham. /n press. Common Nighthawk in The Birds of North America. Edited by A. Poole, P. Stettenheim and F. Gill. Philadelphia: The Academy of Natural Sciences; Washington DC: The American Ornithologists’ Union. Reinertsen, R. E. 1983. Nocturnal hypothermia and its energetic significance for small birds living in the arctic and subarctic regions. A review. Polar Research 1: 269-284. Rust, H. J. 1947. Migration and nesting of Nighthawks in Northern Idaho. Condor 49: 177-188. Received 4 December 1995 Accepted 6 February 1996 1996 NOTES 541 Horsehair Fungus, Marasmius androsaceus, Used as Nest Lining by Birds of the Subalpine Spruce-fir Community in the Northeastern United States K. P. MCFARLAND, and C. C. RIMMER Vermont Institute of Natural Science, RR 2, Box 532, Woodstock, Vermont 05091 McFarland, K. P., and C. C. Rimmer. 1996. Horsehair fungus, Marasmius androsaceus, used as nest lining by birds of the subalpine/spruce-fir community in the northeastern United States. Canadian Field-Naturalist 110(3): 541-543. We examined 54 nests of 10 species on Mt. Mansfield and Mt. Equinox, Vermont and Plateau Mountain, New York in the subalpine Red Spruce (Picea rubens) — Balsam Fir (Abies balsamea) forest bird community in the northeastern United States in 1994 and 1995. Forty-six nexts (85%) were found to contain fine, black, hair-like strands as lining material. This material was defined as the rhizomorphs of Horsehair Fungis (Marasmius androsaceus) and represents the first document- ed use of this material by North American breeding bird species. We believe that many earlier descriptions of nest lining materials used by boreal and subalpine bird species may refer to mis-identified Horsehair Fungus. Our finding that 85% of nests in montane spruce-fir forests of the northeastern United States contained copius amounts of rhizomorphs suggests that this material is an integral component of nest construction in this habitat. Key Words: Horse Hair Fungus, Marasmius androsaceus, nest material, birds, subalpine spruce-fir forest. Several pieces of mushrooms in the genus Marasmius are known to produce copious amounts of thin, black and wiry rhizomorphs as a means of colonizing twigs and leaves (Redhead 1989; Seaver 1994; Singer 1986). The use of marasmioid rhi- zomorphs in the construction of bird nests has not been documented for any species breeding in North America. There are several repors of marasmioid rhi- zomorphs used as nesting material in China (Desjardin, personal communication fide Mu Zang), Cameroon (Desjardin, personal communication), and Brazil (Sick 1957). In Argentina, Golden-winged Cacique (Cacius chrysopterus) and Red-rumped Cacique (Cacicus haemorrhous) construct their nests principally of rhizomorphs from Marasmius crinise- qui (Wright and Ferraro 1986). Hummingbirds col- lect the rhizomorphs of Marasmius nigrobrunneus in Ecuador for use in nest construction (Hedger 1990). In this paper we report the first documented use of Horsehair Fungus (Marasmius androsaceus) rhi- zomorphs as nest lining material by several North American breeding bird species. During demographic studies of the subalpine Red Spruce (Picea rubens)-Balsam Fir (Abies balsamea) forest bird community in the northeast- ern United States in 1994 and 1995 we examined 54 nests of 10 species at three different sites: Mt. Mansfield and Mt. Equinox, Vermont and Plateau Mountain, New York (Table 1). Forty-six nests (85%) were found to contain fine, black, hair-like strands as lining material. While collecting micro- habitat data around nest sites in 1994 we found large amounts of this material on live Balsam Fir trees and dead wood in the understory. These strands, which were found to have small basid- iomes, were compared with those occurring in the nests and found to be identical. Close examination of the strands in the nests revealed that some also had small, dried basidiomes. Samples of the hair-like structures and caps were sent to the North American Marasmius expett, Dennis Desjardin, Director of H. D. Thiers Herbarium, San Francisco State University, for iden- tification. The hair-like structures were identified as rhizomorphs of the Horsehair Fungus (Marasmius androsaceus). The species is saprotrophic on nee- dles, leaves and twigs and parasitic on some erica- ceous plants in mesic and boggy situations (Gilliam 1976; MacDonald 1949; Redhead 1989). Horsehair Fungus is considered common in North America found across the boreal zone and south along the Rockies, Coastal Mountains and the Appalachians (Redhead 1989: Map Fig. 9). Marasmius andro- saceus belongs to a group of closely related species, all of which produce numerous rhizomorphs. Marasmius pallidocephalus also occurs in the Northeast (Gilliam 1976; Redhead 1989) and could be used by birds in nest construction too (Desjardin, personal communication). Various species of the lichen genus Alectoria may resemble black, hair-like structures and could also be used. Many published descriptions of avian nesting material that contain references to hair-like material or fine rootlets may actually represent the rhi- zomorphs that we identified. Wallace (1939) con- ducted life-history studies of Bicknell’s Thrush (Catharus bicknelli) on Mt. Mansfield, Vermont, and thoroughly described the construction of the nest. However, he was unable to identify the inner lining of “fine, black rootlets,” and stated, “they are unquestionably rootlets of some sort ... resembling horsehair, but where the birds get them is a mys- tery.” The Bicknell’s Thrush, formerly a subspecies of the Gray-cheeked Thrush (Catharus minimus), 542 THE CANADIAN FIELD-NATURALIST TABLE 1. Number of bird nests in the subalpine spruce-fir forest found to contain Marasmius androsaceus rhizomorphs used as nest lining. Mt. Equinox, Vermont Present Yellow-bellied Flycatcher (Empidonax flaviventris) - - Brown Creeper (Certhia americana) 0) 1 Bicknell’s Thrush (Catharus bicknelli) ~ — Swainson’s Thrush (Catharus ustulatus) = = Magnolia Warbler (Dendroica magnolia) i 0) Yellow-rumped Warbler (Dendroica coronata coronata) 2 0 Blackpoll Warbler (Dendroica striata) 1 0 White-throated Sparrow (Zonotrichia albicollis) 0 1 Dark-eyed Junco (Junco Hyemalis) - — Purple Finch (Carpodacus purpureus) 0 1 Absent Plateau Mountain, New York Mt. Mansfield, Vermont Horsehair Fungus in Nest Lining Present Absent Present Absent = = 1 0 = _ 15 0) 2 0 2, 0) 1 0) 4 0 1 0) 16 0) = = 0 4 - — 0 1 “_” indicates no nests found at that site. has recently been given full species status (Ouellet 1993; American Ornithologists’ Union 1995). Bent (1949) reported several nest descriptions of minimus. In each case it appeared that the nest lining was composed predominantly of fine grasses, with one nest containing a few dried leaves and rootlets. Bent (1953) described the Magnolia Warbler’s (Dendroica magnolia) nest as “...lined invariably with fine black rootlets, which closely resemble horse-hairs. The lining of black rootlets is present in these and in all other nests of the Magnolia Warbler that I have seen, it seems to be characteristic of the species and will distinguish the nest from those of other warblers.” However, we and several other authors have described the use of this material in large quantities by other warbler species. Similarly, Bent (1953) reported that the Yellow- rumped Warbler (Dendroica coronata coronata) nest was {*...firmly interwoven with black horsehair, or perhaps moose hair, and finer rootlest...”” Brewster (1882) described a nest of the Blackpoll Warbler (Dentroica striata) in the Magdalen Islands, Gulf of St. Lawrence, as having a lining “...of slender, black moss-stems (which curiously resemble horse hair).” Bent (1942) related nest descriptions of Yellow- bellied Flycatcher (Empidonax flaviventris) from several people. In each case the lining material was described as a dark or black hair-like rootlet. Most notable was a description by Fendire, “...the black and shining rootlets of, apparently, ferns, closely resembling horsehair.” Bent (1949) similarly pre- sented Swainson’s Thrush (Catharus ustulatus) nest descriptions, including one from Maine where the lining was described as containing “...black, thread- like parts of the roots of decayed cinnamon ferns.” Finally, Bent (1968) noted that the nest lining of the Purple Finch (Carodacus purpureus) contained “...finer rootlets and horsehair.” We believe that many of these earlier descriptions of nest lining materials used by boreal and subalpine bird species referred to mis-identified Horsehair Fungus. Our finding that 85% of nests in montane spruce-fir forests of the northeastern United States contained copious amounts of rhizomorphs suggests that this material is an integral component of nest construction in this habitat. A number of Marasmius species have been shown to produce antibiotic agents that inhibit the growth of Staphylococcus (Melin et al. 1947). However, the species, Marasmius androsaceus, was practically inactive upon Staphylococcus cultures (Melin et al. 1947). While it is possible that Marasmius androsaceus rhizomorphs may be an effective agent against nest pathogens and parasites of subalpine birds in the Northeast, this is not known. Alternatively, these rhizomorphs may simply pro- vide the best of most easily obtainable material in this habitat for lining nests. A voucher specimen of the fungus has been deposited in the H. D. Thiers Herbarium. Bicknell’s Thrush, Swainson’s Thrush, Blackpoll Warbler, and Yellow-rumped Warbler nests with rhizomorph lin- ing are stored at the Vermont Institute of Natural Science. Acknowledgments We thank D. E. Desjardin and R. Van de Poll for the identification of the voucher specimen. D. 1996 Froelich, J. Goetz, T. Johanssen, J. Chase, K. Bevenstein, D. Lambert, S. Faccio, and C. McFarland helped locate and monitor nests. R. Paradis of the University of Vermont, R. Apple of the Mt. Mansfield Company, and the summit care- takers of the Green Mountain Club all provided logistic support. This project was funded in part by the U.S. Fish and Wildlife Service, the U.S. Forest Service, the Fish and Wildlife Foundation, the Vermont Monitoring Cooperative, the Wharton Trust, and the members and trustees of the Vermont Institute of Natural Science. This paper was improved by the constructive comments of D. E. Desjardin, A. E. Erskine, S. A. Redhead, and an anonymous reviewer. Literature Cited American Ornithologists’ Union. 1995. Fortieth supple- ment to the AOU checklist of North American birds. Auk 112: 819-830. Bent, A. C. 1942. Life histories of North American fly- catchers, larks, swallows, and their allies. United States National Museum Bullein 179. Bent, A. C. 1949. Life histories of North American thrushes, kinglets, and their allies. United States National Museum Bulletin 196. Bent, A. C. 1953. Life histories of North American wood warblers. United States National Museum Bulletin 203. Bent, A. C. 1968. Life histories of North American cardi- nals, grosbeaks, buntings, towhees, finches, sparrows, and allies. United States National Museum Bulletin 237. NOTES 543 Brewster, W. 1881. Notes on some birds and eggs from the Magdalen Islands, Gulf of St. Lawrence. Bulletin of the Nuttall Ornithological Club 7: 253-256. Gilliam, M. S. 1976. The genus Marasmius in the north- eastern United States and adjacent Canada. Mycotaxon: 4: 1-144. Hedger, J. 1990. Fungi in the tropical forest canopy. Mycologist 4(4): 200-202. Melin, E., T. Wiken, and K. Oblom. 1947. Antibiotic agents in the substrates from cultures of the genus Marasmius. Nature 159: 840-841. Ouellet, H. 1993. Bicknell’s Thrush: taxonomic status and distribution. Wilson Bulletin 105: 253-256. Redhead, S. A. 1989. A biogeography overview of the Canadian mushroom flora. Canadian Journal of Botany 67: 3003-3062. Seaver, F. J. 1994. The Horse-hair Fungi. Mycologia 36: 340-342. Sick, H. 1957. Rosthaarpilze als Nestbau-Material Brasilianischer Vogel. Journal fur Ornithologie 98 (40): 421-431. Singer, R. 1986. The Agaricales in Modern Taxonomy. 4th Edition. Koeltz Scientific Books, Koenigsten. 981 pages. Wallace, G. J. 1939. Bicknell’s Thrush: its taxonomy, distribution, and life history. Proceedings of the Boston Society of Natural History 41: 211-402. Wright, J. E., and L. I. Ferraro. 1986. Hebras fungics como principal componente de nidos de boyero en el ne Argentino. Facena 6: 5-16. Received 31 October 1995 Accepted 30 January 1996 News and Comment | The Ottawa Field-Naturalists’ Club 1995 Awards The annual recognition ceremony for outstanding contributions toward Club goals in 1995 was held at The Ottawa Field-Naturalists’ Club wine and cheese party on the evening of 26 April 1996 at the Unitarian Church Hall, 30 Cleary Street, Ottawa. 1995 Member of the Year Award: Bob Bracken The Member of the Year Award is given to the member judged to have contributed most to the Club in the previous year. The recipient this year is Bob Bracken in recogni- tion of his overall generosity and versatility and his specially active and productive 1995. Bob has generously shared his knowledge and enthusiasm for the large variety of natural life repre- sented in the flora and fauna of the National Capital Honorary membership remained at its maximum 0. 25 to the end of 1995. An account by Cendrine Huemer of this Soirée is given in Trail & Landscape 30(3): 89-91 July-September 1996. The tlw awards were presented: | Region with members of the Ottawa Field- Naturalists’ Club for over fifteen years. He has led: numerous club walks covering such diverse groups as ferns, butterflies, molluscs, fish, amphibians, and: birds. He has served on the Rare Birds Sub- Committee, been a co-leader on eight spring birding; trips to Presqu ile, as well three to observe different’ aspects of natural history at Point Pelee in 1993, 1994, and 1995. 1995 Conservation Award to a Member: Jeff Harrison The annual Ottawa Field-Naturalists’ Club Conservation Award is given to a member in recog- nition of a recent outstanding contribution in the cause of conservation. “Conservation” may be defined as “the wise use of natural resources”, and while we often think of conservation as involving the protection of sensitive areas against inappropriate development, it can also involve the management of land for the benefit o wildlife. Accordingly, this year’s Conservation Award is presented to Jeff Harrison for his work as Chairman of the Fletcher Wildlife Garden Committee and for his co-authorship, with his wife Victoria, of “The Urban Naturalist” column in The Ottawa Citizen. The Fletcher Wildlife Garden was created for two main purposes: (1) to establish a wildlife oasis near the heart of Ottawa, and (2) to demonstrate to landowners how they can plant and arrange their property for the benefit of wildlife in the city. Through “The Urban Naturalist”, Jeff, along with Victoria, have helped to nurture a love of wild, liv- ing things in readers of The Citizen, and have given many suggestions about things that individuals can do to help preserve nature in urban settings. Environmentalists have often recommended that lovers of nature should “think globally and act local- ly” when fighting for the wise use of natural resources. Jeff Harrison has certainly taken this advice to heart. 1995 Conservation Award to a Non-member: Kit Chubb Kit Chubb is one of those rare persons who take on a job and fulfil all requirements beyond all expec- tations. That job, in her own words, is “to improve the lot of wild birds”. After a career in nursing, Kit Chubb returned to her first love, the care and protection of wild crea- tures. In 1977, after seeing the desperate need for facilities for dealing with injured birds, she and her husband, Robin Chubb, established the Avian Care and Research Centre in Verona, Ontario, for all birds and especially raptors, herons, and loons. The Centre now receives 400-500 birds a year with almost 200 species represented over the period of its existence. Not content to simply nurse and care and return birds to both health and home, Kit Chubb began a laborious process of observing and record- ing the thousands of birds treated over almost two decades. What makes her work truly outstanding is 544 | | t | a 996 her devotion to sharing her observations, case his- tories and wisdom with veterinarians, other researchers, and the public. Her investigations into bird mortality have proved beyond doubt that lead shot was responsible for the poisoning of countless loons and waterfowl. Thanks to her efforts, great inroads are being made ' into banning lead shot entirely. Not to be deterred { | NEWS AND COMMENT 545 by the little time left with the care of 17 aviaries, she is also a talented writer and artist who uses her skills to rally others to the cause. This award is given in recognition of her untiring efforts to restore the health of hundreds of owls, hawks, eagles and other raptors, both to Kit Chubb and to the Avian Care and Research Centre. _/ 1995 Anne Hanes Natural History Award: Marilyn Light | 7 | =~ ay a ? sr a The Anne Hanes Natural History Award is given in recognition of an outstanding contribution to our knowledge, understanding and appreciation of the natural history of the Ottawa Valley. The 1995 Anne Hanes Natural History Award is presented to Marilyn Light for her eleven-year study of the population dynamics of orchids in Gatineau Park. In the process of research on over 1000 plants of Epipactis helleborine she has dis- covered and named one unique albino form of Platanthera and has determined factors involved in germinable seed yield in the Large Yellow Lady’s Slipper, Cypripedium calceolus; the Lesser Purple Fringed Orchid, Platanthera psycodes, and the helleborine. She recently presented a paper on her work on orchid population biology to the Linnean 1995 President’s Prize: William D. (Dave) Smythe Each year the President selects an active member of the Club whose contribution merits special recog- nition. This year’s selection is Dave Smythe, a rela- tive newcomer to The Ottawa Field-Naturalists’ Club, who has quickly established himself as a very useful member. Dave and his wife Verna joined the Club in 1991. In 1992 they became members of the mailing team for Trail & Landscape. In 1993 Dave joined the Membership Committee and also became a member of Council. In 1994 he began reporting on the Annual Business Meeting for Trail & Landscape. The position of Recording Secretary came open in 1995 and Dave accepted my invitation to add this task to his list of duties. Today he continues to carry all of the responsibilities described above. We have yet to find a limit to Dave’s capacity to serve the Club. Candidates for the President’s Prize must be accomplished and effective. Dave has impressed us in every task he has undertaken. He takes an inde- pendent approach in analyzing problem situations, carefully thinking through the implications, then offering advice if he thinks he has something to con- tribute His writing is concise, yet captures the essence of the subject. Could these qualities be relat- ed to his engineering degree and military training? Society of London. She has also directed an educa- tional video for the Canadian Orchid Congress on the late Joseph Purdon and his conservation efforts with the Showy Ladyslippers on his property in Lanark County. Due primarily to her interest in, and research of, Ottawa Valley flora she is becoming well known worldwide. In addition to her contributions in Trail & Landscape, she is a member of the Garden Writers of America and is a regular contributor to the Orchid Review of the Royal Horticultural Society in England, and Orchids Australia, a publication of the Australian Orchid Foundation. BILL ARTHURS Chairperson, and the Awards Committee of The Ottawa Field-Naturalists’ Club A key function of the Membership Committee is to maintain an up-to-date list of members, particu- larly for mailing Trail & Landscape. As anyone who has maintained a mailing list will testify, this is no simple task; accuracy and timing are critical. For the purpose the Club acquired a personal com- puter. Although he had no experience with person- al computers, Dave undertook to maintain the offi- cial list on this machine. With some good coaching and a computer course, he now competently main- tains the list, prints copies and labels, etc. Refer- ences to computer hardware, software packages, and modems to link with other computers by tele- phone are creeping into Dave’s conversations. The Recording Secretary’s job is onerous in that one must understand the context of each topic dis- cussed, record the essence of each meeting and get the minutes out well before the next meeting. These minutes become the focus for each meeting of the Council. He also is responsible for publish- ing the annual report and administrative details associated with official meetings. Dave makes the job appear easy. As a member of the Council, I have appreciated his thoughtful observations, when I served as President. When not working for the Club, Dave is watching birds and here his characteristic competence again 546 surfaces. His ability to spot and identify birds makes him a welcome member on any birding outing. This interest in birding complements another interest: travel and foreign birds. Before retiring from the Atomic Energy Control Board, Dave had travelled extensively in Europe. Errata and Addenda to A Life with Birds 1875-1947 In “A Life with Birds: Percy A. Taverner, Canadian Ornithologist, 1875-1947” by John L. Cranmer-Byng Canadian Field Naturalist 110(1): 1- 154, due to editorial oversight at late proof stage, the page numbers on the outside back cover contents list for the Selected bibliography, Appendices 1, 2, and 3, are in error and should read 197, 199, 201, and 202; also on page | the reference Charleston AOU meeting photograph in 1937 should be page 179 not 183. Similarly, a few errors, generally by one page, appear in the index. Flagrantly obvious editorial slips include the final entry to the list of illustrations on the inside back cover where Leonard “Street” should be Avenue, and in the photo caption on page 195 where “Hayes” should read Hoyes. Readers wishing to correspond with John L. Cranmer-Byng should note that as of 22 July 1996 his address will be 40 Baif Boulevard, Apartment 303, Richmond Hill, Ontario, Canada L4C 5M9. Rana-Saura: Amphibian Follow-up Project — Rana-Saura Volume 3, number 1, June 1996, the newsletter of the Atlas of Amphibians and Quebec project, contains an updated summary of 1995 survey results. To date of publication 2049 observations col- lected by 277 volunteers had been added to the Quebec Data Bank with more than 100 from 1995 still to be added. A discussion of the most interesting new records and the relative abundance of observations of most observed least observed species is given. Also available is The Atlas of Amphibians and Reptiles of Quebec (reviewed in The Canadian THE CANADIAN FIELD-NATURALIST : Percy A. Taverner, Canadian Ornithologist, i } Vol. 110) This experience introduced him to foreign cultures and gourmet foods, which have remained a continu- | ing interest. Dave and Verna have two sons. FRANK POPE President [1995] of The Ottawa Field-Naturalists’ Club A comprehensive resume of bird study at the Geological Survey of Canada/National Museum of Canada/Canadian Museum of Nature, here particu- larly recommended for its sections on the post- Taverner period as a fitting epilogue to the Cranmer-Byng biography, has appeared as “Ornithology at Canada’s National Museum” by | Henri Ouellet, pages 303-322 in “Contributions to the History of North American Ornithology” Edited by William E. Davis, Jr., and Jerome A. Jackson. 1995. Memoirs of the Nuttall Ornithological Club, Number 12, vii + 501 pages, 108: figures: [Available from the Nuttall Ornithological Club, c/o Museum of Comparative Zoology, Harvard, University, Cambridge, Massachusetts, USA 02138.] FRANCIS R. COOK Atlas of Amphibians and Reptiles of Quebec Field-Naturalist 109(4): 493, 1995) previously pub- lished in french, but, as of the end of June, available in English at $12 a copy plus postage and handling charges of $4.00 for orders up to $24.00, $6.00 for orders up to $120.00 and $8.00 for orders over $120.00 (charges in U.S. funds for U.S. orders). Specify if French or English edition desired. Order from: St. Lawrence Valley Natural History Society 21 125 Chemin Ste.-Marie, Ste-Anne-de-Bellevue, Quebec H9X 3L2 * Phone: 514-457-9449. The Frog Monitor and DAPCAN IV Proceedings The Frog Monitor Volume 1, Number 1, is dated March 1996. This newsletter contains information on the monitoring project for Manitoba amphibians, and lists the names of 19 participants who have monitored for all three seasons of the project, and 17 who have monitored for two. Project growth is demonstrated by the 93 monitoring instruction kits sent out this past season, and the 124 new volun- teers. A capsule survey of the results from the past three years gives which species are observed most frequently and some tentative conclusions on abun- dance trends in some. There is a brief outline of some do’s and dont’s of monitoring technique, a call for questions and a notice of a identification work- shop held by the Manitoba Field Herpetologists 18 April 1996 at the Manitoba Museum. Also available is DAPCAN IV, the Fourth Annual Meeting of the Task Force on Declining Amphibian 1996 Populations in Canada Proceedings held at the Manitoba Museum of Man and Nature October 1-3, 1994 and cosponsored by the Manitoba Department of Natural Resources; Environment Canada, Canadian Wildlife Service, Prairie and Northern Region; and Ducks Unlimited’s Institute for Wet- land and Waterfowl Research. The proceedings include reports internationally (Ron Heyer) and regionally for eastern Canada (Donald F. McAlpine) and western Canada (Stan A. Orchard) as well an assessment by the Historical Database Committee (Wayne F. Weller, Michael J. Oldham, Frederick W. Schueler, and Martyn E. Obbard. A keynote address by Henry W. Wilbur discusses Amphibian Population Biology. Research Reports are presented by David M. Green (Fowler’s Toad)[abstract only], Ronald J. Brooks and Leonard J. Shirose (Bull, Green and Mink frogs), Stephen J. Hecnar (Regional dynamics of amphibian pond communi- ties in southwestern Ontario), Christine Bishop and Karen E. Pettit (Monitoring in Ontario), Peter Taylor (diurnal and seasonal calling patterns of some Manitoba species), Joel Bonin (climatic and landscape changes vs. population declines in Quebec), Joel Bonin, Martin Ouellet, Jean Rodrigue, Jean-Luc Des Granges, Tim R. Sharpel, and Leslie A. Lowcock (measuring health of frogs in agricultural habitats subject to pesticides in Quebec) [abstract only], Kathryn A. Kelsey (effects of logging on stream-breeding amphibian popula- Canadian Association of Herpetologists Bulletin The Canadian Association of Herpetologists Bulletin volume 10, number 1, Spring 1996 is 10 pages and contains notes on “Space Herpetology” by Richard Wassersug and the latest report on “her- petological happenings” from Redpath Museum by Production Editor David M. Green. Also included is a report on the Annual Meeting of the Society for Northwestern Vertebrate Biology by Sheri Watson and on the Fifth International Congress of Vertebrate Morphology by J. M. V. Rayner. Miscellany includes notes on Volunteer Amphibian Monitoring Programs - updates and on Information Bulletins, both by Editor Anthony Russell, and notice that “COSEWIC designates the Wood Turtle a Vulnerable Species in Canada by David M. Green. Abstracts of three 1996 herpetological theses, all from the University of Guelph, are given: Heather Leigh Passmore (M.Sc.) Geographic variation in NEWS AND COMMENT 547 tions in Washington) [abstract only], Jan J. Roth and Reid Westland (ultraviolet and hydrothermal radiation -northwestern Colorado and Kenora, Ontario), Martin Ouellet, Joel Bonin, Jean Rodrique, and Jean-Luc Des Granges (disease investigation, pathological findings and impact on amphibian populations in southern Quebec), Michael Lepage, Rehaume Courtois, Claude Daigle, and Sylvie Daigle (abundance of anurans in Quebec based on volunteer surveys of mating calls). Two posters were presented; one by Christine Bishop, John Struger, and Karen E. Pettit (environmental fate and effects of organophosporus insecticides on amphibians in Ontario) the other by James Duncan, Errol Bredin, Gavin Hanke, Ron Larche, William Preston, Dough Ross, Carol Scott, and Ken Stewart (estimated status of Manitoba amphibians based on criteria used by the Nature Conservancy’s Conservation Data Centre Network. The publication concludes with an account of a workshop on meth- ods and objectives of sampling amphibian popula- tions by Ronald J. Brooks, the discussion and busi- ness meeting as recorded by Carolyn Seburn, and a list of registrants with addresses. WILLIAM B. PRESTON Manitoba Museum of Man and Nature, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2. *Phone (204) 956-2830 *Fax (204) 942-3679 “e-mail: bpreston@mbnet.mb.ca. — life-history in four populations of Chelydra ser- pentina Linneaus; Jacqueline Danielle Litzgus (M.Sc.) Life-history and demography of a northern population of spotted turtles, Clemmys guttata; Megan Leslie Harris (M.Sc.) A characterization of adult and premetamorphic leopard frogs (Rana pipi- ens) and green frogs (Rana clamitans) in wetland influenced by agricultural activities. (Supervisor of the first two, Ronald J. Brooks; of the third, James P. Bogart). Membership in the Canadian Association of Herpetologists is $10.00 (regular) and $5.00 (stu- dent). Application should be made to: PATRICK T. GREGORY Treasurer, Canadian Association of Herpetologists, Department of Biology, University of Victoria, P.O. Box 3020, Victoria, British Columbia V8W 3N5 Notice of the 118th Annual Business Meeting of The Ottawa Field-Naturalists’ Club The 118th Annual Business Meeting of the Ottawa Field-Naturalists’ Club will be held in the auditorium of the Victoria Memorial Museum Building, McLeod and Metcalfe streets, Ottawa, on Tuesday 14 January 1997 at 19:30 h. : DAVE SMYTHE Recording Secretary 548 THE CANADIAN FIELD-NATURALIST Vol. 110 Call for Nominations: The Ottawa Field-Naturalists’ Club 1997 Council Candidates for Council may be nominated by any member of The Ottawa Field-Naturalists’ Club. Nominations require the signature of the nominator and a statement of willingness to serve in the posi- tion for which nominated by the nominee. Some rel- evant background information on the nominee should also be provided. Deadline for nominations 1s 15 November 1996. FRANK POPE Chair, Nominating Committee Call for Nominations: The Ottawa Field-Naturalists’ Club 1996 Awards Nominations are requested from members of The Ottawa Field-Naturalists’ Club for the following: Honorary Membership, Member of the Year, George McGee Service Award Citation, Conservation, and the Anne Hanes Natural History Award. Descriptions of these awards appeared in The Canadian Field-Naturalist 96(3): 367 (1982). The Service Award was renamed the George McGee Service Award for 1993 presentations [see The Canadian Field-Naturalist 108(2): 243-244 (1994)]. With the exception of nominations for Honorary Member, all nominees must be Club members in good standing. Deadline for nominations 1s | December 1996. BILL ARTHURS Chair, Awards Committee Book-Review Editor’s Report for The Canadian Field-Naturalist Volume 109 (1995) Time was something missing from the new titles and reviews in Volume 109. The book-review editor managed to spend about half of his in Africa and thus was not very good at keeping up with all my correspondence and new books. My reviewers have stepped in to save me. I pass my thanks for the patience and understanding they have. I must also echo my annual plea to keep contacting me when a book of interest 1s published or listed in our New Titles. I have over 100 faithful reviewers and many specialist or occasional participants. We count on you all, but the time to keep up with each one of you is just not there. Book Review Statistics for Volume 109: The Book-review statistics for Volume 109 are summarized below. The number of New Titles listed was much smaller than usual. Due to special issues and my own absence, there were only New Titles in the second and fourth issues of 1995. Also due to my lack of time, there were fewer requests for reviews or for books to review. The reviewers and publishers both came through to keep the numbers of books available and the number of reviews published up to our usual numbers. I am always looking for new reviewers and hoping some of the past ones will come back. If you are interested please contact me. Books listed in the New Titles as available will be sent to the first per- son requesting them. New, appropriate titles can be requested from the publishers and will usually be supplied. Book reviews should be | to 2 pages, dou- ble-spaced in length and informative to The Canadian Field-Naturalist readers. Reviewers are Issue 1 Dy 3 4 Total Reviews Requested 8 1] 14 0 33 Books to Reviewer 9 20 13 22 64 Reviews Completed 0) 58 liv 10 85 Reviews Published 16 af 2 43 68 Books Requested 17 6 i 0 30 Books Recieved 6 42 8 27 83 New Titles Listed 0 66 0 120 186 expected to submit reviews within three months of receiving the book. Reviewers get to keep the com- plimentary book supplied. ; WILSON EEDY Book-review Editor, R.R. 1, Moffat, Ontario LOP 1JO Fax: 1-905-333-0798 e-mail: edith @ wat.hookup.net (home) or we @ geomatics.com (work) 1996 Recovery: An Endangered Species Newsletter This is an eight-page news and background leaflet issued by the Canadian Wildlife Service, Environment Canada dated "Spring 1996" but with no volume or issue number included, and no actual address in the issue itself. However, my copy came with a covering note from Linda Maltby, Chief, Endangered Species Conservation Division, Canadian Wildlife Service, Environment Canada, Ottawa, Ontario KIA OH3, dated 10 July 1996. Copies are presumably available through that office. This issue includes an article on attempts to start a new wintering grounds for Whooping Crane: "Flying down to ... New Mexico?" (Brian Johns). [The only current wild self-sustaining population is given as 157 birds and it winters in marshes along the Texas coast, a vulnerable habitat due to the volume of petroleum and chemical products carried shied through adjoining waters.| "Working for endangered The Boreal Dip Net Volume 1, Number 2, Summer-Fall 1996, 4 pages, is the newsletter of the Working Group on Amphibian and Reptile Conservation in Canada [WGARCC] which also includes the IUCN/SSC Task Force on Declining Amphibian Populations in Canada. Articles featured are "Amphibians and the changing atmosphere" (Stan Orchard), "Atmospheric change: themes for discussion" (Andrew Blaustein); "What does climate change mean for North American Amphibians?" (Michael Lannoo); "UV-B and the ecology of northern lakes"; "76th Annual NEWS AND COMMENT 549 species: a provincial perspective" (Arnold H. Boer, New Brunswick Fish and Wildlife Branch); "A red- horse of a different colour" (Alain Branchaud and Rejean Fortin) [The Copper Redhorse, restricted to one or more rivers in southwestern Quebec. |]; Wildlife on the Hill: Parliamentary committee con- siders conservation of endangered species" (Jim Foley); Commentary: Endangered species legislation and land claims: challenge or opportunity?" (John Bailey): Protecting critical habitat for blue racers on Pelee Island (Ben Porchuk and Ron Brooks); an announcement of the RENEW report on the swift fox; a retirement tribute to Bruce Johnson; and notice of the move of Dale Hjertaas to an unspeci- fied position with the Saskatchewan government behind the "front" lines of conservation". FRANCIS R. COOK Meeting of the Society for NW Vertebrate Zoology - A report" (Larry Powell); "Manitoba amphibian monitoring: ‘Wildly successful'" ; New Book: Okoboji Wetlands: A Lesson in Natural History by Michael J. Lannoo, University of Iowa Press, Iowa City. This newsletter bears no address but presum- ably can be obtained from Stan A. Orchard, National Co-ordinator for Canada, WGARCC, Royal British Columbia Museum, Victoria, British Columbia V8V 1X4. FRANCIS R. COOK Great Lakes Fact Sheet: Amphibians and Reptiles in Great Lakes Wetlands: Threats and Conservation This attractive 12-page leaflet, distributed in early July, is published by Environment Canada and authored by Leonard Shirose, Christine Bishop, and Andree Gendron. It focuses on the current threats and conservation initiatives for amphibians and rep- tiles in Great Lakes wetlands and with colour pho- tographs, diagrams, charts, maps, and graphs. It leads off with the example of the Spotted Salamander, then defines a wetland and discusses its value (habitat for wildlife, water purification, food control, recreational activities). It continues with threats (including biological: e.g. Common Carp, Purple Loosestrife); individual species of amphib- ians and reptiles (Blanchard's Cricket Frog, Eastern Spiny Softshell Turtle, Snapping Turtle, Mudpuppy) and "What You Can Do" ( Backyard Surveys, Amphibian Road Call Counts, Marsh Monitoring of the Great Lakes, Amphibian and Reptile Survey). It gives examples of wetland rehabilitation projects in progress (Black Ash Creek pond creation, Cootes Paradise, Oshawa Second Marsh). The newsletter concludes with an outline of the Metro Toronto Zoo's "Adopt-A-Pond" project, Addresses for those wishing to get further involved and a short list of Further Reading. Other fact sheets in this series include: Contaminants in Herring Gull eggs from the Great Lakes, Bringing the Bald Eagle back to Lake Erie, The Fall and Rise of Osprey Populations in the Great Lakes Basis, The Rise of the Double-Crested Cormorant on the Great Lakes: Winning the War against contaminants. This fact sheet series is ideal for school classes and naturalists groups, 1s straightforward, readable, eye-catching. This and other information on wet- lands and habitat loss in the Great Lakes basin are available from: Environment Canada, 4905 Dufferin Street, Downsview, Ontario M3H 5T4. Also visit Environment Canada's Greenlane on the World Wide Web: http://www.cciw.ca/glimr/intro.html. FRANCIS R. COOK 550 THE CANADIAN FIELD-NATURALIST Vol. 110 Joint Annual Meeting Working Group on Amphibian and Reptile Conservation in Canada and Task Force on Declining Amphibians and Populations in Canada (DAPCAN) This joint meeting was a continuation of the highly successful DAPCAN initive as the 6th Annual meet- ing for that group, combined with the new Working Group on Amphibian and Reptile Conservation which contains DAPCAN but has a broader perspective by including reptiles. It was held 5-7 October 1996 at the University of Calgary, Calgary, Alberta, and was co- sponsored by Environment Canada, Canadian Wildlife Service, Burlington, Ontario. As well as annual regional reports the program included a symposium on conservation of reptile populations in Canada and a Global Biodiversity Volume 6, Number 1, Summer 1996, 48 pages, of this Canadian Museum of Nature journal which bills itself as "An International Forum on the Variety of Life on Earth ... research, conservation and wise use" 1s a special marine issue. An initial editorial by Michael L. Smith, Center for Marine Conservation, Washington, D. C., is starkly titled "The sea is changing, and it matters." Following are six papers, each briefly placed in context by an introductory paragraph by Editor D. E. McAllister, on a variety of marine habitats of high concern: "Exploring deep coral reefs: How much biodiversity are we missing?" (Richard L. Pyle, Bishop Museum, Hawai): "Marine reserves: Necessary tools for biodiversity conserva- tion?" (Jack Sobel, Center for Marine Conservation, Washington, D.C.); "Lessons from deep, hot places" (Verena Tunnicliffe, University of Victoria, British Columbia); "The global policy outlook for marine biodiversity conservation" (John Waugh, IUCN-US, Washington, D.C.); "Conservation of marine biodi- versity in the Caribbean: Regional challenges" (Jose A. Ottenwalder, UN Development Program, Dominican Republic); and "Cuba: An island's approach to marine biodiversity" (Diana Ibarzabal and Juan P. Garcia, Institute of Oceanography, Havana, Cuba). This issue's "portrait of biodiversity" workshop on herpetological atlassing. The keynote address this year was given by Dr. R. Bruce Bury, U.S. National Biological Service, Corvallis, Oregon "Amphibian Conservation in Western North America: Progress, Pitfalls and Perspectives". Further information may be obtained from Stan A. Orchard, Chairman, Working Group on Amphibian and Reptile Conservation in Canada, 1745 Bank Street, Victoria, British Columbia V8R 4V7 * phone/fax 604-595-7556 * e-mail: sorchard@ islandnet.com is the sponge Hippospongia lachne. The issue con- cludes with News (Biodiversity, Cyberdiversity, Biodiversity meetings) and Reviews (Book and peri- odical niche). The last item in the latter, the book Biological Systematics: The state of the Art [by A. Minelli, published by Chapman and Hall, 1993] reviewed by Alison M. Murray and Brian W. Coad, Canadian Museum of Nature, concludes with a pithi- ly upbeat epilogue, painfully fitting for all aspects of biodiversity study and conservation but particularly so for the fundamental organizer of this knowledge, pure systematics: "Problems of funding and lack of resources (especial- ly people) plague all areas of systematics and institutions throughout the world. The state of the art may appear to be in a shambles, but progress, while slow, is being made ...." Persons wishing to subscribe, place advertise- ments, or support the publication financially should contact Global Biodiversity Business Manager Dawn Arnold, or Subscription Manager Susan Swan, Canadian Museum of Nature, P. O. Box 3433, Station D, Ottawa, Ontario, Canada KOG 1RO; * Phone (613) 993-5908; * Fax: (613), 990-03i1:8; * e-mail . FRANCIS R. COOK Adopt a Black Rat Snake Program, Charleston Lake Provincial Park, Ontario The Black Rat Snake, Elaphe obsoleta, 1s not only one of Canada's largest snakes (attaining at least seven feet total length here and perhaps as long as nine feet in the United States, at least in the past), but also one of the more restricted in range in this country. In Canada, it occurs in southwestern Ontario, where it is now rare and patchily distributed in surviving islands of woodland, primarily in Norfolk County, and in a disjunct area in eastern Ontario in the "Frontenac Axis", the rocky region lying largely south of Smith Falls to east of Kingston. Within this it can be locally relatively abundant. It has been intensively studied by Dr. Patrick Weatherhead of Carleton University, Ottawa, and his students at the Queen's University Biological Station at Chaffeys Locks since the early 1980s through mark-release and radio transmitter implant monitoring. Additional, later, studies have been on other protected populations in the region at St. Lawrence Islands National Park, Murphys Point Provincial Park, and Charleston Lake Provincial Park. In the latter park, the Adopt-a-Black-Rat- 1996 Snake program has been initiated by the support group, the Friends of Charleston Lake Park. $15 sponsors an identification microchip implant for a Black Rat Snake. Donors are sent their snake's data when it is marked and will be notified each year it is recaptured. As well, they receive an annual report on the project as a whole, whether or not their snake has been found that year. Initiated in 1995, the program has been received enthusiastically by park visitors, local residents, and schools. To the end of June, 1996, 34 snakes had been implanted and six recap- tured. Monitoring data will not only generate data on individual ranges, activity areas, growth, mortality NEws AND COMMENT 551 rates, egglaying areas, overwintering den use pat- terns, and many other details of snake life and habi- tat use, but also, eventually, failitate detection of any changes in these over extended time. Accumulated data from many snakes can be used to develop esti- mates of population size and density as well as for further public education programs. For more information contact Mike Ogilvie, Senior Natural Heritage Education Leader, Charleston Lake Provincial Park, RR 4, Lansdowne, Ontario KOE 1L0. FRANCIS R. COOK Sea Wind: Bulletin of Ocean Voice International Volume 10, Number 2, April-June 1996 of Sea Wind contains information on ongoing and upcom- ing environmental initiatives pertaining to the con- servation of the world's oceans under the headings: the Ocean Voice International Update, The Scoop on Shrimp, Independent World Commission on the Oceans, 8th International Coral Reef Symposium in Panama, and World Survey on Status of Coral Reef Fishes and Habitat. It also includes a "Book Nook", and sections of longer "Sea News" and shorter "Short Sea News" items, as well as an "On the Net" section of computer addresses for information of Ocean Voice and contacts of interest. Subscription and information on back issues and other publica- tions can be obtained from: Ocean Voice International, Inc., P.O. box 37026, 3332 McCarthy Road, Ottawa, Ontario, Canada K1V OWO * Telephone: (613) 264-8986 * Fax (613) 264-9204 * e-mail: mcall@superaje.com Froglog: Y'UCN/SSC Declining Amphibians Populations Task Force Number 18, August 1996, of Froglog contains items on the status of Rana latastei in Croatia and Slovenia, Amphibians Declines in Puerto Rico, The Vanishing Amphibians Exhibit (Smithsonian Institution Travelling Exhibition Service), Amphibians and the Changing Atmosphere (a report on the workshop held in Corvalis, Oregon, 24 February 1996), Re-survey of Yosemite Area Shows Collapse of Anuran Fauna (results of a 1992 survey of 38 sites compared with that of one from 1915 through 1919), Mixed Fortunes of the Frog Fauna of New Zealand, The Effect of UV-radiation on Alpine Newts, Third Meeting of The Central Region Working Group (United States: Iowa, Missouri, Illinois, Indiana, and Ohio), Publications of Interest (14 papers and articles appearing in 1995 and 1996) and New DAPTFE International Coordinator (John Wilkinson replaces John Baker). Froglog is available on request from John Wilkinson, Department of Biology, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom ** Telephone: 01908 (44 1908 if ex-UK) 652274 ** Fax: 01908 (44 1908 if ex-UK) 654167 ** e-mail: DAPTF@open.ac.uk. Froglog can also be found on the World Wide Web at the following URL: http://acsinfo.open. ac.uk/info/newsletters/FROGLOG html Errata: In The Canadian Field-Naturalist 110(2): 335 notice of Froglog Numbers 15, 16, and 17, Stephen Corn was NOT, nor intended to be read as the writer of the notice itself but was singled out as a convenient North American contact for further information. Further, he was not, as stated there, Task Force Chair, but was Monitoring Protocols Chair (see Froglog number 15, page 4, upper right column) at the time, but now is Rocky Mountain Working Group Co-Chair. My apology to both Stephen Corn and to readers for these editorial lapsi and the inconvenience they have caused. The cor- rect contact for Task Force information 1s J. Wilkinson. (address above). FRANCIS R. COOK 552 THE CANADIAN FIELD-NATURALIST Vol. 110 1 Newsletter: Ontario Natural Heritage Information Centre Volume 3, Number 1, Spring 1996 published by the Ontario Ministry of Natural Resources, is 12 pages and contains items on: Interesting Plant Records from Northwestern Ontario, NHIC field- work documents New Mollusc Species for Ontario, Freshwater Mussel Surveys, Ontario's Lost Plants, Rare Communities of Ontario: Perched Prairie Fens, and sections on Stewardship (Natural Areas Manual Files, Natural Area Database, International Alvar Conservation Initiative, Great Lakes Biomonitoring Project, Looking ahead: Delhi to Simcoe Prairies, Gap analysis for the conservation of Ontario rarest elements), NHIC Information Products (list of 21 publications since the last Newsletter authored by NHIC staff), Systems (Database Imports, Software Development, Focus on ... Rob Parry) and News and Notes (NIHC's Future Direction, NIHC on the Move, Survey of Southern Ontario Sand Barrens for Tachysphex pechumani, NHIC-Sir Sandford Fleming College GIS Project, NHIC Logo, Results of Annual COSEWIC Meeting, Publications, Thanks to Don Smith, NHIC Staff List). The NHIC Newsletter can be obtained from the Natural Heritage Information Centre, P. O. Box 7000, Peterborough, Ontario, Canada K9J 8M5 * Fax (705) 745-5575 Picoides: Bulletin of the Society of Canadian Ornithologists Volume 9, Number 1, June 1996, contains — A word from the President: New Pathways, New Horizons (David Nettleship) — S.C.O. Annual Meeting 1996, University of New Brunswick, Fredericton, N. B., 23-26 August 1996 — Canadian Landbird Conservation Program (Henri Ouellet) — S.C.O. Column ("Publish or Perish": an Alternative interpretation: A. J. Erskine: an plea to publish data while it still can be coherently retrieved, "the data now in files may be in landfills soon. Publish or they will perish.") — S.C.O. Student Awards (1995 and 1996) — Recent Literature (Ornithology in Ontario reviewed by A. J. Erskine) — In Press (publications in ornithology within the past six months listed by Canadian centres) — Announcements and News Items — Editor's Musings (A. J. Erskine). Picoides is published by The Society of Canadian Ornithologists, c/o Canadian Wildlife Service, Atlantic Region, P. O. Box 1590, Sackville, New Brunswick, Canada EOA 3C0O. Membership in the Canadian Society of Ornithologists is $10.00 and can be applied for from the Membership Secretary, Dr. Nancy Flood, Department of Biological Sciences, University College of the Cariboo, 900 McGill Road (Box 3010), Kamloops, British Columbia V3C 5N3. Wild Animal and Plant Protection and Regulation of International and Interprovincial Trade Act (WAPPRITTA) On 6 June 1996, Canada formally announced proclamation of the Wild Animal and Plant Protection and Regulation of International and Interprovincial Trade Act. The Act had received royal assent on 17 December 1992 but certain "implementation issues" had to be addressed through regulations for which the Act required agreement by provincial and territorial wildlife ministers. This was received in December 1995, and the Act entered into force 14 May 1996 after the Special Committee of Cabinet gave final approval to the first set of regulations. WAPPRITA attempts to provide a new legislative tool to promote tighter controls on illegal trade of wildlife and plants and to prohibit trafficking in endangered species. Regulations under it will help prevent the illegal possession, trade or commercial sale of restricted wildlife items. Offenders will be subject to fines of up to $150,000 for individuals, and up to $300,000 for corporations, not only for the illegal importation of endangered species, but also for the possession of products made from these species. WAPPRITA compliments existing provin- cial and territorial legislation and improves Canada's compliance with international agreements, such as the Convention on International Trade in Endangered Species of Wild Flora and Fauna (CITES). WAPPRIITA also gives national support to provincial and territorial management programs as it will no longer be possible to break the law in one province and escape prosecution by leaving the juris- diction. Both provincial and foreign permits will be recognized, eliminating duplication. The text of "An act respecting the protection of certain species of wild animals and plants and the regulation of international and interprovincial trade in those species" is published in The Statutes of Canada 1992, Chapter 52, Third Session, Thirty- fourth Parliament 40-41 Elizabeth II, 1991-92, Bill C-42, pages 1-15. The full text of the "Regulations" for the act are published as Statutory Instruments 1996 SOR/96-263 in Canada Gazette Part II, 130 (11): 1758-1808; Ottawa, Wednesday, May 29, 1996. A simplified users guide is published as Your 1996 Guide to WAPPRIITA: Information on Canada's law to control trade in wild animals and plants. Environment Canada, Ottawa, 24 pages. Copies of the User's Guide, The Act and Regulations with the schedules, the Act and Regulations without the schedules, CITES Control List (either the full list or sections on only mammals, birds, reptiles and amphibians, plants or fish and Conservation Groups in Canada "A short field guide to conservation groups — Chapter 1: Canada's National Groups" by Cendrine Huemer appears in Trail & Landscape [The Ottawa Field-Naturalists' Club's regional journal] 30(3): 104-113, July-September 1996. Seventeen groups and five coalitions are listed with a capsule summary of their activities and address and phone number SAMPA III The Third International Conference on the Science and Management of Protected Areas (SAMPA III) will be held in Calgary from 12 to 16 May 1997. Conference participants will consider the linkages between protected areas and the manage- ment of whole ecosystems in both terrestrial and marine environments. Topic areas are: — Partnerships Linking Protected Areas and Working Landscapes — Linking Mountain Landscape Corridors — Science and Values in Decision-making — First Nations Approaches to Protected Areas — Role of Biosphere Reserves in Linking Protected Areas and Working Landscapes — Linking Ecological Monitoring in Working Landscapes and Protected Areas — Greater Ecosystem Concepts — Cross Boundary Issues: Air, Water, Wildlife — Education and the Development of Public Awareness and Support — Defining and Measuring Biodiversity — Biodiversity in Protected Areas — Biodiversity in Working Landscapes — Maintaining Ecological Integrity and Biodiversity — Managing Natural Resources While Protecting Biodiversity — Applications in Landscape Ecology The Call for Papers is out, and abstracts for papers, posters, workshops, panels or exhibits are NEws AND COMMENT 553 invertebrates), and Regulatory Impact Analysis Statement are available from: Publications, Canadian Wildlife Service, Ottawa, Ontario KIA 0H3 * Telephone: (819) 997-1095 * Fax: (819) 953-6283 Information about the Act and Regulations is also available at Environment Canada's home page on the World Wide Web. The address is http://www.ec.gc.ca (with only three omissions). The Society of Canadian Ornithologists is included but the Canadian Association of Herpetologists is omitted as are other professional societies such as the Canadian Society of Zoologists and the Entomological Society of Canada. FRANCIS R. COOK due 17 January 1997. Organizers coach authors to write the abstract to spark the readers’ interest, and provide whatever background is required to under- stand how you reached your conclusions and why they are important. Single-spaced, typewritten abstracts must fit within a box 17 x 9cm(6.5 x 3.5 in), including title of the paper and name of the author(s). Contact the Conference Secretariat for more information. Twice — in Halifax in 1994 and in Wolfville in 1991 — the conference served as a forum for pre- senting and discussing current perspectives on the role of science in managing protected areas and the role of protected areas in the conduct, support and promotion of scientific research. A special marine symposium will again be part of the conference, and pre- and post-conference tours will highlight marine and terrestrial protected areas. In addition, you will soon be able to visit the SAMPA III Webpage — look for a link to us from the Banff or George Wright Society websites. For a link to SAMPA III check: http://www. worldweb.com/ParksCanada-Banff or http://www. portup.com/~gws/home.html. All communications should be sent to Patricia Benson, SAMPA III Conference Secretariat, #552, 220 4th Avenue SE, Calgary, Alberta, Canada T2G 4X3. Tel: (402) 292- 4519, fax: (403) 292-4404, e-mail: sampa3 @ pch.ge.ca. A Tribute To Claude Eugene Garton, 1907-1996 JOAN HEBDEN 2808 Isabella Street, Apartment 1, Thunder Bay, Ontario P7E 5E7 Hebden, Joan. 1996. A tribute to Claude Eugene Garton, 1907-1996. Canadian Field-Naturalist 110(3): 554-557. To see the world in a grain of sand And heaven in a wild flower Hold infinity in the palm of your hand And eternity in an hour. William Blake After several years of confinement due to illness, Claude E. Garton was released by death on | January 1996. His memorial service held a week later was attended by a wide cross-section of the people of Thunder Bay and beyond, who gathered to share memories and rejoice in the long life of a remarkable man. Claude was born on 26 February 1907, on a dairy farm near Aylmer, Ontario, the son of Joseph E. Garton of London, Ontario, and Edna M. Hoffman of Detroit. His three sisters, Mary, Dorothy, and Joan, all predeceased him. He is lovingly remem- bered by his daughter, Lorraine; his grandchildren, Ben and Sarah Martin; Lorraine’s husband, Gordon Morton; and Sarah and Ben’s father, Bill Martin; Bill’s sister Amy Hubert; and nephews Wayne Crawford, Eddie Joe Rothgarber and Bob Rothgarber and their families. He was especially close to his niece, Barbara Boyd, and her children, Bethany and Gordon, who live here in Thunder Bay. Growing up on a farm under the migration flyway, and near Lake Erie, may well have influenced the pattern his life was to follow — one of passionate interest in the natural world. He used to say he began botanizing as a small child. After attending high school in Aylmer and St. Thomas, he went on to graduate from London Normal School in 1926, then taught for two years at S.S. #5 Southwold, Essex County. But, young and ambitious, or curious about the northern part of the province, he accepted a teaching position with the Port Arthur Board of Education, working at St. James Street, Pine Street, and Current River schools, and was soon promoted to principalship, at that time a position requiring considerable dignity and authority. He must have liked Current River school very much, for he stayed there for most of his teaching career, leaving from 1942-1946 only to serve in the Canadian Army in the Personnel Division. He married a Port Arthur girl, Evelyn MacGoldrick, bought a house near Current River school, and became an enthusiastic member of the community in which he taught. It was not a fashion- able part of the city, working class for the most part, but Claude was no snob. He liked the area close to the edge of the city with the “bush” looming on the hills nearby. Like many other ambitious young teachers at the Lakehead, Claude took extra-mural courses from Queen’s University and attended summer school there, earning a degree in Chemistry and Biology in 1942. His intense interest in taxonomy dates from those studies, although he and his friend Lloyd Slichter had started serious collecting as early as 1933. Claude was part of a lively group of friends, most- ly teachers, who enjoyed the out-doors both in sum- mer and in winter. They built a small ski jump on one of the golf courses. With more enthusiasm than skill, Claude tried that jump and knocked himself out! But he returned the following week and mas- tered it. One winter when the freeze-up came well before the snow, he was part of a group which skated the fourteen miles over the harbour ice to the Welcome Islands and back, the same day. Any time free from summer school or teaching he spent roam- ing the wild country beyond the city limits, collect- ing plants in season. By now his circle of friends extended beyond fellow teachers to others concerned with birds and plants. Led by Col. L. S. Dear, an early local naturalist who collected nests, legally, of course, he and several other like-minded people founded the Thunder Bay Field Naturalists in 1933. Except for the war years when energy was needed elsewhere, that club continued to flourish, largely because of the dedication of people like Claude. In those days, conservation was rarely heard of, and birding was regarded with suspicion. When Col. Dear became frail, a triumvirate of friendly rivals carried on. Dr. A. E. Allen, a patholo- gist; Keith Denis, a civil servant: and Claude kept the club going, holding most of the offices in turn, until it became well established and attracted mem- bers who caught their enthusiasm. Claude long out- lived his two old friends, and continued his active membership until he could no longer get out to meet- ings. During those active years he found time to start a Junior Naturalists Club, which, after inevitable ups and downs, is still flourishing under the sponsorship of the Thunder Bay Field Naturalists. 554 1996 HEBDON: TRIBUTE TO CLAUDE EUGENE GARTON 1907-1996 355) Claude E. Garton in the field, Thunder Bay region, Ontario, in retirement. The field trips that were a part of the club’s activ- ities were very popular, and those Claude led always attracted a crowd despite his impatience with late- comers (they were left), and the pace he set. What exciting adventures he led us into! A cedar bog, mosquito infested of course, introduced us to the exquisite Fairy Slipper Orchid, Calypso bulbosa. A long, breathless climb up a heavily forested hill was rewarded by the sight of a magnificent clump of Braun’s Holly fern, Polystichum braunii. A steep, slithery scramble down a gravelly river bank brought us to a pretty waterfall where we actually walked behind the sparkling curtain of water! Claude’s purpose was to show us a rare little fern growing on the wet rocks, but the experience was too exciting for the name of that plant to register — to his annoyance! Then there was the long bush whack into Cavern Lake, where we crawled deep into a strange, lenticular cave to count bats. Those who came on time, and kept up the pace learned a great deal about birds and plants, and the inter-rela- tionships among rocks, soil, climate and life forms. Those who chould not keep up, like two teenagers found sitting on a rock weeping, were quietly res- -cued by more experienced Claude-followers, and brought back to the cars. Evelyn, Claude’s wife, came on some of these outings. A thoughtful woman, she was prepared for emergencies. On one outing to the Slate River to examine the strange concretions that formed in its sedimentary banks, one of the youngsters fell in and got thoroughly soaked. It was Evelyn who produced an extra pair of dry trousers, Claude’s of course, and the boy soon rejoined the hike having to roll up the cuffs only once. Sadly, Evelyn died in 1956, leaving Claude and their eleven-year old daughter, Lorraine, to manage on their own. Despite their heart-break, they did just that. He was a good plain cook, although his jams and jellies made from wild fruits and berries were gourmet fare. Always well orga- nized, he kept house, worked in their garden and brought up a fine daughter. She credits her father’s early feminist views with encouraging her to pursue her own studies with a sure confidence in her own abilities. Ironically, her academic success surpassed her father’s, in earned degrees at least. But then he really did earn his honorary doctorate. Claude continued to teach and fulfill his duties as principal at Current River until 1966, when he retired from the school system and devoted himself full time to his first love, botany. He had been col- lecting for the Department of Agriculture and for the 556 Canadian Museum of Nature, both in Ottawa, as well as building a personal collection that finally outgrew his space at home. So, he gave the 14 000 specimens to Lakehead University where it formed the nucleus of a growing collection. He became curator of the university’s herbarium for a number of years, serv- ing also as an informal counsellor and mentor to many students who soon discovered him to be a source of help and encouragement. The university authorities named the herbarium after him, and began to make use of his obvious abilities as a teacher. He even gave a course in botany to a small class in Geraldton, a little town on Hwy. 11 about 180 miles east of Thunder Bay. Perhaps he remem- bered his own years of extra mural study, and enjoyed giving the personal help that isolated stu- dents appreciate. His circle of friends now included university people, both staff and students and he moved easily in both groups. Claude held membership in a variety of societies and organizations: American Bryological Society, American Institute of Biological Sciences, Canadian Botanical Association, Ottawa Field-Naturalists’ Club, Claude Garton Home and School Association, Federation of Ontario Naturalists (Former Director), New England Botanical Club, Northwestern Ontario Conservation Association, Ontario Public School Men Teachers’ Federation, Port Arthur Horticultural Society (Former Director), Thunder Bay Field Naturalists (Former President), Thunder Bay Fish & Game Association, and the Torrey Botanical Club. During the 1940s and 1950s, Claude had spent some of his summers as Park Naturalist both at Sibley (now Sleeping Giant) and Quetico provincial parks. He identified plants and birds, led nature walks, and generally stirred up interest in both those fascinating places. He started a little museum at Sibley, despite the lack of adequate facilities, to dis- play some of the park’s special habitats. As always he made new friends through this work and through exchanges with other collectors. The correspon- dence he kept includes letters from far and wide — Northwest Territories, many states in the United States, and even Australia. These letters usually refer to plants, or of course, but they invariably include references to good times: in the field, often express- ing thanks for hospitality given and invitations to visit. Claude did visit some of these correspondents, but only on this continent. His new work at Lakehead University must have brought him joy. The students loved him, one class made up a long song about their trials in the field- the refrain of which ran “Just ask Claude.” And he began to look the part. Freed from the obligation of being and looking like a school principal, he gradu- ally adopted the casual dress of his students, wearing tee shirts in summer and plaid shirts in winter along with jeans or work pants and boots. He stopped THE CANADIAN FIELD-NATURALIST Vol. 110 shaving, whether to save time or to make a state- ment, who knows? Soon the rather dapper teacher figure became the white bearded “character” we knew for his last thirty years. He did keep a suit, but rarely wore it, preferring to add a tie to his plaid shirt for formal occasions. The two rarely blended. I think he enjoyed his new persona especially when he began to get fatter. He loved to tell a story about walking along a hallway in the university one January, and seeing a little girl playing with a new doll just outside of an open classroom door. She was probably the child of a student who could not afford a babysitter. Claude stopped to admire the new doll, and asked the child who had given it to her. She looked up at the kindly, white-bearded, rotund old fellow and replied with a big smile, “Don’t you remember? You did.” During the early years of his retirement he contin- ued to collect, extending his territory eastwards to include the shores and islands of Lake Nipigon, and along the coast line of Lake Superior and into Pukaskwa Park, finding many arctic and alpine species that had survived in those cold zones. In 1971 he became involved in an unusual project near Wawa, a small community on Hwy. 17 about halfway between the Lakehead and the Sault. Because of its location, hitch-hikers often became stranded there, and a camp was set up near an archaeological dig run by Professor Ken Dawson of Lakehead University. The hikers were expected to help with the work in return for food and shelter. Claude, who was collecting in the area, soon had them helping him and inevitably becoming interested in the plant life around them. Of course, he and Ken knew each other from their work at the university. In some ways Northern Ontario is like a small town. In spite of all these activities Claude continued to be active with the Thunder Bay Field Naturalists, not leading field trips but arousing support for his many concerns with habitat preservation. Members lis- tened to his appeals, wrote letters, attended Open Houses, and made formal presentations to support such projects as protecting the Bur Oaks, Quercus macrocarpa, at Stanley, a village west of the city; saving the remaining marshes along the shore within the city limits; and preserving an old growth stand of White Pine, Pinus Strobus L., at Greenwood Lake west of Thunder Bay. His enthusiasm and obvious concern inspired the younger members who carried on the struggle when age finally began to slow the old leader down. Unlike many hard workers, Claude lived to be pub- licly recognized for. his efforts and even to see some of his projects succeed. The school he had worked in and for so long was renamed the Claude E. Garton School and its Home and School Association made him an honorary life member. After his memorial ser- vice, the social hour was held at that school with 1996 lunch served by the Home and School members — a very thoughtful choice. Of course the Thunder Bay Field Naturalists had long since made him an hon- orary life member. After naming their herbarium after him, Lakehead University made him first an Honorary Professor of Biology, then an Honorary Doctor of Science, recognizing that he had been too busy doing science to have time for the usual paper work needed to gain degrees. Perhaps that 1s why he published so little; his quiet retirement devoted to writing turned out to be as active as the rest of his life had been. The Federation of Ontario Naturalists awarded him its Conservation Trophy at their annual meeting held in Thunder Bay in 1978, and even the city named him a “Citizen of Exceptional Achievement” in 1979. And the Ottawa Field Naturalists’ Club made him an honorary member in 1985. He saw “his” collection in the herbarium grow rapidly and exchanges with institutions as far away as Europe became common place. Over forty first records for Ontario were accepted and show an achievement in the study of botany that is unmatched anywhere in the north. His old friends in the Thunder Bay Field Naturalists held a dinner to féte him, and he thoroughly enjoyed the good natured “roast” that followed, as a number of old friends told stories of experiences they had shared with him. Perhaps more important than knowing he was appreciated, he lived to see his projects being carried on. The Bur Oaks still stand, sheltering one of the few patches of Poison Ivy, Rhus toxicodentron L. that grow up here. There are two wetland parks along the city’s shoreline. Both attract many casual visitors as well as bird watchers and children learn- ing about marsh life. It is good to have an uncultivat- ed bit of shoreline where one can contemplate the Sleeping Giant and Welcome Islands with no hydro lines or industrial buildings in the way. A handsome Visitors’ Centre now houses a natural history muse- um at Sleeping Giant Provincial Park, and Claude was able to help cut the ribbon at its opening. Even the pines at Greenwood Lake are protected, at least for the present. Dr. W. Carmean, now retired from active teaching at Lakehead University, has contin- ued the fight to save that ancient stand. But perhaps the best reward for a long busy life was the steady stream of visitors who called during the last few years when he could not get out easily. Very old friends like Clark MacDonald recalled with him their early days as teachers here in what must have seemed an alien land. His niece, Barbara Boyd, stood in for Lorraine, who lives in southern Ontario, and with her two children made sure he was well cared for. HEBDON: TRIBUTE TO CLAUDE EUGENE GARTON 1907—1996 SS) By looking at one of his sheets in the herbarium, one can sense some of the character of the man who prepared it. The plant, elegantly arranged on the paper, still retains much of its colour for it was dried and pressed with patient care. The card at the bottom of the sheet, in neat lettering, identifies it completely and states very precisely exactly where and when it was collected. This 1s true even of the specimens that are not showy; the grasses, lichens, and mosses are all conscientiously given their due. But Claude had a side to his character that many of us never suspected. At his memorial service, after the usual tributes and hymns, Barbara’s two chil- dren, Bethany and Gordon, each read a favourite poem of theirs from a collection their “uncle” had copied out in long hand in an obviously much han- dled note book. The young people had shared those poems with Claude. The busy, active outdoorsman so deeply concerned with the natural world that most of us had known so long, was also a lover of poetry. Acknowledgments I appreciate help given by Clark MacDonald; Dr. W. Carmean; Erika North, present Curator of the Claude E. Garton Herbarium; Dr. J. H. Soper, and Dr. Susan Bryan. Publications Dale, H. M., and C. E. Garton. 1984. The aquatic macro- phyte vegetation of an isolated island adjacent to Lake Nipigon, Ontario: A comparative study after a fifty-six year interval. Canadian Field-Naturalist 98: 444-450. Garton, Claude E. 1984. Checklist — Vascular Plants of Thunder Bay District — revised to 1984. Published by Thunder Bay Field Naturalists. 42 pages. Garton, C. E. 1987. Ice age relics, arctic and alpine plants. Nature Northwest [Thunder Bay Naturalists Club newlsetter] 41(1): 2-7. Garton, Claude E., and Joan Crowe. 1993. Checklist — Vascular Plants of Thunder Bay District — prepared by Lake University, based on Garton 1984, revised to 1993. Published by Thunder Bay Field Naturalists. 51 pages. Soper, J. H., C. E. Garton, and D. R. Given. 1989. Flora of the North Shore of Lake Superior. National Museum of Natural Science, Syllogens Number 63. 61 pages. Manuscript reports included a Botanical Inventory of Pukaskwa National Park 1974-1977 (used as source for a booklet produced by the friends of Pukaskwa Park. Also other manuscript reports were made for the Ontario Ministry of Natural Resources on Sibleys, Caven Lake, and Lake Nipigon. Received 12 March 1996 Accepted 18 March 1996 Book Reviews ZOOLOGY Shells of Atlantic and Gulf Coasts and the West Indies By R. Tucker Abbott and Percy A. Morris. 1995. Peterson Field Guide Series, Houghton Mifflin Company, Boston. 350 pp., illus. U.S. $26.95. For the naturalist interested in collecting and iden- tifying sea shells found along the Atlantic coast of Canada, United States as well as the West Indies, the new book by Abbott and Morris represents an essen- tial tool. It is the most complete one-volume guide on marine molluscs yet available for those regions of the Atlantic. This fourth edition comes some 20 years after the last published edition (1973) and almost 50 years since the first edition was published (1947). Faithful to the fomat of the Peterson Field Guides series, this excellent pocket book contains brief yet accurate and informative text and is sup- ported by good quality pictures. The focus of this new edition is on shallow-water species found along the Eastern coasts of North America and the West Indies, especially from Labrador to Texas. The name, description, as well as distribution of approxi- mately 800 molluscs are included in this book. This fourth edition contains a number of signifi- cant changes. As mentioned by the authors, many deep-sea species that were featured in the third edi- tion — and which were out of reach for most amateur or professional malacologists — have been excluded. Also new to this fourth edition is the inclusion of 115 new fine drawings and 74 new beautiful color plates (in the third edition most shells were repre- sented by black-and-white plates). As a professional malacologist I find these color plates very useful as they give the reader a clearer picture of the shell of The Summer Atlas of North American Birds By J. Price, S. Droege, and A. Price. 1995. Academic Press, San Diego. x + 364 pp. U.S. $29.95. The abundance of plants and animals varies geo- graphically. This can be observed by hiking up a mountain or travelling long distances. Although it may seem trivial, this critical fact is not presently reflected on distribution maps available in standard field guides. Instead, species’ ranges are mapped by connecting extreme known localities and filling the range with one colour, or a few colours in the case of migratory species. Recently, the availability of continent-wide databases and the development of spatial statistics have made it possible to illustrate not only the extent each species. Furthermore, the authors have also insisted in adding the scientific authority to each species name given (author, date), which is, as men- tioned by Abbott, useful as it helps to trace original species descriptions and in recognizing homonyms. As in the previous edition, molluscs in this Peterson book are arranged by Classes, Families, Genera, and Species. The nomenclature, however, has been revised according to the extensive and well-accepted classification found in the review work by Kay and Boss (1989) A Classification of the Living Mollusca, edited by Abbott and Boss. The new book by Abbott and Morris also contains a clear and welcome conservation tone in its intro- duction, which should encourage naturalists sensi- tive to the problem of overcollecting by amateur and professional malacologists around the world. In addition, the book includes a well-described listing of tips for proper collecting of shells, and tips on how to prepare live shells after field collecting, including techniques for easy removal-of animal tis- sue from the shells using hot water baths or microwaves; in short, many good tips for the natu- ralist or keen amateur shell collector. I strongly rec- ommend the new fourth edition Shells of the Atlantic and Gulf Coasts and the West Indies by Abbott and Morris to every naturalist interested in molluscs. ANDRE MARTEL Research Division, Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario, Canada K1P 6P4 of a species’ range but also the variation in abun- dance that exists within range boundaries. For exam- ple, Terry Root has produced relative abundance maps in her Atlas of Wintering North American Birds using data from the Christimas Bird Count. In the Summer Atlas of North American Birds, Jeff Price and his collaborators now present relative abundance maps for the summer distribution of 450 bird species and identifiable forms using the North American Breeding Bird Survey (BBS) database. The authors insist that they have produced a summer atlas rather than a breeding atlas since the BBS methodology is aimed at collecting abundance data, not breeding evidence. 558 1996 The authors state that the original goal of this book was to “help birders find birds”. Each colour- coded map illustrates the relative abundance of a species throughout the range it occupied during the 1985-1991 period. The four abundance classes used are illustrated with contrasting colours and bounded by contour lines obtained from kriging (a spatial interpolation technique). In addition, the authors pro- vide the location of the BBS routes where the high- est abundance was recorded for 531 species and identifiable forms (including 81 species whose maps are not provided). The book also includes a table documenting population trends for the 1966-1993 period (based on BBS data) as part of a chapter on conservation issues. Maps of relative abundance are a major improve- ment compared to previously available maps of bird distribution. The authors carefully selected the BBS routes to be included in the data set and they adjust- ed their criterion for mapping the edge of the distri- butions (0.1 to 0.5 birds per route per year) to account for differences in the detectability of species. No text or drawings were included along with the map of each species. This was a wise deci- sion since the targeted audience (“serious” birders) does not need to see drawings of the bird species, BOOK REVIEWS 559 and a text could do little more than describe patterns obvious on the maps. My main criticism pertains to the decision of the authors to crop the maps eastward and northward. Readers from Newfoundland, Prince Edward Island, and Cape Breton will be dismayed to see that their islands are not included in this atlas. Maps only extend northward to a line stretching from the top of Vancouver Island across the middle of the prairie provinces, to the level of Anticosti Island, on the north shore to the St. Lawrence River. The restriction of maps to portions of the continent densely surveyed by the BBS was probably aimed at maximizing the accu- racy of the interpolation technique used to produce the maps. However, a better compromise would have been to include portions of the continent less well covered by BBS routes and to delimit with a line the areas beyond which contour lines are less accurate. In spite of these criticisms, this book is a gold mine of information, not only for birders, but also for “macroecologists” and biogeographers interested in analyzing the variation in species abundance over large spatial scales. MArRc-ANDRE VILLARD Département de biologie, Université de Moncton, Moncton, Nouveau Brunswick E1A 3E9 A Photographic Guide to North American Raptors By Brian K. Wheeler and William S. Clark. 1995. Academic Press, London and New York. 198pp., illus. Contrary to my previous bias that a painting (e.g., by Roger Tory Peterson) can show the diagnostic features of a bird better than most colour pho- tographs, the 377 superb photographs in this book (46 of Red-tailed Hawks alone, including two strik- ing albinos!), have forced me to change my mind. Raptors perhaps have more confusing variants in plumage than any other group of birds; the permuta- tions and combinations at times confound even the most experienced observer. Inconsistencies in plumage have forced Wheeler and Clark, appropri- ately, to make frequent use of hedging words such as “usually” and “often”. After fifty years in the field, one can still encounter a raptor in a plumage never seen before, and be quite unsure as to species identi- fication. General field guides are forced for reasons of space to omit many raptor plumage presentations. If one already owns a copy of the previous Clark and Wheeler book, A Field Guide to [the] Hawks [of] North America, or of Hawks in Flight, by Pete Dunne, David Sibley, and Clay Sutton, does one require this book in addition? The answer in my opinion is a resounding YES. This book offers, in addition to the photographs, many helpful “pearls” or diagnostic clues. Following extensive coverage of every species of raptor, there are fourteen additional chapters dealing with specific raptor identification problems, including perched juvenile large falcons (gyrs have relatively small heads and massive chests); perched juvenile buteos; perched dark-phase buteos; and differentiation between a dark gyrfalcon and a dark peregrine, and between a dark Rough- legged and a Harlan’s Red-tailed Hawk. Some long- standing identification marks are not as specific as we were once told — a raptor with a dark bib may not be Swainson’s; we are shown three other possibili- ties. Similarly, a dark belly-band is not specific for the Red-tailed Hawk, as other species can have one and some redtails do not. A pale-headed buteo can be one of four species. This first edition is not yet perfect. When eponyms, Krider’s, Fuertes’ and Harlan’s, are used freely within the Red-tailed Hawk, even though Krider’s is a color phase and not a subspecies, it seems insupportable to substitute “prairie” for the long-accepted and well-differentiated “Richardson’s” subspecies of merlin; this action could lead one to suspect an anti-British bias and a snub to the Empire’s finest surgeon-naturalist in all history. The Osprey flight photos are a bit disappointing. Figure RT19 shows the underwing of/a juvenile Redtail when the text describes the upper wing. Twice, statements in the text are not shown in adja- cent photos; in Identification Problem 3, a cross-ref- 560 erence to photo RT26, and in problem 2, a cross-ref- erence to photo RS12, would have been useful. A few statements in the text (e.g., the gray blob across the center of the tail in the female Golden Eagle) are not convincingly shown in the photographs. In Problem 3, there is a different shape of the pale panel on the undersides of the primaries; those of the Red-shouldered Hawk are definitely crescent- shaped, but only one of the four other species has a panel that even approximates the geometrical defini- tion of a trapezoid; perhaps a diagram here would have helped the reader. These are minor quibbles. I recommend that you THE CANADIAN FIELD-NATURALIST Vol. 110 buy two copies, one for home and one for your vehile. The points made by Wheeler and Clark are worth referring to again and again: they emphasize, for example, the value of noting relative leg length and relative length of tail versus wing in a perched hawk. Pointers are given to help in differentiating between sexes and ages of some species. If this vol- ume receives the widespread and careful use it deserves, many fewer misidentifications of raptors should find their way into the literature in future. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 Flammulated, Boreal, and Great Gray Owls in the United States: A Technical Conservation Assessment Edited by Gregory D. Hayward and Jon Verner. 1994. General Technical Report RM-253. U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. 214 pp. 3 maps. This publicaion brings together much new material on three little-known species of owls. There is a sepa- rate, large map for each species, depicting, with great accuracy, known sightings within the United States. A table for each species lists presence or absence in each of the 89 National Forests in the United States: the Flammulated Owl is breeding in 24, present in another 30, and suspected in 17 forests; correspond- ing figures for the Boreal are 11, 30, and 3; for the Great Gray, 13, 29, and 6. How are these species threatened by habitat loss or degradation? The Flammulated Owl is the most southerly of the three in distribution, although its range extends into extreme southern British Columbia. It is migratory (though no distant band recovery has yet been obtained). Spring arrival dates are in late April and early May. It hunts exclusively at night, eating noc- turnal arthropods. Pairs shown high site fidelity. In some forests, it is the commonest bird of prey, with 10 owls per 40 hectares in the White Mountains of Arizona; elsewhere there is rarely more than one owl per 40 hectars. It lays a uniformly small clutch of 2 to 4 eggs. Adult survival is probably more critical than annual nest success. Maximum longevity to date is 8 years, 1 month. In order to survive this little owl “simulates greater size and strength by its brava- do and ventriloquial voice, at the same time behav- ing elusively and inconspicuously by night and day.” The Boreal Owl extends much farther south in the Rocky Mountains than had been appreciated. In the 1970s, new roads allowed access; winter recreation in high mountain lands increased, allowing detection of their small and semi-isolated populations. mature and older forest in the spruce-fir zone provides the highest quality habitat for this obligate cavity nester; breeding populations are not found more than 100 m below the spruce-fir zone. Annual survival is esti- mated at 45 to 80% for adults, but only 20 to 50% for juveniles. Winter and summer ranges are both more than 1000 hectares, and up to 3390 ha. Boreal Owls hunt primarily after dark, and usually attack prey only within 10 m of their hunting perch. They eat small mammals (Microtus, Clethrionomys, and Peromyscus), but add the Northern Pocket Gopher, Thomomys talpoides, to their diet in Idaho. They breed in the first year after hatching. Adult annual survival of 25 radio-marked owls in Idaho was 46%. Their main predator is the pine marten (Martes spp.). Because this owl uses already patchy high-elevation forests, global warming could affect it adversely, by driving it to higher elevations. The Great Gray Owl can be the commonest ow! in Black Spruce-Tamarack forest wetlands. In peak vole years, they reach 1.88 pairs per km? in northern Minnesota and adjacent southern Manitoba. They lay their eggs in old goshawk and raven nests, or on bro- ken-topped snags. Mean date of first egg laid is 5 April in Manitoba but 5 May in Idaho and Wyoming. Egg laying is delayed in years of heavy snow cover. In Manitoba and Minnesota, 81% of nests fledged an average of 2.8 young, but 91% of 32 radio-marked fledglings died before they were one year old. Goshawks and Great Horned Owls are common predators, especially when hares and grouse are scarce. Female owls abandon their young after 3 to 6 weeks, but males feed the young until they are three months old. When vole populations crash, the owls may move up to 700 km, the adult females moving earlier and farther. A female banded as an adult was recaptured 13 years later. This up-to-date compendium is highly recom- mended to anyone_interested in these forest owls. New information from James R. Duncan of Winnipeg will be of special interest to readers of this journal. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N OJ8 1996 BOTANY Rare Vascular Plants in the Canadian Arctic By C.L. McJannet, G. W. Argus, S. Edlund and J. Cayouette. 1993. Syllogeus 72, Canadian Museum of Nature, Ottawa. 79 pp., illus. Free. Rare Vascular Plants in the Northwest Territories By C.L. McJannet, G. W. Argus and W. J. Cody. 1995. Syllogeus 73, Canadian Museum of Nature, Ottawa. 104 pp., illus. Free. The recently-retired President of the Canadian Museum of Nature once described the systematic classification of Canadian birds, mammals, plants, amphibians and reptiles as old-fashioned science and stated that the Museum’s fundamental modern man- date was to be an interactive family entertainment device. Long after such smug drivel has been forgot- ten, however, documents like the two considered here will remain testaments to the real fundamental- ly important contribution of this much-beleaguered institution... the systematic collection and authorita- tive analysis of Canadian biodiversity data. These are the latest (final?) elements in a series of fourteen provincial and territorial rare vascular plant treatments going back to 1977. Each has quickly become the “bible” for determining rare plant status in their respective jurisdictions. Various governmen- tal and private initiatives have expanded into a virtu- al industry of investigation and management of regionally and nationally rare species across Canada. In addition to the protection of innumerable rare plant populations, the Rare Plants of Canada Program data presented in these reports have led to the protection of hundreds of sites and the incidental protection of thousands of “non-target” significant organisms. Both reports are utilitarian, staple-bound publica- tions that are produced with clear, readable type and adequate mapping. Each considers those taxa deter- mined to have a small population within their respective region, viz, known from few records or from a very restricted range. Each taxon receives a brief statement on nomenclature, major literature references, phytogeographic affinity, and habitat. The valuable Nature Conservancy status (Global Rank) is noted for most of the taxa described in the Northwest Territories treatment though not with those in the Canadian Arctic study (why not?). Most BOOK REVIEWS 561 importantly, both treatments include small but work- able range maps for each taxon based on a compre- hensive review of authoritatively examined speci- men records in their respective regions. A discussion of the systems of protected areas and the implica- tions of on-going aboriginal land claims for rare plant protection are worthwhile additions to the Northwest Territories study. As one would expect, peripheral species more widespread in other regions make up the bulk of the 206 Northwest Territories and 236 Canadian Arctic taxa treated. Fully a third of these are from the Boreal Forest region. Very few are nationally rare (only 39 in the Northwest Territories) and even fewer (18) are endemic to either area. The Canadian Arctic study is virtually a subset of the Northwest Territories treatment. It includes much of the political territory of the Northwest Territories as well as the relatively small arctic por- tions of Yukon, Manitoba, Ontario, Quebec, and Newfoundland/Labrador - all floras covered by pre- vious rare plant treatments. While it is useful to have an examination of this natural unit (as it would for the Boreal Forest, the Prairie Parkland, etc.), a stand- alone volume seems somewhat extravagant for data which for the most part are also published else- where. There may have been considerable gains in economy and understanding by including this assess- ment in a concise appendix of the Northwest Territories treatment instead. These two studies constitute a solid, comprehen- sive “where we’re at” statement for the investigation and management of the rare flora of northernmost Canada. The series of which they form a part is a fine tribute to the work of recently retired George Argus who was the heart and soul of the Museum’s Rare Plants of Canada program. Congratulations are due, too, to the Ottawa Field-Naturalists for their financial support for the publication of the Northwest Territories report. Old-fashioned science indeed; this is cutting edge stuff. What can be more critical to the advancement of Canadian biological sciences, after all, than the maintenance of our native biodiversity? DANIEL F. BRUNTON 216 Lincoln Heights Road, Ottawa, Ontario K2B 8A8 562 THE CANADIAN FIELD-NATURALIST Vol. 110 Aquatic and Wetland Vascular Plants of the Northern Great Plains By Gary E. Larson. 1993. General Technical Report RM- 238. U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station, Fort Collins, Colorado. 681 pp., illus. The present book deals with over 500 aquatic and wetland vascular plants that occur in the northern Great Plains of North Dakota, South Dakota, Nebraska, Montana, and Wyoming. Though Canada is not covered, most of the aquatic and wetland plants of the prairies of southern Alberta, Saskatchewan, and Manitoba are included. This reviewer grew up with Muenscher’s 1944 Aquatic plants of the United States and with Fassett’s 1957 A manual of aquatic plants. Though of considerable help, these books are rather complex as they include species from all of the United States. Those in the northern Great plains will find the pre- sent book especially helpful as it simplifies things by dealing only with the species of this region. The book has a workable key to Families. This is followed by good family descriptions and helpful keys to genera and species. Each species has an accompanying description, comment on ecology and distribution, a distribution dot map on a county basis, and often a black-and-white line drawing or color photograph. On the negative side, the book contains much empty space; the maps are rather crude and do not show vegetational zones; the illustrations lack descriptive legends and are not original; and there is no spine title. One may also wonder why the author has included some species, such as Solidago gigan- tea and Lilium philadelphicum, as wetland plants. This treatise will be a valuable addition to the bookshelf of anyone involved with the identification of aquatic and wetland vascular plants in central North America. CHARLES D. BIRD Box 22, Erskine, Alberta TOC 1G0 Le symbiose mycorhizienne: Etats des connaissances Edited by J. A. Fortin, C. Charest, and Y. Piché. 1995. Orbis Publishing, Frelighsburg, Québec. vili + 195 pp. $24.95. During the last 30 years Québec achieved interna- tional stature in mycorrhizal research, largely due to the leadership of one man, J. André Fortin. This book records the proceedings of a colloquium on mycorrhizal symbiosis hosted by Dr. Fortin at the Institute for Research on Plant Biology of the University of Montréal, in conjunction with the con- gress of the French Canadian Association for the Advancement of Sciences in 1994. The book contains ten chapters contributed by 17 authors who are predominantly francophones, and who either work in Québec or have strong profes- sional ties with the Québec group. What they offer is a mixed fare of new and not so new; some of the work (most notably that of L. Simon) has been reported elsewhere, in English. Vesicular-arbuscular mycorrhizal fungi (VAM) are the ubiquitous symbionts of plants, including agricul- tural crop plants. Y. Dalpé reviews the classical con- cepts and explores new approaches to the systematics to gain a better understanding of the relationship of VAM fungi as a basis for their more effective exploitation. The influential results of molecular stud- ies by L. Simon promise to fulfil those expectations. M. St-Arnaud et al., and N. Benhamou review the effects of VAM on susceptibility of host plants to pathogens. H. Vierheilig and Y. Piché address the phenomenon of non-host plants, and mechanisms that regulate their resistance to VAM infection. The status of indigenous VAM fungi under present-day agricul- ture, and what it should be if agriculture were to attain sustainability, are reviewed by C. Hamel, and by C. Plenchette and D. Strullu. While ancestors and wild relatives of most agricul- tural crop plants need VAM to prosper, many of the north-temperate forest trees need ectomycorrhizal fungi to survive. J. Gagnon and C. Langlois report on the progress achieved in large-scale commercial “mycorrhization” of nursery seedlings under the aus- pices of the Québec Ministry of Natural Resources. The lesson here is that heady results obtained in a lab- oratory are seldom duplicated in the real world out- side. We still know very little about forest microbiota. This ignorance is laid bare in L. Duchesne and K. Rigal’s discussion of the impact of forest fires on soil microorganisms. Finally, D. Tagu and F. Martin depart into the exotic realm of eucalyptus, in pursuit of questions that only molecular biology can attempt to answer: what genetic interplay between a plant and the fungi underlies the formation of a novel organ that is ectomycorrhiza? This book well represents Québec “mycorrhizolo- gy”: the approach is unabashedly pragmatic. It is not that the authors are unaware of, or downplay, the con- tribution of basic research. On the contrary, it is the appreciation of the fundamental role of mycorrhizal symbiosis throughout the 400 million-year history of terrestrial flora, that has sustained these researchers’ enthusiasm and faith in the vast potential of manipu- lating mycorrhizal associations for human benefit. K. A. PIROZYNSKI 273 Roger Road, Ottawa, Ontario K1H 5C5 1996 Poisonous Plants of Canada By G. A. Mulligan and D. B. Munro. 1990. Publication 1842/E, Agriculture Canada, Ottawa. 96 pp. Plant Alert By D.A. Metzger. 1990. Royal Ontario Museum, Toronto. 26 pp., illus. (English); 29 pp., illus. (French). There is a wealth of literature on so-called “harm- ful” plants but most of this is pseudo-medical regur- gitation of old, anecdotal, and often questionable information. These two publications, though very dif- ferent in their objectives, share a delightfully differ- ent starting point. Both are compilations of the expe- riences and knowledge of active investigators and/or documented cases and strive to offer authoritative reviews of the toxicity of particular taxa in Canada. Poisonous Plants of Canada isthe most ambitious of the two, addressing plants known to have poi- soned humans or domestic animals in Canada as determined from a careful review of the authoritative literature and from the experience of Agriculture Canada personnel. It is a scholarly effort, clearly intended to be more of a reference work for special- ists than a general public “how-to” guide. A brief nomenclatural note commences each of over 200 entries, followed by a Canadian range description. This is followed by a description of the known toxic- ity of that species to human and/or domestic ani- mals. Importantly, detailed reference to documenta- tion supporting these observations is given with each treatment. Separate appendices of those plants found to be toxic for humans and domestic animals are included as summary lists. Unfortunately, no help is offered in the identification of any of these plants, nor is the reader provided with any direction in obtaining such assistance beyond a few lines on making dried samples. While authoritative and infor- mative, Poisonous Plants of Canada could not be considered to be particularly “user friendly” for any but botanically well-informed readers. Haliburton Flora: BOOK REVIEWS 563 Plant Alert, on the other hand, is extremely user friendly. This small bilingual booklet has a much less ambitious botanical scope than Poisonous Plants of Canada, restricting itself only to the most frequently encountered “problem” species reported to Toronto, Ontario, poison control centers. Plant Alert is also clearly aimed at the general public, with clear, excellent quality photographs of almost all of the two dozen or so plant species treated. Each treatment includes a plain language descrip- tion of the appearance and habitat of the plant in question as well as a discussion of its toxicity. No references are provided so the reader is obliged to go on faith here, although an annotated listing of general literature is included at the end. A brief section on “Plants in Other Guises” — jewelry, ornaments, etc. — is a thoughtful and useful addi- tion. Another very helpful feature is the inclusion of treatments of commonly reported non-toxic plants. What a great idea; many a rushed trip to the childrens’ hospital or poison centre may be avoided by this reassuring section. Had the author expanded her selection to include the major species reported by poison centres in other Ontario communities, however, this helpful booklet could have embraced a province-wide scope and enjoyed a wider provin- cial audience. Both publications successfully address their stated target audience and provide useful, well-pro- duced information. One can easily imagine, howev- er, that a hybrid publication combining the user friendly aspects of the one and the comprehensive, authoritative nature of the other would be a tremen- dous reference work indeed. The basis for a very useful and commercially successful publication is clearly indicated. DANIEL F. BRUNTON 216 Lincoln Heights Road, Ottawa, Ontario K2B 8A8 An Annotated List of the Vascular Plants of the County of Haliburton, Ontario By E.G. Skelton and E. W. Skelton. 1991. Life Sciences Miscellaneous Publications, Royal Ontario Museum, Toronto. 142 pp., illus. In late 1983 Steve Varga published an article in the now-defunct Plant Press describing almost 30 regions of the province with on-going floristic com- pilation projects. The publication of such regional studies is widely recognized as important in devel- oping a comprehensive understanding of floristics at a provincial, national and even international scale. Almost 15 years later, however, only a handful of these regional studies have seen the light of day. That may say more for the sorry state of basic field knowledge in Ontario than any bureaucratic and aca- demic wailing. The Skeltons’ Haliburton Flora, however, is a fine addition to that all-too-short list of recently published floras. Eleanor and “Em” Skelton, both deceased shortly after the publication of this list, were amateurs in the most positive sense of that word. They studied botany for the love of it but they undertook their studies seri- ously and with care. For almost 20 years they scoured the rugged, beautiful Canadian Shield country of Haliburton County just south of Algonquin Park, col- 564 lecting over 2000 vouchers to support their observa- tions. Each of their carefully prepared and labelled specimens is deposited in the Royal Ontario Museum herbarium (TRT). The Skeltons took great care to review Haliburton specimens in other collections, including private sources, and sought expert taxonom- ic opinion whenever possible. The result is a fine record of Haliburton’s flora, neatly presented in a well-produced, paper-bound volume with clear, readable type and an attractive colour cover. The species treatments are sparse - one to two lines - but manage to concisely describe the status, habitat, and distribution (the latter represented by a list of codes for the County’s 23 townships) for each of 922 taxa (899 species) then known to occur in the County. The book is also significantly enhanced by an excellent introductory section describing the appropriate background context of geology, soils, vegetation, etc. The Haliburton Flora suffers a bit from a lengthy publication process that did not allow for the inclu- Trees in Canada By John Laird Farrar. 1995. Fitzhenry & Whiteside Limited, Markham, Ontario and the Canadian Forest Service, Natural Resources Canada, Ottawa. x + 502 pp., illus. $40.00. Trees in Canada is a wonderful reference and a steal at $40.00 (Canadian). The layout, type setting, color photographs, and black line drawings make it particularly easy to use. For those of us who relied on its predecessor Native Trees of Canada, the improvements are welcomed and readily apparent. Specifically, Trees in Canada now contains over 300 species and includes for the first time commonly planted or naturalized trees (i.e., Nordmann fir, Siberian larch, Scots Pine, and, to my surprise, Bald- cypress). For added convenience, all trees are strate- fied into 12 Groups, and these groups are identified on the endleaves inside the front and back covers of the book. Red colored tabs are used to show where these Groups appear in the book. The system is effi- cient and helpful. However, as in previous versions, the trees are also stratified by “The Conifers” and “The Broadleaf Trees” in the table of contents, for those who may have found that helpful. The two page per species format has been maintained, at least for all of the native Canadian tree species, and Rowe’s map of the Forest Regions of Canada has been retained. New additions are: maps of the Eastern and Western Plant Hardiness Zones in Canada, “Keys to Groups and Selected Genera’, “Winter Keys to the Genera”, a listing of “Botanical Authors”, and a section titled “Meanings of Tree Names”. Also for the first time, there are “Quick Recognition” tips for all the native THE CANADIAN FIELD-NATURALIST Vol. 110 sion of a relatively large number of additions made by others in the late 1980s. The declining health of the authors contributed to this, no doubt, but it did not prevent them from producing a competent, com- prehensive list. The Haliburton Flora provides and excellent working list and will be a useful reference for any botanist working in the Canadian Shield country of southern and southcentral Ontario. It also constitutes a fine botanical legacy for a delightful and productive couple. The Royal Ontario Museum is also to be con- gratuated for their encouragement and support of the Skelton’s efforts and the publication of such a high quality, reasonably-priced product. The sup- port of the lead provincial institution in the pro- duction of such valuable regional work is appro- priate, important,... and shamefully rare, in Canada. DANIEL F. BRUNTON 216 Lincoln Heights Road, Ottawa, Ontario K2B 8A8 tree species. I found these last three additions particu- larly “neat”, informative and helpful. What I did not like about the book was the fact that species maps do not accurately show the true range of the species. For example, from what is pre- sented in this book one might conclude the range of Pinus contorta Dougl. ex Loud. var latifolia Englem. only extends as far south as Wyoming, which is wrong. However, to the credit of the new version, one should note that the tree distributions in the pre- vious versions of this book used to stop at the bor- ders between the United States and Canada. So the author is trying to be more inclusive. However, I suspect from his perspective one has to decide where to cut these distributions off. Some of the species in this book are also found in Mexico and elsewhere. I am the kind of guy who loves to find and, when possible, identify trees. When visiting a new area of the continent [ take time to view the trees. I find “tree viewing” to be more fun than “bird watching”, because trees tend to remain in place long enough for me to get the job done. However, because of this interest, I have bought more than a few tree identifi- cation books. In my opinion, Trees in Canada 1s now one of the better books in my collection, so I recommend it without reservation to all botanists, greenhouse operators, landscape architects and nature enthusiasts. And I believe it will make an excellent Christmas gift. PAUL M. WOODARD Department of Renewable Resources, 751 General Services Building, University of Alberta, Edmonton, Alberta T6G 2H1 1996 BOOK REVIEWS 565 The Sunflower Family (Asteraceae) of British Columbia Volume 2, Astereae, Anthemideae, Eupatorieae and Inuleae By George W. Douglas. 1995. Royal British Columbia Museum, Victoria, British Columbia. vi + 393 pp., illus. $29.95 In 1982 the British Columbia Provincial Museum published The Sunflower Family (Asteraceae) of British Columbia Volume I -Senecioneae by George W. Douglas and illustrated by Elizabeth J. Stephen as Number 23 of the Occasional Papers Series. In the introduction of this volume the author anticipated publishing the eight tribes set up by Bentham in five volumes. Volume two was to include the Astereae, volume three the Anthemideae, Eupatoreae and Inuleae, Volume four the Cichoreae, and volume five the Cynareae and Heliantheae. The volume just published, however, includes the four tribes indicated in the title and the projected volume three will include the Lactuceae (Cichorieae), Cardueae (Cynareae) and Heliantheae. Hopefully, we will not have to wait so long for this last volume. In the introduction Douglas describes the world- wide distribution of the Asteraceae (Compositae), one of the largest plant families of which at least some species are known to almost everyone. He then discusses the historic and economic uses, floral form and structure, floral biology and pollination, seed dispersal, the arrangement of the tribes and genera in his volumes on British Columbia’s Sunflower family together with information on the location of the over 50 000 specimens he has examined during the writ- ing. He describes his treatment as conservative like those of Cronquist and Welsh, but unlike Cronquist he has treated geographically separate, or mainly separate, taxa of a species at the subspecies level and those with geographically sympatric ranges at the varietal level. In addition, he has not recognized any taxa at the forma level. Here also is a key to the Asteraceae tribes together with a series of excellent line drawings to help the user in working through this key. In the four tribes a total of 27 genera and 132 Species are recognized: Achillea (2), Anaphalis (1), ENVIRONMENT Antennaria (11), Anthemis (3), Artemisia (18), Aster (23), Bellis (1), Brickellia (1), Chamomilla (3), Chrysanthemum (3), Chrysothamnus (2), Conyza (1), Cotula (1), Erigeron (28), Eupatorium (1), Euthamia (2), Filago (2), Gnaphalium (7), Grindelia (2), Haplapappus (3), Heterotheca (1), Inula (1), Leucanthemum (3), Machaeranthera (1), Matricaria (3), Psilocarphus (2), Solidago (6), Tanacetum (3), ° and Townsendia (3). In the main text, like the key to the Asteraceae tribes, the keys to genera within the tribes are pro- vided with drawings to aid progress through the keys. The genera and species in each of the tribes are in alphabetical sequence. Each genus and species has an interesting introductory paragraph which includes numbers, habitats, characteristics, range, and uses, if any. This is followed by a detailed description refer- ences, comments if required, and chromosome num- bers, if known. Each species is accompanied by a full page of drawings depicting habit, heads, florets and other characters as required. These, like the ones in the keys were produced by the careful touch of Elizabeth J. Stephen. Distribution maps are also pro- vided for all species. The work is concluded with a glossary; Appendix 1: British Columbia Astereae, Anthmideae, Eupatoreae, and Inuleae species and their synonyms; Appendix 2: Synonyms of British Columbia Astereae, Anthemideae, Eupatoreae, and Inuleae; Appendix 3: Excluded taxa; Literature cited; and an index. Like the previous volume, this is an excellent treatment which will be welcomed by botanists, stu- dents, and naturalists both in and adjacent to British Columbia. WILLIAM J. Coby Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 Saving Nature’s Legacy: Protecting and Restoring By Reed F. Noss and Allen Y. Cooperrider. 1994. Island Press, Washington, D.C. 416 pp., Cloth U.S. $48.00; Paper U.S. $24.50. Biodiversity, in recent years, has become a topic of concern within the global community. A Convention on Biological Diversity has been creat- ed, resulting in Canada developing a strategy in response to the convention. Saving nature’s legacy is written to provide the land managers, who will be responsible for developing and implementing policy to conserve biodiversity, a guide. The authors “put forth a bold vision of what it might take to maintain all of biodiversity...”. Noss and Cooperrider provide a book complete with a detailed table of contents, a glossary, index, literature cited list, species list, and several tables for comparison of methodologies. Saving nature’s lega- cy also provides (1) historical background to the sub- 566 ject matter, (2) methodology, (3) a close look at management of forests, rangelands, and aquatic ecosystems, and (4) future needs. The book is not meant to be a complete literature review of the topic but provides some key references for further reading. These all complement the thorough and very read- able text, assuring the reader easy access to the infor- mation withing its pages for later referral. The book is one which should be recommended to any one interested in biodiversity issues. Even though the book is mainly American in content it THE CANADIAN FIELD-NATURALIST Vol. 110 should be useful in the Canadian situation. One word of warning, the book challenges and critiques present-day management of natural resources. For those attempting to implement conservation of bio- diversity, Noss and Cooperrider outline a conserva- tion plan which will be challenging, at the very least, to implement. M. P. SCHELLENBERG 434-4th Avenue S.E., Swift Current, Saskatchewan S9H 3M1 Where the Sky Began: Land of the Tallgrass Prairie By John Madson. 1995. Iowa State University Press, Ames. xiv + 326 pp. Cloth U.S. $24.95 In this revised edition Madson presents a magnifi- cent overview of the tallgrass prairies, in which two basic themes are addressed. The book begins with a review of some of the basic biological features of tall- grass prairies, including soil type, moisture regime, fire history, climatic conditions, productivity, flora, and fauna. Unfortunately, there is an inherent and unavoidable danger with using this approach — over- simplification. While there is little doubt that climate and fire history play a significant role in the establish- ment and maintenance of these grasslands, these notably complex ecosystems are still poorly under- stood. Recent discoveries such as plant-fungal co-evo- lutionary relationships (Clay 1988, 1991) indicate that we have much to learn. Nevertheless, as presented and intended, the author has successfully woven sci- ence into a series of anecdotes which I found pleas- antly informative and enjoyable to read. More impor- tantly however, readers not familiar with tallgrass prairie leave with a basic understanding of the unique biology and ecology of this rare type of grassland. In the second part of the volume, anthropogenic effects are considered. Madson makes an excellent point concerning human nature and the economic value of these once vast and untouched prairies over the last century. That is, the prairies were considered to be a resource that could be exploited at will. In our tenacity to tame the prairies and put the land to good use we have almost successfully eradicated the tallgrass prairie. Today it is estimated that less than 1% of the original tall grass prairie exists in small disjunct patches (a number of these patches are listed in the Appendix). Unfortunately, in our haste to clear the land, the number of reliable studies describing the ecology and inventories of the original flora and fauna are few, and provide us with only a partial pic- ture of what the structure and composition of natural tallgrass prairie was like. However, over the last two decades we have made significant advances in restoring prairies to their natural state and it is these prairies that are now allowing us to better understand these complex prairie ecosystems. I found the overall quality of the book to be good. However, I was disappointed with the poor quality of the line drawings. The publishers should have considered colored photographs, such as the one used for the jacket cover or at least watercolors for the illustrations. In addition, I found the consistent use of colloquial names for the flora and fauna, with- out reference to their scientific names frustrating. This problem could have been eliminated by adding the scientific name in brackets the first time the common name was used. After reading this book I felt that the role the prairies have had on our way of life is beginning to come full circle. The native people viewed them- selves as part of the prairie ecosystems, living on what the land naturally provided to them. Over the last 100 years the prairies have been exploited and nearly completely decimated. Floral and faunal species, many of which we will only know through paleoecological studies, are extinct. However, the number of tallgrass prairie restoration projects is growing every year, and with time we may have something more than disjunct patches of the tallgrass prairie that we can pass on as part of our heritage to future generations. References _Clay, K. 1988. Fungal endophytes of grasses: a defensive mutualism between plants and fungi. Ecology 69: 10-16. Clay, K. 1991. Parasitic castration of plants by fungi. Trends in Ecology and Evolution 6: 162-166. BEN A. LEPAGE Department of Geology, University of Pennsylvania, Philadelphia, Pennsylvania 19104-6316 1996 Masterworks of Man and Nature Edited by Robert Osborne. 1994. Facts on File. 2nd edi- tion. Harper-MacRae, Patonga, Australia 402 pp. $43.95 Robert Osborne and Mark Swadling with a host of writers, photographers and production people have assembled this book which documents the World Heritage Convention’s list of natural and cultural sites. Endorsed by UNESCO and IUCN (The World Conservation Union) the book is both a list of sites and an awareness campaign for the World Heritage Convention. As royalties from the sale of these books go to the World Heritage Fund, the publica- tion is a fund-raising scheme for the World Heritage Fund. It is an attractive, large coffee-table picture book which is easy to handle and invites one to look at it immediately. I had an hour of fun with it initial- ly just locating the places which I have visited and reading the pages around them. On closer inspection, I was impressed with many of the sites and the composition of the text. Far from being the ordinary travelogue, the book identifies sites of interest for their cultural or natural heritage. Certainly there is a come-and-see aspect to much of the book, but there is also a come-and-see but don’t touch aspect which is refreshing. Also refreshing are the number of sites which are identified as being wildlife refuges, in particular wildlife refuges for mammals and reptiles. There are a few sites which identify birds as the principle attraction, but of the wildlife sites they are the minority. Rare African pri- mates, elephants and deer are more often identified in the sites than are birds. Some of the sites are iden- tified as important resting places in migration as well as the homes of rare bird species. BOOK REVIEWS 567 This is not another pretty bird book. The whole tone of the book is conservation, history, events, and policies which influence our modern world and how each of these areas are entwined. I kept wondering why other sites were not included and how they could have been missed. For instance of the wonders of the world included, Niagara Falls did not make the list! Have we written off this landform or is there nothing in particular to save there? I have to disagree somehow. Canada did manage to include nine differ- ent sites and an essay by Harold Eidsvik, the Director of the Canadian Parks Services, on old growth forests. It is a measure of our conflicts though that none of the Canadian old growth forests have made the list to date. I find the fact that I am bothered somewhat by the contents of the book to be a good thing. A prophetic stance should bother us and, even better, move us to action. The book has a mildly prophetic character just by telling us that there are valuable natural and historical things slipping through our fingers and we can appreciate the value of holding onto them. On the other hand one can appreciate the book as a beautiful thing in itself, and I would not have any trouble tellng a school to buy it as a picture book or giving it as a gift to someone is who is going travel- ing and looking for exciting places to visit. I can also recommend it to a high school student interested in conservation, knowing that it will be easy reading with a solid message. Jim O’ NEILL St. Mark’s College, 5935 Iona Drive, Vancouver, British Columbia V6T 1J7 The Everglades Handbook: Understanding the Ecosystem By Thomas E. Lodge. 1994. St. Lucie Press, Delray Beach, Florida. xix + 228 pp., illus. paper U.S. $32.50. Because | spent two winters in the wildlands of southern Florida in the early 1970s, I looked forward to a book that promised a comprehensive overview of the biology and ecology of that marvelous region. I was not disappointed; Lodge has compiled a read- able, useful, and enjoyable volume. The book begins with a brief overview of geolo- gy, history, and landscape development. Lively descriptions of “Environments” (freshwater marshes, hardwood hammocks and other tree islands, pinelands, mangroves, coastal levees and saltmarsh- es, estuarine and marine waters) occupy 70 pages. A slightly longer section covers biogeography and the major groups of organisms. The author’s interests in fish biology and zoo- geography are evident. I appreciated the discussion of the role of the alligator in shaping microrelief and vegetation. Coverage of some topics, however, including soils, sawgrass autecology, amphibians, and “coastal lowland” vegetation is curiously brief. The exclusion of the Big Cypress Swamp seems arbitrary. Native American use of southern Florida is scarcely mentioned, although 20th century human impacts and restoration issues are discussed in the final 25 page chapter. There is a 263 item bibliogra- phy and many text citations. A 27-page index enhances the value of the book for reference. Numerous black-and-white photographs are mostly clear and complement the text, especially for non- biologist readers. The author’s charming preface recounts reading the entire manuscript aloud to Everglades conservationist and author Marjory Stoneman Douglas, who (at the age of 99) comment- ed on the text and wrote an introduction. 568 On the whole, The Everglades Handbook is well written and well edited. I found a few statements confusing (e.g., “[The snowy egret] may even feed on the wing, similar to a sea gull”). There are too many instances of subject-verb disagreement. Some readers may find use of the gender-biased “man” uncomfortable (e.g., “accustomed to the presence of man,” page 138). There are few typographic errors (all wetland ecologists should write the words “repellent,” “mosquitoes”, and “gases” on the backs of their hands). In five years a revised edition could fill some of the gaps in coverage, and report on pol- lution control and hydrological restoration. MISCELLANEOUS Evolutionary Naturalism By Michael Ruse. 1995. Routledge, London. 316 + x pp. U.S. $49.95 Naturalism is more than the field biology domi- nating this journal: more broadly it refers, in all of science, the arts, and ethics, to approaches rooted in experience of the natural world. Evolutionary natu- ralism, referring to the approach based on an evolu- tionary perspective, receives a fine treatment in ths book from Michael Ruse, well known, and appropri- ately so, as a philosopher of biology who makes his subject matter understandable and appealing. Nine essays are grouped into three sections, with useful introductions. Under Case Studies, Ruse considers the concept of natural selection as developed by Darwin and Wallace for evidence of the importance of the route in the discovery of the features of a the- ory, and the diagrams of adaptive landscapes by Sewall Wright for the frequently key role of pictures in communicating concepts, resulting in two stimu- lating essays. There is also a third, and almost gos- sipy, essay on the participants, and their impacts, in the paleontological controversy of punctuated equi- libria. Under Evolutionary Epistemology, Ruse examines possible parallelisms in the developments of soci- eties, sciences, and the organic world, including pair- wise comparisons, but more rigorous treatment is needed to achieve definite conclusions. The familiar issue of the historically close linkage of the theory of evolution with the concept of progress is nicely pre- sented, followed by an excellent exposition of evolu- tionary epistemology that Ruse correctly regards as the lynchpin of the book. The adaptive basis of our THE CANADIAN FIELD-NATURALIST Vol. 110 The Everglades Handbook will inform efforts to restore and protect one of the continent’s great wetlands. I hope regulators, managers, politicians, business people, scientists, ecotourists, and con- servationists who work in or visit Florida will study this book. I hope it will also serve as a model for comparable volumes on other wetland regions. ERIK KIVIAT Hudsonia Ltd., Bard College Field Station, Annandale, New York 12504-0217 cognitive abilities, their social and scientific conse- quences, and responses to criticisms are all beauti- fully narrated. Under Evolutionary Ethics there are equally excellent essays on the role of cultural values in evolutionary biology and on a sociobiological approach to ethics. Responding to critics always car- ries the threat of becoming dull, and such is the case here, but particularly valuable attention is paid to the refinement of epistemic values in science, to ques- tioning the cogency of the naturalistic fallacy, and to a view of morality as an adaptive product of altruism. The arguments are made with Ruse’s customary balance and zest, and broad historical and philosphi- cal sources are usefully invoked in a manner highly agreeable to practicing biologists, as might be expected from one who regards evolutionary thought as “one of the greatest testaments to human intellec- tual achievement’. Not that the treatment is free of all redundancies, such as the issue of progress, or of philosophical surprises, such as the rejection of “the notion of a reality beyond our experience”. There is a good bibliography and insufficient index (especial- ly regrettable in such an important book). In conclu- sion, Ruse does an excellent job in promoting the central importance of naturalism as THE approach providing genuine understanding of our world at large. His contribution is especially important in the face of non-natural challenges such as fundamental- ism, occultism, and intellectual sophistry. PATRICK W. COLGAN Canadian Museum of Nature, P.O. Box 3443, Station D, Ottawa, Ontario K1P 6P4 1996 NEW TITLES Zoology Biogeography of the reptiles of south Asia. 1996. By I. Das. Kreiger, Melbourne, Florida. U.S. $27.50. +A birder’s guide to Florida. 1996. By B. Pranty. 4th edition. American Birding Association, Colorado Springs, Colorado. 388 pp., illus. U.S. $18.95. *Birding in metro Halifax: a month-by-month adven- ture guide. 1996. By C. Stevens. Nimbus, Halifax. x + 310 pp. +Bird song: identificaiton made easy. 1996. By E. Jardine. Natural Heritage/Natural History, Toronto. 207 pp., illus. $14.95. +Coloniality in the cliff swallow: the effect of group size on social behavior. 1996. By C. R. Brown and M. B. Brown. University of Chicago Press., Chicago. xiii + 566 pp., illus. Cloth U.S. $95; paper U.S. $95. +The colonial waterbirds of Great Slave Lake, Northwest Territories: an annotated atlas. 1995. By J. Sirois, M. A. Fournier, and M. F. Kay. Occasional Paper No. 89. Canadian Wildlife Service, Yellowknife. 59 pp., illus. Free. Coral snakes of the Americas: biology, identification, and venoms. 1996. By J. A. Roze. Kreiger, Melbourne, Florida. +Duck use of the costal habitats of James Bay. 1996. By A. Reed, R. Benoit, R. Lalumiere, and M. Julien. Canadian Wildlife Service, Saint-Foy, Québec. 47 pp., illus. Free. Available in French. The ecology of a symbiotic community: volume 1: pop- ulation biology of the Japanese lizard Takydromus tachydromoides (Schlegel) (Lacertidae) and volume 2: the component symbiote community of the Japanese lizard Takydromus tachydromoides (Schlegel) (Lacertidae). 1996. By S.R. Telford, Jr. Kreiger, Melourne, Florida. +Female control: sexual selection by cryptic female choice. 1996. By W. G. Eberhard. Princeton University Pess, Princeton. xiv + 501 pp., illus. Cloth U.S. $85; paper WES 29°95) A guide to the frogs and toads of Belize. 1996. By J.R. Meyer and C. F. Foster. Kreiger, Melbourne, Florida. U.S. $24.50. ;Javelinas and other peccaries: their biology, manage- ment, and use. 1996. By L. K. Sowls 2nd edition. Texas A&M University Press, College Station. 352 pp., illus. U.S. $49.95. Kaleidoscope tree boas: the genus Corralus of tropical America. 1996. By P. J. Stafford and R. W. Henderson. Kreiger, Melbourne, Florida. *Landscape approaches in mammalian ecology and conservation. 1995. Edited by W. Z. Lidicker, Jr. University Minnesota Press, Minneapolis. ix + 215 pp., illus. U.S. $35.95. *Larvae of the North American caddisfly genera (Trichoptera). 1996. By G. B. Wiggins. 2nd edition. University of Toronto Press, Toronto. 457 pp., illus. $110. BOOK REVIEWS 569 Lizards. 1996. By M. Rogner. Translated by J. Hackworth (1992, 1994 German edition) 2 volume set. Kreiger, Melbourne, Florida. +Lone star dinosaurs. 1995. By L. Jacobs. Texas A&M University Press, College Station. xiv + 150 pp., illus. U.S. $27.95. (Canadian distributor: University British Columbia Press, Vancouver $39.95) +Messages from an owl. 1996. By M.R. Terman. Princeton U. P., Princton. xi+ 217 pp., illus. U.S. $24.95. Natural history of monitor lizards. 1996. By H. F. Delisle. Kreiger, Melbourne, Florida. U.S. $24.50. +Neotropical birds: ecology and conservation. 1996. By D. F. Stotz, J. W. Fitzpatrick, T. A. Parker II], and D. K. Moskovits. University Chicago Press, Chicago. xx + 478 pp., illus. + plates. Cloth U.S. $100, paper U.S. $37.50. +Ontario birds: a field guide to 125 common birds of Ontairo. 1996. By C. Fisher. Lone Pine, Edmonton. 159 pp., illus. $17.95. +Oppossums, shrews, and moles of British Columbia. 1996. By D. W. Nagorsen. Royal B.C. Museum Handbook. University British Columbia Press, Vancouver. 160 pp., illus. $24.95. +The Princeton field guide to the birds of Australia. 1996. Edited by K. Simpson. 5th edition. Princeton University Press, Princeton. viii + 400 pp., illus. U.S. $39.50. yRuddy ducks and other stifftails: their behavior and biology. 1996. By P. A. Johnsgard and M. Carbonell. University of Oklahoma Press, Norman. xiv + 291 pp., illus. U.S. $49.95. +Shrikes (Laniidae) of the world: biology and conserva- tion. 1995. Edited by R. Yosef and F. E. Lohrer. Volume 6, No. 1. Proceedings of the Western Foundation of Vertebrate Zoology, Camarillo, California. +Spell of the tiger: the man-eaters of Sundarbans. 1995. By S. Montgomery. Houghton Mifflin, Boston. xvii + 232 pp., illus. U.S. $12.95. The thermal warriors: Strategies of insect survival. 1996. By B. Heinrich. Harvard University Press, Cambridge. 332 pp., illus. U.S. $27. Botany +The book of swamp and bog: trees, shrubs, and wild- flowers of eastern freshwater wetlands. 1995. By J. Eastman. Stackpole Books, Mechanicsburg, Pennsylvania. xv + 237 pp., illus. U.S. $16.95. +Catkin-bearing plants of British Columbia. 1996. By T. C. Brayshaw. Revised edition. Royal British Columbia Museum, Victoria. 192 pp., illus. $24.95. +A descriptive forest inventory of Canada’s forest regions. 1996. By S. L. Gray. Information Report PI-X- 122. Canadian Forestry Service, Petawawa. 192 pp., illus. A field guide to the families and genera of woody plants of northwest South America (Columbia, Ecuador, Peru). 1993. By A. W. Gentry. University Chicago Press. Chicago. 918 pp., illus. Cloth U.S. $75; paper U.S. $45. 570 *Field guide to the palms of the Americas. 1995. By A. Henderson. G. Galeano, and R. Bernal. Princeton University Press, Princeton. 1x + 352 pp., illus + 64 plates. WiSo$75: Forest ecosystem classification for Manitoba: field guide. 1995. By C. Zoladeski. University British Columbia Press, Vancouver. 205 pp., illus. $24.95. +Forest plants of central Ontario. 1996. By B. Chambers, K. Legasy, and C. V. Bentley. Lone Pine, Edmonton. 448 pp., illus. $24.95; U.S. $19.95. Plant collecting for the amateur. 1996. By T.C. Brayshaw. Revised edition. Royal British Columbia Museum, Victoria. 48 pp., illus. $8.95. +Plants of southern interior British Columbia. 1996. Edited by R. Parish, R. Coupe, and D. Lloyd. Lone Pine, Vancouver. 463 pp., illus. $24.95; U.S. $19.95. +Trees and shrubs of British Columbia. 1996. By T.C. Brayshaw. Royal British Columbia Museum Handbook. University British Columbia Press, Vancouver. 320 pp., illus. $24.95. *Trees in Canada. 1995. By J. L. Farrar. Fitzhenry & Whiteside, Markham, Ont. x + 502 pp., illus. $40. +Wetland plants of Oregon and Washington. 1995. By B. J. Guard. Lone Pine, Redmond, Washington. 239 pp., illus. U.S. $19.95. + Wild flowers: legends, poems, and paintings. 1996. By N.R. Ranson and M. J. Laughlin. Edited by H. E. Laughlin. Texas A&M University Press, College Station. 102 pp., illus. U.S. $16.95. Environment +Achieving sustainability in Canada. 1996. Edited by A. Dale and J. B. Robinson. University British Columbia Press, Vancouver. c256 pp., $75. +Dealing with risk: why the public and the experts dis- agree on environmental issues. 1996. By H. Margolis. University Chicago Press, Chicago. c200 pp., illus. U.S. $24.95. +Dictionary of natural resource management. 1996. By J. and K. Dunster. University British Columbia Press, Vancouver. c360 pp., illus. $90; U.S. $74.95. +Ecology of the Bering Sea: a review of Russian litera- ture. 1996. Edited by O. A. Mathiesen and K. O. Coyle. Report 96-01. University of Alaska Sea Grant College, Fairbanks. 306 pp., illus. U.S. $25. Global diversity assessment. 1995. Edited by V. H. Heywood. Cambridge University Press, Cambridge. 1140 Pp. +Hiking the ancient forests of British Columbia and Washington. 1996. By R. Stoltmann. Lone Pine, Edmonton. 191 pp., illus. $19.95; U.S. $15.95. +The ice-age history of national parks in the Rocky Mountains. 1996. By S. A. Elias. Smithsonian Institute Press, Washington. xi1.+ 170 pp., illus. U.S. $16.95. THE CANADIAN FIELD-NATURALIST Vol. 110 +Life in 2030: exploring a sustainable future in Canada. 1996. By J.B. Robinson. University British Columbia Press, Vancouver. c224 pp., $25.95. +Mountain bike Nova Scotia: a guide to off-road cycling in Nova Scotia. 1996. By G. Brown and K. Degooger. Nimbus, Halifax. 181 pp., illus. $12.95. Passing the buck: federalism and Canadian environ- mental policy. 1996. By K. Harrison. University British Columbia Press, Vancouver. c244 pp. $70. +People and environment: development for the future. 1996. Edited by S. Morse and M. Stocking. University British Columbia Press, Vancouver. c224 pp. $24.95. +Population dynamics in ecological space and time. 1996. Edited by O. E. Rhodes Jr., R. K. Chesser, and M.H. Smith. University Chicago Press, Chicago. c336 pp.. illus. Cloth U.S. $50; paper U.S. $17.95. Science and ecosystem management in the National Parks. 1996. Edited by W.L. Halverson and G. E. Davis. University Arizona Press, Tuson. 224 pp., illus. Cloth U.S. $45; paper U.S. $15.95. 7+Sea Kayaking in Nova Scotia: a guide to paddling routes along the coast of Nova Scotia. 1996. By S. Cunningham. Nimbus, Halifax. ix + 244 pp., illus. $16.95. +Species diversity in ecological communities: historical and geographical perspectives. 1994. Edited by R. E. Ricklefs and D. Schlater. Chicago University Press, Chicago. 416 pp., illus. Cloth U.S. $98; paper U.S. $32.50. +Status of the World ocean and its biodiversity. 1995. By D.E. McAllister. Ocean Voice International, Box 37026, Ottawa, K1V OWO. Sea Wind 9(4). 72 pp. $7.50 + $2.50 postage. Miscellaneous +Charles Darwin voyaging. 1996. By J. Browne. Princeton University Press, Princeton. xv + 605 pp., illus. U.S. $18.95. *The fossils of the Burgess shale. 1994. By D. E.G. Briggs, D. H. Erwin, and F. J. Collier. Smithsonian Institute Press, Washington. xvil + 238 pp., illus. +Hiking Nova Scotia: thirty of Nova Scotia’s best hiking trails. 1995. By J. Light. Nimbus, Halifax. vi + 106 pp., — illus. Monad to man: the concept of progress in evolutionary biology. 1996. By M. Ruse. Harvard University Press, Cambridge. 640 pp., illus. U.S. $49.95. *River out of Eden: a Darwinian view of life. 1995. By R. Dawkins. Basic Books, Harper Collins, New York. 172 pp., illus. U.S. $20; Cdn. $28. +The shape of life: genes, development, and the evolu- tion of animal form. 1996. By R. A. Raff. University Chicago Press, Chicago. c520 pp., illus. cloth U.S. $55; paper U.S. $29.95. *assigned for review tavailable for review TABLE OF CONTENTS (concluded) redation of an Eastern Chipmunk, Tamias striatus, by a Gray Squirrel, Sciurus carolinensis STEVEN D. FACcIO ate spring arrival, nesting, and fall departure by Common Nighthawks, Chordeiles minor, in Saskatchewan in 1995 Ray G. PouLIN, PAUL A. BRADSHAW, and MarK D. GRAHAM orsehair fungus, Marasmius androsaceus, used as nest lining by birds of subalpine spruce-fir community in the northeastern United States K. P. MCFARLAND AND C. C. RIMMER ews and Comment The Ottawa Field-Naturalists' Club 1996 Awards — 1995 Member of the Year Award: Bob Bracken — - 1995 Conservation Award to a Member: Jeff Harrison — 1995 Conservation Award to a Non- ~ member: Kit Chubb — 1995 Anne Hanes Natural History Award: Marilyn Light — 1995 President's ' Prize: William D.(Dave) Smythe — Errata and Addenda to A Life with Birds: Percy A. Taverner, | Canadian Ornithologist, 1875-1947 — Rana-Saura: Amphibian Follow-up Project for the Atlas of ' Amphibians and Reptiles of Quebec — The Frog Monitor and DAPCAN IV Proceedings — Canadian _ Association of Herpetologists Bulletin — Notice of the 118th Annual Business Meeting of The _ Ottawa Field-Naturalists' Club — Call for Nominations: The Ottawa Field-Naturalists' Club 1997 | Council — Call for Nominations: The Ottawa Field-Naturalists' Club 1996 Awards — Book Review | Editor's Report for The Canadian Field-Naturalist Volume 109 (1995) — Recovery: An Endangered Species Newsletter — The Boreal Dip Net — Great Lakes Fact Sheet: Amphibians and Reptiles in Great Lakes Wetlands: Threats and Conservation — Joint Annual Meeting Working Group on | Amphibian and Reptile Conservation in Canada and Task Force on Declining Amphibians and ; Populations in Canada (DAPCAN) — Global Biodiversity — Adopt a Black Rat Snake Program, 4 | / Charleston Lake Provincial Park, Ontario — Sea Wind: Bulletin of Ocean Voice International Froglog: IUCN/SSC Declining Amphibians Populations Task Force — Newsletter: Ontario Natural Heritage Information Centre — Picoides: Bulletin of the Society of Canadian Ornithologists — Wild Animal and Plant Protection and Regulation of International and Interprovincial Trade Act (WAP- PRIITA) — Conservation Groups in Canada — SAMPA II i r tribute to Claude Eugene Garton, 1907-1996 JOAN HEBDEN Book Reviews - Zoology: Shells of the Atlantic and Gulf Coasts and the West Indies — The Summer Atlas of North American Birds — A Photographic Guide to North American Raptors — Flammulated, Boreal, and Great Gray Owls in the United States: A Technical Conservation Assessment Botany: Rare Vascular Plants in the Canadian Arctic — Rare Vascular Plants in the Northwest Territories — Aquatic and Wetland Vascular Plants of the Northern Great Plains — La symbiose mycorhizienne: Etats des connaissances — Poisonous Plants of Canada — Plant Alert -- Haliburton Flora: An Annotated List of the Vascular Plants of the County of Haliburton, Ontario — Trees in Canada — The Sunflower Family (Asteraceae) of British Columbia Volume 2, Astereae, Anthemidae, Eupatorieae, and Inuleae 2nvironment: Saving Nature's Legacy: Protecting and Restoring — Where the Sky Began: Land of the Tallgrass Prairie — Masterworks of Man and Nature — The Everglades Handbook: Understanding the Ecosystem Miscellaneous: Evolutionary Naturalism New Titles Mailing date of the previous issue 110(2): 4 July 1996 538 538) 541 544 554 558 561 565 568 569 THE CANADIAN FIELD-NATURALIST Volume 110, Number 3 Articles Endemic vascular plants of British Columbia and immediately adjacent regions GEORGE W. DOUGLAS Catatropis lagunae n. sp., Trematoda, Notocotylidae, parasite d'oiseaux de mer CHRISTIANE BAYSSADE-DUFOUR, JEAN-LOUIS ALBARET, HELENE FERMET-QUINET, et KHEMAIS FARHATI Road mortality of amphibians, reptiles and other wildlife on the Long Point Causeway, Lake Erie, Ontario E. PAUL ASHLEY and JEFFREY T. ROBINSON The meningeal worm, Parelaphostrongylus tenuis, a marginal limiting factor for Moose, Alces alces, in southern Québec ANDRE DUMONT and MICHEL CRETE Dietary flexibility of shorebirds in the western hemisphere SUSAN K. SKAGEN and HEATHER D. OMAN Population estimate and habitat associations of the Loggerhead Shrike, Lanius ludovicianus, in southeastern Alberta RONALD R. BJoRGE and DAVID R. C. PRESCOTT Four records of the Chestnut Lamprey, /chthyomyzon castaneus, new to Ontario CLAUDE B. RENAUD, SANDRA C. RIBEY, and FRANCOIS CHAPLEAU Rare and endangered fishes and marine mammals of Canada: COSEWIC Fish and Marine Mammal Subcommittee Status Reports: X ROBERT R. CAMPBELL The status of the Eastern Sand Darter, Ammocrypta pellucida, in Canada ERLING HOLM and NICHOLAS E. MANDRAK The status of the Cutlips Minnow, Exoglossum maxillingua, in Canada E. J. CROSSMAN and ERLING HOLM The status of the Lake Chubsucker, Erimyzon sucetta, in Canada NICHOLAS E. MANDRAK and E. J. CROSSMAN The status of the Blackchin Shiner, Notropis heterodon, in Canada J. HOUSTON The status of the Rosyface Shiner, Notropis rubellus, in Canada J. HOUSTON The status of the Warmouth, Chaenobryttus gulosus, in Canada E. J. CROSSMAN, J. HOUSTON, and ROBERT R. CAMPBELL The status of the Bearded Seal, Erignathus barbatus, in Canada HOLLy J. CLEATOR The status of the Long-finned Pilot Whale, Globicephala melas, in Canada DAWN NELSON and JON LIEN The status of the Pigmy Sperm Whale, Kogia breviceps, in Canada ROBIN W. BAIRD, DAWN NELSON, JON LIEN, and DAvID W. NAGORSEN Notes Tree-climbing by Arctic Ground Squirrels, Spermophilus parryii, in the southwestern Yukon Territory ANNE H. Huss, TIM KARELS, and ANDREA BYROM Do female Sharp-tailed Grouse, Tympanuchus phasianellus, copulate only once during a breeding season? LEONARD J. S. TsuJI Interactions of a White-winged Black Tern, Chlidonias leucopterus, with Arctic Terns, Sterna paradisaea, at Churchill, Manitoba JOSEPH R. JEHL, JR. 199 38 44s 45( 454 462 47( 478 482 486 495 501 Il 325 338 535 530 continued on inside back covel ISSN 0008-3550 The CANADIAN FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Volume 110, Number 4 October—December 1996 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patron His Excellency The Right Honourable Roméo LeBlanc, P.C., C.C., C.M.M., C.D., Governor General of Canada The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse infor- mation on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintain- ing or restoring environments of high quality for living things. Honorary Members Edward L. Bousfield Clarence Frankton Stewart D. MacDonald Loris S. Russell Irwin M. Brodo W. Earl Godfrey Verna Ross McGiffin Douglas B.O. Savile William J. Cody C. Stuart Houston Hue N. MacKenzie Pauline Snure Ellaine Dickson George F. Ledingham Eugene G. Munroe Mary E. Stuart R. Yorke Edwards Thomas H. Manning Robert W. Nero Sheila Thomson Anthony J. Erskine Don E. McAllister Hugh M. Raup 1996 Council President: David W. Moore Ronald E. Bedford Jeff Harrison Vice-Presidents: Michael Murphy ME edo Cotueha) lens Lee Cairnie Ann MacKenzie i : William J. Cody Patricia Narraway Recording Secretary: David Smythe Francis R. Cook Frank Pope Corresponding Secretary: Eileen Evans Ellaine Dickson Tom Reeve ‘ 17: Colin Gaskell Jane Topping Treasurer: Gillian Marston Mane aaR Ghaciicaynee Christine Hanrahan Ken Young Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2. For information on Club activities telephone (613) 722-3050. 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. Editor: Francis R. Cook, R.R. 3, North Augusta, Ontario KOG 1RO; (613) 269-3211 Copy Editor: Wanda J. Cook Business Manager: William J. Cody, P.O. Box 35069, Westgate P.O. Ottawa, Canada K1Z 1A2 (613) 759-1374 Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP 1J0 Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta, Edmonton, Alberta T6G 2E9 Associate Editors: Robert R. Anderson Robert R. Campbell W. Earl Godfrey Warren B. Ballard Brian W. Coad William O. Pruitt, Jr. Charles D. Bird Anthony J. Erskine Chairman, Publications Committee: Ronald E. Bedford All manuscripts intended for publication should be addressed to the Editor with the exception of book reviews which should go directly to Book Review Editor. Subscriptions and Membership Subscription rates for individuals are $23 per calendar year. Libraries and other institutions may subscribe at the rate of $38 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $23 includes a subscription to The Canadian Field-Naturalist. All foreign subscribers (including USA) must add an additional $5.00 to cover postage. 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. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Date of this issue: October-December 1996 (March 1997), Back Numbers and Index Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field-Naturalists’ Club, 1879-1886, and The Ottawa Naturalist, 1887-1919, and Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index compiled by John M. Gillett, may be purchased from the Business Manager. Cover: Dorsal (left) and lateral (right) view of the head of a Night Snake found 5 September 1992 between Oliver and Osoyoos, British Columbia, and photographed 15 January 1993 by Tom Gore, University of Victoria, British Columbia (The same snake is also depicted in Figure 1, page 623). Note the characteristic vertical eye pupils (in bright light) visible in both views. The only other snakes in Canada to have vertical pupils are the rattlesnakes (genera Crotalus and Sistrurus). Photograph courtesy of Patrick T. Gregory, Department of Biology, University of Victoria, Victoria, British Columbia V8W 3N5. See paper on new records and diet of the Night Snake in British Columbia by Howard Lacey, Christopher H. Shewchuk, Patrick T. Gregory, Michael J. Sarell, and Linda A. Gregory, pages 620-625. 1 The Canadian Field-Naturalist Volume 110, Number 4 September—December 1996 The Land Mammal Fauna of Southeast Alaska STEPHEN O. MACDONALD!” and JoSEPH A. Cook? 1P_O. Box 58, Gila, New Mexico 88038 "University of Alaska Museum, Fairbanks, Alaska 99775-6960 MacDonald, Stephen O., and Joseph A. Cook. 1996. The land mammal fauna of southeast Alaska. Canadian Field- Naturalist 110(4): 571-598. Poorly documented distributions for most mammal species in southeast Alaska have long hampered studies of this highly diverse and insular fauna. New information based on 4280 specimens in the University of Alaska Museum and a review of other large collections is reported for 54 species of land mammals that occur or have recently occurred in Southeast Alaska. Based on the presence of endemic taxa or unique combinations of taxa, five subregions can be identified. Patterns of species richness and endemism are similar to other archipelagos. The origin of this fauna has been complex and includes species with Beringian and/or southern refugia affinities. Fossil mammals recovered from Prince of Wales Island and the large number of endemics suggest a coastal corridor along the continental shelf that allowed the movement of a number of mammals into southeast Alaska prior to the Fraser glacial advance. Key Words: mammal, endemic, conservation, island, temperate rainforest, Alexander Archipelago, Alaska. For systematists and biogeographers, Alaska remains one of North America’s last frontiers; nowhere is this sense of frontier more pronounced than amid the hundreds of islands, fiords, and moun- tains that constitute Southeast Alaska, the “panhan- dle” of the 49th state. Indeed, documentation of this complex region’s biological diversity remains at the early stages of exploration and discovery. Even for such high-interest animals as mammals, basic infor- mation on distribution and taxonomic status has been limited, unfocused, or inaccessible, resulting in only broad (Hall 1981; Manville and Young 1965) or pop- ular (Dufresne 1946; Rearden 1981) treatments. An unfortunate consequence is that the tremendous potential of this region for evolutionary, biogeograph- ical, and ecological investigations has not been real- ized. Given the accelerating rate of human-induced stresses and changes on the region’s terrestrial and marine ecological systems, it is even more disturbing that we lack detailed and accurate information for making sound conservation evaluations and wise management decisions. Knowledge of the distribution and taxonomy of the region’s mammals (and particularly its “nongame” mammals) has until recently been based on field col- lections made in the late 1800s and early 1900s, notably by the United States Biological Survey (U.S. National Museum, Washington, D.C.) and the Alexander Alaska Expeditions (Museum of Vertebrate Zoology, Berkeley, California). With few exceptions, these have been our primary sources of information despite their limitations in geographic scope, time spent at collecting localities, and number of specimens obtained. In this paper, we provide new information (gath- ered primarily by UAM through the field season in 1995), corrections, and a current overview of the dis- tribution and taxonomic status of the land mammals that occur or have recently occurred in Southeast Alaska. The marine mammals (24 species of pin- nipeds, cetaceans, and the Sea Otter, Enhydra lutris) of Southeast Alaska are not considered here. Area of Study Southeast Alaska (Figure 1) is defined as the region south of the Malaspina Glacier (59°45'N latitude) and north of Dixon Entrance (54°30’N latitude). With its approximately 16 000 km of shoreline, Southeast Alaska extends some 800 km north to south and 160 km east to west. This region is a naturally fragmented and complex mosaic of hundreds of small to large islands (the Alexander Archipelago), deep fiords, straits and inlets, and a mountainous mainland with numerous and extensive glaciers and ice fields. More than 1000 islands constitute the archipelago with Prince of Wales Island being the largest (> 5000 km/?; Orth 1967). Nelson (1887) and Swarth (1911, 1936) recognized this region as zoogeographically distinct, and termed it the Sitkan district. Along the narrow strip of main- land, the Coastal and St. Elias mountain ranges delim- it the eastern border of the region. They rise precipi- tously to over 5000 m in elevation to form a climatic barrier between the heavy rainfall and relatively mod- Sl D2 erate temperatures of Southeast Alaska and the dry and more extreme temperatures of interior British Columbia. These ranges and their ice fields also act as effective physical barriers to plant and animal disper- sal between the two distinct regions. Six major rivers, the Alsek, Chilkat, Taku, Whiting, Stikine, and Unuk drainages, transect these mountain ranges and may function as dispersal corridors. General overviews and descriptions of the region can be found in Swarth (1936), Klein (1965), Harris et al. (1974), and Mann (1986). Southeast Alaska and coastal British Columbia embody the most extensive temperate rainforest in the world. Together they con- tain a large percentage of the remaining, uncut ancient forests (Alaback 1991). In recent decades this region has been experiencing significant impacts from large-scale logging, roadbuilding, and other human activities. Although much of the international conser- vation focus has been on fragmentation of tropical rainforests (Lovejoy et al. 1986; Saunders et al. 1991), it is quite clear that temperate coastal forests may be experiencing similar effects. Methods and Materials The following accounts incorporate previously unpublished distribution records obtained from field work conducted through the University of Alaska Museum (UAM), University of Alaska, Fairbanks (UAF), over the past thirteen years. As a result of these efforts, VAM houses some 4280 mammal spec- imens (including study skins, skeletons, fluid preser- vations, frozen tissues, karyotypes, and ectoparasites) from across the region (through December 1995). We supplement these specimens with published accounts and verified records from other collections that have specimens and archival information on Southeast Alaska mammals, including those of the American Museum of Natural History (AMNH), Carnegie Museum of Natural History (CMNH), Glacier Bay National Park (GBNP), University of Kansas (KU), University of British Columbia (UBC), University of California Los Angeles (UCLA), University of California Berkeley (MVZ), University of Puget Sound (UPS), United States National Museum (USNM), and the Alaska Department of Fish and Game (ADF&G). Those institutions contain an addi- tional 4600 specimens. Finally, we have relied on the personal observations of several knowledgeable resi- dents of the region (details on file at UAM). Of the 48 native species extant, 25 (52%) are known at the regional scale from fewer than 30 speci- mens. We now have varying levels of specimen rep- resentation from 87 islands (Table 1) of the > L000 that constitute the Alexander Archipelago. The U.S. Biological Survey and the Alexander Alaska Expeditions sampled 22 islands early this century. Generally, coverage has focused on the larger islands. With few exceptions, however, most of our knowl- edge remains rudimentary and poorly represented with specimens. Peromyscus keeni with 2133 speci- THE CANADIAN FIELD-NATURALIST Vol. 110 mens, Sorex monticolus with 1119, Microtus longi- caudus with 655, Ursus arctos with 636, Sorex cinereus with 633, and Mustela vison with 571 are the best documented species. Species that remain poorly represented include bats, many of the larger mam- mals, and most of the species with limited distributions. Scientific nomenclature follows Wilson and Reeder (1993), with recent changes by Hogan et al. (1993). Vernacular names and the arrangement of the taxo- nomic accounts generally follow Jones et al. (1992). Subspecies (Hall 1981, unless otherwise noted) are reported for those taxa that are polytypic in Southeast Alaska. Information on specimens is available from JAC upon request. Annotated List of Species Sorex cinereus Kerr, 1792, Masked Shrew Sorex cinereus occurs along the entire coastal mainland of Southeast Alaska, on islands of the Alexander Archipelago north of Sumner Strait (except Admiralty Island), and on islands in close proximity to the mainland south of Sumner Strait. Two subspecies, S. c. streatori Merriam (type locality = Yakutat) and S. c. cinereus Kerr, have been recog- nized in this region, the latter from White Pass only (Jackson 1925, 1928; Hall 1981; Antell 1987). Mainland specimen records have been reported by several investigators (Merriam 1895; Osgood 1900; Swarth 1911; Jackson 1928; Baker 1951; Antell 1987), in addition to more recent unpublished UAM records. Island specimen records include Baranof, Bell, Black, Chichagof, Douglas, Etolin, Gedney, Grant, Gravina, Hassler, Inian, Kruzof, Kuiu, Kupreanof, Lemesurier, Mitkof, Read, Revillagigedo, and Wrangell islands (Heller 1909; Swarth 1911; Jackson 1928; UAM). The occurrence of Sorex cinereus on Admiralty Island (Hall 1981) has been based on a single speci- men collected during the 1907 Alexander Alaska Expedition (Heller 1909). This specimen (MVZ 68), recently examined by N. Dokuchaev and JAC, is a combination of S. cinereus skull with S. monticolus study skin. This mix-up, along with our failed efforts to turn up this shrew during extensive trapping efforts on the island, lead us to delete Admiralty Island as a valid locality for the species. Sorex monticolus Merriam, 1890, Dusky Shrew Sorex monticolus occurs throughout Southeast Alaska’s mainland and most of the islands of the Alexander Archipelago except, perhaps, Baranof and Chichagof islands. Hall (1981) recognized four sub- species (as S. vagrans) in Southeast Alaska: alascen- sis (type locality = Yakutat), ellassodon, longicauda (type locality = Wrangell) and malitiosus (type locali- ty = east side of Warren Island). Dusky Shrew popu- lations from the head of Lynn Canal were recognized as §. [m]. obscurus (Jackson 1928; Hennings and Hoffmann 1977; Antell 1987). 1996 yal x oe ay cr Sou We Kruzof I. (e) Gulf of Alaska \ CANADA ALASKA \ a © Pacific Ocean Map Area MACDONALD AND COOK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA a3 CANADA Dall I. orrester 1.8 KILOMETERS 40 80 FiGuRE 1. Map of Southeast Alaska and the Alexander Archipelago. Latitudinal and longitudinal coordinates for all islands identified in the species accounts are reported in Table 1. Specimens of this species have been collected along the mainland from Yakutat to Portland Canal (Swarth 1911; Jackson 1928; Hennings and Hoffmann 1977; UAM), and from the following islands: Admiralty, Anguilla, Back, Baker, Barrier, Bell, Betton, Black, Cap, Cat, Coronation, Dall, Douglas, Duke, Eagle, Etolin, Forrester, Gedney, Gravina, Hassler, Heceta, Hoot, Inian, Kadin, Kuiu, Kupreanof, Lemesurier, Long, Lowrie, Lulu, Marble, Mary, Mitkof, Noyes, Owl, Pleasant, Prince of Wales, Revillagigedo, San, San Fernando, Sangeo, Shrubby, Sukkwan, Sullivan, Suemez, Tuxekan, Vank, Warren, Woewodski, Woronkofski, Wrangell, and Zarembo (Heller 1909; Swarth 1911; Jackson 1919, 1928: McGregor 1958; Harris 1968; Hennings and Hoffmann 1977; UAM). Swarth (1911) reported specimens from Shakan, which is on Kosciusko Island, but it remains unclear if he meant nearby on Prince of Wales Island (Hall 1981). The occurrence of S. monticolus on two major islands in the Alexander Archipelago remains contro- versial. A specimen identified as this species from Baranof Island was reported by Hall (1981, as S. vagrans elassodon) in reference to Jackson (1928: 131). Jackson’s (1928) record is a skin without skull (USNM 0238296). Subsequent collecting efforts on Baranof Island have documented only S. cinereus. Most shrew species are difficult to distinguish with THE CANADIAN FIELD-NATURALIST Vol. 110. 574 (panuyuo)) . . ‘ . . (M06 oZEL ‘N07 09S) OGuIAIBT ‘(MAES EET ‘N/ES o$S) UATE AA (M.LT of €1 ‘NUTS oS) SNUOMUAY SNA}10IOPQC ‘SIsUuapoUDI DAJUOT ‘UOSIA “PW ‘DSNjIas ‘2 ‘WW ‘ppuajaa pautuisa vjaysnyy ‘snuvrisawv snsyy ‘sndny sung ‘SMLADUOAOD “YW ‘SNPNVIIBUO] SNJOAII ‘SndiUvaIO “y “gq ‘11Uaay SNIsKMosag ‘SISUAPDUDI LOJSD.) ‘Suodfiasi4s snuLigns skuosnDy]|y ‘(A[WO ‘J OquIaIeZ,) snoIUOSpny snaniospnUuD] ‘snoULof{yvo “ ‘uaay “Wy ‘snsnfion] syodp ‘snsoyyou ‘wu “sg ‘snjooruow xasog ueyoxn |, (M9P oCEL “N,90 o$$) UBMAING “(M17 o€ET ‘N,OT SS) zaulang (M,/6 oZET ‘N,Z0 29S) WOSUIAI}S (MA8S oZET “N/ET 09S) AQqnayg “(A\,LZ o€E1 ‘N,ST oSS) BINA BIBS “(AAT o€ET ‘N00 oSS) ovsuesg (M07 of €1 ‘N,OE $$) OpuBUaT UeS (A\,1Z o£E1 ‘N,9S SS) UBS (MOE oEET ‘N,IZ o$S) SOPRA JO sul (MST o€ EI “NJES o$S) IMO “(MPT of EI ‘N,LS oSS) HIO “(AL,OP o€€1 ‘N,OE o$S) SAXON “(MTT of EL NSO oS) AtBIAT “(AN LT of ET ‘N,8S oSS) AIGIRIN ‘(ALOE EET ‘N,8Z oSS) INT (MTP oCET ‘NA TS of S$) UOT ‘(LEE oEET ‘N, £0 09S) ONSMIDSOY “(AA €Z EET ‘NES 6S) jooH (MOE o€€1 “N/SP SS) BI999H “(ME o€ET ‘NSP ofS) 4OISAAIOT “(AL,SE EET ‘N,8E SS) feqinbsy (M07 of ET 'N, SS o$S) URNED TA “CAL,O€ o€ET ‘NES o$S) A1BeA “(M07 oI EL ‘N,SS obS) a4NG (MPT oT EL “N,6S ofS) “SE ¥9N ‘(M\,00 o€€1 ‘NVLS ofS) HYG “(MP1 ob EI ‘N,£S o$S) UOReUOI0; SANV TS] (CMLBE of ET ‘N97 o$S) AUOD “CAL,ST oT ET ‘N, 10 oS) 38D “(MA,ZZ of€T ‘N,ES 06S) deg YaHLAO (MST oET “NSP ofS) ST are “(M\,9€ o€€1 ‘N,ZZ o$S) OMe ‘(AL,SE EET ‘NOP oSS) BINSUy NUYFHLAOG *¢ (M,TI oZET “N,9T 09S) HOSURAAA “(MOF o€ET ‘N, £7 9S) EYSJOYUOIO AA SaJ]D sa]y ‘snuonuay snapio20pQ ‘SISUAPDUDI DAJUOT ‘0]NS OND ‘UOsIA ‘PF ‘VaunuLa Djaisnyy ‘DUDILIAUD SALAD ‘SNUDILIAWUD SNsAzy ‘sndn] siuvy ‘wnyvssop uozlyjary ‘smiuospny sndvz ‘snoiyjagiz bAjppug ‘snpnva18uo] W ‘snoiupajdsuuad snjosniyy ‘snjos 14addvs sXuouoiyjayy ‘1Uaay SNISKUOAAg ‘SISUaPOUDI AOSV ‘snutaqos sKuoony]H ‘snoquospny snamiospiuD |, ‘suDspayoou SisajakuoisvT ‘subjoa W ‘Muaay “- ‘snénfion] syodyy ‘snjooyuou *g ‘snasauis xa10g (M00 of €1 ‘NPE 09S) FISPOMIOAA “(ML 9E oZET ‘N,8Z 09S) HUBA “(MFI oOET ‘N,9P ofS) ssesuoy, (AOS oLET NTE ofS) ST YsOOB, “(MSE oZET ‘N,OE 09S) JOOMOS ‘(M\,0Z oI EI ‘N,SE o$S) OpasiselplAaay (MOS oZET ‘NOP 09S) JOM “(ALOE o€€T ‘N, Sh 9S) Jourasdnyy “(M00 PEI ‘NSE 09S) HIN (M6S oZEL 'N,9E 09S) BUM “(MLZ oZET ‘N,ZE 09S) WIPE “(M91 o8E ‘N,ES 0S) Ao[sseH “(MALT oSET “Nv ET oLS) PHBH “(M,b0 o€€1 ‘N,LE 09S) PHD (M,9F oIET ‘N,LI 0S) BUIARBID (MEP oLEL “N/E o$S) WRAD “(MTP oT ET ‘N,TS o$S) Loupay ‘(AA,€0 o€E1 ‘NSE 09S) ey (MATT oTET “N,90 09S) UNOVA “(ALO o€E1 ‘N,6Z 09) UOISNJIUOD ‘(ALOT EET ‘N,OP 09S) ‘ST apse SANV TS] YANN] (M0 of €1 ‘N,ZE 09S) WOMAN “CAL, Oh oLET ‘N, PS oSS) MIF (MBP oI ET ‘NIE oSS) U0}Wog NaaHLNOg (CMOE oT EL “NLS oS) HOM “(MSP of EL ‘NUE o$S) WBA “(M87 oIET ‘N,60 0$S) aouUy %Y TIAA *7 SNUDIIAIUD SOUMDAIC) ‘IVP S1AQ ‘snpUdAD] 4afiduoy ‘$2910 sary ‘snuOIUAaY snajloJ0pQ ‘sisuappuDdI XUCT MOJOIUOD DUN, ‘SISUaPDUDI DAJUOT ‘O]NS O]NH ‘uosiA ‘Py ‘syoAU W ‘SiSuadsD]D ‘a " ‘Vaulusa Djasn ‘unuuad ‘py ‘buvoivawup SAAD ‘SOJIAD ‘f) ‘SnuDII4aUID SNS.1-V ‘Sadjna sadn, ‘sndnj ‘D ‘SUD.AID] SIUDD ‘Unjossop UorYJaIy ‘sdaouid "7 ‘sniuospny sndoZ ‘stjvasog skwojdoudy ‘snomjaqiz vappug ‘snpnvoi8u0] W ‘Sisuaipinyvd ‘oP ‘snuouosa0 “pw ‘snoiupayksuuad snjosny ‘Yasupam “3 *D ‘sisuauryys 14addv3 -> ‘symoanps 4D ‘snjuna SKMOUOLIYJA] ‘Dasauld DULOJOAN| ‘aay snasKulodag ‘sisuapouvd 4OISDJ ‘snulqns SKuWOIND]|H ‘snoiuospny snamosvnun yf, ‘kid snpiydousady ‘syisia ‘9 ‘Py ‘vINs1jvI DOWD ‘snuDIIUuaUD sndaT ‘S14v]]09 DUOJOYIC) ‘suBsYANBIOU S14aja{UOIsVT ‘snsnfion] SuOK| ‘snuvysv]p “s ‘siasnjod ‘g ‘snjooijuoUl *s ‘snasaulo xa105 (M61 oS€T “N,€S 08S) UBANTING ‘(AA TT o€€1 ‘N,LO oLS) PRAM “(A,8E oSET ‘N, IZ 08S) WUeseaId “(MST oCET 'N,8E 09S) WAR “(ALOE oZET N,6E 09S) MAC “(ALOT ob EI ‘N,ST 08S) SBIsn0g GNV'INIVIA, ‘T SdYOOdA NMONM HLIM SANVW IST NOIDdaENS§ INASHad VXVL TVNNVIA eo——os———so0sss——oono@®@®>@o9maNNM0m0NN em ‘(poq) exe) d1Wapus Jo soUasaid puv vx} JO suoneuiquios onbrun uo paseq oie suorseiqng “uorSezqns yore UT Ind90 Jey) sjeuTUTeUT pue Spurs! 94} SUIPN[OUT ‘eysePY IseayINOG JO} suoIsaigns o1ydessoasoIg DAL “| ATAVI, 1996 MACDONALD AND CooK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA a5) certainty when only the study skin is available. Jackson’s (1928) specimen may actually have been S. cinereus. Heller (1909: 263) stated that S. monticolus was ‘known to occur on Chichagof Island’; however, we have not been able to confirm this. Sorex palustris Richardson, 1828, Water Shrew The distribution of Sorex palustris in Southeast Alaska is poorly documented. The few published records (all mainland localities) are the Chilkat River near Haines (Jackson 1928; Baker 1951; MacDonald and Elliot 1984; KU; USNM), Rudyerd (sic) Bay, Behm Canal (Swarth 1922; UCLA), Salmon River and Texas Creek near Hyder (UAM), Farm Island (UAM), Stikine River (UAM), and Thomas Bay (UAM), suggesting that this species, while occurring widely along the coastal mainland, is highly localized and rarely encountered. Sorex alaskanus Merriam, 1900, Glacier Bay Water Shrew The only published Glacier Bay Water Shrew record is that of two males (USNM 97712, 97713) taken on 12 June 1899 by A. K. Fisher from Point Gustavus (type locality) and described as a new subspecies, S. navigator (= palustris) alaskanus by Merriam (1900). The subsequent elevation of these two specimens to full species status, S$. alaskanus, by Jackson (1926, 1928) has been questioned by Hall (1981) and Junge and Hoffmann (1981; but see Hutterer 1993; Carraway 1995). One additional specimen of S. alaskanus was collected from Bartlett Cove and is housed in the collection at Glacier Bay National Park (GBNP) at Bartlett Cove. The taxono- my and distribution of S. alaskanus needs clarifica- tion because it is represented by only three specimens. canadensis phaeus Peromyscus keeni, Microtus pennsylvanicus admiraltiae, Microtus longicaudus, Ondatra zibethicus, Ursus Sorex cinereus, Myotis lucifugus, M. keenti, Castor canadensis, arctos, Martes americana, Mustela erminea salva, M. vison, Peromyscus keeni, Microtus oeconomus sitkensis, M. Lontra canadensis, Odocoileus hemionus longicaudus, Ursus arctos, Mustela erminea initis, M. vison, Lontra canadensis, Odocoileus hemionus Sorex monticolus, Myotis lucifugus, M. volans, Castor MAMMAL TAXA PRESENT Myotis lucifugus (Le Conte, 1831), Little Brown Myotis The distribution and occurrence of all bat species are poorly documented in Southeast Alaska, although several species apparently reach their northern range limits in this region (Hall 1981). The Little Brown Myotis is undoubtedly the most numerous and widely distributed bat in Southeast Alaska. Although small bats are often seen through- out the region, relatively few specimens substantiate this assumption. Sitka is the type locality for M. 1. alascensis (Miller 1897). Another subspecies, M. 1. pernox, may occur in the upper Lynn Canal region, westward (inferred from Hall 1981). Records from the mainland include those from the Hyder area, Boca de Quadra, Chickamin River, and the Stikine River (Swarth 1911, UAM). Records from Inian Is. (58° 15’N, 134° 16’W), Krestof (58° 15’N, 134° 16’W), Kruzof (58° 15'N, 134° 16’W), Lemesurier (58° 17'N, 136° 05’W), Lowrie (54° 51'N, 133° 32'W), Moser (57° 41'N, 135° 40'W), ISLANDS WITH KNOWN RECORDS Admiralty (57° 40'N, 134° 20'W), The Brothers (57° 18’N, 133° 50’W), Swan (57° 56'N, 134° 14’W), Myriad Is. (57° 38’N, 136° 13’W), Otstoia (57° 33'N, 135° 26’W), Partofshikof (57° 15'N, 135° 36'W), Baranof (56° 45'N, 135° 10’W), Catherine (57° 22'N, 134° 53'W), Chichagof (57° 30'N, 135° 30'W), Yakobi (57° 56’N, 136° 27'W) TABLE |. (Concluded) Z Z 6 fs R a a8 islands are from Admiralty, Baranof, Chichagof, 3) 2B 2 Ze S Mitkof, Prince of Wales, Revillagigedo, and Wrangell a\ 228 7A (@) (2 islands (Streator 1895*; Heller 1909; Swarth 1911; MY) + Ww UAM). 576 Myotis keenii (Merriam, 1895), Keen’s Myotis The occurrence of Myotis keenii in Southeast Alaska has been based on one record from Wrangell, Wrangell Island (Miller 1897; Anderson 1946). The identity of this specimen, which is housed at the USNM, was recently verified (van Zyll de Jong and Nagorsen 1994). A second and third specimen of Keen’s Myotis were collected in 1993 at Turn Creek, Prince of Wales Island (UAM 23338, male) and at Hoonah, Chichagof Island on 11 July 1994 (UAM 29831, male). The latter came from a maternity roost of M. lucifugus (Parker and Cook 1996). The biology of this species is poorly known and the species is represented by some 59 specimens in museum collections (van Zyll de Jong and Nagorsen 1994). Whether this is an indication that this species is actually rare, and thus a species of concern for conservation, is unknown. Nagorsen and Brigham (1993) suggest that so little information is currently available on this species that little can be said about its habitat affinities. No subspecies are currently rec- ognized (van Zyll de Jong 1985). Myotis volans (H. Allen, 1866), Long-legged Myotis For over 80 years, a single specimen of Myotis volans (MVZ 186) was the only record of this species in Alaska. It was collected by C. Littlejohn on 9 June 1907 at Mole Harbor, Admiralty Island (Miller and Allen 1928). Recently, three specimens from Wrangell Island (UAM 19756, 19757; ADF&G location unknown) and one specimen from Prince of Wales Island (UAM 24822) were recorded (West 1994; Parker et al. in preparation*). The original Admiralty Island record was included under the sub- species M. vy. longicrus (Hall 1981). Myotis californicus (Audubon & Bachman, 1842), California Myotis Miller and Allen (1928) reported two specimens of Myotis californicus from Howkan, Long Island. Recently (Parker et al. in preparation*), two M. cali- fornicus skulls were discovered in El Capitan Cave on Prince of Wales Island (UAM 22143, 22144). A third live animal was collected there in February 1992 (UAM 20498). The Alaska specimens have been included under the subspecies M. c. caurinus. Lasionycteris noctivagans (Le Conte, 1831), Silver-haired Bat This species was known for Southeast Alaska from one specimen (AMNH 213141), a juvenile female found roosting on 4 November 1964 in an old gill net hanging in a shed on Canyon Island, Taku River (Barbour and Davis 1969). Three recent specimens (Parker et al. in prepara- tion*), all collected in January, include a specimen *Reference in Document Cited section. THE CANADIAN FIELD-NATURALIST Vol. 110 (UAM 20738) found in a woodpile 15 km south of Wrangell, a specimen (UAM 30100) found dead clinging to the side of a house in Petersburg, and a specimen (UAM 30099) found alive in a house entryway in Ketchikan. Eptesicus fuscus (Beauvois, 1796), Big Brown Bat Manville and Young (1965) and Barbour and Davis (1969) suggest the occurrence of Eptesicus in Southeast Alaska, presumably in the vicinity of Juneau, but we have not been able to confirm this. We are sure of only one Alaska record, that reported by Reeder (1965) from Shaw Creek, west of Delta Junction in central Alaska. There are no known records of this species anywhere near Southeast Alaska in British Columbia (Nagorsen and Brigham 1993). Ochotona collaris (Nelson, 1893), Collared Pika The Collared Pika is known in Southeast Alaska from only White Pass, at the alpine border of Alaska and British Columbia (Osgood 1900; UAM). This species may be present in other mountain areas of the relatively unexplored mainland, particularly in the Haines and Skagway area. Lepus americanus Erxleben, 1777, Snowshoe Hare Snowshoe Hares have a very limited northern mainland distribution in Southeast Alaska occurring in the Chilkat Valley near Haines, and at Dyea in the vicinity of Skagway (Streator 1895*; Bailey 1920*; Jewett 1941*; UAM; MVZ). Hares are occasionally found in the Taku River valley toward the Canada border (Streator 1895*; J. Owens personal communi- cation 1994). Home (1973*) reported sightings of hares at Glacier Bay and on the Alsek River. A report of hares at the mouth of the Stikine River (Manville and Young 1965) has not been substantiated. Snowshoe Hares now present on Douglas Island were introduced there from Haines stock “a few years previous” (Bailey 1920*; also Wenrich 1922*). The hares on the mainland near Juneau were most likely derived from these animals. In 1923 and 1924, the Alaska Game Commission released Snowshoe Hares from Washington stock to Pt. Retreat, Admiralty Island; Otstoia Island, Peril Strait; and Smeaton Island, Behm Canal. Stock from the Anchorage area were also released in 1924 on Cape Island, Prince of Wales Island; and, on Village Island, Zimovia Strait. All these transplant attempts were considered failures (Burris and McKnight 1973). Oryctolagus cuniculus (Linnaeus, 1758), European or Domestic Rabbit Several residents of Ketchikan have reported (in 1995) the presence of feral rabbits on Betton Island, Clover Pass. Their current status is unknown. 1996 Marmota caligata (Eschscholtz, 1829), Hoary Marmot The Hoary Marmot occurs throughout Southeast Alaska’s mainland coast (tide water up to alpine), with reports from Yakutat, Glacier Bay, White Pass, Juneau area, Taku River, Port Snettisham, Swan Lake, Chickamin River, Boca de Quadra, and Hyder (Osgood 1900; Heller 1909; Swarth 1911; Howell 1915; Hoffmann et al. 1979; UAM). Anderson (1946) and Dufresne (1946) both mention Hoary Marmots along Portland Canal. Specimens collected close to the mainland in 1992 and 1995 on Douglas Island are the only island records for this species in Southeast Alaska. Marmots at Glacier Bay are considered an endem- ic subspecies, M. c. vigilis (Heller 1909; Howell 1915). All others in Southeast Alaska are included under the name M. c. caligata (Hall 1981). Marmots from Juneau were transplanted to the Klawock area, Prince of Wales Island, in 1930 by the Alaska Game Commission (Burris and McKnight 1973). Their pre- sent status is unknown. Of biogeographic interest was the discovery in 1992 of a fossil marmot incisor, dated at > 44 500 yr B.P., and a marmot molar from the same locality in 1994 in a cave on Prince of Wales Island (Heaton 1995). Spermophilus parryii (Richardson, 1825), Arctic Ground Squirrel The Arctic Ground Squirrel is a Holarctic species with a limited distribution in Southeast Alaska. Its known occurrence there is currently restricted to the coastal mountains north of Skagway (Osgood 1900; UAM). The type locality of that subspecies, S. p. plesius, is Lake Bennett, British Columbia, not far from the Alaska border (Osgood 1900; Howell 1938). As with Ochotona collaris and several other alpine species, further inventory work in the relative- ly unexplored mountains along the region’s main- land may change our understanding of the distribu- tion of this species. Tamiasciurus hudsonicus (Erxleben, 1777), Red Squirrel Tamiasciurus hudsonicus is found in suitable for- est habitats throughout the mainland of Southeast Alaska (including Douglas Island) and, before the days of game transplanting, on those islands in the Alexander Archipelago south of Frederick Sound and east of Clarence Strait, including Annette, Betton, Etolin, Gedney, Grant, Gravina, Hassler, Kuiu, Kupreanof, Mitkof, Read, Revillagigedo, Sullivan, Tatoosh, Tongass, Vank, Woewodski, Woronkofski, Wrangell, and Zarembo islands. (Osgood 1900; Heller 1909; Swarth 1911, 1921, 1922; KU; MVZ; UAM; UAM sight records). Squirrels from Glacier Bay to Glacier and White Pass north of Skagway were designated T. h. petulans MACDONALD AND CoOoK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA a7 by Osgood (1900) (type locality = White Pass, south- ern Alaska). From Lynn Canal south, Swarth (1921) considered them a different subspecies, 7. h. picatus (type locality = Kupreanof Island, 25 mi. south of Kake Village, at southern end of Keku Straits). The status of the Red Squirrel as a native resident of Admiralty Island is unclear. Neither Heller (1909) nor Swarth (1921) indicated their presence on this island. Dufresne (1946), however, claimed that the Red Squirrel was indigenous to Admiralty Island. Burris and McKnight (1973) indicated that this ani- mal is “scarce or nonexistent” there. Most evidence suggests that this species was introduced to the north end of the island since the late 1940s or early 1950s. In 1993, the Red Squirrel was reported for the first time as far south as Angoon (R. Carstensen, personal communication 1994). In 1995, a specimen was taken at Hood Bay (UAM). Red Squirrels from the Juneau area were success- fully transplanted to Baranof and Chichagof islands in 1930 and 1931 (Burris and McKnight 1973). Since then, Red Squirrels have been found on Inian, Kruzof, Moser, Partofshikof, Yakobi islands, and on islands in Sitka Sound (UAM; L. Johnson, personal communication, 1994). An apparently unsuccessful transplant occurred on Prince of Wales Island (Fay and Sease 1985*). Glaucomys sabrinus (Shaw, 1801), Northern Flying Squirrel The distribution in Southeast Alaska of this squir- rel of mature-forest habitats is poorly understood and specimen records are few. Flying squirrels are known from the mainland in the Glacier Bay area, near Haines, Chilkoot Inlet, Taiya River, Taku River, Bradfield Canal, and the Cleveland Peninsula and in the Alexander Archipelago from the Barrier Islands (off SW Prince of Wales Island), Betton Island (trapper report), El Capitan Island, Etolin Island, Kosciusko Island (sight record by local resi- dent), Mitkof Island, Prince of Wales Island, Revillagigedo Island, Tuxekan Island, and Wrangell Island (Streator 1895*; Osgood 1900, 1905; Swarth 1911; Walker 1920*; Howell 1934; Fay and Sease 1985*; GBNP; MVZ; UAM). Helm Bay, Cleveland Peninsula, is the type locali- ty for G. s. zaphaeus (Osgood 1905). Flying squirrel populations restricted to Prince of Wales Island are currently recognized as an endemic subspecies, G. s. griseifrons. Manville and Young (1965) cite reports of G. s. alpinus six and nine miles north of Juneau. As is the case with much of the original taxonomic work on the mammals of Southeast Alaska, the description of G. s. griseifrons is based on few speci- mens (Howell 1934). However, preliminary analysis of MtDNA sequence data is revealing fixed basepair differences between subspecies (J. Demboski, per- sonal communication 1995). Given the projected timber harvests on Prince of Wales and adjacent 578 islands in the next ten years, the status of this sub- species needs immediate review. Castor canadensis Kuhl, 1820, Beaver Beaver occur today in suitable habitat along the mainland and on most of the islands of Southeast Alaska, except perhaps Coronation, Warren, Forrester and other small, remote islands. They are especially abundant along the major rivers of the mainland. Around the turn of this century heavy trapping pres- sures temporarily reduced or eliminated Beaver from a number of localities (Gray 1915*; Bailey 1920*). Preserved specimens of Beaver from Southeast Alaska are few. In 1915, Gray noted Beaver were then absent from Etolin, Wrangell and the smaller islands in that area. Heller (1909) reported very old signs of beaver on Chichagof Island. Gary (1915*) was told that Beaver had previously occurred on Chichagof and Baranof islands. In a 1922 Game Condition report, W. Wenrich (1922*) commented, “It was said that at one time beaver were numerous [on Chichagof I.], but for some reason or other there are none left now.” Ten animals from Prince of Wales Island were suc- cessfully “re-introduced” to Baranof Island in 1927 (Burris and McKnight 1973). It remains unclear if Beaver were introduced to Kruzof Island in 1925 (Burris and McKnight 1973). Heller (1909) described, based on six specimens (three adult), the endemic subspecies, C. c. phaeus (type locality = Pleasant Bay) from Admiralty Island and, presumably Chichagof and Baranof islands as well. All others are considered C. c. belugae (Hall 1981). A revision of Beaver in Southeast Alaska with adequate specimens and modern techniques is very much needed (Hafner et al. in press). Peromyscus keeni (Rhoads, 1894), Keen’s Mouse Keen’s Mouse (formerly P. maniculatus and P. sitkensis; Hogan et al. 1993) is widely distributed throughout Southeast Alaska. This species occurs on the mainland at Glacier Bay from Tlingit Point and Muir Inlet (Home 1973*; USNM) eastward to Haines and Skagway southward (Osgood 1900; Heller 1909; Swarth 1911; MVZ; UAM; USNM). In the Alexander Archipelago it is known to occur on the following islands: Admiralty, Anguilla, Annette, Baker, Baranof, Betton, Brothers (East and West), Cat, Chichagof, Cone, Coronation, Dall, Douglas, Duke, Esquibel, Etolin, Forrester, Gravina, Heceta, Inian (“Big”), Kosciusko (Swarth 1911: 156, Figure 3), Kruzof, Kuiu, Kupreanof, Long, Lowrie, Lulu, Marble, Mary, Mitkof, Moser, Noyes, Orr, Partofshikof, Prince of Wales, Revillagigedo, San Fernando, Santa Rita, Shrubby, Swan, Suemez, Sukkwan, Tuxekan, Vank, Warren, Woewodski, Woronkofski, Wrangell, Yakobi, and Zarembo islands (Heller 1909; Osgood 1909; Swarth 1911; McGregor 1958; Harris 1968; Van Horne 1981; KU; MVZ; UAM; USNM). THE CANADIAN FIELD-NATURALIST Vol. 110 Two species and five subspecies of Peromyscus were once recognized in this region (Hall 1981). Recently, Hogan et al. (1993) found chromosome, allozyme, and mitochondrial DNA variation support- ing the hypothesis that Southeast Alaska subspecies of P. maniculatus and P. sitkensis are conspecific. They included them under P. keeni, which was recog- nized as the senior synonym. Currently described sub- species for the region (after Hall 1981) include P. k. algidus, P. k. macrorhinus, P. k. hylaeus (type locali- ty = Hollis, Kasaan Bay, Prince of Wales Island; Osgood 1908), P. k. oceanicus (type locality = Forrester Island), and P. k. sitkensis (type locality = Sitka, Baranof Island). These are in need of revision. Several pelage variants have been collected in Southeast Alaska. Of the adult specimens collected on San Fernando Island, 21 (67%) have partially or entirely brown venters. Two specimens on Shrubby Island and one on Etolin Island exhibited Horner’s black mutation (Conroy et al. 1993*). Neotoma cinerea (Ord, 1815), Bushy-tailed Woodrat The only published specimen records of Neotoma cinerea in this region are those reported by Shaw (1962) from the Taku River (specimen collected in 1940) and the Unuk River (1925). A woodrat col- lected at the head of Lake Bennett, British Columbia, and one seen at Glacier inside Alaska (Osgood 1900) indicate a mainland Alaska distribution at least this far north. Specimens have been taken on nunataks in the Juneau Ice Fields (UPS). Dufresne (1946: 138-139) stated that woodrats were “fairly common near the head of Portland Canal and along the Unuk River. It has also been reported from the Stikine and Taku River watersheds.” Hall (1981), in agreement with Cowan and Guiguet (1965) and Youngman (1975), placed Alaska woodrats under the subspecies N. c. occidentalis. Clethrionomys rutilus (Pallas, 1779), Northern Red-backed Vole Clethrionomys rutilus is currently believed to occur along the northern mainland of Southeast Alaska east and south to Juneau. VAM has recent specimens from Douglas Island, an island in close proximity to the mainland near Juneau. An unpublished revision of Clethrionomys rutilus by Antell (1987) extended the distribution of the subspecies C. r. glacialis to include all populations from Yakutat south to Bartlett Cove (type locality = Glacier Bay; Orr 1945). Populations in the Haines/Skagway area south to Juneau are included under the subspecies C. r. dawsoni. Clethrionomys gapperi (Vigors, 1830), Southern Red-backed Vole Clethrionomys gapperi is believed to inhabit the mainland of Southeast Alaska south of Juneau and on a few islands south of Stikine Strait and east of | 1\ iN ! | | 1996 Clarence Strait, namely Bell, Black, Etolin, Hassler, Read, Revillagigedo, and Wrangell islands (Hall 1981; UAM). Swarth (1922) reported C. rutilus on the Stikine River in British Columbia, downstream as far as Dokdaon Creek. About 45 km further downriv- er at Flood Glacier, he encountered Evotomys wrangeli (= Clethrionomys gapperi). Four subspecies have been recognized in this region (Hall and Cockrum 1952, 1953): C. g. stiki- nensis (Stikine River south from the Flood Glacier, but not including Sergief Island, south to Helm Bay, Cleveland Peninsula); C. g. phaeus (type locality = Boca de Quadra); C. g. wrangeli (type locality = Wrangell Island); and C. g. solus (type locality = Loring, Revillagigedo Island). A possible fifth sub- species is C. g. saturatus, from the upper Portland Canal region. Interspecific and intraspecific taxonomic relation- ships of these species of Clethrionomys should be reviewed. Bee and Hall (1956) and Youngman (1975) considered them conspecific. More speci- mens are needed from areas of contact, specifically along the Stikine River, between Dokdaon Creek and Flood Glacier, and along Southeast’s coastal main- land, from Haines south. Microtus pennsylvanicus (Ord, 1815), Meadow Vole The Meadow Vole, Microtus pennsylvanicus, has a restricted distribution in Southeast Alaska, occur- ring along the mainland valleys of the Chilkat, Taku, and Stikine rivers (including Sergief Island [= “Browns Island”; USNM]); and on Admiralty, Mitkof, and Vank islands (Heller 1909; Dale 1940; Baker 1951; UAM). Dale (1940) and Hall (1981) define three subspecies of Microtus pennsylvanicus for Southeast Alaska: M. p. admiraltiae (Admiralty Island; type locality = Windfall Harbor), M. p. alcorni (mainland valleys of the Chilkat River), and M. p. rubidus (Taku and Stikine river valleys). An analysis of the species by Snell and Cunnison (1983) found no conspicuous groupings, suggesting that subspecific designations may be inappropriate. The identification of a specimen of M. pennsyl- vanicus (as M. drummondii) reportedly taken at Fort Wrangell (= townsite of Wrangell, Wrangell Island) by Mrs. S. H. Young in 1887 (USNM 00190736) was confirmed by SOM; however, this species has not been taken anywhere on the island in subsequent collecting efforts. Microtus oeconomus (Pallas, 1776), Tundra Vole The Tundra Vole inhabits the northern mainland of Southeast Alaska from Yakutat Bay south and east to Glacier Bay and White Pass in the vicinity of Skagway (Hall 1981; Antell 1987; UAM). This species also occurs on Baranof, Chichagof, and Lemesurier islands in the northwestern Alexander Archipelago (Merriam 1897; Heller 1909; UAM). MACDONALD AND COOK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA Dy) Those islands mark the southern extent of the distri- bution of this species in North America. A revision of this species by Antell (1987) did not corroborate an earlier revision by Paradiso and Manville (1961). Antell (1987) described a new sub- species, M. o. littoralis, for populations of Tundra Vole from Bartlett Cove northward along the coast (but not including the area immediately north of Yakutat Bay, which Antell considered M. 0. yaku- tatensis; type locality = north shore of Yakutat Bay). Antell (1987) also concluded that populations of this species (previously included under M. 0. macfarlani) from Haines Junction, Yukon, and in the vicinity of Skagway warrant recognition as a new, and as yet undescribed subspecies. Tundra Voles on Baranof and Chichagof islands, which were not analyzed by Antell (1987), are included in M. o. sitkensis (type locality = Sitka, Baranof Island). Microtus longicaudus (Merriam, 1888), Long-tailed Vole The Long-tailed Vole, Microtus longicaudus, is widely distributed throughout Southeast Alaska. It occurs in suitable herbaceous habitat throughout the coastal mainland, and on islands of the Alexander Archipelago as follows: Admiralty, Anguilla, Chichagof, Dall, Etolin, Hoot, Kosciusko, Kuiu, Kupreanof, Marble, Mary, Mitkof, Moser, Noyes, Orr, Owl, Prince of Wales, Revillagigedo, Stevenson, Suemez, Sullivan, Sukkwan, Tuxekan, Wrangell, and Zarembo islands (Bailey 1900; Osgood 1900; Heller 1909; Swarth 1911, 1933; McGregor 1958; Harris 1968; Van Horne 1982; Antell 1987; MVZ; UAM; USNM). Swarth (1911) reported seeing vole runways, presumably of this species, on Annette Island. Similarly, UAM collec- tors (SOM and D. Pengilly, June 1983) observed runways on Santa Rita and Baker islands. It is curi- ous that Long-tailed Voles have not been document- ed on Baranof Island, despite its close proximity to Chichagof Island populations. Two subspecies have been recognized in this region: M. I. littoralis (type locality = “Shakan”) and M. I. vellerosus (Hall 1981; Antell 1987). Microtus coronarius Swarth, 1911, Coronation Island Vole Microtus coronarius was described as a distinct species by Swarth (1911) from specimens collected on Coronation (type locality = Egg Harbor), Warren and Forrester islands (Swarth 1933). This taxon may be a subspecies of the more widely distributed M. longicaudus. Swarth (1933) distinguished M. coro- narius from M. longicaudus by its larger body size. However, large-bodied voles are not restricted to these three islands, as illustrated by voles (UAM) from other Southeast Alaska localities that fit into the range of body measurements of M. coronarius listed by Swarth (1911, 1933). The close relationship of M. coronarius to M. longicaudus was pointed out by Hall (1981: 809 and Map 457) and needs to be | \\ Vol. 110 | | 580 THE CANADIAN FIELD-NATURALIST Mus musculus Linnaeus, 1758, House Mouse examined further. Warren, Coronation, and Forrester Introduced from Europe, this mouse is commensal | island populations should be examined karyotypical- ly. A molecular systematic study that includes speci- mens from throughout the geographic range of Long-tailed Voles would help clarify the relationship of M. coronarius and M. longicaudus. Such an anal- ysis would provide insight into the significance of the extensive chromosomal variation reported for this species group (Judd and Cross 1980; Musser and Carleton 1993). Ondatra zibethicus (Linnaeus, 1766), Muskrat Muskrats have a limited distribution in Southeast Alaska. Locality records include Yakutat Bay, the Haines area, Taku River (trapper report), Stikine River, Admiralty Island, and Revillagigedo Island. (Swarth 1911; 1922; Bailey 1920*; Hall and Cockrum 1953; Burris and McKnight 1973; MVZ; UAM; USNM). Muskrats have been reported from Kuiu Island, Kupreanof Island, and Mitkof Island (S. Blatt personal communication 1994). There were unsuccessful attempts in 1929 to trans- plant Muskrats from Haines to Klawock Lake, Prince of Wales Island (Burris and McKnight 1973). Synaptomys borealis (Richardson, 1828), Northern Bog Lemming Little is known about this species in Southeast Alaska. Specimens are few, being limited to mainland localities that include the Chilkat River, Taiya River, Turner Lake, Crescent Lake, Port Snettisham, Sumdum, Port Houghton, Thomas Bay, Stikine River, Anan Creek, Unuk River, Chickamin River, Quadra Lakes, and Foggy Bay (Howell 1927; UAM). The type locality for the Southeast Alaska subspecies, S. b. truei, is Wrangell, Wrangell Island (Howell 1927). The only island records are from Wrangell on Wrangell Island (Merriam 1896; Howell 1927) and recently from estuarine meadows on Betton, Back, and Gravina islands (UAM), near Revillagigedo Island. The presence of bog lemmings at the latter three island localities suggest a high probability of this species occurring on Revillagigedo Island as well, although this has yet to be documented. Rattus norvegicus (Berkenhout, 1769), Norway Rat Rattus norvegicus is a commensal, non-native rat associated with urban areas. Preserved specimens are few and include specimens from Petersburg, Mitkof Island; Ketchikan, Revillagigedo Island; and Juneau on the mainland (USNM). They have been seen at the landfill near Sitka (by SOM in 1982), where they are very common at times (L. Johnson, personal com- munication 1994). Hall (1981) listed one subspecies for North America, R. n. norvegicus. Nagorsen (1990) indicated that the taxonomy of North American populations is obscured by multiple introductions. with man, inhabiting urban and agricultural areas, pri- marily. Information on its distribution in Southeast Alaska is nearly non existent and no preserved speci- mens are known. C. P. Streator, in his notes from Juneau in August 1895*, caught three Mus in the for- est near town; he stated that this species was getting common. The form inhabiting northern North America, including Alaska, was believed to be derived from the commensal race M. m. domesticus (Schwarz and Schwarz 1943). Variation among North American populations has not been studied. This subspecies is considered a distinct species by some (Marshall and Sage 1981). Zapus hudsonius (Zimmermann, 1780), Meadow Jumping Mouse Until recently, there were only three locality records for Zapus hudsonius in Southeast Alaska: Yakutat Bay, Chilkat River near Haines (Baker 1951; Hall 1981; UAM) and, surprisingly, a single pregnant female from Portage Cove, Revillagigedo Island (Swarth 1911), 480 km south of Haines. In 1993 and 1995, eight additional Z. hudsonius were collected on Revillagigedo Island: seven at Portage Cove and one at the head of Orchard Lake (UAM). The type locality for the race in Southeast Alaska, Z. h. alascensis, is Yakutat Bay (Merriam 1897). The close morphologic similarity between Z. hudsonius and Z. princeps, coupled with the disjunct distribution exhibited by Z. hudsonius on Revillagigedo Island, suggests that the status and relationship of all jumping mice in this region need to be examined more closely. Zapus princeps J. A. Allen, 1893, Western Jumping Mouse The presence of Z. princeps in.Southeast Alaska was based on one adult male jumping mouse captured by Swarth (1911) on the Taku River. The subspecies of Z. princeps in Southeast Alaska is Z. p. saltator, according to Krutzsch (1954). Jones (1981) consid- ered this taxon to be indistinct from Z. p. princeps. Jumping mice presumed to be this species from dental/cranial characters (Krutzsch 1954) have been collected (all UAM) at mainland sites as follows: Salmon River near Hyder; Gwent Cove, Portland Canal; Kirk Point, Foggy Bay; Wolf Cabin, Chickamin River; Unuk River; Farm Island, Stikine River; and near Canyon Island, Taku River. Erethizon dorsatum (Linnaeus, 1758), Porcupine Porcupines occur throughout the mainland of Southeast Alaska (including Douglas Island) and in the Alexander Archipelago south of Frederick Sound, including Etolin, Hassler, Kupreanof, Mitkof, Revillagigedo, and Wrangell islands (Swarth 1911; Webster 1949; Bailey 1971; Eglitis and Hennon 1996 1986; UAM; L. Carson, personal communication 1994; M. Brown, personal communication 1994). Two subspecies have been recognized in Southeast Alaska: E. d. myops Merriam from the Glacier/White Pass region (Osgood 1900), and probably all popula- tions west to Yakutat Bay (see Hall 1981); and E. d. nigresens from Lynn Canal southward (Hall 1981). Further information is needed to clarify the distribu- tion of this species in the Alexander Archipelago. Canis latrans Say, 1823, Coyote The Coyote is an infrequent visitor to the river val- leys along the southern mainland of Southeast Alaska and a probable resident in some areas along the main- land north from the Taku River. The only island record we are aware of is an adult female Coyote trapped on Mitkof Island near Dry Straits in the win- ter of 1983-1984 (C. Land, personal communication 1994). To our knowledge, no specimens from Southeast Alaska have been preserved. A specimen reported from the Alsek River by J. A. Allen (1908; and given as an Alaska record by Hall 1981) was that of a “Coyote killed near Whitehorse, on the Alsek River, Alaska in February 1907.” We have not been able to locate this specimen and are unsure of exactly where it was taken as Whitehorse is farther north on the Yukon River in Yukon. Canis lupus Linnaeus, 1758, Wolf The Wolf inhabits the entire mainland of Southeast Alaska and islands in the Alexander Archipelago south of Frederick Sound, excluding Coronation, Forrester and, undoubtedly, some of the other small, more isolated islands without an adequate prey base. We are unaware of any records of this species ever occurring on any of the islands north of Frederick Sound. Museum specimens (UAM, except as noted) from the Alexander Archipelago are as follows: Baker, Bell, Conclusion (USNM), Duck, Etolin, Grief, Keene, Kuiu, Kupreanof, Mitkof, Prince of Wales, Read, Revillagigedo, Suemez, Woewodski, Wrangell, and Zarembo islands. The subspecies C. J. ligoni (type locality = head of Duncan Canal, Kupreanof Island) is restricted to Southeast Alaska (Hall 1981; Pederson 1982). Molecular genetic research is currently underway at the Institute of Arctic Biology, UAF, to examine the status of this taxon (G. Shields, personal communica- tion 1994). Vulpes vulpes (Linnaeus, 1758), Red Fox The Red Fox is an infrequent visitor to the river valleys along the southern mainland, and a probable uncommon resident along the northern mainland from the Taku River, north. We know of no reports of this species occurring naturally on any island in the Alexander Archipelago. The only specimen record we are aware of is one in the MVZ from Yakutat. We have not been able to locate a supposedly preserved MACDONALD AND Cook: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 581 specimen of a male Red Fox found poisoned near Haines by E. P. Walker on 9 February 1914 (Field Catalog, USNM); he considered this species “fairly common” there. The fossil jaw of a Red Fox was recently discovered in a cave on Prince of Wales Island (J. Baichtal, personal communication). Red Foxes were introduced for commercial reasons on Cleft, Dry, Kupreanof, Passage, and Sokoi islands between 1894 and 1929; none were successful (Bailey 1993). The subspecies occurring in Southeast Alaska has not been identified (Hall 1981), but it may be V. v. abietorum, the subspecies found in nearby British Columbia. Alopex lagopus (Linnaeus, 1758), Arctic Fox Arctic Foxes (“blue” morph from the Aleutians and Pribilof islands) were commercially stocked on over 170 islands in the Alexander Archipelago of Southeast Alaska between 1899 through 1929 (Bailey 1993: 25-29, for a complete listing). The industry col- lapsed in the 1930s; no Arctic Foxes remain in the region. Ursus americanus Pallas, 1780, Black Bear The Black Bear occurs along the entire mainland coast of Southeast Alaska and on most of the islands in the Alexander Archipelago south of Frederick Sound. We are unaware of Black Bears on Warren, Coronation, or Forrester islands, or on any island north of Frederick Sound except Pleasant Island in Icy Strait (K. Rutledge, personal communication 1994). Mid-Wisconsin and Late-Wisconsin/early Holocene fossil remains of Black Bear have been found in caves on Prince of Wales Island (Heaton and Grady 1992, 1993; Heaton 1995). Ursus americanus pugnax (type locality = Rocky Bay, now Bobs Bay, Dall Island) is the subspecies currently recognized for most of Southeast Alaska (Hall 1981). Ursus a. emmonsii (type locality = St. Elias Alps, near Yakutat Bay) is a race that includes the “glacier bear” color morph and it occurs along the mainland from Lynn Canal west and north to Prince William Sound (Dufresne 1946; Hall 1981). A taxo- nomic revision is needed. Ursus arctos Linnaeus, 1758, Brown Bear Brown Bears occur along the entire coastal main- land of Southeast Alaska (especially along the major river systems; ADF&G 1973), and on most of the islands of the Alexander Archipelago north of Frederick Sound. Brown Bears are occasionally seen but have never become established on islands close to the mainland south of Frederick Sound, specifically Etolin, Mitkof, Revillagigedo, and Wrangell (Jewett 1941*; Klein 1965; S. Blatt, personal communication 1994; J. Gustafson, personal communication 1994). Of biogeographic interest (Klein 1965) is the recent discovery of U. arctos skeletal remains in limestone caves on Prince of Wales Island (Heaton 582 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 2. List of the Recent land mammals of Southeast Alaska, including the number of taxa (subspecies) and the number of these that are endemic to the region, number of museum specimens (museums searched are listed in Methods and Materials), number of islands of known occurrence, and their current conservation status. NUMBER NUMBER OF OF ISLANDS OF NUMBER SPECIMENS KNOWN OF TAxA/ (UAM + OCCURRENCE CONSER- MAMMAL SPECIES NUMBER OTHER (NATIVE + VATION Or SOUTHEAST ALASKA ENDEMIC MUSEUMS) INTRODUCED STATUST INSECTIVORA — Shrews Soricidae Sorex cinereus, Masked Shrew 2/0 356 + 277 D3) = Sorex monticolus, Dusky Shrew 4/2 497 + 622 52 -— Sorex palustris, Water Shrew 1/0 9+4 0 —— Sorex alaskanus, Glacier Bay Water Shrew 1 0+3 0 ESA-C CHIROPTERA — Bats Vespertilionidae Myotis lucifugus, Little Brown Myotis 2/0 85 + 28 cosmopolitan — Myotis keenii, Keen’s Myotis 1/0 2+1 3 B.C. Red List Myotis volans, Long-legged Myotis 1/0 2+1 2 — Myotis californicus, California Myotis 1/0 34+4 2 —— Lasionycteris noctivagans, Silver-haired Bat 1/0 3+1 3 - LAGOMORPHA — Pikas and Hares Ochotonidae Ochotona collaris, Collared Pika 1/0 2+0 0) — Leporidae Lepus americanus, Snowshoe Hare 1/0 04+5 O+1 a= Oryctolagus cuniculus, Domestic Rabbit 1/0 0+0 O+1 non-native RODENTIA — Rodents Sciuridae Marmota caligata, Hoary Marmot Qi 6+ 12 1 IUCN-IK Spermophilus parryii, Arctic Ground Squirrel 1/0 2+5 0 — Tamiasciurus hudsonicus, Red Squirrel PLA 31 +74 21+8 — Glaucomys sabrinus, N. Flying Squirrel PLR AI 46 + 12 12 IUCN-IK Castoridae Castor canadensis, Beaver Pay fe 5+3 cosmopolitan IUCN-IK Muridae Peromyscus keeni, Keen’s Mouse o/3 1350 + 783 Sil — Neotoma cinerea, Bushy-tailed Woodrat 1/0 Oni 0 — Clethrionomys rutilus, N. Red-backed Vole Pui fi 34 + 427 1 — Clethrionomys gapperi, S. Red-backed Vole 4/4 154 + 189 7 IUCN-IK Microtus pennsylvanicus, Meadow Vole 31 40 + 96 3 IUCN-IK Microtus oeconomus, Tundra Vole Bye?) 40 + 231 3} IUCN-IK Microtus longicaudus, Long-tailed Vole 2/0 219 + 436 25 ae Microtus coronarius, Coronation Island Vole efit 20 + 29 3 IUCN-IK Ondatra zibethicus, Muskrat 1/0 17+9 Su — Synaptomys borealis, N. Bog Lemming 1/0 23+ 14 4 (5?) — Rattus norvegicus, Norway Rat 1/0 0+6 0+3 non-native Mus musculus, House Mouse 1/0 0+0 0+? non-native Dipodidae Zapus hudsonius, Meadow Jumping Mouse 1/0 26 + 37 I = Zapus princeps, Western Jumping Mouse 1/0 32-7 0 — Erethizontidae Erethizon dorsatum, Porcupine 2/0 2+ 14 7 = CARNIVORA — Carnivores Canidae Canis latrans, Coyote 1/0 0+0 | (vagrant) = Canis lupus, Wolf iy 310+ 61 Islands S. IUCN-V Frederick Sound 1996 MACDONALD AND COOK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA TABLE 2. (Concluded) OF SOUTHEAST ALASKA extripated Lontra canadensis, River Otter Lynx canadensis, Canada Lynx Rangifer tarandus, Caribou NUMBER OF TAxA/ MAMMAL SPECIES NUMBER ENDEMIC CARNIVORA — Carnivores Canidae Vulpes vulpes, Red Fox 1/0 Alopex lagopus, Arctic Fox — Ursidae Ursus americanus, Black Bear Dil Ursus arctos, Brown Bear BNA (0) Procyonidae Procyon lotor, Raccoon — Mustelidae Martes americana, Marten 2/0 Martes pennanti, Fisher 1/0 Mustela erminea, Ermine 6/5 Mustela nivalis, Least Weasel = Mustela vison, Mink Disa Gulo gulo, Wolverine 1/0 1/0 Felidae Puma concolor, Mountain Lion 1/0 1/0 ARTIODACTYLA — Ungulates Cervidae Cervus elaphus, Elk 2/0 Odocoileus hemionus, Mule Deer 2/0 Alces alces, Moose 2/0 1/0 Bovidae Ovis dalli, Dall’s Sheep 1/0 Oreamnos americanus, Mountain Goat 1/0 583 NUMBER NUMBER OF OF ISLANDS OF SPECIMENS KNOWN (UAM + OCCURRENCE CONSER- OTHER (NATIVE + VATION MUSEUMS) INTRODUCED STATUST 0+1 0) — 0+0 0+ 182 non-native 24+ 213 Islands S. CITES-II of Frederick Sound ESA-C B.C. Blue List 33 + 603 Islands N. of CITES-II Frederick Sound; vagrant S. 0+0 0+3 non-native extripated 347 + 16 9+7 = 1+0 0) B.C. Blue List 10+ 73 cosmopolitan ESA-C none — — 396 + 175 cosmopolitan — 44+5 6 CITES-II 111+ 26 cosmopolitan CITES-I 1+0 1 (vagrant) CITES-II 1+2 0 CITES-II 0+0 0+2 non-native 8 + 94 cosmopolitan — 0+0 5) — none 0) — 0+0 0 — 14+4 0+2 — +Conservation status of these taxa: ESA — Endangered Species Act 1973 (United States Fish and Wildlife Service 1993); IUCN — International Union for the Conservation of Nature, Species Survival Committee (Hafner et al. in press; BC — British Columbia, Canada (1993) Red and Blue Lists (Harper et al. 1994); CITES — Convention on International Trade of Endangered Species, Appendix II. 1995), ranging in age from mid-Wisconsin (35 365 + 800 yr BP) to early Holocene. This island is south of Frederick Sound, where today only Black Bears occur. Brown Bear bones were also recently found in cave deposits on Dall Island (S. Lewis, personal communication 1995). Rausch (1963) included all of the 13 species names originally proposed for Southeast Alaska Brown Bears by Merriam (1918) under the trinomial, U. a. horribilis. Hall (1984) recognized three sub- species for the Southeast Alaska region: U. a. dalli (Yakutat Bay-Dry Bay), U. a. sitkensis (islands north of Frederick Sound, adjacent northern mainland), and U. a. stikeenensis (southern mainland, Canada, Washington). Recent studies of mitochondrial DNA variation (Cronin et al. 1991, Talbot and Shields 1996) indicate that Brown Bears on islands in the Alexander Archipelago are distinct from populations of U. arctos on the Southeast mainland and else- where in Alaska. Procyon lotor (Linnaeus, 1758), Raccoon Raccoons are not native to Southeast Alaska. Natural populations occur as close as Kingcome Inlet, 584 and Vancouver Island and adjacent islands of coastal British Columbia, Canada; they have been introduced to the Queen Charlotte Islands (Nagorsen 1990). Eight melanistic Raccoons from Indiana were released on Singa Island, Sea Otter Sound, in October 1941, spreading to nearby El Capitan and several other islands in this area (Scheffer 1947; Burris and McKnight 1973). Their current status is unknown; however, we know of no recent reports and consider them extirpated. In 1950, Raccoons of unknown origin were released or escaped on Japonski Island near Sitka, with a few eventually spreading to nearby Baranof Island (Elkins and Nelson 1954; Burris and McKnight 1973). Individuals were occasionally seen around the dump at the Sitka airport on Japonski Island up until the early 1970s when the airport was extended and the dump covered. None have been reported in the Sitka area since (L. Johnson, personal communication 1994). Martes americana (Turton, 1806), Marten Marten inhabit forest habitats along the entire coastal mainland of Southeast Alaska and on islands in the Alexander Archipelago in close proximity to the mainland. Natural populations occur on Admiralty (but see below), Etolin, Gravina, Kuiu, Kupreanof, Mitkof, Revillagigedo, Woewodski, and Wrangell islands (Streator 1895*; Swarth 1911; Gray 1915*; Walker 1920*; Dufresne 1946; MVZ; UAM; USNM; S. Blatt, personal communication 1994; L. Carson, personal communication 1994). R. W. Flynn (in Suring et al. 1992*) questioned the natural occurrence of Marten on Admiralty Island, suggesting that their presence there may be the result of an unreported transplant. If true, it would have been done prior to 1903, when W. H. Osgood and N. Hollister (1903*) of the Biological Survey reported that small numbers of Marten occurred on Admiralty Island. Heller (1909: 246) mentioned Marten on Admiralty Island, as did Bailey (1920*) and Cahalane (1940*). Dufresne (1946) stated that Marten were native to this island; however, Gray (1915*) stated they were not found there. Allen Hasselborg, a long-time resident of Admiralty Island, collected Marten specimens from the island in 1910 and 1913 (MVZ), and in 1915 (USNM). Before game laws early this century, Marten num- bers in the region were greatly reduced by overtrap- ping (Gray 1915*; Bailey 1920*; Wenrich 1922*). In 1934, Marten from Behm Canal and Thomas Bay on the mainland were successfully introduced on Prince of Wales Island and Baranof Island. Between 1949 and 1952, Marten were successfully introduced on Chichagof Island with stock taken from Baranof Island, Revillagigedo Island, the Stikine River drainage, Wrangell Island, Mitkof Island, and near Anchorage (Elkins and Nelson 1954; Burris and McKnight 1973). In addition, THE CANADIAN FIELD-NATURALIST Vol. 110 UAM has records of Marten taken in the vicinity of Baranof Island from Kruzof Island, Otstoia Island, Catherine Island, and Yakobi Island. Undoubtedly, all were from undocumented transplants. Clark et al. (1987) separated 14 subspecies of North American Marten into two groups: “ameri- cana” and “caurina.” Hagmeier (1958) and Anderson (1970) have also assessed subspecific dif- ferentiation in the marten. According to Hall (1981), the mainland of Southeast Alaska is occu- pied by two subspecies of the “americana” group, M. a. kenaiensis north and west of Lynn Canal, and M. a. actuosa from northern Lynn Canal south to about Cleveland Peninsula; in the “caurina” group, M. a. caurina occurs on the mainland south of Cleveland Peninsula, while M. a. nesophila inhabits islands in the Alexander Archipelago (but not as outlined in Map 507) and the Queen Charlotte Islands, Canada. Giannico and Nagorsen (1989), in a recent revision, concluded that the subspecies M. a. nesophila should be applied only to Queen Charlotte Islands populations (but see Swarth 1911), and assigned Vancouver Island and coastal British Columbia Martens to M. a. caurina. They suggested that M. a. caurina and M. a. americana may intergrade in Southeast Alaska. Unfortunately, these authors were unaware that the specimens from Baranof and Chichagof islands that they examined came from transplanted stocks. Martes pennanti (Erxleben, 1777), Fisher The earliest report of this species in Southeast Alaska that we are aware of is that of Coues (1877), who stated that the Fisher’s range includes the “southern panhandle of Alaska”, and that he had examined some of these. This report was noted by Allen (1942), Anderson (1946), and Hall (1981). F. H. Gray, in a 1915 report on Southeast Alaska fur- bearing animals to the Bureau of Fisheries, stated: “Fishers are extending their range to the north. But one was even taken on the Stikine River before 1908. Several were taken the last three years from the boundary [of Alaska and British Columbia]. Tracks of this animal were seen not far from here [Wrangell] last winter.” Even earlier, C. P. Streator wrote, in an 1895 summary report to the Biological Survey, “Mr. Chase, a furdealer here [Streator was writing from Loring, Revillagigedo Island], informs me that he has seen skins [of Fisher] that were taken in this region.” Further light is shed by MacLeod (1950) from registered trapline information in adja- cent British Columbia, where trappers reported cap- turing Fisher from the lower Iskut River, the Unuk River, and near Stewart. The occurrence of Fisher in Southeast Alaska has now been confirmed by a voucher specimen collected from the Taku River in 1994 (UAM 24716). This lends credence to the report of a Fisher seen in Southeast Alaska near the mouth of the Unuk River in the winter of 1958-59 1996 (T. Wills, personal communication 1993), and another reportedly trapped at Point Agassiz in the 1950s (S. Geraghty, personal communication 1994). It appears that this species is an uncommon visitor or marginal resident along the mainland of the region, at least as far north as the Taku River. Mustela erminea Linnaeus, 1758, Ermine The Ermine is a Holarctic species widely distrib- uted throughout Southeast Alaska. Islands where Ermine are documented to occur are as follows: Admiralty, Baranof, Chichagof, Dall, Etolin, Kruzof, Kuiu, Kupreanof (Swarth, 1911, described a Native trapper from Kuiu Island taking Ermine on the “Kake islands” [= Kuiu + Kupreanof islands]), Long, Mitkof, Prince of Wales (Fay and Sease, 1985, men- tion, with further detail, Ermine taken on some of the smaller island adjacent to Prince of Wales), Revillagigedo, Suemez (Fay and Sease, 1985, report- ed two Ermine trapped there, but not preserved, in 1981-82), Wrangell, and Zarembo islands (Heller 1909; Swarth 1911; Hall 1944; MVZ; UAM; USNM; L. Johnson, personal communication 1994). Hall (1944, 1951) recognized six subspecies (five are endemic to the region): Mustela erminea arctica (Yakutat Bay and Glacier Bay), M. e. alascensis (type locality = Juneau), M. e. initis (type locality = Saook Bay, Baranof Island), M. e. salva (type locality = Mole Harbor, Admiralty Island), M. e. celenda (type locality = Kasaan Bay, Prince of Wales Island), and M. e. seclusa (type locality = Port Santa Cruz, Suemez Island). A taxonomic revision based on more specimens (e.g., M. e. seclusa is based on a single specimen) is needed. Mustela nivalis Linnaeus, 1766, Least Weasel An unpublished report (Home 1973*) includes a number of credible sightings of the Least Weasel in the Glacier Bay area. Although we are unaware of any specimens of this species taken in Southeast Alaska, we suspect that this species very likely does occur in places along the northern mainland of the region, but not nearly as extensively as suggested by ADF&G (Alaska Department of Fish and Game 1973). Mustela vison Schreber, 1777, Mink Mink are widely distributed throughout Southeast Alaska, occurring in close association with marine and fresh water ecosystems. Documented Mink records in the archipelago are far from complete, but include Admiralty, Anguilla, Baranof, Barrier islands, Butterworth, Castle, Chichagof, Coronation, Esquibel, Etolin, Kruzof, Kuiu, Kupreanof, Long, Mitkof, Myriad islands, Prince of Wales, Revillagigedo, Shrubby, Sitka Sound islands; Sokalof, Suemez, Tongass, Wrangell, and Zarembo (Heller 1909; Swarth 1911; Hollister 1913; MVZ, UAM; USNM; L. Johnson, personal communication 1994). Two subspecies occur in the region (Hall 1981): M. v. energumenos and M. vy. nesolestes (type locality = Windfall Harbor, Admiralty Island). MACDONALD AND COOK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 585 Gulo gulo (Linnaeus, 1758), Wolverine Wolverines inhabit the mainland and occur as resi- dents or vagrants on several islands in the Alexander Archipelago, specifically Fair, Kuiu, Kupreanof, Mitkof, Revillagigedo, and Wrangell islands (Webster 1949; MVZ; UAM; USNM;; C. Land, per- sonal communication 1992). Lontra canadensis (Schreber, 1777), River Otter The River Otter is found along the coastal and inland waters throughout Southeast Alaska. Specimens records are from a number of mainland localities and, in the Alexander Archipelago, from Admiralty, Baranof, Chichagof, Coronation, Dall, Gavanski, Halleck, Krestof, Kruzof, Kuiu, Kupreanof, Long, Marble, Prince of Wales, Rapids, Shrubby, Warren, Woronkofski, and Wrangell islands (van Zyll de Jong 1972; CMNH; MVZ; UAM; USNM). Lontra canadensis mira (type locality = Kasaan Bay, Prince of Wales Island) is a subspecies endemic to Southeast Alaska and coastal British Columbia (Hall 1981) and was once considered a distinct species restricted to Prince of Wales and neighboring islands (Hall and Kelson 1959). Reviews of the taxo- nomic status of Lontra canadensis (formerly Lutra canadensis) indicated that River Otters from Southeast Alaska are morphologically distinct from other populations (van Zyll de Jong 1972; Fagen 1986). Puma concolor (Linnaeus, 1771), Puma or Mountain Lion The rare occurrence of this species in Southeast Alaska was substantiated by the collection of a male on the east side of Wrangell Island along Blake Channel opposite Aaron Creek, 25 November 1989 (UAM 18551). This area is close to the mainland. The animal may have arrived from British Columbia along the trans-coastal corridor of the Stikine River. Charlie Land, ADF&G, Petersburg (personal com- munication 1994), found no indication that this had been a captive animal. Youngman (1975) documented six Puma sightings in Yukon, between 1955 and 1967. An increasing number of recent sightings have been reported to UAM of Pumas in Southeast Alaska (as well as in the Wrangell Mountains of east-central Alaska). Perhaps these mountain lions are expanding their range in relation to the recent westward expansion of Mule Deer into Yukon and east-central Alaska. Lynx canadensis Kerr, 1792, Canada Lynx Canada Lynx are uncommon residents from upper Lynn Canal north and westward, and occasional visi- tors elsewhere along the mainland of Southeast Alaska, occurring infrequently along the major river corridors. Gray (1915*) and Swarth (1936) suggest- 586 ed that when Snowshoe Hares become scarce in the interior of British Columbia, L canadensis are found more frequently in Southeast Alaska. Preserved specimens from the region are limited to one from Yakutat (MVZ), one from Taku Inlet (USNM), and one found in the Hyder landfill (UAM), May 1995 (exact collection locality unknown). Greg Streveler (personal communication 1994) discovered the skeletal remains of a lynx in upper Muir Inlet of Glacier Bay in 1968; the location of that specimen, if preserved, is not known. Bailey (1920*) reported seeing the carcasses of two lynx killed close to the mainland on Douglas Island (where Snowshoe Hares had been introduced), appar- ently the only island record of this species in Southeast Alaska. Cervus elaphus Linnaeus, 1758, Elk or Wapiti Elk were members of Alaska’s large mammal fauna into the early Holocene (Guthrie 1966); howev- er, there is no evidence that they ever occurred in Southeast Alaska. There have been a number of attempts to introduce Elk to Southeast Alaska (Burris and McKnight 1973), beginning in 1926 and 1927 with the release of seven animals (from the state of Washington) on Kruzof Island. Three attempts were made to intro- duce Elk to Revillagigedo Island: in 1937 (Washington stock), 1963, and 1964 (from the Afognak Island herd which was originally from Washington). Animals were also released on Gravina Island in 1962, and on Annette Island in 1963, and, like all of the previous attempts, failed. In 1987, 50 Elk from Oregon were released on Etolin Island. By 1993, they had increased in number and had spread to nearby Zarembo Island and to adjacent areas, with sightings as far away as Cleveland Peninsula (J. Gustafson, personal commu- nication 1994), Odocoileus hemionus (Rafinesque, 1817), Mule Deer Sitka Black-tailed Deer, the most frequently used common name for this subspecies, are found through- out most of Southeast Alaska south of Cape Spencer. They are rare to non-existent along the mainland coast. Deer are strong swimmers and seem to have lit- tle trouble crossing wide expanses of coastal waters. As a result, they occur on nearly every island in the archipelago, except Forrester Island (Heath 1913*; Klein 1965). According to Home (1973*), deer were unknown on the mainland from Cape Spencer to Yakutat (and westward to Prince William Sound) until they were transplanted to islands in Yakutat Bay in 1934 (from Rocky Pass stocks; Burris and McKnight 1973). Other transplants in the region included the Taiya Valley near Skagway in 1951, 1952, and 1956; and on Sullivan Island, Lynn Canal, in 1951-1954. Deer no longer occur in the Yakutat area, and transplants THE CANADIAN FIELD-NATURALIST Vol. 110 to Taiya River Valley and Sullivan Island eventually failed (L. Johnson, personal communication 1994). The type locality for the Sitka Black-tailed Deer, O. h. sitkensis, is Sitka, Baranof Island (Hall 1981 as Dama). There is a report of one record of the larger interior subspecies, O. h. hemionus, taken from the Stikine River (Alaska Department of Fish and Game 1973). Alces alces (Linnaeus, 1758), Moose Since early this century, Moose have been expand- ing their range into Southeast Alaska (Klein 1965), first establishing breeding populations along the major mainland river valleys (specifically, the Alsek, Chilkat, Taiya, Taku, Stikine, and Unuk rivers, and Thomas Bay) and, more recently, on islands in the Alexander Archipelago in close proximity to the mainland (Etolin, Kupreanof, Mitkof, and Wrangell islands; Alaska Department of Fish and Game 1973). In addition, various residents of the region have reported occasional sightings of Moose on a number of other islands in the Alexander Archipelago (north Chichagof, Lemesurier and Pleasant Islands in Icy Strait, Kuiu, Prince of Wales, Revillagigedo, and Woewodski islands). Two subspecies may occur in the area: Alces a. andersoni on Stikine and Unuk rivers may intergrade with A. a. gigas from the Taku River (Klein 1965; Hall 1981). Moose were transplanted to Berners Bay from the Susitna and Matanuska valleys in 1958 and 1960, and to the Chickamin River from Anchorage-area stock in 1963 and 1964. Burris and McKnight (1973) note that a few Moose had occasionally been observed on the Chickamin River prior to their intro- duction. Moose are occasionally seen in the Salmon River and Hyder area (R. Thomas, personal commu- nication 1990). Rangifer tarandus (Linnaeus, 1758), Caribou The probable rare occurrence of Caribou in north- ern Southeast Alaska is based upon three sources. One is Olaus Murie (1935: 69, 71) who wrote, “According to Indian information, Caribou reach the summit of the Coast Range near Skagway but do not occur on the south slope...Certain Indians declared that Caribou used to be at Haines, Alaska, and they thought that the animals were working back that way.” A recent report (about 1990) of a Caribou seen by local residents near the Kelsall River Valley, north of Haines (S. Brewington, personal communication 1994) supports this claim. Finally, Home (1973*) provides two separate reports of Caribou from the area north and inland of Goose Cove, upper Glacier Bay, one in the late 1950s and another in 1967. Recently, a partial metacarpal of a Caribou has been recovered from a cave on Prince of Wales Island (J. Baichtal, personal communica- tion 1995). 1996 Oreamnos americanus (de Blainville, 1816), Mountain Goat Mountain Goats are found in suitable habitat along the entire mainland coast of Southeast Alaska. The only island record of natural occurrence is that of a single animal seen on Wrangell Island for several years (Klein 1965). This species was successfully introduced on Baranof Island in 1923 (Burris and McKnight 1973). Transplant attempts to Chichagof Island in 1953 and again in 1955 were considered failures (Burris and McKnight 1973; L. Johnson, personal communication 1994). The most recent transplant of Mountain Goats in the Alexander Archipelago was on Revillagigedo Island in 1983 (Smith and Nichols 1984) and this population is extant (J. Gustafson, personal communi- cation 1994), Ovis dalli Nelson, 1884, Dall’s Sheep Dall’s Sheep are found adjacent to Southeast Alaska in British Columbia, on the drier western slopes of the Saint Elias Mountains and the Coast Mountains north of Haines and Skagway (Klein 1965; Nichols 1978). The occurrence of Dall’s Sheep in Southeast Alaska is based on a female collected from the Kelsall River Valley, northwest of Haines, by ADF&G biologists (R. Flynn, personal communication 1994; specimen not located). A trapper family living on the Chilkat River occasionally observed sheep along the US/Canada border in the vicinity of Mount Raymond, not far from where ADF&G collected the female sheep (S. Brewington, personal communication 1994). Three O. dalli skulls collected by Allen Hasselborg for the MVZ are said to be from Southeast Alaska, but no other information is provided. C. P. Streator, in his July-September 1895 summary notes to the Biological Survey, stated this about Dall’s Sheep in Southeast Alaska: “A[n] interior [BC] mammal but found at the head of a number of the large inlets”. There was no further elaboration. Faunal Composition Excluding human beings and marine mammals, 54 species of mammals, representing 38 genera, 16 fami- lies, and 6 orders, are known to occur or have recently occurred in Southeast Alaska (Table 2). They consti- tute 86 subspecies and monotypic species. Of these, 27 (excluding Ursus arctos) are essentially endemic to Southeast Alaska. Rodents, with 20 species, consti- tute the most speciose order, followed by carnivores with 14 species. Forty-eight species are native and extant. Four extant species are not native to the region: Oryctolagus cuniculus, Rattus norvegicus, Mus mus- culus, and Cervus elaphus. Procyon lotor and Alopex lagopus, also not native, are now believed extirpated. New species confirmed present in the region since Hall (1981) are Ochotona collaris, Martes pennanti, and Puma concolor. New species, as yet unconfirmed MACDONALD AND CooK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 587 by voucher specimen, are Mustela nivalis, Rangifer tarandus, and Ovis dalli. Distribution Patterns Biogeographic subregions While many gaps in our knowledge of mammals have been filled in recent years, our current under- standing of mammal distribution patterns remains similar to that first outlined by Swarth (1911, 1936). His delineation of biogeographic subregions of Southeast Alaska are in general agreement with the new information, although we further refine his analy- sis below. Based on the presence of endemic taxa and unique combinations of native species, Southeast Alaska can be reduced to five subregions (Figure 2, Table 1). These subregions agree with Swarth (1911, 1936), except that he combined Admiralty, Baranof, and Chichagof islands into a single subregion. The subregions should be viewed as an hypothesis that can be tested by examining relationships among popula- tions of species that are widespread throughout the region. Molecular systematics, in particular, holds great promise for characterizing relationships among island and mainland populations in Southeast Alaska (e.g., Gilbert et al. 1990). 1. Mainland. The mainland coast of Southeast Alaska has several connections to interior Canada through trans-mountain corridors along the major river systems. It forms a definable and species-rich subre- gion. Several islands (e.g., Douglas, Farm) are includ- ed in this subregion due to their proximity to the main- land coast. Mammals indicative of Mainland include Sorex palustris, Marmota caligata, Zapus princeps, and Oreamnos americanus. A number of species such as Ochotona collaris, Spermophilus parryti, Neotoma cinerea, Martes pennanti, Mustela nivalis, Canis latrans, Vulpes vulpes, Lynx canadensis, Rangifer tarandus, and Ovis dalli have been documented only in this subregion. Elevations above 1000 m along the mainland remain some of the least explored areas of the region and the ranges of a number of species need to be clarified. Species like Sorex hoyi, Tamias min- imus, Lemmus trimucronatus, Microtus miurus, and Phenacomys intermedius occur just to the east in British Columbia or Yukon. Specimens of P. inter- medius were collected within 2 km of the Canada bor- der near Hyder in 1995. Six mammals are endemic to this subregion (Table 1) and Hall (1984) considered Ursus arctos dalli endemic to the Yakutat area. 2. Middle and Southern Inner Islands. This subre- gion includes those islands in the Alexander Archipelago south of Frederick Sound that are rela- tively near the mainland. These islands share several taxa that are otherwise found only in the Mainland subregion including Tamiasciurus hudsonicus, Gulo gulo, and Alces alces. The latter species, along with Clethrionomys gapperi, Synaptomys borealis, and Erethizon dorsatum, illustrate a general pattern of 588 BIOGEOGRAPHIC | SUBREGIONS: @) MAINLAND @) MIDDLE & SOUTHERN INNER ISLANDS @) SOUTHERN OUTER ISLANDS NORTHERN INNER ISLANDS 6) NORTHERN OUTER ISLANDS THE CANADIAN FIELD-NATURALIST Vol. 110 FiGuRE 2. Five biogeographic subregions can be identified in Southeast Alaska based on the presence of unique combinations of taxa and endemics (Table 1). westward postglacial colonization into the archipel- ago. Clethrionomys gapperi solus is considered endemic to this subregion (Hall 1981). 3. Southern Outer Islands. Prince of Wales and the numerous small islands to the west form a relatively distinct and insular subregion. Clarence Straits serves as a formidable barrier to faunal movement to the east except, perhaps, in the vicinity of Snow Passage and the Kashevarof Islands at the northeast end of Prince of Wales Island. Those islands may function as a dispersal corridor. Swarth (1911) also included Zarembo Island in this subregion. The absence of Sorex cinereus and Martes americana on Zarembo Island lends support to this hypothesis, although Zarembo Island shares some taxa with sub- region 2. We tentatively include the outer islands of Coronation, Warren, Forrester, Duke, and Mary within this subregion. Six taxa are considered endemic to this subre- gion: Sorex monticolus malitiosus, Glaucomys sabrinus griseifrons, Peromyscus keeni oceanicus, Microtus coronarius, Mustela erminea celenda, and M. e. seclusa. 4. Northern Inner Islands. Admiralty Island is sep- arated from Mainland by Stephens Passage and from the Northern Outer Islands by Chatham Strait. Admiralty Island shares Ursus arctos (considered an endemic subspecies, U. a. sitkensis, by Hall 1984) 1996 MACDONALD AND CooK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 589 Number of Species (@) % of taxa endemic (@) Number of Species (@) Number of Species (@) Number of Species (@) Nv Number of Species (@) % of taxa endemic (m) Number of Species (@) % of taxa endemic (m % of taxa endemic (@ FiGurE 3. Patterns of species richness and endemism of native land mammals along six transects in Southeast Alaska. and an absence of Canis lupus with the Northern Outer Islands. Many features of its mammal fauna set the two subregions distinctly apart, however, including the presence on Admiralty Island of Sorex monticolus (and absence of S. cinereus), Microtus pennsylvanicus, Ondatra zibethicus, and Martes americana. Furthermore, the apparent morphological distinctiveness of Microtus pennsylvanicus admiral- tiae (Heller 1909), Castor canadensis phaeus (Heller 1909), Mustela erminea salva (Hall 1951) and, pos- 590 sibly, Martes americana (“very different looking”, L. Johnson and R. Flynn, personal communications 1994) suggest a relatively high degree of endemism. Thus, we consider Admiralty Island, including sever- al of the smaller adjacent islands, a distinct biogeo- graphic subregion of Southeast Alaska. 5. Northern Outer Islands. Islands in this subre- gion, including Baranof, Chichagof, Yakobi and sur- rounding islands, are relatively isolated from the mainland to the north by Cross Sound and Icy Strait (although Brown Bear and Moose have been known to cross) and from Admiralty Island to the east by Chatham Strait. Microtus oeconomus sitkensis (Merriam 1897) and Mustela erminea initis (Hall 1944, 1951) are endemic to this subregion. Castor canadensis phaeus (Hall 1981) and Ursus arctos sitkensis (Hall 1984) may also be endemic. Species Richness and Endemics Patterns of species richness across the complex landscape of Southeast Alaska were first noted by Swarth (1936). He suggested a general pattern of increasing species richness across the archipelago moving west to east from the Gulf of Alaska toward the mainland. This pattern is consistent along four west-east transects (Figure 3A-D), and by comparing the number of species across the subregions (Table 1). Decreasing species richness with increasing dis- tance from the mainland source populations is a common attribute of archipelagos (Simberloff and Wilson 1969). Contrary to documented latitudinal gradients for mammals (Huston 1994), there is a south-to-north increase in species richness along the mainland. The upper Lynn Canal area supports the highest number of mammal species for Southeast Alaska (Figure 3F). The south-to-north increase in richness along Southeast Alaska’s mainland reflects the history of glaciation in this region. In the Haines and Skagway area, species typical of the coastal fauna (and flora) meet species of both interior British Columbia (species of southern origin such as Lepus americanus, Canis latrans, and Puma concolor) and interior Alaska and Yukon (Beringian origin such as Ochotona collaris, Clethrionomys rutilus, Microtus oeconomus, Mustela nivalis, and Ovis dalli; Youngman 1975). No latitudinal increase in species richness is evident along the outer islands of the Alexander Archipelago (Figure 3E). The naturally fragmented and insular nature of Southeast Alaska suggests a high potential for evolu- tionary divergence and endemism (Anderson 1994) and at least 27 mammalian taxa have been described as occurring only in this region (Hall 1981). Klein (1965) first noted that the proportion of endemic taxa (to total number of taxa present) across the landscape of Southeast Alaska decreases as one moves eastward THE CANADIAN FIELD-NATURALIST Vol. 110 from the more isolated outer islands of the Alexander Archipelago toward the mainland (Figure 3A-D). The pattern is most pronounced across the southern end of the archipelago (Figure 4D). This is not surprising as Prince of Wales Island and the numerous smaller islands that comprise the Southern Outer Islands sub- region (Figure 2) harbor the largest number of endemics on the archipelago (n=6; Table 1). High endemism in that subregion is followed in the archi- pelago by the Northern Inner Islands (n= 3), the Northern Outer Islands (n= 2), and the Middle and Southern Subregion (n = 1). Nonetheless, the highest absolute number of endemics is found in the Mainland subregion, reflecting its extensive latitudi- nal span and relative isolation from the remainder of North America. The relative isolation of the entire region may be the primary factor responsible for ele- vated level of endemism (Anderson 1994), however, the possibility that some endemics are much older than previously suggested (Swarth 1936) is under investigation (Stone et al. 1996*). Whether some members of the current fauna immi- grated to these subregions since the last glaciation (ca. 10-12 000 years ago) and subsequently differentiated (i.e, neoendemics, Myers and Giller 1988) or whether some of the endemics are remnants of a relictual fauna that was more widely distributed prior to the last glaciation (paleoendemics) is basic to our inter- pretation of the evolution of this fauna. Origins An overview of the terrestrial mammals of Southeast Alaska is incomplete without considering the fauna’s origins. First addressed by Swarth (1936), and later expanded and refined (Klein 1965), the history of mammal colonization and evolution in Southeast Alaska has been dynamic and complex. Both Swarth and Klein worked under the assumption that the last glacial maxima (the Fraser or late- Wisconsin glacial) essentially covered all of Southeast Alaska, and that the current distributions of mammals were the result of factors affecting post- glacial colonization from refugia outside the region. The large number of endemic forms now present on a number of outer islands of the archipelago was explained by them within the Holocene time frame (i.e., aS neoendemics). While Swarth (1936: 61) concluded that “whatev- er is peculiar to the Sitkan fauna and flora inevitably has develor 2d subsequent to the Pleistocene time’, and that active colonization is discernible at the pre- sent time, he remained troubled by the peculiar dis- tributions of two insular mammals, Peromyscus sitkensis and Microtus coronarius. To him, these well defined forms afforded corroborative evidence of a relict fauna surviving the last glaciation. Cowan (1935) also concluded that the P. sitkensis group, as well as some of the island endemic forms of Sorex monticolus, were outer island survivors of the late- 1996 MACDONALD AND CooK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 591 ” MID-WISCONSIN INTERSTADIAL (Cirea 45,000 yr B.P.) Woodland Mammals From The South FIGURE 4. Approximate extent of glaciers during the height of the Mid-Wisconsin Interstadial (60 000 - 30000 yr B. P.) including the potential patterns of colonization of mammals along the ice-free corridor of the North Gulf Coast. Tundra-adapted mammals from southern Beringia may have included: Sorex jacksoni, Lepus othus, Marmota camtschatica, M. broweri, Spermophilus parryii, Clethrionomys rutilus, Microtus oeconomus, M. miurus, Lemmus sp., Dicrostonyx sp., Canis lupus, Vulpes vulpes, Alopex lagopus, Ursus arctos, U. maritimus, Mustela erminea, Gulo gulo, and Rangifer tarandus. Possible colonizers from the south include Sorex monticolus, Peromyscus sp.. Microtus longicaudus, Ursus americanus, and Sorex palustris. Wisconsin glaciation. That these taxa possibly repre- sent paleoendemics was again raised (Klein 1965). He expanded the geographic scope of the discussion to include hypothesized refugia outside Southeast Alaska to the south on the Queen Charlotte Islands and northwest along the coast to Prince William Sound, Kenai Peninsula, and Kodiak Island (see also Heusser 1960, 1989; Molnia 1986; Mann 1986; O’Reilly et al. 1993). Klein (1965) also introduced the possibility that the continental shelf, which would have been exposed due to a lowered sea level during the last glacial advance, may have provided refugia. Not considered were the origins of relict populations prior to the last glacial. Along with an increasing appreciation of the dynamic nature of the distributions of mammals throughout the region, two initiatives have consider- able promise for clarifying the history of the region. One is the ability to recover phylogenetic relation- ships from molecular genetic data, providing a view of patterns of colonization and relationships among populations. Another is the recent discovery of hun- dreds of caves in the extensive karst of Prince of Wales and other islands. Some of these caves are yielding a wealth of mammal bones, some of which have dated as far back as the Mid-Wisconsin (over 30 000 y B.P.). These fossil discoveries and ongoing molecular phylogenetic studies of several taxa (e.g., Stone et al. 1996*) are beginning to challenge earlier assumptions and expand our vision of the origin of this fauna both spatially and temporally. Heaton (1995) used radiocarbon dates to docu- ment both Ursus arctos and U. americanus on Prince of Wales Island in the early Holocene and in the pre- vious mid-Wisconsin interstadial (60 000 to 30 000 y B.P.; Mann 1986; Elias 1995). Ursus arctos no longer occurs on Prince of Wales Island and U. americanus was previously thought to have arrived there after the last glacial. Also discovered were teeth of Marmota sp. (> 44 500 y B.P.). Other mam- mal fossils, as yet undated, included Peromyscus sp., Vulpes vulpes, Mustela erminea, and Rangifer taran- dus (Heaton and Grady 1992, 1993; Heaton 1995; J. Baichtal, personal communication). This deep history of occurrence provides an alter- native to the clean-slate hypothesis of postglacial colonization (Swarth 1936; Clague 1989). A scenario that includes the presence of refugia within the region throughout the late-Wisconsin glacial maxima suggests that some mammals may be paleoendemics, with an evolutionary history in the region that dates from at least the last interstadial (Rogers et al. 1991). The possible existence of Ice Age refugia in Southeast Alaska, such as has been hypothesized for the Queen Charlotte Islands (Heusser 1989; O’Reilly et al. 1993), may explain why some members of the Southeast Alaska mammal fauna are distinctive. Recent DNA studies of insular populations of Ursus 592 THE CANADIAN FIELD-NATURALIST Vol. 110 = | ATE-WISCONSIN GLACIAL MAXIMUM (Circa 18,000 yr B.P.) Ficure 5. Approximate extent of glaciers during the Late-Wisconsin (Fraser) about 18 000 yr B. P. Arrows indicate possi- ble ice free refugia found along the North Gulf Coast during this period (Heusser 1960; Hopkins 1967; Heusser 1989; and references therein). arctos in Southeast Alaska are consistent with this hypothesis (Talbot and Shields 1996). The discovery of pre-Fraser marmot teeth found on Prince of Wales Island is of particular interest (Heaton 1995). Today, Marmota caligata occurs only on the mainland, and is thought to be of south- ern origin (Youngman 1975). Heaton (1995) noted that the marmot molar found on Prince of Wales Island was smaller than Marmota caligata, and sim- ilar in size to Marmota flaviventris, found farther south and east. Alternatively, this marmot may be more closely related to M. broweri or M. camtschat- ica, small-bodied species of marmots now occurring on the west and east sides, respectively, of what was once Beringia (see Hoffmann et al. 1979). Or, it may be related to other smaller-bodied forms of marmots including M. c. vigilis, an endemic sub- species from Glacier Bay; or M. c. sheldoni, another smaller-sized endemic, last seen in 1908 on Montague Island, Prince William Sound (Howell 1915; Hoffmann et al. 1979). A northern origin for the Prince of Wales Island marmot, rather than southern as mentioned by Heaton (1995), is consis- tent with the affinities of most of the fossil mam- mals found recently. Since most of these fossils are of Holarctic, tundra-adapted species, we propose that, similar to hypotheses advanced for human migration into the New World (Fladmark 1979), a corridor existed along the coast from southern Beringia to the southern end of Cordilleran ice sheet (Figure 4) from the last interstadial to the early Holocene. Such a corridor implies a connection along the continental shelf which was exposed by a lowered sea level (Figure 4) and would have allowed Beringian species to track (sensu Eldridge 1995) tundra habitat eastward along the coast. Such a corridor may also have allowed southern refugial species adapted to woodland habitat to move northward. The return of the late-Wisconsin glaciation effec- tively fragmented the mammal fauna into isolated refugia along the coast (Figure 5). This hypothesis predicts that some of the numerous endemics found along the Pacific Coast of Alaska (Figure 6) may be paleoendemics or remnants of much more widespread taxa, rather than neoendemics that arrived in their present locations since the last glacial maxima. The arrival of the Holocene warming (Figure 6) resulted in a rise of sea level and the even- tual inundation of the Bering Land Bridge and the continental shelf along the Gulf Coast. While this was occurring, tundra habitats generally retreated to the northwest, while Pacific coastal forests advanced northward along the coast, arriving as far as Kodiak Island approximately 1000 y B.P. (Elias 1995). Thus, the high concentration of endemics noted for Southeast Alaska and areas surrounding the Gulf of Alaska may be the result of deeper historical events than previously envisioned. Alternatively, these endemics may be relatively recent immigrants that have evolved rapidly due to island effects (Foster 1964; Adler and Levins 1994; Lance 1995). At this time, these hypotheses cannot be effectively tested because the evolutionary relationships of the endemics, or their taxonomic status for that matter, are poorly understood. \ 1996 MACDONALD AND COOK: LAND MAMMAL FAUNA OF SOUTHEAST ALASKA 593 HOLOCENE (Recent) Ficure 6. Localities (circled) with concentrations of potential relict faunas (paleoendemics) along the North Gulf Coast. Mammals that occur or have recently occurred at these localities include: Sorex hydrodromus (A), S. cinereus (B, F, G, H, I, J), S. monticolus (B, C, F-I, K, L, M), S. palustris (I, M), Lepus othus (B, C), Marmota sp. (F, G, I, K[fossil], M), Spermophilus sp. (B-F), Peromyscus sp. (J, K, L, M), Clethrionomys rutilus (G-D), Microtus oeconomus (B-E, F- J), Vulpes vulpes (B, C, E-G, K[fossil]), Ursus americanus (F, H, I, K[includes fossil], L, M), U. arctos (B, E-G, J, K[fossil], Mustela erminea (B, E-G, J, K[includes fossil], L, M), Gulo gulo (B, F, M), and Rangifer tarandus (B, C, E[bones in midden], F[extinct circa 1913], K[fossil], L[extinct circa 1910]). Arrow represents postglacial advance of Pacific coastal forest and associated fauna. Conclusions Further excavation of fossil materials from the karst region of Southeast Alaska will continue to clarify our understanding of the composition and test hypotheses related to the origins of this fauna. In addition to paleontology, investigations of the molecular systematics of the extant and extinct mam- mals offers an opportunity to examine the temporal dynamics of a regional fauna that was shaped by late-Pleistocene events. Our ability to characterize the nature of endemicity with molecular data will be particularly valuable for establishing the temporal sequence of events, particularly with respect to sepa- rating paleoendemics from neoendemics. Knowledge of distributions of organisms is fun- damental to our understanding and conservation of biological diversity (e.g., Meffe and Carroll 1994). Well-documented scientific surveys are fundamen- tal to basic theoretical research conducted in this coastal region as well as for applied management decisions related to the evaluation of impacts of perturbations on the biota (Miller 1993; Miller and Scudder 1994; Raven and Wilson 1992). Throughout Southeast Alaska we still lack the base- line data needed to identify and monitor areas of high species richness and heavy concentrations of endemic taxa (biodiversity “hot spots’; Noss and Cooperrider 1994). Controversy surrounds past, present, and projected future human-induced change in this region (United States Department of Agriculture 1991, 1996) and a sustained effort to document the fauna and flora of this region is needed. The recent clear-cut harvest of timber on most of Long Island, an island with good potential for supporting endemics, is an extreme example of what has occurred in this region. Proposed logging on small islands, as well as exist- ing stands of old-growth on the larger islands may significantly impact endemic taxa (e.g., Glaucomys sabrinus griseifrons). In addition to habitat perturbation and change, endemic forms on the islands, in particular, may be negatively affected by species introductions. The drastic effects on insular faunas of the introduction of exotic species has been well documented through- out the world (e.g., Kaufman and Ochumba 1993). A number of the insular taxa noted in this report could be similarly impacted. Such introductions recently have been implicated in the declining seabird colonies of the Queen Charlotte Islands (Gaston 1994; Bertram and Nagorsen 1995). Bailey (1993) reports introductions of Arctic Fox (Alopex lagopus) on over 180 islands in Southeast Alaska. Few data are available for assessing the impact of introduc- tions on insular populations of the Alexander Archipelago, however, numerous introductions of 594 Elk, Mountain Goat, Red Squirrel, Muskrat, Snowshoe Hare, Raccoon, and Marten are known and introductions continue today (Table 2). The Tongass National Forest comprises nearly 80% of Southeast Alaska’s landmass. Considerable public and scientific attention is now focused on the loss of biodiversity on federal lands. We conclude that this inventory of the mammal fauna is but the first step. The need for a comprehensive inventory of the region’s biological diversity is urgent, given the rapid changes that are occurring on many islands with a high potential for endemism. Delay in action may have irreparable consequences that will greatly increase the duration, expense, and probability of failure of recovery efforts (Murphy et al. 1994). Acknowledgments The research, specimens and materials reported here are the result of hard work, support, and collab- oration of many individuals, some of whom may not be listed below. We sincerely thank all those people and in particular the following individuals for their efforts on behalf of the mammalian biota of Southeast Alaska: Private — T. Bethel, R. Bishop, K. Bovee, S. Brewington, L. and J. Carson, R. Carstensen, S. Fiscus, J. Florie, S. Geraghty, D. Harbor, B. J. Hayward, R. Himschoot, R. Hunter, P. Isleib, R. Jahnke, L. Johnson, J. Owens, D. Rice, L. Robertson, T. and K. Wills, and The Friends of the University of Alaska Museum; Alaska Department of Fish and Game — R. Flynn, J. Gustafson, M. Kirchoff, C. Land, D. Larson, M. Robus, M. Sigman, T. Thorton, K. Titus, and the Canyon Island fish crew; U.S. Forest Service — J. Baichtal, S. Blatt, M. Brown, E. Campbell, J. Canterbury, D. Chester, R. Claire, J. Cristner, C. Crocker-Bedford, T. DeMeo, M. Dillman, J. Falk, C. Ford, R. Guhl, T. Hanley, C. Iverson, P. Ianik, S. Jacobson, D. Johnson, L. Kvaalen, M. Laker, W. Martin, K. Post, A. Russell, K. Rutledge, T. Schenck, T. Suminski, D. Walter, S. Wise-Eagle; National Park Service — J. Taggart, H. Lentfer, M. Murray; U.S. Fish and Wildlife Service N. Holmberg, M. Jacobsen, D. Klein, J. Lindell, J. Maniscalo, P. Schempf, E. Grossman, J. McClung.; Sheldon Jackson College — M. Siefert, B. Colthrap, and M. Ahlgren; Alaska Natural Heritage Program — E. West; Institute of Biological Problems of the North (Magadan, Russia) — N. Dokuchaevy, A. Tsvetkova; Curators of Museums — M. Carleton, S. George, D. Nagorsen, J. Patton, R. Timm, C. 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Mammal species of the world: A taxonomic and geographic refer- ence. Second Edition. Smithsonian Institution Press, Washington. 1206 pages. Youngman, P. M. 1975. Mammals of the Yukon Territory. National Museums of Canada, Ottawa, Publications in Zoology 10: 1-192. Received 3 October 1995 Accepted 14 May 1996 Relative Abundance, Migration Strategy, and Habitat Use of Birds Breeding in Denali National Park, Alaska PETER W. C. PATON!” and THOMAS H. POGSON! ‘Alaska Bird Observatory, P.O. Box 80505, Fairbanks, Alaska 99708 Present address: Department of Natural Resources Science, University of Rhode Island, Kingston, Rhode Island 02881 Paton, Peter W. C., and Thomas H. Pogson. 1996. Relative abundance, migration strategy, and habitat use of birds breeding in Denali National Park, Alaska. Canadian Field-Naturalist 110(4): 599-606. The breeding bird community in Denali National Park, Alaska, was studied over a three-year period (1993-1995). Birds were surveyed from nine off-road routes in spruce forests from 1993-1995 and from four on-road routes in 1994 and 1995. Thirty-nine species were detected during off-road routes, whereas 80 species were detected from on-road routes. The most abundant species were sparrows (White-crowned Sparrow [Zonotrichia leucophrys|, American Tree Sparrow [Spizella arborea], and Savannah Sparrow [Passerculus sandwichensis]) and warblers (Wilson’s Warbler [Wilsonia pusilla] and Orange-crowned Warbler [Vermivora celata]). The avian community in Denali National Park was similar to other areas in the region, although there were some distinctive differences. The majority of detected individuals (69%) presumably migrate to the tropics; residents accounted for only 9% of the individuals detected. Shrublands had the greatest relative abundance of any habitats surveyed, whereas species richness was greatest in tall shrub and spruce forest habitats. These findings generally concur with previous research in the region. Both habitats should be important components in any pro- gram that attempts to monitor avian populations in interior Alaska. Key Words: birds, habitat use, abundance, migratory status, Denali National Park, Alaska. Little has been published on avian communities breeding in forests and alpine tundra habitats of central, interior Alaska. West and DeWolfe (1974) calculated densities and energy consumption of birds nesting in forests near Fairbanks. Spindler and Kessel (1980) investigated avian community structure and habitat use in forests and shrub habi- tats near Northway. Farther to the south in Kluane National Park, Yukon Territory, Hoefs (1973) and Theberge (1976) described the abundance and habi- tat use of birds. Although there is some qualitative information available on relative abundance of birds in the Denali area (Dixon 1938; Murie 1963; Kessel and Gibson 1978; Kertell 1988), few quantitative studies have been conducted on the birds in the Park, with the exception of investigations of raptors (e.g., Laing 1985). We investigated the avian community at Denali National Park from 1993 to 1995. We address the following questions: (1) what is the relative abun- dance of birds breeding in Denali National Park? (2) what are their migration strategies, that is, where do birds presumably migrate to after breeding in the Park? (3) which habitats have the greatest species richness and relative abundance, and which habitats do birds from different migratory strategies use? The answers to these questions are important because of avian monitoring programs being initiated in Alaska and the need to determine which habitats and species should be sampled in interior Alaska. Study Area and Methods Denali National Park and Preserve (63°40'N, 149°30'W) is located in the central Alaska Range, approximately 210 km southwest of Fairbanks, Alaska. The Park is 2.4 million ha in size. A 150-km long road bisects the eastern half and our study area includes lands within | km of this dirt road. The climate is characterized by cold, snowy winters and cool summers. Dominant trees are White Spruce (Picea glauca), Black Spruce (P. mariana), Quaking Aspen (Populus tremuloides), Balsam Poplar (P. balsamifera) and Paper Birch (Betula papyrifera). Low, medium, and tall shrub habitats are character- ized by willows (Salix spp.), Dwarf Birch (B. glan- dulosa), Mountain Alder (Alnus crispa), blueberry (Vaccinium uliginosum), and Mountain Cranberry (V. vitis-idaea). Alpine tundra is dominated by sedges, grasses and forbs. Elevations in the road cor- ridor varies from 490 m to 1220 m. Forest habitats are restricted to areas below 600-900 m elevation, alpine tundra occurs above 915 m elevation, and the intervening elevations are dominated by shrub habi- tats. The growing season is relatively short, with most areas in the road corridor free of snow by mid- late June in most years. Permanent snowfields remains on north-facing slopes at higher elevations. We used off- and on-road point counts to deter- mine the relative abundance of birds in Denali National Park (Ralph et al. 1994; Hanowski and Niemi 1995). We established nine survey routes in spruce-dominated forests (hereafter off-road routes). Forests were defined as habitats where trees (i.e., single stemmed-woody plants > 3 m height) consti- tuted > 10% of the crown cover (Viereck et al. 1992). Off-road routes were placed only in forest 599. 600 stands that measured > 100 ha, and were < 0.5 km from the Park road. Each off-road route consisted of 12 stations spaced 250 m apart. Surveys were con- ducted from 10 to 30 June at the same 108 stations in 1993, 1994, and 1995. All birds detected at an unlimited distance from stations were recorded for a 5-minute period (Ralph et al. 1994). We also conducted Breeding Bird Survey (BBS)- type surveys (Ralph et al 1994; hereafter on-road routes) along the entire length of the Park’s main road. Each on-road route was surveyed twice in 1994 and 1995 from 8 to 23 June. Three on-road routes (1.e., Park Highway to the Teklanika River; Teklanika River to Porcupine Forest; and Porcupine Forest to McKinley Bar) were 39.2 km long and had 50 stations. The fourth on-road route, McKinley Bar to Jeholis’ Cabin, was only 31.4 km long with 36 stations. Each station was surveyed for 3 minutes and observers drove between stations. Both on-road and off-road routes were initiated at 0330 ADT (+ 15 minutes). We classified migration strategies for all birds detected during our fieldwork in Alaska based on the previously published classifications schemes of Kessel and Gibson (1978), Gauthreaux (1991), Dobkin (1994), and Rappole (1995). Migration strategies for birds breeding in Denali were catego- rized as follows; resident: majority of the population resides in Alaska the entire year, although some pop- ulations may be irruptive certain years; Nearctic migrant: majority of population winters south of Alaska and north of tropic of Cancer; short-distance Neotropical migrant: all or part of the population winters south of tropic of Cancer, but north of South America; long-distance Neotropical migrant: majori- ty of the population winters in South America; Paleotropical migrant: majority of population win- ters in the tropics of Asia. Two species of redpoll, Carduelis flammea and C. hornemanni, were com- bined into one taxon, redpoll, because of difficulties in identifying the two species (Troy 1985). We used the percentage of stations with detections (hereafter frequency of occurrence; Bart and Klosiewski 1989) and the mean number of individu- als per station (hereafter abundance) as indices to the relative abundance of birds in the Park. Species rich- ness was calculated as the mean number of species detected per station. To determine annual fluctua- tions in abundance, we used the formula: ((year, - year,)/year,) X 100. We calculated annual fluctua- tions for species only that had individuals present both years. Species were categorized as having expe- rienced significant population fluctuations if they increased by over 100% or declined by over 50% between years. We determined habitat use of birds in the Park by measuring vegetation at 100 stations on the two on- road routes (i.e., Park Highway to Porcupine Forest) THE CANADIAN FIELD-NATURALIST Vol. 110 in 1995. Near each station, we placed three 25-m diameter vegetation plots, with plots located 120° apart. Vegetation plots were selectively located 25 to 100 m from the point count station to insure plots did not include the road. Within each vegetation plot, we measured the distance to the four nearest trees and determined the canopy cover of broadleaf and needleleaf trees with a densiometer. We also mea- sured ground cover and shrub height at five points spaced | m apart in each of the four cardinal direc- tions, yielding 20 height measurements per plot. We then categorized each 25 m vegetation plot into the habitat classification scheme of Viereck et al. (1992), using some modifications of Kessel (1979). Six habitats used for analyses presented here include: (1) low-medium shrubs: trees accounted for < 10% of cover, dominated by open or closed canopy (> 25% cover) shrub stands of willow, alder, or Dwarf Birch (< 2.4 m high; (2) tall shrub thicket: trees accounted for < 10% of cover, dominated by open or closed stands of willow or alder > 2.4 m tall, (3) mixed forest: broadleaf and needleleaf frees over 3 m tall are present, both deciduous and spruce con- tribute 25-75% of total tree canopy cover, and total canopy cover is greater than 25%. These forests con- sist of mixed stands of deciduous species (aspen, Balsam Poplar, or birch) and spruce; (4) spruce woodland: trees over 3 m tall are present, over 75% of tree cover is spruce, and canopy closure ranges from 10-24%; (5) spruce forest: trees over 3 m are present, over 75% of the tree cover is spruce, and canopy closure is over 25%; (6) tundra: areas above the upper limit of tree growth that support dwarf shrubs (< 20 cm tall), forbs, and herbs. Each station was classified into one of six habitats based on the dominant habitat at the three vegetation. We com- pared avian abundances between habitats using a random effects analysis-of-variance (ANOVA) model and Tukey’s multiple comparison test (SAS 1988). Statistical significance was set at a= 0.05 for all comparisons. Results Relative abundance On-road routes: We detected 80 species from on- road routes, with 69 species in 1994 compared to 65 species in 1995 (Table 1). Most species were uncom- mon; mean abundance of only 17 of 80 species was > 0.10 individuals per station. In addition, species that were absent one year were generally rare; only 26% had > 0.01 individuals detected per station (n = 27). Warblers and sparrows were the most abun- dant species encountered from on-road routes; White- crowned Sparrow, American Tree Sparrow (Spizella arborea), and Wilson’s Warbler all had detection rates of > 1.0 individual per station in both years (Table 1). Other relatively common species included Savannah Sparrow (Passerculus sandwichensis), 1996 PATON AND POGSON: BIRDS BREEDING IN DENALI NATIONAL PARK 601 TABLE 1. The mean number of individuals detected per station (n = 186 stations per year) for on-road routes in Denali National Park. Migration strategies are given following scientific names: RE = resident, SD = short-dis- tance Neotropical migrant, LD =long-distance Neotropical migrant, NM =Nearctic migrant, PM = Paleotropical migrant. Species 1994 1995 Common Loon (Gavia immer) NM +! Greater White-fronted Goose (Anser albifrons) SD + Green-winged Teal (Anas crecca) SD 0.03 0.01 Mallard (A. platyrhynchos) SD + Northern Pintail (A. acuta) SD 0.01 0.01 American Wigeon (A. americana) SD 0.01 0.02 Ring-necked Duck (Aythya collaris) SD 0.08 Greater Scaup (A. marila) NM + Lesser Scaup (A. affinis) SD 0.08 Oldsquaw (Clangula hyemalis) NM + White-winged Scoter (Melanitta fusca) NM 0.03 Black Scoter (M. nigra) NM + Surf Scoter (M. perspicillata) NM + Barrow’s Goldeneye (Bucephala islandica) NM 0.01 + Bufflehead (B. albeola) NM 0.02 0.01 American Golden-Plover (Pluvialis dominica) LD + 0.02 Whimbrel (Numenius phaeopus) SD 0.02 Lesser Yellowlegs (Tringa flavipes) SD 0.022 0.024 Spotted Sandpiper (Actitis macularia) SD +; + Wandering Tattler (Heteroscelus incanus) SD > Red-necked Phalarope (Phalaropus lobatus) LD + Common Snipe (Gallinago gallinago) SD 0.06 0.03 Least Sandpiper (Calidris minutilla) SD + + Upland Sandpiper (Bartramia longicauda) LD + + Long-tailed Jaeger (Stercorarius longicaudus) LD 0.03 + Bonaparte’s Gull (Larus philadelphia) SD 0.01 + Mew Gull (L. canus) NM 0.23 0.10 Herring Gull (Z. argentatus) SD 0.01 Arctic Tern (Sterna paradisaea) LD + Golden Eagle (Aquila chrysaetos) SD 0.02 0.02 Northern Harrier (Circus cyaneus) SD + Northern Goshawk (Accipiter gentilis) SD + American Kestrel (Falco sparverius) SD + Merlin (Falco columbarius) LD 0.03 + Gyrfalcon (Falco rusticolus) NM 0.01 Rock Ptarmigan (Lagopus mutus) RE 0.02 + Willow Ptarmigan (Lagopus lagopus) RE 0.03 0.04 Short-eared Owl (Asio flammeus) SD + Great Horned Ow! (Bubo virginianus) RE + Three-toed Woodpecker (Picoides tridactylus) RE + + Northern Flicker (Colaptes auratus) NM + + Olive-sided Flycatcher (Contopus borealis) LD + + Say s Phoebe (Sayornis saya) SD 0.02 + Alder Flycatcher (Empidonax alnorum) LD 0.23 0.11 Hammond’s Flycatcher (Empidonax hammondii) SD + + Horned Lark (Eremophila alpestris) SD + 0.01 Violet-green Swallow (Tachycineta thalassina) SD + Cliff Swallow (Hirundo pyrrhonota) LD 0.10 0.30 Gray Jay (Perisoreus canadensis) RE 0.11 0.06 Black-billed Magpie (Pica pica) RE 0.03 0.08 Common Raven (Corvus corax) RE 0.02 + Black-capped Chickadee (Parus atricapillus) RE + + Boreal Chickadee (P. hudsonicus) RE 0.05 0.01 Arctic Warbler (Phylloscopus borealis) PM 0.36 0.25 Ruby-crowned Kinglet (Regulus calendula) SD 0.05 0.02 Gray-cheeked Thrush (C. minimus) LD 0.12 0.10 Swainson’s Thrush (C. ustulatus) LD 0.22 0.15 (Continued) 602 TABLE |. Continued. Species Hermit Thrush (C. guttatus) SD American Robin (Turdus migratorius) SD Varied Thrush (xoreus naevius) NM Northern Wheatear (Oenanthe oenanthe) PM Northern Shrike (Lanius excubitor) NM American Pipit (Anthus rubescens) SD Orange-crowned Warbler (Vermivora celata) SD Yellow Warbler (Dendroica petechia) LD Yellow-rumped Warbler (D. coronata) SD Blackpoll Warbler (D. striata) LD Northern Waterthrush (Seiurus noveboracensis) LD Wilson’s Warbler (Wilsonia pusilla) LD American Tree Sparrow (Spizella arborea) NM Savannah Sparrow (Passerculus sandwichensis) SD Fox Sparrow (Passerella iliaca) NM Lincoln’s Sparrow (Melospiza lincolnii) SD Golden-crowned Sparrow (Zonotrichia atricapilla) NM White-crowned Sparrow (Zonotrichia leucophrys) SD Dark-eyed Junco (Junco hyemalis) NM Lapland Longspur (Calcarius lapponicus) NM Pine Siskin (Carduelis pinus) NM redpoll (Carduelis spp.) RE White-winged Crossbill (Loxia leucoptera) RE '< 0.01 individuals per station. Orange-crowned Warbler (Vermivora celata), red- poll, and Fox Sparrow (Passerella iliaca). Fifteen of 80 species exhibited significant popula- tion fluctuations between 1994 and 1995 (Table 1). Species with particularly large annual fluctuations (1.e., an increase over 200% or a decline greater than 66.7%) were uncommon and included: the Bonaparte’s Gull (Larus philadelphia), Merlin (Falco columbarius), Rock Ptarmigan (Lagopus mutus), Say’s Phoebe (Sayornis saya), Cliff Swallow (Hirundo pyrrhonota), and Boreal Chickadee (Parus hudsonicus). Our surveys were primarily designed to monitor passerines, but other taxa were also detected. Most species of waterfowl were relatively uncommon and tended to be restricted to kettle ponds on the west side of the Park. Diurnal raptors were also uncom- mon, although Golden Eagles (Aquila chrysaetos) were regularly observed both years, as were Merlins in 1994. Willow (Lagopus lagopus) and Rock Ptarmigan (L. mutus) were relatively scarce during on-road routes because surveys were conducted too late to detect displaying males (P. Paton, unpub- lished data). Typical of interior Alaska (Kessel and Gibson 1978), the density of nesting shorebirds (Charadriiformes) was extremely low, with the exception of the Lesser Yellowlegs (Tringa flavipes) and Common Snipe (Gallinago gallinago). The most common corvid in the Park was the Gray Jay (Perisoreus canadensis), where they were found pri- marily in spruce forests. THE CANADIAN FIELD-NATURALIST Vol. 110 1994 1995 0.04 0.02 0.34 0.28 0.12 0.14 + + + + + 0.76 0.88 0.01 0.20 0.22 + + 1.19 1.16 2.05 1.69 0.92 0.70 0.49 0.54 + + 0.07 0.09 3.03 2.33 0.32 0.28 0.02 0.01 + 0.47 0.72 0.07 0.06 Many bird watchers visit Denali National Park to see specific species, most of which were uncommon in the Park during our surveys and included the fol- lowing: seven pairs of Long-tailed Jaegers (Stercorarius longicaudus) nested near the road in 1994 compared to five pairs in 1995. Only three to four pairs of American Golden-Plovers (Pluvialis dominica) were found at higher elevations, where they apparently nested near breeding jaegars. Gyrfalcon (Falco rusticolus) nested at two eyries vis- ible from the road in 1994. One eyrie was active in 1995 near Eielson Visitor Center, although no birds were seen during systematic surveys. The Northern Hawk Owl (Surnia ulula) was not found during our fieldwork, nor were there any reports of this elusive species occurring in the Park in 1993 to 1995. The Northern Wheatear (Oenanthe oenanthe) was only detected at one station on Thorofare Pass in 1994 and was seen at the same location in 1995 after surveys were completed. The Northern Shrike (Lanius excu- bitor) was detected nesting in the Porcupine Forest both years during fieldwork and Lapland Longspur (Calcarius lapponicus) nested in three sites in the tundra zone on Thorofare Pass, Stony Hill, and Highway Pass. Arctic Warblers (Phylloscopus bore- alis) were abundant in higher-elevation tall willow (Salix spp). thickets throughout the Park. Off-road routes: Thirty-nine species were detected in 1993 to 1995 from off-road routes (Table 2). There was little annual variation in species richness 1996 PATON AND POGSON: BIRDS BREEDING IN DENALI NATIONAL PARK 603 TABLE 2. The frequency of occurrence (% stations with detections) and relative abundance (mean number of individuals per station; n =108 per year) for off-road routes in spruce forests in Denali National Park. Frequency of occurrence Abundance Species 1993 1994 1995 1993 1994 1995 Mallard 0.9 +! Greater Yellowlegs 0.9 + Lesser Yellowlegs 2.8 5.6 0.06 0.08 Spotted Sandpiper 3)37/ 0.05 Upland Sandpiper 0.9 + Common Snipe 0.9 + Mew Gull 8.3 6.5 4.6 0.13 0.11 0.06 Spruce Grouse (Dendragapus canadensis) 1.9 0.02 Three-toed Woodpecker 1.9 3.7 0.03 0.04 Northern Flicker 0.9 + Olive-sided Flycatcher 10.2 4.6 13.9 0.12 0.05 0.15 Western Wood-Pewee (Contopus sordidulus) 0.9 + Alder Flycatcher 0.9 + Hammond’s Flycatcher 0.9 + Gray Jay 35.2 24.1 16.7 0.52 0.42 0.20 Black-billed Magpie 0.9 + Common Raven 2.8 0.05 Boreal Chickadee 93) L537 7.4 0.17 0.24 0.11 Ruby-crowned Kinglet 10.2 13.9 11.1 0.15 0.19 0.11 Gray-cheeked Thrush 6.5 2.8 10.2 0.07 0.04 0.14 Swainson’s Thrush 54.6 Wes 47.2 0.89 ilesyil 0.85 Varied Thrush 40.7 57.4 46.3 0.58 1.06 0.75 American Robin 50.9 43.5 43.5 0.67 0.62 0.57 Bohemian Waxwing (Bombycilla garrulus) 4.6 2.8 0.07 0.13 Orange-crowned Warbler 28.7 16.7 29.6 0.38 0.20 0.43 Yellow-rumped Warbler 37.0 65.7 56.5 0.58 1.08 0.82 Blackpoll Warbler 0.9 + Northern Waterthrush 0.9 0.02 Wilson’s Warbler 29.6 36.1 38.9 0.42 0.50 0.52 American Tree Sparrow 29.6 29.6 SiIlES) 0.53 0.74 0.56 Savannah Sparrow 5.6 13.0 93 0.19 0.23 0.12 Fox Sparrow 4.6 O83 2.8 0.03 0.13 0.03 Lincoln’s Sparrow 2.8 4.6 0.03 0.07 White-crowned Sparrow 61.1 72.2 76.9 1.42 2.03 1.94 Dark-eyed Junco 31 80.6 q3s1 ST 1.86 1.45 redpoll 65.7 25.9 36.1 1.59 0.45 0.45 Pine Siskin 0.9 + White-winged Crossbill Ike 39.8 1.9 0.59 4.83 0.07 '+<0.01 individuals per station. on off-road routes: 30 species were detected in 1993, Varied Thrush (/xoreus naevius), and American 26 species in 1994, and 27 species in 1995. Twenty species were found all three years, 4 species were missed only in one year, and 15 species were detect- ed only in one year. The Dark-eyed Junco (Junco hyemalis), White-crowned Sparrow (Zonotrichia leucophrys), and Swainson’s Thrush (Catharus ustu- latus) were consistently the most common species detected from off-road routes, both in terms of fre- quency of occurrence and abundance (Table 2). Two resident species were widespread in only one year; redpolls occurred at over 65% of all stations in 1993, and flocks of 50 White-winged Crossbills (Loxia leucoptera) were encountered in 1994. Three species of thrush (Turdinae) were common throughout Denali’s spruce forests [the Swainson’s Thrush, Robin (Turdus migratorius)|, whereas the Gray- cheeked Thrush (Catharus minimus) were common only on the west side of the Park. The most abundant species of warbler (Parulinae) from off-road routes was the Yellow-rumped (Myrtle) Warbler (Dendroica coronata coronata). Twenty-six percent of the species found on off- road routes exhibited large fluctuations in abundance between years. The White-winged Crossbill experi- enced the greatest annual fluctuation, with a 98.5% decline in individuals detected from 522 in 1994 to 8 in 1995 (Table 1). Other species with large annual fluctuations (i.e., > 200% increase or > 66.7% decline) included the Olive-sided Flycatcher (Contopus borealis), Gray-cheeked Thrush, Hermit 604 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 3. The migration strategies for birds detected in six habitat types during on-road routes in Denali National Park. Migration strategy Habitat Short-distance Long-distance type Resident Nearctic Neotropical Neotropical Paleotropical Relative abundance (% of individuals detected) Spruce forest 9.6 Oo) 63.1 17.1 0.3 Spruce woodland PI 20.7 47.2 18.5 0.9 Mixed forest 5) 9.8 Shei) 25.9 0 Low-medium shrub 6.8 292 46.2 16.6 IK Tall shrub 9.0 20.5 36.3 25.8 8.5 Tundra 12.0 18.4 33.1 23.0 13.4 Species richness (% of species detected) Spruce forest 12 al 53.6 Dies 0.5 Spruce woodland 14.8 IAS} 44.1 22:5) 123 Mixed forest 5.6 2:2 SED 25.0 0 Low-medium shrub 12.2 19.2 42.7 24.3 1.6 Tall shrub 11.7 16.0 35.1 28.6 8.6 Tundra 1333 Fea 32.6 23.6 12.7 Thrush (Catharus guttatus), (Passerella iliaca), and redpolls. Fox Sparrow Migration strategies The majority of species (62.5%) detected during on-road surveys along the Denali National Park road corridor migrate to the tropics; 34 species were short-distance Neotropical migrants (SDM), 14 species were long-distance Neotropical migrants (LDM), 19 species were Nearctic migrants (NM), 11 taxa were residents (RES), and 2 species were Paleotropical migrants (PM) (Table 1). When water- birds (Gaviiformes, Anseriformes, and Charadri- iformes) were excluded from this analysis, 58.8% of total avifauna were tropical migrants; 19 SDM, 9 LDM, 11 RES, 10 NM, and 2 PM. Species richness averaged 5.7 species per station for on-routes in 1994 and 1995, and the majority were Neotropical migrants; 42.5% SDM, 24.3% LDM, 16.9% RES, 12.2% NM, and 4.1% PM (Table 3). Tropical migrants also accounted for the majority of individu- als (71.7%, n= 8594 individuals) detected from on- road stations in 1994 and 1995; 45.1% SDM, 23.9% LDM, 20.6% NM, 7.7% RES, and 2.6% PM. Habitat use We found significant differences in species rich- ness (F = 5.8, p< 0.001) and abundance (F = 19.9, p < 0.001) among the six habitats we surveyed in the Park (Table 4). The relative abundance of all species was greatest in low-medium shrub habitat, where the American Tree Sparrow, White-crowned Sparrow, Wilson’s and Orange-crowned Warbler reached peak relative abundance. Tall shrub habitats also had large numbers of individuals. In contrast, tundra and mixed forest habitats had relatively low abundance. Tall shrub habitats had the greatest species richness of any of the six habitats we sampled; spruce forests and spruce woodlands also had high richness com- pared to the other habitats. In contrast, species rich- ness was relatively low in tundra habitats, where some habitat specialists resided including Lapland Longspurs and shorebirds. Short-distance Neotropical migrants were preva- lent in all habitats (Table 3), both in terms of species richness and relative abundance; they were especial- ly common in spruce forests and low-medium shrub habitats. Residents were common in a variety of habitats, although less so in mixed forest stands. TABLE 4. Habitat use of birds in Denali National Park. Given are the relative abundance (mean number of individuals per station [+ SE]) and species richness (mean number of species per station) in six habitat types for on-road routes. Habitats with the same letter are not significantly different (P > 0.05; Tukey’s multiple comparsion test). Habitat type Low-medium Tall Mixed Spruce Spruce Tundra shrub shrub forest woodland forest R P, Abundance 852 O98 40.38. 12.6 2 Ola OG OIG te iO OiGree nese Ose 10:0 me LOO Richness AIOEE ODEN S61 One 6S O24 SO Ora ee G:0'2'020d 6:2 OF4bed 5.8 <0.001 1996 More residents were detected in spruce woodlands than other habitats. Nearctic migrants, especially the American Tree Sparrow, reached peak abundance in low-medium shrub habitats. Long-distance Neotropical migrants were most abundant in tall shrubs. There were only two species of paleotropical migrants detected during our surveys; the Arctic Warbler was primarily confined to tall shrub habi- tats, whereas Northern Wheatears utilized rocky tun- dra habitats. Discussion The avian community of spruce forests at Denali National Park was generally similar to other areas in central, interior Alaska (West and DeWolfe 1974; Spindler and Kessel 1980). The most abundant species in Black Spruce forests in the Tanana River valley were the Dark-eyed Junco, Swainson’s Thrush, White-crowned Sparrow, and Yellow- rumped Warbler (Spindler and Kessel 1980), which corresponded to our findings in Denali. However, redpolls were apparently more common in Denali (this study) than near Northway (Spindler and Kessel 1980), although West and DeWolfe (1974) found that redpolls were abundant in Fairbanks in 1971. We found that redpoll numbers fluctuated rather dramatically in Denali, and surveys conduct- ed by Spindler and Kessel (1980) might have occurred in years (1975 and 1977) when their num- bers were relatively low. The Townsend’s Warbler (Dendroica townsendii) was abundant to the east of our study area (West and DeWolfe 1974; Spindler and Kessel 1980), whereas we did not find this species during our fieldwork in Denali; it has been reported occasionally in the Park (Kertell 1988). In central Alaska, Townsend’s Warblers are associated with old-growth, dense canopy spruce forests (Spindler and Kessel 1980) and this habitat does not occur in the Park. Several species that were relatively uncommon in shrub habitats in Denali were more abundant farther to the east (Spindler and Kessel 1980). For example, the Lincoln’s Sparrow (Melospiza lincolnii) were the most abundant species in low-medium shrub habitats in the Tanana Valley, whereas they were rare in Denali National Park (this study) and near Fairbanks (West and De Wolfe 1974). In addition, the Common Snipe and Lesser Yellowlegs were much less abundant in Denali compared to farther east. In tall shrubs, the Yellow Warbler (Dendroica petechia) was the most abundant species in the Tanana Valley, but it was not nearly as abundant in Denali National Park (this study) or the Fairbanks area (West and DeWolfe 1974). There is increased interest in birds that breed in the Nearctic region and migrate to the Neotropics due to population declines in a number of species (reviewed by Hagan and Johnson 1992; Rappole and PATON AND POGSON: BIRDS BREEDING IN DENALI NATIONAL PARK 605 McDonald 1994). Yet no published studies have focused specifically on Neotropical migrants in inte- rior Alaska, even though they constitute the majority of birds breeding in the subarctic and boreal regions (Kessel and Gibson 1978; Spindler and Kessel 1980; Erskine and Davidson 1976). Erskine (1977) esti- mated that only 18% of the species breeding in the boreal region were permanent residents, whereas permanent residents accounted for approximately 58% of the breeding densities for sites across the lower 48 states (O’Connor 1992). Our results paral- lel those of Spindler and Kessel (1980) who calculat- ed that permanent residents accounted for only 11% of the individual breeding densities in conifer forests and 4% of the breeding densities in deciduous forests in interior Alaska. We found approximately 8% of the individuals at stations along roadside transects were residents, while 22% of the landbird species detected in the Park were residents. As the majority of birds breeding in interior Alaska are tropical migrants, ecosystem health of the boreal region is clearly dependent on large-scale issues. Residents might be affected by changes solely on their home range (e.g., logging), whereas migrants could be affected by habitat stability on their winter grounds, breeding grounds, or stopover locations (Rappole and McDonald 1994). Therefore, studies hoping to monitor northern avian populations will have consid- er large-scale issues in their study designs. The habitat uses we documented for landbirds in Denali were similar to those found in previous research conducted in interior Alaska. Both Spindler and Kessel (1980) and our results suggest that species richness is greatest in tall shrub habitats. However, Spindler and Kessel (1980) found that rel- ative abundance was greatest in tall shrub thickets, whereas our results suggest that low-medium shrub habitats had the greatest abundance. Our results gen- erally concur with Spindler and Kessel (1980), who found that spruce forests and woodlands appear to be among the most important habitats for residents in interior Alaska because relative abundance was greatest in these habitats. However, we also found that tall shrubs were important breeding and foraging habitats for residents, primarily redpolls. Spindler and Kessel (1980) suggested the high species rich- ness and abundance in tall shrubs was due to the combination of high primary productivity and com- plex habitat structure. In contrast, alpine tundra pos- sibly had low richness and abundance for landbirds due to the relatively simple habitat structure, although most shorebirds are found breeding in alpine tundra. A similar relationship was document- ed in the Kluane Mountains (Theberge 1976). Therefore, our results, coupled with work by Spindler and Kessel (1980), suggest that shrub habi- tats should be an integral component in future moni- toring efforts in interior Alaska because they provide 606 important breeding habitats for a large number of species, particularly tropical migrants. Our analyses of population fluctuations parallel those of Spindler and Kessel (1980). Species that were uncommon had higher fluctuations between years compared to common species. Abundant species, such as the White-crowned Sparrow and American Tree Sparrow, exhibited little annual vari- ation based on both frequency of occurrence and relative abundance estimates. The two exceptions were taxa that were classified as residents, the red- poll and White-winged Crossbill; both are known to be irruptive (Kessel and Gibson 1978). Our surveys did not distinguish whether these annual fluctua- tions represented actual changes in the population sizes of these uncommon taxa, or more plausibly for the irruptive taxa, a redistribution among alternative areas. More importantly, the results of this study can be used to design a monitoring program for cen- tral, interior Alaska. This information on detection probabilities is needed to conduct a power analysis to determine how many survey stations need to be censused annually to monitor avian population trends in this remote region of North America. In addition, it is hoped that this study will help researchers determine what suites of species could be monitored if various habitats were surveyed. Acknowledgments We thank J. Bouton, C. Dooley, A.-M. Barber, T. Eskelin, S. Hoover Paton, J. Mason, A. Wildman and C. Witt for assistance with the fieldwork. E. Rexstad, Institute of Arctic Biology, provided statistical sup- port, and J. Van Horn, National Park Service, provid- ed logistical support. Alaska Department of Fish and Game and the University of Rhode Island, College of Resource Development provided logistical support and computer services. L. Thorsteinson and D. Taylor, National Biological Service, Anchorage, were helpful with all aspects of the project. A. Erskine and two anonymous reviewers made helpful comments on earlier drafts. Financial support for this project was provided by the National Biological Service and National Park Service. Literature Cited Bart, J., and S. P. Klosiewski. 1989. Use of presence- absence to measure changes in avian density. Journal of Wildlife Management 53: 847-852. Dixon, J.S. 1938. Birds and mammals of Mount McKinley National Park, Alaska. Fauna of the National Parks of the United States, Fauna Series Number 3. 236 pages. Dobkin, D. S. 1994. Conservation and management of Neotropical migrant landbirds. University of Idaho Press, Moscow. 220 pages. Erskine, A. J. 1977. Birds in boreal Canada: communi- ties, densities, and adaptations. Canadian Wildlife Service Report Series 41: 1-73. Erskine, A. J., and G. S. Davidson. 1976. Birds in the Fort Nelson lowlands of northeastern British Columbia. Syesis 9: 1-11. THE CANADIAN FIELD-NATURALIST Vol. 110 Gauthreaux, S. A. 1991. Preliminary lists of migrants for Partners in Flight Neotropical migratory bird conserva- tion program. Partners in Flight 2: 30. Hagan, J. M., III, and D. W. Johnston. Editors. 1992. Ecology and conservation of Neotropical migrant landbirds. Smithsonian Institution Press, Washington. 609 pages. : Hanowski, J. M., and G. J. Niemi. 1995. A comparison of on- and off-road bird counts: Do you need to go off the road to count birds accurately? Journal of Field Ornithology 66: 469-483. Hoefs, M. 1973. Birds of Kluane Game Sanctuary, Yukon Territory, and adjacent areas. Canadian Field-Naturalist 87: 345-355. Kertell, K. 1988. Bird checklist for Denali National Park. Alaska Natural History Association, Denali National Park. 3 pages. Kessel, B. 1979. Avian habitat classification for Alaska. Murrelet 60: 86-94. Kessel, B., and D. Gibson. 1978. Status and distribution of Alaska birds. Studies in Avian Biology Number 1: 1—100. Laing, K. 1985. Food habits and breeding biology of Merlins in Denali National Park, Alaska. Raptor Research 19: 42-51. Murie, A. 1963. Birds of Mount McKinley National Park. Alaska. Mt. McKinley Natural History Association, McKinley Park, Alaska. 86 pages. O’Connor, R. J. 1992. Population variation in relation to migratory status in some North American birds. Pages 64-74 in Ecology and conservation of Neotropical migrant landbirds. Edited by J. M. Hagan, III, and D. W. Johnston. Smithsonian Institution Press, Washington, D.C. 609 pages. Ralph, C. J., J. F. Sauer, and S. Droege. Editors. 1994. Monitoring bird population trends by point counts. USDA Forest Service, Pacific Southwest Research Station, PSW-GTR-149, Albany, California. Rappole, J. H. and M. V. McDonald. 1994. Cause and effect in population declines of migratory birds. Auk 111: 652-660. Rappole, J. H. 1995. The ecology of migrant birds: a Neotropical perspective. Smithsonian Institution Press, Washington, D.C. 269 pages. SAS Institute. 1988. SAS user’s guide statistics. SAS Institute Inc., Cary, North Carolina. 554 pages. Spindler, M. A., and B. Kessel. 1980. Avian populations and habitat use in Interior Alaska taiga. Syesis 13: 61-104. Theberge, J. B. 1976. Bird populations in the Kluane Mountains, southwest Yukon, with special reference to vegetation and fire. Canadian Journal of Zoology 54: 1346-1356. Troy, D. M. 1985. A phenetic analysis of the redpolls Carduelis flammea flammea and C. hornimanni exilipes. Auk 102: 82-96. Viereck, L. A., C. T. Dyrness, A. R. Batten, and K. J. Wenzlick. 1992. The Alaska vegetation classification. United States Department of Agriculture, Forest Service, General Technical Report, PNW-GTR-286. 278 pages. West, G. C., and B. B. DeWolfe. 1974. Populations and energetics of taiga birds near Fairbanks, Alaska. Auk 91: 757-775. Received 15 March 1995 Accepted 23 July 1996 Historical Occurrence of Wolves, Canis lupus, in the Maritime Provinces CHRISTINE LOHR and WARREN B. BALLARD! New Brunswick Cooperative Fish and Wildlife Research Unit, P. O. Box 44555, University of New Brunswick, Fredericton, New Brunswick E3B 6C2 'Present address and author to whom correspondence may be addressed: Arizona Game and Fish Department, 2221 West Greenway Road, Phoenix, Arizona 85023-4399. Lohr, Christine, and Warren B. Ballard. 1996. Historical occurrence of Wolves, Canis lupus, in the Maritime Provinces. Canadian Field-Naturalist 110(4): 607-610. We conducted a literature review and contacted several museums to determine the historical occurrence of Wolves (Canis lupus) in the Maritime Provinces. Although there were many anecdotal sightings prior to 1870, no museum specimens originating from the Maritimes were located. Our review suggested that although Wolves were historically present, they were probably not numerous, and were probably extirpated from the Maritime Provinces between 1870 and 1921. Key Words: Gray Wolves, Canis lupus, abundance, distribution, historical, New Brunswick, Nova Scotia. Historically, Wolves (Canis lupus) were distrib- uted across most of North America. They were extir- pated in the southern portion of their range (most of continental United States) by the turn of the century (Nowak 1983; Carbyn 1987). Although healthy Wolf populations exist in Canada and Alaska, the Wolf is considered “vulnerable” by the International Union for the Conservation of Nature (1990). Within recent years there has been considerable interest in restoring Wolves to their former ranges. Reintroduction efforts are currently underway in Wyoming and Idaho (Fritts 1990; Bath 1992; U.S. Department of the Interior 1994). Reintroduction plans are also being designed for the southwestern United States (Bednarz 1988, 1989). An understand- ing of the historical occurrence of extirpated popula- tions is important when considering reintroduction into formerly occupied range. The historical occurrence of Wolves in the north- east portion of the continent has recently been ques- tioned (Kenney in press). If viable Wolf populations have not occurred historically within the Maritime Provinces, then establishment of a Wolf population would be considered a new introduction rather than a reintroduction and as a result, the rationale for eco- logical restoration would disappear. Despite pro and con arguments for ecological restoration, Wolf reintroduction may be prevented if there is sufficient social resistance. Bath (1987) determined that public willingness to reintroduce Wolves to Yellowstone National Park depended on the social group’s interest. For example, although members of the general public were in favor of Wolf reintroduction to Yellowstone, Wyoming Stock Growers were generally opposed, while members of the Wyoming Wildlife Federation were generally in favor of it (Bath 1992). The purpose of this report was to review available literature and evidence to determine if historic Wolf distribution included the Maritime Provinces and in particular, New Brunswick. Methods We conducted a literature search concerning the historic distribution of Wolves within the Maritime Provinces. We also contacted museums within Canada, United States, and United Kingdom to determine if Wolf specimens and records from the Maritime Provinces existed. Since Wolves never occupied Prince Edward Island (Carbyn 1987) it was excluded. Historical Distribution of Wolves Anecdotal Evidence Indian folklore provides the oldest reference to the Wolf in northeastern North America. To the Micmac Indians of New Brunswick, the Wolf was called Malsum, and was the personification of evil (Leland 1884). To the Malecite Indians of New England, a Wolf howl meant death (Wallis and Wallis 1957). One of the earliest non-mythical descriptions of the natural history of Acadia (which includes New Brunswick and Nova Scotia) notably did not men- tion the Wolf in its list of land mammals (Denys 1908). LeClerq (1910) reported that the Wolf was observed in 1691 either on the Gaspe Peninsula, or in northern New Brunswick. In neighboring New England, Wolves were appar- ently abundant in the mid to late-1600s. Josselyn (1672: 15) described the Wolves he encountered in New England as follows: “very numerous, and go in companies, sometimes ten, twenty, more or fewer, and so cunning...”. He observed three Wolves in August 1638 and two in 1664 (Josselyn 1674). 607 608 In Clarke’s (1670: 32) list of the “beasts” of New England he mentioned the “ravenous howling Woolf”. He described their morphology as “big- boned, thin paunched, deep breasted, having a thick neck and head, prick ears, and a long snout, with dan- gerous teeth, long staring hair, and a great bush tail”. Two early writers claimed that Wolves were not indigenous to New Brunswick and became numer- ous only in the mid-1800s (Gesner 1842, 1847; Levinge 1846). Gesner (1842) suggested that Wolves appeared in New Brunswick in 1818. Levinge (1846) also claimed that Wolves were not indigenous to New Brunswick and that they had made their appearance because they were driving White-tailed Deer (Odocoileus virginianus) from the eastern United States. He recounted a story about a local resident who slaughtered a Wolf pack in New Brunswick. Baird (1890) recollected a deer hunt in 1846 near Woodstock, New Brunswick, where he and his companions were awakened by the howling of Wolves at night. Hatheway (1846) also noted an increase in Wolves in the 1840s but he indicated the Wolves had returned to New Brunswick after a period of rarity. Fisher (1838) observed the same low frequency of Wolves before the 1840s. He stated that Wolves were “but seldom seen” in 1836 and 1837. The observed insur- gence of Wolves in the 1840s (Gesner 1842, 1847) may only have been temporary because by the 1870's their numbers were apparently again greatly reduced. Adams (1873) reported that the Grey Wolf only occurred within the northern portion of New Brunswick, and Ganong (1908) reported that Wolves were absent or rare after the 1860s. It was shortly after this time period that Wolves were completely extirpated from New Brunswick (Carbyn 1987). Commercial Records and Museum Specimens Simonds and White supplied settlers on the Saint John River and often received payment with furs (Raymond 1910). From 1764 until outbreak of the revolution in 1774, the Company received two Wolves from Nova Scotia. In 1792, a bounty on Wolves was instituted in New Brunswick. This act of legislation sought “to encourage the destroying of Wolves” on the basis that “many losses have been suffered by sundry inhabitants of this Province, from the destruction of their Sheep by Wolves, to the great discouragement of the increase of that valuable Stock” (Berton 1838: 236). From 1792 up to and including 1793, a total of five Wolves were bountied (Accounts of the Provincial Treasurer, 1793-1794). The last bounties were paid in New Brunswick for three Wolves in 1862 (Ganong 1908). Unfortunately, records of the fur trade by the Hudson Bay Company were incomplete for early French harvests and for the northeastern United States (Novak et al. 1987). Therefore, it was not THE CANADIAN FIELD-NATURALIST Vol. 110 known how many Wolf pelts were acquired in the Maritime Provinces from early human colonization until the Wolf’s extirpation in 1870. Although Wolf pelts harvested in Canada were recorded as early as 1699, it was not until 1919 that these were separated by Province. For example, from 1700 until 1775 the, numbers of Wolf pelts exported to England increased from 483 to 7608, but the origin of these pelts was unknown (Lawson 1943). However, we do know that approximately £78 of fur (all species) was exported from Nova Scotia in 1765. From 1919 to 1984, reportedly four Wolf pelts were recorded from New Brunswick (Novak et al. 1987); three in 1919 and one in 1921. During 1995 we contacted the following museums to determine if any contained Wolf specimens which originated from the Maritime Provinces: American Museum of Natural History, New York, N.Y.; James Ford Bell Museum of Natural History, Minneapolis, Minnesota; Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts; National Museum of Natural Sciences, National Museums of Canada [now Canadian Museum of Nature], Ottawa, Ontario; National Museum of Scotland, Edinburgh, Scotland; Natural History Museum, London, England; New Brunswick Museum, St. John, N.B.; Nova Scotia Museum, Halifax, N.S.; Peabody Museum of Natural History, Yale University, New Haven, Connecticut; and Royal Ontario Museum, Toronto, Ontario. Although all of the museums contained Wolf specimens or records of Wolves, none of the specimens were known to originate from the Maritime Provinces. Discussion Available literature suggested that Wolves may have been more abundant in New England than in New Brunswick during the 1600s. In the late 1700s, Wolves also appear to have been uncommon based on the low numbers that were bountied in New Brunswick and the relatively low numbers in relation to Beaver (Castor canadensis) pelts that were traded to Simonds and White (Raymond 1910). However, fur trade records were biased towards recording Beaver pelts (Innis 1962). Consequently, use of Wolf pelts as an index of Wolf numbers may have been inaccurate. Although there was insufficient evi- dence from this early period to draw firm conclu- sions concerning Wolf abundance, they appeared to be present in relatively low numbers. The notable lack of actual Wolf specimens or records of Wolf specimens from the Maritime Provinces could have been due to several factors: (1) there were no Wolves, or so few, that specimens were not obtained; (2) fur specimens were not kept because the fur was not valuable or they were exported; (3) the disregard for the animal provided no incentive to collect specimens of it; and (4) speci- 1996 mens prior to extirpation have been lost, destroyed, or deteriorated. There appears to have been a consensus that Wolves were low in abundance in New Brunswick prior to the 1840s (Fisher 1838; Hatheway 1846), and that they became more numerous during the 1840s (Gesner 1842, 1847; Baird 1890; Hatheway 1846). The period of Wolf scarcity during the late 1700s and early 1800s through to the 1830s (Hatheway 1846) was likely the reason that some people falsely interpreted Wolf re-appearance in the 1840s as a range expansion. Although we can not verify what canid species early writers actually observed, it appears likely that the identification of the Wolf was probably correct for at least two reasons. First, early colonists were probably familiar with Wolves from their native European countries. Secondly, Coyotes (Canis latrans) though present in New Brunswick approximately 2000 years ago (Sanger 1987) did not significantly recolonize the Maritime Provinces until the 1960s (Moore and Parker 1992). Coyotes probably first re-entered Maine in the 1930s (Nowak 1983), while the earliest known “‘contem- porary” coyote in New Brunswick was reported in 1958 (Wolfram 1964). It was not a moral question to kill Wolves during the early colonization of the Maritime Provinces and New England. Because Wolves killed domestic live- stock such as Goats (Josselyn 1674) and Sheep (Berton 1838) they were hunted with vehemence. This may partially account for Wolves becoming rel- atively scarce prior to the 1840s. Wolf population fluxes in the Maritime Provinces may also have been caused by changes in prey avail- ability. Gesner (1842) and Levigne (1849) suggested that Wolves came into New Brunswick in pursuit of Deer. During the “little ice age” (1350 to 1870), Caribou (Rangifer tarandus) flourished in the Maritime Provinces (F. Scott, Nova Scotia Museum, personal communication). The decline and eventual extirpation of Caribou in New Brunswick in the early 1900s (Squires 1968) may have contributed to the Wolf’s disappearance during this period. Because Wolves were apparently extirpated between 1870 and 1921, evidence for their occur- rence in the Maritime Provinces is based on anecdo- tal observations. However, these types of observa- tions are frequent enough to definitely conclude that Wolves occurred at least sporadically or in low num- bers. Whether they occurred in numbers high enough to constitute a minimum viable population can not be ascertained from existing evidence. Acknowledgments Funding for this study was provided by the New Brunswick Cooperative Fish and Wildlife Research Unit and the Faculty of Forestry and Environmental LOHR AND BALLARD: WOLVES IN THE MARITIME PROVINCES 609 Management. T. Dilworth, G. J. Forbes, G.R. Parker, and M. J. Schousboe provide constructive criticism on early drafts. Literature Cited Adams, A. L. 1873. Field and forest rambles, with notes and observations on the natural history of eastern Canada. Henry S. King and Co., London. 333 pages. Baird, W. T. 1890. Seventy years of New Brunswick life: autobiographical sketches. Press of Geo. E. Day, Saint John, New Brunswick. 358 pages. Bath, A. J. 1987. Attitudes of various interest groups in Wyoming toward wolf reintroduction in Yellowstone National Park. M. A. thesis, University of Wyoming, Laramie. 123 pages. Bath, A. J. 1992. Identification and documentation of public attitudes toward wolf reintroduction in Yellowstone National Park. 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Management of wolves inside and out- side Yellowstone National Park and possibilities for wolf management zones in the greater Yellowstone area. Pages 1-3 to 1-88 in Wolves for Yellowstone? A report for the U. S. Congress, Volume II Research and Analysis. Yellowstone National Park, Mammoth, Wyoming. 610 Ganong, W. F. 1908. On the occurrence of the wolf in New Brunswick. Bulletin Natural History Society, New Brunswick 26: 30-35. Gesner, A. 1842. Fourth report on the geological survey of the province of New Brunswick. Henry Chubb, Market Square, Saint John, N.B. 101 pages. Gesner, A. 1847. New Brunswick; with notes for emi- grants. Simmonds and Ward, London. 388 pages. Hatheway, C.L. 1846. The history of New Brunswick, from its first settlement, containing a geographical description of the province; its boundaries, rivers, lakes, streams, and division into countries and parishes; also its climate, soil, fisheries, mines and minerals, animals, birds, and C.; government, and effects of emigration, capabili- ties for further settlement, trade and exports, internal com- munications, character of inhabitants, and of the aborig- ines, religion, and C. James P. A. Phillips “Head Quarters” Office, Fredericton, New Brunswick. 81 pages. Innis, H. A. 1962. The fur trade in Canada: an introduc- tion to Canadian economic history. The University of Toronto Press. Toronto, Ontario. 446 pages. International Union for Conservation of Nature and Natural Resources. 1990. Foxes, Wolves, Jackals, and Dogs: An Action Plan for the Conservation of Canids. International Union for the Conservation of Nature and Natural Resources. Gland, Switzerland. 116 pages. Josselyn, J. 1672. 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Received 18 July 1995 Accepted 14 August 1996 Keen’s Long-eared Bat, Myotis keenii, Confirmed in Southeast Alaska DOREEN I. PARKER! and JOSEPH A. COOK University of Alaska Museum, 907 Yukon Drive, Fairbanks, Alaska 99775 1Current address: Alaska Department of Fish and Game, Division of Wildlife Conservation, P.O. Box 25526, Juneau, Alaska 99802-5526 Parker, Doreen I., and Joseph A. Cook. 1996. Keen’s Long-eared Bat, Myotis keenii, confirmed in southeast Alaska. Canadian Field-Naturalist 110(4): 611-614. Myotis keenii is apparently restricted to the Pacific coastal forests of northwestern North America. The only documentation of M. keenii in Alaska has been a specimen collected in 1887, causing uncertainty about whether this species normally occurs there. We describe two new records which indicate that M. keenii may be a regular member of the Southeast Alaska fauna and we provide measurements and information on diet for this poorly documented species. Key Words: Keen’s Long-eared Bat, Myotis keenii, Vespertilionidae, southeast Alaska, temperate rainforest. Of the five vespertilionid species occurring in Southeast Alaska (MacDonald and Cook 1996), the Keen’s Long-eared Bat (Myotis keenii) was known from a single specimen taken at Wrangell in 1887 (Miller and Allen 1928). Since then, the possibility persisted that it was an accidental occurrence. The original specimen was preserved in alcohol and the skull was not available to confirm its identification. Consequently, its identity remained uncertain until van Zyll de Jong and Nagorsen (1994) used multiple discriminant analysis of five external measurements to diagnose the specimen as M. keenii. Because M. keenii is distributed throughout coastal rainforests in the Pacific Northwest (van Zyll de Jong and Nagorsen 1994), extensive timber harvest in Southeast Alaska has increased interest in the status of this species. Myotis keenii was previously regarded as conspe- cific with the Northern Long-eared Bat (M. septen- trionalis), but now is considered a separate species (van Zyll de Jong 1979). These species are difficult to distinguish from each other, and from the Western Long-eared Bat (M. evotis), which is sympatric with M. keenii in British Columbia and Washington. The three species can be identified by discriminant func- tion analysis of cranial (van Zyll de Jong 1979) or external (van Zyll de Jong and Nagorsen 1994) mea- surements. We provide information on two new specimens of this apparently uncommon species in Southeast Alaska including morphological measurements and diet data. Previously, only 59 specimens of M. keenii had been collected and deposited in museum collec- tions. Thirty-five are from the Queen Charlotte Islands of British Columbia, nine from other regions of British Columbia, 14 from western Washington, and one from Wrangell Alaska (Figure 1; van Zyll de Jong and Nagorsen 1994). The known range of M. keenii is restricted to Pacific coast rainforests 611 (Nagorsen and Brigham 1993; van Zyll de Jong and Nagorsen 1994), and extends over 2000 km from southwestern Washington to Southeast Alaska (Figure 1). The rarity of this species and lack of eco- logical data have prompted the British Columbia Ministry of Environment to place M. keenii on the provincial “red list” of species under consideration for listing as threatened or endangered (Nagorsen and Brigham 1993). The species has no special con- servation status in the United States. Methods During 18 nights in June, July, and August 1993, mist nets were placed in riparian areas on Prince of Wales and Revillagigedo islands in Southeast Alaska. In 1994 bats were collected from a maternity roost of Little Brown Bats (M. lucifugus) at Hoonah on Chichagof Island (58°06'N, 135°26’ W; Figure 1). Captured bats were prepared as voucher speci- mens. Specimens and frozen tissue samples are archived at the University of Alaska Museum. Stomach contents collected in 1993 were preserved in 70% ethanol and analyzed for prey volume. Species identity was determined using cranial mea- surements (van Zyll de Jong 1985) and verified using multiple discriminant analysis (MDA) of 12 cranial measurements (van Zyll de Jong 1979; van Zyll de Jong and Nagorsen 1994) and 5 external measurements (van Zyll de Jong and Nagorsen 1994) listed in Table 1. Further verification was obtained using MDA of the 7 cranial variables (Table 1) which best discriminate between M. keenii and M. evotis (van Zyll de Jong and Nagorsen 1994). Results On 20 July 1993 an adult male M. keenii (UAM 23338) was collected at Turn Creek, in a karst region of northern Prince of Wales Island (56°10’'N, 133°18’W), approximately 65 km SW of Wrangell 612 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Measurements of new Myotis keenii specimens. Mean + | SD. UAM 23338 UAM 29831 Mean M. evotis from from from +1SD. British Measurements (mm) Turn Creek Hoonah M. keenii* Columbia* Cranial Measurements: Skull length? 14.93 14.38 14.60 + 0.219 15.45 + 0.307 Mastoid width+ 6.91 6.88 7.43 + 0.135 7.76 + 0.167 Orbital width at lacrimal foramina 4.45 4.75 4.30 + 0.130 4.57 + 0.130 Rostral widtht 2.44 2.50 3.33 + 0.093 3.63 + 0.114 Maxillary width at M3+ 5.64 5.80 5:52!+'0/121 5.85 + 0.128 Palatal width at P2+ 3.64 3.65 3.48 + 0.097 3.66 + 0.141 Maxillary width at I3 Dunit 2.63 2.45 + 0.074 2.61 + 0.071 Maxillary tooth row length 5.68 5.63 5.66 + 0.105 6.11 + 0.155 Length of P4M3+ 3.89 3.98 3.96 + 0.072 4.33 + 0.113 Length of M2 1.24 1.26 1.23 + 0.046 137101052 Width of M2 1.68 1.48 1.69 + 0.061 1.85 + 0.052 Upper canine width at cingulumy 0.69 0.62 0.72 + 0.026 0.83 + 0.035 External Measurements: Ear length 17 15 18.37 + 0.491 20.38 + 0.809 Forearm length 37.28 35139 36.70 + 0.986 38.64 + 1.117 Tibia length 17.00 16.08 16.58 + 0.659 17.79 + 0.681 Metacarpal 3 length 31.88 31.98 31.75 + 0.975 34.16 + 1.266 Metacarpal 5 length 31.64 29.66 30.99 + 0.822 33.03 + 1.142 Measurements are defined in van Zyll de Jong (1979). *From van Zyll de Jong and Nagorsen (1994). ymeasurements which best descriminate between M. keenii and M. evotis (van Zyll de Jong and Nagorsen 1994). (Figure 1). This bat was captured in a mist net at 23:20 (2 h 10 min after sunset) within 1 m of a lime- stone cliff and 1 m above the water. Large Western Hemlock (Tsuga heterophylla) and Sitka Spruce (Picea sitchensis) dominate this riparian area; blue- berry (Vaccinium spp.) and Devil’s-club (Oplopanax horridus) dominate the understory. The stomach of the Turn Creek bat contained 40% Trichoptera, 40% Araneae and 20% Diptera; the first indication of dietary habits. A second adult male M. keenii (UAM 29831) was collected on 11 July 1994 from a M. lucifugus maternity roost in the attic of an operating fish can- nery at Hoonah, approximately 160 km N of the Wrangell specimen (Figure 1). That site is surround- ed by large clearcuts and second-growth forest, as well as forested wetlands and riparian areas. Body and cranial measurements for both speci- mens, as well as means (+ 1 standard deviation) for M. evotis from British Columbia and M. keenii are summarized in Table 1. Multiple discriminant analy- sis (MDA) of 12 cranial measurements (van Zyll de Jong 1979) identified the Turn Creek (UAM 23338) and Hoonah (UAM 29831) specimens as M. keenii with probabilities of 88% and 85%, respectively. The MDA of 7 cranial measurements which best dis- criminate between M. keenii and M. evotis (van Zyll de Jong and Nagorsen 1994) identified both speci- mens as M. keenii with probabilities of 98%. MDA of the 5 external measurements (Table 1) identified both specimens as M. keenii (probability 100%), even though the ear of the Hoonah specimen is > 6 standard deviations smaller than the published mean (van Zyll de Jong and Nagorsen 1994). Discussion While only three specimens of M. keenii have been recorded in southeast Alaska, little effort has been expended to investigate bat distribution in that region (Parker 1996). Myotis keenii may be a year- round resident of Southeast Alaska, although winter records are lacking. All specimens of M. keenii have been found in the Pacific Northwest temperate rain- forest ecosystem (Walter 1985), a region with rela- tively mild winters and potentially numerous roost and hibernation sites in caves and trees. Seasonal occurrence of this species, including the documenta- tion of female M. keenii and maternity colonies, should be investigated. Southeast Alaska’s temperate rainforests are structurally complex with abundant live trees, snags, and fallen logs of various sizes (Alaback 1991). Such complexity provides loose bark and tree hol- lows suitable for cavity-roosting species (Bunnell and Allaye-Chan 1984) such as bats (Barclay and Cash 1985; Christy and West 1993; Bradshaw in press; Vonhof in press). Bats use old-growth forests more frequently than second-growth or clearcut 1996 @ Previous sites @® Turn Creek site [x] Hoonah site 0°. 5) Vancouver ————— FicureE 1. Locality records of Myotis keenii in the Pacific Northwest. Adapted from data presented by van Zyll de Jong and Nagorsen (1994). areas in British Columbia, Washington, and Oregon (Barclay and Cash 1985; Lunde and Harestad 1986; Thomas 1988; Christy and West 1993; Bradshaw in press; Vonhof in press), as well as in Southeast Alaska (Parker et al. in press). Caves and crevices are also important bat habitat (Hill and Smith 1984), and over 1769 km? of cave and crevice-containing karst underlie Southeast Alaska forests (United States Department of Agriculture 1996). These habi- tat characteristics may be essential to M. keenii as the species appears to be restricted to coastal forests (Firman et al. 1993). Although limited, our data represent the only diet information available for M. keenii. The mixture of flying insects (60%) and nonflying spiders (40%) consumed suggests M. keenii has a flexible foraging strategy, pursuing prey in flight and gleaning it from surfaces. Similar foraging behavior has been noted for M. evotis (Barclay 1991; Faure and Barclay 1994), a closely related species (van Zyll de Jong and Nagorsen 1994). However, more sampling is required to firmly establish this foraging strategy because bat diets tend to change with season and relative abundance of different prey species (Buchler 1976; Fenton and Morris 1976; Anthony and Kunz 1977). The Turn Creek bat was captured in a riparian area, and its stomach contained a high percentage of Trichoptera, a typically riparian PARKER AND COOK: KEEN’S LONG-EARED BAT IN SOUTHEASTERN ALASKA 613 insect. Other prey occur throughout old-growth forests and riparian areas indicating that M. keenii forages in those areas. Roost requirements of M. keenii are poorly under- stood (Nagorsen and Brigham 1993), but it probably uses snags, hollow trees, rock crevices and caves (van Zyll de Jong 1985). The capture of the Hoonah specimen from a maternity roost of M. lucifugus in an operating fish cannery indicates that this species will roost in a human-occupied building. Myotis keenii roosts in association with M. lucifugus under rocks heated by a hot spring on Hot Spring Island in the Queen Charlotte Islands of British Columbia (Nagorsen and Brigham 1993). The Hoonah speci- men may have been roosting in the noisy cannery due to a lack of undisturbed roosts in the surround- ing area. Nonetheless, this species may tolerate peri- odic disturbance. Bats at the Hot Spring Island colony must abandon their roost periodically when it floods at high tide (Firman et al. 1993). It is uncer- tain whether these observations reflect a tolerance of disturbance or lack of alternate, suitable roosts. Forty-two percent of the highly productive forests in Southeast Alaska were clearcut harvested by 1990 (United States Department of Agriculture 1991; 1993), including over 70% of the karstland forests of Prince of Wales and neighboring islands (Baichtal 1995). These forests contain over 30 000 board feet of useable timber per acre (volume class 6 and 7, approximately 348 m? per hectare; Dilworth 1976). Continued clearcut harvesting may alter forest struc- ture important to bats (Thomas 1988). In fact, bat activity is rare in second-growth forests of Southeast Alaska (Parker et al. in press). In view of the limited knowledge of the habitat requirements of M. keenii, its apparently strong association with old-growth coastal rainforests, and the continuing harvest of these forests in Southeast Alaska, this species and its habitat requirements warrant further and immediate study. Acknowledgments We thank the following individuals for their assis- tance. C. 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United States Department of Agriculture Forest Service, Juneau, Alaska. Pages 1-765. United States Department of Agriculture. 1993. Timber supply and demand 1992. Report Number 12, September 1993. United States Department of Agriculture Forest Service, Juneau, Alaska. 40 pages. United States Department of Agriculture. 1996. Tongass land management plan environmental impact statement. United States Department of Agriculture Forest Service, Juneau, Alaska. van Zyll de Jong, C. G. 1979. Distribution and systemat- ic relationships of the long-eared Myotis in western Canada. Canadian Journal of Zoology 57: 987-994. van Zyll de Jong, C.G. 1985. Handbook of Canadian mammals, volume 2: bats. National Museum of Natural Science, National Museum of Canada, Ottawa. 212 pages. van Zyll de Jong, C.G., and D. W. Nagorsen. 1994. A review of the distribution and taxonomy of Myotis keenii and Myotis evotis in British Columbia and the adjacent United States. Canadian Journal of Zoology 72: 1069-1078. Vonhof, M. Jn press. Roosting ecology and roost-site preference of reproductive Eptesicus fuscus and Lasionycteris noctivagans in the Pend d’Oreille Valley in southern British Columbia. /n Proceedings of the Bats and Forest Symposium 19-21 October 1995. Edited by R.M.R. Barclay and R. M. Brigham. Victoria, British Columbia. Walter, H. 1985. Vegetation of the earth and ecological systems of the geo-biosphere. Third edition. Springer- Verlag, New York. Pages 181-187. Received 16 October 1995- Accepted 5 August 1996 Pumpkin Ash, Fraxinus profunda, in Southwestern Ontario G. WALDRON!, M. GARTSHORE?, AND K. COLTHURST? '7641 Highway 18, RR 1, Amherstburg, Ontario N9V 2Y7 °RR 1, Walsingham, Ontario NOE 1X0 3Essex Region Conservation Authority, 360 Fairview Avenue West, Essex, Ontario NSM 1Y6 Waldron, G., M. Gartshore, and K. Colthurst. 1996. Pumpkin Ash, Fraxinus profunda, in southwestern Ontario. Canadian Field-Naturalist 110(4): 615-619. An addition to the flora of Canada, Pumpkin Ash (Fraxinus profunda), is reported from the southwestern counties of Essex, Kent, Elgin and Haldimand-Norfolk. This is a native tree species of swamp forests. The species is most readily dis- tinguished from other ashes by its relatively large samaras 4 — 7.5 cm long. Key Words: Pumpkin Ash, Fraxinus profunda, tree species, new plant record, southwestern Ontario. Prior to Farrar’s (1995) recent report (based on collections of the authors), the range of Pumpkin Ash, Fraxinus profunda (Bush) Bush, had not included Canada (e.g., Little 1977; Duncan and Duncan 1988; Preston 1989). Nor has the described range included the state of Michigan or northern Ohio. However, in 1992, botanists with the Ohio Department of Natural Resources found the species in northern Ohio, prompting Michigan botanists to look for it successfully, the same year. In Michigan, Pumpkin Ash was found in the southern two tiers of counties adjacent to Ohio. In October 1992, the senior author examined the Michigan collections at the University of Michigan, Ann Arbor, and was advised by the curator, A. A. Reznicek, to look for the species in very wet, wooded sites in southwest- ern Ontario. The first site examined in November 1992, at the Devonwood Conservation Area in Windsor yielded the distinctive large samaras of Pumpkin Ash. Subsequent examinations of swamps across southwestern Ontario have revealed the species to be a common associate of swamp commu- nities. It is remarkable that such a distinctive tree should have escaped attention. Ash trees which, in the past, the authors and others (W. Balkwill, per- sonal communication) found difficult to identify have been found, upon re-examination, to be Pumpkin Ash. The described range of Pumpkin Ash has expand- ed considerably as familiarity with the species has increased. Bush, the accepted author of the species, published the range in 1894 as Dunklin and New Madrid counties in Missouri. In 1897, he extended the range to Arkansas and Florida. Britton, in 1908, described the range as “known to occur from Missouri, Illinois, and eastern Arkansas to Virginia and Florida”. Deam (1921) located it in the south- western counties of Indiana and by 1922 Sargent had it in western New York, southern Indiana and Illinois, western Kentucky and south to Louisiana and Florida. In 1953, Little gave a similar range but included southern Ohio. Braun (1961) mapped it in just two counties of southwestern Ohio near the Ohio River. The Pumpkin Ash range map in Little (1977) shows disjuncts through central and northern Indiana and southwestern Ohio, but nothing in western New York. Apparently, every range map published since 1977 (Elias 1980; Duncan and Duncan 1988; Preston 1989; McCormac 1993) is based on the map Little (1977). In Ontario, the range, based on collections and observations made since 1992, is restricted to an area west of Turkey Point, Regional Municipality of Haldimand-Norfolk (Table 1), in the Niagara Section of the Deciduous Forest Region as defined by Rowe (1977), (Figure 1). The species should be looked for in swamps near Lake Erie east of Turkey Point. The nomenclatural history of Pumpkin Ash is interesting. Bush (1894) described it as a variation of Fraxinus americana, F. americana profunda, based on differences “in the strong pubescence of the shoots, the large size of the leaves, and the very large fruit, the shaft of which is strongly six-sided”. Bush revised his opinion in 1897 (Bush 1897) and made the variety a full species, F. profunda. Synonyms have included F. michauxii Britton (1905) and F. tomentosa Michx. f., the latter proposed by M. L. Fernald in 1938, based on an illustration and descrip- tion published by Francois André Michaux in 1813. F. tomentosa was adopted subsequently by many authors (Fernald 1950; Braun 1961; McCormac 1993), but it is an illegitimate name as noted by Little (1953). Pumpkin Ash is a medium to tall tree, up to 40 metres (Sargent 1933; Fernald 1950). It is a hexa- ploid with n= 138 versus n=46 for most other ash species in eastern North America (Wright 1957). This higher ploidy is reflected in the large size of the species’ organs; the leaves, for example, are 23-45 centimetres in length. Leaflets number 7 or occasion- ally 9, are usually tomentose when unfolding, but later glabrous and often glossy above. The leaflet 615 616 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Pumpkin Ash collections in Ontario. The locations are mapped in Figure 1. Location 1 Springwater Conservation Area, Elgin County UTM 968 322 (map 401/11) 2 Lot C, Conc. II, Gosfield South Township, Essex County UTM 502 581 (map 40J/2) 3 Canard River, 1.4 km SE Gesto, Essex County UTM 458 655 (map 40J/2) 4 Maidstone Conservation Area, Essex County UTM 522 747 (map 40J/2) 5 Rondeau Provincial Park, Kent County UTM 295 835 (map 40/5) 6 West Lorne, Elgin County UTM 516 087 (map 401/12) 7 4km W West Lorne, Elgin County UTM 455 168 (map 401/12) 8 Lot I, Conc. XII, Aldborough Township, Elgin County 9 7km.E. Eagle, Elgin County UTM 547 125 (map 401/12) 10 Jeanette’s Creek Woods, Kent County UTM 916 864 (map 40J/8) 11 Leamington White Oak Woods Environmentally Significant Area, Essex County UTM 736 602 (map 40J/2) 12 Brunet Park, LaSalle, Essex County UTM 312 786 (map 40J/3) 13. 6km NNW Rodney, Elgin County UTM 422 188 (map 401/12) 14 Conc. I, South Walsingham Township Regional Municipality of Halidmand-Norfolk UTM 405 205 (map 401/9) 15 Tilbury West Conservation Area, Essex County UTM 771 776 (map 40J/1) 16 Fish Point Provincial Park Reserve, Pelee Island, Essex County UTM 604 218 (map 40G/10) margin is entire or slightly serrate. Unlike Red Ash, but similar to White Ash, the leaflets have distinct petiolules without the blade decurrent nearly to the base (Figure 2). The twigs are thick and, at least when young, pubescent. The trunk in forest-grown trees is long and slender from a flaring or swollen, and hence pumpkin-like, base from which the tree gets its popular name (Figure 3). The swollen base is seen on some Ontario specimens, but this trait is apparently more commonly exhibited by the species in the swamps of the southern United States. The bark on older trees most commonly develops into solid, continuous ridges similar to those of White Ash and not flaky, broken ridges like those of Collector Collection No. Tan D. MacDonald 14432 UWO 40444 (identified as F. pennsylvanica) G. E. Waldron MICH A. A. Reznicek M. J. Oldham 14665, MICH, TRTE A. A. Reznicek, G. E. Waldron M. J. Oldham 14946, TRTE M. E. Gartshore MICH M. E. Gartshore MICH M. E. Gartshore MICH Wm. Stewart 3684 UWO 43417 Identified as F. tomentosa (= F. profunda) Collection Date 3 September 1983 December 1992 18 May 1993 15 June 1993 22 June 1993 23 June 1993 23 June 1993 27 June 1993 R. Vanderjeugd M. J. Oldham 15917, MICH, TRTE M. J. Oldham 15390 10 July 1993 12 August 1993 MICH M. J. Oldham 15418 14 August 1993 MICH, TRTE G. E. Waldron 15 September 1993 MICH A. A. Reznicek M. J. Oldham 15860, MICH M. E. Gartshore Photo Fax NHIC 29 September 1993 3 June 1994 M. J. Oldham 16413 18 July 1994 S. Reznicek A. A. Reznicek 10064 MICH, DAO 12 July 1995 Red Ash. Trees of Pumpkin Ash can be located by looking for “White” Ashes growing, atypically, in swamps. Pumpkin Ash is dioecious. The staminate flowers are said to have a “campanulate obscurely 4-toothed calyx; stamens two or three; pistillate calyx larger, deeply 4-lobed, accrescent and persistent” (Hough 1936). It is the fruit which provides the best charac- ter for identifying this species in the field. The sama- ras are the largest of any native ash, ranging from 4—7.5 cm in length with a wing 6—14 mm broad and narrowly decurrent to below the middle of the seed- bearing portion. The persistent calyx is similarly large, ranging from 2—5 mm in size (usually 4 mm in 1996 WALDRON, GARTSHORE, AND COLTHURST: PUMPKIN ASH IN ONTARIO 617 pesuunnconouccauncenay PELEELL LULL LTT SOUTHWESTERN ONTARIO FiGurE |. Ontario distribution of Pumpkin Ash, based on collections @, and observations O. ff }\ ! FIGURE 2. Fruiting branch of Pumpkin Ash. ea FiGurE 3. Swollen base of an Ontario Pumpkin Ash. 5cm. a a a en on | 1 2 3 4, FIGURE 4. Fruits (samaras) of three ash species: (A) Pumpkin Ash, F. profunda; (B) Red Ash, F. penn- sylvanica; (C) White Ash, F. americana. THE CANADIAN FIELD-NATURALIST Vol. 110 Ficure 5. A Pumpkin Ash tree growing in an oxbow pond along the Canard River, Essex County, Ontario. Ontario material). The relative size of the samara is shown in Figure 4. In some populations, the samaras are released over a long period from November into April; in others, they are released soon after matur- ing in October. Seed collected in Ontario germinates readily into vigorous seedlings without pretreatment, regardless of the time of collection. Seedlings and small size classes can be found in stands with Pumpkin Ash. Pumpkin Ash usually grows in low areas where water stands at least part of the year (Figure 5). Its associates in Ontario include: Silver Maple, Acer saccharinum; White Elm, Ulmus americana; Pin Oak, Quercus palustris; Swamp White Oak, Q. bicolor; Shumard Oak, Q. shumardii; Red Ash, F. pennsylvanica; Eastern Cottonwood, Populus del- toides; Black Gum, Nyssa sylvatica; willows, Salix spp. and Buttonbush, Cephalanthus occidentalis. Many of these communities would have previously (up to 1950s) been dominated by White Elm. The species is also found on mesic sites in Ontario; a fact which may be explained by the artifi- cial drainage of most of the now largely agricultural landscape. Early surveyors noted much wetter condi- tions. Patrick McNiff at Sandwich (near Windsor) in 1996 1793 writes of a line he ran “S28d, 30E. the distance of Six Concessions, at which distance the land proved so wet and swampy could not continue...”. Thomas Smith (1805-1806) describes what is presently a relatively dry and arable portion of Essex County thus: “In the three townships of Colchester, Gosfield and Mersea not above the third part of them is habitable — extensive swamps and morases [sic] perilous places — thickety and water throughout stag- nant and ruinous.” Mahlon Burwell in 1816 writes of surveying in Essex County: “in the early part of summer it was almost impossible to survey on account of the depth of water in the rear parts of the townships.” Now the areas described above are all much drier but Pumpkin Ash still thrives there. Literature Cited Braun, E. L. 1961. The woody plants of Ohio. Ohio State University Press, Columbus. Britton, N. L. 1901. Manual of the flora of the Northern States and Canada. H. Holt & Co., New York. Britton, N. L. 1908. North American Trees. H. Holt & Co., New York. Bush, B. F. 1894. Notes on a list of plants collected in southeastern Missouri in 1893. Missouri Botanical Garden Annual Report 5: 139. Bush, B. F. 1897. Notes on the botany of some southern swamps. Garden and Forest 10: 514-516. Deam, C. C. 1921. Trees of Indiana. Indianna Department of Conservation Publication 18, Fort Wayne. Duncan, W. H., and M. B. Duncan. 1988. Trees of the Southeastern U.S. University of Georgia Press, Athens. WALDRON, GARTSHORE, AND COLTHURST: PUMPKIN ASH IN ONTARIO 619 Farrer, J. L. 1995. Trees in Canada. Fitzhenry & Whiteside Ltd., Markham, Ontario. Fernald, M. L. 1938. Noteworthy plants of Southeastern Virginia. Rhodora 40: 450. Fernald, M.L. 1950. Gray’s Manual of Botany. Dioscorides Press, Portland. Hough, R. B. 1936. Handbook of the trees of the Northern States and Canada. R. B. Hough Co., Lowville, New York. Little, E. L. 1953. Check list of native and naturalized trees of the United States. U.S. Forest Service, Agricultural Handbook Number 41. Washington. Little, E. L. 1977. Atlas of United States trees. Volume 4. Minor Eastern Hardwoods. U.S. Forest Service. Miscellaneous. Publication Number 1342. Washington, IDWGs McCormac, J.S. 1993. Swamp Cottonwood (Populus heterophylla) in Ohio. Michigan Botanist 32: 35-39. MeNiff, P. 1793. Third report of the Archives of Ontario, 1905. [pages 224-225] Toronto, Ontario. Preston, R. J. 1989. North American trees, 4" edition. Iowa State University Press, Ames. Rowe, J.S. 1977. Forest regions of Canada. Canadian Forestry Service, Ottawa, Ontario. Sargent, C.S. 1922. Manual of the trees of North America. Houghton Mifflin, Boston/New York. Sargent, C.S. 1933. Manual of the trees of North America. Houghton Mifflin, Boston/New York. Smith, T. 1805-1806. Survey records Volume 133. Ontario Ministry of Natural Resources, Toronto, Ontario. Wright, J. W. 1957. New chromosome counts in Acer and Fraxinus. Morris Arboretum Bulletin 8: 33-34. Received 1 December 1995 Accepted 11 September 1996 The Occurrence of the Night Snake, Hypsiglena torquata, in British Columbia, with Comments on its Body Size and Diet Howarb LaAcey!, CHRISTOPHER H. SHEWCHUK’, PATRICK T. GREGORY, MICHAEL J. SARELL?, and LINDA A. GREGORY* 1231 Douglas Avenue, Penticton, British Columbia V2A 2T9 2Department of Biology, University of Victoria, P.O. Box 3020, Victoria, British Columbia V8W 3N5 3RR #2, Oliver, British Columbia VOH 1TO 41087 Briarwood Drive, Cobble Hill, British Columbia VOR 1L0 5Corresponding Author Lacey, Howard, Christopher H. Shewchuk, Patrick T. Gregory, Michael J. Sarell, and Linda A. Gregory. 1996. The occur- rence of the Night Snake, Hypsiglena torquata, in British Columbia, with comments on its body size and diet. Canadian Field-Naturalist 110(4): 620-625. In Canada, the Night Snake (Hypsiglena torquata) is apparently restricted to the hot, dry southern interior of British Columbia and is one of this country’s most poorly known snakes. Its small size, cryptic behaviour, restricted range in the province, and similar appearance to other syntopic species probably account for the small number of observations of this species to date. We summarize data on 14 specimens of the Night Snake found in British Columbia since 1980. Measurements of body size of these specimens are similar to those from Idaho and support the hypothesis that body size in this species is greater in the northern part of its range. Observations of feeding habits in captivity suggest that this species is a generalist predator on amphibians and reptiles. A single feeding record from the field indicates that large prey, up to at least one-third of the snake’s body mass, are sometimes taken. Key Words: Night Snake, Hypsiglena torquata, distribution, body size, food, British Columbia. On 28 September 1980, one of us (HL) found the first Canadian specimen of the Night Snake, Hypsiglena torquata, near Kaleden (49°24’ N, 119°35’ W), British Columbia, about 45 km N of the U.S. border. Since then, to our knowledge, at least thirteen other specimens have been found in British Columbia, all but two south of the first sighting. Although the occurrence of H. torquata in British Columbia, based on HL’s specimen, has been noted in four field guides (Nussbaum et al. 1983; Cook 1984; Gregory and Campbell 1984; Stebbins 1985), it has otherwise been discussed only in a magazine article (Valadka 1992), in an unpublished document (Bufo Inc. 1993. Report on the status of the Night Snake (Hypsiglena torquata) in British Columbia. Submitted to Wildlife Branch, Ministry of Environment, Lands and Parks, British Columbia), and in government reports (e.g., Orchard 1984). Because it has not yet been formally reported in the scientific literature, we herein summarize much of the available data on this species in British Columbia. All but one of the specimens of H. torquata in British Columbia have come from the southern Okanagan Valley, which is shrub-steppe habitat characterized by hot, dry summers, with near-desert conditions in some areas; the remaining specimen was found in the Similkameen Valley, west of the Okanagan, near Keremeos (49°12’ N, 119°49’ W), which has similar vegetation and climate. The most northerly specimens, both found by MJS, have come from the vicinity of Penticton (49°30’ N, 119°35’ W); the most southerly specimens have been caught about halfway between Oliver (49°11’ N, 119°33’ W) and Osoyoos (49°02’ N, 119°28' W). Data on all 14 specimens are summarized in Table 1. These data either were extracted from our own records or from those presented by Bufo Inc. (1993). In two instances, our data were different from those presented in that report: Bufo Inc. recorded SVL and TL (see Table 1) of the 28 September 1980 specimen as 362 and 431 mm and the SVL of the 8 July 1993 specimen as 390 mm. In the latter case, our measure- ment (Table 1) is supported by a photograph and is, in any case, much more consistent with the animal’s large body mass. Also, our record of the 20 July 1991 specimen as a male (Table 1) conflicts with Valadka’s (1992) report of the same specimen as a female. Most live specimens have been found either under rocks or in traps and most have been released. Three specimens have been found dead (Table 1). The specimens retained in collections (Table 1), as well as others that we have found, match the descrip- tion of H. ochrorhynchus deserticola given by Tanner (1944), subsequently treated as H. torquata deserticola (Bogert and Oliver 1945; Dixon 1965), the Desert Night Snake, which is the expected sub- species in this area (Nussbaum et al. 1983). Photographs of the other Night Snakes caught in British Columbia also support this designation (noted in Table 1). Wright and Wright (1957) show a photo- graph of a specimen from Vantage, Washington, 620 621 NIGHT SNAKE LACEY, SHEWCHUK, GREGORY, SARELL, AND GREGORY 1996 panuyuoyd [96€] Joquiny UOISSaD9V | 3104} poasasaid vay) ‘66] Sutids ur yyeop [HUN eLIO}OI A, A}ISIOATUL) 38 UONIAT[O9 dAT] url yday ‘yoRuto}s ul oyeUsTIeI sooA0sQ, (DVI pue ayeuoou yoo pure 10ATO SHO WIM) Pourejuo) B Jopul) UoEMIog XG /C8V d DLd 7661 Joquiaidag ¢ uojonuad yeo Aq uayea ‘dTAWOd Aqjenaed pue J2ATIO JW poasasaig peop punoj JO YON - aytuaant W XS 166] 1sNsNY 6 (Z661 PAPPIRA ul poyioda) posrvojer pure ‘poydeisojoyd dey Jouuny JOATO ‘painseo|| ur yysne JO YON : 89S/ILP W St 166! Aine 07 posrojor pure ‘poydeisojoyd JOATO ‘poinsvoyAl Yoo! B Jopuy) JO YON a IT C/O81 64 C9) 686l SUNT ET (UONROTUNUWIUOS euosiod ‘qH) posrojos pue ‘poydeisojoyd adoyjs (Sf ‘AyAndeo snye) & Ieou oye] UM) UL pourejuleyA YOO B Jopul) xnase A IvaN = 68P/LIT d VS 886] SUNS 9 (1X9] 998) so]poou Owe Aq poinsvouw auid Aap ‘(909] ‘ON ‘99V) UJIM paidAo0o Woda adojs snye (PAT LL SOP) Ul PdAIOSolg uo uado uy uapaye yy IeaN - C6L/STE d TH OS6T Jaquiaidas gz S}UDLUWIOD ainjdeo Jo uoneso'] (3)ssvyy (ww) FL/TAS xas JOIAT[OD aed sjieieqd ayeuntxoiddy ‘syed pue spur] ‘uowuosmAug Jo Ansturpy eriquinjod ysoug = dTAWO ‘umnesnyy viquinjod ysniug [eAoy = WOe ‘WIsue] [kI0} = TL “YSug] JuUaA-jnous = TAS “104d ‘O = GINO ‘Suosied ‘H = dH ‘Y90D ‘4 = OU ‘VorysNeNOW 'D = WO “Ula “S = MS ‘UPTM D = MD ‘WOSITTY “WW = WIN ‘TS IN = SLIN “A1082I1D “T= DVT ANYomoys ‘D = SHO ‘A10801D *q = OLd ‘1opuexopy ‘§ = Vg ‘Aoor'] “H = TH ‘eIquinjod ysniug ur punoy vypnbsoj puajsisddy Jo suswideds ude}NOJ UO UOTRUIOJUT JoyJO puke ved “| ATAV] Vol. 110 THE CANADIAN FIELD-NATURALIST 622 uojonued ‘d THNOE ye poarosaid sq[ea pue pomses]\/ TP{-peod ueSeueyO Ion = SIC/c81 daNO SO6I APN OF sooAosQ posveyar pur dey jouuny pue JaAT[O poydessoioyd ul qysneD uaeMjog 97 SIS/OEr SHO P66I Joquiaidag ¢ pasvayar pure dex jauuny oyeT] poydessol0yd ur qysney xnase A Ivan 6S OLS/S9b StI P66 Jaquiaidag | paseayar pure uojonuad : poydessoj0yg yool & Japup~ IRON - - StIN P66l oune LI pasveayar pure yoor uojonUad poydessoi0yd B Jopuy) IRON : : SI Pool eune ZI ‘[87PI Joquinyy uorssas0V | diay] padrasoid way} ‘9661 Sutds ur Weep [nun BIIOJOIA JO AJISIOATUL) sook0sQ, ye UOTOaTJOO den jauuny pue J9AT[O aaly ul day ur qysned usoMIog El OEr/09E SHO C661 IsNSNY CT uojonuad ‘d THINOd ye uatutoads quoUaINsvaUl os0fog Y TI jSva] 3B peap useq pey yey] pouoyoo.r oym “SHO Aq poarasaid pur poydeisojoyd ‘painsvay TP-peod SOSUIIOY IVAN CSE S09/SCS VN €66I Aine g Sitgel : = uvseuryC Ivan e : WO C661 ABI SZ s}UdTUUUOD) aimdeo jo uoneo0'T (3)ssejy (WW) T.L/TAS 101IaT[OD aed sjiviaq aeurrxoiddy (ponunuo)) “[ ATaV 1996 LACEY, SHEWCHUK, GREGORY, SARELL, AND GREGORY: NIGHT SNAKE 623 FiGurE 1. Specimen of Hypsiglena torquata caught on 5 September 1992 beside posterior portion of Crotalus viridis, which it regurgitated. Photo by Tom Gore on 15 January 1993. which, not surprisingly, closely resembles those seen in British Columbia. Scale counts have been made for only two of the British Columbia specimens: F. Cook (personal communication) examined HL’s original specimen and recorded loreals 1-1, preoculars 1-1, postoculars 2-2, supralabials 9-8, infralabials 11-10, dorsal scale rows 21-23-15, ventrals 194, caudals 49, anal divid- ed; H. Parsons (personal communication) described the second known specimen as having dorsal rows 20-21-17, ventrals 186, subcaudals 47 (the first 19 being single). These values are generally similar to those reported for H. t. deserticola by Tanner (1944) and Wright and Wright (1957). Given the number of specimens now accumulating in collections (Table 1), a thorough study of variation in scale patterns may eventually be possible. Presumably, the range of the Night Snake is con- tinuous from at least Penticton south through the ~ Okanagan Valley to the known part of its range in Washington state. The habitat throughout this area is characteristically dry, with extensive rock out- croppings and talus slopes, and has hot summer weather. Svihla and Cox (1940) reported an 800 km northward extension of the Night Snake’s range to Vantage, Washington, but Nussbaum et al. (1983) show only one additional locality (near Pateros, about 105 km south of the Canada-U.S.A. border) between Vantage and HL’s first specimen in Canada at Kaleden. Since then, MJS has found a specimen, recorded by McAllister (1995), at Keller Ferry (about 90 km east of Pateros and 117 km south of the international border). Thus, a consider- able gap still exists in the species’ known range in British Columbia and Washington. Undoubtedly, the snake’s secretive and mainly nocturnal habits make it difficult to find. Furthermore, its small size and potential resemblance to small specimens of four other syntopic snakes (Western Terrestrial Garter Snake, Thamnophis elegans; Gopher Snake, Pituophis melanoleucus; Racer, Coluber constric- tor; Western Rattlesnake, Crotalus viridis), all of which may be blotched (at least when young), may have caused it to be overlooked by the casual observer. Nonetheless, it is somewhat surprising that the Night Snake was never reported by experi- enced naturalists, including herpetologists, active in the southern Okanagan in the decades preceding the *Pseudacris regilla in one recent phylogeny. See Hedges 1989. 624 first record of the species. It is possible that the Night Snake has expanded its range into British Columbia only recently, but this would represent a rather rapid colonization and therefore seems unlikely. According to Nussbaum et al. (1983), H. torquata rarely exceeds 460 mm in fotal length, although these authors also report that females in southeast- ern Idaho may reach 525 mm SVL. A gravid female found in Idaho by J. Beck (personal communica- tion) was 530 mm SVL (620 mm TL) and weighed 49 g. Diller and Wallace (1986) found maximum SVL’s of males and females in southwestern Idaho to be 400 and 523 mm, respectively; they suggested that northern Hypsiglena are larger than more southerly conspecifics. Our data on body sizes of British Columbia specimens are consistent with this hypothesis. Diller and Wallace (1986) also inferred that males reached sexual maturity at about 290 mm SVL and females at about 400 mm. On this basis, we judge that all but two of the specimens found so far in British Columbia were sexually mature. The natural history of the Night Snake is poorly known, the only substantial report being that of Diller and Wallace (1986). Of particular note here is the observation that the specimen captured by PTG, CHS, and LAG (Table 1) regurgitated the partly digested posterior half of a neonate Western Rattlesnake (Figure 1). From data on numbers of ventral scutes presented for Crotalus viridis by Nussbaum et al. (1983) and Campbell and Lamar (1989) and the length of the remaining part of the rattlesnake (University of Victoria Accession Number 1396), we estimated that this prey item was 236-285 mm SVL, well within the range of sizes at birth reported by Macartney et al. (1990). We used mass-SVL regression equations from Macartney et al. (1990) to estimate that this rattlesnake weighed from about 9-14 g. Thus, the ratio of mass of prey to mass of predator was approximately 0.35-0.54. Although well short of the large relative sizes of prey eaten by some elapids and viperids (Greene 1983), this is a very large prey item, and is substan- tially larger than any reported previously for Hypsiglena, to our knowledge. Diller and Wallace (1986) found that Night Snakes in Idaho ate small lizards and their eggs, anurans, and insects. Although no stomach samples were obtained from any other specimens, we suspect that Hypsiglena in British Columbia is a generalized predator on amphibians and reptiles. Numerous cap- tive-feeding observations were obtained from the two specimens held by PTG and the single speci- mens held by HL and H. Parsons (personal commu- nication). Prey that have been eaten readily by one or more of these snakes are Long-Toed Salamanders (Ambystoma macrodactylum), Western Toads (Bufo boreas), Pacific Tree Frogs (Hyla regilla*), Wall Lizards (Podarcis muralis), and Northwestern THE CANADIAN FIELD-NATURALIST Vol. 110 Garter Snakes (Thamnophis ordinoides). The speci- mens maintained by PTG have eaten either live or dead prey of most types, including chopped-up frogs (H. regilla and Red-Legged Frogs, Rana auro- ra), but they have been especially responsive to live lizards or snakes; although one of these snakes also ate a live recently born mouse shortly after capture, we did not have further success with this kind of food. Other rejected prey include Northern Alligator Lizards (Elgaria coerulea) and nestling swallows (Parsons, personal communication), goldfish, meal- worm larvae, and a dead Ensatina Salamander (Ensatina eschscholtzii). Night Snakes are rear- fanged colubrids with toxic saliva (McKinstry 1978); although they are reported to subdue their prey with this toxic saliva (Goodman 1953; Wright and Wright 1957; McKinstry 1978; Nussbaum et al. 1983), we have not unequivocally observed this aspect of prey capture. A likely prey of Hypsiglena in southern British Columbia is the Western Skink, Eumeces skiltonianus, which is reasonably common in at least two of the sites at which Night Snakes have been found. The range of Hypsiglena torquata in British Columbia is possibly larger than described here. Habitat requirements of Night Snakes seem to over- lap with those of Western Rattlesnakes, Gopher Snakes, and Racers, all of which occur the full length of the Okanagan Valley and also in the Thompson River Valley. We encourage biologists and other naturalists to be vigilant for the Night Snake and to collect basic data (body size, sex, reproductive condition, habitat, stomach contents [obtainable by palpation]) on it so that we can better understand its natural history. We would be pleased to receive additional records of sightings or captures of this interesting snake. Acknowledgments We thank Harry Parsons (Bufo Inc., West Vancouver) and Jonathan Beck (Idaho State University, Pocatello) for their observations, Francis Cook (then of the National Museum of Natural Sciences, now Canadian Museum of Nature, Ottawa) for his confirmation of the specimen caught by HL and scale counts, and Tom Gore for photography. Bill Preston, Manitoba Museum of Man and Nature, Winnipeg, made interesting comments on an earlier draft, particularly on previous lack of records and possible feeding on skinks. Figure | is dedicated to the spirit of Harry Greene, which inhabits all those fascinated by the stomach contents of snakes. PTG thanks the Natural Sciences and Engineering Research Council of Canada for supporting his field studies of snakes. CHS and PTG also thank the World Wildlife Fund and the British Columbia Ministry of Environment, Lands and Parks for their support. 1996 Literature Cited Bogert, C. M., and J. A. Oliver. 1945. A preliminary analysis of the herpetofauna of Sonora. Bulletin of the American Museum of Natural History 83: 301-425. Campbell, J.A., and W.W. Lamar. 1989. The Venomous Reptiles of Latin America. Comstock Publishing Co., Ithaca, New York. Cook, F. R. 1984. Introduction to Canadian Amphibians and Reptiles. National Museum of Natural Sciences, Ottawa. Diller, L. V., and R. L. Wallace. 1986. Aspects of the life history and ecology of the desert night snake, Hypsiglena torquata deserticola: COLUBRIDAE, in southwestern Idaho. Southwestern Naturalist 31: 55-61. Dixon, J. R. 1965. A taxonomic reevaluation of the night snake Hypsiglena ocrorhyncha and relatives. Southwestern Naturalist 10: 125-131. Goodman, J. D. 1953. Further evidence of the venomous nature of the saliva of Hypsiglena ocrorhyncha. Herpetologica 9: 174-175. Greene, H. W. 1983. Dietary correlates of the origin and radiation of snakes. American Zoologist 23: 431-441. Gregory, P. T., and R. W. Campbell. 1984. The Reptiles of British Columbia. British Columbia Provincial Museum, Handbook 44. Victoria, British Columbia. Hedges, S. B. 1986. An electrophoretic analysis of hol- arctic hylid frog evolution. Systematic Zoology 35(1): 1-21. Macartney, J. M., P. T. Gregory, and M. B. Charland. 1990. Growth and sexual maturity of the western rat- LACEY, SHEWCHUK, GREGORY, SARELL, AND GREGORY: NIGHT SNAKE 625 tlesnake, Crotalus viridis, in British Columbia. Copeia 1990: 528-542. McAllister, K. R. 1995. Distribution of amphibians and reptiles in Washington State. Northwest Fauna 3: 81-112. McKinstry, D. M. 1978. Evidence of toxic saliva in some colubrid snakes of the United States. Toxicon 16: 523-534. Nussbaum, R. A., E. D. Brodie, Jr., and R. M. Storm. 1983. Amphibians and Reptiles of the Pacific Northwest. University Press of Idaho, Moscow. Orchard, S. A. 1984. Amphibians and reptiles of British Columbia: a synthesis of literature, life history and dis- tribution, with comments on status and research. Research Branch, British Columbia Ministry of Forests, Victoria, British Columbia. Stebbins, R. C. 1985. A Field Guide to Western Reptiles and Amphibians, Second Edition. Houghton Mifflin, Boston. Svihla, A., and C. Knox. 1940. The spotted night snake Hypsiglena in Washington. Copeia 1940: 52. Tanner, W. W. 1944. A taxonomic study of the genus Hypsiglena. Great Basin Naturalist 5: 25—92. Valadka, A. 1992. Night moves. Nature Canada 21(1): 8-9. Wright, A. H., and A. A. Wright. 1957. Handbook of Snakes of the United States and Canada, Volume 1. Comstock Publishing Associates, Ithaca, New York. Received 7 November 1995 Accepted 14 August 1996 Small Mammals of Even-aged, Red Alder - Conifer Forests in Southeastern Alaska THOMAS A. HANLEY U.S. Department of Agriculture, Forest Service, Pacific Northwest Research Station, P.O. Box 20909, Juneau, Alaska 99802-0909 Correspondence address: Pacific Northwest Research Station; 2770 Sherwood Lane, Suite 2-A; Juneau, Alaska 99801- 8545 Hanley, Thomas A. 1996. Small mammals of even-aged, Red Alder - conifer forests in southeastern Alaska. Canadian Field-Naturalist 110(4): 626-629. Densities, body weights, age and sex ratios of small mammals in three 31- to 36-year-old, even-aged, Red Alder - Sitka Spruce - Western Hemlock (Alnus rubra - Picea sitchensis - Tsuga heterophylla) stands were compared with those in adja- cent old-growth Western Hemlock - Sitka Spruce stands. Mammals were censused by snap-trapping each stand with 100 traps for three consecutive nights in late May-early June and again in late July-early August. Three species of small mam- mals were captured: Sitka Mouse (Peromyscus sitkensis), Long-tailed Vole (Microtus longicaudus), and Common Shrew (Sorex cinereus). Mice were more abundant than voles and shrews in both trapping periods. Significant differences (P<0.05) between even-aged and old-growth stands occurred only during the second period: density of shrews was greater in even-aged than old-growth stands; body weights of juvenile mice were greater in even-aged than old-growth stands (25.3+1.1 vs. 20.5+1.1 g); and voles were more commonly associated with alder than conifer patches within even-aged stands. Mice densities were significantly correlated with estimated total understory biomass across all stands in both trap- ping periods (r.=0.83 in both, P<0.05, one-tailed). Results indicate that even-aged, Red Alder - conifer stands may be valu- able small mammal habitat in southeastern Alaska and that Red Alder may be an especially important component of young stands. Clearcut logging that favors Red Alder might have significantly better consequences for some small mammal species than does high-lead clearcut logging that discourages alder. Key Words: Sitka Mouse, Peromyscus sitkensis, Peromyscus keeni, Long-tailed Vole, Microtus longicaudus, Common Shrew, Sorex cinereus, Red Alder, Alnus rubra, Sitka Spruce, Picea sitchensis, Western Hemlock, Tsuga hetero- phylla, forest management, wildlife habitat, clearcut logging, Alaska. Clearcut logging of old-growth, Sitka Spruce - Western Hemlock (Picea sitchensis - Tsuga hetero- phylla) forests in southeastern Alaska is believed to have broadly negative consequences for wildlife habitat (Schoen et al. 1981, 1988; Samson et al. 1989). The principal reason is because secondary succession following clearcut logging is believed to result in a very depauperate understory once the conifer canopy of the regenerating stand closes (closed canopy from about ages 25 through 150 years; Alaback 1982). Much research has been directed at the implications for Sitka Black-tailed Deer (Odocoileus hemionus sitkensis) (e.g., Wallmo and Schoen 1980; Hanley et al. 1989; Hanley 1993). Less attention has been given to small mammals. Van Horne (1981, 1982a, 1982b) studied the popu- lation dynamics and ecology of the Deer Mouse (Peromyscus maniculatus) and Long-tailed Vole (Microtus longicaudus) in seral stages of Sitka Spruce - Western Hemlock forests on Prince of Wales Island, southeastern Alaska. Her stands were aged 2-5, 7-9, 23-25, and 190+ years but did not include a dense, closed-canopy stage. Van Horne’s 23- to 25-year-old stand had a very high-biomass understory with conifers only beginning to close-in. It was her most productive habitat for mice; the 7- to 9-year-old stand was most productive for voles. Van Horne cautioned, however, that she “would anticipate a drop in population size near the time of [conifer] canopy closure” (Van Horne 1981: page 1058). She based her reasoning on the general pattern of sec- ondary succession described by Alaback (1982). The depauperate understory of closed-canopy, even-aged spruce-hemlock stands is a common phe- nomenon on upland sites in southeastern Alaska (Harris and Farr 1974; Alaback 1982; Deal and Farr 1994). It has become the “typical” pattern in recent decades since the widespread use of high-lead clearcut logging, where logs are transported through the air and soil disturbance is minimized. Earlier clearcut logging, however, involved considerable soil disturbance and resulted in much Red Alder (Alnus rubra) establishment and dominance of logged sites. Those sites have largely been ignored in analyses of secondary succession in southeastern Alaska. Understory of Red Alder-dominated, even-aged second-growth stands appears to be quite different from that of the general pattern for pure conifer stands. Hanley and Hoel (1996) found no significant difference in total understory biomass of 40-year- old Red Alder riparian forests (260 + 9 kg/ha) com- pared with old-growth upland (417 + 93 kg/ha) and 626 1996 old-growth riparian (295 + 3 kg/ha) forests. Similarly, R. L. Deal (unpublished data) found total understory canopy coverage of even-aged, Red Alder - conifer stands more than six times greater than that of a nearby even-aged, spruce-hemlock stand (111 + 22 vs. 18 percent). Few quantitative data are available for understory biomass of Red Alder stands in southeastern Alaska, but general observations indicate that patterns described above are widespread throughout the region. Such stands, therefore, might provide better habitat for small mammals than age, alone, would indicate. The purpose of this study was to census the small mammal populations of three even-aged, Red Alder - conifer stands in comparison with those of three nearby old-growth stands. Similar aged, pure conifer stands were not present in or near the study area. Methods The study took place in the Crab Bay area of Chichagof Island, Alaska (57°45’N, 135°15’ W), within 500 meters of the beach. Three Red Alder - conifer stands, resulting from clearcut logging of predominantly Sitka Spruce on old landslide collu- vial fans, were the focus of the study. The stands were ages 31, 36, and 36 years when the study took place. They were widely separated, two on the south shore about 2 km apart and the third across the bay from the others. Each stand was approximately 30- 40 ha. All three stands were surrounded by old- growth forest except at the beach. The old-growth stands varied from relatively dense canopy, high tim- ber volume to relatively open canopy, low to medi- um timber volume, and they occurred on steeper slopes than did the even-aged stands. Small mammals were censused by snap-trapping with museum-special traps baited with rolled oats. Trapping was conducted twice during the summer of 1994: first 27 May through 5 June, and then again 28 July through 5 August. An old-growth stand near each even-aged stand was trapped simultaneously to provide a reference for comparison. The old-growth stands were not considered paired-samples with their even-aged counterpart, however, because site char- acteristics differed. Trapping in each of the six stands was conducted within a 250 x 60-m grid. The grids were >200 meters from the stand edge and from each other between the two trapping periods. The traps were arranged in a grid of four lines, spaced at 10-m intervals on each line and 15-m inter- vals between lines. Each grid was trapped with 100 traps for three nights. Traps were baited in evening and checked in morning. All snapped traps were counted each morning, and empty snapped traps were excluded from calculations. Animal densities were expressed as number per 300 trap nights. All captured animals were identified to species and collected on-site, taken to the field station and HANLEY: SMALL MAMMALS OF RED ALDER-CONIFER FORESTS 627 weighed, sex determined, and aged (adult or juve- nile, based on pelage coloration) (shrews were not weighed, sexed, or aged). Within-stand variation in animal density was stud- ied in the even-aged stands by assigning each trap station to one of three categories of predominant overstory type (within a 10-m radius): alder, conifer, or mixed. Between-stand variation in habitat condi- tions was assessed by visually estimating total vas- cular understory biomass (kg/ha) of each stand and ranking all six stands within each of the two trapping periods. Admittedly, these were very crude esti- mates, but the ranking of stands was probably rea- sonably accurate. Data from the two trapping periods were analyzed separately. Statistical tests (all conducted at an alpha level of 0.05) were the following (Zar 1974): Student’s t-test for all comparisons between forest types (even-aged vs. old-growth) and between sex or age classes; chi-square analysis for goodness-of-fit for within-stand associations (alder, conifer, or mixed) in even-aged forest; and Spearman’s rank correlation for between-stand comparisons of animal density with total understory biomass. Results and Discussion A total of 319 animals within three species were captured: 200 mice (Peromyscus sitkensis; but note that Hogan et al. [1993] have recently suggested populations previously identified as P. sitkensis and P. maniculatus in southeastern Alaska should be des- ignated as P. keeni), 71 voles (Microtus longi- caudus), and 48 shrews (Sorex cinereus). No signifi- cant differences between even-aged and old-growth stands occurred in densities for mice or voles in either trapping period (Table 1). Densities of shrews, however, were significantly greater in even-aged than old-growth stands in the second period (Table 1). Age and sex ratios of mice and voles did not dif- fer between even-aged and old-growth stands in either period. Adult:juvenile ratios (percentage adult + SD) averaged 68 + 33 for mice and 78 + 24 for voles, both periods combined. Male:female ratios (percentage male) averaged 66+ 25 for mice and 41 +36 for voles, both periods combined. Body weights of juvenile mice were significantly greater in even-aged than old-growth stands in the second trapping period (25.3+ 1.1 vs. 20.5+ 1.1 g). No other differences occurred between forest types. Body weights of adult mice and voles were signifi- cantly greater than those of juveniles during both periods but did not differ between periods. Mouse body weights (grams) averaged 34.9 + 2.1 for adults and 23.0 + 3.5 for juveniles, both periods combined. Vole body weights averaged 48.3 + 10.2 for adults and 27.7 + 6.9 for juveniles, both periods combined. Vegetation within the even-aged stands was highly heterogeneous and clumped with distinct patches of 628 dense spruce and hemlock in areas of undisturbed soils and patches of Red Alder along old skid roads and other areas of much soil disturbance. Other areas were mixed. The heteogeneity was scattered through- out the stands. Overall, trapping stations fell into the following distributions (percentage alder:mixed: conifer): 23:47:30 for the first trapping period; and 14:33:53 for the second period. (Note: expected cap- ture frequencies differed from those distributions slightly, depending on the distribution of snapped traps and species.) Chi-square analysis indicated that voles were more commonly associated with alder than conifer patches during the second period (20:11:12 captures observed vs. 6.4:15.3:21.3 expected). No sig- nificant differences in within-stand associations occurred for other species or trapping period. Ranking of stands by estimated total vascular understory biomass differed slightly between the two trapping periods. Two of the three old-growth stands, however, were consistently ranked the first and sec- ond highest, and the third old-growth stand was con- sistently ranked the lowest (number 6 of 6). Even- aged stands were intermediate. Mice densities (Table 1) were significantly correlated with estimated total understory biomass across all stands in both trapping periods (r,=0.83 in both, P<0.05, one-tailed). Vole and shrew densities were not correlated with under- story biomass in either period (r= -0.07 and 0.54 for voles, periods | and 2, respectively; r, =0.03 and 0.07 for shrews, periods | and 2, respectively). These results indicate that even-aged, Red Alder - conifer stands may be valuable small mammal habi- tat in southeastern Alaska and that Red Alder may be an especially important component in young stands. Van Horne (1983) showed that animal density, alone, may be a misleading indicator of habitat qual- ity because it fails to distinguish between habitats that are capable of producing and maintaining high populations versus those that simply accumulate dis- THE CANADIAN FIELD-NATURALIST Vol. 110 persing subordinate individuals. My data, however, are robust in that sense, because they indicate few or no differences in age, sex, and body weights, in addi- tion to densities. Where differences did occur, they indicated more favorable habitat in the even-aged stands than in the old-growth stands. My overall conclusion of little or no difference, therefore, is conservative. The greatest weakness of my study is that it was conducted in only one year. Nevertheless, the data from the even-aged, alder-conifer stands are surprising in comparison with what might have been expected for similar-aged pure conifer stands. Red Alder stands have also been reported as productive small mammal and amphibian habitat in Oregon (McComb et al. 1993), although habitats there are quite different from those in Alaska. Understory was abundant under Red Alder in the even-aged stands; it was extremely depauperate in the pure conifer clumps. Red Alder, itself, therefore appears to be an important factor contributing to the habitat quality of such stands. It also is important to note, however, that except for voles in the second trapping period, small mammals were distributed throughout the even-aged stands regardless of over- story type. That was true even for mice, which were correlated with understory biomass across stands in both trapping periods. The highly heterogeneous dis- tribution of Red Alder and understory in relation to the home range size of individual animals was likely very important. The Red Alder understory was dominated by Devils-club (Oplopanax horridus), Salmonberry (Rubus spectabilis), Elderberry (Sambucus racemosa), and forbs more characteristic of riparian than upland sites (e.g., Hanley and Hoel 1996). What little understory that occurred in the conifer patches, on the other hand, was mostly blueberry (Vaccinium ovalifolium and V. alaskensis), Fool’s Huckleberry (Menziesia ferruginea), and forbs more characteristic TABLE |. Mean densities (number per 300 trap-nights, + standard deviation) of the Sitka Mouse, Long-tailed Vole, and Common Shrew in even-aged, Red Alder - conifer stands and nearby old-growth stands in two trapping periods. 27 May — 5 June 28 July —5 August Species/age/sex Even-aged Old-growth Even-aged Old-growth Sitka Mouse Adults Males 6.73.1 9.9+9.0 11.0+3.6 9.0+5.2 Females 4.0+4.2 3.9+4.1 4.5+4.3 10.9+7.9 Juveniles Males 3.145.4 0.7+1.2 W3+2al 6.2+7.7 Females 2.5+4.3 0+0 4.5+4.3 8.4+7.6 Total 16.3+10.0 14.5411.9 28.3+6.5 34.9+27.6 Long-tailed Vole* 6.5+4.5 N= eal 22.6+19.4 10.6+6.2 Common Shrew* 1.82221 0+0 2645.32 3.442.10 “Sample sizes for voles and shrews not sufficient for breakdown by sex and age. >Mean density of shrews differed significantly (P<0.05) between even-aged and old-growth stands in 28 July — 5 August. 1996 of upland forests (e.g., Hanley and Hoel 1996). The mixture provided a rich species diversity. My conclusions should be viewed as preliminary because data for Red Alder understories in Alaska are so few, and data for small mammals are even fewer. This study, however, does indicate the need for more research into the potential role of Red Alder in even-aged forest management in southeast- ern Alaska. Clearcut logging that favors Red Alder might have significantly better consequences for some small mammal species than does high-lead clearcut logging that discourages alder. Acknowledgments I thank I. M. Korhonen for help in the field with the trapping and data collection. I thank J.C. Barnard and E. O. Reese for help with subsequent sample preparation. Literature Cited Alaback, P. B. 1982. Dynamics of understory biomass in Sitka spruce-western hemlock forests of Southeast Alaska. Ecology 63: 1932-1948. Deal, R. L., and W. A. Farr. 1994. Composition and development of conifer regeneration in thinned and unthinned natural stands of western hemlock and Sitka spruce in southeast Alaska. Canadian Journal of Forest Research 24: 976-984. Hanley, T. A. 1993. Balancing economic development, biological conservation, and human culture: the Sitka black-tailed deer Odocoileus hemionus sitkensis as an ecological indicator. Biological Conservation 66: 61-67. Hanley, T. A., and T. Hoel. 1996. Species composition of old-growth and riparian Sitka spruce - western hemlock forests in southeastern Alaska. Canadian Journal of Forest Research 26: 1703-1708. Hanley, T. A., C. T. Robbins, and D. E. Spalinger. 1989. Forest habitats and the nutritional ecology of Sitka black-tailed deer: a research synthesis with implications for forest management. General Technical Report PNW- GTR-230. USDA Forest Service, Pacific Northwest Research Station, Portland, Oregon. 52 pages. Harris, A.S., and W. A. Farr. 1974. The forest ecosys- tem of Southeast Alaska. 7. Forest ecology and timber HANLEY: SMALL MAMMALS OF RED ALDER-CONIFER FORESTS 629 management. General Technical Report PNW-25. USDA Forest Service, Pacific Northwest Research Station, Portland, Oregon. 109 pages. Hogan, K., M. C. Hedin, H. S. Koh, S. K. Davis, and I. F. Greenbaum. 1993. Systematic and taxonomic implica- tions of karyotypic, electrophoretic, and mitochondrial- DNA variation in Peromyscus from the Pacific Northwest. Journal of Mammalogy 74: 819-831. McComb, W.C., C. L. Chambers, and M. Newton. 1993. Small mammal and amphibian communities and habitat associations in red alder stands, central Oregon Coast Range. Northwest Science 67: 181-188. Samson, F. B. [and Committee.| 1989. Conservation of rain forests in Southeast Alaska: report of a working group. Transactions of the North American Wildlife and Natural Resources Conference 54: 121-133. Schoen, J. W., O. C. Wallmo, and M. D. Kirchhoff. 1981. Wildlife-forest relationships: is a reevaluation of old growth necessary? Transactions of the North American Wildlife and Natural Resources Conference 46: 531-544. Schoen, J. W., M. D. Kirchhoff, and J. H. Hughes. 1988. Wildlife and old-growth forests in southeastern Alaska. Natural Areas Journal 8: 138-145. Van Horne, B. 1981. Demography of Peromyscus manic- ulatus populations in seral stages of coastal coniferous forest in southeast Alaska. Canadian Journal of Zoology 59: 1045-1061. Van Horne, B. 1982a. Demography of the longtail vole Microtus longicaudus in seral stages of coastal conifer- ous forest, southeast Alaska. Canadian Journal of Zoology 60: 1690-1709. Van Horne, B. 1982b. Niches of adult and juvenile deer mice (Peromyscus maniculatus) in seral stages of conif- erous forest. Ecology 63: 992-1003. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47: 893-901. Wallmo, O. C., and J. W. Schoen. 1980. Response of deer to secondary forest succession in Southeast Alaska. Forest Science 26: 448-462. Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Englewood Cliffs, New Jersey. 620 pages. Received 14 November 1995 Accepted 23 July 1996 Longevity of Wild White-tailed Deer, Odocoileus virginianus, Does in Michigan Timotuy R. VAN DEELEN!:?, HENRY CAmMpPaA III', MAYA HAMaADy?, and JONATHAN B. HAUFLER‘ ‘Department of Fisheries and Wildlife, Michigan State University, East Lansing, Michigan 48824-1222 Present address: Illinois Natural History Survey, 607 E. Peabody Drive, Champaign, Illinois 61820- 6970 3Hiawatha National Forest, 8181 U. S. Hwy 2, Rapid River, Michigan 89878 *Boise Cascade Corp. 111 Jefferson St., P. O. Box 50, Boise, Idaho 83728-0001 Van Deelen, Timothy R., Henry Campa II, Maya Hamady, and Jonathan B. Haufler. 1996. Longevity of wild White- tailed Deer, Odocoileus virginianus, does in Michigan. Canadian Field-Naturalist 110(4): 630-633. In conjunction with a telemetry study of White-tailed Deer in northern Michigan, we aged 110 wild adult deer (96 does, 14 bucks) by means of cementum annuli analysis of the incisorform canine tooth. Aging and subsequent radio-tracking sug- gested that does in this population may live unusually long (>10 years, n = 34). One doe died at 19 years, 10 months - a near record for wild deer. Key Words: White-tailed Deer, Odocoileus virginianus, aging, longevity, Michigan. Most examples of extreme longevity in White- tailed Deer are for captive or supplementally-fed individuals (Ozoga 1969). For example, Popov (1950) and Palmer (1951) reported on tame does that lived to 15 to 19.5 years. Verme (1962) reported a supplementally-fed doe that died at 12 years. Severinghaus (1949) reported that the upper limit for wild deer was 14~-16 years based on the life-spans of captive deer. He speculated that the tooth wear may begin to limit deer after 10 years. Jenkins and Bartlett (1959) reported that few wild deer live past 10.5 years although they report a semi-tame doe that lived 17 years. Hoskinson and Mech (1976) aged two does at 12.9 and 13.8 years and reported that deer of that age were rare in any population. Nelson and Mech (1990) reported on 34 wild deer with ages = 10 years including a 17 year-old buck and a 19 year-old doe. The oldest wild deer reported is a 20 year-old doe that was killed in New York (Sauer 1984). Minimum ages of wild deer have been reported from trap-tag and release studies where deer were initially aged as fawns, yearlings, or adults on the basis of tooth wear and replacement (Severinghaus 1949) and recovered later. Ozoga (1969) reported on the minimum ages for wild deer in northern Michigan based on the subsequent recovery of live- trapped, ear-tagged deer. He reported 11 does whose minimum age was > 10 years. The oldest was 14 years, 9 months. Similarly, Nixon et al. (1991) reported on the minimum ages of previously-tagged Illinois does. The average age for their sample (N=42) was 6.6 years and the maximum was 10 years, 7 months. An alternate method of aging wild deer involves analysis of the annual growth patterns in the cemen- tum of deer teeth (Low and Cowan 1963; Gilbert 1966). Age determination based on cementum annuli analysis has shown nearly universal agreement with the known ages of captive (Low and Cowan 1963; Gilbert 1966; Erikson and Seliger 1969; Lockard 1972) and wild (Thomas and Bandy 1973) Odocoileus spp.; although, as noted by Roseberry (1980), only Thomas and Bandy (1973) reported using a “blind” comparison to eliminate potential bias. Occasionally, “split”, “compound”, or “false” annulations may confound attempts at aging deer, but this problem can be overcome by trained observers with the experience to recognize non-typi- cal patterns (Gilbert 1966; Lockard 1972; Thomas and Bandy 1973; Rice 1980). Methods As part of a telemetry study of deer migration in the Upper Peninsula of Michigan (Van Deelen 1995), we live-trapped deer during the winters of 1991-1992, 1992-1993, and 1993-1994. All deer were manually restrained and initially aged as fawns, yearlings, or adults by means of tooth wear and replacement (Severinghaus 1949). Beginning mid- way through the 1991-1992 field season, we collect- ed canine teeth from the adult deer and submitted them to Matson’s Laboratory (Milltown, Montana) for age determination via cementum annuli analysis. Matson’s is a commercial laboratory that specializes in aging mammals with this technique and has had extensive experience aging White-tailed Deer (N > 10 000). Between 1975 and 1995 Matson’s has aged 62 White-tailed Deer with known ages. Fifty-five (89%) of these were aged correctly, 4 (6%) were off by 1 year, and 3 (5%) were off by >1 year (Matson and Matson 1995). We choose to use the canine teeth instead of the more commonly-used first incisors for two reasons: (1) Matson’s has an aging model for use with White- 630 1996 VAN DEELEN, CAMPA, HAMADY AND HAUFLER: LONGEVITY OF WHITE-TAILED DEER 631 TABLE 1. Estimated ages of deer from Michigan’s Upper Peninsula, 1991-1994. Unless indicated, all age estimation were assigned an “A” certainty code (Table 2). Sex 1 Dear Sn Air me Ol i Female OCR OmaeS Sop wal Sea Obs Ge Male OF as? Age class (years)* Se Oe Solas alone Geely GO SOME SI OPS 2s 1-2 i ) * Age at last birthday based on cementum annuli analysis of the lower canine tooth and an assumed birth date of | June. 'Includes 1 individual with a “B” certainty code (Table 2). “Includes 2 individuals with a “B” certainty code (Table 2). tailed Deer canines, and (2) the incisorform canine teeth are the smallest teeth in the lower jaw and are located immediately anterior to the diastema. We reasoned that removal of a canine tooth would be less traumatic for a deer than removal of a first incisor. Matson’s aging model assumes a | June birth date. Independently of their age, a sub-sample of adult deer (n = 24 does, 2 bucks) were radio-collared for a migration and mortality study. Results We trapped 386 individual deer, 200 of which were classified as fawns (86 female: 114 male), 29 as yearlings (12 female : 17 male), and 157 as adults (146 female : 11 male). We submitted 110 canine teeth to be aged, 96 from does and 14 from bucks. We have no reason to believe that the sub-sample of deer aged though cementum annuli analysis was biased relative to the trapped sample. The sub-sample of deer aged by means of cementum annuli analysis contained some remark- ably old does (Table 1), including 34 (35%) that were > 10 years. Matson’s uses a three-tiered “cer- tainty code” to classify each age estimate as A = results nearly certain, B = some error possible, or C = error likely (Table 2). These rankings represent a subjective judgement based on the histological characteristics of the tooth and the banding pattern in the cementum. Ninety-three percent (N = 102) of the teeth in the sample we submitted received the “A” rating, the remaining 7% (N = 8) received the “B” rating (Table 1). G. Matson (personal commu- nication) noted that the histological condition of the teeth in our sample was excellent and the banding patterns in the cementum were unusually distinct as is the case with deer from northern Minnesota and northern Michigan. Six of the old (>10 years, “A” certainty unless indi- cated) does were radio-collared and monitored between January 1992 and January 1995. One, trapped as an 11-year-old, was shot illegally when 12 years, 6 months. We lost radio contact with two does that were trapped as 12-year-olds. Ages at the time of disappearance were 13 years, 10 months (“B” certain- ty); and 14 years, 2 months. A doe captured as a 13- year-old was killed by Wolves (Canis lupus) the fol- lowing spring (age = 13 years, 10 months). A doe cap- tured as a 14-year-old was still alive at the end of the study (age = 16 years, 9 months). Finally, a doe that was captured as an 18-year-old died of undetermined causes at 19 years, 10 months. With the exception of Sauer (1984), we are unaware of wild White-tailed Deer living longer than this last individual and believe that it is among the oldest recorded. Discussion We believe that recent mild winters, “buck-only” hunting regulations, and relatively low predation have combined to allow does in this population to reach such unusually old ages. Severe winter weath- er is a key determinant in the survival of northern deer (Verme 1968). Northern deer seek areas with protective conifer overstories, known as deer yards (Verme 1965). Starvation occurs when summer- deposited fat reserves are depleted prior to the end of the yarding season (Mautz 1978). Ozoga (1969) reported that over-winter survival was more critical than escape from hunting in allowing longevity in northern Michigan does. TABLE 2. Certainty codes used by Matson’s Laboratory (Milltown, Montana) to express confidence in age estimates based on cementum annuli analysis of White-tailed Deer teeth (G. Matson, personal communication). Estimated Age (years) A 1-7 + 0 year(s) 8-15 a3 I 216 as 2) Certainty Code B (G =a 12, as) +3 en) +4 Verme (1968) combined measurements Of ait chill and snow hazard im a2 winter severity index (WS) thai Gescnbed the effects of wimicr weather on the T2300 'S 100 donne a 20-week winter period were judsed to be severe mm temms of deer survival. in the 20-year penod prior to this siudy (1973/1974 - 1995/1994). WSIS im the central Upper Peamsula averaged $42 (SD = 27.2). in = 10-year period prior to this siady (1983/1984 - 1993/1994). WSIs averaged 67.6 (SD = 19+ ee ra grain o7 Nainral uapublished ). During the three 3 indy = 5 12 - January 1995). there w2s 20 Siarvation-relaiced moriality amons tamho-collaed adult does (N = 36 Geer-years: Van Declen 1995). Hunts morality is a major source of morality for exploited dear populations (Dusek The number of dens used by all Porcupines differed (t = -2.98, df =26, P < 0.006) between areas in the first year. 636 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 2. Percent of Porcupine locations in dens in which den sharing occurred during winters 1991-1992 and 1992-1993 on two study sites, in Massachusetts. 1991-1992 n Area Sex % locations Central Females@> 89 132 Males‘ 17 23 Total4 78 155 East Females 32 114 Malesa¢ 43 108 Totald 37 222 1992-1993 (harsher winter) n n n Porcupines % locations _—_ Porcupines 8 66 100 11 B) 20 5) 3 13 64 105 14 8 8 36 6 7 10 30 5 15 9 66 11 aPercent den sharing differed between years for Central area females (yx? =16.59, df =1, P < 0.001), East area females (2? = 7.68, df =1, P< 0.006) and East area males (x? = 10.89, df =1, P < 0.001). bDen sharing by females differed (x? = 105.9, df = 1, P < 0.001) between areas in both years. ‘Den sharing by males differed (x? = 5.1, df =1, P = 0.024) between areas in the first year. ‘Den sharing differed (x? = 96.37, df = 1, P < 0.001) between areas for all Porcupines in both years. Some pairings of Porcupines remained consistent throughout the winter while others changed. Four individuals, all females, used one den regularly the first winter in the Central area (Table 3). Three returned the second winter (the fourth had died) and used the den regularly. The den was also used regu- larly by another female in the second winter at the same time as the original three were in it and occa- sionally by one other female and two males. On one occasion, one other den in the Central aree contained two Porcupines, one female and one unmarked. No other den sharing was documented in this area. One den was used by three different Porcupines, but not at the same time. Discussion Roze (1987) suggested that lower den density causes Porcupines to share dens, and our data sup- port this notion to some extent. Limited den avail- ability increased den sharing for individuals in the Central area, but den sharing was also common in the East area where Porcupines (at about the same density) had twice as many dens available. The talus slope was the major distinguishing feature of the East area. This provided a greater number of avail- able dens within close proximity to each other, and more opportunity for Porcupine movement between dens. This suggests that sharing also may serve a predator avoidance, social, or thermal function as yet unidentified. While the influence of predator avoid- ance on choice of resting sites by Porcupines has been noted (Sweitzer and Berger 1992; Stricklan 1995), we did not evaluate the possible relation of this to den sharing because none of the 14 deaths of radio-marked Porcupines in our study area were due to predation (Hale and Fuller 1996). In Vermont, Wolfe (1990) found a large number of porcupines denning together during severe weath- er and suggested this may be a factor affecting den sharing. In our study, the second winter was much TABLE 3. Combinations of male and female Porcupine den sharing occurrences and the months in which they occurred during winters 1991-1992 and 1992-1993 on two study sites, in west-central Massachusetts. Area Sexesa> Mating Season Non-mating season Central (9)< EF Oct Dec, Jan, Feb, Mar, Apr FFF Oct, Nov Jan, Feb, Mar, May FFFF Oct, Nov Dec, Jan, Feb, Mar FFFM Oct Jan FFFFM Jan East (16)<¢ FF Jan, Feb, Mar FFF Nov FM Feb, Mar FFM Nov FMM Dec, Jan, Feb MM Dec, Jan, Feb, Mar aDifferences in heterosexual sharing between mating and non-mating season were not significantly different at P <0.001. >The number of Porcupines sharing within one den differed (t = 7.2, df = 115, P <0.001) between areas. “Total marked Porcupines sharing dens. 1996 more severe than the first due to both colder temper- atures and thicker snow, and there was less, not more, shared denning in the East area during that winter. Severe weather may have limited Porcupine movements between dens and consequently decreased incidence of den sharing. Male-female pairing was not limited to mating sea- son (cf Dodge and Barnes 1975). Males and females shared dens as late as March, and many same-sex combinations were found. While mating may be a factor in den sharing during October-November, it does not fully explain the den sharing behavior. In the area with lower den density, a greater pro- portion of locations were shared. Den density may affect den sharing, but it probably is not the single causal factor. Contrary to expectations, we found a more severe winter correlated with less den sharing, possibly due to limited movements. Mating was not shown to be a major factor as male-female den shar- ing occurred throughout the winter and was not lim- ited to the mating season. Acknowledgments We thank E. Amati, M. Hale, E. Jaffee, and N. Nkrumah for field assistance. We received funding and support from the USDA Forest Service Northeastern Forest Experiment Station, the U.S. Department of Agriculture (McIntire-Stennis grant), the College of Food and Natural Resources and the Department of Forestry and Wildlife Management at the University of Massachusetts, Amherst, The Metropolitan District Commission, and The Massachusetts Cooperative Fish and Wildlife Unit, U.S. Fish and Wildlife Service (now National Biological Service). We thank U. Roze and two anonymous referees for helpful comments on the manuscript. Literature Cited Curtis, J. D., and E. L. Kozicky. 1944. Observations on the eastern porcupine. Journal of Mammalogy 25: 137-146. Dodge, W. E. 1967. The biology and life history of the porcupine (Erethizon dorsatum) in western GRIESEMER, FULLER AND DEGRAAF: DENNING OF PORCUPINES 637 Massachusetts. Ph.D. dissertation, University of Massachusetts, Amherst. 162 pages. Dodge, W. E., and V. G. Barnes. 1975. Movements, home range, and control of porcupines in western Washington. U.S. Fish & Wildlife Service, Wildlife Leaflet 507. Griesemer, S. J. 1995. Habitat use of porcupines in west- central Massachusetts. M.S. thesis, University of Massachusetts, Amherst. 79 pages. Hale, M.B., and T.K. Fuller. 1996. Porcupine (Erethizon dorsatum) demography in central Massachusetts. Canadian Journal of Zoology 74: 480-484. Kelly, G. M. 1973. The biology of an isolated porcupine population. M.S. thesis, University of Massachusetts, Amherst. 48 pages. Roze, U. 1987. Denning and winter range of the porcu- pine. Canadian Journal of Zoology 65: 981-986. Schmidt, K.N. 1990. Winter feeding ecology of the North American porcupine (Erethizon dorsatum) in an isolated population of eastern hemlock (T’suga canaden- sis) in northeastern Minnesota. M.S. thesis, University of Minnesota, Minneapolis. 65 pages. Shapiro, J. 1949. Ecological and life history notes on the porcupine in the Adirondacks. Journal of Mammalogy 30: 247-257. Speer, R. J., and T. G. Dilworth. 1978. Porcupine winter foods and utilization in central New Brunswick. Canadian Field-Naturalist 92: 271-274. Stricklan, D., J. T. Flinders, and R. G. Cates. 1995. Factors affecting selection of winter food and roosting resources by porcupines in Utah. Great Basin Naturalist 55: 29-36. Sweitzer, R. A. and J. Berger. 1992. Size-related effects of predation on habitat use and behavior of porcupines (Erethizon dorsatum). Ecology 73: 67-875. Tenneson, C., and L. W. Oring. 1985. Winter food pref- erences of porcupines. Journal of Wildlife Management 49: 28-33. Westveld, M., R. I. Ashman, H.I. Baldwin, R. P. Holdsworth, R. S. Johnson, J. H. Lambert, J. J. Lutz, L. Swain, and M. Standish. 1956. Natural forest vege- tation zones of New England. Journal of Forestry 54: 332-238. Wolfe, D. F.G. 1990. Unusual numbers of porcupines, (Erethizon dorsatum), observed denning together. Canadian Field-Naturalist 104: 585. Received 28 November 1995 Accepted 22 August 1996 Status of the Harlequin Duck, Histrionicus histrionicus, in the Western Northwest Territories MICHAEL A. FOURNIER! and ROBERT G. BROMLEY? ‘Canadian Wildlife Service, P.O. Box 637, Yellowknife, Northwest Territories X1A 2N5 *Department of Renewable Resources, Government of the Northwest Territories, 600, 5102 50th Avenue, Yellowknife, Northwest Territories X1A 3S8 Fournier, Michael A. and Robert G. Bromley. 1996. Status of the Harlequin Duck, Histrionicus histrionicus, in the western Northwest Territories. Canadian Field-Naturalist 110(4): 638-641. Details of historic and recent observations of the Harlequin Duck, Histrionicus histrionicus, in the western Northwest Territories including several possible breeding records are summarized. This evidence suggests a widely distributed popu- lation in the western Northwest Territories numbering in the hundreds of breeding pairs. Key Words: Harlequin Duck, Histrionicus histrionicus, Northwest Territories, distribution, breeding. The Harlequin Duck, Histrionicus histrionicus, has received attention in recent years because of declines in eastern North America. However, indications of declines in western North America also provide rea- son for concern (Breault and Savard 1991; Goudie et al. 1994). Effective conservation of this species may be constrained by a lack of information on basic biol- ogy and distribution (Breault and Savard 1991). Descriptions of the western distribution of the Harlequin Duck in modern ornithological literature invariably fail to recognize adequately the status of this species in the western Northwest Territories (NWT) (e.g., Palmer 1976; Bellrose 1980). A few references record it as a casual or accidental visitor to the region (e.g., A.O.U. 1983; Godfrey 1986). In the following account we provide details of observations of Harlequin Ducks in the western NWT from historic and recent published and unpublished sources. Our objective is to promote awareness of the western NWT breeding population of the Harlequin Duck, so that it may be adequately recognized in future discussions and actions regarding the conser- vation of this species. Historical Records The first written account of the Harlequin Duck in the NWT was that of Richardson (1851) who observed small flocks in the vicinity of Great Bear River and took specimens there. Ross (1862) was the first author to assess the status of this species in the NWT, stating that it occurred north to the Arctic coast but was rare throughout the region. Baird et al. (1884) reported several specimens which were col- lected; near Fort Resolution, at Fort Simpson, at Fort Rae, and on the Barren Lands. Russell (1898) took a female specimen near Fort Rae 27 July 1893 (Preble [1908] also referred to this specimen, stating that it was probably collected in the vicinity of the Yellowknife River). 638 Preble (1908) reported: “Adult males, ..., said to have been shot near the post, were obtained ... at Fort Resolution in 1901 and 1903. ... in 1903 I took a specimen among the rapids on the river north of Lake Hardisty on August 20, and noted another on the lower part of the same stream August 24. ... In the spring of 1904 a pair was secured from a flock of four near Fort Simpson, May 25. ... This duck is said to be often seen on Bluefish Creek, a small rapid stream emptying into the Mackenzie opposite Fort Simpson. ... the bird catalogue of the National Museum shows that skins were received also from Peel River and Fort Liard.” Anderson (1913) noted: “Mr. H.W. Jones reported the first arrival at Hay River, May 16th, 1908.” Harper (1914)* obtained a specimen: “... said to have been shot in the spring of 1912 on one of the channels of the Slave River delta.” Soper (1942, 1957) observed this species only once: “... a full-plumaged male encountered on July 8, 1932, a few miles up Buffalo River south of Great Slave Lake.” Recent Records Grunberg (1979) reported a pair at Tartan Rapids on the Yellowknife River 13 May, and a pair at Mosquito Creek near Rae-Edzo 24 May 1979. Salter (1974) observed four (sex ratio not given) at Fort Providence during fall migration between 9 September and | October 1972. Salter et al. (1974) observed eight Harlequin Ducks at Fort Simpson (one male and a pair 17 May; one female and a pair 18 May; and a pair 22 May); two at Norman Wells (a pair 23 May); and two at Wrigley (a pair 17 May) in 1973 (these may include duplicate sightings). Bromley and Trauger (1981) listed the Harlequin Duck as rare in the Yellowknife area. This status was *See Documents Cited section. 1996 based upon five observations between 1965 and 1979. Two of these observations were published by Grunberg (1979) and noted earlier. The remaining three included an observation on Great Slave Lake 25 May 1965 (sex and number of birds unknown); a pair observed on a roadside pond near Yellowknife 21 May 1965; and two males accompanying a female on the Yellowknife River at Fishing Lake in June 1977. Sirois (1989) reported two Harlequin Ducks on the lower Beaulieu River 26 May 1988, one pair plus one male on the lower Yellowknife River over the period 7-17 May 1988, and a pair plus one male on the Mackenzie River at Fort Providence 23 May 1988. Another pair was observed on the lower Yellowknife River 24 and 26 May 1990 (J. Sirois personal communication to MAF, RGB per- sonal observation). A single bird was observed on the upper Keele River July 1990 (P. Latour personal communication to RGB). A pair was observed on the Johnny Hoe River 3 July 1992, and another pair on Canyon Creek in May 1992 (or 1993?) (P. Rivard personal communication to RGB). Breeding Records Phillips (1926) reported a possible early breeding record of the Harlequin Duck for the western NWT: “... Raine (1892) says he has eggs sent him by mis- sionaries from the mouth of the Mackenzie(!) ...”. However, he does not appear to attach much credibili- ty to this record as in the preceding sentence he remarks: “There is no very good evidence that the Harlequin nests anywhere in the Mackenzie and Athabasca basins.” MacFarlane (1908) provided more details of this record, including the most comprehen- sive description of a possible nesting site of the Harlequin Duck in NWT. He states: “... Mr. Raine, of Toronto, writes that it does breed at the Mouth of the Mackenzie River, where one of his collectors found and sent him eight eggs with the skin of the parent bird. The nest was found on the 19th of June, 1894. It was on a high bank, near some ice floes, under sticks piled up by overflow water in the spring.” Phillips’ scepticism may have been warranted. Peck and Richards (1994) reported that Raine was guilty of errors in identification; the honesty and credibility of his dealings were sometimes suspect; and some professional ornithologists refused to accept any of his records. However, after studying Raines’ Saskatchewan records, Houston (1981) con- cluded that many could be accepted with reasonable confidence. In light of Houston’s review, the great detail of this record provided by MacFarlane, and recent breeding records for this species listed below, we believe it highly probable that this record is valid. Preble (1908) and Phillips (1926) provided details of a second possible early breeding record in their accounts of the Harlequin Duck in NWT. Preble stat- ed: “Reed records eggs from Peel River, taken June FOURNIER AND BROMLEY: THE HARLEQUIN DUCK IN THE NWT 639 13, 1898, ...”. However, Reed’s (1904) own account reported this location as: “Peel River, Alaska, ... Seven eggs in a hollow in river bank, lined with down.” Although not explicitly stated by either Preble (1908) or Phillips (1926), it appears that these authors perceived an error in Reed’s account and corrected it in their own later monographs. There is a Peel River in the Mackenzie River drainage, NWT, but we could find no reference to a Peel River in Alaska (Times Books 1992; NISC 1995*). This appears to be a valid breeding record for NWT. The Harlequin Duck is recognized as a regular breeder in the vicinity of Oldsquaw Lodge (63°30'N 129°00’W) in the Mackenzie Mountains. Miller et al. (in preparation*) state: “ Nesting pairs are com- monly seen near fast water near the Lodge, at Caribou Pass, and along the Tischu and Intga Rivers. Young were seen in the area as late as 27 August in 1982.” The present owners of the lodge report seeing two to three broods of three to five young each year, and male groups of up to six per flock (N. Barichello personal communication to RGB and MAP). Further verification of breeding in the Mackenzie Mountains is provided by the observation of a brood in the Mountain River area in July 1992 (P. Latour person- al communication to RGB) and a brood at Carcajou Lake 7 August 1992 (R. Popko personal communica- tion to RGB). Bromley and Trauger (1981) listed this species as breeding in the Yellowknife area. Although specific details are lacking, this assessment was based on the word of a respected and knowledgeable naturalist, W. L. MacDonald, who resided in the Yellowknife area for many years and travelled extensively in NWT from 1922 to 1970. A synopsis of his knowl- edge of the species stated: “Noted breeding in the Taltson River area from the Alberta border to Slave Lake; on the Cameron River 40 mi. NE of Yellowknife; on the Yellowknife River, formerly in the series of rapids at Con Hydro, lately in rapids just above Quyta Lake; in the series of rapids below the Snare River Hydro; on the Camsell River at White Eagle Falls; on the Bear River Ranids; and on the Carcajou River, 30 miles west of Norman Wells” (unpublished notes in the possession of RGB). Discussion These observations indicate a widespread distribu- tion of the Harlequin Duck in the western NWT. More importantly, they establish the existence of a breeding population in the region, a fact poorly recognized. The Harlequin Duck breeds most often in the vicinity of turbulent streams and rivers where water quality and insect abundance are high, and human activities are limited (Palmer 1976; Bellrose 1980; Breault and Savard 1991). Such habitat exists throughout the Mackenzie Mountains, within the Tundra Cordillera Ecozone, an area of about 152 394 km? (Wiken 640 1986). This area probably comprises the core of the distribution in western NWT. However, there is evi- dence that the fast flowing rivers along the edge of the Precambrian Shield (e.g., Cameron; Snare; Taltson; Yellowknife) may also support modest numbers. These observations do not provide us with enough data to estimate precisely the size of the breeding pop- ulation in the western NWT. However, extrapolation of even the lowest breeding densities reported for the species, to the amount of potential breeding habitat available in the region, suggests a population number- ing in the hundreds of breeding pairs. Acknowledgments We gratefully acknowledge the contributions of unpublished data by N. Barichello, P. Latour, W. L. MacDonald, S. Miller, R. Popko, P. Rivard and J. Sirois. Without these contributions this note would be significantly diminished. T. Fraser of the Environment Canada library, Edmonton, provided able assistance in obtaining literature. A. J. Erskine and an anonymous referee provided comments on the manuscript. Documents Cited (marked with * in text) Harper, F. 1914. Biological observations in the Athabasca and Great Slave Lake region. Unpublished Manuscript Notes, National Museum of Canada. Ottawa, Ontario. 281 pages. [Data later used in Ph.D. thesis, Cornell University, New York, 1925.] Miller, S. J., F. M. Brigham, N. M. Simmons, and L. Carbyn. /n preparation. An annotated checklist of the birds of the Mackenzie Mountains, N.W.T. Canada. Unpublished Manuscript. NISC (National Information Service Corporation). 1995. Arctic and Antarctic regions. Bibliographic data base. National Information Service Corporation, Baltimore, Maryland. Database licensed to Government of the Northwest Territories, Department of Renewable Resources, Yellowknife, Northwest Territories. Literature Cited American Ornithologists’ Union. 1983. Check-List of North American birds. Sixth Edition. 877 pages. Anderson, R. M. 1913. Report on natural history collec- tions of the expedition (1908-1912). Pages 456-494 in My life with the Eskimo. By V. Stefansson. MacMillan Company, New York. 538 pages. Baird, S. F., T. M. Brewer, and R. Ridgway. 1884. The Water Birds of North America. Reprint Edition (1974). Arno Press Inc., Salem, New Hampshire. 1089 pages. Bellrose, F.C. 1980. Ducks, geese and swans of North America. Third Edition. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. Breault, A. M. and J-P. L. Savard. 1991. Status report on the distribution and ecology of Harlequin Ducks in British Columbia. Technical Report Series Number 110, Canadian Wildlife Service, Pacific and Yukon Region, British Columbia. 108 pages. Bromley, R. G. and D. L. Trauger. 1981. Birds of Yellowknife. A regional checklist. Privately Printed. Yellowknife, Northwest Territories. 12 pages. THE CANADIAN FIELD-NATURALIST Vol. 110 Godfrey, W. E. 1986. The birds of Canada. Revised Edition. National Museum of Natural Sciences, Ottawa, Ontario. 595 pages. Goudie, R. I., S. Brault, B. Conant, A. V. Kondratyev, M. R. Petersen, and K. Vermeer. 1994. The status of sea ducks in the North Pacific Rim: toward their conser- vation and management. Transactions of the North American Wildlife and Natural Resources Conference 59: 27-49. Grunberg, H. 1979. The spring migration: Northwestern Canada Region. American Birds 33: 789-790. Houston, C.S. 1981. An assessment of Walter Raine and his Saskatchewan records. Blue Jay 39: 168-181. MacFarlane, R. 1908. List of birds and eggs observed and collected in the Northwest Territories of Canada between 1880 and 1894. Jn Through the Mackenzie Basin. By C. Mair. William Briggs, Toronto, Ontario. 494 pages. Palmer, R. S. Editor. 1976. Handbook of North American birds. Volume 3, Waterfowl (Part 2). Yale University Press, New Haven, Connecticut. 560 pages. Peck, G. K. and J. M. Richards. 1994. The oologists - the era of egg collecting. Pages 90-98 in Ornithology in Ontario. Edited by M. K. McNicholl and J. L. Cranmer- Byng. Special Publication Number 1, Ontario Field Ornithologists. Hawk Owl Publishing, Whitby, Ontario. 400 pages. Phillips, J. C. 1926. A natural history of the ducks. Volume IV. Reprint Edition (1986). Dover Publications Inc., New York. 409 pages. Preble, E. A. 1908. Biological investigation of the Athabaska-Mackenzie Region. North American Fauna Number 27, United States Department of Agriculture, Bureau of Biological Survey, Washington, D.C. 557 pages. Raine, W. 1892. Bird-nesting in north-west Canada. Hunter Rose, Toronto, Ontario. 197 pages. Reed, C. A. 1904. North American birds eggs. Doubleday, Page & Company, New York. 356 pages. Richardson, Sir J. 1851. Arctic searching expedition. Harper & Brothers Publishers, New York. 516 pages. Ross, B. H. 1862. On the mammals, birds, etc. of the Mackenzie River District. Canadian Naturalist and Geologist 7: 137-155. Russell, F. 1898. Explorations in the far north, being the report of an expedition under the auspices of the University of Iowa during the years 1892, *93, and ‘94. University of lowa, lowa City. 290 pages. Salter, R. 1974. Autumn migration of birds through the central and upper Mackenzie Valley region. Chapter II in Bird migrations on the North Slope and in the Mackenzie Valley regions, 1972. Edited by W. W. H. Gunn and J. A. Livingston. Arctic Gas Biological Report Series. Volume 13. LGL Ltd., Environmental Research Associates. 162 pages. Salter, R., W. J. Richardson, and C. Holdsworth. 1974. Spring migration of birds through the Mackenzie Valley, N.W.T. April-May 1973. Chapter I in Ornithological studies in the Mackenzie Valley, 1973. Edited by W. W.H. Gunn, W. J. Richardson, R. E. Schweinsburg, and T. D. Wright. Arctic Gas Biological Report Series. Volume 28. LGL Ltd., Environmental Research Associates. 168 pages. Sirois, J. 1989. Spring migration of waterfowl in the Yellowknife-Thor Lake area Northwest Territories: 1996 1988. Technical Report Series Number 58, Canadian Wildlife Service, Prairie and Northern Region, Yellowknife, Northwest Territories. 39 pages. Soper, J. D. 1942. The birds of Wood Buffalo National Park and vicinity, Northern Alberta and District of Mackenzie, N.W.T., Canada. Transactions of the Royal Canadian Institute 24: 19-97. Soper, J.D. 1957. Notes on wildfowl of Slave River Delta and vicinity, Northwest Territories. Canadian Field-Naturalist 71: 74-81. FOURNIER AND BROMLEY: THE HARLEQUIN DUCK IN THE NWT 641 Times Books. 1992. The Times atlas of the world. Ninth edition. Random House, Toronto, Ontario. 222 pages. Wiken, E. 1986. Terrestrial ecozones of Canada. Ecological Land Classification Series Number 19, Lands Directorate, Environment Canada, Ottawa, Ontario. 26 pages. Received 28 November 1995 Accepted 26 August 1996 Distance Sampling to Estimate Fledgling Brood Density of Forest Birds ERNEST W. Burorp!, DAVID E. CAPEN!, and B. K. WILLIAMS? 'School of Natural Resources, University of Vermont, Burlington, Vermont 05405 *North American Waterfowl & Wetlands Office, U.S. Fish and Wildlife Service, 4401 N. Fairfax Drive, North Arlington, Virginia 22203 Buford, Ernest W., David E. Capen, and B. K. Williams. 1996. Distance sampling to estimate fledgling brood density of forest birds. Canadian Field-Naturalist 110(4): 642-648. Research on the status of avian communities often relies on estimates of abundance, but does not always consider demo- graphic factors such as productivity. We introduce the application of a distance-sampling technique for estimating brood density of fledgling birds in forested habitats. During 1993 and 1994, we conducted 60 line-transect surveys on 10 sites in the Green Mountain National Forest in Vermont. Sites were divided into two groups, which allowed us to test for differ- ences in fledgling density between groups. We detected 508 broods representing 38 species. Using standard distance-sam- pling procedures, we estimated densities of 0.576 (C.V. = 11.89) and 0.513 (C.V. = 12.54) broods per hectare. Density was not statistically different between groups. As with other survey methods, distance sampling favors easily detected species; however, line-transect density of fledglings is less obtrusive and less labor-intensive than mist-netting or nest searches, and can contribute important information to studies of avian communities. Key Words: songbirds, productivity, brood density, distance sampling, line-transect survey, forests, Vermont. Evidence supports the hypothesis that degradation of breeding habitat has caused population declines of neotropical migrant birds by diminishing reproduc- tive success (Sherry and Holmes 1992; Robinson et al. 1995). In order to understand population declines and to implement conservation plans effectively we need to learn more about productivity and survivor- ship (Robinson 1992b; Martin 1992). Data on these demographic factors represent an improvement over such traditional measures as after-hatching-year den- sity. Although many studies have focused on density (e.g., Freemark and Merriam 1986; Askins and Philbrick 1987; Blake and Karr 1987; Derleth et al. 1989; Thompson et al. 1992; Welsh and Healy 1993), this measure reflects many demographic processes, and can be misleading (Robinson 1992a; Vickery et al. 1992). Ralph et al. (1993) emphasized the importance of field monitoring procedures that reflect avian pro- ductivity. At least two major continent-wide pro- grams have been initiated to gather these kinds of data. Monitoring Avian Productivity and Survivorship (MAPS), a mist-netting program, and the Breeding Biology Research and monitoring Database (BBIRD), an ongoing compilation of nest monitoring data, are currently being administered (Martin and Guepel 1993; Ralph et al. 1993). Productivity data are difficult to obtain and often cannot be collected over a large area without a coop- erative effort such as MAPS. Ralph et al. (1993) detailed two methods for monitoring productivity. Using mist nets to capture birds may provide excel- lent demographic data, but yields only an index to productivity. The area sampled using mist nets is constrained by the ability of the crew to move rapid- ly from net to net while processing birds; one 5-10- ha station requires 10 person-days. Also, nets usually must be strategically arranged in choice locations in order to capture a sufficient number of birds (Ralph et al. 1993). Such arrangements, which are usually non-random, often violate statistical assumptions. A second method, nest monitoring, provides a direct measure of productivity but can be impractical with limited resources. Finding nests is labor-inten- sive, and nest searching suffers even greater limita- tions in terms of area sampled and number of species surveyed than does mist-netting. Finding and moni- toring nests also has the potential for causing nest desertion or increased predation, although these impacts often can be minimized or eliminated (Ralph et al. 1993). The utility of nest searches has been demonstrated in studies focusing on restricted areas or on a small number of species (e.g., Patnode and White 1992; Camp and Best 1994). Although our knowledge of species-specific productivity and sur- vival needs to be enhanced, studies of avian habitat degradation need to focus on entire species assem- blages (Martin 1992). Current capabilities for moni- toring productivity must be augmented to meet this objective. We used line-transect surveys to estimate produc- tivity. We believed that recently fledged young were conspicuous enough to be detected in sufficient numbers that their density could be estimated with acceptable precision to provide a comparative mea- sure of productivity. This study was a novel applica- tion of distance-sampling methods, because of the between-group comparison and the composite distri- 642 1996 bution of distances resulting from data collected for many, rather than a single, species. Distance sampling is a class of techniques, which includes line-transect sampling (Burnham et al. 1980) for determining density of biological populations. Estimates of density are computed using a set of dis- tances measured from randomly placed lines to the objects of interest. Objects of interest need not be ran- domly distributed as long as the transects are arranged randomly. Transect sampling represents an improve- ment over strip transects and other classical finite pop- ulation sampling techniques that assume a complete census of objects within the sampled area (i.e., all objects are detected), and calculate density as the num- ber of detected objects divided by the area sampled. It is not necessary to know the size of the survey area if density alone is the parameter of interest. Unbiased BUFORD, CAPEN, AND WILLIAMS: FLEDGLING BROOD DENSITY OF FOREST BIRDS 643 estimates of density can be obtained even when many objects go undetected (Buckland et al. 1993). A central concept of transect sampling is the detection function, the probability that an object is detected given that it is located some specific dis- tance from the transect. The detection function is identical in shape to the probability density function of distances. Measured distances thus contain infor- mation about missed detections, and estimation of density becomes a process of estimating this proba- bility density function. In addition to sound survey design and execution, several assumptions are necessary for successful application of line-transect sampling: (1) objects on the line are detected with certainty, (2) objects are detected at their initial location, and (3) distance measurements are exact. Although violation of TABLE |. Number of fledgling broods detected per species on 30 line transects on the Green Mountain National Forest, Vermont, 1993 and 1994. Species Ovenbird, Seiurus aurocapillus Black-throated Blue Warbler, Dendroica caerulescens Black-throated Green Warbler, Dendroica virens Black-capped Chickadee, Parus atricapillus Blackburnian Warbler, Dendroica fusca Golden-crowned Kinglet, Regulus satrapa Dark-eyed Junco, Junco hyemalis American Robin, Turdus migratorius Yellow-bellied Sapsucker, Sphyrapicus varius Hermit Thrush, Catharus guttatus Blue Jay, Cyanocitta cristata Unidentified fledgling Ruffed Grouse, Bonasa umbellus White-throated Sparrow, Zonotrichia albicollis Brown Creeper, Certhia americana Canada Warbler, Wilsonia canadensis Winter Wren, Troglodytes troglodytes American Redstart, Setophaga ruticilla Yellow-rumped Warbler, Dendroica coronata Least Flycatcher, Empidonax minimus Magnolia Warbler, Dendroica magnolia Red-eyed Vireo, Vireo olivaceus White-breasted Nuthatch, Sitta carolinensis Hairy Woodpecker, Picoides villosus Red-breasted Nuthatch, Sitta canadensis Swainson's Thrush, Catharus ustulatus Wood Thrush, Hylocichla mustelina Chestnut-sided Warbler, Dendroica pensylvanica Downy Woodpecker, Picoides pubescens Rose-breasted Grosbeak, Pheucticus ludovicianus Scarlet Tanager, Piranga olivacea Solitary Vireo, Vireo solitarius Veery, Catharus fuscescens Black-and-white Warbler, Mniotilta varia Broad-winged Hawk, Buteo platypterus Cedar Waxwing, Bombycilla cedrorum Common Grackle, Quiscalus quiscula Pileated Woodpecker, Dryocopus pileatus Tree Swallow, Tachycineta bicolor # broods mR RE RR RE NNN NNNWWAHHKNANANND OA ~~~ CO CO”O 644 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE 2. Estimates of fledgling brood density (#/ha), bootstrap coefficients of variation, and model selection criteria (AIC, and chi-squared goodness-of-fit statistics) for distance data collected on 59 line-transect surveys on the Green Mountain National Forest, Vermont, 1993 and 1994. Sites Density Eivi Group I 0.576 11.89 Group II 0.513 12.54 ‘Probability of a greater chi-squared. assumptions often can be avoided with proper sur- vey design and execution, procedures also are avail- able for addressing violation of assumptions in the analysis stage (Buckland et al.1993). We report on an application of distance-sampling to estimate brood densities of fledgling birds in a forested region of Vermont. Study Area and Methods We conducted this research on the northern two- thirds of the Rochester and Middlebury Ranger Districts of the Green Mountain National Forest (GMNF). This section of the GMNEF contains the central ridge of the Green Mountains and features peaks that exceed 1200 m in elevation. Mountains drop steeply to the east where the forest boundary follows the White River. National Forest land on the western side reaches farther from the central ridge and includes smaller mountains and rolling foothills that rise from the Lake Champlain valley. East Middlebury at 130 m represents the lowest elevation. Forest covers 94% of the northern GMNF. This forest is dominated by northern hardwoods, which make up 77% of the forest area. Approximately 20% of this forested area contains mixed northern hard- woods/spruce-fir, high-elevation spruce-fir, or Paper Birch (Betula papyrifera) cover types. Balsam Fir (Abies balsamea), Eastern Hemlock (Tsuga canadensis), White Pine (Pinus strobus), Red Pine (Pinus resinosa), Red Oak (Quercus rubra), Quaking Aspen (Populus tremuloides), and Red Maple (Acer rubrum) forest types account for about 5% of the forest area. We collected data on 500-ha circular sites strati- fied according to the presence or absence of forest openings. Group I sites contained openings that accounted for approximately 10% of total site area. Group II sites were at least 99% forested. We con- sidered openings to be any 0.5-ha or greater area consisting of non-forest cover, or of forest cover < 8 years old. We surveyed four sites in 1993 and six sites in 1994. Sites encompassed elevations between 340 and 950 m and were located within the mid- slope land-type association on either side of the cen- tral ridge. We established three transects, each approximate- ly 3 km long, on each 500-ha site. Beginning at ran- domly selected starting points, transects consisted of AIC Total x2 P} 790.12 2.07 0.73 813.07 0.74 0.95 straight line segments corresponding to grid points spaced 200 m apart. Transects sometimes contained several 90-degree turns in order to stay within boundaries, or to avoid openings and other transects. Although Buckland et al. (1993) include transect designs that incorporate such turns, we tried to use them sparingly. Transects were flagged at intervals of 25 m or less. Observers surveyed transects during the first three weeks of July, which constitute nearly the full extent of the fledgling period for the birds of interest in the study area. Two observers each walked one transect per day until each observer had covered all transects. To prevent double-counting we usually allowed at least 10 days between transect replicates. Sampling began at 05:30 and usually ended before 11:00. We did not sample on days with substantial precipitation or wind. Observers searched for fledglings while walking slowly along a transect, stopping occasionally, and leaving the transect as necessary for identification and measurement. The few broods encountered dur- ing these periods away from the transect were not recorded unless they were still apparent when sur- veying resumed. All observers were experienced in identification by sight and sound of all species in the study area, but had not received specific training or experience in the identification of fledgling birds. Fledglings were detected by sight or sound. Some detections were readily identified by sound, but oth- ers required more effort. We often made identifica- tions based on the appearance or calls of attending adults. To minimize risk of flushing birds from their original locations, observers attempted to determine fledgling locations at the time of detection and before any further efforts to get a closer look. We measured perpendicular distance from the transect to the closest detected individual in a fledgling brood by pacing back to the transect from the location at initial detection. Early efforts to count individual fledglings were unsuccessful, so we counted broods. A hand-held compass was used to determine perpen- dicular direction. : We used program DISTANCE, a PC-compatible program available from Buckland et al. (1993), to perform comprehensive analyses of line-transect data. The program implements several robust models for the detection function, provides information to 1996 25 20 Frequency Orie SitanlOi wt Sain 20e 25 Distance (m) BUFORD, CAPEN, AND WILLIAMS: FLEDGLING BROOD DENSITY OF FOREST BIRDS 645 801 355 409 45" "501-7 55 FiGureE 1. Frequency distribution of perpendicular distances measured from 29 line transects to fledgling broods on Group I sites on the Green Mountain National Forest, Vermont, 1993 and 1994. (4 values > 60 m not shown.) assist selection of the best-fitting model, and com- putes density and variance estimates. Histograms of distance data were inspected for evidence of violat- ing statistical assumptions and for appropriate cut- points for grouping data. Grouping of distances was an effective method for improving estimator robust- ness when assumptions may have been violated (Buckland et al. 1993). We also truncated outliers (~5% of the distances), which were not helpful in modeling the detection function and could have led to an overparameterized model. We then used pro- gram DISTANCE to fit and select the best of four models for the detection function. Distances were partitioned into groups, with the partition points adjusted as necessary to improve model fit. Data partitioning and truncation steps used were all stan- dard distance sampling procedures described in Buckland et al. (1993). The “half-normal model with hermite polynomi- als” was selected as the best fit to the detection func- tion for both Group I and Group II sites. Model selection was based on Akaike's Information Criterion (AIC), although chi-squared goodness-of- fit statistics were also considered. Buckland et al. (1993) preferred AIC over the chi-squared test for several reasons. Following model selection we used DISTANCE to compute brood density and variance estimates with a bootstrap technique. This analysis procedure was conducted separately for Group I and Group II sites, to test for differences in fledgling brood den- sity between the two groups. Results We detected 508 broods representing 38 species (Table 1). This total greatly exceeded the recom- mended minimum sample size requirement of 60-80 detections (Buckland et al. 1993). Broods of Ovenbirds, Black-throated Blue Warblers, Black- throated Green Warblers, Black-capped Chickadees, and Blackburnian Warblers accounted for 58% of detections. Detections per species ranged from 76 Ovenbird broods to a single brood of several less- common or less-visible species. Nine broods were not identified to species and were thus categorized as “unidentified fledgling”; these instances were often solitary fledglings. Density estimates were not dif- ferent between the two groups of study sites (P > 0.10, Table 2). Our analysis included 59 transects representing a total length of 184 km (one survey of one transect was not included because of a problem with mea- sured distances). Histograms of distance data were comparable between Group I and Group II sites (Figures 1, 2). Approximately 5% of the distances were truncated by specifying a maximum distance of 646 THE CANADIAN FIELD-NATURALIST 42 m during analysis. Detections were high at zero distance, but decreased in a jagged pattern with increasing distance. A particularly large spike occurred at 13 m in the Group II data, but seven par- tition points chosen during model-fitting procedures smoothed distance data from both groups of sites into acceptable distributions. Discussion Representation of species in the distance data cor- responded reasonably well with species composition on the study sites. From point counts conducted along the same transects we knew that Ovenbirds and Black-throated Blue Warblers were the most abun- dant birds in our study area. Because these species nest on or near the ground and are not highly secre- tive, they should account for a high percentage of the fledglings detected. The five species detected most often during fledgling surveys (58% of all detections) accounted for 42% of individuals counted in 50-m- radius point counts conducted earlier in the breeding seasons (E. W. Buford, unpublished data). However, comparison of distance data and point- count data revealed a few cases where species appeared to be misrepresented. Red-eyed Vireos were the third most abundant species on our study area, but accounted for only five fledgling broods. Black-capped Chickadees, on the other hand, were more prevalent in the fledgling surveys than on point counts. We cannot exclude the possibility that poor reproductive success accounted for under-rep- resentation of Red-eyed Vireos in the fledgling data, but poor productivity seemed unlikely in these cir- cumstances. As with other field survey methods, distance sampling over-represents species that are more conspicuous and readily detected. Brown-headed Cowbirds did not appear in our fledgling data. Cowbirds do not occur in heavily forested habitats in Vermont and thus are rare in much of the GMNF (Ellison 1985). Only 4 of 2440 birds detected during point-counts on these transects were cowbirds. A greater abundance of cowbirds might complicate fledgling surveys by introducing the possibility that almost any passerine brood might contain one or more cowbird fledglings. The jagged appearance of the frequency distribu- tions of our distance data (Figures 1, 2) is typical of biological data and is comparable to some examples illustrated by Buckland et al. (1993). Heaping of distances often occurs when estimating distances rather than taking exact measurements, but this problem was not expected here because we did not estimate distances. Evasive movement also can result in abnormally high frequencies at some dis- tance from the transect. This problem may have caused the spike in our data at 13 m, and had the potential to bias the estimates. Grouping distance data by choosing several reasonably-spaced cut- points is one procedure for eliminating the adverse Vol. 110 effects of heaping or evasive movement. Grouping causes little loss of efficiency and increases estima- tor robustness when assumptions may have been violated (Buckland et al. 1993). Our application of transect sampling differs from traditional approaches by representing a suite of species rather than a single species. This feature produces a composite detection function that must meet the same statistical assumptions as a single- species detection function. Shape criteria for the detection function are becoming less stringent as distance sampling theory advances. The main requirement is that objects on the transect are detect- ed with certainty and that detection probability decreases with distance from the transect (Buckland et al. 1993). A multi-species survey properly focused on maximizing detection probability on the transect should meet this requirement. Coefficients of variation were only slightly more than 10% of the density estimates. This precision indicates that the lack of a statistical difference between Group I and Group II study sites did not result from insufficient power of the test. The boot- strap procedure for calculating standard error was preferred because it provides some indication that the jagged distance distributions have biological meaning and do not represent the consequences of a poorly designed or sloppily executed sampling scheme. Buckland et al. (1993) recommended at least 400 bootstraps for computing standard error. Several factors might limit the use of line-transect sampling for the estimation of avian productivity. Our study area represents some of the best forest songbird habitat available in the eastern deciduous forest, as indicated by the high diversity of breeding birds present (Robbins et al. 1986). Poor habitats with very low passerine densities may not produce enough detections for meaningful analysis. Similarly, studies focused on the measurement of productivity in small areas might not have sufficient space to meet sample size requirements. Based on a truncation distance of 42 m as an effective transect half-width, our surveyed area was 770 ha. Many studies are conducted in woodlots much smaller than this. As with other measures of productivity, transect surveys do not perform equally well for all species, and often will not provide enough data to support single-species analyses. In addition, estimating den- sity of fledgling broods — in contrast to nest moni- toring — is only an indirect measure of productivity and does not provide any knowledge of clutch size, hatching success, or number of fledglings per brood. Differences in productivity between years may be expressed through variation in fledgling numbers within broods. Differences in productivity between groups might reflect differences in community com- position coupled with species-specific variability in productivity. Fledgling density could be made more SS 1996 25 20 Frequency OF MUS OMMA SI 20) 25 Distance (m) BUFORD, CAPEN, AND WILLIAMS: FLEDGLING BROOD DENSITY OF FOREST BIRDS 647 30 35 40 45 50 55 FIGURE 2. Frequency distribution of perpendicular distances measured from 30 line transects to fledgling broods on Group II sites on the Green Mountain National Forest, Vermont, 1993 and 1994. (4 values > 60 m not shown.) useful by expressing it relative to adult density determined from point counts or other survey meth- ods. Some limitations of these transect surveys can be avoided by looking at relative brood densities, as we have done by comparing two groups. Despite these limitations, transect sampling is much less obtrusive than nest monitoring or mist- netting, and produces a substantial amount of pro- ductivity data with minimal equipment and few observers. Well-staffed projects could cover large areas with line-transect sampling, and longer breed- ing seasons could result in many more observations per unit area than are reported here. Although sam- pling over a long breeding season might complicate estimates of density, the results of transect surveys may still serve as useful indices to productivity. Line-transect sampling of fledglings can add important information to studies of avian communi- ties. No current method alone provides sufficient data. A productivity monitoring scheme might effectively employ line transects in conjunction with mist-netting or nest searches. Transect sampling needs further evaluation and comparison with other methods of gauging avian reproductive success. Acknowledgments We thank A. W. Ingersoll and J. D. Lloyd, who flagged transects and conducted fledgling surveys. We also appreciate others who helped with field work: D. R. Coker, K. G. Eisenmann, S. S. Germaine, I. D. Martin, H.C. McKenny, R. O'Connell, Y. K. Ortega, and K. Wilkins. J. W. Hughes and F. R. Thompson provided helpful reviews of the draft manuscript. Funding was provided by the Green Mountain National Forest, Cooperative Forest Research Funds administered through the School of Natural Resources, University of Vermont, and the Vermont Cooperative Fish and Wildlife Research Unit. Literature Cited Askins, R. A., and M. J. Philbrick. 1987. Effect of changes in regional forest abundance on the decline and recovery of a forest bird community. Wilson Bulletin 99: 7-21. Blake, J. G., and J. R. Karr. 1987. Breeding birds of iso- lated woodlots: area and habitat relationships. Ecology 68: 1724-1734. Buckland, S. T., D. R. Anderson, K. P. Burnham, and J.L. Laake. 1993. Distance sampling: estimating abundance of biological populations. Chapman and Hall, London 446 pages. Burnham, K. P., D. R. Anderson, and J. L. Laake. 1980. Estimation of density from line transect sampling of bio- logical populations. Wildlife Monographs 72. 202 pages. Camp, M., and L. B. Best. 1994. Nest density and nesting success of birds in roadsides adjacent to rowcrop fields. American Midland Naturalist 131: 347-358. 648 Derleth, E. L., D. G. McAuley, and T. J. Dwyer. 1989. Avian community response to small-scale habitat distur- bance in Maine. Canadian Journal of Zoology 67: 385-390. Ellison, W. G. 1985. Brown-headed cowbird. Pages 368-369 in The atlas of breeding birds of Vermont. Edited by S. B. Laughlin and D. P. Kibbe. University Press of New England, Hanover, New Hampshire. Freemark, K. E., and H. G. Merriam. 1986. Importance of area and habitat heterogeneity to bird assemblages in temperate forest fragments. Biological Conservation 36: 115-141. Martin, T. E. 1992. Breeding productivity considerations: what are the appropriate features for management? Pages 455-473 in Ecology and conservation of neotropi- cal migrant landbirds. Edited by J. M. Hagan, II, and D.W. Johnston. Smithsonian Institution Press, Washington, D. C. Martin, T. E., and G. R. Guepel. 1993. Nest-monitoring plots: methods for locating nests and monitoring suc- cess. Journal of Field Ornithology 64: 507-519. Patnode, K. A., and D. H. White. 1992. Effects of habitat on avian productivity in abandoned pecan orchards in southern Georgia. Journal of Field Ornithology 63: 77-85. Ralph, C. J., G. R. Geupel, P. Pyle, T. E. Martin, and D. F. DeSante. 1993. Handbook of field methods for monitoring landbirds. USDA Forest Service General Technical Report PSW-GTR-144. 41 pages. Robbins, C.S., D. Bystrak, and P. H. Geissler. 1986. The breeding bird survey: its first fifteen years, 1965-1979. U.S. Fish and Wildlife Service Resource Publication 157. 196 pages. Robinson, S. K., F. R. Thompson, HI, T. M. Donovan, D. R. Whitehead, and J. Faaborg. 1995. Regional for- THE CANADIAN FIELD-NATURALIST Vol. 110 est fragmentation and the nesting success of migratory birds. Science 267: 1987-1989. Robinson, S. K. 1992a. Population dynamics of breeding neotropical migrants in a fragmented Illinois landscape. Pages 408-418 in Ecology and conservation of neotropi- cal migrant landbirds. Edited by J. M. Hagan, IH, and D.W. Johnston. Smithsonian Institution Press, Washington, D. C. Robinson, S. K. 1992b. The breeding season: introduc- tion. Pages 405-407 in Ecology and conservation of neotropical migrant landbirds. Edited by J. M. Hagan, III, and D. W. Johnston. Smithsonian Institution Press, Washington, D. C. Sherry, T. W., and R. T. Holmes. 1992. Population fluc- tuations in a long-distance neotropical migrant: demo- graphic evidence for the importance of breeding season events in the American Redstart. Pages 431-442 in Ecology and conservation of neotropical migrant land- birds. Edited by J. M. Hagan, III, and D. W. Johnston. Smithsonian Institution Press, Washington, D. C. Thompson, F. R., UI., W. D. Dijak, T. G. Kulowiec, and D. A. Hamilton. 1992. Breeding bird populations in Missouri Ozark forests with and without clearcutting. Journal of Wildlife Management 56: 23-30. Vickery, P. D., M. L. Hunter, Jr., and J. V. Wells. 1992. Is density an indicator of breeding success? Auk 109: 706-710. Welsh, C. J. E., and W. M. Healy. 1993. Effect of even- aged timber management on bird species diversity and composition in northern hardwoods of New Hampshire. Wildlife Society Bulletin 21: 143-154. Received 12 January 1996 Accepted 30 September 1996 Parasitic Insect Abundance and Microclimate of Gravel Pads and Tundra Within the Prudhoe Bay Oil Field, Alaska, in Relation to Use by Caribou, Rangifer tarandus granti ROBERT H. POLLARD, WARREN B. BALLARD!, LYNN E. NOEL, and MATTHEW A. CRONIN LGL Alaska Research Associates, Inc., 4175 Tudor Centre Drive, Suite 202, Anchorage, Alaska 99508 ‘Present address and author to whom correspondence should be addressed: Arizona Game and Fish Department, 2221 West Greenway Road, Phoenix, Arizona 85023 Pollard, Robert H., Warren B. Ballard, Lynn E. Noel, and Matthew A. Cronin. 1996. Parasitic insect abundance and microclimate of gravel pads and tundra within the Prudhoe Bay oil field, Alaska, in relation to use by Caribou, Rangifer tarandus granti. Canadian Field-Naturalist 110(4): 649-658. During the post-calving period (late June-early August), Caribou (Rangifer tarandus granti) movements, distribution, and behavior are significantly influenced by harassment from parasitic insects such as mosquitoes (Aedes spp.) and oestrids (Warble Fly [Hypoderma tarandi| and Nose Bot Fly [Cephenomyia trompe]). A number of studies have mentioned that Caribou use oil field infrastructure features such as gravel pads and roadbeds during insect harassment but there has been no effort to quantify factors contributing to this behavior. During 1992 and 1993 parasitic insect abundance and several weather parameters were measured on active and inactive gravel drilling pads, and undisturbed adjacent tundra in the Prudhoe Bay oil field, Alaska, in an effort to explain use of these man-made structures by Caribou. Ambient air tempera- tures were lower on gravel pads than adjacent tundra at inactive pads but not at active pad sites. Wind velocities were high- er on inactive and active pads than on adjacent tundra. Mosquitos were more abundant on tundra than gravel pads at both active and inactive sites. Mosquito abundance was positively correlated with temperature and negatively correlated with wind velocity and relative humidity at both inactive and active gravel pads. During periods of high insect harassment Caribou were observed using both active and inactive gravel pads. Key Words: Caribou, Rangifer tarandus granti, insect, harassment, microclimate, Aedes spp., Hypoderma tarandi, Cephenomyia trompe, Alaska. Movements, distribution and behavior of Central Arctic Herd (CAH) Caribou (Rangifer tarandus) during post—calving and dispersal periods are influ- enced by insect harassment (White et al. 1975; Curatolo et al. 1982*!; Lawhead 1988). Caribou are harassed by mosquitoes (Aedes spp.) during late June or early July through late July and then by oestrid flies (Warble Fly [Hypoderma tarandi| and Nose Bot Fly [Cephenomyia trompe]) which emerge 2-3 weeks after mosquito emergence and remain active through August (Dau 1986). Oestrid harassment is most severe after mosquito harassment has abated in late July (Kuropat and Curatolo 1983*; Murphy and Curatolo 1987). Insect harassment can be costly to Caribou health in terms of blood loss, infestation by oestrid fly lar- vae, and movements to coastal insect-relief habitat or other habitats which may not be in optimal foraging areas (Roby 1978; Dau 1986; Downes et al. 1986). When insect harassment ceases, Caribou return inland to preferred feeding habitat. Such movements are thought to reduce overall energy expenditure by increasing feeding and nursing opportunities, thereby ‘References marked with asterisk(*) are listed in separate Documents Cited section following Acknowledgments, all others are in Literature Cited. maximizing energy retention (Cameron et al. 1989*). Insect activity on the Arctic Coastal Plain is influ- enced by weather conditions. Dau (1986) reported that insect activity on CAH summer range was corre- lated with ambient-air temperature. Insects were most active during periods of high temperature, low wind velocity, low humidity, and low cloud cover. Caribou move to the Beaufort Sea because it is cool- er, windier, and more humid than inland areas (White et al. 1975; Dau 1986). Mosquito activity increases with distance from the coast, whereas oestrid fly activity does not (Dau 1986). Others have also evaluated the effects of weather on mosquito activity in Alaska and northern Canada (Hocking et al. 1950; Gjullin et al. 1961). A variety of terrain features provide insect—relief habitat for Caribou. When oestrid flies alone harass Caribou, coastal areas apparently offer little or no relief to Caribou (Dau 1986; Lawhead and Smith 1990*). Promontories, pingos, river deltas, gravel bars, sand dunes, and mud flats are frequently used for insect relief (Roby 1978; Dau 1986; Pollard et al. 1990*). Gravel pads within Alaska’s North Slope oil fields may mimic naturally occurring elevated insect-relief sites such as pingos. Gravel pads and roads provide insect-relief habitat similar to sparsely vegetated gravel bars and are widely used during the insect season (White et al. 1975; Roby 1978; 649 650 Curatolo et al. 1982*; Fancy 1983; Kuropat and Curatolo 1983*; Murphy and Curatolo 1987; Lawhead and Smith 1990*). In spite of numerous accounts of Caribou using roads and gravel pads for insect-relief, there has been no effort to quantify reasons for apparent dif- ferences in insect abundance between gravel pads and adjacent undisturbed areas. The purpose of this study was to quantify insect abundance on gravel pads and adjacent tundra, and attempt to determine factors which influence the use of gravel pads. Our objectives were to quantify mosquito and oestrid fly abundance on gravel pads and adjacent undisturbed tundra; to determine microclimate effects of ambient temperature, wind direction and velocity, and rela- tive humidity on gravel pads and adjacent undis- turbed tundra; and to relate mosquito and oestrid fly abundance to differential microclimate conditions on gravel pads and adjacent undisturbed tundra. Study Area The study area (Figure 1) is on the northern edge of Alaska’s Arctic Coastal Plain (within 147°50'-149°10' N longitude and 70°25’—70°10' W latitude) and is characterized by low relief, many shallow lakes and drained lake basins, and a variety of vegetation types of which wet and moist tundra are dominant (Walker et al. 1980). Summers are short and cool. Summer temperatures are consistently lower along the coast than inland up to 20 km and are influenced by the prevailing east to east—northeast winds (Brown and Hogen 1975; Walker et al. 1980). Our study area encompassed the Prudhoe Bay Oil Field (PBOF) which is characterized by oil produc- tion facilities and supporting infrastructure. Oil Field facilities and roads are built on a gravel base up to 1.6 m thick which insulates the underlying per- mafrost, thus preventing thawing and subsidence. Among the facilities within the PBOF are 53 produc- ing well pads, 31 non—producing well pads, eight gathering centers, two gravel landing strips for jet aircraft, and two base camps. All facilities are con- nected by a network of primary and secondary gravel roads totaling approximately 220 km in length. In 1992, five study sites in the PBOF were select- ed from exploratory gravel well pads, hereafter called inactive pad sites, (Figure 1): Term Well C, Able State 1, Storage Pad, Lake State 1, and Delta State 2. Gravel pads varied in thickness from approximately 0.3 to 1.6m, amount of plant colo- nization (0 to > 50% cover), and proximity to the coast (from 2.4—-11.0 km). Only single capped well heads or steel posts marking the site of plugged wells were on the pads. In 1993, in addition to the five inactive pad sites, we selected five active gravel well pad sites: A Pad, D Pad, K Pad, M Pad, and Y Pad (Figure 1). Gravel pads at these sites are gener- ally much larger and thicker (1.6 m on average) than THE CANADIAN FIELD-NATURALIST Vol. 110 pads at inactive sites. Structures on active pads con- sisted of numerous well houses enclosing active well heads, pipelines, reserve pits, and large elevated pro- duction facilities (modules). All active pads were essentially devoid of vegetation and were from 5.6 to 14.1 km from the coast. Methods We measured mosquito and oestrid fly abun- dance and weather parameters at the five inactive pad sites daily from 26 June to 4 August 1992. During 1993, weather variables and insect abun- dance were measured at the same five inactive pads examined during 1992 and an additional five active pad sites from 27 June to 31 July 1993. Each site was visited once daily between the hours of 1000-1730 Alaska Daylight Time (ADT), and con- tained two sampling stations; one located on the gravel pad (20 m from the edge of the pad) and one located on the tundra (50 m north or northeast of the pad). Because the prevailing winds in summer are from the east—northeast, the sampling station align- ment was designed to minimize effects of the gravel pad on the tundra site. During 1992, ambient air temperatures were recorded every 5 min using Omnidata Datapod® (Logan, Utah) and Dryden R2® (Anchorage, Alaska) electronic data loggers located at each sampling sta- tion. Temperature probes (accuracy +0.25°C), housed in radiation shields, were located at 1.0 m above ground. In 1993, ambient air temperatures were recorded at each sampling station using a hand- held thermometer, equilibrated 15 minutes during each sampling visit. Wind velocity and direction were recorded daily at 1.0 m above ground using a Davis Turbometer® (Hayward, California) wind speed indicator (accura- cy +0.04 m/s) and a hand-held compass for each sampling station. Velocity was averaged over a | min period. Wind direction was recorded as degrees from true north to the nearest 45 degrees. Relative humidity was recorded daily at each sampling station using a sling psychrometer (Forestry Suppliers, Inc., Jackson, Mississippi). At Lake State | site, weather parameters were continuously monitored on the gravel pad and adja- cent tundra in 1992 and 1993 with an Omnidata Easylogger® (Logan, Utah) data logging system which recorded and stored data from the following instruments: (1) an RM Young® (Traverse City, Michigan) wind microvane and anemometer which recorded wind velocity (m/s) and wind direction (degrees from true north); and (2) temperature probes which recorded ambient temperatures. Instruments were placed 1.0 m above the gravel pad and tundra surfaces. All temperature and relative humidity probes were housed in plastic radiation shields which allowed for adequate ventilation yet POLLARD, BALLARD, NOEL AND CRONIN: GRAVEL PADS AND TUNDRA 651 1996 “BYSPLV ‘Paty [IO Avg soypnsg oy} UT sajis Apnys Jo uONVIOT “[ AYN ay Prd-d D ped-V @ Pd-A @ Ped-A @ Ptd-W ® Speg SAToV ZAINS MPA @ 1 VIS Ae] @ PEG AVosy @ 1 IMS I4V COUV @ 1PM WIL @ Speg salou] SALTS XGALS Wopduary auypadt |/PUO Ry ame proy — MUS |[UP JO Pld iy aNaogT XN age eee FN e S-b HNYVIN YBHYOLS ‘Ld Av@ 3OHONYd a QQ FZHAINIOW ‘Ld Ava YAQAMD 3900 isam\ ey — . » ONVIS! dW dS \awnsi SNOT ooo 89 652 protected equipment from direct solar radiation and inclement weather. Insects were sampled daily at each site using sweep nets. Sweeps were conducted at gravel sta- tions first to minimize the chances of insects follow- ing researchers to the gravel pad stations. The sweep net consisted of a 0.5 m diameter net bag attached to a 1.5 m aluminum handle. Each sample consisted of 100 sweeps made in a figure—eight motion at approx- imately one sweep per second; sweeps spanned approximately 0.5 to 2.0 m above ground. The 100 sweeps were composed of 25 sweeps made while walking away from the sampling stations on gravel or tundra in each of the four cardinal compass direc- tions. Mosquitoes and oestrid flies were identified by gross anatomical characteristics, placed in plastic bags and counted daily. Oestrid fly abundance was also continuously sam- pled from 16-30 July 1993 using four sticky traps. Each trap consisted of a dried Reindeer hide, coated with Tangle—trap® adhesive, and draped over a wooden sawhorse. At Term Well C (an inactive pad site), one sticky trap was placed near each sampling station on the gravel pad and adjacent tundra. At S Pad (an active pad site), one sticky trap was placed near the sampling station on the pad, and one sticky trap was placed underneath a large module located on the pad. These traps were checked daily and all captured oestrid flies were removed and counted. Daily temperature, wind velocity, and relative humidity measurements between tundra and gravel pad stations for combined inactive pad sites and combined active pad sites were compared using paired t-tests (Snedecor and Cochran 1980). For 1992 data, we used the average temperature over a three-hour period encompassing each sampling visit, recorded by the elctronic data loggers. For 1993, we used the point-in-time temperature measurements recorded during each sampling visit. Data collected during 1992 and 1993 were ana- lyzed seperately and, where appropriate, pooled. We conducted a single paired t-test (for each weather parameter measured) that included data from all inactive pad sites to compensate for variation among inactive pad sites. We also conducted a single paired t-test that included data from all active pad sites to compensate for variation among active pad sites. Due to the distinct physical differences between inactive pad sites and active pad sites, these data were not combined. Spearman rank correlations were used to determine correlations among weather parameters, mosquito abundance, and distance to coastline. Wilcoxon paired—sample tests were used to determine differences in mosquito and oestrid fly abundance at gravel pad versus tundra stations. Two-sample chi-square tests (Zar 1984: 145) were used to evaluate differences in median wind direc- tion between gravel pad and tundra stations for each site. Raw insect and weather data are contained in THE CANADIAN FIELD-NATURALIST Vol. 110 Pollard and Ballard (1983) and Pollard and Noel (1984). All statistical analyses were conducted using the software packages SYSTAT® (Evanston, Illinois) and STATISTIX® (St. Paul, Minnesota). Results Weather Variables — Gravel Pads vs. Tundra Inactive Pad Sites Ambient air temperatures were higher (P < 0.001) on tundra than on gravel pads in 1992 (x = 6.6°C, SE = 0.3 and x = 6.4°C, SE = 0.32, respectively) and 1993 @ S45. SE = Ol4-and.« = N° Cx SE 0.3, respectively). Similar results were obtained (P < 0.001) when data from both years were pooled. Daily mean temperatures recorded by the data logger at the Lake State 1 study site in 1993 were also high- er (P = 0.008) on tundra (x = 9.3°C, SE = 0.6) than on the gravel pad (x = 9.2°C, SE = 0.5). The range of daily temperature differences between gravel pads and tundra at any one site was 0.0-1.6°C in 1992 and 0.0—6.8°C in 1993. Overall and at individual sites, relative humidity was higher on gravel pads (x = 78.3%, SE = 0.9) than on tundra (x = 77.7%, SE = 0.9) in 1992, but this difference was not significant (P = 0.059). In 1993, daily relative humidity mea- surements were not different (P = 0.54) between gravel pads (x = 82.4%, SE = 1.0) and tundra (x = 82.1%, SE = 1.0). Wind velocities were higher (P < 0.001) on gravel pads than on tundra in 1992 (x = 3.7 m/s, SE = 0.13) and: x ="3:2 m/s.SE = 0!) respectively.) and 1993 (x = 3.6 m/s, SE=0.1 and x = 3.2 m/s, SE = 0.1, respectively). This was also true (P < 0.001) for pooled data and for wind velocity data recorded by weather stations on gravel and tundra at Lake State lin: 1992" Ge=7378 misySE = 0!3"andae =3'2 m/s, SE = 0.3, respectively) and 1993 (x = 3.4 m/s, SE = 0.2 and x = 2.9 m/s, SE = 0.2, respectively). The range of wind velocity differences between gravel pads and tundra at any one site ranged from 0.0 to 2.8 m/s in 1992 and from 0.0 to 8.6 m/s in 1993. There was no significant difference (P > 0.05) in median wind direction between gravel pads and tundra for any of the five sites in both years of study. Temperature and relative humidity were negative- ly correlated on gravel pads (r = -0.6, P < 0.05) and tundra (r = -0.6, P < 0.05) but wind velocity and rel- ative humidity were not. Temperature and wind velocity were negatively correlated (r = -0.2, P < 0.05) on gravel pads, but not on tundra in 1992 and for neither of the sites in 1993. Active Pad Sites — 1993 Temperatures were similar (P = 0.920) on tundra (x = 11.8°C, SE = 0.4) and gravel pads (x = 11.8°C, SE = 0.4). Temperature measurements ranged from 2.3 to 22.9°C, and differences between temperatures 1996 POLLARD, BALLARD, NOEL AND CRONIN: GRAVEL PADS AND TUNDRA 653 TABLE 1. Comparison of sweep net counts of mosquitoes between tundra and gravel sites at five inactive pad sites in the Prudhoe Bay Oil Field, during 1992 and 1993. Number (%) of mosquitoes 1992 Site Tundra Gravel Able State 148(13) 29(5) Storage Pad 70(6) 54(9) Term Well C 5 PPTKOD) 171(29) Delta State 333(30) 79(13) Lake State 1 329(30) 253(43) Total 1107 586 at gravel and tundra stations ranged from 0.0 to 4.1°C. Wind velocities were significantly higher (P = 0.005) on gravel pads (x = 3.1 m/s, SE = 0.1) than on tundra (x = 2.9 m/s, SE = 0.1). Velocity measure- ments ranged from 0.0 to 9.3 m/s, and differences between wind velocity on gravel pads and tundra ranged from 0.0 to 6.3 m/s. There were no signifi- cant differences (P > 0.05) in median wind direction between gravel pads and tundra for any of the five active pad sites. Median wind direction, all sites combined, was northeast (45°) for active pad sites. Daily relative humidity measurements were similar (P = 0.478) on gravel pads (x = 82.2%, SE = 1.0) and tundra (x = 82.5%, SE = 0.1). Relative humidity ranged from 36 to 100%, and differences in humidity between gravel pads and tundra ranged from 0 to 2%. Temperature and relative humidity were negative- ly correlated on tundra (r = -0.8, P < 0.05) and grav- el pads (r = -0.8, P < 0.05) but there were no signifi- cant correlations between temperature and wind velocity or relative humidity and wind velocity. Insect Abundance and Relationships with Weather Inactive Pad Sites Mosquito abundance was low (i.e., 0-4 mosquitoes per day [mpd]) from 26 June through 7 July 1992. On 8 July mosquito abundance peaked (840 mpd). A second peak (268 mpd) occurred on 19 July. Other than these two peaks, mosquitoes were moderately active on 9-11 July (57-169 mpd), 18 July (79 mpd), and 20—23 July (24-100 mpd). After 1993 Tundra Gravel Total 385(8) 39(2) 601(7) 709(15) 461(20) 1294(15) 821(17) 138(6) 1348(15) 1411(29) 526(22) 2349(26) 1557(32) 1190(51) 3329(37) 4874 2354 8921 23 July, mosquito levels were relatively low at all study sites. Mosquitoes were more abundant (P < 0.001) on tundra (x = 31.4, SE = 14.3) than on gravel pads (x = 16.9, SE = 9.7). Sweep net counts of mosquitoes were higher on tundra than on gravel 21 of 25 days (84%) on which mosquitoes were caught. High winds on 4-6 July precluded sweep net sam- pling of insects. In 1993, mosquito abundance was moderate—to— high on 28—29 June (384 and 451 mpd, respectively), 3—4 July (78 and 260 mpd, respectively), 7-9 July (197 and 436 mpd, respectively), 11-12 July (476 and 467 mpd, repectively), and 14 July (194 mpd), with abundance peaking on 5 July (3048 mpd) and 10 July (651 mpd). After 15 July, mosquito levels were relatively low at all study sites. Sweep net counts of mosquitoes were higher on tundra than gravel pads 22 of 29 days (76%) when mosquitoes were caught. Mosquitoes were more abundant (P = 0.002) on tundra (x = 139.3 mpd, SE = 60.6) than on gravel pads (x = 67.3 mpd, SE = 29.5). There was considerable variation among sites rela- tive to total numbers of mosquitoes captured (Table 1). The highest number of mosquitoes occurred at the Lake State | site (37%) followed by Delta State 2 (26%) and Storage Pad and Term Well C (15% each). Able State 1 had the lowest number of mosquitoes overall (7%). Mosquito activity was positively correiated with air temperature (tundra - r = 0.6, P < 0.05; pads - r= 0.5, P < 0.05) and negatively correlated with relative humidity (tundra - r = -0.3, P < 0.05; pads - r = -0.3, TABLE 2. Minimum and maximum weather variable values at which mosquitoes were caught at inactive pad sites in the Prudhoe Bay Oil Field, Alaska, during 1992 and 1993. 1992 Weather Variable Min Max Temperature (°C)* BES) 25.4 Wind Velocity (m/s) 0 53 Relative Humidity (%) 29 97 1993 n Min Max n 89 4.6 21.9 159 100 0.2 6.8 159 99 45 100 152 *averaged over a 3-hour period encompassing sampling visits in 1992. 654 P < 0.05) and wind velocity (tundra - r = -0.2, P < 0.05; pads r = -0.3, P < 0.05). Similar results were obtained for individual years. Periods of maxi- mum mosquito activity coincided with periods of high ambient temperature, low wind velocity and low relative humidity. The minimum temperature at which mosquitoes were caught at any one site in 1992 was 3.5°C and the maximum was 25.4°C (Table 2). Mosquitoes were active with totally calm conditions (0.0 m/s) up to a maximum wind velocity of 5.3 m/s. Minimum relative humidity at which mosquitoes were caught was 29% and the maximum was 97%. The minimum temperature at which mosquitoes were caught at any inactive pad site in 1993 was 4.6°C, and the maxi- mum was 21.9°C. Mosquitoes were active at wind velocities of 0.2—6.8 m/s, and relative humidities of 45—100% (Table 2). During1992, no oestrid flies were caught during sweep net sampling. In 1993, no oestrid flies were caught on inactive pad sites at gravel pad stations during sweep net sampling, and only three Warble Flies were caught in sweep nets on tundra stations adjacent to the pads. In contrast, sticky traps at the Term Well C site were more effective in capturing oestrids. Warble Flies were captured on 13 of 15 days (87%), and Bot Flies were captured on 8 of 15 days (53%). Peaks in catches occurred on 16 July (38 oestids), 19 July (52 oestrids), and 21 July (30 oestrids). On all other days, between 0-19 oestrid per day were captured. More (P = 0.002) oestrids were captured on the tundra sticky trap (total = 152, x = 10.1, SE = 2.9) than on the gravel pad sticky trap (n = 54, x =3.6, SE = 1.4). Active Pads Sites-1993 Mosquito abundance was low on 27 June, 1-2 July, 6 July, 13 July, and 15-31 July (0-49 mpd). Moderate-to—high levels (x =182 mpd) occurred 28-30 June, 3-4 July, 7-10 July, 12 July, and 14 July, with peak abundance on 5 July (852 mpd) and 11 July (652 mpd). After 15 July, mosquito levels were relatively low at all study sites (0-25 mpd). Sweep net counts of mosquitoes were higher (P < 0.001) on tundra (x = 96.2 mpd, SE = 29.8) than on gravel pads (x = 9.7 mpd, SE = 3.1) on all days (n = 27) when mosquitoes were caught. Mosquito activity was positively correlated with air temperature (tundra - r = 0.6, P < 0.05; pads - r = 0.4, P < 0.05) and negatively correlated with wind velocity (tundra - r = -0.2, P < 0.05; pads - r = -0.3, P < 0.05) and relative humidity (tundra - r = -0.5, P < 0.05; pads - r = -0.4, P < 0.05). The minimum temperature at which mosquitoes were caught at any active pad site was 5.4°C, and the maximum was 22.9°C. Mosquitoes were active at wind velocities of 0.0—6.7 m/s, and relative humidi- ties of 47—-100%. THE CANADIAN FIELD-NATURALIST Vol. 110 Only two Warble Flies were collected in sweep nets, one on the gravel pad station at A Pad and one on the tundra station at M Pad on 21 July and 11 July, respectively. Oestrid flies were collected from 16-30 July on sticky traps set at the gravel pad sta- tion and under the module at S Pad. No Warble or Bot Flies were caught on the sticky trap set in the shade of the module. Warble Flies were caught on the gravel pad 5 of 15 days (33%) and Bot Flies were caught 3 of 15 days (20%). Peaks in oestrid catches occurred on 19 July (11 oestrids) and 20 July (5 oestrids). Weather/Insect Relationships Relative to Distance from Coast Mosquito abundance and weather variables on five tundra stations in 1992 and ten tundra stations in 1993 were evaluated in terms of proximity to the Beaufort Sea coast. Tundra stations at inactive pad sites ranged from 2.4—11.7 km from the coast while tundra stations at active pad sites ranged from 5.6-14.1 km from the coast. In 1992, wind velocity and relative humidity were negatively correlated (r = -0.9, P < 0.04) with distance from the coast but there was no correlation between mosquito abundance and distance from coast, or temperature and distance from the coast. In 1993, relative humidity was nega- tively correlated (r = -0.7, P = 0.03) and temperature was positively correlated (r = 0.79, P = 0.006) with distance from the coast. There was no significant correlation between wind velocity and distance from coast or mosquito abundance and distance from coast. Discussion Insect Abundance and Climatic Conditions Several physical differences between inactive and active pads may have effected our results. These included: active gravel pads were higher than inac- tive pads due to a thicker gravel layer; and oil pro- duction structures, including well houses and pro- duction modules, are common on active pads but absent on inactive pads. Structures on active pads provide shade and block winds, both of which proba- bly influence microclimate. For inactive gravel pad sites, ambient air tempera- ture was higher and wind velocity was lower on tun- dra than on gravel pads. Differences in temperature may be rel ted to the reflectance characteristics of the two meaiums. Gravel pads are well-drained and are composed of a mixture of various sizes of gravel and sand. These materials exhibit a characteristically higher reflectance of incoming solar radiation and a lower emissivity than green vegetation at tundra sites (Lillesand and Kiefer 1979). Emissivity is a measure of an object’s ability to radiate energy and is a func- tion of temperature. For active pad sites, there was no difference in ambient air temperatures between 1996 gravel and tundra, possibly due to the influence of structures (well houses, pipelines, and modules) on wind velocities at the pads, even though wind veloci- ties were higher on gravel pads than on tundra. Higher wind velocities on gravel pads were likely due to a “gradient” effect. Wind velocity increases with elevation (Snow 1976) and gravel pads are higher than adjacent tundra. During both years, there were no differences in wind direction or relative humidity between gravel pads and tundra stations at inactive pad sites. This was also true for active pad sites. Relative humidity was negatively correlated with temperature, but not with wind velocity. Identical findings were reported by Dau (1986) and Nixon (1990). In 1992, temperature was negatively corre- lated with wind velocity on gravel pads but not on tundra. However, for both gravel pads and tundra, no correlation was found between these two variables in 1993 or when data were pooled. Dau (1986) and Nixon (1990) also did not find correlations between temperature and wind velocity. During 1993, temperature increased with distance from the Beaufort Sea coast, while relative humidity decreased with distance from the coast for tundra stations at inactive and active pad sites combined. Dau (1986) reported correlations between tempera- ture and humidity with distance from the coast. During 1992 wind velocity and relative humidity decreased with distance from the coast, while tem- perature was not correlated. Dau (1986) and Searby and Hunter (1971) also reported a negative correla- tion between wind velocity and distance to coast. In contrast, there was no correlation between wind velocity and distance during 1993. Dau (1986) reported that mosquito activity increased with distance from the Beaufort Sea coast (for five sites in roughly north-south alignment). We found no significant relationship between mean number of mosquitoes in sweep nets and distance from the Beaufort Sea coast for either 1992 or 1993. Other site attributes, such as proximity to waterbod- ies or distance to the coast measured with regard to prevailing wind direction, may be more relevant. Insect activity in arctic areas is a function of at least two weather variables, i.e., temperature and wind (Curatolo 1975*; White et al. 1975; Roby 1978; Dau 1986). Mosquito abundance was correlat- ed with temperature, wind velocity, and relative humidity for both inactive and active pad sites. Temperature and mosquito abundance had the high- est correlation coefficient for both 1992 and 1993, similar to Dau’s (1986) findings. Nixon (1990) reported that the average mosquito catch was more strongly correlated with wind velocity than tempera- ture. But Nixon (1990) also found a highly significant quadratic regression (r? = 0.92) for the proportion of samples containing mosquitoes with the temperature. POLLARD, BALLARD, NOEL AND CRONIN: GRAVEL PADS AND TUNDRA 655 Peaks in mosquito abundance appear to correspond with periods of high ambient air temperature, low wind velocity, and low relative humidity in both 1992 and 1993. The minimum air temperature (4.6°C) at which mosquitoes were caught in 1993 was similar to the 4.4°C minimum reported by Gyullin et al. (1961) but higher than the minimum recorded in 1992 (3.5°C). Other studies have suggested that mosquitoes are more active at temperatures above 6-7°C (White et al. 1975; Roby 1978; Nixon 1990). The maximum air temperature (21.9°C) at which mosquitoes were caught in 1993 was slightly lower than that recorded in 1992 (25.4°C) and both maximums were lower than the 26°C upper threshold identified by Dau (1986). The maximum wind speed at which mosquitoes were caught was slightly higher in 1993 (6.8 m/s) than in 1992 (5.3 m/s) and higher than the 4.5 m/s maximum reported by Gjullin et al. (1961). However, it is similar to the 6 to 7 m/s range report- ed by Roby (1987), Dau (1986), and Nixon (1990). Mosquito abundance was higher on tundra than on inactive gravel pad stations during both years of study. Active pad sites also had higher mosquito abundance at tundra stations in 1993. Reduced mosquito abundance on gravel pads was probably due to higher wind velocities, lower ambient temper- atures, and poor insect cover habitat on gravel pads. Although plant colonization has occurred on some inactive pads, there were no sizable (i.e., > 5 m7) contiguous stands of vegetation on the gravel pads at any of our study sites except Lake State 1, which was part of an earlier restoration project (Jorgenson 1988*). Interestingly, proportionately more mosquitoes were captured at this site than at any of the other four sites. Vegetation provides resting habi- tat and escape cover for mosquitos during periods of inclement weather (White et al. 1975; Linkswiler and Curatolo 1984*; Dau 1986). Dau (1986) reported annual variation in the pro- portion of days mosquitoes were active. For exam- ple, in 1982 he caught mosquitoes on 64% of sam- pling days and in 1983 on 85% of sampling days. In our study, mosquitoes in any number were captured on 68% of sampling days in 1992 and on 86% of sampling days in 1993. MacLean (1975) suggested that annual variability in mosquito abundance may be related to difference in air temperatures as much as actual differences in abundance. Indeed, daily mean temperatures, on both gravel pads and tundra and over the entire sampling period, were 4.8°C higher, on average, in 1993 than in 1992. Active pad sites generally had fewer mosquitoes than inactive pad sites. Site specific differences in amount of available larval habitat; i.e., temporary marsh pools (dry four to five weeks after spring break up) and semipermanent pools (dry for at least a short period during July or August) (Gjullin et al. 656 1961), may account for some of the differences. Perhaps road dust created by vehicular traffic at active pad sites may also influence mosquito abun- dance by reducing adult resting and cover habitat quality (vegetation dusting) or by coating pool sur- faces and inhibiting larval respiration. Several reasons exist for our low success rate for capturing oestrid flies using sweep nets. In arctic environments oestrids have a clumped distribution, _ occur in low densities, and are much less ubiquitous than mosquitoes (Dau 1986; Curatolo 1975*). Dau (1986) also did not catch oestrids using sweep nets and caught only low numbers in sticky traps relative to mosquito catches. Nixon (1990) was unsuccessful in capturing oestrids using Malaise traps even when using CO, as an attractant. In our study, as with Dau’s (1986), sticky traps were more successful in capturing oestrids. Oestrids were significantly more numerous on the tundra sta- tion sticky trap than on the adjacent gravel pad. No oestrids were collected on the sticky trap placed in the shade of the module at S Pad, but oestrids were collected on the sticky trap placed on the gravel pad nearby. There is some evidence that oestrids are shade-intolerant. Espmark (1968), in fact, noted that Reindeer may stand under the shade of a thick—branched tree to avoid oestrid flies. Dau (1986) suggested oestrid fly distribution was influ- enced by Caribou distribution. Oestrids locate hosts more efficiently than mosquitoes because they use olfaction and warm convection currents associated with warm-blooded animals to locate Caribou (Roby 1978). Our S Pad sticky trap data appear to support Dau’s conclusions. On four of the five days oestrids were caught on the sticky trap at S Pad, Caribou were sighted on or near S Pad during road surveys (Pollard and Noel 1994, unpublished data). On one occasion, a group of Caribou that was observed run- ning from a tundra area up onto the pad, aggregated under the module where our sticky trap was located. Interestingly, the cloud of oestrid flies that had apparently elicited this behavior, stopped short of the module and would not fly under this structure. The highest number of oestrids were caught on the day (19 July 1993) when the most Caribou were observed on the pad. During insect—free periods, movements of Caribou are related to gaining access to preferred forage (White et al. 1975). Caribou disperse inland during oestrid harassment (Lawhead and Curatolo 1984*: Lawhead 1988; Dau 1986) and may expend large amounts of energy avoiding insects (Dau 1986). If insect harassment is particularly severe, energy deficits, resulting from increased running and less time feeding, may affect the survival of animals dur- ing winter and calves during summer (Curatolo 1975*). Availability of man—made insect-telief habi- tats (i.e., gravel pads and roads) may allow Caribou THE CANADIAN FIELD-NATURALIST Vol. 110 to remain in preferred foraging habitat, thereby less- ening the energy demands imposed on Caribou dur- ing the insect season. Dau (1986) indicated that natural insect—relief habitat is limited to within 3 km of the Beaufort Sea coast, and that the area within | km is probably of highest value to Caribou. Apparently, coastal areas do not afford much relief from oestrid harassment (Lawhead and Smith 1990*) and naturally occurring oestrid—relief terrain (e.g., gravel bars, pingos) is limited on the Arctic Coastal Plain (Dau 1986). Furthermore, the quantity and quality of forage may be substantially lower along the coast than inland (White et al. 1975; Dau 1986). Use of oil field facilities by Caribou to ameliorate oestrid harassment is well documented (White et al. 1975; Curatolo et al. 1982*; Fancy 1983; Curatolo and Murphy 1986; Dau 1986; Johnson and Lawhead 1989*; Lawhead and Smith 1990*). During insect harassment Caribou are less sensitive to oil field facilities and activities than at other times (White et al. 1975; Curatolo et al. 1982*; Dau 1986; Murphy and Curatolo 1987). Indeed, the dominating influ- ence of oestrids often overrides any tendency to avoid development related structures and/or activi- ties (Pollard et al. 1992*). Reduced mosquito and oestrid abundance on gravel pads, and absence of oestrids in the shade of a module, indicate that gravel pads and structures on pads offer insect-relief habitat because these sites are windier, drier, and have less vegetative cover. Acknowledgments This study was funded by BP Exploration (Alaska) Inc. (BPX). We would like to thank Chris Herlugson (Supervisor of Environmental Assess- ment for BPX’s Environmental and Regulatory Affairs Department) for his support of this project. We also thank Michelle Gilders (Environmental Scientist, BPX) for her interest and support during this study and for her review of this manuscript. Special thanks go to Benny Galloway who con- ceived the study and to Bob Rodrigues for his invaluable assistance in the field. We also thank Mark Miller, Jeff Wickliffe, Barbara Pierson, Pat Murphy, Scott Mignery, and Virginia Hanebuth for their assistance with data collection. James Dryden and John Kew provided technical assistance with respect to weather instrumentation. Documents Cited [marked * in text citations] Cameron, R. D., W. T. Smith, and S. G. Fancy. 1989. Distribution and productivity of the Central Arctic Caribou herd in relationship to petroleum development. Alaska Department of Fish and Game. Federal Aid in Wildlife Restoration Progress Report, Project W-23-1 and W—23-2, Study 3.35. Juneau, Alaska. 52 pages. 1996 Curatolo, J. A., S. M. Murphy, and M. A. Robus. 1982. Caribou responses to the pipeline/road complex in the Kuparuk oil field, Alaska, 1981. Final report to ARCO Alaska, Inc., Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 62 pages. Johnson, C. B., and B. E. Lawhead. 1989. Distribution, movements, and behavior of Caribou in the Kuparuk oil field, summer 1988. Final report to ARCO Alaska, Inc., and Kuparuk River Unit, Anchorage, by Alaska Biological Research, Inc., Fairbanks, Alaska. 71 pages. Jorgenson, M. T. 1988. Revegetation of the Lake State 1 exploratory well site, Prudhoe Bay oil field, Alaska, 1987. Final report prepared by Alaska Biological Research, Inc., for ARCO Alaska, Inc. 50 pages + appendices. Kuropat, P. J., and J. A. Curatolo. 1983. Caribou. Chapter 4 in Lisburne Development Area: 1983 environ- mental studies. Final report prepared for ARCO Alaska, Inc., Anchorage, by Woodward—Clyde Consultants, Anchorage, and Alaska Biological Research, Fairbanks, Alaska. 52 pages. Lawhead, B.E., and J. A. Curatolo. 1984. Distribution and movements of the Central Arctic Herd, summer 1983. Final report to ARCO Alaska, Inc., Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 52 pages. Lawhead, B. E., and L. N. Smith. 1990. Caribou, 1988 Endicott environmental monitoring program. Draft report to Science Applications International Corp. and U.S. Army Corps of Engineers, Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 45 pages. Linkswiler, C., and J. A. Curatolo. 1984. Distribution and movements of the Central Arctic Caribou Herd within the Lisburne development area, Alaska, July 1984. Final report to ARCO Alaska, Inc., Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 40 pages. Pollard, R. H., R. Rodrigues, and R. C. Wilkinson. 1990. Wildlife use of disturbed habitats in arctic Alaska. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage, Alaska. 111 pages + appendices. Pollard, R. H., M. E. Miller, and R. C. Wilkinson. 1992. Caribou distribution in the Prudhoe Bay oil field, sum- mer 1990. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage, Alaska. 40 pages + appendices. Pollard, R. H., and W. B. Ballard. 1993. Parasitic insect abundance and microclimate on gravel pads and tundra, and observations of caribou in the Prudhoe Bay oil field, Alaska, summer 1992. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage, Alaska. 18 pages + appen- dices. - Pollard, R. H., and L. E. Noel. 1994. Caribou distribution and parasitic insect abundance in the Prudhoe Bay oil field, summer 1993. Report by LGL Alaska Research Associates, Inc. Prepared for B.P. Exploration (Alaska) Inc., Anchorage, Alaska. 53 pages + appendices. Literature Cited Brown, J., and R. K. Hogen. 1975. Selected climatic and soil thermal characteristics of the Prudhoe Bay Region. Pages 1-11 in Ecological investigations of the tundra biome in the Prudhoe Bay Region, Alaska. Edited by J. POLLARD, BALLARD, NOEL AND CRONIN: GRAVEL PADS AND TUNDRA 657 Brown. Biological Papers, University of Alaska, Special Report Number 2. University of Alaska, Fairbanks, Alaska. Curatolo, J. A. 1975. Factors influencing local move- ments and behavior of barren—ground Caribou (Rangifer tarandus granti). M.S. thesis. University of Alaska, Fairbanks, Alaska. 145 pages. Curatolo, J. A., and S. M. Murphy. 1986. The effects of pipelines, roads, and traffic on the movements of Caribou, Rangifer tarandus. Canadian Field—Naturalist 100 (2): 218-224. Dau, J. 1986. Distribution and behavior of barren—ground Caribou in relation to weather and parasitic insects. M.S. thesis. University of Alaska, Fairbanks, Alaska. 149 pages. Downes, C. M., J. B. Theberge, and S. M. Smith. 1986. The influence of insects on the distribution, microhabitat choice, and behavior of the Burwash caribou herd. Canadian Journal of Zoology 64: 622-629. Espmark, Y. 1968. Observations of defense reactions to Oestrid flies by semi—domestic forest reindeer (Rangifer tarandus L.) in Swedish Lapland. Zoologische Beitrage. 14: 155-167. Fancy, S.G. 1983. Movements and activity budgets of Caribou near oil drilling sites in the Sagavanirktok River floodplain, Alaska. Arctic 36: 193-197. Gjullin, C. M., R. I. Sailer, A. Stone, and B. V. Travis. 1961. The Mosquitoes of Alaska. U.S. Department of Agriculture., Agricultural Handbook Number 182. 98 pages. Hocking, B., W. R. Richards, and C. R. Twinn. 1950. Observations on the bionomics of some northern Mosquito species (Culicidae: Diptera). Canadian Journal of Research 28: 58-80. Lawhead, B. E. 1988. Distribution and movements of Central Arctic Herd Caribou during the calving and insect seasons. Pages 8—13 in Reproduction and calf sur- vival: Proceedings Third North American Caribou Workshop. Edited by R. D. Cameron and J. L. Davis. Wildlife Technical Bulletin Number 8, Alaska Department of Fish and Game, Juneau, Alaska. Lillesand, T. M., and R. W. Kiefer. 1979. Remote Sensing and Image Interpretation. John Wiley and Sons, New York. 612 pages. MacLean, S. F. 1975. Ecology of tundra invertebrates at Prudhoe Bay, Alaska. Pages 115-123 in Ecological investigations of the tundra biome in the Prudhoe Bay region, Alaska. Edited by J. Brown. Biological Papers, University of Alaska, Special Report Number 2. University of Alaska, Fairbanks, Alaska. Murphy, S. M., and J. A. Curatolo. 1987. Activity bud- gets and movement rates of Caribou encountering pipelines, roads, and traffic in northern Alaska. Canadian Journal of Zoology 65: 2483-2490. Nixon, W. A. C. 1990. Group dynamics and behavior of the Porcupine Caribou Herd during the insect season. M.S. thesis. University of Alaska, Fairbanks, Alaska. 109 pages. Roby, D. D. 1978. Behavioral patterns of barren—ground Caribou of the Central Arctic Herd adjacent to the Trans—Alaska Pipeline. M.S. thesis. University of Alaska, Fairbanks, Alaska. 200 pages. Searby, H. W., and M. Hunter. 1971. Climate of the North Slope of Alaska. NOAA Technical Memorandum. U.S. Department of Commerce. National Oceanic and 658 Atmospheric Administration, National Weather Service. Washington, D. C. 54 pages. Snedecor, G. W., and W. G. Cochran. 1980. Statistical Methods. 7th edition. Iowa State University Press, Ames, Lowa. 507 pages. Snow, W. F. 1976. The direction of flight of Mosquitoes (Diptera, Culicidae) near ground in West African savan- na in relation to wind direction, in the presence and absence of bait. Bulletin Entomological Research 65: 555-562. Walker, D. A., K. R. Everett, P. J. Webber, and J. Brown. 1980. GeoBotanical atlas of the Prudhoe Bay region, Alaska. CRREL Report Number 80-14, U.S. Army Corps of Engineers, Hanover, New Hampshire. THE CANADIAN FIELD-NATURALIST Vol. 110 White, R. G., B. R. Thomson, T. Skogland, S. J. Person, D. E. Russell, D. F. Holleman, and J. R. Luick. 1975. Ecology of Caribou at Prudhoe Bay, Alaska. Pages 151-187 in Ecological investigations of the tundra biome in the Prudhoe Bay region, Alaska. Edited by J. Brown. Biological Papers, University of Alaska, Special Report Number 2. University of Alaska, Fairbanks, Alaska. Zar, J. H. 1984. Biostatistical analysis. Second edition. Prentice Hall, Englewood Cliffs, New Jersey. 718 pages. Received 9 January 1996 Accepted 20 August 1996 Summer Distribution of Caribou, Rangifer tarandus granti, in the Area of the Prudhoe Bay Oil Field, Alaska, 1990-1994 ROBERT H. POLLARD!, WARREN B. BALLARD’, LYNN E. NOEL, and MATTHEW A. CRONIN LGL Alaska Research Associates, Inc., 4175 Tudor Centre Drive, Suite 202, Anchorage, Alaska, 99508 'Present address: 9424 Canton Loop, Anchorage, Alaska 99515 Present address: Arizona Game and Fish Department, 2221 West Greenway Road, Phoenix, Arizona 85023 Pollard, Robert H., Warren B. Ballard, Lynn E. Noel, and Matthew A. Cronin. 1996. Summer distribution of Caribou, Rangifer tarandus granti, in the area of the Prudhoe Bay oil field, Alaska, 1990-1994. Canadian Field-Naturalist 110(4): 659-674. Aerial surveys were conducted within the Prudhoe Bay oil field area in northern Alaska during 1990-1994 to document summer distribution of Caribou (Rangifer tarandus granti). Numbers of Caribou observed per survey ranged from 34 to 13058. When insect activity was moderate to high, more Caribou were observed within the oil field than when insect activ- ity was low. Severe mosquito (Aedes spp.) harassment often resulted in large post-calving aggregations of Caribou in coastal areas. When mosquito harassment subsided, aggregations often moved inland through the oil field or laterally along the coast. On several occasions, large groups of Caribou (2000-4000 individuals) were observed feeding or bedded down in the central portion of the oil field. Caribou were observed on, and used, oil field gravel pads and roads as insect relief habi- tat during the mosquito season in late June through mid-July, and also used the shade of oil field structures when oestrid flies (Hypoderma tarandi and Cephenomyia trompe) were abundant from mid-July to early August. Five years of observa- tions document that Caribou use habitat within and travel through oil fields during summer during periods of high insect activity. Key Words: Caribou, Rangifer tarandus granti, Prudhoe Bay, Alaska, insect harassment, oestrid flies, mosquitoes, post- calving, oil field, aerial survey. The potential impacts of oil field development on Caribou (Rangifer tarandus granti) of the Central Arctic Herd (CAH) has been the focus of intensive research in Alaska's North Slope oil fields since the early 1970s (Clough et al. 1987). Effects of oil fields on movements and habitat-use patterns have been of particular concern (Cameron and Whitten 1976*, 1980; Klein 1980; Curatolo and Murphy 1986; Cameron et al. 1992). The CAH has grown from about 5000 animals in 1975 (Cameron and Whitten 1979*) to 23444 animals in 1992 (Cameron 1993*) and numbered 18093 in 1995 (P. Valkenburg, Alaska Department of Fish and Game, personal communication). The CAH generally win- ters in the northern foothills of the Brooks Range and migrates each spring to calving grounds and summer range on the Arctic Coastal Plain between the Itkillik and Colville rivers (151°00' W longitude) on the west to the Sadlerochit River (145°00' W lon- gitude) on the east (Clough et al. 1987). The Prudhoe Bay oil field (PBOF) is located within the summer range of the CAH. During the calving period, the CAH segregates into segments east and west of the Sagavanirktok River (Cameron and Whitten 1977*; Lawhead 1988). *Unpublished, see Documents Cited section following Acknowledgments. All other dates are to published refer- ences given in Literature Cited. The western segment uses the area in and around the Kuparuk oil field during the post-calving period. The eastern segment uses areas to the east of the PBOF. A 1992 photocensus of the CAH estimated 8602 ani- mals in the eastern segment and 14 842 animals in the western segment (Valkenburg 1993*). The most recent photocensus in 1995 indicated 11 634 animals in the eastern segment and 6459 in the western seg- ment (P. Valkenburg, Alaska Department of Fish and Game, personal communication). During the calving period female Caribou with calves may distance themselves from oil field facili- ties such as roads (Whitten and Cameron 1985; Dau and Cameron 1986; Klein 1991; Cameron et al. 1992). However, the Kuparuk and Milne Point oil field areas west of the PBOF have not been aban- doned, as calving has continued in these areas con- current with expanding oil field development (Cameron et al. 1992). There is little information on use of the PBOF for calving prior to oil field devel- opment in 1970, but it is believed that it was not a concentrated calving area (Child 1973*; Gavin 1983*; Whitten and Cameron 1985; Klein 1991). Furthermore, Whitten and Cameron (1985) suggest- ed that the generally wet conditions in the PBOF area (between the Kuparuk and Sagavanirktok rivers) provides suboptimal calving habitat. CAH Caribou use snow-covered lowlands or well-drained uplands over river valleys or deltas for calving (Carruthers et al. 1987). 659 660 148° 55° 148° 50° 148° 45' 148° 40' 148° 35' 148° 30' 148° 25' 70° 25' 70° 20° 70° 15'f 70% 10) 70° 05'|__ THE CANADIAN FIELD-NATURALIST 148° 20' Vol. 110 148° 15' 148° 10' 148° 05' 148° 00° 147° 55' 147° 50' 147° 45' MAP LOCATION ‘0° 25' 70° 10' 3 4 5 MILES Sie 4.5 KILOMETERS + 70° 05" A ; rd I Ks bs Le ‘aul Q) 148° 25' 148° 20' 148° 15° 148° 10° ee Nes 148° 00" 147° 55' 147° 50° | 148° 55 | 148° 50 [ee L 148° 45' 148° 40' 147° 45' pli 148° 0S' 148° 35' 148° 30° FicureE |. Location of strip-transect centerlines for aerial surveys conducted between 26 June and 8 August, 1990-1994, Prudhoe Bay oil field, Alaska. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities. In contrast to the calving period, the PBOF may have been used frequently by Caribou during the post-calving period (late June through early August) prior to oil field development (Child 1973*; Whitten and Cameron 1983a). Child (1973*:7) identified extensive “traditional trail systems of Caribou” in the area that is now the PBOF. Whitten and Cameron (1985) and Johnson and Lawhead (1989*) suggested that, since the mid-1970s, movements by large post- calving aggregations through the PBOF have become rare or ceased entirely. However, harass- ment by mosquitoes and oestrid flies (warble flies, Hypoderma tarandi and nose bot flies, Cephenomyia trompe) have a large influence on Caribou post-calv- ing distribution and movements (White et al. 1975; Curatolo et al. 1982*; Lawhead 1988), and there is evidence that the oil field areas may provide relief habitat from insects (Pollard et al. 1996). There is interest in expanding oil field develop- ment in arctic Alaska, including areas within the Arctic National Wildlife Refuge used as calving and post-calving habitat by the Porcupine Caribou Herd (Clough et al. 1987). Because impact assessments at existing oil fields will be used to infer potential impacts at new developments, it is important to docu- ment accurately the extent of post-calving use of the PBOF. The objectives of this study were to quantify the post-calving distribution and number of Caribou in the area of the PBOF over five years and relate caribou distribution and numbers to insect activity. Study Area The study was conducted on the northern edge of the Arctic Coastal Plain of Alaska north of 70°05'N latitude between the Kuparuk and Sagavanirktok rivers (Figure 1). The 1394 km? study area contains the PBOF and is characterized by little elevational relief, many shallow lakes and drained lake basins, and a variety of vegetation types dominated by wet and moist tundra (Walker et al. 1980). The PBOF includes 53 producing well pads, 31 exploration pads, 8 gathering centers, 2 gravel land- 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 661 TABLE |. Estimated numbers and sex/age composition of Caribou in relation to relative insect activity observed during aer- ial surveys conducted in the Prudhoe Bay study area, Alaska, during the post-calving period, 1990-1994. Year Relative Date Bulls Cows Calves Yearlings Unclassified Total Insect Activity* 1990 4,5 July 3151 4203 2947 563 3 10867 high 12 July 60 14 + 2 2 82 low 19 July 253 372 179 58 66 928 low 26 July 44 35 9 1 20 109 low 1 August 77 45 12 1 24 159 low 1991 26 June 137 27 23 29 51 267 low 3 July 259 25 25 8 53 370 low-moderate 10 July 268 156 121 6 230 781 low 16 July 2151 1801 1797 0 1330 7079 moderate 25 July 522 108 96 if G32 1465 low 31 July 35 12 7 2 76 132 low 1992 9 July 2535 2746 2641 427 1982 10331 high 10 July 4074 3242 3186 346 2210 13058 moderate 13, 14 July 72 24 23 9 19 147 low 19 July 2503 1579 1572 452 1538 7644 high 23 July 566 428 405 91 446 1936 high 25 July 826 541 530 78 528 2503 high 28 July 254 96 92 10 141 593 moderate 31 July 44 23 18 D 60 147 low 1993 1 July 1315 434 209 171 113 2242 low 6 July 60 10 3 2, 0 75 low 10 July 438 727 713 0 1210 3088 high 17 July 190 454 252 32 215 1163 moderate 19 July 2055 625 542 145 430 3797 high 26 July 52 31 10 0 3 96 low 1994 5 July 129 55 40 11 4] 276 » 11 July 804 922 562 54 DT 2599 bp 19 July 138 97 64 6 44 349 Y 26 July 18 12 2 0 2 34 Y 31 July 198 496 386 18 119 1217 P 7,8 August 32 36 9 1 1 719 b “Quantified in detail by Pollard et al. (1996). Insect activity not monitored. ing strips for jet aircraft, and 2 base camps. Other service facilities are present both in the oil field and the nearby community of Deadhorse. All facilities are supported by elevated gravel pads (approximate- ly 1.5 m) and are connected by a network of gravel roads (totaling approximately 220 km in length) and above-ground pipelines. Our study area included the entire PBOF complex, and also considerable area surrounding the oil field (Figure 1). For example, the southeast, southwest, and northwest portions of the study area have large areas which are distant from oil field infrastructure. A detailed assessment of Caribou distribution in relation to infrastructure is beyond the scope of this report, and will be described elsewhere. The results of that analysis show no evidence of avoidance of infrastruc- ture by Caribou. Here we are concerned with the numbers, distribution, and movements of caribou rela- tive to the PBOF as a whole. For this paper, we will loosely define the PBOF as the area within 10 km of roads and oil field infrastructure. Methods During 1990-1994 between 26 June and 8 August, we conducted a series of aerial surveys of the study area from fixed-wing aircraft (Cessna 206) with two 662 THE CANADIAN FIELD-NATURALIST Vol. 110 Es REAPER E AE LR 12 July 1990 No. of Groups No. of Groups Range inRange Range in Range 27 a <10 39 @ <10 @ 10-100 4 A 10-100 0 | @ 100-1000 7 A 100-1000 0 , 1 @>1000 1 A 210000 is AGE a A A A A } : \ e | 5 Kilometers e a. ! L P 19 July 1990 26 July 1990 No. of Groups No. of Groups RB in Ran in Ran @ <10 59 <0) 63 @ 10-100 ) A 10-100 0 @ 100-1000 1 A j00-1000 0 @ > 1000 ) A +1000 o | | | | if | Oo | ae: e | GT} Wf) @ | 5 Kilometers A Voy iP H la soa Ve f py Ll FiGureE 2 a-b. Distribution of Caribou observed in the study area during five aerial surveys conducted during 4 july — 1 August 1990. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities. observers and pilot flying at 130 km/h and 90 m above ground level. Each survey consisted of flights along 29 systematically spaced, fixed-width strip transects (Caughley 1977). Transects were oriented north-south and centered on longitudinal section and township lines mapped on 1:63 360-scale U.S. Geological Survey (USGS) topographic maps. Survey transects were 1.6 km wide and spaced 1.6 km apart. During surveys, each observer was respon- sible for searching an 800 m-wide swath on one side of the transect centerline. Aluminum rods attached to the aircraft wing struts enabled visual control of tran- sect strip-width. Surveys were conducted between 0800-1700 Alaska Daylight Time. During surveys, when Caribou were sighted, we slowed the survey aircraft, circled each group, and estimated total number and sex/age classes (i.e., bulls, cows, calves, yearlings, and unclassified 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 663 5 Kilometers re Cc. 1 Aug 1990 : No. of Groups in Ran Ran. 87 @ <10 @ 10-100 1 FIGURE 2 c. See caption on facing page. adults). Data were entered into a laptop computer linked to the GPS receiver in the airplane. Locations of Caribou groups were determined by using GPS coordinates recorded on the transect line corrected with the visual estimation of their distances from the transect line. Caribou data were combined with digi- tal base-map data in MapInfo® (MapInfo Corp., 200 Broadway, Troy, New York 12180) and mapped for each survey. For mapping purposes Caribou group size was categorized into four size classes: < 10, 10- 100, 101-1000, and > 1000 individuals. During 1992 and 1993, we systematically moni- tored weather patterns and insect activity on a daily basis (Pollard et al. 1996). Weather variables were measured using automated weather stations and hand- held instruments (Pollard et al. 1996). Insect abun- dance was measured using sweep nets, insect traps, and ground observations of Caribou. Insect activity during 1990 and 1991 was subjectively estimated based on ground observation before and after surveys, and by interviews with field technicians located on the ground during aerial surveys. We classified relative insect activity as relatively low, moderate, or high. Insect activity was not measured during 1994. Results Distribution and abundance of Caribou within the study area was highly variable within and among years (Table 1). Numbers of Caribou observed on any survey ranged from 34 to 13 058. Movement of Caribou through the oil field appeared to be directly related to insect activity. In general, when insect activity was low, most Caribou moved inland away from the coast and insect-relief habitat within the oil field. However, when insect activity was moderate or high, Caribou either were present within or were moving through the oil field to reach insect-relief habitats along the coast or on sandbars within major river systems (Table 1, Figures 2-6). When Caribou moved through or were present within the oil field, it was necessary for them to cross pipelines, roads, and associated infrastructure. We describe Caribou distri- bution and abundance for each year of study below. More detailed descriptions of each survey are given in unpublished reports available from the authors (Pollard et al. 1992a*, b*; Pollard and Ballard 1993*: Pollard and Noel 1994*, 1995*). 1990. — The 4-5 July survey revealed large groups of Caribou in the coastal region of Simpson Lagoon and the Kuparuk River Delta (Figure 2a). These groups comprised most of the 10 867 Caribou in the study area. Insect activity was high during this survey (Table 1). Smaller groups were cistributed throughout the central portion of the PBOF within 18 km of the coast. On 12 July, insect activity was low and the coastal aggregations of Caribou had dis- persed (Figure 2a). Most Caribou had left the study area and only 82 individuals were observed in small groups spread throughout the PBOF. On 19 July, insect activity was again low and most Caribou were in small groups distributed along 30 km between the west channel of the Sagavanirktok River and the Putuligayuk River, and within the PBOF (Figure 2b). One large group of 400 individuals was observed near the Trans-Alaska Pipeline, about 11 km south of the central portion of the PBOF. On 26 July, insect activity was still low and 109 individuals were distributed in small groups along the 664 THE CANADIAN FIELD-NATURALIST Vol. 110 5 Kil@meters ———— | a. 26 June 19914 3 July 19914 No. of Groups No. of Groups Range inRange Range in Range @® <10 49 a <10 45 @ i0-100 7 A @ 100-1000 0 @ > 1000 10-100 12 A 100-1000 o A >1000 ie} me | @ 1 | | 5 Kilometers [ce ces eee 10 July 19914 16 July 1991 No. of Groups No. of Groups Range in Range Range in Range @ <10 97 <10 21 @ 10-100 15 @ 100-1000 1 100-1000 8 A A A @ > 1000 0 «6B >1000 3 10-100 6 zie FiGure 3 a-b. Distribution of Caribou observed in the study area during six aerial surveys conducted during 26 June—31 July 1991. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities. Sagavanirktok River and along the southern limit of the study area (Figure 2b). Some of these groups were close to roads and infrastructure. The 1 August survey was also during low insect activity and revealed small groups of Caribou scattered along the southern portion of the study area and throughout the PBOF (Figure 2c). 1991. — During the 26 June survey, insect activi- ty was low and a total of 267 Caribou were observed along the Kuparuk, Putuligayuk, and Sagavanirktok river channels and in the PBOF (Figure 3a). Twelve small groups were observed southeast of the eastern channel of the Sagavanirktok River. On 3 July, insect activity was classified as low to moderate and 370 individuals were observed within 16-18 km of the coast. Small groups were found across the study area, including areas along the Sagavanirktok and Kuparuk rivers and in the PBOF (Figure 3a). 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 665 | § ° 5 Kilometers SS Cc. N EEE 31 July 1991 No. of Groups inR Lo 25 July 1991 No. of Groups 1 R @ <10 151 a <10 37 @ 40-100 33 A 10-100 1 @ 100-1000 1 @ > 1000 10) A 100-1000 0 A >1000 () FIGURE 3 c. See caption on facing page. Over twice as many Caribou (n = 781) were observed during the 10 July survey as on 3 July, although insect activity was again low (Table 1). Small groups were scattered throughout the study area, including the PBOF, and there was a large con- centration inland, south of the PBOF and west of the Trans-Alaska Pipeline (Figure 3b). Relatively high winds and low temperatures suppressed insect activi- ty on 10 July (Pollard et al. 1996). The 16 July survey indicated a large influx of Caribou into the study area as we observed 7079 individuals (Table 1). Most were located along the Kuparuk River and near the coast in the western portion of the PBOF (Figure 3b). Smaller groups were observed along the coast in the Sagavanirktok River Delta. Mosquitoes were moderately active during the early part of the day, but activity dimin- ished in the afternoon as winds increased (Pollard et al. 1996). Ground-based observations conducted at 1700 hours, after the aerial survey, revealed that many of the large groups observed during the aerial survey had combined and were moving south through the center of the oil field. A group of approximately 4000 Caribou moved clockwise through the entire PBOF and was observed finally on the floodplain of the Kuparuk River. To make these movements, this group crossed at least six roads and ten pipelines. On 25 July, with low insect activity, there were 1465 Caribou in the study area. Most were distribut- ed east of the west channel of the Sagavanirktok River, and there were small groups in the PBOF (Figure 3c). The 31 July survey, also with low insect activity, indicated that even fewer Caribou were in the study area; only 132 Caribou were sighted. Most of these individuals were inland in the southwestern portion of the study area, with some small groups in the PBOF (Figure 3c). 1992. — During the aerial survey on 9 July, 10 331 Caribou were observed in the study area (Table 1). Mosquito activity was high on this date. An influx of Caribou from the west resulted in high concentrations in coastal areas between the Kuparuk River and the western shore of Prudhoe Bay (Figure 4a). Ground observations conducted on the evening of 9 July, after the aerial survey, suggested that mosquito activity had decreased and several large groups had combined and traveled south through the oil field. This is the same pattern observed on 16 July 1991 with moderate mosquito activity. By 2200 hours on 9 July, a group of approximately 4000 Caribou was observed bedded down in the central portion of the oil field. On 10 July, 13 058 Caribou were observed in the study area. Mixed-sex Caribou groups were widely distributed across the oil field (Figure 4a). Approximately 3000 Caribou were observed on grav- el drilling pads and roads, probably seeking relief from mosquitoes which were moderately active. We observed several groups from the ground which encountered roads with high levels of vehicular traf- fic. The animals were delayed in crossing until traffic subsided, and then crossed or walked along the roads. The 13-14 July survey was completed over a two-day period due to mechanical problems with the aircraft. Only 147 Caribou were observed in the 666 THE CANADIAN FIELD-NATURALIST | < 5 Kilometers p>) AE ON ASR 9Ji iy 4 9 92 EA ERS UA I 10 July 1992 No. of Groups No. of Groups Range inRange Range in Range © <10 15 <10 30 @ 10-100 «15 10-100 29 @ 100-1000 9 100-1000 20 A A A A >1000 2 t a: , 13-14 1 19 July 1 No. of Groups No. of Groups Range inRange Range in Range : \ Pi @ <10 46 aA <10 31 @ 10-100 2 4 10-100 9 {x @ 100-1000 0 A 100-1000 13 | b ) (5p | @>1000 0 A >1000 (1 UD \ ON i f @ oy Py YI i e | | 1 | | j | | 5 Kilometers | b. \—— ec FicurE 4 a-b. Distribution of Caribou observed in the study area during eight aerial surveys conducted during 9 July—31 July 1992. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities. study area during the survey, most of which were in small groups south of the PBOF near the Sagavanirktok River (Figure 4b) and scattered throughout the PBOF. Insect activity was low dur- ing this survey. During the 19 July survey, 7644 Caribou were observed in the study area (Table 1). Observations from this survey, and ground-based observations conducted during 20-22 July, indicated large groups of Caribou moved through the PBOF on these dates. Insect activity (mosquitoes and oestrid flies) was moderate to high on 19-22 July. Many Caribou were on gravel roads and pads, or in the shade of build- ings and pipelines. These Caribou displayed signs of insect harassment (e.g., shaking their heads, bodies, and twitching their tails, and kicking) (Pruitt 1960; Espmark 1968; Kelsall 1968; Skoog 1968; Curatolo 1975; Dau 1986). Caribou remained on the gravel Vol. 110 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 667 y\ 5 Kilometers 25 July 1992 No. of Groups Range in Range 23 July 1992 No. of Groups Range in Range e <10 69 MyRe10 75 @ i0-100 §=618 4 10-100 14 @ 100-1000 4 A 100-1000 4 @ > 1000 OA Sicc0. 0 5 Kilometers be} d Miberermeroe 28 July 1992 31 July 1992 No. of Groups No. of Groups Range inRange Range in Range ® <10 83 a <10 63 @® 10-100 1 A 10-100 0 A 00-1000 0 A >1000 «OO @ 100-1000 3 @ > 1000 0 FiGurE 4 c-d. See caption on facing page. structures from several minutes to >2 hours. Many large Caribou groups were observed in the Sagavanirktok River delta, or on barren mud flats and gravel river bars, probably seeking insect relief from coastal winds (Figure 4b). Several groups of 100-1000 animals were in the center of the PBOF. On 23 July, 1936 Caribou were observed in the study area (Table 1). Mosquito abundance was low, but oestrid activity remained high. Approximately 27% of the Caribou observed were on gravel pads, roads, or under buildings seeking relief from insects. The highest concentrations of Caribou occurred in the Kuparuk and Sagavanirktok river channels and deltas (Figure 4c). Many small groups were scattered throughout the PBOF. A total of 2503 Caribou were observed on 25 July within the study area (Table 1). A majority were in large groups on gravel bars in the Sagavanirktok River. Many smaller groups were widely distributed across the southern half of the study area. Cooler temperatures, fog, and moderate winds from the northeast suppressed mosquito 668 & 5 Kilometers a THE CANADIAN FIELD-NATURALIST Vol. 110 6 July 1993 No. of Groups No. of Groups i in Ran @® <10 63 a <10 15 @ to100 330 A @ 100-1000 6 @ > 1000 0 10-100 2 A 100-1000 0 A >1000 Oo 5 Kilometers —_—_— b. UTERO SRT Sas 17 July 1993 No. of Groups in Ri 10 July 1993 No. of Groups oR Ran ia @ <10 15 <10 35 @ i0-100 0 10-100 14 @ 100-1000 4 400-1000 2 R A A A A >1000 0 | | i | L FiGurE 5 a-c. Distribution of Caribou observed in the study area during six aerial surveys conducted during | July — 26 July 1993. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities activity but not oestrid fly activity on this day (Pollard et al. 1996). During the 28 July survey, 593 Caribou were observed in the study area (Table 1). The majority were located in small groups in the southern half of the study area. There were a few large groups along the Sagavanirktok River (Figure 4d). A similar dis- tribution was observed on 31 July for 147 Caribou, except several small groups were in the PBOF (Figure 4d). Insect activity was moderate on 28 July and low on 31 July, and many Caribou exhibited behavior characteristic of animals experiencing oestrid fly harassment. 1993. — During the 1 July survey, insect activity was low, and 2242 Caribou were observed (Table 1). 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 669 j “5 Kilometers \ eI Cc. 19 July 1993 No. of Groups in Range Range Range __in Range @ <10 25 a <10 51 @ 10-100 16 4 10-100 1 @ 100-1000 9 A 100-1000 0 @ > 1000 1 A 2100 oo FIGURE 5c. See caption on facing page. The majority of these were in the western portion of the study area (Figure 5a). Most of the larger groups were along the Kuparuk River. Many moderately sized groups were observed in the drainages of the Kuparuk, Putuligayuk, and Sagavanirktok rivers and in the PBOF (Figure 5a). We observed only 75 Caribou on the 6 July survey. Most of these animals were in small groups widely dispersed in the central portion of the PBOF (Figure 5a). Few mosquitoes were active on either 1 July or 6 July (Pollard et al. 1996). On 10 July, insect activity was high, and 3088 Caribou were observed. The majority of Caribou were in four large groups, three of which were in or adjacent to the Kuparuk River (Figure 5b). The fourth large group was located on the coast in the northwest portion of the study area. There were a ‘few small groups in the PBOF. On 17 July, 1163 Caribou were observed (Table 1). The majority were scattered across the northern half of the study area, primarily in the northwest por- tion. Most Caribou occurred in small groups, several of which were observed on gravel pads and roads in the PBOF. The two largest groups (140 and 300 ani- mals) were moving northeast through the PBOF. Observations of Caribou behavior and quantitative measures of insect abundance suggested that oestrid flies were moderately active and mosquito activity was low (Pollard et al. 1996). The number of Caribou (n =3797) observed on 19 July had more than tripled since 17 July, probably as a result of an influx of Caribou from the Kuparuk oil field to the west, as insect activity increased over those two days. As on 17 July, the majority of Caribou were located in the northwest section of the study area (Figure 5c). Sixty-two percent of the ani- mals were in four large groups that were moving north along the Kuparuk River. Other groups of 10-1000 were scattered throughout the PBOF. Oestrid fly activity was high on 19 July, but mosqui- to activity was low. On 26 July, only 96 Caribou were observed (Table 1). These animals were widely scattered across the study area in small groups pri- marily south of the PBOF (Figure 5c). Insect activity was low on 26 July. 1994. — During the 5 July survey, 276 Caribou were observed (Table 1, Figure 6a). Sixty-two percent of the animals were located in the northwest section of the study area. The remainder were in small groups scattered throughout the PBOF and near the Sagavanirktok River. On 11 July, we observed the largest number of Caribou (i.e., 2599) for any survey of 1994. Ninety-four percent of the Caribou were within four groups. Three of these groups were in the south and west side of the PBOF, and one was on the coast near the mouth of the Kuparuk River. Relatively small numbers of Caribou occurred in the study area on 19 and 26 July (Figure 6b). On the 31 July survey, 1217 Caribou were observed in the study area (Table 1). Caribou were widely-scattered across the study area, including the PBOF. Eighty- two percent of the Caribou were located in the south- west corner of the study area (Figure 6c). In the 7-8 August survey, only 79 Caribou were observed widely scattered throughout the PBOF. Discussion It was suggested that large post-calving aggre- gations stopped moving through the PBOF as a 670 THE CANADIAN FIELD-NATURALIST Vol. 110 4 5 Kilometers _-_—— @ > 1000 EE SES EE OE ESTER / RR CBI EST TO 5 July 1994 11 July 1994 No. of Groups No. of Groups Range in Range __in Range @® <10 37 A <10 24 @ i0-100 «8 A @ 100-1000 0 10-100 3 A 100-1000 3 A 1 >1000 5 Kilometers --——-— ' 19 July 1994 26 July 1994 No. of Groups No. of Groups Range inRange Range in Range @ <10 55 a 1000 0 @ 100-1000 0 @>1000 0 of: FiGureE 6 a-b. Distribution of Caribou observed in the study area during six aerial surveys conducted during 5 July — 8 August 1994. Solid lines are roads or pipelines and rectangles are gravel pads which support oil field facilities. result of oil field development (Whitten and Cameron 1985; Johnson and Lawhead 1989*). However, from 1975-1989, post-calving surveys of the PBOF were limited in intensity and number (Cameron and Whitten 1976*; Whitten and Cameron 1983b; Cameron et al. 1986; Carruthers and Jakimchuk 1986; Carruthers et al. 1987). Because Caribou frequently move into and out of the PBOF in response to changing environmental conditions, infrequent, non-systematic, and/or low- intensity aerial surveys may be of limited value for examining Caribou distribution in relation to oil field infrastructure. Perhaps, more importantly, the relatively small numbers of Caribou observed in the PBOF in the mid-1970s to mid-1980s (Whitten and Cameron 1985; Lawhead 1988; Johnson and Lawhead 1989*), and the large numbers we observed in the 1990s (Table 1), may be related to 1996 POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 671 | @e A , 4 | 286 | Kilometers c | -——4 7, 8 August 1994 No. of Groups Range in Range a <10 51 31 July 1994 No. of Groups Range in Range @ <10 87 g @ 10-100 19 A 10-100 0 @ 100-1000 2 A 100-1000 0 A >1000 10) FIGURE 6 c. See captions on facing page. the increasing number of Caribou within the CAH. The CAH grew from 5000 to 23 000 animals between 1975 and 1992 (Whitten and Cameron 1983b; Valkenburg 1992*; Cameron 1993). It is also possible that Caribou have habituated to the oil fields over the last two decades. There is no hunting or deliberate harassment of animals in the PBOF, so this is a likely possibility. Weather conditions are generally cooler and windier, and there is less vegetation near the Beaufort Sea coast than inland in the PBOF area (Dau 1986; Linkswiler and Curatolo 1984*; White et al. 1975). These factors are probably responsible for fewer mosquitoes on the coast than inland. Caribou often use the coast as mosquito-relief habitat on warm days, and move inland with cooler evening temperatures (Roby 1978). This occurred on 16 July 1991 and 9 July 1992. Caribou are not always on the coast when mosquitoes are active, however. On 10 July 1992, mosquito activity was moderate, and few Caribou were on the coast. Most Caribou were wide- ly distributed across the PBOF, and about 3000 were on gravel pads and roads, probably seeking relief from mosquitoes. Lawhead and Curatolo (1984*) suggested that, under moderate levels of insect harassment, Caribou will move only as far toward the coast as needed to reach ameliorating tempera- tures or wind. In the Kuparuk oil field to the west of the PBOF, Dau (1986) suggested forage was of high- er quality inland compared to that near the coast, and that Caribou sought such forage after expending large amounts of energy avoiding insects. When only oestrid flies are present (from late July through mid-August), coastal areas apparently offer limited relief to Caribou (Lawhead and Smith 1990*). Unvegetated sites, such as gravel bars and mud flats, and elevated sites, such as pingos and sand dunes, are used as oestrid fly-relief habitat (White et al. 1975; Fancy 1983; Kuropat and Curatolo 1983*; Murphy and Curatolo 1987; Pollard et al. 1996). We observed Caribou using oil field gravel pads and roads as well as gravel bars in the Sagavanirktok and Kuparuk rivers when oestrid flies were active. The influence of oestrid flies can over- ride any tendency to avoid oil field structures or activities (White et al. 1975; Curatolo et al. 1982*; Dau 1986; Murphy and Curatolo 1987). Indeed, gravel pads and roads appear to provide insect-relief habitat similar to sparsely vegetated gravel bars (Murphy and Curatolo 1987; Pollard et al. 1996). Caribou often are observed lying, standing, or travel- ing on gravel pads and roads or in the shade of build- ings and pipelines (Roby 1978; Fancy 1983; Murphy and Curatolo 1987; Johnson and Lawhead 1989; Pollard and Noel 1994*). During our survey on 19 July 1992, 2400 of the 7644 Caribou observed were on gravel pads and roads in the PBOF, and the rest were on mud flats and gravel bars in river deltas. Bergerud et al. (1984: 7) examined the impacts of human disturbances on Caribou and concluded that, “Caribou apparently have a high degree of resilience to human disturbance, and seasonal movement pat- terns and extent of range occupancy appear to be a function of population size rather than of extrinsic disturbance.” Our study of post-calving use of the PBOF from 1990-1994 also indicates that Caribou distribution and movements are influenced by weather and insects more than human disturbance. 672 Our observations are similar to those reported 20 years ago, prior to extensive oil field development. White et al. (1975) found that insect-harassed Caribou moved into the Prudhoe Bay area, across the Kuparuk River Delta, or down the drainages of the Sagavanirktok River to gain access to coastal insect relief habitat. When insect levels diminished, Caribou dispersed inland, moving in a southwesterly direction through the area now occupied by the PBOF. Recently, Cameron et al. (1995) examined the dis- tribution and movements of 6 to 40 radio-collared Caribou annually during 1980-1993 in relation to a quadrant surrounding the PBOF complex. They relo- cated each Caribou at least once during the period 25 June through 10 August. They concluded that few Caribou were in the general area of the PBOF com- plex. Our data suggest that Caribou distribution and use of the PBOF area may be high or low, and is highly variable and can change within hours. Our periodic daily surveys may even have under-repre- sented the occurrence of Caribou in the PBOF. Daily surveys and/or relocation of radio-collared Caribou every 3 to 4 hours would better document the use of the PBOF by Caribou. Our observations show that Caribou can and do traverse oil field areas during periods of insect harassment. Others have also shown that Caribou cross under elevated pipelines and over roads to access insect-relief habitat (Curatolo and Murphy 1986; Lawhead 1988). Roads, pipelines, and human activity may block, delay, or deflect Caribou as they move through the PBOF, but movements of large groups do occur. In summary, our observations show the use of, and movement through, the PBOF and surrounding area by Caribou in small and large groups. Caribou occurrence is variable, and depends on weather and insect conditions. Acknowledgments This research was funded by BP Exploration (Alaska) Inc. (BPXA). We would like to thank C. Herlugson, S. Taylor, D. Beaubien, and M. Gilders of BPXA for their support during this study. Digital basemaps were provided by AeroMap U.S., Inc. Special thanks go to M. McDonald, D. Cornell, and J. Talcott for their logistical support and piloting exper- tise. R. Skoog, M. Miller, B. Rodrigues, J. Luginbuhl, and D. Tremaine provided able assistance in the field, C. Wilkinson provided assistance in GIS analyses and map production, and A. Bishop provided assis- tance in report production. We also thank W. Wilson, D. Esler, B. Pierson, and two anonymous reviewers for their comments on the paper. Documents Cited [marked * in text citations] Cameron, R. D. 1993. Distribution and productivity of the Central Arctic Caribou Herd in relation to petroleum THE CANADIAN FIELD-NATURALIST Vol. 110 development: case history studies with a nutritional per- spective. Alaska Department of Fish and Game, Federal Aid in Wildlife Restoration Progress Report, Project W- 24-1 Study 3.35. December 1993. Juneau, Alaska. 34 pages. Cameron, R. D., and K. R. Whitten. 1976. First interim report on the effects of the Trans-Alaska Pipeline on Caribou movements. Special Report Number 2. Joint State/Federal Fish and Wildlife Advisory Team, Anchorage, Alaska. 38 pages. Cameron, R. D., and K. R. Whitten. 1977. Second inter- im report on the effects of the Trans-Alaska Pipeline on Caribou movements. Special Report Number 8. Joint State/Federal Fish and Wildlife Advisory Team, Anchorage, Alaska. 10 pages. Child, K. N. 1973. 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Final report to ARCO Alaska, Incorporated, and Kuparuk River Unit, Anchorage, by Alaska Biological Research, Incorporated, Fairbanks, Alaska. 71 pages. Kuropat, P., and J. A. Curatolo. 1983. Caribou. Chapter 4 in: Lisburne development area: 1983 environmental studies. Final report to ARCO Alaska, Incorporated, Anchorage, by Woodward-Clyde Consultants, Anchorage, and Alaska Biological Research, Fairbanks, Alaska. 52 pages. Lawhead, B. E., and J. A. Curatolo. 1984. Distribution and movements of the Central Arctic Herd, summer 1983. Final report to ARCO Alaska, Incorporated, Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 52 pages. Lawhead, B. E., and L.N. Smith. 1990. Caribou, 1988 Endicott environmental monitoring program. Final report to Science Applications International Corporation and United States Army Corps of Engineers, Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 106 pages + appendices. Linkswiler, C., and J. A. Curatolo. 1984. Distribution and movements of the Central Arctic Caribou Herd within the Lisburne development area, Alaska, July 1984. Final report to ARCO Alaska, Incorporated, Anchorage, by Alaska Biological Research, Fairbanks, Alaska. 40 pages. - Pollard, R. H., and W. B. Ballard. 1993. Caribou distri- bution in the Prudhoe Bay oil field, summer 1992. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage. On file at BP Exploration (Alaska) Incorporated, Anchorage, Alaska. 24 pages + appendix. 1996 Pollard, R. H., P. C. Lent, M. E. Miller, and R. C. Wilkinson. 1992. Caribou distribution in the Prudhoe Bay, Kuparuk, and Milne Point oil fields, Summer 1991. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage. On file at BP Exploration (Alaska) Incorporated, Anchorage, Alaska. 25 pages + appendix. Pollard, R. H., M. E. Miller, and R. C. Wilkinson. 1992. Caribou distribution in the Prudhoe Bay oil field, Summer 1990. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage. On file at BP Exploration (Alaska) Incorporated, Anchorage, Alaska. 40 pages + appendices. Pollard, R. H., and L. E. Noel. 1994. Caribou distribution and parasitic insect abundance in the Prudhoe Bay oil field, summer 1993. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., Anchorage. On file at BP Exploration (Alaska) Incorporated, Archorage, Alaska. 49 pages + appendices. Pollard, R. H., and L. E. Noel. 1995. Caribou distribution in the Prudhoe Bay oil field, Alaska, summer 1994. Final report by LGL Alaska Research Associates, Inc. Prepared for BP Exploration (Alaska) Inc., On file at BP Exploration (Alaska) Inc., Anchorage, Alaska. 23 pages + appendices. Valkenburg, P. 1992. Central Arctic Caribou Herd Caribou/Subunits 26B and 26C. Pages 187-198 in Caribou. 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Redistribution of calving Caribou in response to oil field development on the arctic slope of Alaska. Arctic 45: 338-342. Cameron, R. D., and K. R. Whitten. 1980. Influence of the Trans-Alaska Pipeline corridor on the local distribu- tion of caribou. Pages 475-484 in Proceedings Second International Reindeer/Caribou Symposium. Edited by E. Reimers, E. Garner, and S. Skjenneberg. Direktoratet For Vilt Og Ferskvannsfisk, Trondheim, Norway. Cameron, R. D., K. R. Whitten, and W. T. Smith. 1986. Summer range fidelity of radio-collared Caribou in Alaska’s Central Arctic Herd. Rangifer, Special Issue 1: 51-55. POLLARD, BALLARD, NOEL, AND CRONIN: SUMMER DISTRIBUTION OF CARIBOU 673 Carruthers, D. R., S. H. Ferguson, and L. G. Sopuck. 1987. Distribution and movements of Caribou, Rangifer tarandus, in the central arctic region of Alaska. Canadian Field-Naturalist 101: 423-432. Carruthers, D. R., and R. D. Jakimchuk. 1986. Caribou of the central arctic region of Alaska in relation to adja- cent Caribou herds. Rangifer, Special Issue 1: 65-71. Caughley, G. 1977. Sampling in aerial survey. Journal of Wildlife Management 41: 605-615. Clough, N. K., P. C. Patton, and A. C. Christiansen, Editors. 1987. Arctic National Wildlife Refuge, Alaska, coastal plain resource assessment-report and rec- ommendation to the Congress of the United States and final legislative environmental impact statement. Volume 1. United States Fish and Wildlife Service, United States Geological Survey, and Bureau of Land Management, Washington, D.C. 208 pages. Curatolo, J. A. 1975. Factors influencing local move- ments and behavior of barren-ground Caribou (Rangifer tarandus granti). M.S. thesis, College of Biological Science and Renewal Resources, University of Alaska, Fairbanks, Alaska. 145 pages. Curatolo, J. A., and S. M. Murphy. 1986. The effects of pipelines, roads, and traffic on the movements of Caribou, Rangifer tarandus. Canadian Field-Naturalist 100: 218-224. Dau, J. R. 1986. Distribution and behavior of barren- ground Caribou in relation to weather and parasitic insects. M.S. thesis, College of Natural Science, University of Alaska, Fairbanks, Alaska. 149 pages. Dau, J. R., and R. D. Cameron. 1986. Effects of a road system on Caribou distribution during calving. Rangifer, Special Issue 1: 95-101. Espmark, Y. 1968. Observations of defense reactions to oestrid flies by semi-domestic forest reindeer (Rangifer tarandus L.) in Swedish Lapland. Zoologische Beitrage 14: 155-167. Fancy, S. G. 1983. Movements and activity budgets of Caribou near oil drilling sites in the Sagavanirktok River floodplain, Alaska. Arctic 36: 193-197. Kelsall, J. P. 1968. The migratory barren-ground Caribou of Canada. Canadian Wildlife Service Monograph Number 3. Queen’s Printer, Ottawa, Ontario. 339 pages. Klein, D. R. 1980. Reaction of Caribou and reindeer to obstructions — a reassessment. 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Parasitic insect abundance and microcli- 674 mate of gravel pads and tundra within the Prudhoe Bay oil field, Alaska, in relation to use by Caribou, Rangifer tarandus granti. Canadian Field-Naturalist 110(4): 649-658. Pruitt, W.O. 1960. Behavior of the barren-ground Caribou. Biological Papers of the University of Alaska, Number 3: 1-44. Roby, D. D. 1978. Behavioral patterns of barren-ground Caribou of the Central Arctic Herd adjacent to the Trans-Alaska oil pipeline. M.S. thesis, University of Alaska Fairbanks, Alaska. 200 pages. Skoog, R.O. 1968. Ecology of the Caribou (Rangifer tarandus granti) in Alaska. Ph.D. thesis, University of California, Berkeley, California. 669 pages. Walker, D. A., K. R. Everett, P. J. Webber, and J. Brown. 1980. Geobotanical atlas of the Prudhoe Bay Region, Alaska. Cold Regions Research and Engineering Laboratory Report 80-14, United States Army Corps of Engineers, Hanover, New Hampshire. 69 pages. White, R. G., B. R. Thomson, T. Skogland, S. J. Person, D. E. Russell, D. F. Holleman, and J. R. Luick. 1975. Ecology of Caribou at Prudhoe Bay, Alaska. Pages THE CANADIAN FIELD-NATURALIST Vol. 110 151-187 in Ecological investigations of the tundra biome in the Prudhoe Bay region, Alaska. Edited by J. Brown. Biological Paper, University of Alaska, Special Report Number 2, University of Alaska, Fairbanks, Alaska. Whitten, K. R., and R. D. Cameron. 1983a. Population’ dynamics of the Central Arctic Herd, 1975-1981. Acta Zoologica Fennica 175: 159-161. Whitten, K. R., and R. D. Cameron. 1983b. Movements of collared Caribou, Rangifer tarandus, in relation to petroleum development on the arctic slope of Alaska. Canadian Field-Naturalist 97: 143-146. Whitten, K. R., and R. D. Cameron. 1985. Distribution of Caribou calving in relation to the Prudhoe Bay oil field. Pages 35-39 in Proceedings of the First North American Caribou Workshop. Edited by A. M. Martell and D. E. Russell. Canadian Wildlife Service Special Publication, Ottawa, Ontario. Received 29 January 1996 Accepted 17 September 1996 Tree Species Composition, Structure, and Carbon Storage in Stands of Urban Forest of Varying Character in Halifax, Nova Scotia B. FREEDMAN, S. LOVE, and B. O’ NEIL Department of Biology, Dalhousie University, Halifax, Nova Scotia B3H 4J1 Freedman, B., S. Love, and B. O’Neil. 1996. Tree species composition, structure, and carbon storage in stands of urban forest of varying character in Halifax, Nova Scotia. Canadian Field-Naturalist 110(4): 675-682. Characteristics of stands of urban forest in Halifax, Nova Scotia, were analyzed with regards to tree-species composition and carbon sequestration. Older stands in residential and institutional neighbourhoods are dominated by non-native species of trees, thus limiting their usefulness in terms of protection of indigenous biodiversity values. These stands do, however, provide substantial ecosystem service in terms of carbon storage, because they are dominated by relatively large trees with an almost closed canopy (averaging 83% cover), and supporting a relatively large stand biomass (averaging 131 t/ha of above-ground tree biomass). In comparison, younger residential neighbourhoods have larger frequencies of native tree species, reflecting a recent trend to preserving some natural forest as buffer strips between properties. However, the propor- tion of native species of trees in the younger neighbourhoods is expected to decrease with time, because non-indigenous species are the strongly dominant choice among recent plantings by homeowners. Because of recent disturbance, urban for- est in younger neighbourhoods has a relatively small biomass, but this should increase with succession. Natural-forest urban parks have a tree-species composition and stand structure that are similar to that of more remote, natural forests. Key Words: urban forests, tree species, structure, carbon storage, Halifax, Nova Scotia. Urban forests consist of trees growing in areas where the dominant land use is urban or suburban, that is, in the vicinity of homes and commercial, institutional, and industrial buildings. Urban forests are important because they provide many important environmental benefits, the most notable of which are the following: (1) provision of habitat for urban wildlife, including some elements of indigenous bio- diversity, (2) improvement of urban aesthetics, (3) provision of out-of-doors shade and ameliorated microclimates, (4) energy savings through smaller demands for seasonal cooling and heating of the interior of buildings (trees cool buildings by provid- ing shade, and decrease heating costs by reducing wind speeds near buildings, thereby diminishing air- infiltration rates), (5) reduction of air pollution through direct uptake and physical filtering of gases and particulates, coupled with smaller emissions of pollutants associated with smaller energy demands for heating and cooling buildings, and (6) fixation of atmospheric carbon dioxide in the aggrading biomass of trees (Heisler 1986; Rowntree and Nowak 1991; McPherson 1994; Nowak 1994b; Nowak et al. 1994; Freedman and Keith 1995). Urban forests can develop from trees and stands that remain when a forested area is converted to resi- dential use, or they may develop from saplings plant- ed after homes or other buildings are constructed, or from plants that self-seed into the habitat. If the trees are planted, the species are commonly non-indige- nous to the local region, and even to the continent. Because of the non-indigenous nature of many of their species of trees, urban forests commonly sup- port markedly different biodiversity values than the more-natural forests that may occur as remnants within the urban environment, or more extensively in the surrounding area. The fixation of atmospheric CO, in urban forests is significant in view of the well-documented, increasing concentrations of this gas, from about 280 ppm in the mid-1800s to 355 ppm in 1995, with fur- ther increases projected into the future (Wigley and Raper 1992; Boden et al. 1993; Freedman 1995a). The increased concentrations of CO, and other radia- tively active gases (or RAGs; notably methane, nitrous oxide, ozone, and chlorofluorocarbons) have resulted from emissions associated with human activities, and may cause an intensification of Earth’s greenhouse effect (Marland and Rotty 1985; Ramanathan 1988; Anonymous 1995; Freedman 1995a). At least one-half of the potential enhance- ment of the greenhouse effect is estimated to be associated with increased concentrations of CO, (Schneider 1989; Lashof and Ahuja 1990; Rodhe 1990; Bolin 1991). This environmental change could cause an increase in Earth’s average surface temper- ature and other climatic changes, potentially result- ing in ecological disruptions of great consequence to natural biodiversity and to human socio-economic systems (Bolin et al. 1986; Schneider 1989a,b; Houghton et al. 1990; Wetherald 1991; Bolin 1991; Rosenzweig and Parry 1994; Freedman 1995a). One action that would contribute to mitigation of this potential problem is to offset some emissions of RAGs by increasing the rate of fixation and storage of atmospheric CO, by vegetation. The most sensible 675 676 option towards CO,-carbon sequestration is to increase the amount of organic-carbon stored in the biomass of trees, in both rural and urban forests (Marland 1988; Sedjo 1989a,b; Houghton 1990, 1991; Moulton and Richards 1990; Nagle 1990; Freedman et al. 1992; Nabuurs and Mohren 1993; Trexler and Haugen 1995; Freedman and Keith 1995). In addition to being a potential source of CO,-offset credits through fixation of this gas in their accumulating biomass, urban trees help to reduce CO, emissions by decreasing energy require- ments to cool and heat buildings (Nowak 1993). The purpose of the present study is twofold: (1) to examine variations of tree-species composition in an urban forest, and to interpret the patterns in terms of indigenous biodiversity values, and (2) to contribute to the information base required for evaluation of the potential role of urban trees in carbon sequestration. To these ends, this study has assessed the species composition of trees, stand structure, and quantities of carbon stored in trees in neighbourhoods of vari- ous character in the City of Halifax, Nova Scotia, including areas in which the predominant land-use is residential, institutional, or park. Methods 1. Study Area The City of Halifax is located on the Atlantic Coast of Nova Scotia, Canada, at about 44°39’N and 63°37’ W. Halifax is the largest city in Atlantic Canada, with a population of about 114-thousand in its 79 km? area, and 296-thousand in its 2.5-thousand km? metropolitan area, which includes Dartmouth, Bedford, Sackville, and built-up areas of Halifax County (McCann 1988). The local economy is heav- ily dependent on government services, financial institutions, the armed forces, the transportation industry, universities, and other regional services — there is relatively little manufacturing, fishing, or other primary industrial activities (McCann 1988). Halifax has a temperate maritime climatic regime, with a mean annual precipitation of 137 cm/yr, of which 86% arrives as rain and the remainder as snow (Anonymous 1993). The mean annual temperature is 6.5°C, mean daily temperature is 13.7°C during the growing season of May to October, and there are 4254 degree-days above 18°C each year (Anonymous 1993). Soils in the City of Halifax are primarily shaly loams developed from surficial tills of Precambrian slates, with some bedrock exposures, generally good drainage, and a rolling topography (MacDougall et al. 1963). The area is in the East Atlantic Shore sec- tion of the Acadian Forest region, in which the typi- cally dominant natural vegetation is forest dominated by Black Spruce (Picea mariana), White Spruce (P. glauca), and Balsam Fir (Abies balsamea). Red Maple (Acer rubrum), white and Yellow Birches THE CANADIAN FIELD-NATURALIST Vol. 110 (Betula papyrifera and B. lutea), Red Spruce (Picea rubens), and White Pine (Pinus strobus) are less prominent, although they can be locally abundant within this forest section (Rowe 1972). Within the City of Halifax, however, most of the natural forest has been cleared and converted into urban land-uses, including an urban forest dominated by non-indige- nous species of trees. 2. Study Sites Residential neighbourhoods in Halifax represent a continuum of ages and housing types. For the pur- poses of this study, representative study areas were chosen to illustrate the following classes of land-use: (1) Residential areas with homes less than 8 years old. There were three sampling sites in this cate- gory, located in a neighbourhood known as Clayton Park, where urbanization is relatively recent. The sample sites were named Warwick (sampled as a 1.3-ha plot), Turnmill (0.89 ha), and Canterbury (1.0 ha). The homes in Warwick and Turnmill were less than two years old, while those in Canterbury were less than eight years old. Residential areas with homes 40-50 years old. The three sampling sites were located in a neigh- bourhood known as Fairview. The sample sites were named Central (2.4 ha), Coronation (2.2 ha), and Birch (0.71 ha), and all contained homes that were 40-50 years old. (3) Residential areas with homes older than 70 years. There were three sampling sites, located in a neighbourhood known as the South End. The sample sites were named South (1.1 ha), Chestnut (0.51 ha), and Cedar (0.82 ha), and had homes that were 70 or more years old. Institutional. The three sampled sites consisted of a hospital complex and two universities, with their associated lands. The sample sites were parts of the campuses of Dalhousie University (8.2 ha), Saint Mary’s University (4.2 ha), and an area containing the Killam Children’s Hospital (2.3 ha). (5) Natural-area urban parks. Two natural-forest, urban parks were sampled: Fleming Park (45 ha) and Point Pleasant Park (75 ha). Both of these have second-growth, conifer-dominated forests that are essentially natural in character, although there has been some removal of dead trees, par- ticularly in Point Pleasant Park. In addition, data on tree-species composition were obtained for the Public Gardens, a horticultural urban park managed in the Victorian style (Stewart and Harvey 1982), and for several boulevards which are essentially managed as horticultural strips. (2 wa (4 wm 3. Measurement of Trees The fieldwork for this study was conducted during the growing season of 1995. All trees within the resi- 1996 dential and institutional sites were sampled (with permission of landowners, where private property was involved), including boulevard trees. All trees with a diameter (at breast height; DBH) greater than 5.0 cm were identified, and the DBH was measured. Because of the large areas of Point Pleasant and Fleming Parks, these were each sub-sampled using eight, widely spaced plots of 20m xX 20m. The field data were used to compute standard descriptors of forests, including basal area and density. The canopy areas of the entire plots were estimat- ed using aerial photographs with a scale of 1:10*. A grid of 2e:m X 2cm was mounted over the airphoto, and the numbers of intersections of grid points with the tree canopy was recorded, and converted to per- cent cover. 4. Calculations of Tree Biomass and Carbon Storage The above-ground biomass of trees was estimated using published regressions of DBH versus dry weight (Ker 1980; Freedman et al. 1982; Freedman 1984). Species-specific equations were used where available, as follows: Abies balsamea, Acer rubrum, A. saccharum, Betula lutea, B. papyrifera, Picea glauca, P. mariana, P. rubens, Populus tremuloides, and P. grandidentata (Freedman et al. 1984), and Fagus grandifolia, Larix laricina, Pinus banksiana, P. resinosa, P. strobus, and Tsuga canadensis (Ker 1980). For other species, biomass was estimated using generic equations for coniferous or angiosperm trees (Freedman 1984). Carbon content was estimat- ed to be one-half of dry weight (Nowak 1994b). Results and Discussion 1. Species Composition Species composition of the stands of urban forest in Halifax varied tremendously, from a relatively natural mixture of native species in Point Pleasant and Fleming Parks, to a more species-rich assem- blage of non-indigenous species in areas managed for horticultural purposes, such as the Public Gardens. The frequency of non-native species of trees was small in the natural-area parks, but much greater in other stands of urban forest, particularly in the oldest stands, such as the South End and the three sampled lengths of boulevard (Table 1). The most species-rich stand of urban forest is the Public Gardens, which is managed to include a diverse assemblage of horticultural plants, and con- tained 74 species of trees within an area of 4 ha. However, three non-indigenous species comprised 46.7% of the 604 trees in the Public Gardens: Linden (Tilia * europaea) 20.0%, Scotch Elm (Ulmus glabra) 20.0%, and Norway Maple (Acer pla- tanoides) 6.7%. Of the 604 trees in the Public Gardens, 91% are non-indigenous to North America, and only 3.6% are native to Nova Scotia. FREEDMAN, LOVE, AND O’ NEIL: TREE SPECIES OF URBAN FOREST 677 TABLE |. Frequency of species of trees non-native to North America in various neighbourhoods in the city of Halifax. The data for the residential areas are each weighted aver- ages of three sample plots, with the cumulative area indi- cated in parentheses. All trees in the horticultural park were surveyed, but the natural-area parks were each sub-sampled using eight quadrats of 0.04 ha. Percent Number Age of trees of trees Neighbourhood (years) non-native sampled 1. Residential Neighbourhoods Clayton Park (3.2 ha) 0-8 8% 791 Fairview (5.3 ha) 40-50 25 447 South End (2.4 ha) 70-80 iD. 187 2. Boulevard Strips Boulevard 1 (3.0 km) >70 93 S11 Boulevard 2 (3.0 km) >70 78 279 Boulevard 3 (5.5 km) >70 67 560 3. Horticultural Park Public Gardens (4 ha) mature 9] 604 4. Natural-area Urban Park Point Pleasant (75 ha) mature 0.2 476 Fleming (45 ha) mature 0.0 382 Among the three residential neighbourhoods, the relatively older South End had the greatest frequency of non-indigenous species, which accounted for 72% of the tree density. Among the 29 species of trees in the total sampled area of 2.4 ha, the most frequent were: Norway Maple (32.5% of density), American Elm (Ulmus americana; 12.6%), Linden (10.4%), and European Ash (Fraxinus excelsior, 7.4%). The intermediate-aged Fairview plots had 49 species in the sampled area of 5.3 ha, with 25% of the tree den- sity comprised of non-indigenous species. The most frequent species were Red Maple (34.4%), White Birch (11.5%), Norway Maple (5.6%), and European Mountain Ash (Sorbus aucuparia; 5.5%). The most- recently developed neighbourhood, Clayton Park, had 37 species in the 3.2 ha sampled area, with 8% of the density being non-indigenous species. The most frequent species were Red Maple (24.4%), Red Spruce (21.2%), Balsam Fir (14.9%), and White Birch (11.1%). The relatively large proportion of native species in the most-recently developed residential neighbour- hood (Clayton Park) results from the survival of many trees from the natural forest that previously occupied that area. In part, this results from relative- ly large lot sizes in these newly developed subdivi- sions, coupled with the relatively “natural” approach taken in much of the landscaping. This was intended to preserve many of the existing trees, thereby decreasing the costs of site preparation and post-con- struction landscaping, while also catering to the 678 desires of many new homeowners to have a well- treed lot. It is notable, however, that non-native species of trees remain the strongly dominant choice of homeowners in their horticultural plantings in this young neighbourhood, so the frequency of non- indigenous species will likely increase over time in this developing, urban forest. The 14.7 ha of institutional lands contained 36 species of trees, the most frequent of which were Red Oak (Quercus rubra; 12.8%), Red Maple (12.4%), American Elm (11.5%), and Norway Maple (11.5%), a species mixture that complements that found elsewhere in older parts of the city. Almost all of the species in the natural-area urban parks are native trees, the most frequent of which are Red Spruce (36.0%), Red Maple (19.7%), White Birch (11.0%), and White Pine (9.4%). These are also dominant species in much of the natural forests elsewhere in central Nova Scotia. The overall char- acter of the stands in the natural-area urban parks is of a mature, conifer-dominated mixedwood. This is in marked contrast to the character of the stands in residential and institutional neighbourhoods of Halifax, which are strongly dominated by non- indigenous species of angiosperm trees. Therefore, except for the natural-area parks, the urban forest of Halifax is dominated by a diverse assemblage of non-indigenous species of trees, rep- resenting a miscellany of horticultural selections from far-flung places having a temperate climate. It is typical of cities in North America (and elsewhere) that extensive areas of urban forest contain non- indigenous species of trees, shrubs, and ground veg- etation, and are often dominated by these alien species (Whitney and Adams 1980; Dorney et al. 1984; McBride and Froehlich 1984; Miller and Winer 1984; McBride and Jacobs 1986; Nowak 1994a). This fact is symptomatic of a larger element of the global biodiversity crisis — the ever-increas- ing homogenization of Earth’s biotas and ecosys- tems, characterized by huge increases in the rates by which certain species of plants, animals, and microorganisms are being distributed by humans beyond their natural ranges, either deliberately or accidentally (Bratton 1982; Harty 1986; Drake et al. 1989; Goudie 1990; Freedman 1995a). Introductions associated with horticulture are intended to achieve a culturally desired, out-of-doors aesthetic. This objec- tive is accomplished through the cultivation of a variety of non-native plants which mainstream horti- culturalists have come to know and prefer, have therefore become widely available commercially, and are commonly utilized in preference to local species, against which there is often a negative, hor- ticultural bias. Unfortunately, introductions of some alien species, including trees, can cause many indirect, ecological damages, such as deterioration or elimina- THE CANADIAN FIELD-NATURALIST Vol. 110 tion of the habitat of native species, accidental releases of pathogens and pests, and sometimes the establishment of self-maintaining populations of species that turn out to be aggressively invasive (Bratton 1982; Harty 1986; White et al. 1993; Freedman 1995b; Ruesink et al. 1995). To the extent that it is dominated by non-native species of trees, the urban forest of Halifax is likely to have limited value in providing habitat for indigenous elements of biodiversity. In view of this biodiversity-related consideration, it is encouraging that the tree-species composition of the youngest of the studied residential neighbour- hoods reflects an emerging trend to preserving boundary-strips of natural forest and individual native trees in new housing developments in Halifax. It is less heartening, however, to note that almost all new plantings in this young neighbourhood are of non-indigenous species, and that native species are exceedingly difficult to purchase in horticultural out- lets in the study area (as is common in most of North America). Nevertheless, the retention of some native vegetation in the newest residential neighbourhoods represents a substantial improvement over the fash- ion of urban-forest management in older areas of Halifax, where virtually all existing trees are of planted, non-indigenous species. 2. Stand Characteristics The three residential neighbourhoods can be con- sidered to represent a chronosequence of stands of increasing age since their conversion from natural forest into a housing-dominated land-use. Broader elements of the successional pattern can be inferred from structural attributes of the urban forests of these neighbourhoods (Table 2). The youngest neighbour- hood (Clayton Park, <8 years old), averaged a rela- tively large density (241 stems/ha) of smaller trees (average DBH range of 10-15 cm among major species), with a small canopy cover (13.9%), and a small above-ground tree biomass (16.9 tonnes/ha). The intermediate-aged neighbourhood (Fairview, 40- 50 years old) had a smaller tree density (158 stems/ha) but larger trees (range of average DBH of major species of 13-25 cm), a more extensive canopy cover (31.9%), and a greater stand biomass (18.8 t/ha). The oldest neighbourhood (South End, >70 years old) had the smallest tree density (145 stems/ha), the largest trees (range of average species DBHs of 30-62 cm), a relatively complete canopy cover (82.6%), and the largest stand biomass (131 t/ha) among the residential neighbourhoods that were studied. The three institutional plots are all located in older areas of the city of Halifax. Although the average tree density (50 stems/ha) and canopy cover (19.9%) were relatively small (because of the large buildings and extensive parking lots on institutional lands), the individual trees are relatively large (average range of 1996 FREEDMAN, LOVE, AND O’ NEIL: TREE SPECIES OF URBAN FOREST 679 TABLE 2. Characteristics of stands of urban forest in Halifax, Nova Scotia. Data are averages (+ standard error), for only the most prominent species of trees. Number Mean DBH Density Biomass Carbon Species Sampled (cm) (no./ha) (t/ha) (t/ha) 1. South End (>70-year-old residential neighbourhood; 3 plots, 2.4 ha) Acer platanoides 106 31.9 45.6 (8.1) 39.4 (9.0) 19.7 (4.5) Ulmus americana 41 41.6 17-2 2) 28.8 (10.9) 14.4 (5.4) Tilia x europaea 34 42.3 14.8 (3.7) 16.4 (4.2) 8.2 (2.1) Fraxinus excelsior 24 34.7 13.0 (9.2) 13.2 (10.9) 6.6 (5.5) Acer rubrum 22 30.4 10.0 (4.4) 6.7 (3.2) 3.3 (1.6) Quercus rubra 10 62.3 Slee) 15.6 (8.5) 7.8 (4.2) 23 other species 89 - 39.4 5.0 25 TOTAL 326 - 145.1 (39.7) 131.0 (43.9) 65.5 (21.9) 2. Fairview (40-50-year-old residential neighbourhood; 3 plots, 5.3 ha) Acer rubrum 289 18.3 47.9 (14.6) 8.0 (3.6) 4.0 (1.8) Betula papyrifera 97 sil PTAs!) 1.6 (0.3) 0.8 (0.2) Acer platanoides 47 24.5 Sen Ge) 3.0 (0.9) 1.5 (0.4) Sorbus aucuparia 46 12.8 1WRIE(/a10) 0.5 (0.2) 0.3 (0.1) 45 other species 361 - 60.2 al) 29 TOTAL 840 - 158.0 (20.7) 18.9 (5.3) 9.5 (2.6) 3. Clayton Park (<8-year-old residential neighbourhood; 3 plots, 3.2 ha) Acer rubrum 186 11.9 59.7 (6.4) 7.1 (0.3) 3.6 (0.2) Picea rubens 162 12.9 51.7 (9.3) 4.6 (1.6) 2.3 (0.8) Betula papyrifera 85 11.0 PATTIE (O}S)) 1.5 (0.8) 0.8 (0.4) Abies balsamea 114 10.2 3523) (i759) 1.2 (0.6) 0.6 (0.3) Pinus strobus 34 14.9 10.7 (4.9) 0.8 (0.3) 0.4 (0.2) 32 other species 172 - 55.8 ey 0.9 TOTAL 763 - 240.9 (31.2) 16.9 (2.1) Seed) 4. Institutional Neighbourhoods (3 plots, 14.9 ha) Quercus rubra 97 30.6 ANIL (5)) 3231253) Ie GIat8) Acer rubrum 94 27 4.6 (2.6) 1.6 (1.0) 0.8 (0.5) Acer platanoides 87 29.1 35) (1-4) 2.5 (1.3) L2i(OsM) Ulmus americana 87 28.3 10.1 (7.1) 6.9 (3.6) 3.5 (1.8) Acer pseudoplatanus 20 28.9 1.8 (1.1) eA (OM) 0.5 (0.3) 29 other species 351 - 22.8 3M) 1.9 TOTAL ST: - 50.2 (5.9) NS} (OAS)) 10.6 (1.2) ' 5. Natural-area Urban Park (Point Pleasant, 75 ha, 8 sub-plots of 0.04 ha) Picea rubens 259 17.3 809.4 (204.6) 106.2 (15.8) 53.1 (7.9) Picea snags 35) 11.8 109.4 (52.8) 4.1) (2:5) 2.1 (1.4) Picea glauca 27 18.3 84.4 (68.3) A GUL) 6.1 (5.6) Pinus strobus 26 31.4 81.3 (20.5) 26.9 (16.3) 13.5 (8.2) 9 other species Ws) - 234.3 27.4 1337 TOTAL 422 - 1318 (165) 176.8 (12.4) 88.4 (6.2) 6. Natural-area Urban Park (Fleming, 45 ha, 8 sub-plots of 0.04 ha) Acer rubrum 169 8.5 528.1 (87.7) 13.9 (1.9) 6.9 (1.0) Betula papyrifera 96 10.3 300.0 (91.1) 13.9 (3.9) 7.0 (1.9) Picea rubens W 18.0 234.4 (70.6) 34.5 (16.4) 17.3 (8.2) Pinus strobus 61 19.9 190.6 (65.1) 26.5 (8.9) 13.3 (4.5) Tsuga canadensis 13 21.4 40.6 (31.3) 5.7 (4.6) 2.9 (2.3) 12 other species 91 - 284.5 13.7 6.9 TOTAL 505 - 1578 (219) 109.5 (11.5) 54.8 (5.7) DBHs of major species was 22-50 cm), similar to those in the South End residential neighbourhood. The data for residential neighbourhoods in Halifax indicate a somewhat better-developed urban forest than has typically been reported in the United States, where the average urban area supports a tree density of only 52 stems/ha, and a canopy cover of 28% (Rowntree and Nowak 1991), comparable to the 680 institutional data for Halifax. The Halifax data are more similar, however, to cover values in northeast- ern U.S. cities, which average 24%-37% overall, and 46% in residential neighbourhoods (Rowntree 1984). The stands of forest in the two natural-area urban parks have values of tree density (average 1448 stems/ha), canopy cover (95.0%), basal area (34.3 m’/ha), and biomass (average 143 t/ha) that are with- in the range of mature forests in rural Nova Scotia. Freedman et al. (1986) studied four stands of natural, mature, spruce-dominated forest in central Nova Scotia using methods similar to those in the present study, and found a range of tree density of 1.71- 2.56 X 103 stems/ha (average 1.92 x 103 stems/ha), basal area of 28-45 m?/ha (average 37 m7/ha), and above-ground biomass of 112-233 t/ha (average 159 t/ha). However, the natural-area urban parks and rural forests differ in some other aspects of forest structure. In particular, the stands in Point Pleasant and Fleming Parks have relatively few snags (that is, dead but still-standing trees) and little coarse-woody debris, because these are regularly “cleaned” from the stands as part of routine park management. Overall, however, within the limitations of this rather coarse comparison, the stands in the natural-area urban parks in Halifax are structurally similar to natural conifer-dominated forests elsewhere in Nova Scotia. Although the tree density of the oldest residential neighbourhood (South End, 145 stems/ha) is consid- erably smaller than in the natural-area urban parks (1448 stems/ha), the biomass of these two stands of urban forest is similar (131 t/ha and 143 t/ha, respec- tively). This is because of the large sizes of average trees in the South End, compared with the natural- area stands, which have a heterogenous size spec- trum containing both smaller and larger trees. 3. Carbon Density in the Urban Forest The average carbon content of the above-ground tree biomass of the oldest residential neighbourhood (South End; 65.5 t/ha) is only slightly smaller than that of the mature stands of the two natural-area urban parks (Point Pleasant and Fleming; 71.6 t/ha). The younger residential neighbourhoods are in earlier suc- cessional stages, and have correspondingly smaller carbon densities in their tree biomass (Clayton Park, 8.5 t/ha; Fairview, 9.5 t/ha), as do the relatively sparsely treed institutional areas (10.6 t/ha). These data for Halifax compare favourably with data for urban forests in the United States, where average car- bon storage has been estimated to be about 27 t/ha (Rowntree and Nowak 1991). Somewhat smaller car- bon densities were found in a study of urban forests in the greater Chicago area, where trees store an average of 17 tC/ha in the city, and 22 t/ha in a nearby urban- izing area that still supports some natural tree cover (McPherson 1994; Nowak 1994b). THE CANADIAN FIELD-NATURALIST Vol. 110 For perspective, it is useful to note that the amount of carbon stored in trees in the mature residential neighbourhood (i.e., 65.5 t/ha; South End) is equiva- lent to the carbon content of approximately 104 x 10° litres of gasoline, 125 < 10% m3 of nat- ural gas, 71.2 < 10° kg of anthracite coal, and 81.9 x 10° kg of bituminous coal (these data assume the following carbon densities of fuels: gaso- line, 0.63 kgC/l; natural gas, 0.525 kgC/m?; anthracite coal, 0.92 kgC/kg; bituminous coal, 0.80 kgC/kg; Freedman and Keith 1995). It would be possible to increase the quantities of carbon stored in tree biomass in the urban forests of Halifax. Such a change might be desired as a compo- nent of an integrated strategy of reducing net emis- sions of radiatively active gases, by increasing forest biomass in rural and urban lands. In urban areas, this objective would best be approached by optimizing tree spacing and longevity to maximize carbon stor- age in the mature urban forest, while also attaining the many non-carbon benefits of increasing numbers of trees in urban aréas, as previously noted (Rowntree and Nowak 1991; McPherson 1994; Nowak 1993, 1994b; Nowak et al. 1994; Freedman and Keith 1995). In the United States, for example, it has been estimated that urban areas contain about 225-million tree-planting opportunities, in which sub-optimal tree densities could be subjected to fill- planting (Sampson et al., 1992). It has also been esti- mated that the planting of 10-million urban trees each year over a ten-year period in the U.S. would offset the emissions of 363 X 10° tonnes of CO,- carbon over the next 50 years, including 77 x 10°t by direct sequestration and 286 * 10° t through reduced energy requirements for cooling and heating buildings (Nowak 1993). There would, of course, be a need to accommodate potential problems associated with increased numbers of urban trees, such as obstructions of views and access, increased leaf and branch litter, interference with utility lines, shading of lawns and gardens, heaving of paving, obstruction of sewers and rain gutters, and seasonal emissions of pollen, to which some people have allergies. However, these draw- backs should be viewed as being much less substan- tial to society at large, in comparison to the many important benefits of the urban forest. Moreover, most problems with urban trees can be easily dealt with by sensibly planning their locations, and by planting appropriate species, particularly native ones. Acknowledgments This research was supported by a NSERC Individual Operating Grant to B.F., and a NSERC Summer Assistantship to B.O. We gratefully acknowledge the comments of M. Johnston and D. 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Received 12 January 1996 Accepted 13 September 1996 i} Malocclusion in the Jaws of Captive Bred Arctic Wolves, Canis lupus arctos N. E. FEDEROFF National Biological Service, Endangered Species Research Branch, Patuxent Wildlife Research Center, 11510 American Holly Drive, Laurel, Maryland 20708. Present address: Environmental Protection Agency, Mail Code 7507C, 401 M St. SW, Washington, D. C. 20460. Federoff, N. E. 1996. Malocclusion in the jaws of captive bred Arctic Wolves, Canis lupus arctos. Canadian Field- Naturalist 110(4): 683-687. Similar abnormalities in the skulls of captive Arctic Wolves (Canis lupus arctos) and a wild Arctic Wolf found dead on Ellesmere Island, Northwest Territories, in 1986 are described. The malocclusion is likely to be recessively inherited and would be expressed more frequently in association with increased levels of inbreeding. A re-shaping of the skulls may have occurred due to the effects of the malocclusive trait. The Ellesmere skull was short and wide in comparison to the captive skulls which were long and narrow. The focus of effect was in a foreshortening of the rostrum and the resulting shortened toothrow. Key Words: Arctic Wolf, Canis lupus, skull abnormalities, captive, wild, recessive, inbreeding, malocclusion, Ellesmere Island, Northwest Territories. Clutton-Brock et al. (1994) described an anom- alous (reported as a 24 year old adult male) Arctic Wolf (Canis lupus arctos) skull taken from one of two dead Wolves found on Ellesmere Island, Canada, in 1986. The skull was reported as exhibit- ing severe abnormalities of the jaws (malocclusion). The authors suggested possible explanations, including the hypothesis that the skull was from a wolf-dog hybrid. The authors later concluded that the Ellesmere skull showed no evidence of hybridization but instead was from a Wolf with severe jaw abnormalities. Mating is possible between Arctic Wolves and domestic Dogs [Canis familiaris| (Maagaard and Graugaard 1994). According to assertions made by Arctic natives, Soper (1940) wrote that crosses between Arctic Wolves and Eskimo Dogs appeared to be a fairly regular phenomenon. Although inter- breeding and production of fertile offspring between Wolves and Dogs is well documented (Young and Goldman 1944), gene flow has been mostly uni- directional, from Wolves into Dogs. Limited postu- lated evidence exists of introgression from Dogs into wild Wolf populations, particularly in Italy (Boitani 1984), although Randi et al. (1993) and Lorenzini and Fico (1995) have questioned introgression as a substantial threat in that country. However, inter- breeding, in itself, would not necessarily cause such reported abnormalities (Iljin 1941). I describe similar abnormalities for skulls of captive bred Arctic Wolves and suggest a possible cause for the abnor- mality reported by Clutton-Brock et al. (1994). In the United States, a line of Arctic Wolves has been bred in captivity since 1983 and all existing captive arctos from this line are directly descended from a founding pair obtained from the Stanley Park Zoo in Vancouver, British Columbia (S. Smith and F. W. Halvorson, USDA permit #42-B-059 Iowa and #41-A-301 Minnesota, personal communication, 1995). The founders were directly descended from a brother and sister pair taken from Axel Heiberg Island (5 km west of Ellesmere Island) in 1961, stud- ied and bred by Kuyt (1969, 1972), and donated to the zoo in 1968. No breeding data from the zoo, which may now be closed, could be obtained. In 1983 the founding pair produced a litter of two pups. A female from this breeding was then repeated- ly bred back to her father, producing several litters (Figure 1). The founding female had since died and no other Arctic Wolves could be found to facilitate an outcross. Most of the pups from the father/daughter cross were donated to other facilities, although a brother and sister had an accidental breeding, produc- ing four offspring, three (75%) of which exhibited malocclusion of the jaws (F. W. Halvorson, personal communication, 1995). Another facility, also in pos- session of a donated brother and sister pair from this line, also produced a litter of four offspring, two (50%) of which exhibited the malocclusion (F. W. Halvorson, personal communication, 1995). The pups from the accidental breeding were allowed to mature into adulthood and two females of the three wolves exhibiting the abnormality were then donated back to the original breeding facility where they were euthanized and the skulls removed. The malocclusion expressed itself most often when brother and sister matings from this line of arctos occurred. However, some of the offspring (10%) from the father/daughter cross also exhibited the abnormality. 683 684 THE CANADIAN FIELD-NATURALIST Vol. 110 P1 F2 F3 (expressed: slight) (FXexpressed) Skull (RH1) (FYexpressed) Skull (RH2) F (non-expressed carrier) @lexwresseq) M (non-expressed carrier) Founder Male (non-expressed) carrier ( ) Female (non-expressed) (FYexpressed: severe) Male (non-expressed) F (non-expressed carrier) ilies: (P\expressed) M (non-expressed carrier) @upexpressed) pup (non-expressed pup (non-expressed Alaskan Wolf Outcross Female (non-expressed) no pups exhibiting malocclusion (all should be carriers) M (non-expressed) (never bred Female (non-expressed) ( Re ( ) F (non-expressed) (never bred) Female (non-expressed) (carrier) F (non-expressed) Founder Female (non-expressed) M (DIED: non-expressed) M (non-expressed) F (non-expressed) M (non-expressed) M (non-expressed) F (non-expressed) F (non-expressed) M (non-expressed) F (DIED: non-expressed) M (non-expressed) M (non-expressed) 1/4 ee (carrier) Cpupyexpressed) F (non-expressed carrier) FicureE 1. Lineage of malocclusive captive Arctic Wolves (expressed and non-expressed). Degree of malocclusion varied from slight to severe, initially exhibiting itself in some wolves as young as 5 weeks of age and in others as old as 5 to 6 months. Malocclusive pups at other facilities have since been outcrossed with other Wolf subspecies. No offspring were produced exhibiting the abnor- mality (S. Smith, personal communication, 1995). However, when outcrossed Wolves (carriers) were bred back to one another, the malocclusion was evi- dent once again in the offspring (S. Smith, personal communication, 1995), indicating that the malocclu- sion may be the result of a recessive gene or poly- genic trait. The skulls removed from the two adult (approx. 2 years old) malocclusive female captive Arctic Wolves were sent to the author, cleaned, and mea- sured by the author and R. M. Nowak (USFWS, Office of Scientific Authority). Ten cranial mea- surements were taken as described by Nowak (1996). Both skulls exhibited overall wolf morphol- ogy (Table 1). Ratios (greatest length to zygomatic width and alveolar length of toothrow (P1 to M2) to maximum width across outer sides of crowns of P4) calculated from cranial measurements of a series of male (n=14) and female (n=7) arctos were com- pared with those of the male Ellesmere and the two female captive skulls. The captive skulls appeared to be longer and more narrow than the male Ellesmere skull. Ratios (greatest length to zygomat- ic width) from the captive skulls were greater than 2 standard deviations above the female series ratio mean. The ratio calculated from the male Ellesmere skull was exactly 2 standard deviations below the ratio mean of the male skull series. Ratios (alveolar length of toothrow to maximum width across outer sides of crowns of P4) from the captive skulls were just beyond 2 standard deviations below the female series ratio mean. Normally in Wolves, values for alveolar length of toothrow are slightly greater than values for maximum width across outer sides of crowns of P4, in which the opposite was true in regards to the abnormal skulls, although this condi- tion has also been observed occasionally in other specimens, especially in Wolves from the arctic (R. Nowak, USFWS, personal communication, 1996). The characteristics of the malocclusion were evident in the portion of the rostrum anterior to the fourth premolars. The facial regions of the skulls were foreshortened and undershot by the mandible. A mis-alignment of the upper third premolars was evi- dent as well (Figure 2). Buchalczyk et al. (1981) have also reported a bilateral skewness of P3 in 1996 FEDEROFF: MALOCCLUSION IN THE JAWS OF ARCTIC WOLVES 685 TABLE |. Skull measurements (mm) of two malocclusive captive adult female Arctic Wolves (RH1, RH2) and a series of female arctos (n=7). Skull Measurements (mm) Greatest length* Zygomatic width" Alveolar length of toothrow (P1 to M2)’ Maximum width across outer sides of crowns of P4° Maximum width across inner edges of alveoli of P1” Width of frontal shield” Height from alveolus of M1 to lowest point of orbit” Depth of jugal” Crown length of P4° Maximum crown width of M2* Greatest diameter of tympanic bullae Anteroposterior length of Cl Crown length of P3 Maximum crown width of M1 Alveolar length from p1 to m3 Mandibular depth taken between p3 and p4 Crown length of p4 Crown length of m1 *Measurements used by Nowak (1996). Skull Series‘ RH 1 RH 2 (mean) (SD) 237.0 237.0 244.40 + 6.45 121.0 125.0 ISSHfOP 2287 72.0 73.4 8426-2103 75.0 74.5 78.90 + 1.42 28.8 29.8 SiR OFiie = O'60 64.1 63.0 61.49 + 2.64 39.8 39.0 Ass ess alga) 15.1 16.1 18.41 + 0.87 24.9 25.0 26.37 + 0.64 12.6 15) 13.49 + 0.49 Biles 32.0 — — 13.4 13.5 — — 16.2 16.4 — — 21.0 21.5 — — 97.6 95.7 — — 24.6 24.5 = == Se; 525 — — 29.2 28.6 —_— — ‘Nowak, R. M. 1973. North American Quaternary Canis. Ph.D. dissertation (unpublished data). University of Kansas. skulls collected from Wolves in Poland, although the occurrence of malocclusion was not mentioned. The malocclusion was roughly 15 mm (anteroposte- FIGURE 2. (a) Upper: Left side view of a captive female Arctic Wolf skull (RH2) exhibiting the malocclu- sion. (b) Lower: Palatal view showing compacted third premolars in the captive skull. rior distance between upper and lower incisors). Each of the malocclusive skulls exhibited the same abnormalities and, after review of the abnormal skull ratios in relation to the series ratio means, it seems that a re-shaping of the skulls may have occurred due to the effects of the trait. Similar jaw abnormalities have also been observed in a series of skulls from captive Red Wolves (Canis rufus) at The Slater Museum, University of Puget Sound, Tacoma, Washington (USA) (R. M. Nowak, person- al communication, 1995). These observations may indicate that malocclu- sion in captive Arctic Wolves is probably associated with a recessive gene or a polygenic trait that may occasionally be expressed in the wild population. Due to the fact that the Wolf found on Ellesmere Island was an adult, it is apparent that malocclusive Wolves can survive into adulthood. Wolves with such abnormalities could probably kill Arctic Hare (Lepus arcticus) but probably not larger prey such as Musk Oxen (Ovibos moschatus), although the entire pack would share in consuming the kill, thereby allowing a malocclusive Wolf to survive (L. D. Mech, NBS, personal communication, 1995). Arctos is probably the least studied North American subspecies of Wolf. There are no data available regarding population density or genetics for this subspecies. Inbreeding may play a role in the expression of this abnormality, although more research is needed. If hybridization was the cause of the malocclusion in the Ellesmere Wolf, both par- ents, Wolf and Dog, would had to have been carriers 686 E + aise £ 3 Sin 30]; 2 = oO € 125 e RH2 S > e RH1 N 420} 115} 110 T T T T T T 225 230 235 240 245 250 255 260 Greatest length of skull (mm) FIGURE 3. Bivariate position comparison of skull measure- ments (greatest length and zygomatic width) in a series of female Canis lupus arctos (n=7) and two malocclusive captive Arctic Wolves (RHI, RH2). + = series mean. of the recessive trait. If this was the case, the gene(s) would nevertheless still have been evident in the wild population prior to the hybridization event. Whether the trait was originally brought into the Arctic Wolf population through hybridization or was always evident in the wild can only be speculated upon. Clutton-Brock et al. (1994) have suggested that recent changes in the skulls of Arctic Wolves, especially a reduction in size and change in shape, may be the result of interbreeding between Wolves and Dogs. However, Nowak (in press) has ques- tioned whether there have been any such changes. Should there indeed have been such changes, they may be due to subtle influences of the malocclusive 90 85 804 75 4 Alveolar length of tooth row (mm) 70 + ; r 1 70 75 80 85 90 Maximum width across outer sides of crowns of P4 (mm) FIGURE 4. Bivariate position comparison of skull measure- ments (Alveolar length of toothrow and maximum width across outer sides of crowns of P4) in a series of female arctos (n=7) and two malocclusive cap- tive Arctic Wolves (RH1,RH2). + = series mean. THE CANADIAN FIELD-NATURALIST Vol. 110 trait, which may be multi-factorial in relation to effects on cranial dimensions (Figure 3 and 4). Although all of the abnormalities in the skulls of the captive females apparently resulted from the maloc- clusive condition, the extent to which factors involy- ing captivity may have influenced the skulls is unknown. The malocclusive trait may be inherent in other Wolf populations as well (Vila et al. 1993), although it may express itself more frequently under captive breeding conditions where inbreeding may be a factor. Wolves are not immune to the effects of inbreed- ing depression. Lairke and Ryman (1991) suggest that deleterious homozygous alleles may be fairly common in natural Wolf populations. Results from a pedigree analysis using the SAS program INBREED (1996) imply that this abnormal condi- tion is genetically based and is most probably the result of a recessive gene or polygenic trait possibly expressed more frequently through increased levels of inbreeding in captivity. However, due to finding this condition in the wild Ellesmere Wolf, the abnormality itself was not the result of captive breeding, although inbreeding may increase the probability and frequency of expressing the abnor- mality. Inbreeding coefficients were fairly high for the Wolves expressing the abnormality. Coefficients for captive Arctic Wolves expressing the abnormali- ty ranged from 0.250 for animals resulting from the father/daughter cross to 0.375 for animals resulting from the brother/sister cross. Coefficients are very likely to be higher depending on the exact related- ness of the founders. The wild Ellesmere Wolf and the captive Wolves exhibiting the malocclusion originated from two separate but nearby islands. However, Arctic Wolves can freely travel across the ice between the islands, which would facilitate gene flow (L. D. Mech, personal communication, 1995). The maloc- clusive trait may still occur in parts of the Arctic Wolf population, due in part to gene flow between the island populations, and may be expressed more frequently in association with increased levels of inbreeding within the population. Further research is needed concerning population dynamics, long range movements to and from other Arctic islands, pat- terns of dispersal, geographic isolation from other Wolf subspecies, and levels of inbreeding and hybridizati n with Dogs, if any, in the Arctic Wolf population. Acknowledgments The author would like to thank F. W. Halvorson, S. Smith, C. Burkett, and C. Murphy for important captive lineage information without which this manuscript could not have been written; L. D. Mech and R. M. Nowak whose involvement and peer review greatly assisted the preparation of the manuscript; J. Kelham for captive care facilities; 1996 M. Holmes for typing the manuscript; K. Boone for excellent figure preparation; J. Eisemann for insightful comments during manuscript develop- ment; H.R. Perry Jr., I. Moore, and L. Lammons for administrative support; F. Kueppers, J. Eisemann, D. Pennock, R. Pierotti, D. W. Sparling, D. Hoffman, I. L. Brisbin, and G. Olsen for critical peer review of the manuscript. Literature Cited Boitani, L. 1984. Genetic considerations on wolf conser- vation in Italy. Bollettino di Zoologia 51: 367-373. Buchalezyk, T., J. Dynowski, and S. Szteyn. 1981. Variations in the number of teeth and asymmetry of the skull in the wolf. Acta Theriologica 26(2): 23-30. Clutton-Brock, J., A. C. Kitchener, and M. M. Lynch. 1994. Changes in the skull morphology of the Arctic wolf, Canis lupus arctos, during the twentieth century. Journal of Zoology, London 223: 19-36. Kuyt, E. 1969. Feeding ecology of wolves on Barren ground caribou range in the Northwest Territories. M.A. thesis. University of Saskatchewan, Saskatoon, Saskatchewan. Kuyt, E. 1972. Food habits of wolves on Barren ground caribou range. Canadian Wildlife Service Report Series: Number 21. Ijin, N. A. 1941. Wolf-dog genetics. Journal of Genetics 42: 359-414. Lorenzini, R., and R. Fico. 1995. A genetic investigation of enzyme polymorphisms shared by wolf and dog: FEDEROFF: MALOCCLUSION IN THE JAWS OF ARCTIC WOLVES 687 Suggestions for conservation of the wolf in Italy. Acta Theriologica 3: 101-110. Maagaard, L., and J. Graugaard. 1994. Female Arctic wolf, Canis lupus arctos, mating with domestic dogs, Canis familiaris, in Northeast Greenland. Canadian Field-Naturalist 108(3): 374-375. Nowak, R. M. 1996. Another look at wolf taxonomy. Pages 375-398 in Ecology and conservation of wolves in a changing world. Edited by L. N. Carbyn, S. H. Fritts, and D. R. Seip. Proceedings of the 2nd North American Wolf Symposium, August 24—27, 1992. Canadian Circumpolar Institute; University of Alberta, Edmonton, Canada. Nowak, R. M. Jn press. Hybridization: The double-edged threat. Canid News (Newsletter of the IUCN, Canid Specialist Group) Number 3. Randi, E., V. Lucchini, and F. Francisci. 1993. Allozyme variability in the Italian wolf (Canis lupus) population. Heredity 71: 516-522. SAS 1996. SAS Institute Inc., version 6.11, Cary, North Carolina. Soper, J. D. 1940. Eskimo dogs of the Canadian Arctic. Canadian Journal of Geography 20(2): 101-103. Vila, C., V. Urios, and J. Castroviejo. 1993. Tooth losses and anomalies in the wolf (Canis lupus). Canadian Journal of Zoology 71: 968-971. Young, S. P., and E. A. Goldman. 1944. The wolves of North America. American Wildlife Institute, Washington, D.C. 636 pages. Received 29 January 1996 Accepted 20 August 1996 Impacts of Habitat Fragmentation on Pairing Success of Male Ovenbirds, Seiurus aurocapillus, in Southern New Brunswick DWAYNE L. SABINE, ARNOLD H. BOER!, and WARREN B. BALLARD? New Brunswick Cooperative Fish and Wildlife Research Unit, University of New Brunswick, P. O. Box 44555, Fredericton, New Brunswick E3B 6C2 'Present address: Fish and Wildlife Branch, Department of Natural Resources and Energy, P. O. Box 6000, Fredericton, New Brunswick E3B 5H1. Present address: Arizona Game and Fish Department, 2221 West Greenway Road, Phoenix, Arizona 85032. Sabine, Dwayne L., Arnold H. Boer, and Warren B. Ballard. 1996. Impacts of habitat fragmentation on pairing success of male Ovenbirds, Seiurus aurocapillus, in southern New Brunswick. Canadian Field-Naturalist 110(4): 688-693. Populations of some species of nearctic-neotropical migrant birds have been declining in recent years, and much attention has focused on fragmentation of North American forests by agriculture, urbanization, and forest harvesting as one of the causes for the declines. This study evaluated the hypothesis that fragments of mature forest in a managed-forest landscape in southern New Brunswick were suboptimal habitat for Ovenbirds (Seiurus aurocapillus) compared to large tracts of simi- lar forest. There were no significant differences in pairing success or territorial density of male Ovenbirds between frag- ments and contiguous forest sites in 1992 and 1993. However, when only smallest fragments were considered mating suc- cess and density were lower but differences were not significant. Potential food abundance for Ovenbirds did not differ between fragment and large forest sites, nor between mated and unmated male territories. This study suggests that frag- mentation of mature forest in a largely forested landscape may not cause the adverse effects reported for Ovenbirds else- where. Key Words: Ovenbird, Seiurus aurocapillus, nearctic-neotropical migrant, habitat fragmentation, New Brunswick. Concern has been expressed that in recent years a number of nearctic-neotropical migrant songbird species, especially those breeding in forested habi- tats, have been declining (Aldrich and Robbins 1970; Briggs and Criswell 1979; Robbins 1979; Terborgh 1989; Hagan and Johnston 1992). Forests throughout the North American breeding range of these species are becoming increasingly modified by human activ- ities. Forest fragmentation is one such modification and, in recent years, impacts of forest fragmentation on nearctic-neotropical migrant birds have attracted attention. Several studies have suggested that some species tend to be intolerant of habitat area reduc- tion: 1.e., they are “area-sensitive” (Galli et al. 1976; Whitcomb et al. 1981; Robbins et al. 1989). Although some forest bird species appear to be area-sensitive, occurrence or density of birds may be an inappropriate measure for estimating minimum area requirements. Population density is not always a suitable indicator of habitat quality (Van Horne 1983; Vickery et al. 1992). Patch sizes corresponding to the area of minimum occurrence may be in the range of marginal rather than preferred habitat for area-sensi- tive species (Gibbs and Faaborg 1990). Studies by Ficken and Ficken (1966) Best (1977) and Probst and Hayes (1987) suggest that marginal habitats for some species were occupied by populations with low levels of reproductive success. These populations may be maintained by an influx of individuals from more productive habitats (Askins and Philbrick 1987; Askins et al. 1987; Pulliam 1988). Forest fragments may represent marginal habitat for area-sensitive birds for a variety of reasons, including increased rates of nest predation (Andren and Angelstam 1988; Wilcove 1985; Gates and Gysell 1978), brood para- sitism (Brittingham and Temple 1983), interspecific competition (Ambuel and Temple 1983; Whitcomb et al. 1981), decreased insect abundance and diversity (Whitcomb et al. 1981), and a reduced likelihood of maintaining critical microhabitats in small fragments for species which rely on a range of conditions (Wilcove et al. 1986). Forests in New Brunswick are currently undergo- ing alteration due to intensive forestry practices. This alteration differs from the type of fragmentation which has attracted most of the research effort to date, namely fragmentation of the eastern deciduous forest due to agriculture and urbanization. Instead of agricultural and/or urbanized areas, mature forest fragments in eastern Canada exist in a mosaic of regenerating forest, and tend to exhibit comparative- ly low isolation and high interconnectivity. Effects of habitat fragmentation on forest birds in this type of landscape are poorly understood. However, some species shown to be intolerant of fragmentation in the eastern United States are apparently unaffected by fragmentation of mature boreal forest through forest harvesting (Welsh 1987). Our study evaluated the hypothesis that fragments of mature forest in a managed-forest landscape rep- 688 1996 resent suboptimal habitat for Ovenbirds (Seiurus aurocapillus), as compared to large contiguous tracts of similar forest. Pairing success of male birds was chosen as an index of habitat quality, as resource quality is an important factor influencing mate selec- tion by females for those species in which the male provides a resource (e.g., territory) prior to mating (Searcy 1982). There is some evidence that females exhibit a greater sensitivity to habitat quality, insofar as it affects reproductive success. Females may be more likely to select territories based on features enhancing offspring production, such as nest site availability and quality, whereas males may be more likely to select territories based on features which enhance territory defense or mate attraction, such as song perch availability and quality (e.g., Zimmerman 1982; Sedgewick and Knopf 1992). To test the hypothesis, we predicted that pairing success of male Ovenbirds would be lower on mature forest fragments than on contiguous forest sites. Ovenbirds were selected for intensive study because they are a forest-interior species (Whitcomb et al. 1981; Robbins et al. 1989), and they were abundant in the mature tolerant hardwood forests of southern New Brunswick. Study Area The study was conducted in Albert and Saint John counties in southern New Brunswick, within and west of Fundy National Park (FNP) (45°65’N, 65°15’W). The park is a 207 km* contiguous tract of mature forest. The forest consists largely of toler- ant hardwood (Sugar Maple, Acer saccharum, and Yellow Birch, Betula allegheniensis, with a minor Red Spruce, Picea rubens, component) and mixed- wood stands (Red Spruce - tolerant hardwoods). The landscape surrounding FNP is largely forested and is subject to intensive forest harvesting, resulting in extensive fragmentation of mature forest. A large proportion of the coniferous and mixedwood stands have been harvested, and replaced with either natu- rally regenerating stands or plantations of Black Spruce (Picea mariana), White Spruce (Picea glau- ca), or Jack Pine (Pinus banksiana). Tolerant hard- wood stands have been subject to little or no harvest pressure. Criteria for selecting study sites were as follows: (1) presence of a mature (>40 yr), closed, tolerant hardwood canopy; (2) absence of well-developed subcanopy and shrub layers; (3) lack of recent distur- bance by logging in fragments; and (4) presence of a well-defined edge on all sides of fragments. The third criterion could not be attained for all fragments. Scattered Red Spruces were often selectively removed from fragments during recent harvests of surrounding stands. Other fragments appeared to have been selectively logged for Red Spruce at some time within < 30 years. As a result, most fragments contained a nearly pure hardwood canopy, while SABINE, BOER, AND BALLARD: OVENBIRDS IN NEW BRUNSWICK 689 FNP sites invariably contained a few scattered, mature Red Spruces. Although all fragment sites were enveloped by abrupt edges, several sites were surrounded by regenerating stands on only three sides, and separated from other mature forest tracts by road openings. We located 40 tolerant hardwood fragments between 1.0 ha and 51.5 ha in size (average = 11.1 ha) throughout a 25354 ha area west of FNP. Five forest fragments near FNP were selected for study in 1992, with two others added in 1993. Three plots within FNP were selected for study in 1992, with one more added in 1993. The seven fragments meet- ing our criteria ranged from 2.5 ha to 38.5 ha (mean = 12.6 ha), and six of these were only 2.5 ha to 16.3 ha in size (Table 1.). Methods To determine pairing status, male Ovenbirds were followed and closely observed for evidence of being mated. Observations commenced approximately one week after female birds were detected in breeding habitat, with all observations occurring between 3 June and 30 June. Males were classified as paired if they were observed interacting with a female or fledglings (Probst and Hayes 1987). The presence of a female responding to the male’s song with a “tsip- series” vocalization (Lein 1980) was also assumed to indicate a mated pair. Males were followed until such evidence of pairing was observed, or for a maximum of 90 minutes. We assumed that pairing status determinations would be valid within 90 min- utes, since Gibbs and Faaborg (1990) reported that curves describing the cumulative percentage of male ovenbirds classified as paired versus total elapsed time reached 100% at 67 minutes. Territories were marked for ease of relocation by flagging singing perches as males were followed. Food Availability Five 0.02 ha sample plots were located in each of 20 Ovenbird territories (10 in FNP, 10 in fragments) by randomly selecting compass coordinates and dis- tances from a point at the approximate center of each territory. Habitat variables were measured in these plots following the example of Smith and Shugart (1987). Distance to the nearest edge was also measured for each sample plot. Five 0.1 m< lit- ter samples were collected from each 0.02 ha plot. Invertebrates were extracted from these litter sam- ples by hand sorting and placed in a 70% ethyl alco- hol solution, and were later air dried at 50°C for 24 hours. Masses and numbers of individuals were recorded. Total mass of all invertebrates collected per sample was assumed to be a suitable index of potential prey availability, since most invertebrate groups are consumed by Ovenbirds in amounts pro- portional to their availability on the forest floor (Stenger 1958). 690 Statistical Tests Pairing success of males on fragments and FNP sites were compared using 2 x 2 contingency tables and the Fisher exact test (Zar 1984). Ovenbird densi- ties and invertebrate biomass were compared using the Student’s ¢-test (two-tailed). Probability levels < 0.05 were considered significant. Results The proportion of unmated male Ovenbirds was similar between fragment and FNP sites (Table 1). Pairing success of male Ovenbirds did not differ between years on fragment sites (P = 1.00), FNP sites (P = 1.00), or all sites combined (P = 0.71). Therefore, years were combined to compare pairing success between fragment and FNP sites. There was no difference between pairing success of male Ovenbirds on fragments (82%) versus FNP sites (84%) (P = 1.00). However, one large fragment (Number 7, 38.5 ha.) contained most (67-80%) of the male Ovenbirds observed on fragments (Table 1). When this large fragment was removed from the sample, the difference between pairing success of males on fragments (40%) and FNP sites (84%) was more pronounced (P = 0.07). Mean density of territorial male Ovenbirds on fragment sites (1.1/10 ha) was somewhat lower than on FNP sites (1.9/10 ha)(t = -1.35, P = 0.19). Lack of statistical difference is likely due to small sample size. Cumulative densities (total number of males on all sites divided by total area of all sites) were similar between fragment (2.0/10 ha) and FNP sites (2.0/10 ha) (t = -0.05, P = 0.96). When the largest THE CANADIAN FIELD-NATURALIST Vol. 110 fragment was removed from the sample, mean density was lower on fragment sites (0.7/10 ha) than on FNP sites (1.9/10 ha) (t = -2.39, P = 0.03), but cumulative density on the smallest fragment sites (1.0/10 ha), although lower, was not statistical- ly different from FNP sites (2.0/10 ha) (t = 1.14, Pi=\Oi3e)) Mean leaf-litter invertebrate biomass did not differ between Ovenbird territories on fragment sites (0.012 g/0.1 m*) and FNP sites (0.013 g/0.1 m’) (t= 0.14, P = 0.89). There was also no difference between mean invertebrate biomass on territories of mated males (0.012 g/0.1 m?) and unmated males (0.016 g/0.1 m7) (t = -0.79, P 0.43). Leaf-litter invertebrate biomass was significantly correlated only with one habitat variable (i.e., canopy cover) (Table 2). Discussion Reduced rates of pairing success of male Ovenbirds in forest fragments have been reported from New Jersey (Wander 1985), Missouri (Gibbs and Faaborg 1990; Van Horn et al. 1995), Pennsylvania (Porneluzi et al. 1993), Quebec- Ontario (Villard et al. 1993), and Maine (Hagen et al. 1996). Pairing success in these studies ranged from 19%-59% in fragments, and from 55%-85% in large forest plots. The high levels of pairing success of male Ovenbirds in the forests examined in this study (82% on fragments, 84% on FNP sites), and the lack of difference in pairing success between fragments and large, contiguous forest sites (FNP), contrast sharply with these studies. However, with the exception of Hagen et al. (1996), the other stud- TABLE 1. Number of territorial male Ovenbirds found on fragment and Fundy National Park sites during 1992 and 1993 in southern New Brunswick. Fragment Plot Distance to Percentage of Number of Site size size mature forest* regenerating forest” territorial males° (ha) (ha) (m) within 1-km radius 1992 1993 Fragment | DS DES 40 43.4 0 0 Fragment 2 5.6 5.6 20 61.3 0 0 Fragment 3 7.6 7.6 25 40.6 1P,1N 0 Fragment 4 8.7 5.9 40 95.2 IN IP, Fragment 5 9.3 9.3 200 65.0 ns? 0 Fragment 6 16.3 121 80 70.1 ns IN Fragment 7 38.5 27s 130 41.8 8P,1N 8P subtotal - 70.1 ~ - 9P,3N OP,1N Park 1 20700 14.9 0 0 4P,1N 3P Park 2 20700 16.6 0 0 3P,1N 0 Park 3 20700 22.8 0 0 5P,1N 3P,1N Park 4 20700 27.0 0 0 ns 3P,1N subtotal - 81.3 _ = 12B;3Nj 4 9PsUN “stand edge-to-edge distance to the closest mature forest block *clearcuts and plantations <15 years old © P=paired, N=non-paired dnot surveyed ( 1996 TABLE 2. Correlation coefficients between leaf-litter inver- tebrate biomass and habitat variables during June 1992 and 1993 in southern New Brunswick. Habitat variable r P ground cover -0.07 0.49 density of shrubs <1.5 m in height -0.01 0.96 density of shrubs >1.5 m in height -0.09 0.40 canopy cover 0.20 0.05 density of trees 3.0-6.9 cm dbh 0.04 0.69 density of trees 7.0-14.9 cm dbh 0.03 0.76 density of trees 15.0-22.9 cm dbh -0.02 0.87 density of trees >22.9 cm dbh -0.08 0.41 relative basal area of coniferous trees -0.16 0.12 distance to nearest opening 0.03 0.80 ies were conducted in areas where forest fragments were embedded in a mosaic of agricultural land. Hagen et al. (1996) examined an industrial forest landscape in north central Maine. They observed higher densities of male Ovenbirds in fragmented tracts, a reverse of the trend reported in the other studies. They suggest these higher densities are pos- sibly the result of a crowding effect following the loss of habitat surrounding the fragments. They pro- pose that behavioural dysfunction resulting from abnormally high densities may explain the lower pairing success in fragments in their study. We observed no crowding effect in our fragment sites, as densities were either similar to or lower than densi- ties in contiguous forest plots. However, our frag- ment sites were surrounded by young clearcuts or plantations ranging in age from 3 - 15 years old. It seems plausible that any crowding effect may have dissipated in the years following harvest of the mature stands. Our fragment sites were chosen to represent the extreme with regard to size in an effort to better identify potential impacts. Although our smaller fragments contained few Ovenbirds, density and pairing success for male Ovenbirds in the largest fragment (38.5 ha) were very similar to values recorded for the FNP plots. In contrast, several of the other studies reported lower densities and rates of pairing success in fragments much larger than our largest fragment. For example, Van Horn et al. (1995) recorded pairing success rates ranging from 0.0% to 27.7% in five fragments between 150 ha and 350 ha in size. Of the six smallest fragments studied by Porneluzi et al. (1993) (9.2 ha to 41.9 ha), only one plot (19.4 ha) contained territorial Ovenbirds. These results suggest that the threshold for negative impacts of fragmentation might occur at a much lower fragment size in forested landscapes than in the more modified agricultural landscapes. Although the absence of Ovenbirds from many smaller fragments resulted in a low mean density for SABINE, BOER, AND BALLARD: OVENBIRDS IN NEW BRUNSWICK 691 fragments, cumulative densities, which essentially weight each fragment or plot relative to its size, were almost identical between FNP and fragment sites. Study sites in FNP usually contained extensive sec- tions which were unoccupied by Ovenbirds but which appeared identical to those parts of the stands which were occupied. Hence, it is likely that if plots similar in size to our smaller fragments were ran- domly placed throughout the FNP study sites, many would have been found to be devoid of Ovenbirds. However, the absence of Ovenbirds on small frag- ments may have been due to a reduced likelihood of a bird finding and using a small patch, and not an inherent deficiency of habitat quality resulting from fragment size. It should be noted that pairing success is one of many factors contributing to overall reproductive out- put. Nesting success and fledgling survival might also be affected by habitat fragmentation. For exam- ple, nest predation and parasitism have been shown to be significant causes of reduced reproductive success for many bird species within forest fragments located within agricultural or suburban landscapes. Robinson (1992) reported nest predation rates of 80% and nest parasitism rates of 76% for neotropical migrants in forest fragments in Illinois. Nest parasitism would not be a factor in forested landscapes, because Brown-headed Cowbirds (Molothrus ater), the only obligate nest parasite in eastern North America, are extremely uncommon in these habitats. No Cowbirds were encountered during the two years of our study. However, nest predation was increased by fragmenta- tion of mature forest in forested landscapes in Maine (Small and Hunter 1988) and Pennsylvania (Yahner and Scott 1988). Unfortunately, our sample of nests was too small to estimate nest predation rates for our study area. Only five Ovenbird nests were located; of these one was abandoned shortly after it was found, one was destroyed by heavy rain, and three success- fully fledged four young each. Food availability is an important factor in deter- mining habitat quality. Ovenbirds feed mainly on leaf-litter invertebrates. It was conceivable that abundance of leaf-litter fauna in forest fragments might be affected by increased evaporation rates and decreased litter moisture rates caused by reduction of forest area (Lee 1978). However, we found no differ- ence in invertebrate biomass between fragment sites and FNP sites, or between territories of mated and unmated males. Smith and Shugart (1987) found that a variety of habitat features were important in pre- dicting prey abundance for Ovenbirds. We replicated their study, adding distance to the nearest large opening as a habitat variable. However, we found no correlation between invertebrate biomass and open- ing distance, nor between invertebrate biomass and most of the other habitat variables which Smith and Shugart (1987) found to be significant. 692 It appears that fragmentation of mature forest with- in a forested landscape may not cause the deleterious effects reported for Ovenbirds where forest fragmen- tation is caused by non-forest habitats. Alternatively, negative effects may occur but at a much lower threshold of fragment size. Unfortunately there is lit- tle information available concerning habitat fragmen- tation in largely forested landscapes (but see Rosenberg and Raphael 1986; Keller and Anderson 1992; Hagen et al. 1996). The current forest harvest- ing pattern in New Brunswick results in relatively large, irregular, interconnected blocks of unharvested, mature forest. Small, distinct, mature forest frag- ments are uncommon, and represent a small propor- tion of the available mature forest. If this pattern con- tinues, then fragmentation of mature forest as it occurs in New Brunswick may not be detrimental to all forest birds. Acknowledgments This project was funded by the Integrated Resource Management component of the Canada - New Brunswick Cooperation Agreement on Forest Development. We thank S. Makepeace, New Brunswick Department of Natural Resources, and Energy, and L. D. Morton, Canadian Forest Service, for their constructive comments and D. M. Keppie for insightful discussions. E. Sullivan and R. Otto assisted with fieldwork. Lodging was provided by the New Brunswick Department of Natural Resources and Energy. Additional assistance was provided by various people at Fundy National Park, particularly P. Deering and S. Flemming, and by J. Gilbert, J. D. Irving Ltd. Literature Cited Aldrich, J. W., and C. S. Robbins. 1970. Changing abun- dance of migratory birds in North America. Smithsonian Contributions in Zoology 26: 17-26. Ambuel, H., and S. A. Temple. 1983. Area-dependent changes in the bird communities and vegetation of southern Wisconsin forests. Ecology 64: 1057-1068. Andren, H., and P. Angelstam. 1988. Elevated predation rates as an edge effect in habitat islands: experimental evidence. Ecology 69: 544-547. Askins, R. A., and M. J. Philbrick. 1987. 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Population dynamics of breeding Neotropical migrants in a fragmented Illinois landscape. Pages 408-418 in Ecology and conservation of Neotropical landbird migrants. Edited by J. M. Hagan III and D. W. Johnson. Smithsonian Institution Press, Washington, D. C. Rosenberg, K. V., and M. G. Raphael. 1986. Effects of forest fragmentation on vertebrates in Douglas-fir forests. Pages 263-272 in Wildlife 2000: Modelling habitat relationships of terrestrial vertebrates. Edited by J. Verner, M. L. Morrison, and C. J. Ralph. University of Wisconsin Press, Madison, Wisconsin. 470 pages. Searcy, W. A. 1982. The evolutionary effects of mate selection. Annual Review of Ecology and Systematics 13: 57-85. Sedgewick, J. A., and F. L. Knopf. 1992. Describing wil- low flycatcher habitats: scale perspectives and gender differences. Condor.94: 720-733. Small, M. F., and M. L. Hunter. 1988. Forest fragmenta- tion and avian nest predation in forested landscapes. Oecologia 76: 62-64. Smith, T. M., and H. H. Shugart. 1987. Territory size variation in the ovenbird: the role of habitat structure. Ecology 68: 695-704. 1996 Stenger, J. 1958. Food habits and available food of Ovenbirds in relation to territory size. Auk 75: 125-140. Terborgh, J. 1989. Where have all the birds gone? Princeton University Press, Princeton, New Jersey. Van Horne, B. 1983. Density as a misleading indicator of habitat quality. Journal of Wildlife Management 47: 893-901. Van Horn, M. A., R. M. Gentry, and J. Faaborg. 1995. Patterns of Ovenbird (Seiurus aurocapillus) pairing suc- cess in Missouri forest tracts. Auk 112: 98-106. Vickery, P. D., M. L. Hunter, Jr., and J. V. Wells. 1992. Is density an indicator of breeding success? Auk 109: 706-710. Villard, M-A, P. R. Martin, and C. G. Drummond. 1993. Habitat fragmentation and pairing success in the ovenbird (Seiurus aurocapillus). Auk 110: 759-768. Wander, S. A. 1985. Comparative breeding biology of the ovenbird in large vs. fragmented forests: implica- tions for the conservation of neotropical migrant birds. Ph.D dissertation, Rutgers University, New Jersey. 153 pages. Welsh, D. A. 1987. The influence of forest harvesting on mixed coniferous-deciduous boreal bird communities in Ontario, Canada. Acta Oecologica/Oecologia Generalis 8: 247-252. SABINE, BOER, AND BALLARD: OVENBIRDS IN NEw BRUNSWICK 693 Whitcomb, R. F., C. S. Robbins, J. F. Lynch, B. L. Whitcomb, M. K. Klimkiewicz, and D. Bystrack. 1981. Effects of forest fragmentation and avifauna of eastern deciduous forest. Pages 175-205 in Forest island dynamics in man-dominated landscapes. Edited by R. L. Burgess and D. M. Sharpe. Springer-Verlag, New York. Wilcove, D. S. 1985. Nest predation in forest tracts and the decline of migratory songbirds. Ecology 66: 1211-1214. Wilcove, D. S., C. H. Mclellan, and A. P. Dobson. 1986. Habitat fragmentation in the temperate zone. Pages 237-256 in Conservation Biology: The Science of Scarcity and Diversity. Edited by M. E. Soulé. Sinauer Associates, Sunderland, Massachusetts. Yahner, R. H., and D. P. Scott. 1988. Effects of forest fragmentation on depredation of artificial nests. Journal of Wildlife Management 52: 158-161. Zar, J. 1984. Biostatistical analysis. Prentice Hall, New Jersey. 718 pages. Zimmerman, J. L. 1982. Nesting success of Dickcissels (Spiza americana) in preferred and less preferred habi- tats. Auk 99: 292-298. Received 8 March 1996 Accepted 30 September 1996 Renesting Intervals in Sprague’s Pipit, Anthus spragueii GLENN C. SUTTER, DAN J. SAWATZKY, DAWN M. Cooper, and R. MARK BRIGHAM Department of Biology, University of Regina, Regina, Saskatchewan S4S 0A2 Sutter, Glenn C., Dan J. Sawatzky, Dawn M. Cooper, and R. Mark Brigham. 1996. Renesting intervals in Sprague's Pipit, Anthus spragueii. Canadian Field-Naturalist 110(4): 694-697. Many bird species will attempt to raise more than one clutch in a given breeding season. For grassland songbirds, the inter- val between nesting attempts is usually measured indirectly because individual birds are difficult to track once a nest has failed or their chicks have fledged. Here we report observations of renesting by Sprague's Pipits (Anthus spragueii) carry- ing radio transmitters. As part of a larger study of pipit nesting ecology, we monitored activity patterns of radio-tagged adult females in 1994 and 1995. Three birds started replacement nests 10-15 days after losing a clutch, and one started a second nest 21 days after successfully fledging chicks. Such long intervals between nesting attempts support the impres- sion that the frequency of renesting in pipits is relatively low. Key Words: Sprague's Pipit, Anthus spragueii, renesting, grassland birds, radio-telemetry. Many bird species are able to increase their repro- ductive output by raising more than one brood per breeding season. Passerines breeding at temperate latitudes will frequently attempt more than one nest per season, with some grassland species averaging 1.5-2.0 clutches per female per year (Maher 1973; Martin 1995). For most grassland species, estimates of renesting (or double-brooding) frequency are based largely on indirect evidence, such as clutch initiation dates and the length of a species’ breeding season. The interval between nesting attempts is rarely known for grassland species because few marked populations have been studied and birds can be difficult to follow (Harris 1933; Maher 1973). Here we report observations of renesting by Sprague’s Pipit (Anthus spragueii) based on informa- tion collected via radio telemetry. Sprague’s Pipits are small (< 25 g), ground-nesting passerines that breed in temperate grasslands of North America (Owens and Myres 1973). They tolerate a wide range of grazing pressure on their breeding grounds, but tend to be less common in heavily grazed areas (Dale 1983; Owens and Myres 1973) and in introduced vegetation (Cody 1974; Wilson and Belcher 1989; Sutter 1996). Pipits arrive in southwestern Saskatchewan in late April or early May, and females lay clutches of 4-5 eggs beginning in mid May (Maher 1973). Based on clutch-initiation dates, breeding pairs produce an aver- age of 1.5 clutches per year and have mean incubation and nestling periods of 14 d and 11 d, respectively (Maher 1973). More information is required on the breeding biology of this species because pipit popula- tions appear to be declining with the loss of suitable breeding habitat (Knopf 1994). Methods From 13 May - 9 August 1994 and 9 May - 8 July 1995, we studied Sprague’s Pipits breeding in 256 ha of native mixed-grass prairie at the south end of the Matador Provincial Community Pasture (50°41'N, 107°44'W), 30 km SE of Kyle, Saskatchewan, Canada. The general area has a flat to rolling topog- raphy and is grazed by cattle annually. Native vege- tation on the study site is dominated by northern and western wheat grass (Agropyron dasystachyum and A. smithii, respectively), Junegrass (Koeleria gracilis) and Green Needle Grass (Stipa viridula). Other abundant plant species include Snowberry (Symphoricarpos occidentalis), sedges (Carex spp.), Pasture Sage (Artemisia frigida) and numerous forbs (Coupland et al. 1973). Plant names are taken from Looman (1982) and Vance et al. (1984). We located pipit nests by dragging a weighted 30- m rope over the study site and carefully searching wherever the rope caused birds to flush. We also found nests after encountering incubating birds by chance, and by tracking signals from birds carrying radio-transmitters. We marked each nest by center- ing the site along a 10-m line and flagging the vege- tation at both endpoints. Stakes or pin flags were not used because they tended to attract predators and cat- tle (personal observation). Nest initiation dates were estimated by candling eggs or back-dating from actual or estimated hatch dates, based on a mean incubation period of 14 d (Maher 1973; Sutter 1996). Active nests were checked every two days. To monitor incubation activity, we captured birds by flushing them off the nest into a mist net. Each bird was outfitted with a back-pack style, motion- sensitive radio transmitter (model BD-2 or BD-2G, Holohil Systems Ltd., Woodlawn, Ontario) by pass- ing the loop of an elastic thread harness over each wing (Brigham 1992). The mass of the transmitter and harness was 1.4-1.7 g, which represents 6-7% of a breeding female pipit’s body mass (23.1-25.0 g, Sadler and Maher 1974). Each transmitter had a flex- 694 1996 ible 17.5-19 cm antenna which extended beyond the bird’s tail by 2-3 cm. Females were distinguished from males by the presence of a brood patch and the absence of a cloacal protuberance. We monitored day-time activity patterns by track- ing signals from radio-tagged birds with either hand- held Merlin 24 (Custom Electronics, Urbana, Illinois) or automated Lotek SRX-400 (Lotek Engineering, Aurora, Ontario) receivers attached to 3- or 5-element yagi antennas. Birds that had either successfully fledged young or abandoned their nests were tracked to determine the interval between nest- ing attempts. Results Over the two years of this study, we found 51 pipit nests and put radio-transmitters on 32 birds. All but one of the birds we caught were clearly females, suggesting that females are primarily responsible for attending the nest in this species. The exceptional bird was within the size range of the others we caught (culmen = 9.2 - 12.7 mm; tarsus = 23.4 - 30.5 mm; right wing chord = 74 - 81 mm) and had a large brood patch, but it also had a small cloacal protuber- ance and may therefore have been a male. Disturbance effects, predation and technical prob- lems reduced the number of samples available for estimating the time interval between pipit nests. In 1994, 12 of the 19 birds we caught (63%) aban- doned their nests after being disturbed at the nest site and (or) because of irritation due to the trans- mitter. One of these birds died during inclement weather (heavy rain), another was obviously taken by a predator, and three removed their transmitters by cutting the elastic thread harness. Signals from the other seven disappeared either because the trans- mitter battery died or because the bird had left the study area or been captured by a predator. Two birds subsequently returned to the study area and started new nests 10-14 days after their signals had disappeared. Disturbance and transmitter effects were less problematic in 1995, presumably because of increased experience on our part. Only three of the 13 birds we caught that year abandoned their nests (one because of injury) and two were obvious- ly depredated. Overall, 15 birds (seven in 1994 and eight in 1995) accepted the transmitter and showed apparent- ly normal incubation behaviours. Two tended nests for more than two weeks before losing their trans- mitters, two lost their nests to predators, and seven raised chicks to fledging age. The other four birds were among nine that we followed beyond their first nesting attempt. Two of these birds were tracked to measure the time required for a bird to renest after successfully raising a clutch. One removed her trans- mitter eight days after her chicks had fledged, with- out having started a second nest. The other started a SUTTER, SAWATZKY, COOPER AND BRIGHAM: RENESTING IN SPRAGUE’S PIPIT 695 new nest 21 days after her first clutch fledged and successfully reared four more chicks. Thus, assum- ing this bird was behaving normally, the interval between successful pipit nests appeared to be rela- tively long. The other seven birds were tracked to measure the time required to replace failed nests. One of these birds lost her transmitter and three either left the study area or were depredated within 4-7 days of los- ing their clutch. No evidence of renesting was observed in each case. The other three birds built replacement nests 10-15 days after abandoning their first clutch, providing additional evidence that the interval between nesting attempts is relatively long in this species. Discussion Our results show that some Sprague’s Pipits attempt to raise more than one brood per year at Matador, requiring 10-15 days to replace lost clutches and up to three weeks to initiate a second clutch after nesting successfully. Our estimate of the time needed to replace a lost nest is substantially longer than the two days required by Horned Larks (Eremophila alpestris) (Beason and Franks 1974), implying that pipits require more time to secure additional copulations, choose a new nest site and build a new nest. The interval we observed between successful nests is also longer than the seven days reported for Horned Lark (Beason and Franks 1974) and the 1-8d reported for Baird’s Sparrow (Ammodramus bairdii) (Cartwright et al. 1937; S. Davis, personal communication), suggesting that pipits may extend their parental care well beyond the nestling stage, as reported in the congeneric Rock Pipit (A. spinoletta) (Askenmo and Unger 1986). Our estimates of pipit renesting intervals are based on the assumption that the radio-tagged birds we studied were behaving normally, which may be incorrect. Information based on radio-telemetry can be biased due to behavioural effects associated with the presence of the transmitter, especially when the transmitter is attached as a harness-style backpack (Sykes et al. 1990; Rappole and Tipton 1991). Waterfowl carrying harness transmitters will contin- ue to incubate, for example, but their reproductive behaviour can be altered to a point that affects nest initation patterns (Rotella et al. 1993; Pietz et al. 1993). Also, the transmitter and harness we used amounted to slightly more than 5% of the animal’s body mass, which has been suggested as an upper limit for radio-tracking studies of flying animals (Aldridge and Brigham 1988). That being said, we have several reasons to believe that telemetry and harness-style transmitters are valid techniques for studying incubation behaviour in this species. We rarely saw untagged female pipits in 696 flight, presumably because they spend much of their time tending the nest and foraging for ground-based prey (Harris 1933; Maher 1974). Also, 15 of 32 radio- tagged birds were able to carry out presumably nor- mal incubation behaviour, and those that retained their transmitters and avoided predators were able to con- struct second or replacement nests. Moreover, the intervals we detected support estimates of renesting based on the frequency distribution of clutch intiation dates, which indicates that female pipits may only produce 1.5 clutches per year (Maher 1973). There are also several advantages to gathering information on pipit incubation behaviour using radio telemetry. First, telemetry provides high-reso- lution data on the location and activity patterns of individual birds (see Sutter 1996). There is simply no other way to collect this information for birds such as pipits because they rarely fly and are virtual- ly impossible to observe directly as they forage in dense grassy vegetation (personal observations). Second, the use of radio-tags leaves no doubt as to the identity of an individual, or which of the parents is tending the nest. Again, this information is diffi- cult to obtain through direct observation because identifying leg bands are often hidden by the grass and the sexes of many grassland species are morpho- logically similar. Finally, the use of telemetry means there is no observer or blind to disrupt bird behav- iour or attract predators. The fact that we found renesting in pipits was not surprising. The breeding season for pipits at Matador can last 3 months, and individuals need only 25 days to raise a clutch to fledging (Maher 1973). Moreover, renesting has been reported in many other grassland species breeding at temperate latitudes (Beason and Franks 1974; Cartwright et al. 1937; DeSmet and Conrad 1991; With 1994) and in several congeners (Hendricks 1991; Askenmo and Unger 1986, and references therein). The frequency of renesting in pipits may be affect- ed by a combination of factors, including predation, energetic constraints and competition. The risk of predation can be extremely high for grassland birds (With 1994) with individuals breeding at Matador encountering nest predation rates as high as 69% (Maher 1973). The link between energy costs and renesting behaviour stems from the fact that birds generally need to invest large amounts of energy to breed at temperate latitudes, often directing resources away from their own survival and mainte- nance requirements (Martin 1987; but see Winkel and Winkel 1995). Finally, birds attempting to renest may have to compete for food supplies and nest sites, especially if breeding habitats are saturated (Wiens 1974, 1977). Our results confirm that Sprague's Pipits will attempt to maximize their sea- sonal reproductive output by renesting, despite the potential predation risk and energetic penalties. THE CANADIAN FIELD-NATURALIST Vol. 110 Acknowledgments Our sincere thanks to the many volunteers who helped with nest searches, and especially to T. Troupe and M. Pallin. Thanks also to the Saskatchewan Department of Agriculture and Food for allowing us to work on the Matador pasture, and to the University of Saskatchewan for the use of the Matador Grassland Field Station. Comments from J. W. Belcher, S. D. Grindal, R. G. Poulin, A. J. Erskine and an anonymous reviewer greatly improved earlier versions of the manuscript. This research was supported by the Natural Sciences and Engineering Research Council of Canada (NSERC) through a postgraduate scholarship to GCS and a research grant to RMB. Literature Cited Aldridge, H. D. J. N., and R. M. Brigham. 1988. Load carrying and maneuverability in an insectivorous bat: a test of the 5% “rule” of radio-telemetry. Journal of Mammalogy 69: 379-382. Askenmo, C., and U. Unger. 1986. How to be double- brooded: trends and timing of breeding performance in the Rock Pipit. Ornis Scandinavica 17: 237-244. Beason, R. C., and E. C. Franks. 1974. Breeding behav- ior of the Horned Lark. Auk 91: 65-74. Brigham, R. M. 1992. Daily torpor in a free ranging goat- sucker, the common poorwill (Phalaenoptilus nuttallii). Physiological Zoology 65: 457-472. Cartwright, B. W., T. M. Shortt, and R. D. Harris. 1937. Baird’s sparrow. Transactions of the Royal Canadian Institute 46: 153-198. Cody, M. L. 1974. Competition and the Structure of Bird Communities. Princeton University Press, Princeton, New Jersey. Coupland, R. T., E. A. Ripley, and P. C. Robins. 1973. Description of site: I. Floristic composition and canopy architecture of the vegetative canopy. Matador Project Technical Report Number 11. Canadian Committee for the International Biological Programme. The National Research Council and the University of Saskatchewan, Saskatoon, Saskatchewan. Dale, B. C. 1983. Habitat relationships of seven species of passerine birds at Last Mountain Lake, Saskatchewan. M.Sc. thesis, University of Regina, Regina, Saskatchewan. DeSmet, K. D., and M. P. Conrad. 1991. Management and research needs for Baird’s sparrow and other grass- land species in Manitoba. Pages 83-86 in Proceedings of the Second Endangered Species and Prairie Conservation Workshop, January 1989. Edited by G. L. Holroyd, G. Burns, and H. C. Smith. Provincial Museum of Alberta Natural History Occasional Paper Number 15. Harris, R. D. 1933. Observations on a nest of Sprague’s Pipit (Anthus spragueii). Canadian Field-Naturalist 47: 91-95. Hendricks, P. 1991. Site fidelity and renesting of female American Pipits. Journal of Field Ornithology 62: 338-342. Knopf, F. L. 1994. Avian assemblages on altered grass- lands. Studies in Avian Biology 15: 247-257. Looman, J. 1982. Prairie grasses identified and described by vegetative characters. Agriculture Canada publication 1413, Supply and Services Canada, Ottawa. 1996 Maher, W. J. 1973. Birds: I. Population dynamics. Technical Report Number 34 of the International Biological Programme Matador Project. University of Saskatchewan and the National Research Council of Canada, Saskatoon, Saskatchewan. Maher, W. J. 1974. Birds III. Food habits. Matador Project Technical Report Number 52. Canadian Committee for the International Biological Programme. The National Research Council and the University of Saskatchewan, Sasktoon, Saskatchewan. Martin, T. E. 1987. Food as a limit on breeding birds: a life-history perspective. Annual Review of Ecology and Systematics 18: 453-487. Martin, T. E. 1995. Avian life history evolution in rela- tion to nest sites, nest predation, and food. Ecological Monographs 65: 101-127. Owens, R. A., and M. T. Myres. 1973. Effects of agricul- ture upon populations of native passerine birds of an Alberta fescue grassland. Canadian Journal of Zoology 51: 697-713. Pietz, P. J.. G. L. Krapu, R. J. Greenwood, and J. T. Lokemoen. 1993. Effects of harness transmitters on behaviour and reproduction of wild mallards. Journal of Wildlife Management 57: 696-703. Rappole, J. H., and A. R. Tipton. 1991. New harness design for attachment of radio transmitters to small passerines. Journal of Field Ornithology 62: 335-337. Rotella, J. J.. D. W. Howerter, T. P. Sankowski, and J. H. Devries. 1993. Nesting effort by wild mallards with 3 types of radio transmitters. Journal of Wildlife Management 57: 690-695. Sadler, D. A. R., and W. J. Maher. 1974. Birds: IV. Bioenergetics and simulation of energy flow. Technical Report Number 63 of the International Biological SUTTER, SAWATZKY, COOPER AND BRIGHAM: RENESTING IN SPRAGUE’S PIPIT 697 Programme Matador Project. University of Saskatchewan and the National Research Council of Canada, Saskatoon, Saskatchewan. Sutter, G. C. 1996. Habitat selection and prairie drought in relation to grassland songbird community structure and the nesting ecology of Sprague’s Pipit, Anthus spragueii. Ph.D. thesis. University of Regina, Regina, Saskatchewan. Sykes, Jr., P. W., J. W. Carpenter, S. Holzman, and - P. H. Geissler. 1990. Evaluation of three miniature radio transmitter attachment methods for small passer- ines. Wildlife Society Bulletin 18: 41-48. Vance, F. R., J. R. Jowsey, and J. S. McLean. 1984. Wildflowers across the prairies. Western Producer Prairie Books, Saskatoon, Saskatchewan. Wiens, J. A. 1974. Climatic instability and the “ecologi- cal saturation” of bird communities in North American grasslands. Condor 76: 385-400. Wiens, J. A. 1977. On competition in variable environ- ments. American Scientist 65: 590-597. Wilson, S. D., and J. W. Belcher. 1989. Plant and bird communities of native prairie and introduced Eurasian vegetation in Manitoba, Canada. Conservation Biology 3: 39-44. Winkel, W., and D. Winkel. 1995. Costs and benefits of second broods in Coal Tits (Parus ater). Journal fiir Ornithologie 136: 29-36. With, K. A. 1994. The hazards of nesting near shrubs for a grassland bird, the McCown’s Longspur. Condor 96: 1009-1019. Received 3 June 1996 Accepted 27 September 1996 Notes First Nesting of the Razorbill, Alca torda, in the Wolves Archipelago, New Brunswick. KIMBERLEY MAWHINNEY!, and DAN SEARS? ‘Atlantic Cooperative Wildlife Ecology Research Network, University of New Brunswick, P.O. Box 45111, Fredericton, New Brunswick E3B 6E1 P.O. Box 1702, Sackville, New Brunswick EOA 3CO Mawhinney, Kimberley, and Dan Sears. 1996. First nesting of the Razorbill, Alca torda, in the Wolves Archipelago, New Brunswick. Canadian Field-Naturalist 110(4): 698-700. Two pairs of Razorbills, Alca torda, were observed nesting on South Wolf Island in the Wolves Archipelago, Bay of Fundy, New Brunswick, during 1995. One pair had nested and successfully hatched one young in 1994 as evidenced by a hatched membrane from the previous year. Both pairs nested and successfully fledged one young each in 1995. This is the first nesting record for this species in The Wolves Archipelago. Other long-standing breeding colonies of Razorbills known to exist in New Brunswick are at Machias Seal Island and Yellow Murr Ledge. Key Words: Razorbill, Alca torda, breeding, the Wolves Archipelago, Bay of Fundy. Razorbills, Alca torda, breed in mainly small colonies throughout Atlantic Canada, with the bulk of the North American population centred in south- ern Labrador (Nettleship 1980). Of the five small colonies known to exist in the Maritimes, two are in New Brunswick at the southern limit of its breeding range (Erskine 1992). These occur at Yellow Murr Ledge, south of Grand Manan Island, and Machias Seal Island. Razorbills were recorded as occurring in The Wolves Archipelago as early as 1979 (MacKay and Bosien 1979) and non-breeding Razorbills were regularly observed in the area during the summer months between 1986 and 1993 (Kehoe, personal communication). On 14 May 1995, five Razorbills were observed flying around the southernmost tip of South Wolf Island in The Wolves Archipelago (44° 58'N, 66°55'W). On 22 May, two nest sites were located on the south-facing granite cliff of South Wolf Island approximately 10 m above the high tide mark and immediately adjacent to a recently estab- lished Black-legged Kittiwake, Rissa tridactyla, colony (Kehoe 1994). One nest site had a single pale blue egg with red- dish brown markings laid on the bare rock. Dried droppings and other bits of vegetation from the immediate vicinity were placed directly beneath it. This site also contained a leathery shell membrane, likely from a successfully hatched egg laid at this site in 1994. Razorbill pairs generally keep the same breeding site from season to season (Nettleship and Birkhead 1985) and shell membranes are indicative of successfully hatched eggs (Girard 1939). The 698 Kittiwake colony on South Wolf was visited regu- larly during the summers of 1992 and 1993, and although Razorbills were observed in the area there was no evidence of nest sites (Kehoe, personal com- munication). This suggests that 1994 was the first year that breeding activity could have taken place. These Razorbills were regularly monitored throughout the 1995 breeding season. By 27 May both nest sites contained eggs and were being incu- bated. On 9 July one pair was observed with a fully feathered pre-fledged chick and the other pair was still incubating an egg. By 22 July one pair had suc- cessfully fledged a chick and the other pair was observed with a fully feathered pre-fledged chick. By 18 August both pairs had successfully fledged their one young. The North American Razorbill population was persecuted by humans through egging and hunting, and human occupation of many former nesting islands, up to the early 1900s. By this time its num- bers were substantially reduced throughout its range and it was locally close to extinction (Bent 1919). Since that time, with better protection of nesting birds and the regulation of hunting under the Migratory Birds Convention in 1916, populations began to increase. While Razorbills do breed in Nova Scotia, the Gulf of Maine colonies make up the major portion of the Razorbills at the southern edge of their range in the eastern Atlantic. A. R. Locke (1971 Census of seabirds nesting in Nova Scotia May 18 to June, 1971. Unpublished Reprint Canadian Wildlife Service, Ottawa) reported a long 1996 standing colony 51 pairs of Razorbills nesting in eastern Nova Scotia on colonies at Hertford/Ciboux Islands. Evidence of breeding at Pearl Island and at Margaree Island, in Nova Scotia was documented in 1971 and 1981, respectively (Erskine 1992). Of the over 3000 islands in the Gulf of Maine six were known historically to support Razorbills, but almost nothing is known of the size of these former colonies (Drury 1973-1974; Korschgen 1979). With the exception of The Wolves Archipelago, five of these islands in the Gulf of Maine are currently breeding sites for Razorbills. Only two of these islands have been reported as nesting colonies in southern Canada. Breeding records for the Razorbill colony on Machias Seal Island, in New Brunswick, date back to a single egg in 1886 (Squires 1945) and another single egg in 1922 (Pettingill 1939). However, Razorbills were not regularly observed on Machias Seal Island until 1937 (Pettingill 1939) and seventy- five pairs were reported in 1940 (Squires 1945). This population declined dramatically to 20 pairs in 1947 (Palmer 1949) and disappeared altogether in the late 1940s and early 1950s. Razorbills returned to Machias Seal Island by 1955 (Squires 1976), and they are presently estimated at 150-200 pairs (1995, _ K. Amey, personal communication). The long stand- ing razorbill colony at Yellow Murr Ledge in New Brunswick has remained relatively stable over the past 80 years. Squires (1945) reported a colony of 150 birds in 1924 and Pettingill (1939) found 200 in 1935. They are currently estimated at 130 breeding pairs (1993, S. Kress, personal communication). It seems likely that Razorbills formerly bred more widely on islands in the Gulf of Maine and any evi- dence of breeding today may be the suggestions of re-occupation of former sites. The first documentation of Razorbills, re-appear- ing in Maine was reported in 1952 on Matinicus Rock in outer Penobscot Bay (Drury 1973-1974) with most recent estimates reported at 40 nesting pairs in 1986 (Podolsky 1989). The second Maine site reported to be colonized was on Old Man Island. Drury (1973-1974) reported 40 individuals around the island in May of 1973 but was unable to confirm nesting. Korschgen (1979), however, esti- mated 10 nesting pairs on the island in 1974 and more recent estimates reported 26 breeding pairs in 1986 (Podolsky 1989). The third and most recent Razorbill colony established in Maine is located on Freeman Rock. Two nesting pairs were first report- ed in 1982; however, current numbers are not known (Podolsky 1989). Further prospecting by Razorbills has been observed at several former alcid colonies in Gulf of Maine (Drury 1973-1974), but gulls now occupy all of them, and gull interference may have inhibited or prevented recolonization (Nettleship 1972). NOTES 699 Whether or not Razorbills are re-establishing themselves in The Wolves Archipelago is unknown. This is the first nesting record for this species in The Wolves Archipelago, Bay of Fundy and is the third known breeding location for this species, near the southern limit of its range in New Brunswick. It adds to a short but growing list of recent occupation of breeding sites by seabirds in a region from which most species were extirpated a century ago. In recent years, new records for breeding birds in the Gulf of Maine/Bay of Fundy indicate that several seabird species may be extending their breeding ranges and/or recolonizing traditional breeding grounds. In 1992, 12 pairs of Black-legged Kittiwakes were discovered breeding in The Wolves Archipelago (Kehoe 1994) and have increased to more than 100 pairs in 1995 (personal observation). Common Murres (Uria aalge) and Great Cormorants (Phalacrocorax carbo) recently established them- selves as breeding birds on at least one site each, in the same general area, after nearly a century’s absence (Erskine 1992). Further monitoring of the presence and activities of Razorbills in The Wolves Archipelago should provide insight into the dynam- ics of range expansion and/or recolonization of this species in New Brunswick. Acknowledgments Personal communications of records were provid- ed by F.P. Kehoe, New Brunswick Department of Natural and Energy; S.W. Kress, National Audubon; and K. Amey, University of New Brunswick, Fredericton, New Brunswick. Special thanks to A. J. Erskine for his comments which helped to improve the text. Literature Cited Bent, A. C. 1919. Life histories of North American diving birds. U.S. National Museum Bulletin 107: 1-239. Drury, W. H. 1973-1974. Population changes in New England seabirds. Bird-Banding 44: 267-213; 45: 1-15. Erskine, A. J. 1992. Atlas of Breeding Birds of the Maritime Provinces. Nimbus Publishing Limited and Nova Scotia Museum, Halifax. 270 pages. Girard, G. L. 1939. Notes on the life history of the Shoveller. Transactions of North American Wildlife Conference 4: 364-371. Kehoe, F. P. 1994. A New Brunswick Black-legged Kittiwake, Rissa tridactyla, colony. Canadian Field- Naturalist 108(3): 375-376. Korschgen, C. E. 1979. Coastal waterbird colonies: Maine. U.S. Fish and Wildlife Service, Biological Service Program FWS/OBS-79/09. MacKay, A. R., and R. K. Bosien. 1979. Bay of Fundy Resource Inventory, Volume 5: The Wolves Islands. Final report to the New Brunswick Department of Fisheries, Fredericton, New Brunswick. Reference NB78-1B. Nettleship, D. N. 1972. Breeding success of the Common Puffin Fratercula arctica L. on the different habitats at Great Island, Newfoundland. Ecological Monographs 42: 236-268. 700 Nettleship, D. N. 1980. A guide to the major seabird colonies of eastern Canada: identity, distribution and abundance. Canadian Wildlife Service “Studies on Northern Seabirds” Manuscript Report Number 97: 1-133. Nettleship, D. N., and T. R. Birkhead. Editors. 1985. The Atlantic Alcidae. The evolution, distribution and biology of the auks inhabiting the Atlantic Ocean and adjacent water areas. London: Academic Press. 574 pages. (Including D. N. Nettleship, and P. G. H. Evans. Pages 53-154). Palmer, R.S. 1949. Maine Birds. Bulletin Museum Comparative Zoology 102: 1-636. THE CANADIAN FIELD-NATURALIST Vol. 110 Pettingill, O. S. Jr. 1939. The bird life of the Grand Manan Archipelago. Proceedings Nova Scotia Institute Science 19: 293-372. Pedolsky, R. H. 1989. The Razorbill in the Gulf of Maine. American Birds. 43(1): 14-16. Squires, W. A. 1945. The James S. Lord Collection of Birds’ Eggs. Acadian Naturalist 2: 67-80. Squires, W. A. 1976. The birds of New Brunswick. Second edition. The New Brunswick Museum, Saint John, Monograph Series. Number 7. Received 17 October 1995 Accepted 10 September 1996 Anomalies in the Eggs of Diving Ducks of the Genus Aythya MICHAEL A. FOURNIER and JAMES E. HINES Canadian Wildlife Service, P.O. Box 637, Yellowknife, Northwest Territories X1A 2N5 Fournier, Michael A., and James E. Hines. 1996. Anomalies in the eggs of diving ducks of the genus Aythya. Canadian Field-Naturalist 110(4): 700-701. We observed anomalies among the eggs of three species of diving ducks, including a shell-less egg outside the nest of a Canvasback (Aythya valisineria) and dwarf or runt eggs in the nests of Lesser and Greater scaups (Aythya affinis and A. marila). Shell-less eggs are rarely documented in wild birds. Runt eggs are more frequently documented, but we found no published record of their occurrence in scaups. Our observations indicate a frequency of occurrence of runt eggs in scaups of 0.10 %, similar to that reported for a number of other species. Key Words: Canvasback, Aythya valisineria, Greater Scaup, Aythya marila, Lesser Scaup, Aythya affinis, shell-less eggs, runt eggs, dwarf eggs. Impairment in the efficiency of the avian repro- ductive system may result in malformations of the egg (Romanoff and Romanoff 1949). Shell-less eggs, frequently produced by domestic birds (Romanoff and Romanoff 1949), are “little known” among wild species (Terres 1982). Abnormally small eggs, commonly referred to as dwarf or runt eggs, have been reported for a variety of species, both domestic and wild (e.g,. Romanoff and Romanoff 1949; Berger 1961; Rothstein 1973; Koenig 1980; Terres 1982; Mulvihill 1987) includ- ing at least one diving duck, the Bufflehead (Bucephala albeola) (Erskine 1971). Although runt eggs are not considered rare, there remains a need for reliable frequency data based on large samples of runt and normal eggs from wild populations (Mulvihill 1987). While conducting studies of breeding waterfowl in the Yellowknife area (62°27'N 114°22'W) and on the nearby North Arm of Great Slave Lake, Northwest Territories, we observed anomalies among the eggs of three species of diving ducks. On 2 June 1994, we observed a shell-less egg lying in the water next to the nest of a Canvasback (Aythya valisineria). The membrane had split at one end and the contents apparently leaked into the sur- rounding water. The membrane was leathery in appearance and had the texture of very fine sandpa- per on the outside. It also had a small (about 1 cm) curled and tapered projection at one end. The nest contained four normal eggs which the female was incubating. This seemed a small clutch, as mean clutch size for Canvasbacks in the Yellowknife area was 7.05+0.16 (S.E.) eggs (n=60, Fournier and Hines unpublished data). Shell-less eggs may be otherwise normal, but are often abnormal both in size and shape (Romanoff and Romanoff 1949). This shell-less egg, although devoid of contents and thus flaccid, appeared normal in both categories. Terres (1982) stated that shell- less eggs had been reported for the Traill’s Flycatcher (Empidonax traillii) and Common Grackle (Quiscalus quiscula). No further records were found in a review of literature for the period 1980-1994. Shell-less eggs may be more common than has been reported but, being very fragile, are likely most often destroyed before they can be observed (Romanoff and Romanoff 1949). Romanoff and Romanoff (1949) concluded that the cause of shell-less eggs was either a failure of the glands in the shell-secreting portion of the oviduct or premature laying due to violent peristalsis, rather 1996 than calcium starvation. However, the small clutch size in conjunction with the shell-less egg produced by this female could perhaps be considered indica- tive of nutrient (calcium?) limitation. On 21 June 1994, we discovered a dwarf egg in the nest of a Lesser Scaup (Aythya affinis), which also contained eight normal eggs (Case #1). The dwarf egg measured 28.0 X 24.4 mm, whereas the eight normal eggs averaged 58.5 X 41.0 mm. On 2 August 1994, we observed two dwarf eggs in the nest of a Greater Scaup (Aythya marila) (Case #2). There were no other eggs present in the nest. The dwarf eggs measured 37.5 < 28.6 mm and 38.1 X 28.4 mm. In comparison, Bent (1923) gave the average size of Greater Scaup eggs as 62.4 X 43.7 mm (n=180). On 28 June 1995, we observed one dwarf egg in the nest of a Greater Scaup which also contained seven normal eggs (Case #3). The dwarf egg mea- sured 44.7 X 31.8 mm, whereas the seven normal eggs averaged 63.2 X 43.9mm. This dwarf egg occurred on the same island as Case #2 and could have been produced by the same (philopatric) female. On this same date, we observed a single runt egg in an incomplete scaup nest on an island approx- imately 200 m distant (Case #4). This egg measured 39.9 X 29.3 mm. Given the dates of observation and presence of other normal-sized eggs, the dwarf eggs discovered in Cases #1 and #3 were undoubtedly products of first nesting attempts. The lack of any other eggs accompanying the dwarf eggs in Case #2, as well as the fact that the hen was still incubating in August, suggested that those eggs resulted from a renesting attempt. Case #4 may have been the product of either a first nest or a renest. With the exception of Case #4, all these runt eggs were being incubated at the time of discovery, as indicated by female behaviour, egg warmth, and amount of down in the nest. During the course of our study we examined 5169 scaup eggs. This value is not easily divided between the two species because; 1) there is overlap in the size range of the eggs of the two species, 2) many nests were located without the female being observed, and 3) there appears to be a relatively high rate of parasitism between the two species on our study area (Fournier and Hines, unpublished data). Therefore, the data were pooled to provide an overall frequency of occurrence for scaups. The five dwarf eggs represented a frequency of occurrence of 0.10%. This lies within the range of values observed in other field studies, as reported by Koenig (1980) (0.02 to 0.15, average 0.08 percent — omitting woodpeckers) with the exception of Manning and Carter’s (1977) cbservations for Canada Geese (Branta canadensis) (0.60 percent). Frequency of NOTES 701 occurrence in studies of museum specimens is higher (0.23 to 0.54, average 0.36 - omitting woodpeckers) (Koenig 1980), undoubtedly for reasons outlined by Rothstein (1973) (e.g., selective collecting). We could find no previous published record of the occur- rence of runt eggs in scaup. The cause of dwarf eggs was succinctly summa- rized by Romanoff and Romanoff (1949) as “the result of temporary disturbance or accident, rather than a permanent abnormality or disorder of the reproductive organs.” This explanation has been reiterated by others (e.g., Rothstein 1973; Koenig 1980) but, as Mulvihill (1987) pointed out, the caus- es of this phenomenon are not understood. Continued careful documentation of occurrences of dwarf eggs may eventually provide evidence for a more satisfactory explanation. Acknowledgments Librarian A. Welch of the Department of Renewable Resources, Government of the Northwest Territories, conducted the search of recent literature for information on shell-less eggs. Librarian T. Fraser of Environment Canada provided assistance in obtaining much of the literature reviewed for the production of this note. J. Roosdahl provided able field assistance. A. J. Erskine and an anonymous ref- eree provided comments on the draft manuscript. Literature Cited Bent, A. C. 1923. Life histories of North American wild- fowl. Part I. United States National Museum Bulletin 126. Reprint Edition. Dover Publications Inc. New York. 244 pages. Berger, A. J. 1961. Bird study. John Wiley and Sons Inc. New York. 389 pages. Erskine, A. J. 1971. Buffleheads. Monograph Series Number 4, Canadian Wildlife Service, Ottawa. 240 pages. Koenig, W. D. 1980. The incidence of runt eggs in wood- peckers. Wilson Bulletin 92: 169-176. Manning, T. H. and B. Carter. 1977. Incidence of runt eggs in the Canada Goose and Semipalmated Sandpiper. Wilson Bulletin 89: 469. Mulvihill, R.S. 1987. Runt eggs: a discovery, a synopsis and a proposal for future study. North American Bird Bander 12: 94-96. Romanoff, A. L. and A. J. Romanoff. 1949. The avian egg. John Wiley and Sons Inc. New York. 918 pages. Rothstein, S. I. 1973. The occurrence of unusually small eggs in three species of songbirds. Wilson Bulletin 85: 340-342. Terres, J. K. 1982. The Audubon Society encyclopedia of North American birds. Alfred A. Knopf Inc. New York. 1109 pages. Received 15 November 1995 Accepted 31 July 1996 702 THE CANADIAN FIELD-NATURALIST Vol. 110 Range Extension of the Hairy-tailed Mole, Parascalops breweri, in Northern Ontario STEPHEN J. HECNAR! and DARLENE R. HECNAR? 'Department of Biological Sciences, University of Windsor, Windsor, Ontario N9B 3P4 °5720 Oxley Avenue, LaSalle, Ontario N9H 1N3 Hecnar, Stephen J., and Darlene R. Hecnar. 1996. Range extension of the Hairy-tailed Mole, Parascalops breweri, in northern Ontario. Canadian Field-Naturalist 110(4): 702-703. In July 1995 we observed and photographed a Hairy-tailed Mole (Parascalops breweri) near Agawa Bay in Lake Superior Provincial Park in Ontario (47°22'N, 84°38'W). This represents a new record approximately 45 km north from the previ- ous peripheral location of Pancake Bay, Ontario (46°58’N, 84°42’W). Numerous tunnels of the species observed in the area suggested that populations were viable. Key Words: Hairy-tailed Mole, Parascalops breweri, distribution, range extension, Ontario. On 30 July 1995 we observed and photographed a Hairy-tailed Mole (Parascalops breweri) near the north end of Agawa Bay (47°22’N, 84°38’W) in Lake Superior Provincial Park (Figure 1). This sight- ing represents a new record of approximately 45 km north of the previous peripheral record of Pancake Bay, Ontario (46°58’N, 84°42’W; Anderson 1946; van Zyll de Jong 1983; Dobbyn 1994). The sighting also adds a new species to the mammal list of Lake Superior Provincial Park. The Hairy-tailed Mole is readily distinguished from other moles that occur in Ontario by its short, hairy tail and lack of protuberances on the snout (Dobbyn 1994). The specimen we observed was recently deceased and near the exit of its tunnel at the edge of an abandoned logging road in the Awausee Trail area (topographic map 41 N/7, UTM 783485). The specimen was a female, uniformly slate grey in color, and 15.8 cm in total length (tail length 2.5 cm). We observed no apparent injuries on the body. We did not collect the specimen because we were in the area for other purposes and did not have means of preservation on hand. However, we photographed the specimen and submitted prints to the Canadian Museum of Nature (D. Campbell) and the Royal Ontario Museum. The Hairy-tailed Mole occurs in the temperate Eastern Deciduous Forest of North America and ranges from Ontario and Québec southward into the mountains of Virginia and Tennessee (van Zyll de Jong 1983). It is now found mainly in second- growth hardwoods, edge habitats, and nearby mead- ows, where soils are light and moist, but well- drained (van Zyll de Jong 1983). The species is con- sidered to be generally uncommon (van Zyll de Jong 1983). There are very few records of this species north of Lake Huron and east of Lake Superior in Ontario (Dobbyn 1994). The dearth of records from the area may indicate either genuine absence or rari- ty, or it may reflect the lack of thorough surveys. Lake Superior Provincial Park has a modified continental climate, granitic bedrock, and rugged topography. The park is representative of the eco- tone between the Boreal Forest and the Great Lakes- St. Lawrence Forest (Rowe 1972). The specimen and tunnels of the species were found in areas domi- nated by hardwoods such as Sugar Maple (Acer sac- charum) and Yellow Birch (Betula alleghaniensis), 85° W 84° w 47° 30’ N AGAWA RIVER MONTREAL RIVER PANCAKE BAY LAKE /7 SUPERIOR ] 46° 30’ N FiGurE |. Study Area. Circle = Hairy-tailed Mole speci- men and tunnels observed, Triangle = area of numerous tunnels, Square = former peripheral pop- ulation. Solid line = Trans-Canada Highway, Crossed line = Algoma Central Railway, Dashed line = north wall of the Agawa Canyon, Dotted line = south boundary of Lake Superior Provincial Park. Shaded area indicates former species range within the study area (redrawn from van Zyll de Jong 1983). 1996 trees typical of the Great Lakes St. Lawrence Forest. The location where we found the mole consists of approximately 70-year-old second-growth hardwood on sandy loam and gravelly soils. We observed numerous earthworms and other soil invertebrates under woody debris in the area. We observed a large number of surface tunnels, all in upland forest and well-drained locations south of the north wall of the Agawa Canyon in the Awausee Trail area and around Crescent, MacGregor, Mud, and Kenny Lakes. Although Lake Superior Provincial Park was established in 1944, no previous records of Parascalops breweri for the park exist, and the species was not recorded in a small mammal survey conducted by the Royal Ontario Museum in the 1980s (Pasitschniak-Arts 1985). The only other mole in central Ontario is Condylura cristata, the Star- nosed Mole (van Zyll de Jong 1983). Although the ranges of both the Star-nosed Mole and Hairy-tailed Mole are broadly sympatric in Ontario, they would rarely be syntopic given the Star-nosed Mole’s pref- erence for wet habitats and that of the Hairy-tailed Mole for upland habitats (van Zyll de Jong 1983). NOTES 703 Acknowledgments We thank D. Campbell of the Canadian Museum of Nature, Ottawa for verifying the Pancake Bay peripheral record. R. M’Closkey made valuable comments on the manuscript. Funding was provided by NSERC to R. M’Closkey. Literature Cited Anderson, R. M. 1946. Catalogue of Canadian recent mammals. National Museum of Canada, Bulletin Number 102, Ottawa. 238 pages. Dobbyn, J.S. 1994. Atlas of the mammals of Ontario. Federation of Ontario Naturalists, Don Mills. 120 pages. Pasitschniak-Arts, M. 1985. Distribution, abundance and natural history of small mammals in Lake Superior Provincial Park: final summary report. Department of Mammalogy, Royal Ontario Museum, Toronto. 83 pages. Rowe, J.S. 1972. Forest regions of Canada. Environment Canada, Canadian Forestry Service, Publication number 1300. 172 pages. van Zyll de Jong, C. G. 1983. Handbook of Canadian mammals: | marsupials and insectivores. National Museums of Canada, Ottawa. 210 pages. Received 22 November 1995 Accepted 25 July 1996 Field Anesthesia of Striped Skunks, Mephitis mephitis, Using Halothane SERGE LARIVIERE and FRANCOIS MESSIER Department of Biology, University of Saskatchewan, 112 Science Place, Saskatoon, Saskatchewan S7N 5EZ Lariviere, Serge, and Francois Messier. 1996. Field anesthesia of Striped Skunks, Mephitis mephitis, using halothane. Canadian Field-Naturalist 110(4): 703-705. From April to August, 1993-1995, Striped Skunks (Mephitis mephitis) were captured in southcentral Saskatchewan, and immobilized using halothane. Eighty-one trials were performed on 47 adult skunks (11 males, 36 females) and seven juve- niles (4 males, 3 females). An initial dose of 8 mL (n = 78) provided complete, acceptable, and inadequate anesthesia in 51 (66%), 19 (24%), and 8 (10%) trials, respectively. First awakening behavior, stand-up posture, and recovery occurred after a mean (SD) of 52 s (34s), 71 s (28 s), and 87 s (15 s), respectively. Halothane provided rapid induction and recovery when used on Striped Skunks. Key Words: anesthesia, halothane, immobilization, Mephitis mephitis, Striped Skunk. Striped Skunks, Mephitis mephitis, have been anesthetized using injectable drugs such as sodium pentobarbital (Verts 1960), phencyclidine hydrochloride (Seal et al. 1970), a mixture of tile- tamine hydrochloride and zolazepam hydrochloride (Lariviere and Messier 1996), or ketamine hydrochloride (Rosatte and Hobson 1983). Most often, injectable anesthetics are administered by means of a pole syringe (Verts 1960) or following physical restraint (Crabb 1941). Solid-sided traps are now commercially available, and may be advanta- geous when working with skunks. Advantages may include reduction of scenting, reduction of stress lev- els in captured animals, and protection from weather. However, opaque traps do not permit the use of pole syringes (Verts 1960), and commercial traps do not possess a pushing barrier to force animals out of the trap for physical restraint (Crabb 1941). In contrast to injectable drugs, volatile anesthetics permit immo- bilization without physical contact, and can be used to immobilize animals captured in opaque traps. Propane gas has been used for the anesthesia of Striped Skunks. However, its safety and effective- ness has not been detailed (Rosatte 1987), and the 704 THE CANADIAN FIELD-NATURALIST Vol. 110 TABLE |. Levels of immobilization of Striped Skunks exposed to a single dose of 8 mL of halothane gas for various exposure times and trap types. Exposure time (min) Trap type Anesthesia 2 3 4 5 Total Homemade Complete 11 9 6 2 28 wooden box Acceptable 3 dl 0 13 Inadequate 0 0 4 0 4 Trials 14 12 17 2 45 Minnesota Complete 0 D 9 3 14 plasti-catch Acceptable 0 0 0 1 1 Inadequate 0 0 0 0 0 Trials 0 2 9 4 15 Tomahawk Complete 3 ps 2 2 9 wire-mesh Acceptable 2 0 Y 1 5 Inadequate 1 0 3 0 4 Trials 6 2 7 3 18 All traps Complete 14 13 ily 7 51 Acceptable 5 3 9 2 19 Inadequate 1 0 i @) 8 Trials 20 16 33 9 78 high inflammability of this gas makes it potentially dangerous to use. Halothane gas (M.T.C. Pharmaceutics, Cambridge, Ontario) has been used for the field immobilization of mammals such as Muskrats (Ondatra zibethicus; Blanchette 1989), weasels (Mustela erminea; Murphy and Dowding 1994), and American Martens (Martes americana; Herman et al. 1982). Its advantages include speed of induction and recovery, and ease in anesthesia proce- dures (Blanchette 1989). We tested the effectiveness of halothane gas for the chemical immobilization of Striped Skunks. From April to July, 1993-1995, Striped Skunks were live trapped in the Thickwood Hills, south- central Saskatchewan (52°N, 107°W). Skunks were captured in (1) a mesh-wire live trap (model 204, 19x 19x 51cm, Tomahawk Live Trap Co., Tomahawk, Wisconsin); (2) an opaque-sided plastic trap (23 X 23 X 60 cm, Minnesota Plasti-Catch, Mitlyng Development, Minnesota); or 3) a custom- built wooden live trap (19 X 18 X 71 cm). Traps were baited with canned sardines and checked daily, in the morning. A wood box (25 X 25 X 53 cm) was made to cover mesh-wire traps to provide a closed environ- ment for anesthesia. Because the edges of the box rested on the ground surrounding the trap during anesthesia, limited air penetrated the handling box and prevented an hypoxic condition. Wooden traps were ventilated through three 1-cm holes in the back, as well as a 0.5-cm space around the trap door. Similarly, ventilation holes in the door of the plastic trap allowed ventilation. All procedures were performed ouside to minimize exposure of humans to halothane (Short 1974: 17). Halothane (8 mL) was applied directly under the door of wooden and plastic traps. For wire traps, halothane was applied to cotton balls and placed inside the handling box. Animals were left undis- turbed for 2-4 min, after which their state of immobi- lization was qualified by their response to mechanical stimuli. State of anesthesia was classified as complete (no reaction to mechanical stimuli and no muscular resistance), acceptable (impaired movements and muscle resistance), or inadequate (no recumbancy, complete awareness, unimpaired movements). Anesthetized skunks were immediately removed from traps for handling. Inadequately anesthetized skunks were left in the trap for an additional 2-4 min, after which their state of anesthesia was reassessed. Failure to detect any sign of anesthesia after the sec- ond waiting period led to the second application of 8 mL of halothane. Handling procedures often began by an injection of Telazol (Lariviére and Messier 1996), or were limited to eartagging, sexing, and weighing. For most recap- tures, halothane anesthesia was sufficient. Recovery from halothane was evaluated only for skunks not injected with Telazol. Three characteristics of recovery were monitored: first awakening behavior (first non-stimulated head movements), stand-up posture, and complete recov- ery (unimpaired locomotion, gait normal). Recovery characteristics were calculated from the time of ces- sation of exposure to halothane. Excitation levels of captured skunks were evaluated before and after 1996 application of halothane, and classified as low (limit- ed movements in trap, no scenting) or high (panting, scenting, constant movements in trap). Eighty-one trials were performed on 11 adult males (mean body mass = 2.60, SD = 0.42), 36 adult females (mean body mass = 2.06 kg, SD = 0.31, n= 33; Lariviére and Messier 1996), and seven juve- niles (4 M, 3 F; mean body mass = 1.00 kg, SD = 0.36 kg, n = 6) in the three different trap types (Table 1). Of these, 71, 6, and 3 trials involved use of a single, double, and triple application of halothane, respectively. One trial required five applications. Seven of 20 trials in mesh-wire traps resulted in scenting during the covering of the trap, while only three of the 61 skunks captured in opaque traps scented. Similarly, skunks captured in wire traps were excited more often (11/20) than skunks cap- tured in opaque traps (15/61). Skunks were exposed to halothane between 2- 5 min. Initial applications of 8 mL of halothane lead to complete, acceptable, and inadequate immobiliza- tion in 51, 19, and 8 trials, respectively (Table 1). Fifty-eight trials were followed by an injection of Telazol (Lariviére and Messier 1996). Other trials (23/81) involved no use of injectable anesthetics. Following halothane anesthesia, skunks exhibited first awakening behavior quickly (mean = 52 s, SD = 34 s, n=17), whereas stand-up posture and complete recovery occurred after 71 s (n= 14, SD = 28 s) and 87 s (n= 16, SD = 15 s), respectively. Halothane applications seemed less effective in mesh-wire as compared to opaque traps. Additional stress is imposed on skunks captured in mesh-wire traps through the observation of nearby researchers, and scenting frequently occurs when covering wire- traps. Excited animals can develop life-threatening cardiac arrhythmias under halothane anesthesia (Sawyer 1982: 75), and excitement of skunks cap- tured in wire-traps often appeared high. Therefore, the use of opaque traps is more suitable for handling Striped Skunks. Halothane anesthesia on Striped Skunks was characterized by rapid induction and rapid recovery. The risk of overdose under uncontrolled field appli- cation of halothane presumably increases with pro- longed exposure. Maintenance of halothane anesthe- sia Over periods exceeding ~5 min requires a small concentration of anesthetic (1.5%; Sawyer 1982: 75) with the use of accurate and bulky delivery systems (Herman et al. 1982). Field conditions may not allow precise anesthesia equipment and procedures to be adopted. Nonetheless, we conclude that, regardless of sex or body mass, halothane is an ade- quate anesthetic for short (~1 min) handling proce- dures such as eartagging and recaptures. NOTES 705 Acknowledgments This study was financed mainly by the Prairie Habitat Joint Venture through a research grant from Ducks Unlimited (Institute for Wetland and Waterfowl Research), and the Canadian Wildlife Service. SL wishes to thank Fonds pour la Formation de Chercheurs et l'Aide 4 la Recherche (FCAR), Québec, for a postgraduate scholarship. Field assis- tance was provided by J. Bantle, B. Dinter, E. Howard, G. Poon, L. Walton, B. Fry, H. Royer, K. Eade, P. Toner, and S. Daviduik. L. Marinelli, J. A. Virgl, N. Caulkett, R. G. Clark, and R.C.N. Penner reviewed earlier drafts of this manuscript. Literature Cited Anonymous. 1991. Guidelines for the use of animals in research. Animal Behaviour 41: 183-186. Blanchette, P. 1989. Use of halothane to anaesthetize muskrats in the field. The Journal of Wildlife Management 53: 172-174. Boever, W. J., J. Holden, and K. K. Kane. 1977. Use of Telazol™ (CJ-744) for chemical restraint and anesthe- sia in wild and exotic carnivores. Veterinary Medecine/Small Animal Clinician 72: 1722-1725. Crabb, W. D. 1941. A technique for trapping and tagging spotted skunks. The Journal of Wildlife Management 5: 371-374. Herman, M. F., J. F. Pepper, and L. A. Herman. 1982. Field and laboratory techniques for anesthetizing marten with halothane. Wildlife Society Bulletin 10: 275-277. Jacobson, J. O., E. C. Meslow, aud M. F. Andrews. 1970. An improved technique for handling striped skunks in disease investigations. Journal of Wildlife Diseases 6: 510-512. Lariviere, S., and F. Messier. 1996. Immobilization of striped skunks using Telazol®. Wildlife Society Bulletin 24: in press. Murphy, E. C., and J. E. Dowding. 1994. Range and diet of stoats (Mustela erminea) in a New Zealand beech for- est. New Zealand Journal of Ecology 18: 11-18. Rosatte, R. C. 1987. Skunks. Pages 599-613 in Wild furbearer management and conservation in North America. Edited by M. Novak, J. A. Baker, M. E. Obbard, and B. Malloch. Ontario Ministry of Natural Resources, Toronto. 1150 pages. Rosatte, R. C., and D. P. Hobson. 1983. Ketamine hydrochloride as an immobilizing agent for striped skunks. Canadian Veterinary Journal 24: 134-135. Sawyer, D. C. 1982. The practice of small animal anes- thesia. Volume 1. W.B. Saunders Company, Philadelphia, Pennsylvania. Seal, U. S., A. W. Erickson, and J. G. Mayo. 1970. Drug immobilization of the Carnivora. International Zoo Yearbook 10: 157-170. Short, C. E. 1974. Clinical veterinary anesthesia: a guide for the practitioner. The C. V. Mosby Company, Saint Louis, Missouri. Verts, B. J. 1960. A device for anesthesizing skunks. The Journal of Wildlife Management 24: 335-336. Received 6 March 1996 Accepted 23 September 1996 706 THE CANADIAN FIELD-NATURALIST Vol. 110 European Beaver, Castor fiber, Pinned by a Felled Tree NILS B. KILE and FRANK ROSELL Telemark College, Department of Environmental Sciences, N-3800 Bg, Norway Kile, Nils B., and Frank Rosell. 1996. European Beaver, Castor fiber, pinned by a felled tree. Canadian Field-Naturalist 110(4): 706-707. The death of an adult European Beaver (Castor fiber) caused by a felled tree in Southeast Norway is reported. The trunk fell on the beaver’s tail pinning it to the ground. Key Words: European Beaver, Castor fiber, Southeast Norway, felled aspen, Populus tremula. On the morning of 21 November 1987 we found the remains of an adult European Beaver (Castor fiber) in Southeast Norway (58°39'N, 7°58'E). The animal apparently died after being trapped by a felled tree. It is uncertain whether this beaver, or another, actually felled the tree, though beaver reportedly most often fell trees alone (Wilsson 1971). Owesen (1979) reported that beaver run quickly to the side when a tree starts to fall. The felled aspen (Populus tremula) was about 40 cm in diameter at the base. The trunk had fallen across the Beaver’s tail about 10 cm from the tip and 2 m from the stump of the tree. In an attempt to escape, the beaver had scraped a circular channel to the front and sides of where it lay. The car- cass was frozen and still intact, suggesting that death had occurred recently. Beaver apparently cannot pre- dict where a tree will fall. Wilsson (1971) observed that trees fall in all directions, though more frequently towards shore simply because they often lean water- wards, or have better developed crowns on that side. North American Beaver (C. canadensis) have been reported trapped or killed by felled trees on four occasions (Table 1). Scotter and Scotter (1989) reported two, and Hitchcock (1954) reported one such death. These Beavers were dead when found. Ellarson and Hickey (1952) described the trapping TABLE |. A review of beaver (Castor canadensis and C. fiber) trapped or killed by trees. Source Locality Time of year Ellarson and Dane County-lowa October Hickey, 1952 County, USA Hinze, 1950 Revier GroBkiihnau, August Dessau, Germany Hinze, 1950 Germany ? Hinze, 1950 Germany ? Hitchcock, Bristol, Vermont, September 1954 USA Owesen, 1979 Norway ? Piechocki, Germany Between 1977 21 March — 7 June Scotter and Scotter, 1989 Sturgeon, Alberta, Canada ? Scotter and Edmonton, Alberta, October Scotter, 1989 Canada Stocker, 1978 Kanton Thurgau, August Schweiz This study Aust-Agder county, November Norway * Two separate individuals ** Later released *** Tater killed Part of body Age Species trapped Status Adult Castor Right hind Alive** canadensis foot i Castor fiber ? Dead ? Castor fiber ? Dead ? Castor fiber Right hind Alive foot Adult Castor Head Dead canadensis 2 Castor fiber Hind foot Alive*** Adults* Castor fiber u ? Adult Castor Across the Dead canadensis upper back Adult Castor Across the Dead canadensis shoulders x Castor fiber Right hind Dead foot Adult Castor fiber Across Dead the tail 1996 of a North American Beaver when the tree it had felled landed on one of its hind feet. This beaver was released unharmed. Four specific references to European Beaver being trapped or killed by trees have been reported previously. Hinze (1950) men- tioned three incidences of beaver trapped under felled trees, of which one survived. Piechocki (1977) mentioned two and Stocker (1978) one inci- dent. Owesen (1979) described how a beaver sur- vived being trapped when a felled tree pinned its hind foot to the ground. Most tree-felling by beavers occurs during autumn at the peak season for dam and den building, and six of eight animals were reported killed between August and November (Table 1). Only adult beaver were involved and only one (a male) had been sexed (Scotter and Scotter 1989). Felled trees were not considered as a significant mortality factor in North American Beaver (Hill 1982). What was once thought to be a freak event now appears to occur, although infrequently, with some regularity in Beaver populations in both Europe and North America. Acknowledgments We thank H. Parker for reviewing our manuscript. NOTES 707 Literature Cited Ellarson, R.S., and J. J. Hickey. 1952. Beaver trapped by tree. Journal of Mammalogy. 33: 482-483. Hill, E. P. 1982. Beaver (Castor canadensis). Pages 256-281 in Wild mammals of North America — Biology, management, and economics. Edited by J. A. Chapman and G.A. Feldhamer. John Hopkins University Press, Baltimore and London. Hinze, G. 1950. Der Biber. K6rperbau und Lebensweise, Verbreitung und Geschichte. — Akademie verlag, Berlin. [In German]. Hitchcock, H. B. 1954. Felled tree kills beaver (Castor canadensis). Journal of Mammology 35: 452. Owesen, A. 1979. I beverskog. Gyldendal Norsk Forlag Oslo. 120 pages. [In Norwegian] Piechocki, R. 1977. Okologische Todesursachenforschung am Elbebiber (Castor fiber albicus). Beitr. Jagd- und Wildforsch 10: 332-341. [In German]. Scotter, G. W., and E. Scotter. 1989. Beaver, Castor canadensis, mortality caused by felled trees in Alberta. Canadian Field-Naturalist 103: 400-401. Stocker, G. 1978. Tod eines Bibers, Castor fiber Linné, 1758, durch gefallten Baum. Saugetierk. Mitt. 26: 237-239. Wilsson, L. 1971. Observations and experiments on the ethology of the European beaver (Castor fiber L). Viltrevy 8: 115-266. Received 9 May 1996 Accepted 26 September 1996 Common Eider, Somateria mollissima, Incubates Gadwall, Anas strepera, Eggs: A Case of Clutch Adoption Due to Human Disturbance? DONALD F. MCALPINE Natural Science Department, New Brunswick Museum, 277 Douglas Avenue, Saint John, New Brunswick E2K 1E5 McAlpine, Donald F. 1996. Common Eider, Somateria mollissima, incubates Gadwall, Anas strepera, eggs: A case of clutch adoption due to human disturbance? Canadian Field-Naturalist 110(4): 707—708. A female Common Eider (Somateria mollissima) was observed on a hatching clutch of Gadwall (Anas strepera) eggs on an island in the Bay of Fundy. This aberrant behaviour may have been related to eider egg predation by gulls, rates of which had been increased by human activity on the island. Key Words: Common Eider, Somateria mollissima, disturbance, eco-tourism, Gadwall, Anas strepera, New Brunswick. Associations between nesting Common Eiders (Somateria mollissima) and gulls (Larus spp.) have been widely studied (Bourget 1973, Gotmark and Ahlund 1988, Swennen 1989 and references cited therein). Although Bellrose (1980) concluded that the larger gull species are the main predators of eider eggs and ducklings over most of the species’ range, Swennen (1989) noted that eiders have evolved a num- ber of behavioural adaptions that diminish the risk of gull predation on these life-history stages. Eider nest success has been reported to be higher within gull colonies than outside (Gotmark and Ahlund 1988). Perry (1982) documented cases in which Common Eider were observed taking over, and even defending from the original occupant, nests of both Herring (Larus argentatus) and Lesser Black-backed Gulls (L. fuscus). This nest stealing occurred prior to egg laying by gulls, but on one occasion a gull was observed to deposit an egg in a nest after being oust- ed, with the result that the eider incubated a mixed clutch. The outcome in this case was unknown. However, eiders have not been reported to incubate and eventually hatch the eggs of another species of bird, although eiders frequently nest in mixed-species 708 associations with larids and sometimes other ground- nesting waterfowl. Here, I document a case in which a female Common Eider appropriated the nest of a Gadwall (Anas strepera) containing a clutch of nine eggs. These eggs may have been incubated by the eider for a maximum of five days at the end of the incubation period, with the result that all eggs hatched and the ducklings were brooded by the eider. I suggest that this aberrant behaviour may have been the result of unintentional disturbance by naturalists visiting the island during a bird-banding programme. On 19 June 1986 I discovered a Gadwall nest con- taining 10 eggs on Manawagonish Island, New Brunswick (45°12' N 66°06’ N), an island of about 20h 1.5 km offshore in the lower Bay of Fundy. Principal species nesting on the island include more than 4000 Double-crested Cormorant (Phala- crocorax auritus), several hundred Herring and Great Black-backed Gulls (L. marinus) and a few Great Blue Herons (Ardea herodias) and Common Eiders (Astle and McAlpine 1985). The Gadwall nest was 5 m into a thicket of 1.5 m high Spiraea lat- ifolia in an abandoned field about 100 m from the water’s edge. No parent bird was on the nest when it was discovered. As few Gadwall nesting records in New Brunswick were known, a single egg was col- lected as documentation (McAlpine et. al. 1988; NBM 5994). The egg contained a well-developed embryo, and I expected to find the remaining eggs had hatched or were pipping when I returned to the nest on 23 June. At that time a female Common Eider was on the nest brooding seven ducklings while the last two struggled from the shell. The eider remained on the nest at my approach, and I watched her for about 1 minute. As I backed away, the bird left the nest, moved quickly through the undergrowth for about 30 m, and then took to the air. As the chances for survival of the young seemed diminished, and as I would not be returning to the island to monitor their fate, I retrieved the young. Three birds died within 72 hours; the remaining six were deposited at a local zoo, where one perished just prior to fledging, with the remaining five eventually released. The observation reported here is likely the result of egg predation by gulls after the incubating eider was flushed from it’s own nest by bird-banders; incubating female eiders are almost always on the nest during the day, although they may leave for short periods at dusk (Andersson 1975, cited in Goétmark 1989). Bird-banding activities for gulls and cormorants in the days prior to the discovery of the adoption of the Gadwall clutch were responsible for three known incidents of gull predation on eider clutches near the Gadwall nest. Paynter (1951) and Gilliland (1990) reported sig- nificant predation on eider eggs and ducklings by gulls on islands in the lower Bay of Fundy. Gilliland (1990) believed disturbance during field research on THE CANADIAN FIELD-NATURALIST Vol. 110 nesting eiders might be responsible. Dwernychuk and Boag (1972) attributed increased predation rates on duck eggs by gulls to disturbance by researchers. Likewise, Gétmark and Ahlund (1984) noted that disturbance by field workers led to much higher pre-. dation rates on clutches of eggs in eider colonies when eggs were left exposed. Swennen (1989) attrib- uted increasing eider nest success at a gull colony to reduced disturbance by researchers. It is clear that the presence of observers may increase predation on eider clutches, leading perhaps to unusual behaviour such as that noted here. In the past two decades there has been an increase in eco-tourism in the Bay of Fundy, with emphasis on whales and seabirds. Documenting incidents of aberrant behaviour of birds at nest sites, particularly when there are links to observer presence, may be useful in assessing the impact of possible eco-tourist activity on wildlife in the future. Literature Cited Andersson, A. 1975. The nesting ecology of the eider and the Velvet Scoter in the archipelago of Stockholm. A preliminary report. Statens Naturvardsverk PM 1009: 107-111. Astle, W. O., and D. F. McAlpine. 1985. Observations on the seabirds of Manawagonish Island, New Brunswick: Movements and population changes 1940- 1983. Proceedings of the Nova Scotia Institute of Science 35: 21-25. Bellrose, F.C. 1980. Ducks, geese and swans of North America. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. Bourget, A. A. 1973. Relation of eiders and gulls nesting in mixed colonies in Penobscot Bay, Maine. Auk 90: 809-820. Dwernychuk, L. W., and D. A. Boag. 1972. Ducks nest- ing in association with gulls - an ecological trap? Canadian Journal of Zoology 50: 559-563. Gilliland, S. G. 1990. Predator prey relationships between Great Black-Backed Gull and Common Eider popula- tions on the Wolves archipelago, New Brunswick: a study of foraging ecology. M.Sc. thesis, University of Western Ontario, Hamilton, Ontario. 96 pages. Gotmark, F., and M. Ahlund. 1984. Do field observers attract nest predators and influence nesting success of common eiders? Journal of Wildlife Management 48: 381-387. : Gétmark, F., and M. Ahlund. 1988. Nest predation and nest site selection among Eiders, Somateria mollissima: the influence of gulls. Ibis 130: 11-123. McAlpine, D. F., J. Finne, M. Phinney, S. Gilliland, and S. Makepeace. 1988. Breeding records for the Gadwall (Anas strepera) in New Brunswick. Le Naturaliste cana- dien 115: 95-96. Paynter, R. A. 1951. Clutch-size and egg mortality of Kent Island Eiders. Ecology 32: 497-507. Perry, P. 1982. The use of gull nests by Eiders. British Birds 75: 360-365. Swennen, C. 1989. Gull predation upon Eider, Somateria mollissima, ducklings: destruction or elimination of the unfit? Ardea 77: 21-45. Received 17 May 1996 Accepted 28 August 1996 1996 NOTES 709 Présence de la Puccinellie étroite, Puccinellia angustata, au Nunavik, Québec JACQUES CAYOUETTE! et MARCEL BLONDEAU? ‘Centre de recherches de l'Est sur les céréales et oléagineux, Programme de protection des cultures, Agriculture et Agroalimentaire Canada, Ferme expérimentale centrale, Edifice Wm Saunders, Ottawa, Ontario K1A 0C6 22400 chemin Ste-Foy, Sainte-Foy, Québec G1V 1T2 Cayouette, Jacques, et Marcel Blondeau. 1996. Présence de la Puccinellie étroite, Puccinellia angustata, au Nunavik, Québec. Canadian Field-Naturalist 110(4): 709-711. Aprés avoir révisé les récoltes du Québec précédemment identifiées a la Puccinellie étroite, Puccinellia angustata, les auteurs rapportent maintenant les seuls spécimens authentiques de cette espéce pour cette province. II s’agit de leurs récoltes faites 4 l’embouchure de la riviére Lacoudray (61°46'N — 72°35'W), dans la portion septentrionale du Nunavik. Un dénombrement chromosomique de 2n = 42 a été effectué sur un individu provenant de cette station. Cette localité marque une extension d’aire de plus de 500 kilométres vers le sud pour cette graminée du Haut-Arctique. Elle est comparée a P. deschampsioides et a P. macra a qui elle ressemble superficiellement. P. angustata devrait étre ajouté a la liste des plantes vasculaires susceptibles d'étre désignées menacées ou vulnérables au Québec. With the elimination of specimens from Quebec that had previously been identified as Narrow Alkali-Grass, Puccinellia angustata, the authors now report the only authentic specimens of this species from the province. These were collected by them at the mouth of Lacoudray River (61°46'N, 72°35'W) in the northernmost part of Nunavik. A chromosome number of 2n = 42 has been found on one specimen from that site. This locality represents a range extension of more than 500 kilome- ters to the south for this High-Arctic grass. It has been found to be distinct from the superficially related P. deschamp- sioides and P. macra. P. angustata should be added to the list of vascular plants likely to be listed as vulnerable or threat- ened in Quebec. Key Words: Narrow Alkali-Grass, Puccinellie étroite, Puccinellia angustata, P. deschampsioides, Nunavik, Québec, chro- mosome number, range extension. La Puccinellie étroite, Puccinellia angustata (R. Br.) Rand & Redf., est une Poacée circumpolaire du Haut-Arctique (Figure 1; Hultén 1968; Porsild et Cody 1980). Elle est mentionnée par Scoggan (1978) pour la Gaspésie et le nord du Québec. Selon cet auteur les mentions méridionales seraient des erreurs d'identification. Quant aux citations de Polunin (1940) et de Dutilly et Lepage (1951) pour le Nouveau-Québec, Scoggan précise que les spéci- mens de référence n’ont pas été vérifiés. Polunin (1940) inclut Puccinellia vaginata (Lange) Fern. & Weath. dans sa définition de P. angustata. Rouleau (in Marie-Victorin 1964) semble suivre cette opinion puisqu'il mentionne P. angustata pour le Québec et non pas P. vaginata. Toutes les récoltes de “P. angustata” sensu Polunin pour la région de la baie Wakeham que nous avons retracées ont été révisées a P. vaginata (Blondeau et Cayouette sous presse), mais le spécimen correspondant a la mention de “P. angustata” pour Port Burwell, au nord du Labrador, n’a pu étre retrouvé. Une mention pour cette derniére région apparait sur la carte de Porsild (1964) mais n’a pas été retenue par Porsild et Cody (1980). En consultant une carte inédite de Porsild (Cody com- munication personnelle), on retrouve une mention “Rousseau 1951” localisée un peu plus au sud sur la cote est de la baie d’Ungava, Nunavik. Le spécimen correspondant a été retracé (Ungava, poste de la riviere George, 9 aott 1951, J. Rousseau 1162, QUE) mais il a été révisé a P. laurentiana Fern. & Weath. par Sgrensen. Dutilly et Lepage (1951) citent P. angustata pour la région de la Pointe Louis-XIV, au sud de la baie d’ Hudson (ilot 4210 milles au nord- est du cap Jones: Dutilly et Lepage 12789); le spéci- men correspondant retracé a QFA a été révisé a P. pumila (Vasey) Hitchc. par Sgrensen. Dans le cadre d’une étude floristique au Nunavik, Nouveau-Québec, Blondeau et Cayouette (sous presse) ont exploré en 1994 le bras sud-est du havre Douglas; prés de 1’embouchure de la riviére Lacoudray, les auteurs ont récolté des individus qui correspondent bien a la description de P. angustata (Sgrensen 1953; Sgrensen in Hultén 1968), aux récoltes authentifiées par Sérensen dans les herbiers CAN et DAO et a la description d’individus de l’Arctique canadien (McLachlan et al. 1989) sauf pour la taille qui est un peu plus grande (> 30 cm). La citation complete de la récolte du Nunavik est la suivante: Québec, Nunavik, [havre Douglas, bras sud-est], riviére Lacoudray, rive est, environ 61°46'N — 72°35'W — UTM: 18VXD274514, berge argileuse, 6-VIII-1994. J. Cayouette et M. Blondeau C7914 (DAO), Blondeau et Cayouette HD94264 (C, QFA, Herb. Blondeau). En raison de leurs tiges dressées et de leurs pani- cules étroites, les individus du havre Douglas ressemblaient a P. deschampsioides T.J. Sérensen dont il existe des récoltes de la région de Salluit au Nunavik (Sugluk Inlet ou Anse Sugluk, 62°15'N — 75°28'W : Dutilly 6980v (QFA), 6980x (CAN, 710 THE CANADIAN FIELD-NATURALIST Vol. 110 FiGure |. Répartition nord-américaine de Puccinellia angustata. Sources: Cercles - Porsild et Cody (1980), Hultén (1968); triangle - nouvelle station. QFA), 6980y (CAN), Duman 2432 (CAN, QFA)); Sgrensen (1953) lui-méme citait la récolte de Dutilly 6980x qu'il avait authentifiée comme la seule men- tion de cette espéce connue a I'époque hors du Groenland. P. angustata differe de P. deschampsioides par les principaux caractéres suivants mesurés sur notre récolte du havre Douglas (P. angustata) et sur les récoltes de Salluit (P. deschampsioides) : longueur de la glume supérieure (angustata: 2.9-3.2 mm vs deschampsioides: 2.0-2.5 mm), longueur du lemma (3.6-4.0 mm vs 2.5-3.2 mm), nombre de fleurons par épillet (3-5 vs 4-8), pubescence a la base du lemma (abondante vs dispersée); de plus la présence de longs cils a la base des carénes du paléa caractérise P. angustata, alors que ces cils sont moins longs et moins abondants chez P. deschampsioides. Bent Fredskild du Grénlands Botaniske Unders@gelse de Copenhague (in litt. 1995) confirme que notre récolte du havre Douglas ne correspond pas 4a P. deschampsioides. Un dénombrement chromosomique effectué sur des mitoses racinaires provenant d’un individu de la popu- lation du havre Douglas (Cayouette C7914-1) a révélé 2n = 42 chromosomes, le seul nombre connu jusqu’a maintenant pour P. angustata (Love et Love 1975). Par contre, le nombre de 2n = 56 est le seul connu pour P. deschampsioides (Love et Love 1975). P. angustata ressemble sous certains aspects a l'espéce boréale nord-américaine P. macra Fern. & Weath., surtout par son inflorescence étroite, par le degré de pubescence a la base du lemma et par les longs cils a la portion inférieure des carénes du paléa. Il s’en distingue par les caractéres suivants mesurés sur P. angustata du havre Douglas et cer- taines récoltes de P. macra de la baie James (DAO): longueur de la panicule (angustata: 3-7 cm vs macra: 8-11 cm), longueur du lemma (3.6- 4.0mm vs 2.5-3.0 mm), nombre de fleurons par épillet (3-5 vs 5-6), couleur habituelle des épillets (violacée vs verdatre), longueur du caryopse (1.8- 2.0 mm vs 1.4-1.6 mm); de plus la nervure dorsale du lemma de P. macra porte fréquemment des aci- cules alors que celle de P. angustata est glabre. Il semble n'exister aucun dénombrement chromo- somique connu pour P. macra. A la riviére Lacoudray, P. angustata a été retracé a un seul endroit, soit un peu en amont de V’embouchure, a la base d’une berge argileuse, un habitat qui correspond a ce que rapportent McLachlan et al. (1989). Les panicules violacées, étroites et dressées de P. angustata contrastaient vivement avec les grosses touffes prostrées et glauques de P. vaginata qui croissait 4 proximite. Cette extension d’aire de P. angustata est remar- quable, du fait qu’il atteint rarement le sud du cercle 1996 polaire, notamment en Amérique du Nord; il se retrouve maintenant a plus de 500 kilométres au sud de la plus proche localité sur Vile de Baffin (Figure 1). Méme si a premiére vue la présence d’une espéce du Haut-Arctique peut sembler étonnante 4a la lati- tude du havre Douglas, elle ne représente pas le seul cas connu pour cette région. Des espéces du Haut- Arctique comme Deschampsia brevifolia R. Br. et Poa hartzii Gand., ainsi qu’ Erigeron compositus Pursh, une espéce arctique-alpine présente dans le Haut-Arctique, ont déja été signalées pour le bras sud-ouest du havre Douglas, 4 quelques kilométres du site de la riviére Lacoudray (Cayouette 1984). Puccinellia angustata n’apparait pas sur la liste des plantes vasculaires rares du Québec (Bouchard et al. 1983) ni sur la liste des plantes vasculaires sus- ceptibles d’étre désignées menacées ou vulnérables au Québec (Lavoie 1992). Comme il semble que notre récolte soit la premiere authentifiée au Québec, cette espéce devrait faire partie d’une prochaine ver- sion de la liste des plantes rares du Québec. Remerciements Merci a Sue Porebski et a Walter Wojtas (DAO) pour la confection de la carte de répartition, 4 Walter Wojtas pour son assistance technique au dénombre- ment chromosomique, a William J. Cody (DAO) pour le prét des cartes inédites de A. E. Porsild et ses commentaires sur le manuscrit, a Claude Roy (QFA) pour le prét de spécimens, a Bent Fredskild (C) pour son expertise sur les Puccinellia du Groenland, a André Ouellet d’Ammuumaajuq Adventure (Kangiqsujuaq) pour les facilités de transport et a Harry Okpik notre guide inuit de Kangiqsujuaq. Littérature citée Blondeau, M., et J. Cayouette. Sous presse. La flore vas- culaire de la baie Wakeham et du havre Douglas, détroit d’ Hudson (Nunavik, Québec). Provancheria. NOTES 711 Bouchard, A., D. Barabé, M. Dumais et S. Hay. 1983. Les plantes vasculaires rares du Québec. Syllogeus 48. Cayouette, J. 1984. Additions et extensions d’aire dans la flore vasculaire du Nouveau-Québec. Le Naturaliste canadien 111: 263-274. Dutilly, A., et E. Lepage. 1951. La traversée de |’ Ungava en 1945. II. Liste annotée des plantes vasculaires. Le Naturaliste canadien 78: 5—77. Hultén, E. 1968. Flora of Alaska and neigboring territo- ries. Stanford University Press, Stanford, California. Lavoie, G. 1992. Plantes vasculaires susceptibles d’étre désignées menacées ou vulnérables au Québec. Direction de la conservation et du patrimoine écologique, Ministére de l1’Environnement du Québec, QEN/SP00014. 180 pages. Léve, A., and D. Love. 1975. Cytotaxonomical atlas of the Arctic flora. J. Cramer, Vaduz. Marie-Victorin, F. 1964. Flore laurentienne. Deuxiéme édition. Révisée par E. Rouleau. Les Presses de l'Université de Montréal. McLachlan, K.I., S. G. Aiken, L. P. Lefkovtich, and S. A. Edlund. 1989. Grasses of the Queen Elizabeth Islands. Canadian Journal of Botany 67: 2088-2105. Polunin, N. 1940. Botany of the Canadian eastern arctic. Part 1, Pteridophyta and Spermatophyta. National Museum of Canada, Bulletin Number 92. Ottawa. Porsild, A. E. 1964. Illustrated flora of the Canadian Arctic archipelago (1957). National Museum of Canada, Bulletin Number 146. Ottawa. [Amended version with a supplement. ] Porsild, A. E., and W. J. Cody. 1980. Vascular plants of continental Northwest Territories, Canada. National Museum of Natural Sciences, Ottawa. Scoggan, H. J. 1978. The flora of Canada. Part 2, Pteridophyta, Gymnospermae, Monocotyledonae. National Museum of Natural Sciences Publications in Botany 7. Ottawa. Sorensen, Th. 1953. A revision of the Greenland species of Puccinellia Parl. Meddelelser om Grgnland 136(3): 1-179. Recu 6 février 1996 Accepté 1° octobre 1996 2 THE CANADIAN FIELD-NATURALIST Vol. 110 Red Squirrel, Tamiasciurus hudsonicus — Short-tailed Shrew, Blarina brevicauda, Brief Interaction ROBERT W. NERO Wildlife Branch, Box 24, 200 Saulteaux Crescent, Winnipeg, Manitoba R3J 3W3 Nero, Robert W. 1996. Red Squirrel, Tamiasciurus hudscnicus — Short-tailed Shrew, Blarina brevicauda, brief interac- tion. Canadian Field-Naturalist 110(4): 712. A Red Squirrel, Tamiasciurus hudsonicus, tussled briefly with a Short-tailed Shrew, Blarina brevicauda, beneath a bird feeder. Key Words: Manitoba, Red Squirrel, Tamiasciurus hudsonicus, Short-tailed Shrew, Blarina brevicauda. In the winter of 1995-1996, Red Squirrels fre- quently visited bird feeders at my Winnipeg home, one individual in particular aggressively driving away both other Red Squirrels and the much larger Grey Squirrels (Sciurus carolinensis). By November 1995, only two Red Squirrels were present, one at each of our two main feeders. Unexpectedly, from the first snowfall of October 1995 up to the end of January 1996, Short-tailed Shrews were also visible. From our living room window, which faces our well wooded backyard, my wife and I watched shrews come out onto the snow regularly at five different sites. These places were far enough apart to suggest that different individuals were involved. On a few occasions, we saw two shrews out at the same time. The fondness of Short-tailed Shrews for sunflower and other seeds has been pointed out earlier (Nero 1995). One shrew foraged on mixed birdseed beneath a feeder attached to a large spruce in front of our window. The tree lacks branches for several feet upwards, so we had a good view. Initially, I had scattered birdseed on the ground near the base of the tree to attract migrant sparrows and other birds; then, after it snowed, I cleared an area to expose the seed. The shrew, attracted by the seeds, kept coming out from burrows in the snow to feed, even in the day- time. I saw it as early as 05:00 h and as late as 22:00 h. Almost daily, especially beginning at about 15:00-16:00 h, it would come out into the open, feed briefly, then scurry back out of sight. Sometimes, it repeated this pattern for half an hour. I couldn’t tell if it was just eating or taking seeds away to store, but its occasional appearance seconds later some dis- tance away suggested that it was storing seeds. Occasionally, the shrew would crouch and feed for several minutes. On 9 December 1995, at about 13:45 h, a sunny day with a temperature at -24°C, my wife saw a Red Squirrel tussling with the shrew in the food patch. The two scrambled about for a few seconds, then the shrew ran off over the snow, pursued by the squirrel which caught up with the shrew about 2 feet (ca 61 cm) away. Again there was a struggle, then, sur- prisingly, considering their size difference, the shrew once again escaped, darting down one of its several burrow openings. The squirrel then climbed back into the spruce tree. Apparently, no injury resulted to the shrew (or the squirrel!), for a day later we saw the shrew come out to feed as usual. The carnivorous tendencies of the Red Squirrel, along with its habit of defending a food source, have previously been noted (Nero 1987, 1993; Taylor 1988; Sullivan 1991). Had the Red Squirrel been able to kill the shrew, if that was even its intention, it might have eaten it. The robust and aggressive nature of this venomous insectivore, however, make it an unusually tough and dangerous opponent. It seems more likely, however, that the Red Squirrel was simply defending its food source from a poten- tial competitor. Given the notable scarcity of acorns in fall 1995, our squirrels seemed more aggressive and even desperate in their efforts to obtain food. Acknowledgments Ruth Nero is thanked for taking note of this inci- dent and describing it to me shortly after seeing it take place. Gordon G. Graham kindly typed the manuscript. Literature Cited Nero, R. W. 1987. House Sparrow killed by Red Squirrel. Blue Jay 45: 180-181. Nero, R. W. 1993. Another Red Squirrel bird-kill. Blue Jay 51: 217-218. Nero, R. W. 1995. Short-tailed Shrew apparent cause of mortality of Eastern Screech-Owl. Blue Jay 52: 176-178. Sullivan, B. D. 1991. Additional vertebrate prey items of the Red Squirrel, Tamiasciurus hudsonicus. Canadian Field-Naturalist 105: 398-399. Taylor, P. 1988. Predatory Red Squirrels. Blue Jay 46: 97. Received 29 April 1996 Accepted 27 September 1996 1996 NOTES 713 Changed Status of the Hooded Merganser, Lophodytes cucullatus, in the Yellowknife area, Northwest Territories MICHAEL A. FOURNIER and JAMES E. HINES Canadian Wildlife Service, P.O. Box 637, Yellowknife, Northwest Territories X1A 2N5 Fournier, Michael A., and James E. Hines. 1996. Changed status of the Hooded Merganser, Lophodytes cucullatus, in the Yellowknife area, Northwest Territories. Canadian Field-Naturalist 110(4): 713-714. Increased observations of Hooded Mergansers, Lophodytes cucullatus, have occurred in the Yellowknife area in recent years. These observations indicate that Hooded Mergansers now utilize this region during the post-breeding, moulting, and migration periods. The occurrence of paired birds in spring suggests the possibility of breeding. There are two plausible explanations for this increase, population increase and subsequent range expansion or displacement due to environmental disturbance. Key Words: Hooded Merganser, Lophodytes cucullatus, Northwest Territories, increased numbers, changed status, moulting. The breeding range of the Hooded Merganser encompasses the southern and central portions of most Canadian provinces, but the northern limits of its range are poorly defined (Godfrey 1986) and migratory movements, dispersal, and moulting distri- bution are largely unknown (Bellrose 1980, Dugger et al. 1994). In summer and fall non-breeding or post-breeding birds occur northward of breeding areas across cen- tral and northern Canada and Alaska (Harper 1953, Mossman 1957, Nero 1963, Palmer 1976, Bellrose 1980, Hooper 1989). In British Columbia, the Hooded Merganser is distributed, in spring and sum- mer, north to the Peace Lowlands and the Liard River drainage (Campbell et al. 1990). In the Yukon it is considered rare, indicating only one to three observa- tions occur annually (Anonymous 1992). In Alberta, Hooded Mergansers were recorded as possible, prob- able, or confirmed breeders at five locations in the northern third of the province between 1987 and 1991 (Semenchuk 1992). Historically, the Hooded Merganser has been con- sidered rare in the Northwest Territories (Baird et al. 1884, MacFarlane 1908, Preble 1908). In the Yellowknife area it was classified as a rare summer transient (Bromley and Trauger 1981) based on observations of six individuals between 1962 and 1968. More recently, the Hooded Merganser has been classified as a migrant and summer resident in the mainland portion of the Northwest Territories (Sirois and McRae 1994). The assignment of this status was based largely on data presented here. Many of the observations we discuss, including the largest groups for each year, are the product of ran- dom sightings. Therefore, the factor of increased observer effort as a partial explanation for our obser- vations cannot be eliminated. However, some of these observations are the product of systematic sur- veys conducted by the Canadian Wildlife Service, since 1985, as part of a study of boreal forest ducks in the Yellowknife area. Thus, the increase has been substantiated with systematically collected data. We observed Hooded Mergansers sporadically between 1985 and 1991 in the Yellowknife area, including sightings of lone males in May 1985, June 1989, and May 1990. Five Hooded Mergansers (four males and one female) were observed near Yellowknife in October 1990 and a pair was observed in May 1991. In the summer and fall of 1992 increased numbers of observations of this species began to occur in the Yellowknife area, and continued through the spring of 1995. Observations of 91, 131, and 125 individu- als, from a variety of locations around Yellowknife and the North Arm of Great Slave Lake, were report- ed in 1992, 1993, and 1994 respectively. Although some of these may have been repeat observations of the same birds, these numbers represent a significant increase over the next highest yearly total of six indi- viduals, observed in 1990. The largest single groups observed were: approxi- mately 35 (sex ratio unknown) in 1992, 46 (40 males and six females) in 1993, and 58 (42 males and 16 females) in 1994. All of these large groups were observed in the fall. Hooded Mergansers were observed as late as 22 October in 1993. Observations of two flightless males in July 1992 and a single moulting male in June 1993 provided the first evidence of a probable moult migration of this species to the Yellowknife area. All other observa- tions were of birds in breeding plumage and/or capa- ble of flight. Although most of our observations occurred in late summer and fall, increased observations in spring and early summer began to occur in 1994 and continued in 1995. Between 16 May and 1 June 1994, 23 indi- viduals were observed, including five pairs, whereas in 1995, 26 individuals were observed between 10 May and 11 June, including at least five pairs. These numbers represent a substantial increase. Prior to 714 1994, the highest total number of Hooded Mergansers observed in May and June of any single year was two, which occurred in 1962 (two lone males), 1991 (one pair), and 1993 (two lone males). The earliest recorded arrival of this species in the Yellowknife area occurred on 10 May 1995. Increased observations of Hooded Mergansers near Yellowknife may be indicative of an overall population increase and/or range expansion of the species. We could find no direct evidence to support this hypothesis but this is not surprising. A popula- tion increase would be difficult to detect, as Hooded Mergansers are generally not specifically identified, but grouped with all other merganser species, during surveys designed to detect waterfowl population trends. However, an apparent population increase and range expansion in the northwestern portion of the range was reported previously by Palmer (1976), and a recent review of current knowledge for this species (Dugger et al. 1994) suggested that popula- tions are possibly increasing over much of the range. Our observations could also be a result of a change in migrational or moulting distribution, perhaps relat- ed to loss of suitable habitat in other areas. One major environmental perturbation which could explain a northward shift in distribution of this species during the breeding and/or post-breeding seasons is recent increases in forest exploitation in northern portions of the western provinces. The Hooded Merganser is closely tied to forested wetland systems (Dugger et al. 1994) and a number of authors have related popula- tion declines in this species with deforestation (Phillips 1926, Salt and Salt 1976; Dugger et al. 1994). Displacement of birds (declines) in one region may result in increases elsewhere. Our observations add to the knowledge of this species in the northern portions of its range. First they indicate that an increasing (but probably small) portion of the population migrates as far north (at least) as the Yellowknife region in spring and sum- mer (c.f., Dugger et al. 1994). Further, some of these birds remain to moult and apparently utilize the area for staging prior to fall migration. Finally, paired birds now occur in the Yellowknife area at the appropriate time and in sufficient numbers to suggest the possibility of breeding, although this seems unlikely given the currently recognized breeding range of the species (e.g. Palmer 1976, Bellrose 1980, Godfrey 1986; Dugger et al. 1994, but see Semenchuk 1992). Acknowledgments The authors gratefully acknowledge the assistance of R. Brook and M. Kay during field studies and the contribution of observations by D. Kay of Ducks Unlimited, Yellowknife, and J. Sirois (CWS). A.J. Erskine and an anonymous referee provided com- ments on the manuscript. THE CANADIAN FIELD-NATURALIST Vol. 110 Literature Cited Anonymous. 1992. Birds of the Yukon - field check list. Yukon Department of Renewable Resources, Whitehorse, Yukon Territory. 4 pages. Baird, S. F., T. M. Brewer, and R. Ridgway. 1884. The water birds of North America. Reprint Edition (1974). Arno Press Inc., Salem, New Hampshire. 1089 pages. Bellrose, F.C. 1980. Ducks, geese and swans of North America. Third Edition. Stackpole Books, Harrisburg, Pennsylvania. 540 pages. Bromley, R.G., and D. L. Trauger. 1981. Birds of Yellowknife: A regional checklist. Privately Printed, Yellowknife, Northwest Territories. 12 pages. Campbell, R. W., N. K. Dawe, I. McTaggart-Cowan, J. M. Cooper, G. W. Kaiser, and M.C. E. McNall. 1990. The birds of British Columbia. Volume 1, Nonpasserines. Royal British Columbia Museum, Victoria, British Columbia. 514 pages. Dugger, B. D., K. M. Dugger, and L. H. Fredrickson. 1994. Hooded Merganser (Lophodytes cucullatus). In The Birds of North America, Number 98. Edited by A. Poole and F. Gill. The Academy of Natural Sciences, Philadelphia, and The American Ornithologists’ Union, Washington, D.C. 24 pages. Godfrey, W. E. 1986. The birds of Canada. Revised Edition. National Museum of Natural Sciences, Ottawa, Ontario. 595 pages. Harper, F. 1953. Birds of the Nueltin Lake expedition, 1947. American Midland Naturalist 49: 1-116. Hooper, D. F. 1989. Hooded Merganser in east-central Saskatchewan. Blue Jay 47: 111-112. MacFarlane, R. 1908. List of birds and eggs observed and collected in the Northwest Territories of. Canada between 1880 and 1894. Pages 287-446 in Through the Mackenzie basin. By C. Mair. William Briggs, Toronto, Ontario. 494 pages. Mossman, A.S. 1957. Hooded Mergansers at Afognak Island, Alaska. The Condor 59: 341. Nero, R. W. 1963. Birds of the Lake Athabasca region, Saskatchewan. Special Publication Number 5, Saskatchewan Natural History Society, Regina, Saskatchewan. 143 pages. Palmer, R. S. 1976. Handbook of North American birds. Volume 3, Waterfowl (Part 2). Yale University Press, New Haven, Connecticut. 560 pages. Phillips, J. C. 1926. A natural history of the ducks. Volume IV. Reprint Edition (1986). Dover Publications Inc., New York, New York. 409 pages. Preble, E. A. 1908. Biological investigation of the Athabaska-Mackenzie region. U.S. Department of Agriculture, Bureau of Biological Survey, Washington, D.C. 557 pages. Salt, W. R., and J. R. Salt. 1976. The birds of Alberta. Hurtig Publishers, Edmonton, Alberta. 498 pages. Semenchuk, G. P. Editor. 1992. The atlas of breeding birds of Alberta. Federation of Alberta Naturalists, Edmonton, Alberta. 391 pages. Sirois, J.. and D. McRae. 1994. The birds of the Northwest Territories. Canadian Wildlife Service, Yellowknife, Northwest Territories. 27 pages. Received 29 November 1995 Accepted 20 August 1996 1996 NOTES Ws Extralimital Occurrences of Willow Ptarmigan, Lagopus lagopus, in Maine ROGER D. APPLEGATE Wildlife Division, Department of Inland Fisheries and Wildlife, 650 State Street, Bangor, Maine 04401 Present address: Department of Wildlife and Parks, P. O. Box 1525, Emporia, Kansas 66801 Applegate, Roger D. 1996. Extralimital occurrences of Willow Ptarmigan, Lagopus lagous, in Maine. Canadian Field- Naturalist 110(4): 715. Five previous Willow Ptarmigan records in Maine, south of the usual species range, from 1892 to 1988, are summarized. A new record, from Brooks, Waldo County, taken in May 1990, is documented with a specimen. Key Words: Willow Ptarmigan, Lagopus lagopus, distribution, Maine. The Willow Ptarmigan, Lagopus lagopus L., win- ters primarily on the southern edge of the species breeding range, in the arctic tundra throughout the Palearctic and Nearctic, including the British Isles and Newfoundland . It is reported to wander casually south in winter to Montana, North Dakota, Minnesota, Wisconsin, central Ontario and Maine (AOU 1983). Records from Maine include a Willow Ptarmigan in winter plumage shot in Kenduskeag, Penobscot County (44°50’'N, 68° 75’W), on 23 April,1892 (Merrill 1892). Other reports that are considered reli- able are for T6 R7 WELS, Penobscot County (46° 09'N, 68° 38’ W), 1951 [Dolley 1951, reported by Vickery (1978), as Sherman Mills], Freeport, Cumberland County (43° 55'N, 70° 05’ W), 6 December 1954; Bailey Island, Cumberland County (43°40’N, 70°00’ W), May 1977 (Vickery 1978); and Petit Manan National Wildlife Refuge, Washington County (44° 25’ N, 61°54’ W), April 1988 (Widrig 1989). In May 1990 an adult male in winter plumage was taken captive in Brooks, Waldo County (44° 32’N, 69°07’ W), Maine. The ptarmigan later died in cap- tivity and was turned over to Maine Department of Inland Fisheries and Wildlife officials. The specimen was submitted to the Wildlife Collection, Department of Wildlife Ecology, University of Maine, Orono, Maine 04473. Willow Ptarmigan are considered to be cyclic with an approximate 10-year periodicity of peaks and lows (Johnsgard 1973). Keith (1963: 57) charted data (jargely from Williams 1954) on ptarmigan population peaks and found an 8-10 year grouping of peaks. The Maine occurrences of Willow Ptarmigan in 1892, 1951, 1954, 1977, 1988, and 1990 may be related to these peaks. Some populations of Willow Ptarmigan are con- sidered migratory and this dispersal may be the result of high ptarmigan densities on the breeding range (Watson and Jenkins 1964). Territorial defence from February to May results in dispersal of excess non-territorial ptarmigan in the population (Mossop 1988). This could account for the prepon- derance of records of ptarmigan south of the breed- ing range during April and May. Acknowledgments C. K. Williams and A. J. Erskine provided helpful suggestions on the manuscript. K. Kemper collected the Ptarmigan specimen and supplied details. M. McCollough encouraged publication of this note. Literature Cited AOU. 1983. The A. O. U. checklist of North American birds. 6th edition. American Ornithologists Union, Washington, D. C. 877 pages. Dolley, R. G. 1951. Canadian willow ptarmigan (Lagopus lagopus albus) in Maine. Bulletin of the Maine Audubon Society 7: 79-80. Johnsgard, P. A. 1973. Grouse and quails of North America. Lincoln: University of Nebraska Press. 553 pages. Keith, L. B. 1963. Wildlife’s ten year cycle. University of Wisconsin Press, Madison. 201 pages. Merrill, H. 1892. Lagopus lagopus in Maine. Auk 9: 300. Mossop, D. H. 1988. Winter survival and breeding strate- gies of willow ptarmigan. Pages 330-378 in Adaptive strategies and population ecology of northern grouse. Edited by A. T. Bergerud and M. W. Gratson. University of Minnesota Press, Minneapolis., 809 pages. Vickery, P. D. 1978. Annotated checklist of Maine birds. Falmouth: Maine Audubon Society. 20 pages. Watson, A., and D. Jenkins. 1964. Notes on the behaviour of the red grouse. British Birds 57: 137-170. Widrig, R. S. 1989. The birds and plants of Petit Manan National Wildlife Refuge. Privately published by R. S. Widrig, Box 265, Milbridge, Maine 04658. Williams, G. R. 1954. Population fluctuations in some northern hemisphere game birds (Tetraonidae). Journal of Animal Ecology 23: 1-34. Recieved 9 January 1996 Accepted 6 September 1996 News and Comment Froglog: 1'UCN/SSC Declining Amphibians Populations Task Force Number 19, November 1996, of Froglog, the newsletter of the Declining Amphibian Populations Task Force of the World Conservation Union’s Species Survival Commission, contains items on The Ghost of Pesticides Past? [evidence of persis- tant anthropogenic residues in certain Great Lakes basin frog populations], The Amphibians of Bangladesh, SSAR/DAPTF Symposium Report, Amphibians Under Pollution Impact in Ukraine, Amphibian Decline and Environmental Alterations, Ecosystem Breakdown in Paraguay, Green Frogs Exploited in Former Yugoslavia, Froglog Shorts, Publications of Interest. Recovery: An Endangered Species Newsletter The Fall 1996 issue of this newsletter published and distributed by the Canadian Wildlife Service, Environment Canada, has 12 pages. Although an ISSN number is given (0847-0294), neither vol- ume or issue number appear. On page 7 the Coordinator is given as Chuck Dauphine, and edit- ing and design credited to West Hawk Associates. Copies are free. There is no indication intended frequency of publication, but presumably it is at least twice a year (see previous notice in News and Comment The Canadian Field-Naturalist 110(2): 549). A french edition is available under the title SAUVEGARDE. For subscription send name, address, and language of choice, to Recovery, Canadian Wildlife Service, Environment Canada, Ottawa, Ontario K1A 0H3. This issue’s contents are: Broadening the scope of recovery [greater public participation and emphasis on ecosystem management] (Chuck Dauphine); Recovery Watch — Garry oak ecosystems under siege (Erich Haber, Chair, COSEWIC) — Scotland’s national emblem threatened in Canada [Pitcher’s Thistle] (Anwar Mann, Department of Plant Sciences, University of Western Ontario) — COSEWIC Update [Committee assesses first mol- lusc: Eelgrass limpet]; Saving the loggerhead shrike (condensed from a proposal by the Avian Science Froglog is available on request from John Wilkinson, Department of Biology, The Open University, Walton Hall, Milton Keynes, MK7 6AA, United Kingdom ** Telephone: 01908 (44 1908 if ex©UK) 652274 ** Fax: 01908 (44 1908 if exOUK) 654167 ** e©mail: DAPTF@open.ac.uk. Froglog can also be found on the World Wide Web at the following URL: http://acsinfo.open.ac.uk/info /newsletters/FROGLOG.html FRANCIS R. CooK and Conservation Centre located at McGill University); RENEW Update: A new home on the range [Hook Lake bison calves to foster healthy Northwest Territories herd] (Cormack Gates, chair, Wood Bison Recovery Team); CITES considers tur- tles, bison for 1997 [proposed additions to Appendix II are the softshell, snapping and map turtles, trans- fers from Appendix I to I of the North American gyrfalcon population, and Canadian authorities are considering the preparation of a proposal to transfer the wood bison from Appendix I to II to allow trade in farm-bred animals] (Charles Dauphine); Ranking: A Proposal): A new designation system [an outline distributed at the national endangered species work- shop in June in Toronto suggests new Red, Yellow and Green and Status Undetermined lists based on seven simple criteria] (Bill Harper, Ray Halladay, B.C. Ministry of Environment, Lands and Parks; Gordon Court, Steve Bechtel, Bill Hall, and Bob Andrews, Alberta Department of Environmental Protection); Editorial — Pigeonholing Nature [on the value of status designation even for peripheral species] (Chris Shank, Department of Resources, Wildlife and Economic Development in the Northwest Territories). FRANCIS R. Cook 716 Book Reviews ZOOLOGY Polygyny and Sexual Selection in Red-winged Blackbirds By William A. Searcy and Ken Yasukawa. 1995. Princeton University Press, Princeton. xviii + 312 pp., illus. Cloth U.S. $55; paper U.S. $29.95. The August Krogh Principle states that for every question in biological research, there is at least one species that is ideally suited to study. When it comes to polygyny and sexual selection in birds, few species can rival the Red-winged Blackbird as an “August Krogh” species. Not only is it one of the most polygynous of all bird species, it is also widespread in North America and it breeds in open habitats where capture, nest finding, and direct observations are easy. Consequently, “redwing” behaviour has been studied in many places by many people (a good proportion of which are Canadian). This book is an excellent review of the extensive body of research carried out so far on redwing reproductive behaviour, with a discussion of how results fit with current views of polygyny and sexual selection. Chapter | is an introduction to mating systems and sexual selection. Chapter 2, on parental care, tells us that males are mostly responsible for defence against predators, while females do most of the food provisioning. Chapter 3 describes the male territorial behaviour. Chapter 4 examines the vari- ous factors that affect female reproductive success. Avoidance of predation, by means of strategic nest location or guarding by the male, seems paramount. Chapter 5 shows where females choose to settle, based on various territory attributes; surprisingly, the number of females already present on the territo- ry does not seem to matter. Chapter 6 collates the previous chapters in a test of various models to explain the maintenance of polygyny in redwing populations. It wisely uses an approach where alter- native models are first falsified before support for the remaining model is provided. Here as in the rest of the book, more weight is given to experimental, rather than correlational, evidence. Chapter 7 turns to sexual selection and the extent to which we can see it operating in present popula- tions (not very much, it turns out). Sexual selection was still a potent force in the past, and chapter 8 pre- sents the evidence that many male traits, most notably epaulets and song, are the adaptive result of sexual selection. Chapter 9 extends this adaptationist treatment to females and their behaviour in a polygy- nous system. Chapter 10 recapitulates the conclu- sions from the different chapters, and compares red- wings with other polygynous birds. The writing style is clear, with precise wording. Topics are presented in a smooth logical order, and in a cogent way. In places where evidence is ambiguous, the authors give their opinion about the most valid interpretation, but they never fail to pre- sent the alternatives. Each chapter ends in a conclu- sion that summarizes the main findings while point- ing to future research. (There is still room for research in this busy field, if only because some interpretations are based on negative results, which need to be replicated to increase statistical power.) The authors are both very active in redwing research, and they seem to benefit from regular contact with other heavy hitters, as evidenced by the inclusion of some unpublished results and personal communica- tions. An impressive 435 references are listed. This book may be too specialized for the field nat- uralist, notwithstanding the fact that almost all stud- ies on redwings are conducted in the field. However, the book is a must for students of polygyny and sex- ual selection; in fact, every professional ornithologist should have it, given that redwings have provided many of the textbook examples on which our views of polygyny and sexual selection in birds are based. August Krogh would approve. STEPHAN REEBS Département de biologie, Université de Moncton, Moncton, New Brunswick E1A 3E9 Artificial Nest Structures for Ospreys: A Construction Manual By P.J. Ewins. 1994. (Aussi disponible en frangais). Canadian Wildlife Service, Toronto. 39 pp., illus. Free. This is a publication of potential interest to hand- skilled field naturalists. Indeed, this concise and well-illustrated construction manual was written pri- marily to encourage and help people construct, install, and maintain artificial nest platforms for Ospreys. Designs presented are those deemed best suited to habitats found in Canada but are by no means restricted to those, as evidenced by the large proportion of the designs depicted that were devel- oped for habitats found in the United States. TAG 718 Site selection considerations and general notes on construction are presented in point form, as a pream- ble to descriptions of artificial nest platforms. Here, proximity of such platforms to sensitive commercial interests sites such as fish ponds and pisciculture sta- tions would be a worthwhile addendum to the manu- al’s site selection considerations. The majority of nest platform designs are grouped under single pole, tripod/quadrapod, or power line structures. Because installation of the latter structures requires significant assistance from utility personnel, such information should be of rather limited value to most naturalists. Nonetheless, there still remains an array of thirteen nest platform designs for the more typical field naturalist to choose from. Most types of platforms are depicted by means of a self-explanatory diagram and a complete list of materials is provided as well. Although diagrams vary in quality, important details always remain discernible. I found only one minor typographical error in the manual. The dia- Sibley Fishes By S. A. Stephenson and W. T. Momot. 1994. Occasional Paper No. 15, Centre for Northern Studies, Lakehead University, Thunder Bay. iv + 133 pp. illus. $10.00. Sibley Fishes is an intriguing title which perhaps needs some explanation. Sibley is the name of the peninsula jutting out into Lake Superior just east of Thunder Bay. Habitat on the peninsula was protected by the creation of Sibley Provincial Park in 1944. The park, now known as Sleeping Giant Provincial Park, protects a remnant of the original flora and fauna, including some White Pine forest. This attractive small book has two main purposes: to provide a guide to the fish fauna of the park and to record present distribution so that changes may be recognized. It does both tasks well. It begins by describing the area and the methods used to sample the fish community. Various factors that influence distribution are considered, especially the ecological requirements of fishes and the geolog- ical processes that have limited or aided their disper- sal. Events since the retreat of the last Wisconsin glacier, about 9700 years ago, and those associated with draining of glacial lake Agassiz determined fish distribution on the Sibley peninsula. Problems today concern preservation of the naturally established fish communities from deliberate or accidental introduc- tions and from undue fishing pressure. The body of the book consists of brief descriptions of each of the 36 species recorded within the park. Each description is accompanied by an accurate sketch showing the form, fin placement, scale tex- ture, and contour pattern. A map shows the streams and lakes where each species has been recorded. THE CANADIAN FIELD-NATURALIST Vol. 110 gram of the optional inspection step for the Sanibel Tripod is referred to as Figure 13 instead of Figure 11 on page 17. As a biologist, I wished that some esti- mates of manpower and construction costs as well as information on the relative durability and vulnerabili- ty to vandalism of the main types of nest platform structures would have been provided. Such details might be obtainable through raptor organizations list- ed at the end of the manual. This manual should prove particularly useful to personnel working for raptor organizations, wildlife agencies, and power utilities as well as to naturalists who want to play a more active role in Osprey prop- agation and those summer cottage owners who asked for the installation of a nest platform within sight of their dwelling but were placed on waiting lists. SYLVAIN R. POIRIER P.O. Box 1103, Dalhousie, New Brunswick EOK 1BO As well as its obvious educational value, there is a third purpose to the book which should interest readers of The Canadian Field-Naturalist. This is to stimulate debate about conservation of aquatic resources within parks. We allow, even encourage, sports fishing in our parks, while we object to all other forms of exploita- tion. Why do we accept this illogical practice with its risks of introductions of exotic species for put-and- take fishing, introductions of parasites and diseases, and the danger of destabilising established native fish communities? The authors do not think we should ban fishing in parks but seem ambivalent about it because of the risks. Arguments for allowing the practice to continue are that it is a valid outdoor experience which pays handsome dividends in creating a love for the outdoors, promotes a conservation ethic and establish- es a group of unpaid, wise watchdogs, keeping their eyes on the state of the aquatic environment. An unresolved problem associated with banning fishing in parks is dealing with the question: what good are the fish since they are generally invisible unless caught? In this respect, to realise the second purpose of the book the authors (page 9) invite per- sons collec*ing specimens in the park to share the species dist.'butional information with them, which suggests fish sampling be allowed in the park for sci- entific reasons. It seems to me it would be best to treat fish like the rest of the flora and fauna and allow only obser- vation. This can be done by promoting snorkelling, underwater photography, or by the use of glass bot- tomed boats. Providing the fish are not trained to appear at feeding sites, as occurs in some places, this 1996 use of fish in parks should not alter natural commu- nities and undisturbed aquatic communities should be one of the assets of parks. This book is likely to appeal most to park visitors and residents of northwestern Ontario. BOOK REVIEWS 719 GEOFF POWER Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1 The Rise of Fishes: 500 Million Years of Evolution By John A. Long. 1995. The Johns Hopkins University Press, Baltimore and London. 223 pp. 352 illus. Cloth US. $49.95. The Rise of Fishes is a magnificent book, written by the Curator of Vertebrate Palaeontology at the Western Australian Museum, Perth, Australia. It pro- vides a chronology of fish-like vertebrates from their origins to the present day. It explains how they learned to swim, manipulate and consume large prey, respire efficiently in water and in air, and how the organ systems of their bodies evolved. It is written from the perspective of someone who lives and works near the centre of some of the action, Gondwanaland, perhaps the cradle of fish evolution. The author knows many of the fossils first hand and has discovered and described enough of them to be able to write about them with confidence and authority. The story begins with the earliest evidence of life on earth and a description of the fossilization process, evolution, continental drift and the geological time scale. The origins of fish-like vertebrates are consid- ered next, along with their near-vertebrate relatives and bone, the characteristic vertebrate tissue. Then follow chapters on the Agnatha, Chondrichthyes, Acanthodii, and Placoderms. The origins, classifica- tion and diversity of each of these classes of verte- brate is presented in an interesting and lively way. The latest finds, many from Australia and China, are included. The author has the skill to make these ancient fishes seem real and alive. The extraordinarily good, coloured photographs of the actual fossils help, as do the many diagrams and illustrations of the fish- es. What I liked about these chapters was the feeling, after reading them, that I knew something about the inhabitants of the ancient seas and freshwaters. The bony fishes, Class Osteichthyes, include almost all living fish species and a lot of fossil repre- sentatives. I liked the description of the advantages these fish acquired from their teeth, bony scales, skeleton, and lungs, which became the hydrostatic organ, and their fins. I also liked the picture of them struggling to survive amongst the other more heavily armoured Devonian predatory fishes. The competi- tion was apparently worth it for their descendants radiated out to fill almost every aquatic habitat on earth and gave rise to the immense variety of bony fish we know today. The most successful group, the Actinoptergii, are traced from their origins in the Devonian from which deposits a variety of “palaeoniscoids”, including genera from the tropical Late Devonian Gogo reefs of Western Australia, are described. Radiation of the group is followed through the Mesozoic with a selec- tion of fossil fish, many from Australia, that were contemporaries of the great reptiles. Finally, in the late Cretaceous and through the Cenozoic., we get a glimpse of ray-finned fish diversity from the fine fossil beds of Brazil (Santana formation), USA (Green River, Wyoming), and Italy (Monte Bolca). Lungfishes get extensive treatment with a view of Devonian diversity, a good treatment of their denti- tion, feeding ecology, and the origin and use of lungs. The Crossopterygians are likewise given thor- ough treatment, with emphasis on the Coelacanths. In this group it has been possible to check deduc- tions from fossils against observations on the only extant member, Latimeria chalumnae, discovered off the coast of South Africa in 1938. John Long’s sense of humour shines through when relatives of the Coelacanths are labelled fat-headed, beady-eyed predators (Porolepiformes) and, one of them, a fossil from Quebec, is named Quebecius quebecius. He comments, “where else with a name like that?” Hlustrations of other relatives, the dagger-toothed fishes, Onychodontiformes, and the giant killers, Rhizodontiformes, are life-like to the detail of blood gushing out of prey impaled on their teeth. The greatest step in vertebrate evolution, the move onto land, provides a climax to the book. It is described eloquently, logically, and with a sense of excitement and discovery. Like a good detective story, the plot should not be revealed until all the clues have been considered. The book comes to a fascinating and somewhat unexpected end. I thoroughly recommend The Rise of Fishes to anyone with any interest in fish or vertebrate palaeontology. The more than 300 beautiful illustra- tions are worth the price of the book alone. They are colourful, reflecting the brilliant colours of most of Australia’s present ichthyofauna. Would they have been so bright if they had been prepared by a Canadian? Perhaps not. This book was badly needed to make more generally known many new discover- ies and to provide a counter balance for popularity of the dinosaurs. The real action was earlier. Read this book and discover for yourself. GEOFF POWER Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1 720 THE CANADIAN FIELD-NATURALIST Vol. 110 Masters of the Ocean Realm: Whales, Dolphins, & Porpoises By John E. Heyning. 1995. University of British Columbia, Vancouver. 112 pp., illus. $24.95. When I was researching the East Coast whale watching industry for a federal government report, several of the operators I interviewed told me that their client’s knowledge had changed dramatically in the last few years. It was no longer possible to satis- fy them with a boat ride and a few sightings of big black shapes as clients now demand much more. The increase in awareness and interest in whales has been promoted by many fine new books and television programs. Most recently these books have tended to be more specialised accounts of single species and groups of whales or more detailed field and site guides. This new contribution reverses that trend and goes back to the most fundamental basics. Indeed, in many ways it is a book on the basics of mammal, and not just whale biology. A non-biologist could read this book without difficulty and, with reason- able effort, could gain an understanding of many of today’s environmental issues. The author begins by describing what a whale is (and is not) and gives some of the remarkable facts that hold us in awe. To illustrate and clarify impor- tant points he uses diagrams, charts, drawings and photographs. For example, he has used a drawing of a dolphin, followed by a diagram of the same dol- phin to explain the skeletal structure. Similar outline and skeleton drawings of a dog and a human follow to show the relationship between the species. Each group of arm bones is colour coded to make these associations easier to follow. To capture special ideas that are peripheral to the flow of the text he has included inset boxes. His first use of this device is to explain parasitism in general and the particular para- sites some whales carry. Heyning uses an inset in his chapter on whale evo- lution to outline the major points of general evolu- tionary theory. This complements his account of the development of whales from small hoofed land mammals to giants of the sea. In describing the life of a whale he informs the reader of such whale traits as filter feeding with baleen, deep diving without the bends, sonar and echolocation, and the physical acts of spyhopping, lobtailing, and breaching. As well, he also covers more generic concepts such as intelli- gence, the food pyramid, and migration. The chapters on conservation and research sum- marize the problems, needs, and status as these top- ics relate to whales, dolphins, and porpoises. He also provides the reader with a sense of these issues as they relate to our world in general. For example, the latest techniques in DNA fingerprinting, satellite tracking, photo-identification, and population census methods are covered with great simplicity and clari- ty. The author is primarily concerned with the appli- cation of these techniques to whale research but as they are more widely used in scientific investigations the reader is introduced to their general application. The first and last chapters of the book cover the rela- tionships, both real and mythical, between people and whales. While these are described without an overlay of sensational emotion they leave us with a clear sense of our responsibilities for the whales and other creatures; responsibilities created by our own misuse of the world in which we live. The text begins with very straightforward lan- guage and anybody from teenager to a senior will have no difficulty reading this book. The author pro- gressively introduces scientific expressions as they are required and explains each one in simple terms. Once introduced, he continues to use the new term for the rest of the text, so the language becomes a lit- tle more complex as you progress through the book. The combination of readily understandable text and top rate illustrations make this book a winner for me. It is ideal for anyone newly interested in whales and other marine mammals or is about to go on their first whale watching trip. There is enough information to make a trip more rewarding but not enough to give information overload. However, the book is not, and is not meant to be, an identification guide as this type of material should be provided by the trip oper- ator or one of the field guides dedicated to whales. This book would be an ideal gift for a budding naturalist of any age. Teachers could also use it effectively as it is such an excellent introduction to many aspects of biology that uses a fascinating group of animals as its core. I have been studying whales for over 20 years and I have read many books on these mammals. Despite the basic nature of this new addition to whale literature I did learn some- thing new and I had some long-unused segments of my memory dusted off and updated. For whatever reasons you choose you will not go wrong in buying this fine book. Roy JOHN 1-613 McPherson Avenue, Saskatoon, Saskatchewan S7N 0X7 1996 BOTANY BOOK REVIEWS 7?) A Checklist of Vascular Plants for Bruce and Grey Counties Compiled by Bruce-Grey Plant Committee. 1995. 41 pp. Available from Own Sound Field Naturalists, Box 401, Owen Sound, Ontario N4K 3T1 or The Saugeen Field Naturalists, Box 20156, Hanover, Ontario N4N 3T1. $5.00 + $1.00 postage and packing. Checklists of the vascular plants of any region are always most useful for visitors and the local inhabi- tants interested in what is growing around them. This checklist for Bruce and Grey Counties, which includes all of Bruce Peninsula (but excludes Keppel, Sarawak, and parts of Derby Townships of Grey County), is an excellent one. It includes 1420 taxa (species, subspecies, and hybrids) representing 134 families. The list is divided into three groups: Ferns and Fern Allies, Conifers, and Flowering Plants. The lat- ter is again divided into Dicotyledons and Monocotyledons. Within each of these groups, the families, genera, and species are listed alphabetical- ly, followed by a common name. What makes it spe- cial is the detail that is provided. The region is divid- ed into three areas, Bruce Peninsula (p), southern Bruce County (b), and Grey County (g). The area from which each taxon is known follows the latin name, e.g., p,b,g, or P,B,G, if the distribution of a native species is understood and is considered to be Sonoran Desert Plants: An Ecological Atlas By Raymond M. Turner, Janice E. Bowers, and Tony L. Burgess. 1995. The University of Arizona Press, Tucson. xvi + 504 pp., illus. U.S. $70.00. The Sonoran Desert Region occupies the southeast corner of California, the southwest quarter of Arizona, and, in Mexico, much of Baja California and about two-thirds of the State of Sonora. In 1972, J. R. Hastings, R. M. Turner, and D. K. Warren pub- lished An Atlas of Some Plant Distribution in the Sonoran Desert. This contained maps of 238 species. It was their intention, at that time, to compile “a more complete atlas showing the distribution of the major perennial plants of the Sonoran Desert”. The present volume includes 339 species repre- senting 54 families. It is greatly enlarged from the earlier treatment and now includes such information as scientific name and authority, selected common names, description, diagnostic characters of similar species, taxonomic problems, habitat, distributional patterns in the maps and profiles, biogeography, phe- nology, physiology, reproductive ecology and polli- nation, seedling establishment, growth rate and life- locally rare. In addition, marginal annotations have been made as follows: Introduced from outside Ontario Escaped from planting Rare in Ontario — Natural Heritage Information Centre (S-ranking 1-3 only) Historical Record — not recorded for 30 years or more, in some cases less if a single colony is known to be extirpated. ma Am * Pages 40 and 41 are Instructions for Reporting New or Rare Plants and Appendix A: Counties of Bruce and Grey Report of New or Rare Plant Species. Reports of new information should be sent to: Ministry of Natural Resources, 611 9th Avenue East, Owen Sound, Ontario N4K 3E4. They will in turn forward the information to the Bruce-Grey Plant Committee. All of this makes this checklist one of the best yet produced and those who contributed to it are to be congratulated. WILLIAM J. CoDYy Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-food Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A 0C6 span, horticulture, ethnobotany, and economic botany. In addition, beside each distribution map, there is a scale which depicts the altitudes of the localities on the map, a very unique feature. Scattered throughout the volume are a collection of photographs of various species, mainly taken by J. R. Hastings. These have a tendency to be rather dark, but still are most useful. A glossary, literature cited, and an index complete the work. This publication represents a tremendous amount of work over a long period of time to ascertain the distributions, habitats, and other information on this group of most interesting plant species. It will be most useful both to those interested in their local vegetation and those who visit parts of the Sonoran Desert for the first time. WILLIAM J. Coby Eastern Cereal and Oilseed Research Centre, Agriculture and Agri-food, Canada, Wm. Saunders Building, Central Experimental Farm, Ottawa, Ontario K1A OC6. 722 The Private Life of Plants By David Attenborough. 1995. Princeton University Press. Princeton, New Jersey. 320 pp. + illus. The book starts with “Plants can see. They can count and communicate with one another. They are able to react to the slightest touch and to estimate time with extraordinary precision.” This statement succeeded in getting this reader’s attention, curiosity. fascination. whatever you call it. The writing is lucid and concise. Technical terms are kept to a minimum. The numerous, excellent color photographs occupy about 50 per cent of the book Attenborough says “This book is an attempt to see the natural world [of plants], not from our point of view, but from theirs.” I had the feeling that the pupose of the book was to raise our level of aware- ness of the importance of plants in the World and how they are essential for our survival. It is not exaggeration to say that all animals, and that includes humans, depend upon plants for survival. What would our life be like without wheat. corn. beans, potatoes, grapes, etc. for food and drink? What would our diet be like without the grasses, clovers, grains, etc. that eventually appear in the markets as steaks, hams, and chops? Attenborough makes the point that we exploit plants, not only for food, but for clothes, building materials, and decora- tion. And the books shows how we do this and how, to put it in Attenborough’s intriguing view, the plants use us. ENVIRONMENT Peterson First Guides: Forests By John C. Kricher. 1994. Houghton Mifflin Co., Boston and New York. 128 pp.. illus. U.S. $4.95. This is truly a pocket guide being 9.5 x 18.5 cm. It is designed for adults (teenagers and up) and is an introduction to the various types of forests in North America with a distinct emphasis on the United States. The introduction discusses the identification of plants and animals, and the characteristics of for- est types. The major part of the book treats 48 forest types. The format, with a couple of exceptions, devotes two pages to each forest type with a page of coloured illustrations of the plants and animals fac- ing a page of descriptive text. The illustrations are very good but there are some problems: 1) Most people will not be able to separate Red Crossbills from Pine Grosbeaks, page 70, because the distinctive differences are not clear. 2) The names Red-headed Woodpecker and Rose- breasted Grosbeak are reversed on page 54. 3) The Red-tailed Hawk illustration, page 14, shows a bird THE CANADIAN FIELD-NATURALIST Vol. 110 The book is composed of six chapters that are titled: Travelling, Feeding and Growing, Flowering, The Social Struggle, Living Together, and Surviving. The format of each chapter is a series of detailed examples which illustrate the various aspects of the lives of plants. Attenborough trav- elled the World to assemble the fascinating exam- ples that illustrate his story. Many of the items are from the exotic parts of the World where man has not trashed the landscape, thus destroying the nat- ural areas where the plants and animals had devel- oped, or evolved if you prefer, the specialized rela- tionships on which a plant may depend for its sur- vival. Some of the examples are marvels of intrica- cy. the reasons for which have escaped the eyes of man until recently. Sex, in all sorts of permutations and moods, inevitably appears in every plant’s private life. Sexual deception is a favourite theme, such as orchids that look like bees. And there are more than just plants. Whether you are interested in birds, bees or bats, this book has something for you. Attenborough has produced a television series of the same name. The two overlap as well as comple- ment each other. J. GINNS Cantley, Quebec J8V 3G5 in a nest with only the head and tip of the back showing. It does not show the characteristic features needed to identify the bird. 4) Working with such a restricted number of pages and trying to deal with hundreds of organisms, it is perplexing to see some plants and animals illustrated more than once, for example, Sub-alpine Fir is shown on pages 74, 76, and 122. The writing is sometimes wordy and sometimes repetitious. Thus some pertinent facts about the organisms could not be included. The content is sometimes confusing. For example, the phrase “Hickory seeds are contained in thick hickory nuts,...” is misleading, because the hickory seed is the nut. The nut is-composed of a thick outer husk and a thinner inner shell which contains the kernel. The section on“Widespread North American mam- mals” includes the Pronghorn, the Collared Peccary, and the Grizzly Bear, three species which are restricted in numbers and geographical distribution, 1996 but does not mention the Beaver, Porcupine, and Raccoon, three common and widespread mammals. The text describes the Great-horned Owl’s call as “a resounding hoot,” but a simple hoot is not the dis- tinctive multinote call that is characteristic of this owl in the northeast. The distinctive features ofthe Pileated Woodpecker (page 113) should have includ- ed the red cap. And we are told that “Birds tend to be vocal and active during the daylight hours,...” But such a statement seems a poor use of crucial space which could have been used to tell readers that the hours of dawn and dusk are the best times to see and Terrestrial Ecosystems Through Time: BOOK REVIEWS V3} hear birds. Thus the quality is lower than that we associate with the Peterson Field Guides. Despite some weaknesses, this is a good introduc- tory guide to forest types and the plants and animals that inhabit those forests. I think most people will be impressed to see how many distinct types there are. It is recommended to all naturalists because of its broad overview of the plant communities which we commonly refer to as forests. J. GINNS Cantley, Quebec J8V 3G5 Evolutionary Paleoecology of Terrestrial Plants and Animals By A. K. Behrensmeyer, J.D. Damuth, W. A. DiMichele, R. Potts, H. D. Sues and S. L. Wing. 1992. University of Chicago Press. xix + 568 pp., illus. In search of the universal we find impossibility. There can be no bible, a single book describing all there is to know of the past. Yet, we try, if for the only reason as to hint at the possibilities and com- plexities of our understanding of the past. Most geologic or primary paleontological texts provide brief outlines of the major events in time and the changes geological or biological with perhaps a simple synthesis of what the ecosystems may have looked like. It is feared that the prospective student’s next jump after learning the parochial is complete into whatever small field of taxon or time he or she choses, at least for a time, forgetting the interrela- tionships of groups at the higher level. Terrestrial Ecosystems Through Time not only bridges this gap, slowly leading the student along deeper into the cav- erns of knowledge but attempts to place the taxon or time in a global, continuous area, inflating the two dimensional faunal lists into three dimensional ecosystems. The 541 pages of text and voluminous references introduce the ways of inferring ecology from mor- phology without neglecting the importance of sedi- mentological or taphonomic information. The com- plex and still growing data from today’s ecosystems are even more mysterious yet wonderfully inciteful the further back in time (chapters 1-2). If we examine specifically the fossil plants (intro- duced in chapter 3) and animals (introduced in chap- ter 4), each have their own pitfalls in paleoecological interpretation but many strengths as well. The floral record “preserves information about ancient vegeta- tion on very fine temporal and spatial scales” (page 140). Morphology of species may indicate a particu- lar role in the ecosystem (ecomorphs) to which infor- mation may be enhanced by locomotor capabilities, body size, and behaviour (e.g. dinosaur nests) to name a few. The following three chapters are summaries of major biological events through time, Paleozoic, Mesozoic and early Cenozoic, and late Cenozoic. And the closer to the present we get, the complexity of species and their possible interactions increase, which is illustrated by the depth of discussions. Each chapter includes paleotopographic and plate position maps. The scope of these chapters is grand and the authors have relied on discussions with major participants in their respective fields. To syn- thesise the interelationships of species from bivalves, to plants, to man is herculean but the authors succeed in providing the essential hints of ecosystem reconstruction. Though I used the term “student” as a potential user of this book I should underscore my meaning as a idealistic term. We are all students and no mat- ter what level of our personal understanding we must be drawn back and reminded of the place in the grand scheme of things when an extant or extinct species belongs. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO 724 A Primer of Ecology By Nicholas J. Gotelli. 1994. Sinauer Associates, Inc., Sunderland, Massachusetts. xiv + 206 pp., illus. U.S. $18.95. Ecology is a tremendously varied and complex subject. If it weren’t, life on earth would not be what it is. Most of us are interested in a relatively small area of ecology. Nevertheless we complain that too little is understood and there is too much to do with- in our field. A wide range of methods or techniques are available to help us understand the relationships between organisms which, after all, is what ecology is all about. The techniques range from a hand lens to DNA sequencing. This book is an introduction to the mathematical models taught in university ecology courses. Models are necessary tools to the understanding of plant and animal communities, to describe past events, to pre- dict the effects of natural events, and to the applica- tion of management techniques on existing popula- tions. It is the author’s contention that ecology text- books do not give enough explanation to the simpler mathematical models, in particular the exponential model of population growth. As a result the students lack the detailed understanding necessary to use and manipulate the more complex models. The goal of the author was “to present a concise but detailed exposition of the most common mathematical mod- els in population and community ecology.” An understanding of continuous differential equations and calculus is assumed. The seven chapters are titled: exponential popula- tion growth, age-structured population growth, metapopulation dynamics, competition, predation, THE CANADIAN FIELD-NATURALIST Vol. 110 and island biogeography. Each chapter is divided into sections on model presentation and predictions, model assumptions, model variations, empirical examples, problems, and solutions. ; The empirical examples section of each chapter focusses on a few field studies and how mathemati- cal models either explain trends or could have been applied to predict trends. In one example the Song Sparrow population of Mandatre Island, British Columbia, is used to illustrate the concepts of density-dependent and density-independent. Island populations are a favored situation for population studies because the isolation reduces the factors affecting fluctuations; 1.e., emigration and immigra- tion are often nil. The Song Sparrow population has varied greatly but its variation did not conform to a simple logistic growth model. Analyses indicated that space and food limitations determined breeding success, because only a limited number of breeding males could establish territories. The excess male population or “floater males” would move in if a res- ident male was lost. Food, when artificially supplied, resulted in a fourfold increase in chicks. Some other examples are Grizzly Bear populations, Red Grouse and its parasites, hare-lynx cycles, and competition between intertidal sandflat worms. A well-written book treating a specialized aspect of ecology, this is certainly not everyone’s piece of cake because the mathematics is too detailed. However, the basic premises are parts of ecology that we should, at least, be aware of. J. GINNS Cantley, Quebec J8V 3G5 The Wilderness Condition: Essays on Environment and Civilization Edited by Max Oelschlaeger. 1992. Sierra Club Books, San Francisco, California. 345 pp. One of the most positive results of society’s heightened environmental awareness is our increased knowledge and appreciation of wilderness. For many of us, wilderness can be a tonic for everything that creates stress in us and that separates us from the earth. The media have also discovered the environ- ment, providing extensive coverage of certain high- profile issues. While it is vital to be aware of current environmental topics, it is most refreshing to occa- sionally read a well-written book that avoids dis- cussing any of these more transitory and newsworthy environmental topics in favour of more fundamental issues. The Wilderness Condition is such a book, an impressive anthology of essays that reflect on the subject of wilderness philosophy. That subject may not seem to be the most natural combination, but, in this book, the tripartite marriage of wilderness, phi- losophy, and history seems to work quite well. The essays comprising this volume were orginally pre- sented at a 1989 conference in Colorado, then pub- lished in both hard cover and softcover versions. The Wilderness Condition concerns some of the more underlying and often-neglected environmental issues of our society. The simple subject of the essays in this book is the tortured relationship between civilization and the environment. So, instead of just covering the more current environ- mental issues, this book provides readers with some necessary background reading on this subject. Max Oelschlaeger, the editor of this volume and the author of both the introduction and the last chap- ter, states that this book finds its role primarily in the “re-greening of the American consciousness.” What 1996 an ambitious goal!! Towards this end, the writers address some of the more subtle values of wilderness and the relationship between wild nature and civi- lization in a manner designed to facilitate cross-dis- ciplinary discussion. Three themes unite this collec- tion of essays: the evolutionary perspective that con- firms why humankind and nature are necessarily interdependant, how our language reflects our under- standing (or lack thereof) of environmental issues, and finally how we can initiate a new recognition of the environment as a “vital entity”. For those inter- ested in these aspects of wilderness, this volume cer- tainly contains a wealth of valuable information. The authors explore the dynamic tension between wild nature and civilization, offering insights into why this relationship has become so tortured. They argue that no amount of technology will ever dis- place our primal connection to nature. Rather than simply deploring the prevailing attitudes towards our imperiled environment, the authors also offer fresh, realistic and inspiring ideas for alleviating the crisis. Oelschlaeger emphasizes the diversity of the writ- ers’ backgrounds. Most of the essayists are also uni- versity instructors and their choice of topics and ‘writing style relects their varied levels of academic expertise, often with a resultant prose that is a little too pedantic. Nevertheless, all the authors have impressive writing credentials in wilderness philoso- phy and environmental ethics, including Gary Snyder and Paul Shepard. As with many anthologies, there is some unevenness in the style of writing, but this can also be viewed as a opportunity to enjoy this variety of rhetoric. Many of the chapters concern notable topics. Shepard presents his theory of a primitive develop- ment of wilderness. Michael Zimmerman looks at the MISCELLANEOUS BOOK REVIEWS 725 relationship between wilderness and the human con- dition, as seen through the eyes of other writers. There are also predictable discussions of the eco- philosophies of Aldo Leopold and John Muir (by Curt Meine and Peter Michael Cohen, respectively), per- haps not too surprising seeing as how the hard-cover version of this book was published by the Sierra Club. Finally, Oelschlaeger himself explores the use of lan- guage by some of the great wilderness writers. Perhaps a result of the topics and authors, many of this book’s chapters are heavy and extensive reading. Many of the essays carry catchy titles, but one brief Appendix is laborously titled “Exegesis of Claude Levi-Strauss’s Idea of Nature as Totemic Metaphor and Caste Metonym” ..., not exactly something you can sit down and casually read. At the end of the vol- ume, there are two appendices and a detailed series of references and notes for each of the chapters. In addition, I was particularly impressed with the effu- Sive prose on the dust cover of the hard cover ver- sion of the book. Our society has become so enamoured with wilder- ness that many people will travel for hours to find the last vestiges of anything remotely wild or will read virtually anything on the subject. Unfortunately, many of us wilderness lovers lack a strong back- ground in wilderness philosophy. It is books like The Wilderness Condition that can provide some valuable reading to help us develop that much-needed philoso- phy and environmental ethic. Our love for wilderness suggests that this is a worthwhile first step. MIke Morris Mono Cliffs Outdoor Education Centre, R.R. #1. Orangeville, Ontario LOW 2Y8 Evolutionary Archaeology: Methodological Issues Edited by Patrice A. Teltser. 1995. The University of Arizona Press, Tucson, Arizona. 216 pp., illus. Cloth U.S.$45; Paper U.S. $21.95. The contributors to Evolutionary Archaeology examine the applicability of Darwinian evolutionary theory to the evolution of archaeological material culture. Their main premise is that artifacts are part of the human phenotype; therefore, Darwinian evo- lutionary processes act upon material culture in the same way as they act upon the human phenotype. The chapters consist either of theoretical and methodological discussions or of examples in which these principles are used to explain archaeological phenomena. The introductory chapters (Teltser; Jones, Leonard, and Abbott; and Dunnell) provide a stimulating discussion of the differences between traditional cultural evolutionary theory (which they refer to as “essentialist”) and an evolutionary theory more closely aligned with Darwinian origins and principles, which they call “selectionist”. The major difference, as they see it, is that selectionist evolu- tion makes no assumptions about the “progressive” nature of evolutionary changes, whereas traditional cultural evolutionary theory has assumed that change is directional and progressive. My reaction to this book is definitely mixed) Un the positive side, they argue for more rigour in selecting units with which to measure evolution. For example, Dunnell (Chapter 3) appropriately criti- cizes the use of artifact “type”, a discontinuous phe- 726 nomenon, as the unit by which to measure evolution, which is a continuous phenomenon. On the other hand, I find the authors’ strict focus on the artifact to be both reductionist and mechanis- tic. It ignores the role of knowledge as the means by which technology is used to enhance the potential reproductive success of the human phenotype (see Ridington 1988). In fact, survival involves a com- plex interaction of knowledge, technology, and actors (Hakken 1993). Granted, archaeologists find artifacts and not “knowledge”; nevertheless, to ignore this complex interaction weakens the theory. In fact, the more I read of the book, the more I began to question the explanatory power of selec- tionist theory to explain this complex interaction. For example, both Jones, Leonard, and Abbott (Chapter 2) and Ramenofsky (Chapter 7) discuss the non-role of intent in shaping human adaptation. Their propo- sition is that intent is solely a source of variation and not a mechanism by which traits are selected (true enough). But the selectionist explanation seems to be too simplistic to descibe the complex course of events that follow an initial act of intent, especially when we see in our contemporary society the contin- uation and proliferation of technology that appears to provide anything BUT reproductive success (see for example, Lutz 1996). Given the complex nature of the interaction among technology, knowledge, and society, chaos and complexity theory (cf., Saperstein 1995) seem to be more appropriate than selectionist theory for explaining cultural evolution. While I found the theoretical chapters stimulating (even though I didn’t agree with everything), I found the chapters describing examples of applied selec- tionist theory quite unconvincing. They did not pro- Huxley: The Devil’s Disciple By Adrian Desmond. 1994. Michael Joseph, London. 475 pp.. photographs. Adrian Desmond’s first foray into biographical accounting, in book form, was hand in hand with J. Moores and the object of their desire was the 19th century centre piece of biological awareness, Charles Darwin. Despite the richness of the Darwinia indus- try the success of Darwin is two fold; the incorpora- tion of the multitude of original source (Darwin was one of the most avid 19th century note takers); and the continual connection of the purple prose of the Victorian era with contemporary feeling. The book reads like a novel, the center character, fully inflated from the pages is a person, affected by the social, political, scientific, and personal. Huxley: The Devil’s Disciple is Desmond’s newest contribution, done in the same vein; 379 pages of flowing text, with nearly a hundred pages of references. THE CANADIAN FIELD-NATURALIST Vol. 110 vide me with any new insight into artifact change and the adaptive advantages it provided. Furthermore, they made several questionable assumptions. Graves and Ladefoged (Chapter 8), for example, propose that activity which is NOT direct- ed toward food-gathering or reproduction is more likely to originate in places where environmental perturbations are more likely to be severe, namely at the “geographic margins of a regionally distributed archaeological complex” (page 162). They provide neither rational nor example to explain and demon- strate why this would be so; and I could not think of any examples out of my own experience that would support such a proposition. In O’Brien and Holland’s conclusion, they state that their intent was to convince archaeologists of selectionist evolution’s “powerfulness relative to explaining variation in the archaeological record” (page 176). They haven’t convinced me. References Cited Hakken, D. 1993. Computing and Social Change: New Technology and Workplace Transformation, 1980-1990. Annual Review of Anthropology 22: 107-132. Lutz, Diana. 1996. No Conception. The Sciences 36(1): 12-15. Ridington, Robin. 1988. Trail to Heaven: Knowledge and Narrative in a Northern Native Community. Douglas & McIntyre, Vancouver. Saperstein, Alvin M. 1995. War and Chaos. American Scientist 83(6): 548-557. MARGARET G. HANNA Curator, Aboriginal History Unit, Royal Saskatchewan Museum, 2340 Albert Street, Regina, Saskatchewan S4P 3V7 The term “bull dog” is often associated with T. H. Huxley (1825-1895). By definition a bull dog, is sub- servient to his master (in popular perception to Darwin) and who would do the nasty business when called upon. Desmond’s new accounting shows in vivid detail that Huxley was not the usual bull dog (“if I had a wish to live thirty years, it is that I may see the foot of Science on the necks of her Enemies”, page 253) but for the most part he was his own master, serving his own needs. In his early days his frustration with patronage appointments and the low value of sci- ence in universities forged his fight to correct these trends and in turn making his voice a (paid) sound of reason (and to think that he might have come to Canada to hold a post at the University of Toronto in 1851). He disliked the political and rich appointments to the few chairs in science by the old guard but felt that a position should be earned on merit and determi- 1996 nation. A philosophy that extends past the halls of academia. His fights, and there were many, spanned the spectrum of 19th century ideas from external poli- tics of science to evolution, human origins and human rights, religion, and education for the masses. All these were no where so cleverly, and clearly put than out of the mouth or with pen than of T. H. Huxley. We read or hear tales of the giants whose shoul- ders we stand on and at times, due to our own igno- rance, don’t even know whose shoulders they are. Huxley fought hard for the rights of the down trod- den, though many of these rights and ideas were not conceived by him no one enjoyed being in the wrong with him. His rival, Richard Owen, the “Cuvier of England” found that out the hard way when the issue of human origins and our species relationships were seriously, for the first time, examined. Owen, who wasn’t allergic to the idea of evolution, had a severe reaction to it when our species was dragged into the discussion. Huxley’s role as a scientist was two fold: not only illustrate to your learned colleagues what you are doing as a scientist but also open up to the public, no matter at what station in life they may be, and let them rejoice in your discovery, no matter how seemingly mundane. Make them believe that ideas are not strictly for the rich or powerful. That NEw TITLES Zoology *Amphibians of Oregon, Washington, and British Columbia: a field identification guide. 1996. By C. C. Corkran and C. Thomas. Lone Pine Publishing, Edmon- ton. 173 pp., illus. $21.95; U.S. $16.05 *Animal evolution: interrelationships of the living phyla. 1995. By C. Nielsen. Oxford University Press, Don Mills, ix + 467 pp., illus. $67.50. *Atlantic coastal birds. 1996. Edited by K. Kaufman. Peterson Flash Guides. Houghton Mifflin, New York. 24 panels. U.S. $7.95; $10.95. *Atlas of Saskatchewan Birds. 1966. By A. R. Smith. Special Publication No. 22. Saskatchewan Natural History Society, Regina. 456 pp., illus. $45. *Atlas of the distribution of fish within the Canadian tributaries of western Lake Superior. 1996. By W. T. Momot and S. A. Stephenson. Centre for Northern Studies, Lakehead University, Thunder Bay. 383 pp., illus. $30. *A birder’s guide to Trinidad and Tobago. 1996. By W.L. Murphy. 2nd edition. Peregrine Enterprises, Parkersburg, West Virginia. U.S. $15.95. The bird garden. 1995. By S. W. Kress. Dorling Kindersley, New York. 176 pp., illus. U.S. 24.95. Birding. 1995. By J. Forshaw, et al. Time-Life Books, Alexandria, Virginia. 288 pp., illus. U.S. $29.95. BOOK REVIEWS Vy coachmen and housemaids have the right to learn too. No one did it better. Unlike financially secure gentiemen of science like Darwin, Huxley was a man of his times, continually in debt, family obligations demanding time, finding a way to survive the personal pain of the loss of a child, and yet keep the strength and focus to swing the sword of the righteous. In some ways it was a differ- ent time than today. But in many ways things haven’t changed all that much. All the issues and feelings of Huxley and his time are still prevalent today. Desmond does not document the latter years of Huxley’s life (1871-1895) when a person’s seniority must bare witness for the new breed of bull dogs, sinners and saints. A transition that no one likes. By 1870 when the book ends, Huxley had succeeded, to the benefit of his country and science as a whole, in either initiating, guiding, or witnessing great change in a century that we are beginning to see only through the fog of time. Like Darwin, Desmond’s Huxley dissipates the fog of the mythical, and por- trays the real. At least as real as a hundred years of retrospection can serve up. TIM TOKARYK Box 163, Eastend, Saskatchewan SON OTO *Birds of the Besnard Lake area. 1996. By G. G. Bortolotti and K. Wiebe. Special Publication No. 20. Saskatchewan Natural History Society, Regina. 456 pp., illus. $12. *+Birds of Kenya and northern Tanzania. 1996. By D. A. Zimmerman, D. A. Turner, and D. J. Pearson. Princeton University Press, Princeton. 740 pp., illus. U.S. $65. Bombardier beetles and fever trees: a close-up look at chemical warfare and signals in animals and plants. 1996. By W. Agosta. Addison-Wesley, New York. vii + 224 pp., illus. U.S. $25. +A checklist of the birds of Chile. 1996. By S.N. G. Howell. American Birding Association, Colorado Springs. 31 pp. U.S. $4.95. +Ecology and conservation of wolves in a changing world. 1995. Proceedings of the Second North American Symposium on Wolves, Edmonton, 25—27 August, 1992. Canadian Circumpolar Institute, Edmonton. 620 pp., illus. +Encyclopedia of Canadian Fishes. 1996. By B. Coad. Canadian Museum of Nature, Ottawa. 928 pp., illus. $100. The fascinating world of beetles. 1995. By M. A. Julivert. Baron’s Hauppauge, New York. 31 pp., illus. U.S. $6.95. +The forgotten pollinators. 1996. By S. L. Buchmann and G. P. Nabhan. Shearwater Books. Island Press, Washington. xx + 292 pp., illus. U.S. $25. +Foundations of animal behavior. 1996. Edited by L. D. Houck and L. C. Drickamer. University of Chicago Press, 728 Chicago. xvi + 843 pp.. illus. Cloth U.S. $95: paper U.S. $34.95. *The garter snakes: evolution and ecology. 1996. Ani- mal Natural History Series volume 2. University of Oklahoma Press, Norman. xx + 332 pp.. illus. U.S. $65. Gorilla. 1995. By I. Redmond. Knopf, New York. 64 pp.. illus. U.S. $19. +Handbook of birds of the world. Volume 3: Hoatzin to Auks. 1996. Edited by J. del Hoyo, A. Elliott and J. Sargatal. Lynx Editions. Barcelona; illus. U.S. $175. Handbook of marine fauna of north-west Europe. 1995. Edited by P. Hayward and J. S. Ryland. Oxford University Press. London. 316 pp.. illus. Cloth £60: paper £29.50. Oxford University Press. Don Mills. Cloth $162: paper $79.95. *Hybrid ducks: a contribution towards an inventory. 1996. By E. and B. Gillham. Published by B. L. Gillham, Box 563, Wallington, Surrey. SM6 9DX. 88 pp.. illus. £16. Insects through the seasons. 1996. By G. Waldbauer. Harvard University Press. Cambridge, Massachusetts. Xili + 289 pp.. illus. U.S. $24.95. An introduction to invertebrate conservation biology. 1995. By T. R. New. Oxford University Press. Don Mills. 208 pp.. illus. Cloth $94.50: paper $47.50. Mammals of Australia. 1996. Edited by R. Strahan. Smithsonian Institute Press, Washington. 756 pp.. illus. U.S. $75. The octopus’s garden: hydrothermal vents and other mysteries of the sea. 1996. By C. vanDover. Addison- Wesley. New York xiv + 183 pp.. illus. U.S. $20. +Prairie night: black-footed ferrets and the recovery of endangered species. 1996. By B. Miller. R. P. Reading. and -S. Forrest. Smithsonian University Press. Washington. xvii + 254 pp.. illus. U.S. $34.95. Readings in animal cognition. 1996. By M. Beckoff and D. Jamison. MIT Press, Cambridge, Massachusetts. xv + 379 pp.. illus. U.S. $30. *Red fox: the catlike canine. 1996. By J. D. Henry. New edition. Smithsonian Institute Press, Washington. 174 pp.. illus. U.S. $15.95. *Red-winged blackbirds: decision-making and repro- ductive success. 1996. By L. D. Beletsky and G. H. Orians. University of Chicago Press, Chicago. c288 pp.. illus. Cloth U.S. $54: paper U.S. $21.95. Sea stars, sea urchins, and allies: echinoderms of Florida and the Caribbean. 1996. By G. Hendler. Smithsonian Institute Press, Washington. xi + 390 pp.. illus. U.S. $39.95. The Snow geese of La Perouse Bay: natural selection in the wild. 1995. Oxford University Press, Don Mills. xv + 297 pp.. illus. $106.95. The species composition of peracarid fauna (Crustacea, Malacost race) of the northwest water polynya (Greenland). 1996. By A. Brandt, S. Vassilenko. D. Pipenburg. and M. Thurston. Meddelelser om Gronland, Bioscience 44. 30 pp., DKK 80. THE CANADIAN FIELD-NATURALIST Vol. 110 7Studies of high-latitude seabirds, 4: trophic relation- ships and energetics of endotherms in cold ocean sys- tems. 1996. Edited by W. A. Montevecchi. Papers from the Marine Birds and Mammals in Arctic Food Webs Symposium, St. John’s, Newfoundland, 1992. Occasional Paper No. 91. Canadian Wildlife Service, Ottawa. 73 pp-. illus. Free. Views from the bridge: a memoir of freshwater fishes of Trinidad. 1995. By Julian S. Kenny. Rapid Environmental Assessments, 74 Ana Street, Port of Spain. Trinidad. U.S. $20. Watchable birds of the southwest. 1995. By M. T. Gray. Mountain Press. Missoula. ix + 187 pp.. illus. U.S. $14. 7 Whitetail spring. 1996. By J. Ozoga. Seasons of the Whitetail Book 3. Willow Creek Press. Minocqua, Wisconsin. 144 pp.. illus. U.S. $29.50. Wild bears of the world. 1996. By P. Ward and S. Kynaston. Facts on File, New York. 192 pp.. illus. U.S. $25.95. The wisdom of the hive: the social ecology of honey bee colonies. 1996. By T. D. Seeley. Harvard University Press, Cambridge, Massachusetts. xiv + 295 pp., illus. U.S. $49.95. Botany Agnes Chase’s first book of grasses: the structure of grasses explained for beginners. 1996. By L. G. Clark and R. W. Phol. 4th edition. Smithsonian Institute Press, Washington. 176 pp.. illus. U.S. $16.95. Caliciales. 1996. By L. Tibell. The New York Botanical Garden, Bronx. 79 pp.. illus. U.S. $20. Conifers: the illustrated encyclopedia. 1996. By D. M. van Gelderen and J. R. F. van Hoey Smith. Timber Press, Portland, Oregon. 2 volumes, 706 pp.. illus. U.S. $125. +Ecology and management of Sitka Spruce: emphasiz- ing its natural range in British Columbia. 1996. By E. B. Peterson, N. M. Peterson. G. F. Weetman. P. J. Martin, and W. Stanek. c240 pp.. illus. $65. Ecology of the southern conifers. 1996. Edited by N. J. Enright and R. S. Hill. Smithsonian Institute Press, Washington, 352 pp.. illus. U.S. $60. Gladiolus in Africa: systemics, biology, and evolution. 1996. By P. Goldblatt. Timber Press, Portland, Oregon. 256 pp.. illus. U.S. $39.95. How to draw plants: the techniques of botanical illus- tration. 1996. By K. West. Timber Press, Portland. Oregon. 152 pp.. illus. U.S. $19.95. +Leaf venation patterns, volume 7: the classification of leaf venation patterns. 1995. By E. P. Klucking. J. Cramer, Stuttgart. 337 pp.. illus. DM 270. Nevada vascular plant types and their collectors. 1996. By A. Tiehon. New York Botanical Garden, Bronx. 103 pp. Juncaceae. 1996. By H. Balslev. Flora Neotropica Volume 68. New York Botanical Garden, Bronx. 163 pp.. illus. U.S. $29.50. 1996 Orchids of Kenya. 1996. By J. Stewart. Timber Press, Portland, Oregon. 144 pp.. illus. U.S. $55. Palms throughout the world. 1996. By D. L. Jones. Smithsonian Institute Press. Washington. 410 pp.. illus. USS. $49. *Plant collecting for the amateur. 1996. By T. C. Bray- shaw. U.B.C. Press. Vancouver. 44 pp.. illus. *Plants of southern interior British Columbia. 1996. Edited by R. Parish, R. Coupé. and D. Lloyd. Lone Pine, Edmonton, 463 pp., illus. $24.95: U.S. $19.95. *Plants of the western boreal forest and aspen park- land. 1995. By D. Johnson. L. Kershaw, A. Mackinnon, and J. Pojar. Lone Pine, Edmonton. 392 pp., illus. $24.95. Silk tree, guanacaste, monkey’s earring: a generic sys- tem for the synadrous Mimosaceae of the Americas, part 1: Abarema, Albizia, and allies. 1996. By R. C. Barnaby and J. W. Grimes. New York Botanical Garden, Bronx. 292 pp., illus. U.S. $45. Trees of the northern United States and Canada. 1995. By Iowa State University Press, Ames. x + 502 pp.. illus. U.S. $39.95. Trees of Nova Scotia: a guide to native and exotic species. 1996. By G. Saunders. Nimbus, Halifax. 112 pp.. illus. $9.95. 7 Wildflowers: legends, poems, paintings. 1996. Reprint of 1989 edition. By N. R. Ranson and M. J. Laughlin. Texas A & M University, College Station xii + 82 pp.. illus. Environment +The abstract wild. 1996. By J. Turner. University Arizona Press, Tuscon. c180 pp. Cloth U.S. $32.50: paper U.S. $15.95. Agricide: the hidden farm and food crisis that affects us all. 1996. By M. W. Fox. 2nd edition. Krieger. Melbourne, Florida. U.S. $33.50. yAn ancient forest of the Lake Temagami site region (4E). 1996. By P. Quinby and T. Lee. Canadian Nature Federation, Ottawa. 72 pp., illus. + map. $10. yBalancing act: environmental issues in forestry. 1996. By H. Kimmins. 2nd edition. UBC Press, Vancouver. c272 pp.. illus. $29.95. 7The biology of rocky shores. 1996. By C. Little and J. A. Kitching. Oxford University Press, London. 272 pp.. illus. Cloth £35; paper £14.95. Oxford University Press. Don Mills. Cloth $94.50; paper $40.50. +The Boundary Waters wilderness ecosystem. 1996. By M. Heinselman. University Minnesota Press, Minneapolis. 384 pp., illus. U.S. $24.95. Bring back the buffalo: a sustainable future for Amer- ica’s great Plains. 1996. By E. Callenbach. Island Press, Washington. 280 pp., illus. U.S. $22.50. Dinosaurs, diamonds, and things from outer space: the great extinction. 1995. By D. B. Carlisle. Stanford Uni- versity Press, Stanford. viii + 241 pp., illus. Cloth U.S. $45; paper U.S. $17.95. BOOK REVIEWS 729 Discover America: the Smithsonian book of the nation- al parks. 1996. By C. E. Little. Smithsonian Institute Press, Washington. 224 pp.. illus. U.S. $34.95. 7Ecological identity: becoming a reflective environmen- talist. 1995. By M. Thomashow. MIT Press, Cambridge. Massachusetts. xxii + 228 pp.. illus. U.S. $12.50. *Ecology of a polar oasis: Alexandra Fiord, Ellesmere Island, Canada. 1996. Edited by J. Svoboda and B. Freedman. Captus Press, North York. Ontario. 286 pp. $38.50. +Evolution and environment in tropical America. 1996. Edited by J. B. C. Jackson, A. F. Budd. and A. G. Coates. c400 pp.. illus. Cloth U.S. $65: paper U.S. $27.50. Far from tame: reflections from the heart of a conti- nent. 1996. By L. Allman. University of Minnesota Press, Minneapolis. 176 pp.. illus. U.S. $16.95. *Field guide to ecosites of the mid-boreal ecoregions of Saskatchewan. 1996. By J. D. Beckingham, D. G. Nielsen, and V. A. Futoransky. U.B.C. Press, Vancouver. xv + 446 pp.. illus. *Field guide to ecosites of northern Alberta. 1996. By J. D. Beckingham and J. H. Archibald. U.B.C. Press. Vancouver. xiv + 515 pp., illus. + map. +Field guide to ecosites of Southwestern Alberta. 1996. By J. H. Archibald, G. D. Klappstein, and I. G. W. Corn. U.B.C. Press, Vancouver. xiii + 505 pp., illus. + map. $29.95. yField guide to ecosites of west-central Alberta. 1996. By J. D. Beckingham, I. G. W. Coms, and J. H. Archibald. Canadian Forestry Service Special Report 9. U.B.C. Press, Vancouver. xiv + 622 pp., + map. Floods of fortune: ecology and economy along the Amazon. 1996. By M. Goulding, N. J. H. Smith, and D. J. Mahar. Columbia University Press, New York. vi + 193 pp. illus. U.S. $29.95. +The food web of a tropical rain forest. 1996. Edited by D. P. Reagan and R. B. Waide. University Chicago Press. Chicago. 11 + 616 pp., illus. Cloth U.S. $110: paper U.S. $39.95. Imperiled waters, impoverished future: the decline of freshwater ecosystems. 1996. By J. N. Abramovitz. Worldwatch Institute, Washington. 80 pp.. illus. $5. “In search of nature. 1996. By E. O. Wilson. Island Press, Washington. x + 214 pp., illus. U.S. $19.95. The magic ground. 1996. By H. Horwood. Nimbus, Halifax. 196 pp., illus. $16.95. Marine biology: function, biodiversity, ecology. 1995. By J. S. Levinton. Oxford University Press, London. 448 pp.. illus. £21.50. Oxford University Press, Don Mills. $71. +The marshes of southwestern Lake Erie. 1995. By L. W. Campbell. Ohio University Press, Athens. xii + 233 pp.. illus.