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Rr TANNOY fray cab } ae eke ETE ne ie ain) Ca) xs Meese atel ahd . oN 4 at ie Rey | The CANADIAN 4% FIELD-NATURALIST Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada Voi. XXXII, No. 5. NOVEMBER, 1918. Zz & oS < GER ae UN ox \ VE RRR NRE SS PO SOSA EA) ‘S S/F N= | ey . ~~: | EY, | By = | yy 1 : % wu yd, * . a ransaclions * Pe. oppaws, CADADA- rin ht and Publishing Company: Ginae Printing ae oF 4855. Ae hid | te es 4 ‘ Wi A i } : hi i ISSUED DECEMBER 18, 1918, Entered at Ottawa Post Office as second class matier. ) A %) ek ee eee ees taf i { i X (\ ey e I Centennial Year Volume 93, Number 1 January-March 1979 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Madame Jules Léger The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things. The Members of Council are listed on the inside back cover. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors C. D. Bird A. J. Erskine David P. Scott E. L. Bousfield Charles Jonkel Stephen M. Smith Francis R. Cook Charles J. Krebs Robert E. Wrigley George H. La Roi Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang Production Manager: Pauline A. Smith Business Manager: W. J. Cody Subscriptions and Membership Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The Canadian Field- Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879- 1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 4J5 Book Review Editor: Dr. J. Wilson Eedy, R.R. !, Moffat, Ontario LOP 1J0 Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon, USA 97731 (address valid until August 1979). Address manuscripts on birds to the Associate Editor for Ornithology: Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick E0OA 3C0 All other material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0 Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231- 4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461). Cover: Since the Club was founded in 1879, it has published a journal under three successive names. Covers bearing the two names preceding the present one are illustrated. Left, Ortawa Field- Naturalists’ Club, Transactions, published for 1879 to 1886. Right, The Ottawa Naturalist, published from 1887 to 1919; it bore this cover design only between April 1918 and March 1919. THE CANADIAN FIELD-NATURALIST Volume 93 1979 CENTENNIAL YEAR THE OTTAWA FIELD-NATURALISTS’ CLUB OTTAWA CANADA iF ’ ae The Canadian Field-Naturalist Volume 93, Number | January-March 1979 One Hundred Years in Perspective — the Changing Roles and Objectives of The Ottawa Field-Naturalists’ Club ROGER A. FOXALL President, The Ottawa Field-Naturalists’ Club, January 1977-January 1979 Following an invitation to members of the Ottawa Literary and Scientific Society, “fully forty gentlemen attended the meeting held on the 19th March, 1879. After a lengthy discussion as to the form the organization should take, the Ottawa Field-Naturalists’ Club was born.” The quotation above is from the first historical sketch of the Club, taken from the 1880 Annual Report when the Club was fully one year old. Since then there have been several articles describing the many memorable people and events that add up to the varied and fascinating 100-year history of our Club. A series of historical articles is currently being published in Trail & Landscape. The aim of this message is not to provide another summary of people or events, but to review instead the ways in which the Club’s functional role and objectives have changed over time and to present a personal view of our present role and objectives in terms of future needs. In the beginning, the objective of the Club was stated simply as “the study of the Natural History of this locality.” The raison d’étre, however, was more explicit in an 1884 summary: “The value of these studies cannot be over-estimated for, from their very nature, they are most beneficial both to mind and body, while, from an economical point of view they are no less important . . . . The future of Ottawa is in large measure dependent upon the development of the vast stores of minerals contained in the surrounding districts, and which are as yet almost unexplored. The manifold uses and demand for all vegetable products make it necessary that the nature and habits of plants should be closely studied, as well as those of the countless myriads of insect foes which deprive the cultivator of so much of his hard-earned gains... . It gives very great satisfaction to the members of the Council to find that the Club is being gradually recognized as a source of reliable reference on all these matters.” So the objective in part was the study of natural history to assist the economical development of the Ottawa area. Times have indeed changed! A specific example of success in this direction was the advice given to the Ottawa Granite Company that within a few miles of the city there existed an outcrop of quartzite deposits just as good as quartzite then being imported from New York State. This advice led to successful commercial exploitation of the deposits. Although economic development of the Ottawa area was one aim of the Club, another was to popularize the study of the different branches of natural history and thereby increase appreciation of nature. On the local scene this entailed programs of organized studies, excursions, lectures, and generally spreading the good word. By the thirtieth anniversary, Club membership had increased from the initial 40 to over 300. Increasingly, however, the Club’s successes led to recognition throughout the Dominion, and its role began to broaden from that of a strictly local club towards 2 THE CANADIAN FIELD-NATURALIST Vol. 93 that of a national society. In an 1893 The Ottawa Naturalist, the editor found it necessary to state, “Although the scope of the Club’s work has been gradually widened to receive the benefits of investigations made by its members wherever they may be located, the special work for which it was organized must stili merit the chief attention, and although much has been observed and recorded of the Natural History of Ottawa, there still remains vastly more to be done.” But the transformation accelerated! A series of 42 nature study articles was written for The Ottawa Naturalist by leading Club members, and between 1903 and 1908 a total of 238000 separate copies was distributed to schools and colleges across Canada. Such was the success and expanding influence of the Club that Ottawa’s Evening Journal (although perhaps a little biased) found it appropriate in 1908 to state that “today The Ottawa Field-Naturalists’ Club is the greatest institution of the kind in the entire continent of North America.” Partly because of the Club’s campaigns and the travels of Club members, but mainly because of a general increasing interest in nature, similar organizations were established in several cities across Canada. Increasingly, the need developed for a publication covering the natural history of all of the Dominion, until in 1918 the Foreword to the last volume (32) of The Ottawa Naturalist announced: “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 standing, seems a logical nucleus from which such a publication should be developed.” The first issue of The Canadian Field- Naturalist was published in April 1919. This event in the Club’s 40th year marked the end of a lengthy transition and formalized the dual role of the Club — a local natural history club and a national society. Somewhat earlier, in October 1912, the objectives of the Club had been expanded significantly to reflect the two roles: To foster an acquaintance with and a love for nature; to study especially the natural history of the Ottawa District; to encourage investigation and to publish the results of original research in all departments of natural history; to arrange for out-of-door excursions during the summer months; to provide free lecture courses during the winter months; and ina general way to render assistance to students or others interested in Nature Study. With one significant exception, to which I will refer below, these objectives remained unchanged until 1972. Unfortunately the success of the Club’s first 40 years was to be followed by a 30-year period during which many difficulties were encountered. Until about 1923, the Club received a grant from the Ontario Department of Education that contributed significantly to the sustenance of The Ottawa Naturalist and later The Canadian Field-Naturalist. \ronically, post-war economies led to cancellation of the grant when it was needed most, for the creation of The Canadian Field- Naturalist placed a significant burden of responsibility on the Club and all its members. Without the grant, many more members were needed to provide financial stability. But the growth was too slow. In 1934 P. A. Taverner, Chairman of a Special Committee on Membership, issued an appeal to all Canadian naturalists to boost membership. A reduction in size, from 24 to 16 pages an issue, and other parsimonious economies had been tried, but still the year-end balance was negative. The journal survived, of course, but for many years the fight for The Canadian Field-Naturalist’s survival represented the principal objective of The Ottawa Field-Naturalists’ Club. The difficulties in maintaining the national role were paralleled by difficulties with the local role. Here, analysis is more difficult. We can perhaps rationalize the decreasing enthusiasm in terms of two gradual developments. The early years of systematic, detailed studies of local natural history must have represented a frontier-like challenge — new knowledge was easily acquired and each discovery, be it a new species of plant, insect, bird, or butterfly, contributed to the developing picture. But the acquisition of new knowledge makes further discoveries just that much more difficult. Members would have to travel further and look harder, and greater expertise was needed. But why 1979 FOXALL: PRESIDENT’S MESSAGE 3 try so hard? The initial objective, to assist in the economic development of Ottawa, was surely less important now. The urgency of acquiring knowledge in support of conservation and preservation to slow down development was not yet recognized. For whatever reasons, it is a fact that emphasis on studying the local natural history declined to the point that in 1947, the objective “to study especially the natural history of the Ottawa District” was removed from the Constitution. Although excursions and lecture programs continued to foster and sustain members’ interests in local natural history, the membership grew slowly and even declined for several years. In retrospect it seems likely that the Club’s surviving the difficult 30 years can be attributed to its dual role. For without the local membership, The Canadian Field-Naturalist might have failed financially, and without the responsibility of continuing The Canadian Field-Naturalist, the local club might have failed for lack of sufficient motivation. The last 30 years, 1949-1979, have also been characterized by significant changes, but the story is a happier one. The Canadian Field- Naturalist has steadily developed into a scientific journal of both national and international repute — a journal almost unique in the world for its breadth and standard of scientific natural history reporting. The content has evolved also. The studies reported have progressively become more sophisticated as new knowledge requires increasingly more detailed examination of natural phenomena. Other, less formal journals and club magazines are now published and these provide media for reporting studies and findings of more local importance. There is no doubt that these evolutions have resulted in The Canadian Field- Naturalist becoming of lesser interest to many local members. But the need for The Canadian Field-Naturalist is at least as great today as it was in 1919, and, through its continuing support, The Ottawa Field- Naturalist’s Club is playing a very valuable role in the reporting of Canadian natural history. The continuing decrease of Ottawan content in The Canadian Field- Naturalist and the need to regenerate the local Club, led to the start, in 1949, of a Club Newsletter for local members. Steadily the pendulum swung back as successive Presidents and Councils worked very hard to rebuild. It was not easy, as illustrated by the anguished appeals in the Newsletter of Chairmen of Excursions and Lectures urging members to attend the many walks and talks that were being arranged. But the rebuilding succeeded and by the late 1960s the local Club had regained much of the status and drive of the early period. As a Canadian Centennial project, the Club upgraded the Newsletter to become the magazine Trail & Landscape. For the last eleven years, the staff of Trail & Landscape has succeeded in producing an excellent, informative magazine that focusses on local sightings, events, and issues. Sometimes serious, sometimes lighthearted, but always highly readable, Trail & Landscape plays an important role in generating and maintaining the enthusiasm of local members. Somewhat surprisingly, in retrospect, natural history organizations throughout North America were slow to recognize that man’s rapid development and exploitation represented a serious threat to all natural environments and living things. In the beginning, The Ottawa Field-Naturalists’ Club had ~ worked in support of economic development. When, later, favorite areas such as Dow’s Swamp were taken from us, no organized howls of protest were heard. Such things were taken for granted as a normal result of development. But by the mid-1960s organizations everywhere recognized the threat to what had seemed before to be an apparently infinite environment. The battle began to preserve for future generations those areas that had so far survived exploitation, to control thoughtless pollution, and to prevent the indiscriminate use of chemicals that threatened wildlife. In 1972, the Club’s objectives were rewritten to reflect the new motivation: To promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible: to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things. On the national scene, the new emphasis is being supported by the publication in The Canadian Field- Naturalist of, for example, excellent papers on the effects of chemicals such as PCBs in the environment, and the internationally acclaimed issues on the status of the Peregrine Falcon. The 4 THE CANADIAN FIELD-NATURALIST Vol. 93 local Club established a Conservation Committee of Council and became heavily involved in the battles to defend the Mer Bleue and Gatineau Park, and more recently to defend many other natural areas of the Ottawa-Carleton Region. Today, the Club continues its dual role, publishing The Canadian Field- Naturalist on behalf of all Canadian naturalists, and performing the functions of a typical local natural history club — arranging excursions and lectures, publishing a local magazine, and attempting strenuously to protect from further development many areas of significance to natural history. A successful one hundred years? Certainly! The Club’s founders would surely be pleased with what transpired from that first meeting on 19 March 1879. The Club has evolved to meet the needs of naturalists over a period of immense change in Canada, thousands of people have benefited directly from their association with it, and many more thousands have benefited indirectly from its achievements. More could have been done, particularly on the local front, for it is surely ironic that a club formed one hundred years ago to study Ottawa’s natural history has been forced to scramble hurriedly during the last few years trying to acquire sufficient knowledge of local areas (some only ten miles from Parliament Hill) to support their preservation and conversation. Although the celebration of our Centennial Year will be largely retrospective, some time should be spent thinking about and discussing our current roles and objectives in terms of future needs. Some predictions can be made with a reasonable degree of confidence. These are: — the need for The Canadian Field- Naturalist, or an equivalent journal, will continue into the foreseeable future: — the pressure on natural environments and living things will continue to increase, in the short term at least; — the potential for increasing public interest in and concern for natural history will increase as the amount of leisure time continues to increase: and — in the longer term, leisure activities, if uncontrolled, will become a serious threat to natural environments. Let us now examine our objectives in these terms. The second objective, “to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible,” is certainly appropriate. Although several local Club members have expressed the opinion that The Canadian Field- Naturalist has become too “professional” or too “scientific” to be published by a club largely made up of amateur naturalists, I remain convinced that the Club should continue to publish The Canadian Field-Naturalist on behalf of all Canadian naturalists at least until it is demonstrated that another organization could provide it a better home. Similarly, the third objective, “to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things,” will continue to be appropriate. Our recent performance in support of this objective has not been outstanding and the Club should concentrate on becoming more involved with other organizations. To attempt to do this effectively on a national scale would be unrealistic, but more could and should be done to promote effective cooperation with organizations in Ontario and Quebec. The first objective, “To promote the appreciation, preservation and conservation of Canada’s natural heritage,” sounds impressive but is as a separate objective, the least realistic for The Ottawa Field-Naturalists’ Club today or in the future. We do support it to some extent by publishing The Canadian Field-Naturalist and by cooperating with other organizations, but some modification of the objective appears necessary if the Club is not to pretend it is trying to duplicate the function of a large national conservation organization such as the Canadian Nature Federation. This is not to say that I feel The Ottawa Field-Naturalists’ Club should become overly parochial, but I do think that the stated objectives of any organization must represent specific, realistic challenges that can be pursued effectively. Finally, | would urge the Club to reintroduce the objective “to study especially the natural history of the Ottawa District,” with this or some similar wording, to reflect the localized nature of most of 1979 FOXALL: PRESIDENT’S MESSAGE 5 our activities and the need to continue increasing our knowledge of the district. During the conservation battles of the last few years, local planning bodies have come to recognize the Club as the authoritative source for much of the data on which to base decisions affecting the natural environment. This recognition is due entirely to the hard work of relatively few members. Past appeals to members to help in this endeavor by turning out for inventory-type excursions have largely failed. If the Club is to succeed in meeting the challenges for the future based on the predictions listed above, it must surely focus again on the original objective, chosen on 19 March 1879 — “the study of the Natural History of this locality.” Biology — The Unknown Science? YORKE EDWARDS British Columbia Provincial Museum, Victoria, British Columbia V8V 1X4 Biology is a science largely unknown to the people of the world. If life is the most important “thing” on Earth, this is a strange condition. The disturbing result is that not only is the man in the street largely ignorant in all things biological, but also that decisions on how man will treat our life- support system — Planet Earth — are made without biological understandings. Biology is largely missing as general knowledge, hence also as an automatic aid to intelligent decisions. When, on occasion, biology does happen to be given its day, there is no background of accepted status to give it weight. Biology has little hope of being valued or even understood if it is to surface in public affairs only at moments of crisis. Scientists do a remarkably good job of keeping the public uninformed. This is partly because the public is rather narrow-minded about what is important, which creates some reluctance on the part of scientists to become too visible. But then, of course, a more informed public might be more understanding. When we add to this picture the high proportion of scientists unable to be very informative about their work to almost everyone, the reasons for science’s isolation from the man in the street are clear and appear to be formidable. The poor record of scientists being intelligible and convincing in public seems to be programmed that way by the nature of both today’s science and today’s scientists. Science today is largely concerned with exploring smaller and smaller bits of our world, with things beyond the awareness of everyday human experience. Scientists lead this trend, but few are helping to put the bits together into more understandable wholes. It is the era of the specialist. By contrast the generalist in science has become rare, which I am convinced endangers both science and the world. The generalist in science is a necessary companion to the abundant specialist who is traditionally learning “more and more about less and less.” The need is not just to assemble what we know about the world into forms meaningful to many, it is also a matter of possessing the resulting wisdom necessary for survival. Man’s wildly compounding knowledge has him more and more in control of his support system Earth in an accelerating use of this control that is quite out of control. Science feeds the largely destructive trend. Few people, including most scientists, can glimpse what is really happening, for the quality and quantity of the effects are beyond the sensory capacities of most people. Understanding, acquired from those few who do understand, is the only hope of self control, understanding that comes from broad insights into the world and how it functions. Science must do a much better job of arranging “more and more of less and less” so that bit plus bit becomes a somewhat visible, somewhat accurate, somewhat understandable whole. Many scientists consider the discovery of truth to be their sole justification. For example, they regard the basic discoveries in atomic physics as standing alone; atomic warheads are therefore other men’s burdens, not those of the pure scientists. This assumes it not true that achieving atomic fission made Hiroshima’s devastation probable, even inevitable in the political climate of the time. In most respects all of us have minds not much changed from those of ancestors in Ice Age caves. The products of science available to us all are power in the hands of ignorance. If science put the power there, and continues to do so without adding very effectively the controls of understanding, we are surely building systems for our own destruction. Only science can correct this situation. We need a new morality in the sciences, which would result in major efforts to make science and its influence on Earth understandable to those outside the ivory towers, and to those in them too, for most scientists do not understand most science. There is a modern myth in our culture that the scientist is among the most brilliant and wise of men, understanding the world as few others do. Some scientists believe this. But there are many 6 1979 EDWARDS: GUEST EDITORIAL 7 people who do not, and in fact, from his specializations, and from his frequent mental and physical isolation from the worlds that most people experience, there is an emerging image in our society of the scientist being rather dangerous. Inevitably, dedicated specialization results in tunnel vision. Neither the specialization nor the resulting narrow view of the world are by any means confined to scientists in our society, nor is the trend for these many specializations to be increasingly specialized. But they are relatively unimportant outside of science. It is specialization within science itself that can keep people awake at night. Science has given us formerly unbelievable powers and capabilities at the price of mushrooming humanity in a disintegrating support system. If it is not already too late, the end of irresponsible science is the only cure, and then only if irresponsibility is replaced by responsibility, not just for tomorrow’s new problems but for yesterday’s legacy of old ones as well. It is fashionable to say that “It was the side-effects that got us.” There can be no clearer evidence of our narrow views. “Side-effects” are a tunnel-vision illusion, for the broad view will see only BCILCCtS: I have begun this essay with the broadest possible sweeps quite by plan. My resulting generalizations contain many of the “part truths” and “too general” sorts of “facts” that drive most scientists into ever deeper specializations with their attractively simple foci of study. In disciplines properly requiring the greatest possible degrees of accuracy, the uncontrolled drift of scientists toward the high accuracy areas is perhaps predictable. But I wonder if this is both science’s blind alley, and Earth’s sentence to increasing impoverishment of its life. While the sciences choke on ever increasing flows of new detail from examining the parts of the world, not many scientists are putting the pieces together into meaningful wholes of knowledge. We need more syntheses by more generalists, more generalization even at the calculated risk of less accuracy. The need is not so much for fewer specialists as it is for more generalists, not so much for less publishing of science as for more interpretation of science where there is recognition of the value of generalization that may be accurate enough to be of great value, while also being uncertain and perhaps therefore inaccurate to some degree. Conquering the tsetse fly “problem” in Africa is receiving much research. The tunnel vision approach would focus simply on eradicating flies as a relatively simple biological problem. The generalist might be aghast at such efforts in view of the vast ignorance of what the power to destroy the tsetse would really accomplish through chain reaction and time. Tunnel vision sees only flies, with perhaps another vision of people happily eating more cattle. A broader view might see over a quarter of a continent an end result of human poverty and misery in rapidly expanding deserts that mark the end of rich faunas and floras once containing reasonably successful people. The innocence | of dropping sucha powerful force as easy tsetse eradication into the lap of a humanity largely blind to the terrible powers involved has many parallels in history, where seemingly small additions to the capability of man has resulted in world changes too big for most people to see, and too drastic for those touched by it to understand. Science has rarely been in the arena of public debate, expounding its discovered truths. Without science participating in human affairs with a vigor equal to its unique capability for discovery, what hope is there for wise choices? What hope for future life in high quality environments? Not always do the issues have the potential inflation factor of tsetse flies or nuclear fission. But the effects of lesser tinkerings add up, with overharvesting here, mercury waste there, disease eradication somewhere else, each with their chain reactions. Not long ago I read an enthusiastic article froma Canadian botany department assessing the harvest potential from the kelp growing along the British Columbia coast. A new industry was about to be born, and the facts were proclaimed as if from the local Chamber of Commerce. On impulse, I penned a note to the author asking to what extent the intended harvest would be also habitat removal to the rich and specialized fauna of the kelp “forests,” and what animal species were involved. The reply was a refreshingly candid admission that the thought had not occurred to him. Tunnel vision was perhaps quite properly his view of the kelp beds, 8 THE CANADIAN FIELD-NATURALIST Vol. 93 but there seemed to be no other view involved. No one had considered effects and chain reactions. I work in one of Canada’s larger museums with specialists in history, zoology, botany, ethnology, archaeology, and other fields. When I first examined this small community of mixed disciplines, I was impressed by the isolation from the others in which each group worked; but I became more impressed later on those rare occasions when cross-communication between two of them triggered an exciting and enlightening experience for both. Acommon language of biology bound together the life sciences, and an ecological dimension common to biology and anthropology gave them common ground from which to exchange ideas. But history was different; for traditional reasons it was unable to comprehend either biological or ecological concepts, so was unprepared to consider them of use. This was the most impressive museum experience ofall, and I have tried to understand it, not just for reasons of improving a museum, but for its possible deeper meanings. The new and popular interest in ecology has not touched the historians about me, at least not as historians. To converse with them I must drop my ecological view of the world and lean heavily on my everyday experiences with people interested simply in experiences with people. This sort of superficial observation of the human scene has been man’s need and entertainment since man became man, perhaps before, and it seems that traditional history as a discipline is largely a formalized version of this trait. A scientist might conclude that attempts to explain history without using the knowledge of biology and ecology must result in very limited understandings. A historian may counter that to record the passing scene 1s enough; let others try to explain it. The science-less view of history must be largely the view of the world as seen by most people. There is little concern with cause and effect, the only part of the world worth noting being people. Their successes and failures, if explained at all, result from their own interactions, chance, and the supernatural. Modern history of course has its ecologically oriented historians, or so the literature would suggest unless they are instead scientists trespassing in history. Nevertheless, science is not a useful part of traditional history, not to the extent of its being a constant means of understanding the past. This ignoring of man’s supreme accomplishment in recent times — scientific knowledge — in understanding the world is the standard human condition. We gladly accept the wealth and ease that science has given us. We eagerly learn to apply the directions of science for achieving health and comfort and riches from our environments. We spray the aphids on a bush, and thereby unknowingly poison a hundred other kinds of creatures, including ourselves. We pour chemicals of unknown capability into a sewer, for safety, and thereby take food in the form of fish from the mouths of a thousand people. We remove a rich forest with machines from an endless list of discoveries, and for quick profit burden future generations with square miles of sun-baked wastelands. To learn to exterminate a fly may be, unknowingly, to learn how to create deserts over half a continent. We are given power far greater than we realize, far more given to unexpected chain reactions than we can guess, by a science not doing the important part of its job. The brilliance that gives inventions and discoveries must come from a system discovering and disseminating also some understanding of the impact of those inventions and discoveries upon the face of Earth. The world needs generalists in science, using new approaches in science, to discover the true meanings of its discoveries, and to report them in useful form giving the sure, the probable, and the possible effects of their use. Only such synthesizers, at all levels of science, can scientifically reconstruct whole animals, rebuild whole habitats and whole landscapes, putting aggregates of knowledge back into the public reach so that whole things, or at least meaningful parts of things, are understood as far as men understand them. This is not easily done. That is why science fails to do it. By the standards of accuracy demanded in most sciences these reconstructions will have some inaccuracy. If this is unavoidable, it must become acceptable insofar as its accuracy is useful, because there is not just a general intellectual need for non-scientists to see more sums of knowledge, there is some urgency since our science and technology seem to have us programmed in several ways to self- destruct. The challenge of evolving effective methods for such a prevalent new dimension to science may 1979 EDWARDS: GUEST EDITORIAL 9 not be the main problem. Present scientific circles frequently regard as somehow second class those of its scientists that put much effort toward being understood in lay society. It is almost as if there was a cult for obscurity. Perhaps we have a long way to go. Science must of course be encouraged to pursue the obscure. But let it also learn to reassemble the parts. Biology is not presently a force in world affairs because it does not communicate well, at least not at the level of public understanding. Economists (and merchants, and lawyers, and others) make decisions instead. Until biologists begin to replace economists because they have a superior ability to correct errors and to predict events (which should not be too difficult) there is not much hope for realistic decisions in human affairs. If life is indeed Earth’s most important attribute, biology is then the discipline best able to guide man’s decisions concerning his (and Earth’s) welfare. If this is even partly so, we need a biology not Just active in knowing more of less and less, but also at knowing more of less plus less. Why? So that people may understand their world. And so that humanity may survive with dignity. The Canadian Field-Naturalist — the Status Quo or a New Direction? To some of us the most significant and lasting contribution that The Ottawa Field-Naturalists’ Club has made to society is the publication of The Canadian Field- Naturalist. From almost the very beginning, since 1880, the club has published a scientific journal and from the early years (1889 onwards) its scope has included papers on the natural history of Canada. This centennial year of the club is a time for contemplation, reflection, and, by the express wishes of Council, a reassessment of The Canadian Field-Naturalist’s réle as a publication of the club and its relationship to the club. Do Problems Exist? In general, the present members of the Council of The Ottawa Field-Naturalists’ Club are reported to be “uncomfortable” about The Canadian Field-Naturalist. Some would prefer more papers by amateur naturalists, more items of local interest, more club content, and less emphasis on professional scientific papers. Some, reflecting comments made by local members, question why a portion (40% in 1978) of their membership fees should be allotted to a publication that appears to have minimal relevance to the club and why those members who are not interested in The Canadian Field- Naturalist should receive it at all. Fortunately, others understand that The Canadian Field- Naturalist is providing a much needed service to Canadian natural history. Most people agree, however, that the club is the most appropriate publisher of The Canadian Field- Naturalist, at least until a more suitable alternative organization is evident. There is no doubt that the relationship between The Canadian Field- Naturalist and The Ottawa Field-Naturalists’ Club as its publisher is a very important one. Unfortunately, however, uncertainty regarding this relationship continues to arise from time to time. Ina way the publication ofa national scientific journal by a basically local natural history club is an anomaly, especially as The Canadian Field- Naturalist is recognized as the “official” publication of the Ottawa club. It is the only publication specifically mentioned in the club’s constitution and is sent to all club members. Until recent years many of the members of Council were senior scientists (the Ottawa area with many federal government departments has a high concentration of scientists) with strong interests in the journal. They recognized its broad scope and scientific significance. Today most Council members are not professional field-research scientists, hence the composition of Council is probably more comparable to those of other local natural history societies. Many of the professionals (mainly field-biologists) who were once active members of the club and served on its Council are now involved in different activities. Therefore, is it unexpected that some members of the present Council don’t really understand The Canadian Field- Naturalist? Their uneasiness about it is evident in their current re-evaluation of its rdle and in the stated view of the Chairman of the Publications Committee that the club should publish a journal that is in keeping with the wishes of the membership. But exactly what is meant by “membership”? Is it considered to be the “local” (arbitrarily defined geographically), largely non-professional members, the people who can most easily communicate their opinions to Council? Or does it also include, as it should in the true sense, the local and non-local professionals; the former local members who have moved away but still want to keep in touch with the club; the distant people who became members because they wanted to receive The Canadian Field-Naturalist (prior to 1976 this was the only way an individual could receive the journal); and possibly others who, although they may dwell far from Ottawa, wish for other reasons to belong to the club? And shouldn’t the opinions of our subscribers and other readers also be considered? The members of Council are voted into office at the Annual Business Meeting. The slate drawn up by a Nominating Committee each year from the local members of the club has always been acclaimed into office by the local members who attend. To my knowledge there has not been a single club election in recent decades. Considering that the club’s Council is assumed to represent the 10 1979 SMITH: EDITORIAL Ui membership, one can interpret the “wishes of the membership” to be equivalent to the wishes of the Council members after they have taken into account the expressed opinions of local members. In forming his/her opinion will each member of Council think of his/her own personal preferences or will he/she consider the broader outlook, recognizing The Canadian Field-Naturalist’s role in the advancement of scientific knowledge of Canadian natural history? And is recognition being given to all those the journal now serves beyond the interested local members, namely the subscribers, external members, authors, and the many readers who consult it in libraries? The major reasons for my editorial then are: first, to alert those concerned about and interested in The Canadian Field- Naturalist to be informed of the present uneasiness of the Council and the steps it is planning to take; second, to solicit informed input from our readers to help the Council decide whether the status quo should be maintained or The Canadian Field- Naturalist should take a new direction; and third, to state my personal opinion regarding the journal’s present status and réle and my thoughts regarding its future. Functions of The Canadian Field-Naturalist To me The Canadian Field- Naturalist fills the important need fora national field-oriented natural history journal publishing, and hence recording for posterity, original and significant information with relevance to Canadian plants and animals. For some years now there has been a drop-off in club activity in geology and palaeontology and there has been a concomitant dearth of manuscripts submitted to The Canadian Field- Naturalist on these subjects. This is unfortunate because these areas of natural history were once very strong ones in the club and the journal. Thus The Canadian Field- Naturalist currently plays a role in advancing knowledge in the biological sciences as authors record, analyze, and interpret their data. In recent years this function has become more important because several journals formerly available to field-biologists will no longer accommodate purely observational (non-experimental) data nor descriptive passages outlining the results of field investigations. The Canadian Field- Naturalist, by acting as a model for content and format, also serves to stimulate amateurs and young biologists to record their observations and to start their own investigations leading to fruitful lifetime hobbies or careers as field-biologists. There is no doubt for many of us that the scientific quality of The Canadian Field- Naturalist is currently high. Otherwise it would not attract submissions from a wide variety of natural scientists, nor would respected scientific experts serve as Associate Editors or referees, nor would the National Research Council of Canada have supported its publication by awarding substantial grants when funds were needed. In earlier years, however, The Canadian Field- Naturalist was neither completely rejected nor completely accepted by the scientific community because it did not use the peer review system and was only partially scientific. Today it is accepted as a scientific journal and is widely distributed, reaching many lay and professional readers including those in Canadian and foreign: libraries. It serves to varying degrees both writers and readers by permanently preserving valuable information of national importance and by helping to build the reputation of professionals who must publish in a refereed journal of high standards and quality. Although from time to time The Canadian Field- Naturalist publishes significant scientific papers concerning Ottawa biota, I agree that in general it currently does not relate very much to local naturalists with purely local interests other than in its publication of the reports and notices of the club. Current editorial policy does exclude papers that are of only regional or local interest, i.e., those that are not nationally significant, but these papers are appropriate for and can be accommodated in society newsletters and in regional or local natural history publications, many of which are of excellent quality. If The Canadian Field- Naturalist were to publish these items, it would again be ina gray area and eventually it would lose its current status and reputation as a primary scientific publication. I feel that most naturalists should find something of interest in every issue of The Canadian Field- Naturalist although I recognize that it does not fill the needs of all the local members. But the club does publish five times a year an excellent magazine, 7rail & Landscape, containing items of local | THE CANADIAN FIELD-NATURALIST Vol. 93 natural history interest, matters of conservation in the Ottawa area, plus announcements of lectures, excursions, and special meetings of the club. As far as many local members are concerned Trail & Landscape in fact is the voice of their club, yet it has no official status as a club publication and its continued existence is not completely assured. Certainly re-evaluation of Trail & Landscape’s status, réle, and relationship with the club seems urgently required in the present context. Clearly The Canadian Field-Naturalist is filling the need for a field-oriented national natural history journal; it serves both amateur and professional naturalists and field-biologists, maintains high scientific standing, and has assured itself of a place in history as it records the current state of the discipline. In doing so it is also fulfilling the second objective of the club, “to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible.” Are Amateur Naturalists Considered? One of the serious concerns of the Council of The Ottawa Field-Naturalist Club is the position of amateurs vis-a-vis The Canadian Field-Naturalist. Various members have recently expressed particular concern that the highly professional nature of the content and the more prestigious format of the journal have intimidated amateur naturalists and thus prevented them from submitting manuscripts. At Council’s request a survey was conducted of two volumes from each of the last three decades and all authors were classified as “professionals” if they had an institutional address and as “amateurs” if they had a home address. The survey clearly indicated a gradual decade-to-decade decrease in the number of papers by authors classed as amateurs. But these results are open to several interpretations. The question of whether the number of submissions from amateurs with data worthy of publication has decreased because of the journal’s content and format cannot yet be answered. I sincerely hope, however, that no potential author has been deterred from submitting a paper that he or she felt was of national significance. Let us consider some possible factors that could have contributed to the results obtained in the survey. a) Is an institutional address an adequate criterion for identifying an author as a professional? It must be realized that a professional in one field of science may be an amateur in another discipline even within the biological sciences. Also many authors prefer to use an institutional address even if the paper they submit has no relevance to their particular profession. Thus the survey could not identify all the amateurs in the true sense of the word. b) Are the amateurs of today sending their manuscripts to other publications that were either not in existence or not of sufficient interest or quality for amateurs of earlier years? In general, over the last few decades most scientific publications, including The Canadian Field- Naturalist, have changed to become less anecdotal, more structured, and more polished. At the same time there has been a dramatic increase in the number and quality of alternative sources of publication for natural history notes. Some of the regional and local publications of natural history societies have changed from mimeographed newsletters, where often virtually any material submitted was accepted, to a printed booklet format, where raised standards are evident. The high calibre of these publications has enhanced the image of the organizations publishing them and surely is appreciated by their members. Trail & Landscape is an excellent example. c) Isamateur science recognized as the initial training ground for many of those who will eventually make their living as professional field-biologists? Perhaps these young people with their special enthusiasm for natural history are our most inspired amateurs. Although many manuscripts submitted to The Canadian Field- Naturalist are from authors with institutional addresses, often the authors are really students, 1.e., advanced amateurs who are not yet professionals. d) Are today’s amateurs motivated to make their own observations systematically and rigorously and to write them up for publication? In the past this motivation was often supplied by interested 1979 SMITH: EDITORIAL 13 professionals who spent considerable time answering questions, identifying organisms, and interpreting often routine natural phenomena for members of the public, including amateur naturalists. Amongst these were occasionally people with exceptional latent talent who needed only a bit of interest to be shown in them by such notables as Jim Baillie and A. F. Coventry in Toronto, Clifford Carl in Victoria, Earl Godfrey in Ottawa, and many others. Although these exceptional professionals published valuable papers on their own research, perhaps equally important although often unsung contributions on their part were in providing encouragement to beginning naturalists to observe and record significant data. Because of our increasingly complex society, and the drives to establish their own reputations and empires, it is my impression that fewer and fewer professionals in universities, museums, and government departments take time to communicate with amateurs. Therefore, often amateurs now cannot readily discern whether their observations, sometimes just important single observations, are of particular interest and significance. Moreover, the incentive for amateurs to report (write up for publication) their observations may be lacking without this professional encouragement. é€) Dosome of the few particularly keen amateurs who want to help advance knowledge prefer to work under the direction of a professional field-biologist rather than on their own? Certainly Opportunities to do this are available and advertisements specifically outlining projects are fairly widely circulated. Undoubtedly any publications resulting from this type of joint amateur- professional collaboration bear an institutional address. Jf) Overall, do more amateurs today pursue natural history for the particular pleasures it brings to them rather than through a desire to carry out a particular study or to contribute to knowledge? With the increased leisure time in our changing society as well as increased incomes, people are travelling more extensively, not only in Canada but all over the world. They enjoy adding new species to their life lists, and taking photographs and making sound recordings using the array of technical equipment now available to them. The greatly increased mobility and opportunities for fleeting but exciting glimpses of different biomes present a far different situation than in earlier years. Then people used to be more sedentary and amateurs were more interested and persistent in devoting time to the valuable study of a single species or a few local species. Their repeated, often tedious, observations over long periods of time provided data that could be synthesized and interpreted. New trends and even definite changes were uncovered. Of course, with the advancement of knowledge, it is no longer so easy to find new fields to conquer, e.g., to describe species or habits or extensions of ranges. Are many of today’s amateurs still willing to do potentially valuable long-term studies in the field and to analyze and interpret their data? Or are they preoccupied with spotting rare species here or travelling to distant lands to identify, check off, then move on to the next, rather than studying, watching, and recording? g) Are amateurs reluctant to send a manuscript to The Canadian Field- Naturalist because it is too “scientific”? Those of us currently responsible for the content of the journal are anxious to have submitted by amateur and professional naturalists and field-biologists papers on results of sound original investigations in any field of natural history that is of significance and relevance to Canada. The pages of the journal are open to all amateurs whose papers meet these criteria. Reluctant amateurs should be particularly aware that important data not put on record may be lost to science. Although we endeavor to maintain the journal as a first-class scientific one, we also emphasize that scientific writing does not need to be, and preferably is not, pretentious but rather is simple and straightforward. Ideally it should be clear, concise, accurate, logical, and interesting. The journal isa broad general one and it is important that the papers we publish be understood by as many as possible of our readers. Although we expect trained biologists to write their manuscripts in the accepted style (and not all do), I emphasize here that we are particularly willing to help amateurs by giving advice and guidance on writing to improve their submitted manuscripts. The major criterion for acceptance is the importance of the information and its relevance to Canadian natural history. 14 THE CANADIAN FIELD-NATURALIST Vol. 93 In the past, amateurs played an extremely important réle in the advancement of knowledge in the natural sciences, particularly in field biology. Are amateurs still doing so? My questions and possible interpretations of the results of the survey regarding amateurs may help to illustrate that answers to seemingly simple questions are often complex and involve many interrelationships. Future Role of The Canadian Field-Naturalist The existence of The Canadian Field-Naturalist as a primary scientific and hence archival publication depends on several factors. These include the submission of manuscripts that, after review by competent referees, are found acceptable for publication; the availability of sufficient funds to cover publication and distribution costs; and the agreement of the publisher to maintain the journal as such. The Articles and Notes published in The Canadian Field-Naturalist are refereed and almost all undergo minor or major revision before they are accepted. But there are also several non-refereed pages in each issue containing general information considered to be of wide interest. Should the number of these pages be increased? Should they contain more items of interest only to the local club members? For instance, would items that now appear in Trail & Landscape (sent to all local members and any other members who request it) or in The Shrike (the club birding newsletter, available only on subscription) be suitable? If so, would these other publications still be maintained as they are now? At present the major expenses of the journal are met by page charges levied to authors and the subscription fees from institutions. Individual subscribers number just over 300 but their fees plus the portion allotted from the membership fees are not insignificant. The results of a questionnaire circulated to the club’s membership with the renewal forms for 1974 (see editorial in Volume 89(1), 1975) indicated that most people were members in order to receive The Canadian Field- Naturalist, but at that time only institutions, not individuals, could subscribe. Those who were club members in 1975 were given a choice starting in 1976 of maintaining their membership in the club witha portion of their membership fees (currently 40%) going to the journal, or of becoming subscribers for exactly the same annual fee but with the entire fee (100%) going to the journal. I wonder how many through knowledgeable choice, apathy, reluctance to change the status quo, or perhaps lack of knowledge of the financial implications to the journal, chose to remain members of the club. The opportunity fora member to become a subscriber was offered only once although there is no reason anyone wishing to do so shouldn’t change from being a member to become a subscriber at the start of any year. Because the club does have members across Canada and elsewhere, whether in name only or not, it considers that it speaks for a national and international membership. Although all members potentially have votes at the Annual Business Meeting and can elect the Council and introduce new business at the meeting, it is the local members, now mostly non-professionals, who come to the meeting, vote, and serve on the Council. But these are the people who will control the future of The Canadian Field- Naturalist. The Council of The Ottawa Field-Naturalists’ Club intends to produce within a year or soa statement regarding the desired réles for all the club’s publications and their relationship to the club. Thus the specific positions of Trail & Landscape and The Shrike as well that of The Canadian Field- Naturalist will be clarified and the réle of each as a club publication defined. Because this is a complex matter, the Council executive has assured me that any changes in direction will be implemented slowly. Therefore, after reading the background material in this editorial, but not being constrained by it to matters covered therein if there are other points to be brought out, interested persons are invited to state their views. I hope that favorable as well as critical comments, if justified, will be made because it is important to reaffirm where the journal is performing satisfactorily as well as where it may need modifying. Subscribers have one way to influence the journal directly, and that is by maintaining or discontinuing their subscriptions, but they can also express their views regarding The Canadian Field Naturalist’s role as a publication of The Ottawa Field Naturalists’ Club. I hope 1979 SMITH: EDITORIAL 15 all interested readers will put forth their thoughts because there is still time to have input into the deliberations of Council on this matter. Indeed, I invite them to do so by writing directly to the club (Box 3264, Station C, Ottawa, Ontario K1Y 4J5); carbon copies to the Editor would be appreciated. More formal comments as Letters to the Editor for publication are also solicited. The 100-year history of the club and the journal is perhaps unique and it is desirable that the two maintain a good relationship. Although the Editor and the editorial board are currently responsible for the editorial policy and content of the journal, ultimately it is the Council’s decision that will shape the future of The Canadian Field-Naturalist. This editorial is a distillation of many hours of discussion and analysis of the past and the current situation as interpreted by my husband and me. Therefore, by expressing our concerns, I hope that the opinions solicited from readers of all categories will help to influence the outcome of the current re-evaluation of the rdle of this journal. Furthermore, I hope the Council’s written statement will solve the misunderstandings and misinterpretations, calm the Council’s current uneasiness, clarify the club’s policy about its publications, and strengthen the position of The Canadian Field- Naturalist by formal reaffirmation of its present rdle as a primary scientific natural history publication published in Canada and relevant to Canada. LORRAINE C. SMITH Editor Demographic and Dietary Responses of Red-tailed Hawks during a Snowshoe Hare Fluctuation ROBERT S. ADAMCIK,! ARLEN W. TODD,? and LLOYD B. KEITH! 'Department of Wildlife Ecology, University of Wisconsin, Madison, Wisconsin 53706 2Department of Recreation, Parks and Wildlife, Edmonton, Alberta Adamcik, Robert S., Arlen W. Todd, and Lloyd B. Keith. 1979. Demographic and dietary responses of Red-tailed Hawks during a Snowshoe Hare fluctuation. Canadian Field-Naturalist 93(1): 16-27. Red-tailed Hawks (Buteo jamaicensis) responded functionally but not numerically to a cyclic fluctuation of Snowshoe Hares (Lepus americanus) near Rochester, Alberta. During 1966-1975, 89% of the resident pairs laid eggs, a nesting density of | pair per 8.3 km?. Mean clutch size (overall 2.2) ranged between years from 1.7 to 2.6, partly in response to prey-density changes. Mean hatching dates varied from 30 May to 10 June. Marked differences in annual productivity, 0.28 to 1.90 fledged young per breeding pair, primarily reflected nestling mortality. About 50% of all nestling losses were associated with food shortage. Seventy percent of annual variation in mortality through age 4 wk was jointly attributable to the frequency of rain and the weight of food brought to nestlings. The latter varied directly with Snowshoe Hare densities. The stationary population of breeding adult redtails contrasted with the numerically and reproductively cyclic horned owls (Bubo virginianus), but resembled redtail and other temperate-zone raptor populations further south. Food reductions during hare declines in the boreal forest ecosystem do not seem to be critical to survival of redtail nestlings unless combined with above-average rainfall. Key Words: Red-tailed Hawk, Buteo jamaicensis, Snowshoe Hare, Lepus americanus, population dynamics, cyclic fluctuations. A study of Red-tailed Hawks (Buteo jamai- censis) was conducted in central Alberta during 1966-1975. Its objective was to examine changes in raptor demography, food habits, and pred- atory-prey interactions during one complete population cycle of the Snowshoe Hare (Lepus americanus). Early results of this work were reported by Meslow and Keith (1966) and Luttich et al. (1970, 1971). McInvaille and Keith (1974) summarized findings through 1971, the year of peak spring densities of hares on the study area; they examined intraspecific relation- ships among redtails and interspecific relation- ships between redtails and Great Horned Owls (Bubo virginianus). They also assessed the effect of increasing hare densities on redtail breeding biology, and estimated rates of predation on prey populations. The most recent paper, that of Keith et al. (1977), examined the role of Red- tailed Hawks and other predators in the cyclic fluctuations of Snowshoe Hares and Ruffed Grouse (Bonasa umbellus). The hare population at Rochester declined after spring 1971, reaching the lowest point in its cycle by spring 1975. Over those 4 yr, significant changes occurred in reproduction, nestling sur- vival, and food habits of redtails. This paper describes such changes and examines their relationship to prey densities. It also presents a general analysis of redtail productivity since 1966. Methods Study Area The 162-km? study area (130 km?in 1966) was a block of mixed agricultural and forested land near Rochester, Alberta, about 100 km N of Edmonton. Luttich et al. (1971) and Rusch etal. (1972) described the vegetation, topography, and land-use practices on the area; McInvaille and Keith (1974) classified the major habitats and outlined their distribution as of 1971. Except for secondary growth in fire-killed areas, and some further clearing of forested land for pasture, the study area changed little over the next 4 yr. The breeding season for redtails in central Alberta is early April through July. Mean temperature at Rochester during this period is 12°C, with average minimum and maximum temperatures of 6°C and 18°C, respectively. Temperatures near freezing have been recorded in all summer months; and during extended periods of cloudy or rainy weather in July and 1979 ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 17 TABLE !—Summary of weather data during 10 breeding seasons of the Red-tailed Hawk near Rochester, Alberta! Periods in nesting cycle and types of weather data 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 Mean Courtship Days in period? 26 31 31 Ds 23 DY 36 31 22 24 28 Mean daily temperature (°C) -l 2 4 9 5 5 2 3 6 5 4 Days with precipitation (%) 31 19 29 52 9 15 3] 29 32 46 29 Mean daily precipitation (mm) het Ost Ox WA Oa 0.8 AO Oss 1.1 O09) Mean snow depth (cm) 7 25 | 2 9 25 24 6 23 8 13 Days with snow on ground (%) 65 97 29 40 52 78 83 58 82 46 63 Incubation Days in period 32 32 32 32 32 sy) 32 32 32 32 32 Mean daily temperature (°C) 12 12 10 11 10 13 12 1] 9 11 1] Days with precipitation (%) 44 25 34 34 22 22 DD) 38 50 53 34 Mean daily precipitation (mm) 2.1 1.2 0.8 Ie 0.4 1.0 0.9 2.1 0.7 ES) 1.2 Nestling (pre-tethering) Days in period 28 28 23 28 26 30 28 34 24 27 28 Mean daily temperature (°C) 13 14 14 14 17 13 14 15 15 13 14 Days with precipitation (%) 39 43 57 32 39 57 50 53 42 70 48 Mean daily precipitation (mm) 1.1 0.9 1.9 ha eal 4.2 SHO AN (0) O) eles 2.6 Tethering Days in period 20 28 27 22 28 21 23 16 31 22 2 Mean daily temperature (°C) 17 18 16 16 16 15 13 15 15 19 16 Days with precipitation (%) 60 43 67 55 64 67 61 69 74 55 62 Mean daily precipitation (mm) a) WHO Se) 4.2 6.7 5.0 IES 16 4.8 305) 3,5) 'Weather data were collected at the Meanook Meteorological Observatory, 26 km N of Rochester. Total days in each period were determined as follows: courtship—date of first observation of a red-tail on the study area to mean date of start of incubation; incuoation—mean date of start of incubation to mean date of hatch (32 d); nestling—mean date of hatch to mean date of tethering: tethering—mean date of tethering to mean date of release. August, daytime maximums around 7°C are not uncommon. Although occasional snow flurries occur as late as mid-May, most preci- pitation from late April to August falls as rain. Total spring-summer (15 August — 31 August) rainfall is variable, ranging from 11 to 44cm during our study. Heavy showers and rainstorms are common, but precipitation more often occurs as light rain over several days. Demographic Methods Methods of censusing raptors, tethering nest- lings, and estimating prey densities were iden- tical to those described by McInvaille and Keith (1974). Snowshoe Hare and Ruffed Grouse population estimates were revised slightly after additional analyses. We expanded the small- mammal census to include a spring (April-May) index for 1973-1975, and determined summer (June-July) densities of juvenile Richardson’s Ground Squirrels (Spermophilus richardsonii) on two study areas beginning in 1969. Waterfowl data used in the present paper were based on counts along a 0.4-km-wide aerial transect between the towns of Cold Lake and Swan Hills. This transect passed about 11 km N of Roches- ter. Weather data were collected at the Meanook Meteorological Observatory, 26km N _ of Rochester. Examination of Weather Data To analyze relationships between weather and productivity of redtails, we arbitrarily divided their reproductive cycle at Rochester into four stages: courtship, incubation, nestling (pre- tethering), and tethering (Table 1). The court- ship stage began with the first date of observa- tion on the study area each spring. The tethering stage ended with release of the last young we had tethered to obtain food data. Mean dates of first observation, start of incubation, hatch, tethering of nestlings, and release of fledged young during 1966-75 were 5 April, 3 May, 4 June, 2 July, and 25 July, respectively. Within the courtship stage we examined five weather variables: mean daily temperature, percent days with precipitation, mean daily precipitation, mean snow depth, and percent 18 THE CANADIAN FIELD-NATURALIST days with snow on the ground. The last two variables were irrelevant in the three later stages of the reproductive cycle. Results Prey Populations Spring densities of adult Snowshoe Hares peaked in 1971 at 510 per 100 ha of habitat, and thereafter declined to only 1% of peak numbers by 1975 (Table 2). Because the birth rate was higher in 1970 than in 1971, young hares were more numerous in 1970, and the total spring- summer (May-July) population was conse- quently higher that year. As with adults, total hare numbers fell markedly after 1971 to about 1% of peak numbers by 1975. Richardson’s Ground Squirrel populations rose between 1968 and 1970, fell to about 25% of their maximum by 1974, and increased again in 1975. During the highest year, adult densities averaged 988 per 100 ha of habitat (closely grazed pastures) in May, and juveniles averaged 3350 per 100 ha in June (Table 2). Populations of voles (Microtus pennsylvan- icus) continued to fluctuate during 1971-1975, as in previous years. amplitudes of change between high and low populations were 7- to 32- fold in late summer trap-night indices. Lowest numbers occurred in August 1968, 1971, and 1975. May populations were lower in 1973 and 1975 than in 1974 (Table 2). Ruffed Grouse were most abundant at 61 per 100 ha of upland forest in May 1968. A major decline took place between the springs of 1970 and 1972, as numbers fell 65% to 16 per 100 ha. The population remained low through 1974, then more than doubled by May 1975. Sharp-tailed Grouse (Pedioecetes phasian- ellus) increased from 1966 to 1970, but decreased abruptly within the next year and were scarce thereafter (Table 2). Waterfowl trend data for 1971-1975 indicated a general decline of about 50%. Red-tailed Hawk Numbers, Reproduction, and Nestling Mortality During 1972-1975, the Red-tailed Hawk population on our study area maintained the approximate numerical stability of earlier years (Table 3). Total individuals varied from 53 in 1967 to 39 in 1975, while breeding pairs ranged Vol. 93 from 24 (1 per 6.8 km2) in 1966 to 16 (1 per 10.1 km2) in 1973. There was no significant difference between years in portion of the population paired (mean 94%), the proportion breeding (mean 84%), or the proportion of pairs breeding (mean 89%). Mean hatching dates differed significantly (P <0.05) between years (Figure 1), being latest in 1967 (10 June) and earliest and most variable in 1972 and 1973 (2 June and 30 May, respectively). Significant (P < 0.05) annual variation also occurred in mean clutch size and mean brood size during 1966-1975. Mean clutch sizes were largest in 1970 and 1972 (2.6) and smallest in 1966 (1.7) and 1975 (1.9). Egg counts were made only once per nest, ranged over the incubation period, and included nests off the study area (Table 3, footnote 2). There was a marked change in the causes and rates of nestling mortality among redtails during 1972-1975 compared with earlier years (Table 3). Losses attributable largely to horned owls averaged 43% in 1968-1969 (Luttich et al. 1971), but dropped to 14% in 1970-1971, as sharply rising hare densities apparently buffered such predation (McInvaille and Keith 1974). Preda- tion by owls recurred in 1972 coincident with a major decline in hares. During 1973-1975, we recorded no horned owl predation on nestling redtails, as Snowshoe Hares became scarce and owl numbers declined through egress and non- breeding (Adamcik et al. 1978). But total nestling mortality on the study area rose from 37% in 1972 to a mean of 66% during 1973-1975. Almost all of such losses occurred before tethering (i.e., with 3-4 wk after hatching) in 1973-1975, whereas only about half had occur- red by that time in earlier years. Of the 30 young lost prior to tethering during 1966-1972, 18 disappeared completely, and the remains of 12 (40%) were found in or below the nest. Four of the latter had been cannibalized by siblings. In 1973, 8 of 18 (44%) dead young were found in or below the nest, and 6 of these had been cannibalized. In 1974, 4 of 14 dead young starved in the nest, 2 disappeared, | was cannibalized, and 7 (50%) died after falling from the nest. Three of the latter seven birds survived their initial fall and were replaced in the nest; two 19 ADAMCIK ET AL.. RED-TAILED HAWKS AND SNOWSHOE HARES 1979 (DIAS AYIIPIEAA PUR YSI4 “S'A ‘UBULION “CY Jo Asoqinoo P1BP) JaIsayIOY JO N WY |] noqe passed JOOSUBI} OY) SSITLH UBMS PUB IAB] P[O_. 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C€7 £01 /us0g ‘ou Ing Ae Ane 6e VC ey (O01 / ou ABI PV pte H FOYSMOUS L961 9961 XOpul potiad sasseyo saiads uoijejndod oul | aay jo adc] BAG|Y ‘1o]S9yIoY Jesu SYMEH] payiei-poy Aq uaye} satoads Aaid swOs 10} saoipul uoNeindog —Z A1aVL 20 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 3—Some population and productivity statistics for a Red-tailed Hawk population on a 162-km? study area near Rochester, Alberta 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 Population statistics Breeding pairs! 24 20 19 18 19 19 21 16 17 18 Non-breeding pairs ? 3 3 3 2 | 2 4 2 1 Resident singles ? 7 4 0 | 2 2 4 2 1 Productivity statistics? Mean clutch size NWS 2.00 AMT 2.14 2.56 2.28 2.61 2.00 2.42 1.9 Mean brood size 1.71 2.00 2.07 2.14 yen y) DEG D3) 1.96 2.26 1.71 Nestling mortality (%)3 0-4 weeks 0 12 24(10) 11 5 11(16) 29(18) 64(45) 45(23) 72(78) 5-8 weeks 5 0 33(27) 28 13 0(38) 12( 6) 0(46) 17(22) 25(40) Total 5 12 49(35) 36 17 11(48) 37(23) 64(70) 54(40) 79(87) Young fledged per breeding pair4 1.58 1.60 0.68 1.22 1.90 37) 1.29 0.63 0.94 0.28 ‘Breeding pairs laid eggs; nonbreeding pairs did not. ?Mean clutch and brood sizes include nests off the 162-km2 study area, but within 40 km. There were 4, 1, 24, 10, 15,8, 15, and 16 off-study-area clutches in 1966-1968 and 1971-1975, respectively; and 5, 5, 18,9, 23, 10, 14, and 13 broods. Total nestling mortality (%) equals 100 times the complement of the product of the two-interval survival rates: thus for 1969 total mortality was 100 [1 — (0.89) (0.72)] = 36. ’Mortality rates shown in parentheses are for 18, 11, 24, 11, 13, and 13 off-study-area broods in 1968 and 1971-1975, respectively. 4Includes nests off the 162-km2 study area. fell out again, and the third was cannibalized by a sibling which later starved. In 1975, 15 of 18 (83%) dead nestlings died during a single 32-h period of heavy rain, high wind, and low temperatures. In an earlier year, 1972, 8 of 12 deaths had occurred within | wk of a heavy rainstorm. The immediate causes of death were thus notably different between years. We determined pre-tethering losses among nestlings in nests situated off the study area during 6 of 10 yr (Table 3). In no single year did such losses on and off the study area differ significantly, but the overall averages of 44 and 32%, respectively, were different (P< 0.05). We suspect that our more frequent intrusions ad- versely affected brooding and feeding of young at study area nests. On the other hand, as discussed in a later section, mortality during the tethering period (age 5 to 8 wk) on the study area averaged significantly less (P <0.05) than untethered young elsewhere (1.e., 15 vs. 30%). The end result of these time- and area-specific differences in mortality was a similar mean rate of loss over the entire nestling period of about 50% on both areas Dietary Responses to Changing Hare Densities Food Habits Snowshoe Hares, Richarson’s Ground Squirrels, and waterfowl constituted between 59 and 89% of the weight of food for young Red- tailed Hawks at Rochester each year (Table 4). The remainder consisted of a variety of small-to medium-sized mammals and birds, of which Franklin’s Ground Squirrels (Spermophilus franklinii), voles and sometimes grouse were important. Snowshoe Hares were the dominant food of redtails during 1970-1973. Their peak use in 1970-1971 reflected the hawks’ strong func- tional response to increasing hare densities (MclInvaille and Keith 1974). During the next two years hare densities declined sharply, but there was a notable lag in the redtails’ functional response to declining hare numbers. The con- comitantly low ground squirrel population (Table 2) may have been partly responsible for this lag. By 1974, however, hares were near their cyclic low, and ground squirrel densities were the lowest we recorded; redtails then switched to waterfowl, which became the most important single food item, comprising 36% of their diet Consumption of Richardson’s Ground Squirrels was lowest (10% biomass) during 1972, coincident with continued heavy utilization of hares and a sharp decline in the ground squirrel population (Tables 2 and 4). By 1975, with hare densities at their cyclic low and ground squirrel 1979 ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 2) 20 MAY 25 3 ©090000096 JUNE 20 10 IS FiGure 1. Annual distribution of Red-tailed Hawk hatching dates on the 162-km? study area near Rochester, Alberta. Dates of hatch of 63 to 84% (mean 72%) of study-area nests were determined yearly. Circles indicate nests first visited during incubation and/or within | wk after hatching. Mean date is given at left. densities up four-fold, ground squirrels comprised 40% of the redtails’ diet. Biomass of Prey Brought to Tethered Young The amount of food supplied to tethered nestlings varied significantly (P < 0.01) between years (Table 5). During 1966-1971, the mean daily biomass of prey per brood rose steadily from 345 to 768 g, while that per individual young increased more irregularly from 212 to 444 g owing to annual differences in brood size. Between 1971 and 1973, the amount brought daily to broods and individual young declined by 66 and 48% to means of 260 and 231 g, respectively, and remained low through 1975. Eighty-nine and 86% of the foregoing annual variation for broods and individual young, respectively, was attributable to changes in mean daily biomass of Snowshoe Hares in the redtail diet. Annual utilization of hares was a direct function of their densities: regressions of mean daily biomass on total hare population estimates (Table 2) yielded coefficients of determination 2D THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 4 — Spring-summer (1 June - 31 July) food habits of nestling Red-tailed Hawks near Rochester, Alberta! Percent frequency Percent biomass Prey species? 66) "67" 68 69 70) Jl 72 #73 74 75 66° 67 68 69-70) S/5 SZ See eS Snowshoe Hare 2 4 7 6 4a Bh — 2 9 tr 0 8 S17 24") 25 46 2a eaesS 4 0 Richard’s Ground Squirrel Ai 3 DS DR BP) 2109 6G Ul 7 V4 42) 25 34 939) 38° 28) sO Sees, Franklin’s Ground Squirrel 4 5 6 Al {ivr 3 3 2 2 2 5 7 7/ 5 l 3 5 4 Volesand mice 32 41 13 34 #40 30° 36 42 44 #23 5 6 DD 5 4 y) 4 6 10 4 Other mammals 7 10 16 13 I 5) 6 7 G@ DW 6 8 8 9 | 5 4 5 Total mammals 6 2 6/7 7 87 8 6) Wl SO. 6 G © 7 9 8) 90 GG 77 54 74 Waterfowl 8 12 Wl Gemedte eo alOh BR eae ae 27a, alle Bel 5 1 205 Wee SOmen9 Ruffed Grouse 2 | Ore a 2 | l | tir One | SA ats) I Phar), am Ot 0 Sharp-tailed Grouse l | | | I itr OR OR OY 2a De ne? | | ltr Or O- @ Unidentified Grouse I I 0 I 0 eet OR Om OL reed 822 SO) 2a er Omer) | O-= Oo Other birds 3 12 IS 10 © FT IG IS Io Bl IW »d Tile at RD! gD ee eo NO Total birds 25m Di Boer WIS) 3675 29" 405535) 850372 25- <2) NN cls WS le ae Ome Totals4 101 100 100 100 100 100 99 100 99 99 101 100 100 100 101 101 99 101 101 101 'Total number of food items was 695, 1063, 585, 563, 879, 545, 1224, 387, 811, and 299 in 1966-1975; biomass total (in kilograms) was 167.2, 255.0, 199.9, 161.6, 298.9, 215.6, 308.0, 85.8, 107.5, and 47.6, respectively. *Prey not specifically identified in the table were “Voles and mice”: Microtus pennsylvanicus (average > 80% of small-mammal biomass). Clethrionomys gapperi (7%), Peromyscus maniculatus (4%), Zapus hudsonius (1%), and Sorez cinereus (tr): “Other Mammals”: Ondatra zibethicus, Tamiasciurus hudsonicus, Thomomys talpoides, Glaucomys sabrinus, Mustelafrenata, M. nivalis, M. erminea, Mephitis mephitis; “Waterfowl”: Podiceps grisegena, Anas platyrhynchos, A. acuta, A. strepera, A. discors, A. carolinensis, A. americana, A. clypeata, Aythya americana, A. collaris, A. affinis, Bucephala albeola, Porzana carolina, Fulica americana; and “Other birds”: Accipter cooperi, Falco sparverius, Perdix perdix, Charadrius vociferus, Columba livia, Colaptes auratus, Sphyrapicus varius, Picoides villosus, Perisoreus canadensis, Pica pica, Turdus migratorius, Sturnus vulgaris, Dendroica petechia, and unidentified songbirds and domestic chickens. 3Less than 0.5. 4Insects and amphibians constituted less than 1% of the total biomass in all years. TABLE 5—Weight of prey items brought to tethered nestling Red-tailed Hawks near Rochester, Alberta! 1966 1967 1968 1969 1970 1971 1972 1973 1974 1975 Mean daily biomass (g) of food per brood Snowshoe Hare 28 Wi 121 122 289 399 229 101 14 0 Other prey species 317 375 383 457 340 369 280 159 331 316 Total (+ SE) 345 +30 452445 504+76 579+114 629+66 768 +80 5S09+53 260423 345+55 316+48 Mean daily biomass (g) of food per individual Snowshoe Hare 17 47 92 68 174 231 170 90 10 0 Other prey species 195 229 292 257 204 213 207 141 228 173 Total (+ SE) DIDS AN6 2762 30) S84ee al 325 22 Sl, S78 245) 7444 se 38) 3772 83" 23 sa DRS eee OMe laws 1The number of tethered broods was 14, 15, 13, 11, 17, 12, 17,7, 7, and 3 during 1966-1975, respectively. Prey biomass totals during 1966-1971 have been recalculated since the study by McInvaille and Keith (1974), using better estimates of mean prey weights; trends remained the same as reported in that earlier paper. 1979 (r2) of 0.89 and 0.79 for broods and individuals. The above relationships reflect both the hare’s periodic domination of the prey base, and the redtail’s strong functional response to fluc- tuating hare densities. Nest Distribution Using Clark and Evans’ (1954) _nearest- neighbor test (see McInvaille and Keith 1974), we tested for randomness the spacing within annual nesting populations during 1972-1975. As in previous years, the distribution of redtail pairs on the study area was significantly (P< 0.05) regular, suggesting that territoriality continued to function as a spacing mechanism. General Analysis of Factors Affecting Productivity The most notable demographic change among Red-tailed Hawks from 1966 to 1975 was the lower fledging rate during the last 3 yr: viz., 0.62 young per breeding pair in 1973-1975 vs. 1.38 in 1966-1972. Although the main immediate reason for this decline was increased nestling mortality (Table 3), there were significant annual variations in mean date of nest initiation, and mean clutch and brood size, which might also have affected productivity. We therefore examined each component of productivity to determine what affected it and whether it interacted importantly with others to influence rates of fledging. We considered the following demographic variables: (1) mean date of nest initiation (= start of incubation); (2) mean clutch and brood size; (3) hatching success; (4) nestling mortality to age 3-4 wk (pre-tethering); and (5) nestling mortality from tethering to age 7-8 wk. Where appropriate, and where data were adequate, we examined each of the above in relation to weather; prey density and/or biomass brought to young; and frequency of observer visits to nest sites during courtship, incubation, nestling, and tethering periods. Mean Date of Nest Initiation Mean date of nest initiation was analyzed in relation to weather factors and prey population levels through simple and multiple regression. Annual variation in mean date of nest initiation was not significantly related to percent days with precipitation, average daily precipitation, average snow depth, percent days with snow on ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES 23 the ground, or mean daily temperature during the courtship period. Nor was there any significant relationship between annual dates of nest initiation and spring densities of Snowshoe Hares, Richardson’s Ground Squirrels, or previous-fall densities of mice and voles. Dates of nest initiation were likewise unrelated to various combinations of prey population indices and weather data. Clutch and Brood Size, Hatching Success, and Nest Desertion We initially suspected that weather conditions during courtship affected availability of prey and/or redtail hunting activity, and thus might influence clutch size. Annual variations in mean clutch and brood sizes, however, were unrelated to weather at Rochester. Years with largest clutches (1970, 1972, and 1974) had a broad spectrum of weather conditions during the courtship period (Tables | and 3). For example, 1970 and 1972 had mean daily temperatures during courtship of 5° vs. 2°C, rain or snow on 9 vs. 31% of the days, and snow depths of 9 vs. 24cm. In 1966, when clutches were smallest, mean temperature and percent days with precipation resembled conditions in 1972, while snow depth was close to that in 1970. Additionally, we could find no correlation between clutch size and average daily precipi- tation or percent days with snow on the ground. Although our data suggest that prey abundance may influence clutch size, the relationship was certainly not clear-cut. The large-clutch springs of 1970 (2.6) and 1974 (2.4) were each associated with high vole populations (MclInvaille and Keith 1974; Table 2); hare and > ground squirrel densities were also high in the first case but low in the second. In 1972, when clutches were equally large (2.6), adult hare densities were about 70% of those in 1970, adult ground squirrels about 30%, and voles were little utilized and probably scarce. The two springs in which mean clutch size was smallest, 1966 (1.7) and 1975 (1.9), were characterized by low hare and vole populations; we have no information on adult ground squirrels in 1966, but they too were low in 1975. It is, of course, possible that conditions on winter ranges or during migration influence the size of redtail clutches. There was a positive and nearly significant correlation (r = 0.60, P = 0.07) 24 THE CANADIAN FIELD-NATURALIST between annual dates of first arrival at Rochester and mean dates of egg laying, but mean clutch sizes were largely unrelated to the latter (r = -0.34, P > 0.25). The cause of major annual variations in mean clutch size probably resided on the breeding grounds since largest clutches occurred coincidently during 1970 and 1972 among redtails and non-migratory Great Horned Owls at Rochester (Adamcik et al. 1978). In calculating rates of hatching and nest desertion prior to hatching, we excluded nest desertions attributable to climbing at the onset of laying or incubation. The significant between- year difference in hatching and desertion rates was entirely owing to data from 1968. In May, a major fire swept through part of the study area and likely prompted four nest desertions. Only 50% of all eggs hatched that year, and 6 of 19 (32%) active nests were abandoned prior to hatching. During the other nine years, hatching success averaged 92% (range 79 to 100%) and nest desertion only 6% (range 0 to 17%). Pre-tethering Mortality Pre-tethering losses of nestlings varied between years from 0 to 75% (Table 3). We analyzed such mortality through stepwise regression, in which the initial independent variables were mean daily temperature, percent days with precipitation, mean daily precipi- tation, mean numbers of days between our visits to nests, and mean prey biomass brought daily to tethered broods (expressed both as biomass per brood and per nestling). All statistics except the last were obtained during the pre-tethering period. The amount of food brought to young was measured during the tethering period, but we considered it also indicative of relative amounts provided before tethering. We chose a discriminant probability level of 10%, and constructed two separate models, one incor- porating mean daily biomass of prey per tethered brood, the other per individual. The above described regression analysis discarded mean daily temperature, mean daily precipitation, and our frequency of visits during the nestling stage from those models which best accounted for the annual variance in _pre- tethering mortality. The resulting multiple regression equations and coefficients of determination were these: Vol. 93 (1) Y; = 3.88 + 1.36A; — 0.14B; r = 0:67; FQ,7) = 7, Bs (2) Y,; = 8.49 + 1.25A; - 0.09C, r =0.73; F2,7) =9:49, P1005 where Y; = estimate of pre-tethering mortality in year 1; Ai; = percent days with precipitation during the nestling stage in year 1; B; = mean biomass (g) of prey brought daily to individual tethered young in year 1; C, = mean biomass (g) of prey brought daily to tethered broods in year 1. These models leave approximately 30% of the variance in early nestling mortality unaccounted for. We mentioned previously that Luttich et al. (1971) attributed higher tethering-period mortality in 1968 and 1969 to horned owl predation, and MclInvaille and Keith (1974) suggested that such predation was buffered by the rising hare population in 1970 and 1971. A further horned owl kill, the first since 1969, occurred in 1972, when hares had declined sharply and owls were still abundant. Owl numbers fell rapidly during 1973-1975 (Adam- cik et al. 1978), and no further predation on tethered redtails was observed. Losses to horned owls, either before or after tethering, would likely be independent of weather and biomass of prey brought to young redtails, and might thus account for some if not most of the remaining unexplained variance in our models. Tethering-period Mortality Variations in tethering-period mortality were irregular, and independent of weather factors, amount of food supplied by adults, or prey population densities. This contrast with pre- tethering mortality doubtless reflects an in- fluence of our tethering methods, as two important causes of pre-tethering mortality, sibling aggression and exposure, were largely prevented by tethering. Tethered nestlings could not reach one another; and beginning in 1971, tethering sites were kept dry with a suspended plastic sheet. Thus losses of study-area young were probably minimum estimates of normal tethering-period mortality. This conclusion is supported by the fact that in 2 of 6 yr tethering-period mortality was significantly IDS) lower (P< 0.05) on the study area than off (Table 3); and in at least 1 of the remaining years (1968), the main mortality factor was owl predation, which tethering did not prevent. Discussion Comparative Demography Reported densities of nesting redtails have ranged widely from | pair per 1.3 km? in California (Fitch et al. 1946) to | pair per 39.5 km? in Utah (Smith and Murphy 1973). The mean of | nesting pair per 8.3 km? at Rochester over 10 yr was near that of | pair per 11.3 km? reported in nine studies, including the above two, during 18 yr (Craighead and Craighead 1956; Gates 1972; Hagar 1957; Johnson 1975; Orians and Kuhlman 1956; Seidensticker and Reynolds 1971). Eleven percent of 188 pairs at Rochester did not lay eggs during 9 yr, 1967-1975. This was close to the 14% of 278 pairs reported on six other study areas during 13 yr (Craighead and Craighead 1956; Hagar 1957; Johnson 1975; Orians and Kuhlman 1956; Smith and Murphy 1973). Such lack of nesting applied to adults, 1.e., birds almost 2 yr and older (Luttich et al. 1971), because in only two cases (Gates 1972; Luttich et al. 1971) were yearlings (brown-tailed indi- viduals) paired. Each of these yearlings had mated with an adult. The annual frequency of resident singles varied from 0 to 13% of the total Red-tailed Hawk population at Rochester. When concerted efforts were made in earlier studies to obtain a total census, unmated birds were likewise often found, but in variable numbers (Craighead and Craighead 1956; Fitch et al. 1946; Smith and Murphy 1973). The percentage of yearlings among singles apparently differs markedly between areas: all singles were yearlings in southern Michigan (Craighead and Craighead 1956), whereas none was at Rochester. Fitch et al. (1946). described the harassment of single birds by territorial pairs on a California study area, and it seems likely that a major portion of the yearling cohort may frequent localities where adult breeding densities are low but food 1s plentiful. Henny and Wight (1972) calculated that maintenance of redtail populations was depen- dent upon a fledging rate of about 1.35 young ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES D5) per breeding pair. This calculation was based on the assumptions, among others, that the 22% nonbreeders reported in studies published through 1970 were yearlings, and that all adults nested. We have noted above that non-nesting occurs among adults, averaging 13% of 466 resident pairs. This is undoubtedly a con- servative estimate since unpaired adults are also frequently present. Yearlings are so rarely a part of the paired resident population that they can be largely ignored as potential breeders. Reduction of adult nesting from 100 to 87% increases the estimated required fledging rate from about 1.35 to 1.56 young per breeding pair. These calculations are particularly sensitive to errors in age-specific estimates of annual mortality. As Luttich et al. (1971) pointed out, first-year mortality may have been overesti- mated, thereby elevating the rate of fledging apparently needed for population maintenance. The stationary redtail population at Rochester, for example, fledged a mean of 1.38 young per breeding pair during the first 7 yr of our study. Although we may not know precisely the fledging rate that will balance the population, the much lower rate that occurred during the last 3 yr (mean of 0.62) was doubtless inadequate. In the absence of compensatory changes in survival and/or ingress, the earliest that such decreased productivity might have been detected by us was in 1975; but our population data are really insensitive in this respect, because an ap- proximately 50% drop in fledging rate would theoretically reduce by only 2 the number of pairs present 2 yr later. Henny and Wight’s (1972) survey of redtail clutch sizes implied east-west and south-north gradients of increase. Their mean of 2.9 for 20 nests in southern British Columbia, Alberta, and Saskatchewan was among the highest noted. Our data from Rochester do not fit this model. Annual means recorded by us ranged from 1.7 in 1966 to 2.6 in 1970 and 1972 (Table 3); and the overall average of 2.2 during 1966-1975 was comparable to the lowest regional means shown by Henny and Wight. Our egg counts, conducted only once during incubation and at no consistent stage, provided an estimate of mean clutch size over the incubation period; consequently, they may be biased low if partial losses or clutches occurred before hatching. We suspect, however, 26 THE CANADIAN FIELD-NATURALIST that most earlier studies of redtails were similarly biased. Factors Affecting Nestling Survival to Age 3-4 Weeks Fluctuations in nestling mortality influenced annual productivity of redtails at Rochester more than did all other demographic variables. About 70% of the variance in pre-tethering mortality was jointly attributable to frequency of rainfall and amount of food provided by parent birds. These two factors are not likely independent, because redtail hunting activity, and probably hunting success, were reduced by rain. The amount of food brought to nestlings was, of course, also affected by prey densities. Direct adverse effects of rain on survival of nestling redtails was earlier observed in California (Fitch et al. 1946) and New York (Hagar 1957), but not among Red-shouldered Hawks (Buteo lineatus) in Maryland (Henny et al. 1973) or Buzzards (Buteo buteo) in England (Tubbs 1974). Several times at Rochester entire broods were found dead, wet, and abandoned after heavy rains. Although it was not always possible to isolate the individual roles of inclement weather and low prey densities in nestling mortality, we could at least identify situations where malnutrition rather than exposure per se appeared mainly responsible. We conclude that emaciation, cannibalism, and most falls from nests were largely symptomatic of food shortage. An unknown but substantial portion of total disappearances was probably also caused by cannibalism and falls, and thus likewise ascribable to food shortage. From the above, we estimate that approximately 50% of pre- tethering losses resulted directly or indirectly from a shortage of food. Reproductive Strategy of Northern Redtails Luttich et al. (1971) noted that breeding populations of arctic- and temperate-zone rap- tors responded differently to changes in prey densities. The former, largely dependent on widely fluctuating microtines, are highly mobile and annually concentrate in areas where lemmings and voles have attained temporary abundance. Temperate-zone raptors, on the other hand, exploit a much more diverse and stable prey base, and local breeding populations of these Vol. 93 raptors are often remarkably stationary from year to year (see recent summary by Newton 1976, p. 280). Newton (1976) concluded that, where nest sites are not limiting, regional differences in breeding densities of diurnal raptors reflect differences in average levels of prey abundance. This apparent adjustment to the food resource that usually prevails is sometimes inadequate, for as Craighead and Craighead (1956, p. 226) observed, “There are many examples of hawks and eagles returning to previous nesting territories and attempting to raise broods where there was not sufficient food to support them.” Redtail breeding populations are distributed throughout most wooded regions of the United States and Canada south of the tundra. As a predator, this raptor is a generalist, taking a great variety of prey. One might thus expect that it seldom experiences food shortage. Our studies at Rochester suggest, however, that this may not be true for northern redtails occupying habitats in which Snowshoe Hares are strongly cyclic. We estimated earlier, for example, that malnutrition caused about 50% of the nestling losses within 3-4wk after hatching. We concluded too that approximately 70% of the annual variance in such mortality could be attributed to differences in the biomass of food supplied to nestlings and the frequency of rain. The former was largely determined (r? = 0.89) by the biomass of Snowshoe Hare, which was in turn largely determined (r2=0.89) by hare population densities. Despite the fact that their prey base has a major cyclic component, breeding populations of northern redtails exhibit the same numerical stability and reproductive persistence that characterizes redtails and other temperate-zone raptors further south. The lack of any notable reproductive response to changing hare densities suggests to us that reductions in food during hare declines become critical to nestling survival only when associated with above-average rain- fall after hatching. The latter is a post facto random event for which an accommodating reproductive strategy could not likely evolve. In contrast, Great Horned Owls at Rochester have a breeding strategy that is clearly adapted to the hare cycle (Adamcik et al. 1978) and differs sharply from the evident constancy of 1979 reproduction among horned owl populations further south. As McInvaille and Keith (1974) noted, northern horned owls unlike redtails, “... are year-round residents, subject to the stresses of cold and potential food-shortage during winter. When hares and grouse are scarce there are few alternative and available prey species. Most small-mammal activity is sub- nivean, and there is a paucity of overwintering songbirds.” Under these circumstances, cyclic lows in hare populations are certain to affect the owls’ physical and/or physiological condition and hence their rate of breeding in late winter (March). When Snowshoe Hares were scarcest, as in 1974 and 1975, horned owls at Rochester laid no eggs, whereas during the 1968-1972 period of moderate-to-high hare populations all pairs produced and incubated a clutch. Such variable reproduction combined with ap- preciable ingress and egress generated a four- fold change in numbers of resident pairs over the 10-yr cycle of the hare (Adamcik et al. 1978). The population dynamics of horned owls in the boreal forest ecosystem thus resemble those of arctic raptors (high mobility, conspicuous density, and reproductive changes), while breeding populations of northern redtails retain the characteristic stability of temperate-zone raptors. Acknowledgments The financial support for this study was provided by the University of Wisconsin, College of Agricultural and Life Sciences; the Research Council of Alberta; the Canadian Wildlife Service; the National Science Founda- tion (Grant GB-12631); and the Green Tree Garden Club, Milwaukee. Literature Cited Adamick, R.S., A. W. Todd, and L.B. Keith. 1978. Demo- graphic and dietary responses of Great Horned Owls during a Snowshoe Hare fluctuation. Canadian Field- Naturalist 92(2): 156-166. Clark, P. J. and F.C. Evans. 1954. Distance to nearest neighbor as a measure of spatial relationships in popu- lations. Ecology 35(4): 445-453. Craighead, J.J. and F.C. Craighead, Jr. 1956. Hawks, owls and wildlife. The Stackpole Company, Harrisburg. 443 pp. Fitch, H.S., F. Swenson, and D.F. Tillotson. 1946. Behavior and food habits of the Red-tailed Hawk. Condor 48(5): 205-237. ADAMCIK ET AL.: RED-TAILED HAWKS AND SNOWSHOE HARES Di Gates, J.M. 1972. Red-tailed Hawk populations and ecology in east-central Wisconsin. Wilson Bulletin 84(4): 421-433. Hagar, D.C., Jr. 1957. Nesting populations of Red-tailed Hawks and horned owls in central New York state. Wilson Bulletin 69(3): 263-272. Henny, C. J. and H.M. Wight. 1972. Population ecology and environmental pollution: Red-tailed and Cooper’s Hawks. /n Population ecology of migratory birds. United States Department of the Interior, Bureau of Sport Fisheries and Wildlife, Research Report Number 2. pp. 229-250. Henny, C. J., F. C. Schmid, E. M. Martin, and L. L. Hood. 1973. Territorial behavior, pesticides, and the population ecology of Red-shouldered Hawks in central Maryland, 1943-1971. Ecology 54(3): 545-555. Johnson, S. J. 1975. Productivity of the Red-tailed Hawk in southwestern Montana. Auk 92(4): 732-736. Keith, L. B., A. W. Todd, C. J. Brand, R. S. Adamcik, and D.H. Rusch. 1977. An analysis of predation during a cyclic fluctuation of Snowshoe Hares. 13th International Congress of Game Biologists. pp. 151-175. Keith, L.B. and L. A. Windberg. 1978. A demographic analysis of the Snowshoe Hare cycle. Wildlife Mono- graphs Number 58. 70 pp. Luttich, S.N., L. B. Keith, and J.D. Stephenson. 1971. Population dynamics of the Red-tailed Hawk (Buteo jJamaicensis) at Rochester, Alberta. Auk 88(1): 75-87. Luttich, S. N., D. H. Rusch, E. C. Meslow, and L. B. Keith. 1970. Ecology of Red-tailed Hawk predation in Alberta. Ecology 51(2): 190-203. McInvaille, W. E. and L. B. Keith. 1974. Predator-prey relations and breeding biology of the Great Horned Owl and Red-tailed Hawk in central Alberta. Canadian Field- Naturalist 88(1): 1-20. Meslow, E. C. and L. B. Keith. 1966. Summer food habits of Red-tailed Hawks near Rochester, Alberta. Canadian Field-Naturalist 80(2): 98-100. Newton, I. 1976. Population limitation in diurnal raptors. Canadian Field-Naturalist 90(3): 274-300. Orians, G. and F. Kuhlman. 1956. Red-tailed Hawk and horned owl populations in Wisconsin. Condor 58(5): 371-385. Rusch, D. H., E. C. Meslow, P. D. Doerr, and L. B. Keith. 1972. Response of Great Horned Owl populations to changing prey densities. Journal of Wildlife Management 36(2): 282-296. Seidensticker, J.C., IV and H.V. Reynolds, III. 1971. The nesting, reproductive performance, and chlorinated hydrocarbon residues in the Red-tailed Hawk and Great Horned Owl in south-central Montana. Wilson Bulletin 83(4): 408-418. Smith, D. G. and J. R. Murphy. 1973. Breeding ecology of raptors in the eastern Great Basin of Utah. Brigham Young University Science Bulletin, Biological Series 18(3): 1-76. Tubbs, C. R. 1974. The buzzard. David and Charles Ltd., London. 199 pp. Received 8 May 1978 Accepted 18 September 1978 The Oriskany Sandstone Outcrop and Associated Natural Features, a Unique Occurrence in Canada DIANNE FAHSELT,! PAUL MAYCOCK,? GORDON WINDER,? and CRAIG CAMPBELL4 'Department of Plant Sciences, University of Western Ontario, London, Ontario N6A 3K7 2Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6 3Department of Geology, University of Western Ontario, London, Ontario N6A 3K7 4421 King Street North, Waterloo, Ontario N2J 3Z4 Fahselt, D., P. Maycock, G. Winder, and C. Campbell. 1979. The Oriskany sandstone outcrop and associated natural features, a unique occurrence in Canada. Canadian Field-Naturalist 93(1): 28-40. An unusual type of oak-hickory forest not reported previously in Ontario has been found growing on sandstone in the southern part of the province. The sandstone is the middle Lower Devonian Oriskany Formation (approximately 380 million years old), a unit which crops out to no significant extent anywhere else in Canada and is found only toa very limited degree in the subsurface. This highly fossiliferous outcrop has a total areal extent of less than 250 ha (1 mi’), most of it either under a thin cover of sandy soil or exposed as actual outcrop. Although the site is very dry it is unusually rich in plant species composition; at least 40 tree species grow in the area including an unusually large representation of oaks. There is an unexpectedly large number of understorey species for a dry forest. Some species in the understorey are more northerly elements that are quite uncommon in southern Ontario; others are southern species which are rare throughout their range in Ontario. A number of prairie associates are present as well. Twenty-two of the plants are rare in Ontario. The site serves as habitat for Elaphe obsoleta obsoleta(Black Rat Snake), one of the largest species of snake in Canada, whose southern Ontario populations are declining. Key Words: Oriskany, sandstone, oak-hickory forest, big-shell community, brachiopods, Black Rat Snake. The objective of this paper is to outline some outcrops of Oriskany sandstone are indicated on of the outstanding natural features of the older geological maps (Stauffer 1915; Caley Oriskany outcrop in southern Ontario. The 1940) but just one, the largest of these, has been biologically important aspects of the site, which — verified as Oriskany using modern petrographic have been identified only recently, are in criteria (J. F. Cowan, 1977, unpublished data, imminent danger of being lost owing to quarry- University of Western Ontario). Only the one ing by the Flintkote Company of Canada. An authenticated exposure is shown on recent maps “environmental protection area” has been selec- (Sanford 1969; Hewitt and Liberty 1972; Telford ted by the company and will remain under 1975). This outcrop of flat-lying sediments, company ownership, but it contains little of located about 3.0 km NE of the village of Nelles biological significance. Quarrying of the area is Corners, Regional Municipality of Haldimand- recommended by the Ontario Municipal Board Norfolk, Ontario (42°56’N, 79°57’W) is shown and approved by the Ontario Ministry of as an irregularly shaped area with an average Natural Resources. Zoning by-laws were modi- diameter of approximately 1.5 km. Air photos fied in June 1978 to permit aggregate extraction and field observations, however, reveal that the at the Oriskany site, although the Regional outcrop is actually even more limited than the Municipality of Haldimand-Norfolk now desig- recent maps suggest. The greater part of the nates it as an environmentally unique ecological formation does not occur as an actual outcrop, area. but is covered either by the younger Bois Blanc Formation or glacial till. Figure | shows an Geology estimate of the areal extent of the outcrop at the The Oriskany Formation occurs to a very — surface and in the subsurface. This sketch was limited extent in southern Ontario but in no drawn on an air photo base using observations other part of Canada. A number of small made by ground reconnaissance. 28 1979 , aypiiee RB 1 O- STN Pn a5 vat ¥ 4 ! ~ 6 Loos) ; = pay 1 wee Py yee, sete | Sq 3 + FAHSELT ET AL — - SESS Ay . . Rae te RRAN RD. McMO * a ‘ cu rani Mens: ths i Since brad oe Seni a (Z = ’ rt (tom aE ye al Cc ‘: SP Cee Ee to Oreeeceoen 12 AlS as Pare ee ye me, pose | STU ee Go - CT weep che Core of Sai ¢ < , - ORISKANY SANDSTONE OUTCROP FEATURES deve th) mT NE RD * -—-~) CS \ EO 3S ne a T a a ST a PO NSO ann SLD BA Roth ES TS a4 eee gory ES ‘ron ay e existing quarry with subsurface Oriskany in west wall ee ee may: FIGURE 1. Sketch based on an air photo showing the Oriskany outcrop area near Nelles Corners, Ontario. Estimated limits of Oriskany sandstone in the subsurface are shown by a broken line. Vertical hatching represents Oriskany overlain by the Bois Blanc Formation; the unhatched portion represents Oriskany at the surface or directly under the soil. The locations of study sites I-IV (Tables | and 2) are indicated, as well as the open prairie area (P). 30 THE CANADIAN FIELD-NATURALIST There is one unmapped occurrence of authen- tic Oriskany at “Shoap’s Farm” about 2.4 kmSE of Springvale, Ontario (42°57’N, 80°97’W) (Parks 1913), which consists of a vertical face only a few metres in length. As it has no horizontal exposure, it reveals only limited geological information and has little influence on native plants growing on the surface. Thus, the small Springvale outlier is of relatively little significance. The Oriskany sandstone outcrop near Nelles Corners is a flat-lying erosional remnant which is now separated by a distance of 100 km from the edge of the continuous Oriskany in the subsurface south of Lake Erie (Figure 2). The EL AnKoE Vol. 93 type locality is at Oriskany Falls, southwest of Utica, New York. The New York correlation chart (Rickard 1975) shows the formation as part of the Deerparkian Stage (or the Siegenian, in European terminology) which is middle Lower Devonian or in absolute terms, about 380 million years old. This is an exceptionally coarse-grained sandstone consisting mostly of quartz with some feldspar. The quartz grains are well rounded and closely packed with a mini- mum of cement, which near the top of the formation may be calcite. The degree of cemen- tation is variable and the rock tends to be friable at the surface where the rock is subject to frost action. The unit is very massive occurring in beds ONTARIO ISOLATED ja EES ORNS ANY ee WR te OUTCROPS GEOGRAPHIC EXTENT (Omir mie ORISKANY SANDSTONE PENNSYLVAMA ORISKANY OUTCROP ar 2 FiGURE 2. Geographic extent of the Oriskany Sandstone Formation. The stippled area represents Oriskany in the subsur- face south of the Great Lakes. Isolated erosional remnants are indicated where they crop out in New York State and Ontario. Constructed from Rickard (1975), Kreidler (1964), and Lytle (1964). 1979 up to 100cm thick, which break out in large blocks along vertical joint planes. The rock is porous but rain falling on the surface runs laterally and disappears down joints or passes through to the underlying Bass Islands Forma- tion. The permanent water table is 16-20 m below the surface. The imminent quarrying operation would be carried out above the water table. Although fossil preservation 1s rather unusual in a coarse sandstone, parts of the Oriskany are highly fossiliferous. There are internal as well as external molds which show both interior and exterior characters of the shells. Fossils at the Nelles Corners site have been identified by Stauffer (1915), Caley (1940), and Best (1953) and include brachiopods, corals, trilobites, gastropods, pelecypods, and miscellaneous forms. Index fossils characteristics of the Oris- FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 31 kany sandstone include Acrospirifer, Costispiri- fer, Hipparionyx, Rensselaeria, and Oriskania (all brachiopods) (Figures 3-7). Some genera can be found in both North America and Europe (e.g., Acrospirifer), and others are reported only from eastern North America (e.g., Hip- parionyx). The large brachiopods that dominate the fossil fauna and are referred to as a “big shell community” (Boucot and Johnson 1967) are not found in any other Canadian sandstone. Of the more than 75 fossil species reported, only about one third occur elsewhere in Ontario or Quebec. Unconsolidated rock materials over the out- crop are mainly of glacial origin and are either nonexistent or very thin. The site is therefore not suitable for agriculture and 1s largely covered by open woods known locally as the Clanbrassil Forest. The biological significance of the area was not realized until relatively recently. FIGURES 3-7. Index fossils characteristic of the Oriskany 6, Rensselaeria; 7, Costispirifer. Formation. 3, Hipparionyx; 4, Acrospirifer; 5, Oriskania, By THE CANADIAN FIELD-NATURALIST Vegetation The first indications that the vegetation was in any way unusual were the observations of F. S. Cook (University of Western Ontario, personal communication 1976) who noted the presence of saxicolous mosses which are rarely seen in southern Ontario because their required sub- strate is essentially unavailable. Outcrops most common in southern Ontario are associated with the Niagara Escarpment and they are mainly limestone and dolostone. These rocks differ markedly from sandstone in their calcium content, the pH of materials derived from them by weathering, and their suitability for certain kinds of plant growth. Open oak-hickory forest growing on sand- stone bedrock is very unusual for Ontario. In order to develop a precise idea concerning the structure and composition of the forest, the vegetation was examined in detail and part of the area was sampled quantitatively. Methods Five homogenous areas ranging in size be- tween about 2 and 6 ha were selected. Four were forested and one was an opening within the forest. A list was compiled of all trees, shrubs, and herbs in each forested area. Two of the larger areas were sampled quantitatively using the Point Quarter Method (Curtis 1959); 30 sampling points were scattered at random throughout each site. Around each point four trees and four saplings were randomly selected in each of four quadrants, 1.e., the nearest speci- mens to the center point in each quadrat were selected. At every other point a sample of the ground layer was taken by recording presence (frequency) in metre-square quadrats. A number of environmental features as well as additional structural and compositional details of the vegetation were recorded when the sampling procedure was completed. A frequency value was determined for each tree species on the basis of its percentage occurrence at points. These values were summed for all tree species and the contribution of an individual species to that sum was calculated asa percentage (= relative frequency). Similarly, the number of stems contributed by a species at points of occurrence was calculated as a per- centage of the contribution of all species at all Vol. 93 points (= relative density). The total basal area for one species was calculated as a percentage of basal area for all species (= relative dominance). The three values for a given species, relative frequency, relative density, and relative domin- ance, were combined to give its importance value. This is a precise measure of the ecological influence of a tree species in the forest. In the forest openings lists were taken of all vascular plants present, and general observa- tions on community structure and environ- mental features were recorded. Parts of the outcrop along Townline Road (Figure 1) were badly disturbed with old foundations, old buildings, small abandoned quarries, and road systems. Because much of the associated vegetation was weedy, it was not examined in the same way as other sites within the general area. Results and Discussion Open oak-hickory communities (Figure 8) of the type located near Nelles Corners on sand- stone bedrock had not previously been found during an extensive survey of forest vegetation of southern Ontario (Maycock 1963). At least 28 different tree species are found in the stands sampled on the Oriskany outcrop area (Table I). This is considerable diversity for sucha poor dry site. At Site I (Figure 1) where the soil is very shallow there are no clear-cut dominants but Quercus velutina (Black Oak), Q. rubra (Red Oak), Prunus serotina (Black Cherry), Quercus alba (White Oak), and Carya ovata (Shagbark Hickory) are the major tree species. The source of names used in the text and iables is Fernald (1950). All 28 tree species are found in Site 1, but more than half of stand importance is accounted for by oaks which together have an importance sum of 163 (Table 1). In the sapling’ layer Fraxinus americana (White Ash), Prunus sero- tina, Amelanchier arborea (Serviceberry), and Acer saccharum (Sugar Maple) are well repre- sented. All of the oaks are represented to a limited extent in the sapling layer (approxi- mately 20% of sapling-density) but are less important here than in the tree layer. Hickory reproduction appears to be maintaining the status of this species. A rich assortment of tall shrubs is found including (listed in order of is y Ne ; nie me j 9 @ FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 33 FIGURE 8. Open oak-hickory forest on sandstone at the Oriskany outcrop in southern Ontario. The best of these stands are presently scheduled for quarrying by King Paving and Materials of Burlington, Ontario, a Division of the Flintkote Company of Canada. decreasing dominance) Cornus racemosa (Race- mose Dogwood), Amelanchier arborea, Vibur- num rafinesquianum (Downy Arrow-wood), Prunus virginiana (Choke-cherry), and Vibur- num acerifolium (Maple-leafed Viburnum). Pyrus coronaria (Wild Apple) is one uncom- mon species of note. Amelanchier arborea does not often attain tree sizes but in this stand there are a number of large specimens. There is also a rich tall herb layer which includes Preridium aquilinum (Brake), Solidago caesia (Blue-stem Goldenrod), and Aster sagittifolius (Arrow- leafed Aster), and a low herb layer with Carex pensylvanica, Galium aparine (Cleavers), and G. circaezans (Wild Licorice). In total 108 species of herbs and shrubs are present. Lichens are uncommon in forests of the deciduous forest regions but both lichens and mosses are locally abundant in Site I. Site II is immediately below Site I on a gradual south-facing slope and also has shallow soils. The dominant trees are Red Oak and White Oak with a lesser representation of Shagbark Hickory and Black Cherry. Black Cherry, Shagbark Hickory, and Red Oak are well represented in the sapling layer, while Cornus racemosa and Prunus virginiana are important elements in the shrub layer. In the medium-height herb layer Solidago caesia and Geranium maculatum (Wild Cranesbill) are found, and in the low herb layer are Maian- themum canadense (\wo-leafed Solomon’s- seal), Carex pensylvanica, and Aster macro- phyllus (Large-leafed Aster). This is also a dry complex with considerable diversity including 24 tree species and 87 herbs and shrubs. Site III is just south of Site II toward the bottom of the gentle slope. In Site III the 34 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE |—Tree species growing in selected stands at the Oriskany outcrop area. Locations of the four sites are shown in Figure |. Presence is indicated by plus signs and importance values (defined in text) for the two larger sites are given in parentheses Species ] Acer nigrum (Black Maple) + Acer rubrum (Red Maple) + (4) Acer saccharum (Sugar Maple) + (9) Amelanchier arborea (True Serviceberry) + (20) Carpinus caroliniana (Ironwood) + (3) Carya cordiformis (Bitternut Hickory) + Carya ovata (Shagbark Hickory) Celtis occidentalis (Hackberry) + Crataegus chrysocarpa (Hawthorn) + Crataegus sp. (Hawthorn) ste Fagus grandifolia (Beech) - Fraxinus americana (White Ash) + (17) Juglans nigra (Black Walnut) + Juniperus virginiana (Red Cedar) + Ostrya virginiana (Hop-hornbeam) + Pinus strobus (White Pine) + Populus grandidentata (Large-tooth Poplar) + Populus tremuloides (Trembling Aspen) P Prunus cerasus (Cherry) + Prunus serotina (Black Cherry) tp Pyrus coronaria (Wild Apple) at Quercus alba (White Oak) Quercus macrocarpa (Bur Oak) F Quercus muehlenbergii (Chestnut Oak) Quercus rubra (Red Oak) Quercus velutina (Black Oak) Tilia americana (Basswood) Ulmus americana (White Elm) + Total trees 28 Total herbs and shrubs 108 Soil pH 4.6-7.0 Depth A, (cm) 6 Depth B (cm) 13 Moisture dry Sites I] Il] IV + (3) + (3) + + + (18) a + (3) + + + + (7) + (21) + + fi A + + (1) + + i 2 + (15) + + (4) + + + (6) + + + (5) + (19) + + 4 + (78) + + + (4) + + (86) + + se (112) 7 + (8) + + 24 13 12 87 45 27 8.0 Usd) 6.4-7.8 8 10 10 10 48 18 dry—mesic dry—mesic mesic unconsolidated surface soil is deeper (see Table 1) than in either Site I or I]. White Oak and Shagbark Hickory are the two dominant trees and scattered Black Cherry is also found. In the sapling layer cherry and White Oak are well represented. Prominent tall shrubs are Cornus racemosa and Viburnum acerifolium, while herbs include Aster macrophyllus, Carex pen- sylvanica, Galium aparine, and Potentilla sim- plex (Old-field Cinquefoil). In total there are 13 tree species in Site II] and 45 species of herbs and shrubs. Site IV, like Site III, is on deeper soil. The dominants are Sugar Maple and Red Maple with some White Oak, White Ash, and Red Oak. This stand is heavily shaded and the saplings are mostly maple. The low shrub layer is very sparse and includes scattered Viburnum acerifolium. The only herbs are low and include Carex pensylvanica, C. pedunculata, and Solidago caesia. There are 12 tree species represented in Site IV and 27 herbs and shrubs. There are few mosses or lichens. In the forest openings, common dominants are Cornus racemosa, Danthonia spicata (Pov- erty Grass), Rhus typhina (Staghorn Sumac), Prunus virginiana, Desmodium canadense (Canadian Tick-trefoil), Hypericum perforatum 1979 (St. John’s-wort), Solidago canadensis (Can- adian Goldenrod), and S. juncea (Stiff Golden- rod). One of the openings (P in Figure 1) is of particular interest because of its high component of dry prairie elements, notably Lespedeza intermedia (Intermediate Bush-clover), L. capi- tata (Headed Bush-clover), Polygala verticillata (Whorled Milkwort), P. pol/ygama(Polygamous Milkwort), Desmodium paniculatum (Panicu- late Tick-trefoil), D. rotundifolium (Round- leafed Tick-trefoil), D. dillenii, Galium pilosum, Linum vriginiana (Virginian flax), Andropogon gerardi (Gerard’s Beardgrass), Monarda fistu- losa (Wild Bergamot), Physalis heterophylla (Variable-leafed Groundcherry), Solidago nem- oralis (Woodland Goldenrod), Panicum lineari- folium (Linear-leafed Panic Grass), and Aster ericoides. The Oriskany outcrop is considered to be a drier-than-average site for southern Ontario. If sites in the province were ranked along a continuum according to water availability, the Oriskany outcrop would be the driest of those capable of supporting forest. The trees are rather widely spaced approaching a condition inter- mediate between forest and savanna. The open condition of the canopy results in high light intensities within the stand and permits repro- duction of species such as Black Cherry, hickory and oaks, as well as heavy shrub and herb layers. Populus grandidentata (Large-toothed Pop- lar) is one species found at the site that 1s usually characterized by a very shallow rooting system. Many of the dominants, however, are species able to form long tap roots. For example Quercus velutina, Q. alba, and Carya ovata all produce vigorous primary roots early in devel- opment enabling the seedlings or young saplings to withstand drought conditions. Quercus rubra is another species represented at the site that is capable of producing deeply penetrating tap roots (Fowells 1965). Pinus strobus (White Pine) typically produces just the vestige of a tap root but has several large laterals extending outwards and down (Fowells 1965). White Pine root systems have been shown to penetrate toa depth of 4.5 m (Brown and Lacate 1961). Many trees and shrubs are first- or second- collection records for the Haldimand County which close gaps in the distributional patterns FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 35 for species both east and west. Thus the site has phytogeographical significance. In the Oriskany outcrop area, the favorable sites with deeper soil are definitely less diverse in tree species. On better sites more dominance is exerted by fewer species. Greater moisture, more favorable pH, and perhaps better nutrition contribute to this pattern of tree occupancy. Species diversity is much greater in sites with shallow soils (e.g., I and II). At these sites there are a number of major tree species but no clear dominants, and there are far greater numbers of shrub and herb species. Diversity is undoubtedly related to the great variety of microhabitat available for plants. Some rock surfaces are exposed, others are covered with thin soil. The soil depth varies and the shallower soils have a low pH due to sandstone and absence of carbonate rocks providing opportunities for acid-loving plants, conditions which are seldom available in southern Ontario. The maple stand (Site IV) is on deeper soil and is representative of a type of vegetation very common in southern Ontario. The interesting species of the sandstone outcrop type are absent. The sites of most value botanically are very heterogenous at the ground surface, 1.e., those with shallow acidic soil over sandstone. The age of older trees in the maple stand is approxi- mately the same as the age of older trees in the other forested sites studied. The openings adjacent to the forest stands have allowed the invasion of dry prairie ele- ments, further adding to the diversity of the area. The high diversity indicates that there may be other unusual species or communities in ad- jacent areas. For example, the Dry Lake area nearby has an interesting assortment of thicket and seasonal meadow communities. Quercus bicolor (Swamp White Oak) is one uncommon species that has been found in the lowland fringes of Dry Lake. One group of rare and notable plants occur- ring in the outcrop area consists of more northern species that are quite uncommon in southern Ontario, although they may be wide- spread northward, and the other is composed of southern species that are rare throughout their range in Ontario. Northern species such as Hairgrass (Deschampsia flexuosa) and Running 36 THE CANADIAN FIELD-NATURALIST Clubmoss (Lycopodium clavatum) are without doubt present here owing to the sandstone substrate and are otherwise quite rare in south- ernmost Ontario. Because of its southern loca- tion, the site also contains a number of un- common species that are members of the Carolinian flora of Ontario. This flora occupies the region directly north of Lake Erie to a sinuous line from Toronto through London to Port Franks on Lake Huron. Examples of such species in this area are Wild Crab (Pyrus coronaria), Black Walnut (Jug/ans nigra), Hack- berry (Celtis occidentalis), and Chestnut Oak (Quercus muhlenbergii). In summary, the unique vegetational status of the area is com- plemented by the presence of a number of rare and interesting species. Twenty-two of the species found in the Oriskany outcrop area are among those that have been listed as rare by Argus and White (1977) (Table 2). It is most unusual to find such a large number of rare plants concentrated in so small an area. Habitat for the Black Rat Snake The presence of unusual forest associations and unusual understorey plants provides a particular habitat for wildlife. The most notable animal known to inhabit the Oriskany site 1s the Vol. 93 Black Rat Snake (Elaphe obsoleta obsoleta), the largest species of snake in Canada. The Black Rat Snake is considered rare, threatened, or endangered in Ontario and Canada by several authors (Campbell 1969; Cook 1970a; Anony- mous 1970; Froom 1972; Stewart 1974; Parsons 1976; Cook 1977; Gregory 1977). Observations of Elaphe obsoleta at the out- crop were made by W. W. Judd (University of Western Ontario) in 1976 and J. Webber (Erindale College, University of Toronto) in 1977, and by at least four local residents during the past two years. Judd observed a snake at close range for about | min; Webber was able to sketch anatomical details from a distance of 2 ft. One local person observed many snakes (of varying sizes) simultaneously in the spring of NSE It should be noted that the combination of rocky terrain and extensive relatively undis- turbed forested area such as that found at the Oriskany site constitutes ideal habitat for the Black Rat Snake. Knudsen (1955) stated that the Black Rat Snake inhabits areas that have an abundance of rocky areas and crevices, and from these sites they venture out into nearby agri- cultural lands. The snake is sometimes con- sidered a forest species (Hay 1892; Morse 1904; TABLE 2—Rare plants (Argus and White 1977) found at the Oriskany outcrop. Areas are those shown on Figure | Species Asplenium platyneuron Panicum lanuginosum var. praecocius Carex laxiflora var. gracillima Disporum lanuginosum Polygonatum biflorum Juglans nigra Quercus bicolor Quercus muehlenbergii Ranunculus hispidus Arabis canadensis Prunus americana Desmodium rotundifolium Lespedeza intermedia Linum virginianum Viola pedata var. lineariloba Thaspium barbinode Vaccinium pallidum (V. vacillans) Asclepias exaltata Conopholis americana Galium pilosum Swertia caroliniensis Aster pilosus Common name Area found Ebony Spleenwort I Woolly Panic Grass P V Fairy Bells II Solomon’s Seal II Black Walnut I Swamp White Oak V Chinquapin Oak II Stiffly-hairy Buttercup I Sickle-pod ] Wild Plum 111 Round-leafed Tick-trefoil I Intermediate Bush-clover Pp Virginia Flax le Bird-foot Violet IP Meadow Parsnip Vv Hillside Blueberry I II Squawroot I I American Columbo V P 1979 Wright and Wright 1957; Fitch 1963); Wright and Wright (1957) and Fitch (1963) both mention occurrences associated with oak-hick- ory forests. Nearly half of the location records for radio-implanted specimens (Fitch 1963) were in trees while others were in burrows, low vegetation, or buildings. At the Oriskany site approximately 300 ha (3 km2) including the outcrop and immediately adjacent areas are considered suitable habitat. In Ontario, there are now two main isolated ranges for the Black Rat Snake. The one in the Kingston ~ Rideau Lakes area is maintaining itself to some degree, but the range in south- western Ontario is clearly discontinuous despite maps (e.g., Conant 1975) indicating a continu- ous range along Lake Erie. In southwestern Ontario, there are only a few small isolated locations which are considered by C. A. Camp- bell to be sites of Black Rat Snake populations. These with dates of sightings in parentheses are (1) Essex County: Point Pelee, Pelee Island (1977), (2) Kent and Middlesex County: Skunk’s Misery (1968, 1974), (3) Haldimand-Norfolk Regional Municipality: the western part of the former Norfolk County (the main center of abundance) (1940-1978) and other locations including the Oriskany site (1976-1978), and (4) Niagara Region: Fonthill in the Shorthills area (1928-1976). Even at these locations the snakes are not abundant; they are threatened by agriculture and a multitude of other human activities. Clearly, the number of large snakes in southwestern Ontario is seriously declining, as 1s the available habitat. There is no evidence that the Oriskany population is continuous with others in Ontario. Further details on the Black Rat Snake in Ontario are in an unpublished report by C. A. Campbell (1977) entitled “The status of the Black Rat Snake Elaphe obsoleta obsoleta in Ontario and particularly in the Haldimand-Norfolk Region” (available from C. A. Campbell or D. Fahselt). Fitch (1963) calculated that the home range of adults in Kansas is 12 ha for males and 9 ha for females. In Maryland, the average home range of E. obsoleta was estimated to be 18+ha (Stickel and Cope 1947). Small populations are always particularly vulnerable, so the entire forest and outcrop complex must be maintained in its present condition in order to maximize FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 37 numbers. Restriction of habitat is listed as one of the three major factors contributing to the decline of Elaphe obsoleta (Cook 1970a, b). General Discussion Threats to the Area As the area is unique and interesting in several ways it is useful to consider the impact of likely types of disturbances. First, there is abundant evidence that fires have occurred in the past. Second, it is obvious that exposed bedrock of this nature is an attractive commercial source of mineral aggregate. (a) Fire Certainly dry sites such as this are quite susceptible to fire. Within the area blackened stumps or charcoal in the soil are evident at various locations. Though some scattered trees are older than 90 yr, it is clear that neither the oak nor maple forest represents an advanced state of succession. Few of the trees are large, most having basal areas of 516 mm2 (80 in2) or less. Fire may be one of the factors responsible for maintaining the oak-hickory forest in its present secondary successional state; burning would permit oaks, hickories, and Large- toothed Poplar to persist and predominate. Y oung oaks and hickories, and to a lesser extent even mature trees, sprout vigorously after the tops are killed back to the ground by burning (Fowells 1965); and fire may help to perpetuate Populus grandidentata as well as some of the prairie elements. Periodic burning may be partially responsible for maintaining the present successional state of the stand and contributing to the diversity of the interesting plant species. Selective logging at the site may have produced a similar effect. (6) Commercial quarrying Beneath the Oriskany formation at the Nelles Corners site is approximately 15m of dolo- stone, a calcilum-magnesium carbonate (the Bass Islands Formation), which is desirable as aggre- gate for paving and building. Quarrying would remove the sandstone formation upon which the oak-hickory forest depends. The forest could not re-establish itself afterward on the freshly ex- posed rock surfaces of the quarry floor because soil chemistry and water availability would be 38 THE CANADIAN FIELD-NATURALIST quite different. Also there would be cold air drainage into the quarry bottom. If commercial removal of dolostone does take place an excava- tion with sheer vertical faces would result and this would have adverse effects on any peripheral remnants of forest. The edge effect associated with a disturbance as drastic as quarrying would likely be much more pronounced in a dry area such as this than it would in a mesic site. The establishment of a preserve for the fossiliferous portion of the Oriskany outcrop would not have a sufficiently large geographic area to support the unique plant communities such as those in Sites I and II, nor would it be suitably placed to include them. Another problem associated with a possible quarry operation would be the effects of wind- blown particulates. Dust reduces available light and in some cases reacts with water to form toxic solutions. Trees growing near a source of carbonate dust were reported to have reduced terminal growth (Manning 1971) or to be ina generally poor condition (Brandt and Rhoades 1972). Particulate interference with stomatal behavior seriously affects the diffusion resis- tance of leaves and changes the rates of gaseous exchange (Ricks and Williams 1974). This could aggravate moisture stress on sites that are exposed or dry (Smith 1974). The rates of degradation of photosynthetic pigments in the leaves of Quercus petraea were significantly changed owing to particulate pollutants (Ricks and Williams 1975) and senescence occurred earlier at the polluted sites. Leaves with moder- ate limestone dust deposits had a greater incidence of fungal leaf spots (Manning 1971), and plants dusted with cement-kiln dust were more susceptible to fungus leaf spot disease (Schonbeck 1960). A numerical model which predicts the rate of dispersion of atmospheric particulates and the amounts of deposition in wooded areas was described by Belot et al. (1976). A forest canopy significantly increases the concentration of par- ticulates deposited near the source. It is pre- dicted that maximal deposition would occur within | km of dust-producing activities. Tree species respond differentially to dust accumulation. Brandt and Rhoades (1973) have shown that while lateral growth of Liriodendron Vol. 93 tulipifera (Tulip Tree) was increased as a result of deposition of dust from nearby limestone quarries and processing plants, Quercus prinus, Q. rubra, and Acer rubrum underwent a reduction in lateral growth. Therefore, in a mixed stand involving these species, importance values would be altered with time. In fact Brandt and Rhoades (1972) documented significant differences in the seedling-shrub and sapling strata between two comparable sites, one witha heavy accumulation of limestone from quarries and processing plants and the other a control area with no dust accumulation. The dominant species at the dusty site would therefore change with continued dust accumulation. The papers by Brandt and Rhoades indicate that it would be difficult to maintain a natural balance among tree species in a forest adjacent to a sustained source of heavy limestone dust. It cannot be imagined that quarrying would have anything but a detrimental effect on Elaphe obsoleta. First, the snake is susceptible to road- kill (Fitch 1963). Second, it retreats with the advent of disturbance to wooded areas (Morse 1904; Minton 1968). Quarrying would lead to an obvious loss of suitable habitat. Synopsis We have emphasized the geological impor- tance of the Oriskany Formation in Canada and provided some indication of the unusual nature of this forest tract and its richness in tree and other plant species as well as its unusual structural and compositional features. We have also attempted to show its value as a habitat for the Black Rat Snake. The Oriskany site repre- sents the on/y example of oak-hickory forest on sandstone in Ontario. It is decidedly unique in representing a dry upland type of oak-hickory forest not recognized in Ontario. The oak- hickory type in Ontario is usually associated with heavy clay soils which have peculiar drainage and moisture features and never seems to approach a classical dry oak-hickory type as does the existing example at Nelles Corners. This feature coupled with the large number of unique and interesting plant occurrences, es- pecially the unusually large number of oak species, combines to produce a natural area of great value and interest, and one which should 1979 be given complete protection. Preservation of the entire area would be consistent with the adopted policy statement of the Canadian Institute of Forestry (Weetman 1972) that 123-410 ha (300-1000 acres) is a highly desir- able size for forested natural areas. Interesting natural features, geological and biological, oc- cur throughout the Oriskany outcrop area, so it is particularly crucial in this case that a large preserve be set aside. The Flintkote Company has selected an “environmental protection area” of 19 ha (47 acres) to be located along Townline Road, the southern boundary of the quarry site. This selection has been approved by the Ontario Municipal Board and by the Ontario Minister of Natural Resources. The area does contain features of geological interest but, because of its very small size and the disturbances there, it is of virtually no importance biologically. Acknowledgements Maycock, Winder and Fahselt acknowledge the financial assistance of the National Research Council of Canada. Literature Cited Anonymous. 1970. Table of extinct, rare and endangered vertebrates in Ontario. Ontario Fish and Wildlife Review 9: 15-21. Argus, G. W. and D.J. White. 1977. The rare vascular plants of Ontario. National Museums of Canada, Syl- logeus Number 14. Belot, Y., A. Baille, and J.L. Delmas. 1976. Model numérique de dispersion des polluants atmosphériques en présence de couverts végétaux. Atmospheric Environment 10: 89-98. Best, E. W. 1953. Pre-Hamilton Devonian stratigraphy of southwestern Ontario. Ph:D. thesis, University of Wis- consin, Madison, Wisconsin. Boucot, A. J. and J. G. Johnson. 1967. Paleography and correlation of Appalachian Province Lower Devonian sedimentary rocks. Tulsa Geological Society Digest 333 35-67- Brandt, C. J. and R. W. Rhoades. 1972. Effects of lime- stone dust on lateral growth of forest trees. Environ- mental Pollution 4: 207-213. Brandt, C. J. and R. W. Rhoades. 1973. Effects of lime- stone dust accumulation on composition of a forest community. Environmental Pollution 3: 217-225. Brown, W.G. E.and D.S.Lacate. 1961. Rooting habits of white and red pine. Forestry Research Branch Technical Note 108. (Available from Department of Fisheries and Environment, Ottawa.) Caley, J. F. 1940. Palaeozoic geology of the Toronto- Hamilton area, Ontario. Geological Survey of Canada, Memoir 224. pp. 83-87. FAHSELT ET AL.: ORISKANY SANDSTONE OUTCROP FEATURES 39 Campbell, C. A. 1969. Who cares for the Fowler’s Toad? Ontario Naturalist 4: 24—27. Conant, Roger. 1975. A field guide to reptiles and am- phibians of eastern and central North America. 2nd edition. Houghton Mifflin Co., Boston. Cook, F. R. 1970a. Rare or endangered Canadian amphib- ians and reptiles. Canadian Field-Naturalist 84(1): 9-16. Cook, F. R. 1970b. Endangered wildlife—reptiles and am- phibians. /n Endangered wildlife in Canada. Canadian Wildlife Federation, Ottawa. pp. 10-11. Cook, F. R. 1977. Review of the Canadian herpetological scene. Proceedings of the Symposium on Canada’s Threatened Species and Habitats. Canadian Nature Fed- eration, Special Publication Number 6. Curtis, J. T. 1959. The vegetation of Wisconsin. PEON of Wisconsin Press, Madison. Fernald, M. L. 1950. Manual of botany. American Book Company, New York. Fitch, H.S. 1963. Natural history of the black ratsnake (Elaphe o. obsoleta) in Kansas. Copeia 1963 (4): 649- 658. Fowells, H. A. 1965. Silvics of forest trees of the United States. Forest Service, United States Department of Agriculture, Handbook Number 271. Froom, Barbara. 1972. The snakes of Canada. McClelland and Stewart, Toronto. Gregory, P. T. 1977. Rare and threatened snake species of Canada. /n Canada’s threatened species and habitats. Canadian Nature Federation, Special Publication Num- ber 6. Hay, O. P. 1892. The batrachians and reptiles of the state of Indiana. Indiana Department of Geology and Natural Resources Annual Report 17: 409-602. Hewitt, D.F. and B.A. Liberty. 1972. Map 2254. In Paleozoic geology of southern Ontario. Edited by D. F. Hewitt. Ontario Division of Mines, Geological Report 105. Knudsen, G. J. 1955. Pilot blacksnake (Elaphe o. obso- leta). Wisconsin Conservation Bulletin 1955 (November): 12: Kreidler, W. L. 1964. Gas and oil developments in New York State 1963. American Association of Petroleum Geologists Bulletin 48: 778-783. Lytle, W.S. 1964. Developments in Pennsylvania. Ameri- can Association of Petroleum Geologists Bulletin 48: 784-800. Manning, W. J. 1971. Effects of limestone dust on leaf condition, foliar disease incidence and leaf surface microflora of native plants. Environmental Pollution 2: 69--76. Maycock, P. F. 1963. The phytosociology of the deciduous forests of extreme southern Ontario. Canadian Journal of Botany 41: 379-438. Minton, S.A., Jr. 1968. The fate of amphibians and reptiles in a suburban area. Journal of Herpetology 2(3--4): 113-116. Morse, M. 1904. Batrachians and reptiles of Ohio. Pro- ceedings of the Ohio State Academy of Science 4(3), Special Paper Number 9: 93-144. Parks, A. W. 1913. The palaeontology of the Guelph, Onondaga and Hamilton formations in western Ontario. Canada Geological Survey, International Geological 40 THE CANADIAN FIELD-NATURALIST Congress Field Guide Book Number 4. Parsons, H. 1976. Foul and loathsome creatures. Parks Canada, Ottawa. Rickard, L. V. 1975. Correlation of the Silurian and Dev- onian rocks in New York State. New York Science Service Geological Survey, Chart Series 24. Ricks, G. R.and R. J. H. Williams. 1974. Effects of atmos- pheric pollution on deciduous woodland. Part 2. Effects of particulate matter upon stomatal diffusion resistance in the leaves of Quercus petraea (Mattuschka) Leibl. En- vironmental Pollution 6: 87-109. Ricks, G. R.and R. J. H. Williams. 1975. Effects of atmos- pheric pollution on deciduous woodland. Part 3. Effects on photosynthetic pigments of leaves of Quercus petraea (Mattuschka) Leibl. Environmental Pollution 8: 97-106. Sanford, B.V. 1969. Geology: Toronto—Windsor area. Geological Survey of Canada, Map 1263A. Schonbeck, H. 1960. Beobachtungen zur Frage des Ein- flusses von industriellen Immissionen auf die Krank- bereitschaft der Pflanze. Berichte der Landesanstalt Bodennutzungsschutz (Bochum) |: 89-98. Vol. 93 Smith, W. H. 1974. Air pollution — effects on the struc- ture and function of the temperate forest ecosystem. Environmental Pollution 6: 111-129. Stauffer, C. R. 1915. The Devonian of southwestern On- tario. Geological Survey of Canada, Memoir 34. Stewart, Darryl. 1974. Canadian endangered species. Gage Publishing, Toronto. Stickel, W. H. and J. B. Cope. 1947. The home ranges and wanderings of snakes. Copeia 1947(2): 127-136. Telford, P.G. 1974. Dunnville area. Ontario Department of Mines, Map P988. Weetman, G. F. 1972. Canadian Institute of Forestry poli- cy for selection, protection and management of natural areas. Forestry Chronicle 48: 1-3. Wright, A.H. and A.A. Wright. 1957. Handbook of snakes of the United States and Canada. Volume I. Comstock Publishing Associates, Ithaca, New York. Received 19 June 1978 Accepted 19 October 1978 Breeding Areas and Overnight Roosting Locations in the Northern Range of the Monarch Butterfly (Danaus plexippus plexippus) with a Summary of Associated Migratory Routes FRED A. URQUHART and NORAH R. URQUHART Life Sciences, Scarborough College, University of Toronto, West Hill, Ontario MIC 1A4 Urquhart, F. A. and N. R. Urquhart. 1979. Breeding areas and overnight roosting locations in the northern range of the Monarch Butterfly (Danaus plexippus plexippus) with a summary of associated migration routes. Canadian Field- Naturalist 93(1): 41-47. Key Words: Monarch Butterfly, breeding areas, overnight roosting locations, northern range, Lepidoptera, Danaidae. Asa result of over 40 years of investigation dealing with the ecology of the Monarch Butterfly (Danaus plexippus plexippus). with special reference to its migratory habits, it is now possible to outline the major breeding areas in its northern range in Canada and to explain the absence of such areas in the western provinces. The characteristics of the overnight roosting clusters located along the migratory routes are described and compared to those of the overwintering clusters. Asummary of the migratory routes for these northern populations is presented, based on previous published records for North America. For the past 41 yr, commencing in the summer of 1937, we have been involved in a long-term study of the biology of the Monarch Butterfly, Danaus plexippus plexippus (Danaidae: Lepi- doptera), with special reference to its migratory habits. Although most of our field studies and alar tagging (Urquhart 1941, 1960) have been centered in Ontario, observations have been made in all provinces from Vancouver, British Columbia to Cape North, Nova Scotia. These field surveys were for the purpose of investi- gating the presence, or absence, of breeding areas and, where adults were found, to alar tag them for migratory studies. It is the purpose of the present paper to summarize our field data for the northern range in Canada, correlating this information with the migratory habit. Because the program has been carried out over a long period of time it has been possible to witness the change in the northern range of the Monarch Butterfly and the increase in its abundance where at one time larvae were unknown or of rare occurrence. Methods Field trips were carried out in various locali- ties in each Canadian province and notes made concerning the presence or absence of milkweed plants (Asclepias spp.), the source of food for the 41 larvae of the Monarch Butterfly, and of the larvae and adults of the Monarch Butterfly. Members of the Insect Migration Association (IMA) sent observations concerning the pres- ence of the larvae and adults of the Monarch Butterfly, together with notes as to their abun- dance in each area. An alar-tagging program (Urquhart 1960, 1965; Urquhart and Urquhart 1976b) was carried out along with field investigations and by members of the IMA in order to follow the movements of the Monarch Butterfly. Publicity arising from the recapture of alar- tagged specimens resulted in further observa- tions being sent to us by interested individuals. All observations and recapture data are on permanent file at Scarborough College, Univer- sity of Toronto. Results and Discussion Breeding Range Field surveys carried out from 1937 to 1940 indicated that breeding areas of the Monarch Butterfly in Ontario were mostly confined to the lower Great Lakes regions extending from Gananoque and Kingston in the east to Goderich in the west and south to Leamington. Large concentrations, owing to the presence of 42 THE CANADIAN FIELD-NATURALIST dense growths of the Common Milkweed (Asc/e- pias syriaca), were concentrated in the Sarnia- Leamington, Brantford-Hamilton, Toronto- Oshawa, and Belleville regions. Although larvae were collected further north at Barrie and Midland they did not occur in large numbers because growths of the Common Milkweed were not as dense as those found further south. As a result of the increase in the number of roads and highways and the clearing of forested areas for power lines, the Common Milkweed has gradually spread northward thus increasing the amount of this host plant for the larvae. By 1975 records of large concentrations of both larvae and adults were reported from various locations along the north shore of Lake Superior, particularly at Sudbury, Sault Ste. Marie, and Thunder Bay. Inthe early summer of 1977 larvae were more abundant in the Sault Ste. Marie area than in the T oronto-Oshawa area; we had larvae mailed to our laboratory in Toronto from Sault Ste. Marie in order to carry out various research projects. The reason for this difference in the numbers of Monarch Butterfly larvae in the Lake Superior regions in the early summer as com- pared to the Toronto-Oshawa area can be explained. We have previously shown (Urquhart and Urquhart 1976d, 1977) that Monarch Butterfly migrants travel in a northwesterly direction in the autumnal migration and north- easterly in the vernal. In this way the migrants reach the north shore of Lake Superior, via Michigan, before arriving in the Ontario penin- sular regions. Since the Common Milkweed 1s now abundant in the Lake Superior regions larger populations of butterflies have occurred here in early summer than further south. Field surveys each summer from 1962 to 1977 from Toronto north to the northeast shore of Georgian Bay disclosed increasing abundance of the Common Milkweed covering hundreds of hectares along roads, highways, and unculti- vated fields. Also, as a result of the clearing of forested areas, with particular reference to power lines, ever increasing areas have been made available for the further spread of the larval food plant. One experimental plot of 0.5 ha located in a forested area, that had been cleared during lumbering operations, revealed a population of 536 larvae on 15 July 1977. Vol. 93 For other Canadian provinces records accumulated over the past 41 yr (including 4 yr of field investigations of 1940-1944 in the western provinces and surveys in the eastern provinces together with numerous reports from various interested individuals and members of the IMA) indicate conclusively that there are no breeding areas in British Columbia and Alberta. A few scattered larval populations have been reported for southern Saskatchewan (Duval) and southern Manitoba (Transcona, Furness). Although there have been numerous records of adults seen in Nova Scotia, New Brunswick, Prince Edward Island, and occasionally in Newfoundland, we have no records of any breeding populations there. Three reports have been received from Quebec (Montreal, Drum- mondville, Quebec). The marked differences among the population numbers in the various provinces are due in part to the migration routes and in part to the distribution of the species of milkweed. Since the migration tends to a northeast-southwest direc- tion from the overwintering Mexican Site (Urquhart and Urquhart 1976d) the migrants miss the western provinces with only a few stragglers being reported. Similarly, the eastern provinces are outside the regular migratory route. Although species of milkweed of the genus Asclepias are found in all provinces, except Prince Edward Island, Labrador, and Newfoundland, the majority of species are found in Ontario where Asc/epias syriaca occurs in the greatest abundance. One species is reported for British Columbia (A. speciosa); there are no records from Alberta: six occur in Manitoba (A. incarnata, A. verticillata, A. ovalifolia, A. syriaca, A. speciosa, A. viridiflora); two occur in Saskatchewan (A. ovalifolia, A. speciosa): 10 occur in Ontario (A. incarnata, A. verticillata, A. tuberosa, A. exaltata, A. quadrifolia, A. syriaca, A. sullvantii, A. purpurescens, A. hirtella, A. veridiflora); two species occur in Quebec (A. incarnata, A. syriaca); one species 1s recorded from New Brunswick (A. syriaca); and one species from Nova Scotia (A. incarnata). From the above records of the distribution of species of the larval food plant and the direction of migration, it is obvious why there 1s by far the largest population in the northern range located in the province of Ontario. SU) Overnight Roosting Clusters During the autumnal migration the Monarch Butterflies cluster on trees of various species along the migratory routes to remain during the night period (Urquhart 1960). Of the species of deciduous trees chosen Red Maple (Acer rub- rum), Sugar Maple (Acer saccharum), Mani- toba Maple (Acer negundo), and willow (Salix sp.) were most common. Pine (Pinus sp.) and spruce (Picea sp.) were most frequently chosen since it was possible for the butterflies to obtain a secure hold with the sickle-shaped tarsal claws (Urquhart 1960). During periods of calm weather individuals of a cluster are widely spaced (Figure 1). During stormy conditions with strong winds the migrants cluster closer together and there are fewer clusters on the trees. The massing together URQUHART AND URQUHART: MONARCH BUTTERFLY 43 under conditions of strong winds has the distinct advantage of causing the weighted branch of the tree to sway in the wind rather than being whipped about, which would dislodge the migrants. The leeward side is always chosen as a protection against the wind. Since migrants tend to cluster on certain trees year after year, it has been thought that perhaps an odor of some kind may have been left the previous year thus acting as a guide for future generations of migrants. Numerous observa- tions in many localities, however, have not produced evidence to substantiate such a sug- gestion. For example, when a particular group of roosting trees was removed, the migrants selected another group that had not been used previously. As has been demonstrated, the direction of migration, the presence of suitable FIGURE |. Overnight roosting Monarch Butterflies during the autumnal migration showing the widely scattered nature of the clusters. 44 THE CANADIAN FIELD-NATURALIST trees en route located near nectar-producing flowers such as the Canada Goldenrod (Soli- dago canadensis) and the New England Aster (Aster novae-anglia), and topography are the factors determining the choice of roosting trees (Urquhart 1960). Although overnight clusters may occur in numerous localities throughout southern On- tario, they are most abundant and composed of larger clusters along the north shores of Lake Ontario and Lake Erie. This is owing to the migrants’ antipathy to fly over large bodies of water beyond sight of land; hence they move in ever increasing numbers along the lake shores (Urquhart 1960, 1966, 1976a). The manner in which the migrants cluster ona particular part of the roosting tree or trees is as follows. The first arrival flies around the tree, testing wind direction and a suitable site upon which to land. Many tests are made before a final location is chosen. Having finally landed the butterfly opens the wings to display the more brilliantly colored dorsal surfaces. This acts as a beacon to others that in turn land and open their wings. If one migrant lands too close to one already at rest the latter snaps its wings in a warning gesture which acts as a further stimulus for others to land upon the particular branch. Eventually a cluster is formed. When the migrants reach their final over- wintering destination in the Neovolcanic Moun- tain site of Mexico (Urquhart 1976b; Urquhart and Urquhart 1977), they form dense clusters that are quite unlike those of the overnight roosts (Figure 2). Migrants on the overnight roosts, as mentioned previously, occur as small scattered clusters, the individuals widely spaced; they do not cluster on the trunks of the trees or upon adjacent bushes or upon the ground. Inthe overwintering clusters, the roosting trees are so densely covered that it is not possible to see the foliage of the trees; the trunks are also densely covered as well as the neighboring bushes. In some loci! the ground is also covered (Urquhart 'Site refers to the geographic location of an overwintering site, such as the Mexican Site, the California Site, etc.; Area refers toa particular geographic location where a number of clusters are to be found, such as a volcanic mountain in Mexico or the Monterey Peninsula in California; Locus (1) refers to a particular location within an area where a cluster exists which may change from year to year, suchas lociinan area in Mexico or in the Monterey Peninsula. Vol. 93 and Urquhart 1977: Urquhart and Urquhart 1976a, b, d, 1977). This difference between the two clusterings is due to the massed millions of migrants in the overwintering roosts as com- pared to the smaller clusters scattered over a wide area along the migratory routes. Migratory Routes The Monarch Butterflies migrate from the breeding areas in Canada mainly to Mexico where they overwinter in various loci in various moun- tain areas of the Mexican Site in the Neovolcanic Mountains, sometimes referred to as the “Cross Range” (Urquhart and Urquhart 1979). Migrants from west of Lake Superior move SSW (190°): those from the north shore of Lake Huron move SSW (200°) and those from extreme southwestern Ontario move SSW (215°) (Figure 3, route D). Migrants from the rest of Ontario and western Quebec indicate two flight patterns. The greater number move SSW (200°) to the coast of the Gulf of Mexico and thence westward (270°) (Figure 3, route C) following the Gulf coast eventually towards the overwintering site (Figure 3, route E) on a tra- jectory SSW (195°). Others, perhaps as a result of strong westerly winds during the flight period, move SSE (110°) (Figure 3, route A) to the Atlantic coast. Following the coast line SSW (195°-210°) the majority reach the coast of the Gulf of Mexico where, following the coast line, they travel westward (Figure 3, route C). Others, representing a small proportion of the migrating population, termed an “aberrant” population (Urquhart 1976a, b: Urquhart and Urquhart, 1976c, 1977), move down the Florida peninsula SSE (140°), thence to Cuba and Yucatan SSW (240°) (Figure 3, route B) or, continuing SSE (120°-140°) become scattered over the islands of the Antilles. The final destination of this aberrant population is as yet unknown. Vernal Migration Vernal migrants leave the Mexican Site in late February and March (Urquhart and Urquhart 1979). Mating takes place at the site, when the clusters break up, and along the migratory routes as far as central Texas — males rarely proceed further. They enter the various Cana- dian provinces, especially Ontario, commencing in the last week of May through June and early July. During late June and early July second- 1979 URQUHART AND URQUHART: MONARCH BUTTERFLY 45 FIGURE 2. Overwintering roosting Monarch Butterflies on a tree in the Neovolcanic Plateau Site of Mexico showing the compact nature of the clusters completely covering the branches and trunks of one of over a thousand trees. 46 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE 3. Migration routes of the Monarch Butterfly from the breeding areas in Canada to the overwintering site in Mexico and the aberrant migration routes to Yucatan and the Antilles. @ breeding areas; * major overnight roosting areas; Aoverwintering site;—sdirection of flight. A, flight pattern to the Atlantic coast: B. flight pattern through Florida to Yucatan: C, flight pattern along the north coast of the Gulf of Mexico; D, flight pattern from the western provinces, north of Lake Huron and southwestern Ontario; E, overwintering Mexican Site; F, possible flight direction towards the Antilles. 1979 generation adults, resulting from eggs deposited on species of milkweed by migrating females, enter Canada. These are readily identified by their brighter coloration, those from the Mexi- can Site being decidedly faded with tattered wing margins (Urquhart 1966). First-generation adults from breeding areas in Canada appear from mid-July through August. A second generation occurs in the more southern sections from mid-August through September. Autumnal migration commences in mid- August, the numbers increasing through late August and September and, under conditions of high temperature, through October. A few stragglers have been collected in November and three specimens during the first week of Decem- ber. It is highly unlikely that these late migrants reach the overwintering sites. Acknowledgments Financial support for the studies of the biology of the Monarch Butterfly has been received from the National Research Council of Canada; the National Geographic Society, Committee for Research and Exploration; and the Insect Migration Association. Literature Cited Urquhart, F. A. 1941. A proposed method for marking migrant butterflies. Canadian Entomologist 1941 (February): 21-22. Urquhart, F. A. 1960. The Monarch Butterfly. University of Toronto Press, Toronto. 361 pp. Urquhart, F.A. 1965. Monarch Butterfly (Danaus plexippus) migration studies: autumnal movement. URQUHART AND URQUHART: MONARCH BUTTERFLY 47 Proceedings of the Entomological Society of Ontario 5: 23-33. Urquhart, F. A. 1966. A study of the migration of the Gulf Coast populations of the Monarch Butterfly (Danaus plexippus L.) in North America. Annales Zoologici Fennici 3: 82-87. Urquhart, F. A. 1976a. Migration of butterflies along the gulf coast of northern Florida. Journal of the Lepidop- terists’ Society 30: 59-61. Urquhart, F. A. 1976b. Found at last: the monarch’s winter home. National Geographic Magazine 150: 161-173. Urquhart, F. A. and N.R. Urquhart. 1976a. Monarch Butterfly (Danaus p. plexippus L.) overwintering popula- tion in Mexico (Lepidoptera: Danaidae). Atalanta 7: 56-60. Urquhart, F. A. and N.R. Urquhart. 1976b. Ecological studies of the Monarch Butterfly (Danaus p. plexippus L.). National Geographic Society Research Report 1976: 437-443. Urquhart, F. A. and N.R. Urquhart. 1976c. A study of peninsular Florida populations of the Monarch Butterfly (Danaus p. plexippus L.; Danaidae). Journal of the Lepidopterists’ Society 30: 73-87. Urquhart, F. A. and N.R. Urquhart. 1976d. The over- wintering site of the eastern population of the Monarch Butterly (Danaus p. plexippus L., Danaidae) in southern Mexico. Journal of the Lepidopterists’ Society 30: 153-158. Urquhart, F. A. and N. R. Urquhart. 1977. Overwintering areas and migratory routes of the Monarch Butterfly (Danaus p. plexippus; Lepidoptera: Danaidae) in North America, with special reference to the western popula- tion. Canadian Entomologist 109: 1583-1589. Urquhart, F. A. and N. R. Urquhart. 1979. Vernal migra- tion of the Monarch Butterfly (Danaus p. plexippus L.; Lepidoptera: Danaidae) in North America from the overwintering site in the neovolcanic plateau of Mexico. Canadian Entomologist (In press). Received 25 May 1978 Accepted 14 August 1978 Reproductive Biology of the Big Brown Bat (Eptesicus fuscus) in Alberta DAVID B. SCHOWALTER and JOHN R. GUNSON Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2 Schowalter, D. B. and J. R. Gunson. 1979. Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta. Canadian Field-Naturalist 93(1): 48-54. Data on Big Brown Bat (Eptesicus fuscus) populations in Alberta were collected from 1972 to 1977 during surveys to determine the prevalence of rabies. The 60 maternity colonies located included 56 in older buildings; many of these sites were used as hibernacula. Most young were born during the latter part of June; parturition was estimated to extend from at least 5 June to 12 July. Fifteen of 115 pregnancies were twins. Ages of a sample of bats were determined by counts of dental annuli; those ages generally correlated with tooth wear. Yearling females had a lower pregnancy rate than older females. Key Words: Big Brown Bat, life history, dental aging, Alberta. The Big Brown Bat (Eptesicus fuscus) 1s relatively abundant and widely distributed over North America (Hall and Kelson 1959; Barbour and Davis 1969). Most studies of the species have been in eastern North America (Christian 1956; Brenner 1968; Davis et al. 1968; Barbour and Davis 1969; Kunz 1974; Mills et al. 1975) where the usual litter size is two. Comparatively few data are available for the species in western North America, particularly western Canada, where the usual litter size is evidently one (Cockrum 1955; Christian 1956; Kunz 1974). A bat-rabies monitoring program in Alberta was prompted by the first diagnosis of a rabid bat in the province in 1971, and an outbreak of rabies among other animals in 1970 and 1971. Epidemiological results of this program have recently been presented by Dorward et al. (1977); biological data on Ef. pallidus in Alberta collected during the course of the study are presented here. Methods Colonies were located through the following means: complaints of bats relayed by govern- mental personnel and systematic surveys during 1971 to 1977; newspaper advertisements in 1971 and 1972; results of a questionnaire mailed to rural homeowners in 1973; and reviews of histories of rabies-suspect bats. Collections of 10 to 20 bats for rabies testing were made at a number of maternity roosts; larger collections 48 were made at two maternity roosts and a fall roost, all of which were known to have harbored rabid individuals. Bats were collected within colonies by hand, forceps, and a modified Constantine trap (Constantine 1958); at colony exits by a variety of enclosure traps; and by mist-nesting away from colonies. Colonies were generally visited only once, although two or three visits per year were made to one colony to band bats. Available for analysis were 256 Big Brown Bats submitted as rabies-suspect indi- viduals from 1974 to 1977. Age (adult or juvenile) as determined by closure of the epiphyses of the fingers, and sex were noted for each bat. Reproductive status of adult females was recorded, as was the number and sex of fetuses of parous individuals. Ages of adult bats taken at two maternity roosts, a fall roost, and a sample of rabies-suspect individuals were determined from counts of dental annuli (Schowalter et al. 1978). The first of January was arbitrarily used as the date on which bats became a year older: thus a bat classified as a l-yr-old was taken the summer after the summer of its birth. Tests for significance were by the simple chi-square test. Results and Discussion Big Brown Bats appeared to be more abun- dant in southern than central Alberta (Figure 1). We discovered 32 colonies in southern Alberta and 28 colonies in the larger area of central 1979 HINTON Colony Size @® 1-25 @ 26-250 ee Over 250 (0) 50 100 Km +4 FIGURE |. Distribution of known Alberta. Although estimates of the number of bats were approximate, colonies were evidently larger in the south, where the average number of adults was 80. In central Alberta average colony size was 44. Mist-net captures indicated that Big Brown Bats were relatively abundant in natural habitats along river valleys in the southern area of the province. Much of this apparent greater abundance in the south is related to the occurrence of maternity roosts in the cities of Medicine Hat and Lethbridge. We have seldom located maternity roosts of Big Brown Bats, or other species, in more northerly cities in Alberta. A search of 28 old buildings in Edmonton, mostly schools, which were similar to buildings SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 49 -— _—_— — = aw = =, =- EOMONTON a e e C ) S O CAMROSE e @ ° @ @ e® ® e @ g RK) CALGARY Big Brown Bat colonies in Alberta. frequently colonized in Medicine Hat and Lethbridge, produced no evidence of active maternity colonies. A strong preference was evident for the formation of maternity colonies in older build- ings; 56 of 60 buildings with Big Brown Bat maternity colonies were built prior to 1925. Banding studies (Beer 1955; Davis et al. 1968; Barbour and Davis 1969) have demonstrated strong site attachment of Big Brown Bats. Failure to colonize newer buildings may in- dicate a decreased or stable population that is strongly attached to roosts currently in use. Alternatively the variable environment in older buildings may meet the physiological needs of 50 THE CANADIAN FIELD-NATURALIST the bats better, by permitting both summer and winter roosting. Many nurseries were evidently utilized as hibernacula by at least a few (and often many) bats in Alberta, as noted by others (Mills et al. 1975; Hitchcock 1949). Environ- mental conditions in newer buildings are likely to be different as a result of recent changes in heating systems and insulation standards. Big Brown Bat roosts were similar to those described elsewhere (Barbour and Davis 1969; Mills et al. 1975); those we observed were generally cooler than Little Brown Bat (Myotis lucifugus) maternity roosts inspected. Big Brown Bats were extremely tolerant of light in some roosts. Many owners of the buildings that were used as maternity roosts by Big Brown Bats believed that bats were resident during the winter, although only 19 reported having seen bats in this period. Other buildings, particularly in the city of Edmonton, that were not maternity roosts were utilized by wintering bats. Our winter captures of Big Brown Bats, and citizen submissions of bats for rabies testing from Edmonton, far exceeded those of summer. On the other hand, numerous summer submissions were made of Little Brown Bats and a few Silver- haired Bats (Lasionycteris noctivagans) (Dor- ward et al. 1977; this study, unpublished data). These observations suggest that Big Brown Bats moved into Edmonton to hibernate. Five buildings in Edmonton and one non- maternity roost building in Camrose and Hinton regularly were occupied by bats in winter. As many as six bats captured in one winter at two of these sites were known to us; however, it is unlikely that most bats encountered in these buildings were reported. Winter occurrences were frequently associated with the onset of cold weather. In addition Big Brown Bats have been discovered during the course of building demoli- tion and remodelling; and in one case, loud rock bands in a high school gymnasium were con- sidered almost certain to cause bat activity. As noted by Barbour and Davis (1969) E. fuscus is extremely cold-tolerant. Two indi- viduals were captured outside a building on 3 December 1975. Temperature at time of capture was near —15°C, and had been even lower earlier in the day. One bat was alert and shivering; the other was torpid, but quickly became active Vol. 93 when warmed. The bats had roosted near a recently filled crack and may have been attempt- ing to enter the building. Timing of events at maternity roosts was difficult to define as visits were made ir- regularly. It was evident, however, that there was variation between colonies, and probably be- tween years. Bats were active at one location, which was thought to be a hibernaculum as well, on 8 April 1976 and 13 April 1977. The nature of bat activity at these times was unknown as the apparent food supply was extremely limited. One maternity site was determined to be without bats as late as 10 May 1977. Roosting in four types of groups (pregnant females, females with naked young, furred young, and lactating females) was observed in one maternity roost. Similar separate roosting, although usually in only two or three types of groups, was noted in other colonies. This grouping created uncertainty as to the timing of parturition, as generally not all the bats in a roost were known to have been observed. These groupings were thought to be related to differ- ing thermal requirements of individuals at different stages of reproduction and growth. Time of parturition varied considerably both in, and between, colonies. Fetal examination and observation of neonates indicated that most young were born in the latter part of June. The earliest estimated birth date was 5 June. Near- term pregnant females were taken as late as 12 July. Of 114 adult females taken from maternity roosts during late May and early June, 105 were pregnant. Of 115 bats examined in advanced pregnancy, 15 had twins, which corroborates Christian’s (1956) finding that one is the usual litter size in western North America. Sixty-two of 84 single young had implanted in the right horn of the uterus. Kunz (1974) also noted a tendency towards implantation in the right horn. Fetal sex ratio was near 50:50 (Table 1). During 1974 and 1975 significantly more adult females (P< 0.05) than adult males were sub- mitted for rabies testing (Table 1). During these two years public concern about bat rabies was high; more than 400 bats were submitted per year. It appeared that many normal-acting bats were submitted as rabies suspects from mater- nity roosts. Most bats submitted in 1976 and 1979 SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 5] TABLE |—Age-specific sex ratio in samples of Epresicus fuscus from Alberta, 1972 to 1977 Sample Adults Juveniles Fetuses No. % Male No. % Male No. % Male Colony survey 20 May-31 July 261 3.1 100 45.0 73 S23 1 Aug-30 Sept 69 23.2 79 26.6 Rabies suspect 1974-1975 107 36.4 46 60.9 1976-1977 58 S522 45 71.1 1977, when approximately 200 bats a year were submitted, were either grounded or found in other atypical situations. In those two years the numbers of adult males and females submitted were similar (Table 1). Juvenile males tended to leave colonies earlier than juvenile females during late July and August. They occurred significantly less fre- quently (P< 0.01) than juvenile females in our samples from buildings during August and September (Table 1). This earlier departure may have subjected them to higher mortality, as they were submitted for rabies testing more frequently (P< 0.05) than juvenile females (Table 1). Results of dental aging (Figure 2) indicated a life-expectancy similar to that found by Goeh- ring (1972) from a 20-yr banding study in a Minnesota hibernaculum. Our samples varied in age structure, particularly in the proportion of I- yr-old bats (Figure 2). One-year-olds constituted 39% of the rabies-suspect sample; the sample was comprised of bats from the greatest variety of situations and capture dates, and may be the most representative of the population age structure. It is, however, made up of bats that died from natural or human causes. One-year- old females from maternity colonies made up only 16% of the sample (Figure 2). That only 15 of 31 l-yr-old rabies-suspect and survey females were parous or had suckled young demonstrated that many lI-yr-old females are non-parous as noted by Christian (1956) and Barbour and Davis (1969). Non-parous 1|-yr-old females may tend not to roost in maternity colonies and may be more likely to be submitted as rabies-suspect bats than older females. Ages of bats from Connaught School (Figure 2) have been discussed by Schowalter et al. (1978). Younger animals appeared to be under- represented in that sample compared to the other samples and what would be expected from banding results (Goehring 1972). Estimates of the total number of adults in the Connaught colony in June ranged from 500 to 1000, comparable to the largest colonies noted by Mills et al. (1975). Those authors determined that there was an unknown density-dependent mechanism functioning to regulate populations in large colonies. The relatively small number of individuals in some of the younger age-groups in the Connaught sample suggests low survival of young or that a large proportion of juvenile females disperse from the colony prior to their second year. The pregnancy rate of adult females from Connaught, 45 of 51, while less than that of females from other colonies (P< 0.05), is not low enough to suggest that the regulatory mechanism operates primarily through limiting reproduction. Tooth wear has been considered as a means of aging bats (Twente 1955; Christian 1956; Stege- man 1956). Mills et al. (1975) found a highly: significant relationship between tooth wear and ages of 208 Big Brown Bats banded as im- matures. They observed wide variation in wear rates of individual bats. One of us (JRG) recorded tooth wear of bats in the Connaught School sample. Although analysis was limited by the subjective nature of the estimates of tooth wear and the possibility of errors in aging by annuli, there was general agreement between the two methods (Figure 3). Tooth wear would appear to be potentially useful as a method of determining relative population-age structures of Big Brown Bats in Alberta. The possibility of a population decline and the factors governing the growth and size of colonies S52 THE CANADIAN FIELD-NATURALIST Vol. 93 Connaught School Maternity Sample n=51 Stony Plain Maternity Sample White Elephant Fall sample =43 Percent of Sample Rabies Suspect n= 49 12 Si waa eal 7emoie > elOm lal l2ilS ue lAghlow Onl Zale Age Group FIGURE 2. Distribution of ages of samples of Big Brown Bats from Alberta (determined by counts of dental annuli). Specimens from Connaught School, Medicine Hat; Stony Plain; and White Elephant Theatre, Bow Island. Numerals at base of bars indicate number of bats. 1979 —~-~NWAU—NWEUNUAN DWDODO,H NW EN AN © DO O Number of Bats EF] EI = eo es SCHOWALTER AND GUNSON: BIG BROWN BAT, ALBERTA 53 Light wear n=19 x=2.9 Moderate wear n= 19 x= 5.8 Heavy wear n= 13 x= 9.) De Si Ae Ol 72a, LOnill 2) 1S 42 lo Onl7a18" 19-20, Dental Age Group FIGURE 3. Comparison of amount of tooth wear and ages, as determined by counts of dental annuli, of Big Brown Bats from Connaught School. in central and southern Alberta need further investigation. Dental aging appears to offer a powerful tool in such investigations, although this technique requires further evaluation. Acknowledgments The interest and support of H. Vance and G. Whenham of the Veterinary Services Division, Alberta Department of Agriculture are greatly appreciated. The Agriculture Canada Animal Diseases Research Institute (Western) at Leth- bridge, Alberta has provided continued co- operation in allowing access to specimens and records of rabies-suspect animals; W. Dorward, J. Bradley, H. Boumans, B. Prins, and D. Meyers of that institution contributed in various ways to this study. Field and laboratory assis- tance were provided by L. Harder, B. Treichel, L. Dube, P. Cole, W. Johnson, and W. Wynnyk. N. Previsich contributed generously of his time and energy. We thank A. Todd and L. Harder for comments on an earlier draft. Literature Cited Barbour, R. W. and W. H. Davis. 1969. Bats of America. University Press of Kentucky, Lexington. 286 pp. Brenner, F. J. 1968. A three-year study of two breeding colonies of the Big Brown Bat, Eptesicus fuscus. Journal of Mammalogy 49: 775-778. 54 THE CANADIAN FIELD-NATURALIST Beer, J. R. 1955. Survival and movements of banded Big Brown Bats. Journal of Mammalogy 36: 242-248. Christian, J.J. 1956. The natural history of a summer aggregation of the Big Brown Bat, Eptesicus fuscus fuscus. American Midland Naturalist 55: 66-95. Cockrum, E. L. 1955. Reproduction of North American bats. Transactions of the Kansas Academy of Sciences 58: 487-511. Constantine, D. G. 1958. An automatic bat-collecting de- vice. Journal of Wildlife Management 22: 17-22. Davis, W. H., R. W. Barbour, and M.D. Hassell. 1968. Colonial behavior of Eptesicus fuscus. Journal of Mam- malogy 49: 44-50. Dorward, W.J., D.B. Schowalter, and J. R. Gunson. 1977. Prelimary studies of bat rabies in Alberta. Can- adian Veterinary Journal 18: 341-348. Goehring, H.H. 1972. Twenty-year study of Eptesicus fuscus in Minnesota. Journal of Mammalogy 53: 201- 207. Hall, E.R. and K.R. Kelson. 1959. The mammals of North America. Volume |. Ronald Press, New York. 546 pp. Vol. 93 Hitchcock, H. B. 1949. Hibernation of bats in southeastern Ontario and adjacent Quebec. Canadian Field-Naturalist 63: 47-59. Kunz, T. H. 1974. Reproduction, growth and mortality of the vespertilionid bat Eptesicus fuscus in Kansas. Journal of Mammalogy 55: 1-13. Mills, R. S., G. W. Barrett, and M. P. Farrell. 1975. Pop- ulation dynamics of the Big Brown Bat ( Eptesicus fuscus) in southwestern Ohio. Journal of Mammalogy 56: 591— 604. Schowalter, D. B., L. D. Harder, and B. H. Treichel. 1978. Age composition of some vespertilionid bats as deter- mined by dental annuli. Canadian Journal of Zoology 56: 355-358. Stegeman, L. C. 1956. Tooth development and wear in Myotis. Journal of Mammalogy 37: 58-63. Twente, J. W., Jr. 1955. Aspects of population study of cave-dwelling bats. Journal of Mammalogy 36: 379-390. Received 14 June 1978 Accepted 18 September 1978 Wild Mallard Stocking in a Large Marsh Habitat ROBERT O. BAILEY Macdonald College of McGill University, Ste. Anne de Bellevue, Québec HOA !CO Bailey, R.O. 1979. Wild Mallard stocking in a large marsh habitat. Canadian Field-Naturalist 93(1): 55-62. During 1971 and 1972, 1204 female and 214 male wild-strain Mallard (Anas platyrhynchos) ducklings were released on the Delta Marsh to test Mallard stocking as a method to increase breeding populations. Hand-reared yearling hens arrived after most of the unmarked hens in spring. Homing to the release site was observed in 26-28% of yearling hens and 53% of 2-yr-olds. Yearling marked hens initiated nests later and were less successful than unmarked hens in producing broods. Differences between the marked yearlings and unmarked breeders were attributed to the presence of the adult hens in the unmarked portion of the breeding population. In view of the poor reproductive success of hand-reared birds and apparently high potential for natural immigration and production, Mallard stocking is a questionable procedure on the Delta Marsh. Key Words: Mallard, stocking, spring arrival, homing, nest initiation, productivity, breeding success. Early propagation efforts involved mainly the release of semi-domestic strains of Mallard (Anas platyrhynchos) ducklings. Lincoln (1934), Errington and Albert (1936), Benson (1939), Foley (1954), Hunt et al. (1958), and Schlad weiler and Tester (1972) concluded that releases of game farm birds were generally unsuccessful, because of the inability of these birds to survive and reproduce in the wild, and because they were extremely vulnerable to hunting. Stocking pro- grams using wild-strain birds, however, have been successful in establishing Mallards (Foley et al. 1961; Lee and Kruse 1973) and Wood Ducks (Aix sponsa) (Doty and Kruse 1972) on vacant or understocked habitat. In 1969 the Delta Waterfowl Research Station began to study the potential of releasing hand- reared wild Mallard ducklings to increase local breeding populations. Sellers (1973) made sev- eral releases into pothole habitat near Minne- dosa, Manitoba, and the breeding population increased from 12 pairs in 1969 to 66 pairs in 1971 on his 2.56-km2 study area. But only 9% of the homing (marked, hand-reared) females at Minnedosa produced broods in 1971. Sellers attributed poor production to severe nest preda- tion associated with a lack of nesting cover. Upland nesting cover had been removed by agricultural tilling and burning. Based on those findings, the present study was initiated in 1971 on the Delta Marsh, where agricultural land use is negligible and nesting cover appeared ade- quate. The objective of this study was to determine whether hand-reared Mallards would home to a specific release site in the Delta Marsh and, if so, would reproduce at a level to maintain their numbers. When it became evident in 1972 that yearling hens were not producing broods, the emphasis of the study was changed to a comparison of productivity in hand-reared and wild Mallards on the same areas. Study Area The Delta Marsh is a 140-km?2 expanse of shallow bays, creeks, and potholes at the south end of Lake Manitoba (Figure 1). The north edge of the marsh is separated from the lake bya sandy wooded ridge. Dense stands of Yellow Cane (Phragmites australis) interspersed with patches of White-top (Scolochloa festucacea) meadow cover most of the dry surface from the ridge to the bay edges. Cattail (Typha /atifolia) grows in dense stands around potholes, creeks, and on flooded bay shorelines. Bulrush (Scirpus spp.) is common in large shallow bays. Addi- tional description of topography, climate, and vegetation is given by Bird (1961, p. 19), Love and Love (1954), and Anderson and Jones (1976). In 1971 duckling releases were made at the Diversion, 4.8 km W of the village of Delta. This study area is 2.56 km? in size, divided by the Assiniboine River Diversion. Creeks, potholes, and borrow pits are typical wetlands on the area. In 1972 an additional release site was estab- 56 THE CANADIAN FIELD-NATURALIST ASSINIBOINE Voles93 STUDY AREAS | DIVERSION 2 AIKENS BAY 3 COOKS CREEK @PAIRS OF YELLOW SADDLED MALLARDS 1 KM +—4 FiGure |. Map of the Delta Marsh illustrating the study areas and the location of marked females established outside the release sites in 1973. lished at Aikens Bay, a 0.8-km? area, 8 km E of Delta. The aquatic and upland habitat at Aikens Bay was managed for waterfowl (mainly by water-level regulation) and was considered to be high quality. An unknown number of wild Mallards occupied each release site prior to stocking. A 2.56-km? area at Cooks Creek Meadow, 4.8 km E of Delta, was chosen for a control (1.e., no birds released) in 1972. That area is typical marsh habitat. In 1973 water levels were lower than usual on parts of the Delta Marsh. Wind tides from Lake Manitoba maintained water levels in the larger bays, but lack of precipitation left many nearby potholes and shallow marshes dry. Precipitation at Portage la Prairie was 7.62 cm lower than the long-term mean for 6 mo prior to 30 March 1973 (Atmospheric Environment Monthly Reports). Methods Ducklings were hatched 1n an incubator, from eggs taken from a captive flock originating from eggs gathered in the wild and hatched at Delta. Ducklings were hand-reared (Ward and Batt 1973) until 4-5 wk of age before release. Each duckling was fitted with an individually num- bered plastic nasal saddle and a standard U.S. Fish and Wildlife Service leg band immediately prior to release. Birds were liberated in groups of 10-15 throughout all wetlands on the study areas. In 1971, 456 females and 180 males were released at the Diversion. In 1972, 503 females and 34 males were liberated at Aikens Bay and 245 females at the Diversion site. Mallard breeding densities were estimated by checking on foot each wetland on the study area. Each Mallard observed was checked for a nasal saddle. Counts were conducted as suggested by Dzubin (1969). A 12.87-km roadside transect was run 5-6 times a week on the Diversion and surrounding area each year and several walking transects were conducted at Aikens Bay. These transects provided information on the ratio of marked to unmarked females, homing hens, and the male:pair ratio. The ratio of marked to unmarked females was used to obtain an estimate of marked hens represented by drakes. Sex ratio of Mallards counted prior to 15 1979 April was applied to lone males and groups of five or less to correct breeding pair estimates for the unmated drake cohort. I assumed that all Mallards present were counted during each census. Several square kilometres of marsh sur- rounding the study areas were searched each year for marked birds. Locations of marked hens were plotted on a map and revisited until the hen was identified or had disappeared. In 1973, nest searches were carried out 2-3 times a week between 09:00 and 12:00 from the second week of April to the beginning of July. Two observers and a Labrador retriever partici- pated in each search. Nests were located, plotted on a map, and marked witha piece of fluorescent tape on vegetation or a stake 3-4 m away. Nest initiation date was estimated for each nest by back-dating eggs in incomplete clutches (Dane 1966). Eggs in nests containing down were floated (Westerskov 1950) to estimate the ap- proximate stage of’ incubation. Nests were revisited after the anticipated hatching date and nests containing finely crushed egg-shells and membranes were considered hatched. Brood searches consisted of walking the emergents surrounding wetlands. The number and age of young (in weeks) and the marking, if any, on the female were used to avoid duplica- tion in counts. This technique was supplemented with morning and evening road transects, walking transects, and observation periods wherein one block of habitat was observed for 2-3 h. Broods in Aikens Bay were observed from a 6-m tower in a central location on the site. In spring 1973, 15 paired Mallard females were collected on the marsh, at least 2 km away from the study areas, in an attempt to estimate the proportion of adults in the unmarked population. Only females from isolated pairs were shot. Hens were qualitatively classed as yearling or adult based on examination of wing feathers (Carney and Geis 1960; Hopper and Funk 1970). Wings from known adults were examined for comparison. Results Spring Arrival and Breeding Densities The first Mallards (six) in 1972 arrived on the Diversion on 7 April, and did not include any BAILEY: WILD MALLARD STOCKING 57 TABLE I—Mallard breeding pair counts (with 95% con- fidence intervals) on the Diversion and Aikens Bay, Delta, Manitoba. Pairs are corrected for sex ratio. Pairs Study area 1971 1972 Total released released unmarked Diversion 5 May 1972 7 ae 7/ — 1947 36 17 May 1972 19+5 = 6} a8 5) ay Means 18.0 13.5 32 1 May 1973 644 9+4 1145 26 17 May 1973 9+5 10+6 WD) ae 7/ 30 Means ES 9.5 LES 28 Aikens Bay 26 April 1973 —— 1444 il ae 5 35 8 May 1973 — 1344 9+5 22 Means 13.5 15.0 29 marked hens. On 13 April the first marked female and her mate were observed in a flock of five pairs on the study area. Most of the marked Mallards did not arrive until the week starting 21 April, when the mean number of marked hens observed per road transect increased to 3.85 from 0.58 the previous week. Over the same time period the mean number of unmarked hens observed per road transect decreased from 17.4 to 10.6. Spring 1973 was phenologically very early, and the first Mallards arrived at Delta on 21 March. On 27 Marcha marked yearling hen was seen at Aikens Bay (Peter Ward, personal communication). When I arrived on 12 April, all hens were back. The breeding pair censuses (Table 1!) were timed to correspond with Mallard nest initiation dates as indicated by increases in the male: pair ratio along roadside transects. Prior to 15 April 1972, 496 Mallards were seen with a sex ratio of 57.3 males to 42.7 females. Prior to 15 April 1973, 543 Mallards showed a sex ratio of 54.7 males to 45.3 females. Mallard breeding pairs on the Diversion averaged 32 per 2.56 km? in 1972 and 28 in 1973 (Table 1). Aikens Bay contained 29 pairs of Mallards (93 pairs per 2.56 km?2). Surveys commencing 14 and 21 May 1973, on Cooks Creek Meadow revealed a population of 54 and 55 pairs of Mallards per 2.56 km? respectively. 58 THE CANADIAN FIELD-NATURALIST Homing of Mallards Released in 1971 and 1972 High pre-fledging mortality of an unknown number of ducklings in 1971 severely limited the number of hens alive to return in 1972 (Bailey, unpublished data). In all, 28 females were individually identified on the Diversion in 1972 (see below for number that actually settled on the area). All homing females were accompanied by unmarked drakes and no marked males were observed. Extensive searches of the surrounding marsh did not reveal additional marked birds. Breeding pair counts indicated that about eight 1971-released birds returned to the Diver- sion in 1973, (Table 1). It was not possible to recognize individually all 2-yr-old females in 1973 because the paint had worn off many saddles. Assuming a 52% annual survival rate for adult females in southwestern Manitoba (Anderson 1975) and considering that 28 hens returned to the Diversion in 1972, then the eight birds observed in 1973 represented a homing rate of 53% for 2-yr-old females. This adult homing rate is considered minimal since the Delta Marsh is a major Mallard harvest area within south- western Manitoba. In 1973, 17 hens released in 1972 were identified on the Diversion (see below for number that settled). Marked hens from 1972 releases there appeared to have homed solely to the release site, because they were not found elsewhere in the marsh. Females homing to Aikens Bay encountered high densities of marked and unmarked Mal- lards (Table 1). At Aikens Bay, 14 returning marked hens and | unmated marked male were identified. An additional 17 marked hens were observed scattered widely throughout the marsh E of Delta and W of Clandeboye Bay (Figure 1). Numerous marked females from Aikens Bay releases only established themselves on the lakeshore opposite Aikens Bay. Another marked male was observed with an unmarked hen at Marshy Point, 30.4km E of Delta (Robert Blohm, personal communication). I estimated that 186 of the females released at the Diversion and 366 females in Aikens Bay fledged in 1972. Robert Jones (personal com- munication) calculated that the first-year mor- tality rate of hand-reared Mallards released at Delta between 1954 and 1970 was 67.2%. To Vol. 93 calculate homing rate I assumed this mortality rate for first-year fledged females in this study. The 17 returning hens at the Diversion in 1973 represented 28% of the possible yearling sur- vivors. There was a 26% homing rate (n = 31) of marked yearling hens returning to Aikens Bay and the east marsh in 1973. Comparison of Breeding Pair Counts and Homing Individuals There was a discrepancy between the number of identified yearlings homing to the Diversion and the corresponding breeding pair estimates (28 identified homing in 1972 vs. pair estimate of 18; and in 1973, 17 identified homing vs. pair estimate of 9.5). This suggested that many homing marked hens were missed during the census or a certain proportion of marked yearlings visited the study area only briefly each year. In 1972, 13 (46%) of 28 homing hens were identified only once during 51 roadside transects on the Diversion whereas the remainder (15) were recognized an average of 4.7 times each. The mean residency bout for females sighted more than once was 32 d in 1972, as determined by the number of days between the first and last sighting. In 1973, 7 (41%) of 17 hens were seen only once during 53 transects and the remainder (10) recognized an average of 2.5 times. The mean residency bout was 31.3 d in 1973. Nest Initiation and Productivity Two of the earliest nests of unmarked hens were back-dated to 16 and 20 April 1973, whereas the earliest marked females (two) began nesting on I5 May. A second peak of nest initiation by unmarked hens occurred from 27 May to 2 June (Figure 2), and was apparently due to renesting and initial nesting by Mallards moving into newly-formed water areas after 3.2cm of precipitation received during this period. A larger proportion of unmarked hens than of marked yearlings had started nesting prior to 15 May (x° = 4.8, P< 0.05, n = 36). Generally poor reproductive success was recorded for released birds nesting as yearlings. Nest predators, chiefly Striped Skunks (Mephi- tis mephitis), Raccoons (Procyon lotor), and Red Foxes (Vulpes vulpes), were common on both study areas and accounted for most nest failures (Table 2). No marked Mallard hens with broods were discovered on the Diversion in 1979 PERCENT OF TOTAL NESTS INITIATED = I) Os £ i) o Oo (e) Oo eo) eo) Oo Oo IS 2l | APRIL | MAY M= MARKED YEARLINGS n:=!! Ml UNMARKED MALLARDS n= 24 MR 28% 9.7 12 BAILEY: WILD MALLARD STOCKING 59 Pi biba 19°26. 25° 79 | JUNE I6 23 FIGURE 2. Distribution of nest initiation dates for marked vearlings and unmarked Mallards on the Delta Marsh study areas, 1973. 1972. In 1973, one brood of five ducklings was found with a 1972-released hen; however, no broods led by 1971-released hens were observed. Two marked hens with broods were found at Aikens Bay and two more marked hens with broods were located in the vicinity of the release site. One marked hen with a brood was observed on Cooks Creek, 1.6 km S of Aikens Bay. Mallard breeding pair counts for the Diversion and Aikens Bay indicated that the ratio of marked yearling hens to unmarked females (41:40) was approximately I to I each year. Significantly more broods accompanied by unmarked hens (P = 0.005, binomial test, Siegel ° 1956), however, were discovered on the release sites. TABLE 2—Nest success records for Mallards and other dabblers, Delta Marsh, 1973 Year Hatched Not of Species release Nests No. OK Destroved revisited! Deserted Mallard marked 1972 1] 2 20 8 | 0 marked 1971 3 0 0 3 - 0 unmarked 28 1] 42 IS) 2 0 Other dabblers 81 23 36 32 19 7 'Not included in the calculation of the percentage of nests hatched. 60 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 3—Percentage of marked and unmarked Mallard pairs producing broods at the Diversion and Aikens Bay in 1972 and 1973 1972 1973 Study area Year of Number of Percentage Number of Percentage release breeding pairs producing broods breeding pairs producing broods Diversion 1971 18.0 0.0 V3 0.0 1972 _— —_ QS 10.5 unmarked IBES 22. 15) 34.3 Aikens Bay 1972 _ — 13.5 14.8 unmarked — a 15.0 50.0 Fourteen nests of marked Mallards were Discussion discovered during nest searches conducted in 1973 (Table 2). Eleven of these were nests of yearling hens, and three were initiated by 2-yr- old hens at the Diversion. Two of the 11 yearling nests hatched, 8 were destroyed by predators and 1 was not relocated. All three nests of 2-yr-old hens were destroyed. Nesting success was 20% (n= 10) for nests initiated by marked yearling Mallards, whereas unmarked Mallard hens hatched 42% (n = 26) of nests initiated (no significant difference (P = 0.150) using a Fisher exact probability test (Siegel 1956)). The low probability obtained, however, suggests that the number of yearling nests found may not constitute a large enough sample to show a real difference, especially when a substantial proportion (see below) of the unmarked population was also yearling hens. When the combined nesting success of I- and 2- yr-old marked hens was compared with nesting success of unmarked hens, the Fisher exact probability decreased to 0.070. Nesting success of other dabblers encountered during nest searches was 36% (n = 81). Fifteen paired Mallard females were shot in spring 1973. These females included nine adults and six yearlings, based on wing-feather exami- nation. Evidently there was a large proportion of adult females in the unmarked Mallard popula- tion. The percentage of unmarked hens producing broods was much higher than that of marked hens on each study area in both years (Table 3). During the 2-yr study, 76 Mallard hens were identified as first-year breeders; of these, only 6 hens (8%) were observed with broods on orinthe vicinity of the release sites. Most of the marked yearlings arrived well after the majority of unmarked hens. Spring arrival of hand-reared yearlings may extend over several weeks. Sellers (1973) concluded that yearling marked hens did not lag behind adults, based on the appearance of marked birds with the first arrivals in his area; however, he did not determine when the majority of marked year- lings arrived in relation to adults. Wild-strain hand-reared Mallard females were able to return to specific release sites within the marsh. High pre-fledging mortality of 1971 releases did not prevent some survivors from returning to the study area in spring 1972. Homing of 53% (estimated) of surviving 2-yr-old hens to the Diversion showed that nesting failure during the previous year was probably not a deterrent to homing in subsequent years. A high percentage of adult hens homed in spite of low water levels in 1973. Lower homing rates of yearlings compared to those of adults have been reported for Mallard, Gadwall (A. strepera), Pintail (A. acuta), Blue- winged Teal (A. discors) (Sowls 1955); Wood Duck (Bellrose et al. 1964); Shoveler (A. clypeata), (Poston 1974); and Tufted Duck (A. fuligula) (Mihelsons et al. 1970). The pro- portion of yearling marked Mallards homing was approximately one-half that of marked adults. Furthermore, only about 60% of homing yearling hens resided on the area for extended periods each nesting season. The remainder were sighted only once, usually prior to nest initiation or after nesting in July. These observations support the contention of Hochbaum (1955, p. 124) that some hens may home to their natal area but then move elsewhere to nest. On 29 July 1979 1973, a flightless, 1972-released Mallard was observed 0.8km S of Aikens Bay, so an unknown proportion of marked hens also used the marsh to molt. Males did not generally home to release sites. In Mallards, pairing usually takes place on the wintering grounds and males follow their mates back to their natal area (Sowls 1955). But the appearance of one homing drake on Aikens Bay indicated that some drakes return to natal areas if unmated in the spring. Numbers of Mallards seen in May and June aerial surveys of the marsh in 1973 increased 2.7 times over the previous three years’ average (Delta Waterfowl Research Station, unpub- lished data). This increase may have also been partially due to low water levels causing birds to be more visible than usual from the air. The extremely high density of Mallard pairs at Aikens Bay in 1973 was partly due to the homing of release hens since one-half of the females found there were marked. The unstocked Cooks Creek Meadow also contained a relatively high density of Mallard pairs. It is possible that higher than usual numbers of Mallards on wet areas of the marsh in 1973 masked the effects of the stocking effort. In contrast, the Diversion held comparatively few Mallards after two years of stocking. The number of pairs on the Diversion remained the same each year, and there was evidence that marked yearlings were homing to, but not remaining on, the site. Marked yearlings also were found along the lakeshore at Aikens Bay. It is likely that the study areas were filled to capacity with breeding pairs. Reproductive behavior and success of first- time breeders differs from those of adults in many birds (Lack 1966). Bellrose et al. (1964) found that yearling Wood Ducks nested later than adults. Grice and Rogers (1965) showed that yearling Wood Ducks were less successful than adults in obtaining nesting sites when breeding populations were high. Heusmann (1975) indicated that limited nestings, smaller clutch size, and lower brood survival led to poorer annual production by yearling Wood Ducks. Gates (1962) found that adult Gadwall females established home ranges and began nesting earlier than yearlings in the spring. Stotts and Davis (1960) showed that adult Black Ducks BAILEY: WILD MALLARD STOCKING 61 (Anas rubripes) were paired before yearlings and suggested that adults reach breeding “tempo” before hens nesting for the first time. Clutch size was also larger in adult Black Ducks. Mihelsons et al. (1970) found that yearling female Tufted Ducks started nesting later than the older birds and had comparatively poor success. Yearling marked hens in this study arrived later on the release sites than most unmarked females, initiated nesting later, and were less successful than unmarked hens in producing broods. Lower productivity of released hens was apparently not caused by a lack of nesting cover, as found by Sellers (1973) because unmarked hens on the same areas were much more successful. Breeding by yearling hand-reared Mallards resembles a pattern observed in first- time breeders of other duck species, and lower productivity may be partly due to the fact that they are all breeding for the first time. The assumption that a considerable proportion of the unmarked Mallard population were adults was supported by the collecting of 15 hens. Poor nesting success was also documented for a small sample of adult marked hens. Hence, the possible effects of hand-rearing on future breed- ing success remain unknown. It is important from a management standpoint to determine whether the differences observed between hand- reared and wild Mallards in this study are due to hand-rearing or age. The fact that no similar age- related breeding biology studies of wild Mal- lards are available to compare with these results points out an important research need. In view of the poor reproductive success of hand-reared - birds and the apparently high potential for natural immigration and production, Mallard stocking 1s of questionable value on the Delta Marsh. Acknowledgments I gratefully acknowledge the advice of J.R. Bider received during this study. Thanks are also due to B. D. J. Batt, P. Ward, and R. E. Jones for suggestions during the field study, and to them and A. J. Erskine, A. Dzubin, and R. D. Titman for review of the manuscript. The help of students and assistants at the Delta Waterfowl Research Station was appreciated. The com- ments and criticism of R.W. Stewart were invaluable throughout this study. R. McCulloch 62 THE CANADIAN FIELD-NATURALIST assisted with the illustrations. Support of the Delta Waterfowl Research Station and Ducks Unlimited Foundation is acknowledged. I grate- fully acknowledge scholarships from the Na- tional Research Council of Canada and the Province of Québec. Literature Cited Anderson, D. R. 1975. Population ecology of the Mallard: V. Temporal and geographic estimates of survival, recovery and harvest rates. United States Fish and Wildlife Service Resource Publication 125. 110 pp. Anderson, M. G. and R. E. Jones. 1976. Submerged aqua- tic vascular plants of the east Delta Marsh. Manitoba Department of Renewable Resources and Transportation Services. 120 pp. Bellrose, F C., K. L. Johnson, and T. U. Meyers. 1964. Relative value of natural cavities and nesting houses for Wood Ducks. Journal of Wildlife Management 28: 661-676. Benson, D. 1939. Survival studies of Mallards liberated in New York State. Transactions of the North American Wildlife Conference 4: 411-415. Bird, R. D. 1961. Ecology of the aspen parkland of western Canada in relation to land use. Canada Department of Agriculture Contribution 27. 155 pp. Carney, S. M. and A. D. Geis. 1960. Mallard age and sex determination from wings. Journal of Wildlife Manage- ment 24: 372-381. Dane, C. W. 1966. Some aspects of breeding biology of the Blue-winged Teal. Auk 83: 389-402. Doty, H. A. and A. D. Kruse. 1972. Techniques for esta- blishing local breeding populations of Wood Ducks. Journal of Wildlife Management 36: 428-435. Dzubin, A. 1969. Assessing breeding populations of ducks by ground counts. /m Saskatoon wetland seminar. Canadian Wildlife Service Report Series 6. pp. 178-230. Errington, P.L. and W.E. Albert, Jr. 1936. Banding studies of semi-domesticated Mallard ducks. Bird- Banding 7: 69-73. Foley, D. D. 1954. Studies on survival of three strains of Mallard ducklings in New York State. New York Fish and Game Journal |: 75-83. Foley, D. D., D. Benson, L. W. DeGraff, and E. R. Holm. 1961. Waterfowl stocking in New York. New York Fish and Game Journal 8: 37-48. Gates, J. M. 1962. Breeding biology of the Gadwall in northern Utah. Wilson Bulletin 74: 43-67. Grice, D. and J. P. Rogers. 1965. The Wood Duck in Massachusetts. Massachusetts Division of Fish and Game, P-R Report Project W-19-R. 96 pp. Vol. 93 Heusmann, H. W. 1975. Several aspects of the nesting biology of vearling Wood Ducks. Journal of Wildlife Management 39: 503-507. Hochbaum, H. A. 1955. Travels and traditions of water- fowl. University of Minnesota Press, Minneapolis. 301 pp. Hopper, R. M. and H. D. Funk. 1970. Reliability of the Mallard wing age-determination technique for field use. Journal of Wildlife Management 34: 333-339. Hunt, R.A., L.R. Jahn, R.C. Hopkins, and G.H. Amelong. 1958. An evaluation of artificial Mallard pro- pagation in Wisconsin. Wisconsin Conservation Depart- ment, Technical Wildlife Bulletin 16. 70 pp. Lack, D. 1966. Population studies of birds. Clarendon Press, Oxford. 341 pp. Lee, F.B. and A.D. Kruse. 1973. High survival and homing rate of hand-reared wild strain Mallards. Journal of Wildlife Management 37: 154-159. Lincoln, F. C. 1934. Restocking of marshes with hand- reared Mallards not proved practicable. United States Department of Agriculture Yearbook 1934: 310-313. Love, A. and D. Love. 1954. Vegetation of a prairie marsh. Bulletin of the Torrey Botanical Club 81: 16-34. Mihelsons, H., G Lejins, A. Mednis, and V. Klimpins. 1970. Attachment to the territory and reproduction effectiveness of the Tufted Duck on Lake Engure (Latvian SSR). International Congress of Game Biologists 30: 82-85. Poston, H. J. 1974. Home range and breeding biology of the Shoveler. Canadian Wildlife Service Report Series 25. 49 pp. Schladweiler, J.C. and J. R. Tester. 1972. Survival and behavior of hand-reared Mallards released in the wild. Journal of Wildlife Management 36: 1118-1127. Sellers, R.A. 1973. Mallard releases in understocked prairie pothole habitat. Journal of Wildlife Management 37: 10-22. Siegel, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-Hill Book Company, New York. 312 Pp. Sowls, L. K. 1955. Prairie ducks. Stackpole Company, Harrisburg, Pennsylvania, and Wildlife Management Institute, Washington, D.C. 193 pp. Stotts, V. D. and D. E. Davis. 1960. The Black Duck inthe Chesapeake Bay of Marvland: breeding behavior and biology. Chesapeake Science I: 127-154. Ward, P. and B.D. J. Batt. 1973. Propagation of captive waterfowl. North American Wildlife Foundation and Wildlife Management Institute, Washington, D.C. 64 pp. Westerskoy, K. 1950. Methods tor determining the age of game bird eggs. Journal of Wildlife Management 14: 56-67. Received 12 June 1975 Accepted 2 August 1978 Notes Recent Collections of the Black Redhorse, Moxostoma duquesnei, from Ontario EDWARD KotT,! ROBERT E. JENKINS,2 and GREGORY HUMPHREYS! ‘Department of Biology, Wilfrid Laurier University, Waterloo, Ontario N2L 3C5 2Department of Biology, Virginia Commonwealth University, Richmond, Virginia, USA 23284 Kott, Edward, Robert E. Jenkins, and Gregory Humphreys. 1979. Recent collections of the Black Redhorse, Moxostoma duquesnei from Ontario. Canadian Field-Naturalist 93(1): 63-66. The Black Redhorse, Moxostoma duquesnei, a species considered endangered in Canada, was recently collected from two localities on the Nith River, a tributary of the Grand River, Ontario. It has not been collected in Canada since 1938. This species is often confused with M. erythrurum, the Golden Redhorse. A comparison between these species is included. Key Words: Moxostoma duquesnei, redhorse suckers, Ontario, morphology, endangered species. The Black Redhorse, Moxostoma duquesnei, 1s ROM9637 Ontario, Elgin County, Catfish Creek. 26 generally found in clear streams of moderate width March 1937. H. C. White. 1 adult. that are not subject to appreciable siltation. Because it © ROM10364 Ontario, Elgin County, Catfish Creek south is susceptible to siltation, it is becoming rare in many of Aylmer. 6 March 1938. H.C. White. I adult or E Underhill 1974; C d Colli supae ge cone ee Sacske: 1969) Dea tes aa The Lake Champlain system of the St. Lawrence . ; . drainage was included in the range of this species by This species has previously been recorded in SES e Canada from only two localities in Ontario, both in Hubbs (1930; p. 23) onthe basis of “Greeley (MIS) Its the Lake Erie drainage: a tributary of the Grand River poppe ae musucalice! by Uaioe (ER BHT Cre) WO (Hubbs and Brown 1929) and from Catfish Creek miclnge mul OH UNS Sie Lanitenge re RV: Gresley (Hubbs and Brown 1929; Scott and Crossman 1973). CHG ioe (pHIOLS SWieln ers! aun ee The last recorded capture of this species in Canada SOND fia VINE Lorene dramage) Woearmutcd was in 1938 and repeated attempts to collect it from ee tate ele Crain eheavine ise (On) the Conese: Catfish Creek since then have failed (Scott and oe a ee le ae ne, Crossman 1973). For this reason the species has been peas ) als@ lundhealios (ie MeN ete So included in lists of rare and endangered Canadian Grune Wes not neal peste) Byjthespeoe fishes (Anonymous 1970: McAllister and Gruchy Three specimens of this species were recently 1977). horn La eren collected by two of us from two localities in the Nith ys eae: Beene ie rae one. River, a tributary of the Grand River, in the Lake Erie McAllister and Gruchy 1977). drainage basin, at least 35 km upstream from the » Specific locality records of previous collections of sae ey 28) soe ne ue ee pace this species based on specimens in the University of SOaTe So olantorn, coe) lana Sete ue! Melati (Cg Michigan, Museum of Zoology (UMMZ) and Royal 1), was collected on 21 October 1976 ESE woWmn ee Ontario Muweun GROMD ake as Holla: Plattsville, Oxford County, Ontario, (43 18 30 N; : 80°37'30”W) using a 9-m, 6.4-mm-mesh seine. This UMMZ85887_ Ontario, Catfish Creek, | mi W of Jaffa. part of the river consists of a stretch of swift-flowing 21 July 1927. Brown and Rupp. Total 45 spms., 1 ad — shallow water (about 15 cm deep at the time) over a (202 mm SL) others young (reported by Hubbs and gravel bottom. This section ends in a poo] about | m Brown 1929), as 56 specimens. deep with a rubble bottom. The specimen, which was UMMZ89075 Ontario, Oxford County (near Brant Coun- —_—co}lected in the shallows, is in the Wilfrid Laurier ty line) Cedar Creek, tributary of Grand River between University Museum as number WLU5594. Paris and Princeton. 5 Sept. 1928. C. L. Hubbs. 3 speci- Two specimens (81mm and 79 mm_ standard mens 75, 215, 215 mm SL (3 Aug. 1927 according to : : 5 length) were collected using the same technique on 29 Hubbs and Brown 1929) [no stream in the area indicated ; Sipe currently bears the eae Cheeta June 1977 (Figure 1) at the second locality just north ROMI975 Ontario, Elgin County, Catfish Creek near Of Ayr, Waterloo County, (43° 17'45"N; 80°28°10°N) Aylmer. 21 March 1926. H. C. White. 1 specimen, 253. and 24 km downstream from the first. This segment of SL. the river is similar to the previous in that it consists of 63 64 THE CANADIAN FIELD-NATURALIST ) ith il cm ] AIL a | sl | al ie Vol. 93 | FIGURE 1. Moxostoma duquesnei collected from the Nith River: upper specimen, NMC 78-1; middle, WLU5233: lower, WLU5594. a zone of rapids followed by a deep pool. The rapids had a boulder bottom with a depth up to | m. At low water levels, the pool was effectively divided bya gravel bar into two smaller pools. At the time of capture the depth of the pool was about 75 cm. The sides were gravel and the bottom was covered in fine silt. Both specimens were collected from the pool. One specimen is deposited in the Wilfrid Laurier Uni- versity Museum as WLU5233, and the other at the National Museum of Natural Sciences, Ottawa as NMC78-1. Moxostoma erythrurum, Golden Redhorse, was collected with M. duquesnei at the first locality and M. macrolepidotum, Shorthead Redhorse, was col- lected at the second locality. The earlier and recently collected specimens of M. duquesnei agree in morphology and coloration with the species from elsewhere in the Lake Erie basin (tributaries in the United States) and from through- out most of the wide range of the species in the United States, from the southern parts of the lakes Ontario, Huron, and Michigan basins and southward. The species in the Nith River system that most closely resembles M. duquesnei is M. erythrurum. Many workers encounter difficulty in distinguishing them. The following meristic data (taken by methods of Hubbs and Lagler 1958) are from specimens from the Lake Erie basin of Canada and the United States unless otherwise stated (Jenkins 1970 and subsequent study). The best character for separation of these two species is the lateral-line scale count: M. duquesnei ranges 45-48, x = 46.4, N = 28: M. eryvthrurum 39-44, x = 41.0, N = 46. Slightly overlapping counts may be expected from additional specimens. Other meristic differences are the following: body circumferential scales (axis just anterior to dorsal fin), M. duquesnei 33-36, M. erythrurum 29-34; predorsal scale rows, M. duquesnei 16-20, M. erythrurum 13-16; pelvic fin rays, M. duquesnei usually one or both fins with 10 rays (in 19 of 28 specimens; nine with 9 as the highest count), M. erythrurum 9-9 or 9-8 in 50 of 52 specimens, 10-9 or 10-10 in the other two; post- Weberian vertebrae (counts from throughout total range of species), M@. duquesnei 39-43 (40 in largest Nith specimen), M. erythrurum 35-39. 1979 NOTES 65 TABLE 1—Some characteristics of three specimens of Moxostoma duquesnei and 11 specimens of M. ervthrurum from the Nith River Specimen Standard. Head length/ Eye diameter/ Caudal peduncle Caudal peduncle Number of Number of and museum length, snout length snout length depth/ depth/ lateral pelvic rays number mm caudal peduncle standard length line Slctimnrichit length scales ia M. duquesnei WLU5594 49 2.84 0.81 0.6 0.094 47 10 10 WLU5233 79 2.56 0.55 0.67 0.090 46 9 9 NMC78-1 81 Dee) 0.61 0.7 0.095 46 9 9 Mean 69.7 2.65 0.66 0.66 0.093 46.3 OS OS Range 49-81 2.55-2.84 0.55-0.81 0.6-0.7 0.090-0.095 46-47 9-10 9-10 M. erythrurum WLU4763 41 3.3 0.94 0.68 0.102 40 9 9 45 3323 0.94 0.67 0.104 42 9 9 47 3.29 0.94 0.68 0.094 42 9 9 49 3.18 0.85 0.69 0.100 43 9 9 49 323 0.89 0.68 0.090 43 9 9 WLU4556 54 2.62 0.69 0.72 0.106 44 9 9 57 Des 0.65 0.66 0.093 42 10 10 59 2.90 0.76 0.71 0.102 43 9 9 62 2-38) 0.64 0.71 0.103 42 10 9 62 2.62 0.68 0.68 0.094 4] 9 9 WLUS5305 75 3.02 0.87 0.91 0.111 40) 9 9 Mean 54.6 2.92 0.80 0.71 0.100 41.8 O22 ON Range 41-75 2.53-3.3 0.64-0.94 0.66-0.91 0.090-0.111 40-44 9-10 9-10 The species also differ in certain aspects of morphometry (Hubbs and Lagler 1958; Jenkins 1970; Phillips and Underhill 1971; Scott and Crossman 1973; Pflieger 1975) but greater overlap occurs with these characters than in at least most of the former. Some of the overlap is due to allometric growth of body parts and to comparison of samples of speci- mens with dissimilar body lengths (Jenkins 1970). Table | expresses the interspecific differences, with overlapping, based on the young and small juveniles from the Nith River. A recently discovered difference, that is particularly useful for sorting large series of small specimens, concerns coloration of the upper body. The difference is best developed in the anterior dorsolateral area. In M. duquesnei the melanophores of each scale are more uniformly distributed, the total effect being an evenly dusky to darkly colored back. In M. ery- thrurum the posterior margin of each scale usually is obviously darker than the somewhat pale central area of the exposed portion of the scales, hence the scales appear somewhat bicolored. The difference is usually slight, but is discernible using syntopically collected and similar-sized specimens of the two species. The size of the specimens collected, apparently young and small juveniles, indicates that a breeding population of this species still exists in Canada. The Nith River follows a series of gravelly spillways until it unites with the Grand River (Chapman and Putnam 1966). For this reason it has a considerable gradient, with a minimum summer flow of about 2.8 m>s!. The region drained is rural in nature, with New Hamburg (population 2990 in 1974) being the largest center. For part of its course it follows valleys 15.2-30.5 m in depth: the other stretches are bounded by flood- plains. These floodplains are largely cleared and are used for growing grain or as pasture land; however, in some areas natural reforestation 1s occurring on abandoned or little-used farmland. The river system is relatively clear, except during . the spring flood period. Turbidity in Formazin Turbidity Units, for June and July, taken at a location near Ayr collection site was 2.70 and 2.40 (Anony- mous, no date). The Nith River system should be protected to enable the continued existence, in Canada, of the ecosystem that has harbored this species and other more typical elements of the biota of Canada. This species, itself, should be protected as an endangered species in Canada. Acknowledgments We thank Frank Mallory, Department of Biology, Wilfrid Laurier University and Bruce M. Buckland, Ministry of Natural Resources, Cambridge, for assisting us in collecting the specimens. Additional specimens were kindly loaned by E. J. Crossman, 66 THE CANADIAN FIELD-NATURALIST Royal Ontario Museum, Toronto, and R. M. Bailey, University of Michigan, Museum of Zoology, Ann Arbor. D. E. McAllister provided the X-ray negative used for the vertebral count. This study was sup- ported, in part, by National Research Council of Canada Grant No. A-0575 to E. Kott. Literature Cited Anonymous. Undated. Water quality data for Ontario lakes and streams. 1974. Volume IX. Water Resources Branch, Ontario Ministry of the Environment. xxxiii + 324 pp. Anonymous. 1970. Table of extinct, rare and endangered vertebrates in Ontario. Ontario Fish and Wildlife Review 9(1-2): 15-21. Chapman, L. J. and D. F. Putman. 1966. The physiog- raphy of southern Ontario. Second edition. University of Toronto Press, Toronto. xiv + 386 pp. Cross, F.B. and J.T. Collins. 1975. Fishes in Kansas. University of Kansas Museum of Natural History, Public Education Series Number 3. 189 pp. Eddy, S. and J.C. Underhill. 1974. Northern fishes. University of Minnesota Press, Minneapolis. 414 pp. Harlan, J. R. and E.B. Speaker. 1969. lowa fish and fishing. State Conservation Commission, Des Moines, lowa. 365 pp. Hubbs, C. L. 1930. Materials for a revision of the cato- stomid fishes of eastern North America. University of Vol. 93 Michigan Museum of Zoology Miscellaneous Publica- tion 20. 47 pp. Hubbs, C. L. and D. E.S. Brown. 1929. Materials for a distributional study of Ontario fishes. Transactions of the Royal Canadian Institute 17. 56 pp. Hubbs, C.L. and K.F. Lagler. 1958. Fishes of the Great Lakes region. University of Michigan Press, Ann Arbor, Michigan. 213 pp. Jenkins, R. E. 1970. Systematic studies of the catostomid fish tribe Moxostomatini. Ph.D. thesis, Cornell University, Ithaca, New York. 818 pp. McAllister, D.E. and C.G. Gruchy. 1977. Status and habitat of Canadian fishes in 1976. Jn Canada’s threatened species and habitats. Canadian Nature Federation, Special Publication 6. pp. 151-157. Pflieger, W.L. 1975. The fishes of Missouri. Missouri Department of Conservation. 343 pp. Phillips, G. L. and J. C. Underhill. 1971. Distribution and variation of the Catostomidae in Minnesota. Bell Museum of Natural History, University of Minnesota, Occasional Papers, Number 10: 1-45. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada Bulletin 184. 966 pp. Trautman, M. B. 1957. The fishes of Ohio with illustrated kevs. Ohio State University Press, Columbus, Ohio. 683 PP- Received 8 June 1978. Accepted 8 August 1978. Xanthochroism! in the Evening Grosbeak CHRISTOPHER W. HELLEINER Nova Scotia Bird Society, c/o Nova Scotia Museum, 1747 Summer Street, Halifax, Nova Scotia B3H 3A6 Helleiner, Christopher W. 1979. Xanthochroism in the Evening Grosbeak. Canadian Field-Naturalist 93(1): 66-67. Key Words: xanthochroism, Evening Grosbeak (Hesperiphona vespertina). This note describes two unusually pigmented Evening Grosbeaks, Hesperiphona vespertina, and summarizes several other reports between 1970 and 1977 in Nova Scotia. Xanthochroism in this species has previously been reported by Saunders (1958) and Godfrey (1967). L.B. Macpherson (Maitland, Hants County, Nova ‘According to Harrison (Bird-Banding 1966, 37: 121) the use of the term “xanthochroic” should be limited to replace- ment of normal color by yellow. Most examples reported are more probably non-melanic schizochroic, where the absence of melanin leaves carotenoid pigments pheno- typically apparent. I have therefore inserted the word “apparent” to modify xanthochroism in the text. Scotia) described an almost completely lemon-yellow and white male Evening Grosbeak in winter 1969-70. The primaries and tail were entirely white, except for the tip of one tail-feather, which was dark brown or black. The head was faintly tinged with olive, but the rest of the plumage was yellow. The legs and feet as well as the lower mandible were flesh-colored. The upper mandible was gray. The eyes were dark. The same bird, or one very like it, reappeared the following winter. I saw a xanthochroic male Evening Grosbeak with somewhat more dark pigmentation in Halifax for a few days on and around 19 March 1977. The dark on the head, breast, and back were largely absent, except for a little coarse brown mottling on a yellow ISS) background. The dark color in the wings and tail appeared normal, but the white area seemed some- what more extensive than usual, overlapping the dark primaries a little. Both the Maitland and the Halifax bird behaved normally in the company of others of their species, and competed aggressively for food. Less detailed reports of several other apparently xanthochroic Evening Grosbeaks have appeared in winter bird reports in the Nova Scotia Bird Society Newsletter in 1972, 1973, and 1977. These represent a total of about seven such individuals since the first specimen was collected in this province in 1958. The Evening Grosbeak has become much commoner in Nova Scotia during the last two decades. Average numbers recorded on Christmas counts have in- creased from 330 to 1160 to 1610 per count in successive 5-yr periods starting in 1963 (Nova Scotia Bird Society Newsletter). One might therefore expect aberrant individuals to be seen more often. Neverthe- less the frequency with which this otherwise very rare NOTES 67 color variation is being reported in this species is surprising. Smith (1966) states that xanthochroism is rarer than other color variations in wild birds; Gross (1965) lists many examples of melanism, several of erythrism, but only six of xanthochroism. Perhaps in the Evening Grosbeak affected individuals do not suffer significant competitive disadvantage, and live to reproduce. Literature Cited Godfrey, W.E. 1967 Xanthochroism in the Cape May Warbler and Evening Grosbeak. Canadian Field-Natural- ist 81: 226-227. Gross, A. O. 1965. Melanism in North American birds. Bird-Banding 36: 240-242. Saunders, A. A. 1958. A yellow mutant of the Evening Grosbeak. Auk 75: 101. Smith, R. W. 1966. A case of xanthochroism in the Yellow- faced Grassquit, Tiaris olivacea. Ibis 108: 627-628. Received 19 May 1978 Accepted 18 September 1978 Interaction between a Long-tailed Weasel and a Snowy Owl PETER C. BOXALL Department of Biology, University of Calgary, Calgary, Alberta T2N IN4 Boxall, Peter C. 1979. Interaction between a Long-tailed Weasel and a Snowy Owl. Canadian Field-Naturalist 93(1): 67-68. A Long-tailed Weasel (Mustela frenata) was observed rushing towards a Snowy Owl (Nyctea scandiaca) and its recently- captured prey. This interaction is interpreted as attempted food piracy. The fearlessness of weasels may account for their presence in the diet of owls. Key Words: Mustela frenata, Nyctea scandiaca, food piracy, predator. Little is known about the relationships of weasels (Mustela spp.) to other predators. But the aggressive predatory behavior and curiosity of weasels have been described (Armitage 1961; Hansen 1952). While conducting research on the behavioral ecology of wintering Snowy Owls (Nyctea scandiaca), | wit- nessed an interaction between a weasel and a Snowy Owl. At 11:24 on 15 December 1976, I was observing a large female Snowy Owl perched on a fence-post at the edge of a stubblefield, 6 km E of Calgary, Alberta. I noted a Long-tailed Weasel (M. frenata) moving slowly through the field approximately 30 m from the owl. At 11:25 the owl flew into the stubblefield, captured a small mammal, and remained on the ground with the prey item in its feet. During this time, the weasel was sitting upright about 20 m from the capture location, and had apparently witnessed the hunting attempt. After the capture, the weasel slowly moved to within 10m of the owl. As the weasel approached, the owl picked up the prey in its bill, shook it several times, and started to swallow it. The weasel then ran quickly to within 3 m of the owl, whereupon the owl flew back to its original perch. The weasel continued forward and investigated the area where the hunting attempt occurred. Subsequently, the owl flew 500-600 m away and perched in a large tree. Both owls and weasels are dependent upon rodents for food during the winter. Weasels, however, are 68 THE CANADIAN FIELD-NATURALIST sometimes utilized as food by wintering Snowy Owls. Gross (1944) reports the remains of three weasels (species not given) in one Snowy Owl stomach. Catling (1973) found the stomach of one Snowy Owl in Ontario to contain an entire Ermine (M. erminea). In addition, | have found the remains of one Least Weasel (M. nivalis) and three Long-tailed Weasels in a sample of 100 pellets from the Calgary region. Although the vision of weasels appears to be quite poor (Murie 1935), their ability to locate moving objects, and to detect prey by scent, is good (Ewer 1973). The weasel, during the interaction described above, behaved as if the owl’s prey was detected by scent. I believe that the weasel’s actions towards the owl were probably motivated by hunger. The sudden rush by the weasel at the owl as the latter started swallowing the prey suggests a possible attempt at food piracy. The unusual behavior of the weasel, and the fact that the owl had just ingested a prey item, probably allowed the weasel to escape unscathed. Fearless behavior toward a larger pred- ator, as described above, could explain in part the observed levels of predation upon weasels in nature. Vol. 93 I thank M. Ross Lein, Dick Braithwaite, and Tom Dickinson for criticism of the manuscript. This observation was made while I was conducting research supported in part by grants from the National Research Council of Canada to M. Ross Bem: Literature Cited Armitage, K. B. 1961. Curiosity behavior in some muste- lids. Journal of Mammalogy 42: 276-277. Catling, P.M. 1973. Food of the Snowy Owl wintering in southern Ontario, with particular reference to the Snowy Owl hazard to aircraft. Ontario Field Biologist 27: 41-45. Ewer, R.F. 1973. The carnivores. Cornell University Press, Ithaca, New York. Gross, A.O. 1944. Food of the Snowy Owl. Auk 61: 1-18. Hansen, R.M. 1952. Notes on behavior of long-tailed weasel in Utah. Journal of Mammalogy 33: 492. Murie, A. 1935. A weasel goes hungry. Journal of Mam- malogy 16: 321-322. Received 23 May 1978 Accepted 3 August 1978 Black-necked Stilts Nesting near Edmonton, Alberta Dick DEKKER,! ROBERT LISTER,2 TERRY W. THORMIN,*.D. V. WESELOH,? and LINDA M. WESELOH4 13819-112A St., Edmonton, Alberta T6J 1K4 210823-80 Avenue, Edmonton, Alberta T6E 1V9 3John Janson Nature Centre, Edmonton Parks and Recreation, Edmonton, Alberta. Present address: Provincial Museum of Alberta, Edmonton, Alberta TSN 0M6 4Provincial Museum of Alberta, Edmonton, Alberta T5N 0M6. Present address: Canadian Wildlife Service, Canada Centre for Inland Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6 Dekker, Dick, Robert Lister, Terry W. Thormin, D. V. Weseloh, and Linda M. Weseloh. 1979. Black-necked Stilts nesting near Edmonton, Alberta. Canadian Field-Naturalist 93(1): 68-69. A series of sightings of Black-necked Stilts, Himantopus mexicanus, near Edmonton, Alberta, in the spring of 1977 culminated in the finding of two nests with eggs, which provide the first authenticated records of this species breeding in Canada. Key Words: Himantopus mexicanus, breeding record, Black-necked Stilts, Alberta. On 1 May 1977, when checking birds with Kathleen Ball and Bruce and Karen Heming at the southeast end of Beaverhill Lake some 80 km (50 mi) SE of Edmonton, RL was told by Ron Slagter of Edmonton that he had found two Black-necked Stilts on a shallow bay of the lake. They proceeded to the spot and saw the birds feeding with about 20 American Avocets, Recurvirostra americana. The stilts were viewed for about 20 min with binoculars and telescope at a distance of 50 m, but the birds were readily identified with the naked eve. Later it was learned that the birds had also been seen by Gerry and Betty Bulmer of Edmonton, who took recognizable photographs of them (Bulmer 1977). On 3 May, RL was informed that R. C. Hodgins, of Stony Plain, Alberta, had seen another pair of stilts ona pond near St. Albert, about 10 mi(16 km) NW of Edmonton, but a later search of the area failed to relocate the birds. Hodgins is familiar with the stilt and was positive of his identification. 1979 On 24 May, Dick Dekker reported two Black- necked Stilts on the north shore of Beaverhill Lake, and on 4 June, he found four of the birds in a marshy area on the west side of the lake. On the following day, Dekker and Lister went to view the birds. They were joined at the lake by the Weselohs and Thormin. The area had been dry in April, but heavy rains and some snow later had flooded it, so in June it was mostly mud and water, in parts heavily overgrown with Marsh Ragwort (Senecio congestus var. palustris). At the marsh were three Black-necked Stilts and several avocets. One of the stilts seemed to run from a clump of ragwort, and resorted to a “broken-wing” dis- traction display. We withdrew for a time to give the bird a chance to return to a nest if it had one. After about 20 min we moved in, and again the bird was seen to run from a mound of mud. DVW waded through the mire and was rewarded by finding a nest and four eggs. He collected one egg which is now in the Provincial Museum of Alberta, and pictures were taken of the nest. The nest was made of weed stems, apparently the stalks of ragwort, and was located just above the water level. Avocets’ nests in the vicinity were all on dry land some distance from the water, which may have receded since the eggs were laid. All the avocet nests we saw had less nest material in them than that of the stilt. On 17 June DVW and Eric Tull discovered a second stilt nest approximately 50 m west of the first one. The second nest contained seven eggs, one of which was also collected and preserved at the museum. This nest, except for the greater number of eggs, was similar in construction, content, and placement to the first. Three days later the first nest contained only one egg, which was pipped. By 07:45 on 21 June the chick had successfully emerged and was observed in the nest (DVW and Eric Mills). The fate of the young from the other two eggs is unknown, but they probably hatched on 19 or 20 June and were led to the nearby marsh. By 23 June the first nest was completely vacated and the second nest still contained six eggs. Hatching in the second nest was first noted at 06:10 on 4 July when the nest contained one young and five eggs. By 20:15 on 4 July, two young had emerged and a third could be heard peeping inside its cracked egg; the two young were banded by Martin McNicholl. On 7 July the second nest still contained three eggs which were cold and addled, and the remains of a unbanded young stilt were found only a few feet from the nest. Presumably this was the young that was heard peeping on 4 July. The three addled eggs were collected and are preserved at the museum. The two nests of Black-necked Stilts near Edmon- ton, in 1977 contained I 1 eggs, of which four definitely hatched, two probably hatched, two were collected NOTES 69 early in incubation, and three were addled (and subsequently collected). Three young are known to have left the nest, two others may have, and one was found dead. In the spring of 1977 there were far more avocets around Beaverhill Lake than RL could remember in about 50 years of visiting the lake (see also Dekker 1977). Dry weather in the western states as well as in sourthern Alberta, accompanied by very little run-off from the meagre snowfall of winter, had left many marshes dry, and it is possible birds had travelled far north of their normal breeding grounds to find suitable nesting habitat. There were also several sightings of Black-necked Stilts in Montana during the first week of May, and that state too recorded its first breeding records (two) of the stilt in 1977 (Paul Skaar, personal communication to DVW). Salt and Salt (1977) list three previous sightings of the Black-necked Stilt in Alberta. Parts of a Black- necked Stilt are said to have been found (by Tom Randall) near Brooks in the mid-1950s but they were not preserved. On 12 May 1970, a bird was seen and photographed near Calgary (Weseloh 1972) and on 24 May 1972, another was seen and photographed near Irricana (Weseloh and McKay 1972). Godfrey (1966) remarks “A set of eggs in the National Museum of Canada was, according to the label, taken at Qu’Appelle, Saskatchewan, on June 13, 1894, by Edward Arnold. This is so far from the known breeding range that the possibility of an error in the data seems probable.” In light of the current Alberta and Montana nesting records, perhaps the Saskatchewan record should be given more credence. Literature Cited Bulmer, B. 1977. Black-necked Stilts observed at Beaverhill Lake. Edmonton Naturalist 5(6): 145. Dekker, D. 1977. Avocets and habitat. Edmonton Naturalist 5(6): 144. Godfrey, W.E. 1966. The birds of Canada. National . Museum of Canada Bulletin 203. 428 pp. Salt, W.R. and J. R. Salt. 1977. The birds of Alberta. Hurtig Publishers, Edmonton. Weseloh, D. V. 1972. First verified record of the Black- necked Stilt for Alberta. Canadian Field-Naturalist 86:165. Weseloh, D. V. and W.G. McKay. 1972. The second report of a Black-necked Stilt in Alberta. Calgary Field- Naturalist 4(5):; 90-91. Received 10 April 1978 Accepted 6 August 1978 Addendum No Black-necked Stilts were seen at Beaverhill Lake in 1978. Received 8 November 1978 70 THE CANADIAN FIELD-NATURALIST Vol. 93 Common Garter Snake Predation on Robin Nestlings KATHY MARTIN Department of Zoology, University of Alberta, Edmonton, Alberta Present Address: Biology Department, University of Prince Edward Island, Charlottetown, Prince Edward Island CIA 4P3 Martin, Kathy. 1979. Common Garter Snake predation on Robin nestlings. Canadian Field-Naturalist 93(1): 70-71. Key Words: American Robin, Turdus migratorius, nests, nestlings, Common Garter Snake, Thamnophis sirtalis, predation, prey selection. During a two-year study of American Robins (Turdus migratorius) on Comox burn, a Douglas Fir (Pseudotsuga menziesii) plantation on Vancouver Island, British Columbia, a large percentage of nests were unsuccessful. Of 175 active nests found during 1971 and 1972, at least 59 (34%) were destroyed owing to predation (Martin 1973). The only predation I actually witnessed involved a Common Garter Snake (Thamnophis sirtalis) removing young Robins froma nest on the ground. On 11 July 1971 at 16:15 I found a Robin nest well camouflaged under a fallen log. Three nestlings approximately 9 d old begged vigorously when approached, and I banded, weighed, and measured them. Their weights were 32, 35, and 54 g. On 12 July at 09:40 I found that a garter snake, approximately 90 cm long, had removed the smallest nestling and was ingesting it. Another young was out of the nest and the third remained in it. As I approached, the snake began to move away. It had ingested the nestling’s head and neck, but the abdomen and legs were still dangling from its mouth. I wished to detain the snake for Martin McNicholl (MKM) to photo- graph, so I tied a string to the nestling’s leg, attached it to a Red Huckleberry (Vaccinium parvifolium) stem and left the site. At 10:05 the snake had regurgitated the Robin and disappeared. I returned the live young to the nest, collected the dead one, and entered a small observation blind. At 11:30, I heard a nestling squawking and saw a snake pull the second lightest young (35 g) from the nest. The snake was biting or grasping the neck of the nestling as MKM and I approached. When the snake saw us, it tried to move away with the young Robin, but the nestling prevented this by grabbing vegetation with its feet. MKM took several photographs and we left the nest site. As I watched from a blind, the snake pulled its prey a short distance away and out of sight. At 12:00 the female Robin returned with food in her beak and remained near the nest chirping softly until 12:20 when she fed the remaining young (54 g) and flew away. The snake crawled out of a hole near the nest at 12:30 at which time it had ingested only the head and neck of the second nestling. At 12:50, the male Robin returned with food, but by then the snake had disappeared. At 13:45 I revisited the site and found the second nestling dead 6 m north of the nest. As there was no interference by humans or adult Robins after 11:30 it may be that the snake was unable to swallow this nestling. At 15:35, the third and largest young was still in the nest, and on 13 July at 11:10 found it fledged about | m north of the empty nest. At 12:00 on 13 July, the second dead young Robin was still near the nest and I saw a snake nearby of the same species and size as seen earlier. On 15 July at 15:45, the dead nestling was gone. While I witnessed the predation, the adult Robins did not observe the snake, because they spent relatively long periods away from the nest and the snake was not present when they returned. The literature refers to snakes as occasional predators of Robin eggs and young (Bent 1949). On my study area, predation on Robin nests by garter snakes was probably infrequent because most Robin nests were situated approxmately | m above the ground in trees (Martin 1973). The major prey species of T. sirtalis on Vancouver Island are amphibians and earth worms (P. Gregory, personal communication): however, T. sirtalis does occasionally eat nestling birds, sometimes climbing small bushes and trees (Gregory 1975). There have been a number of reports of snakes raiding passerine nests (Cink 1977: Best 1974, 1978: Ervin and Rose 1973). Both Cink (1977) and Best (1974) reported that snakes ate the complete clutch of eggs or young. Best (1974) reported Blue Racers (Coluber constrictor) returning to the nest after they had removed all the contents. I have found no reports that snakes have shown selectively in killing nestlings. My observations indicated that a snake first killed the lightest and smallest nestling. It then returned and killed the second-lightest nestling, possibly indicating prey selection by size. Because the snake abandoned the second nestling, perhaps these young Robins were larger than normal prey and too large to ingest. I thank P.T. Gregory, M. Harris, M. K. Mc- Nicholl, and F.C. Zwickel for comments on the manuscript. 1979 Literature Cited Bent, A. C. 1949. Life histories of North American thrush- es, kinglets, and their allies. United States National Museum Bulletin Number 196. 452 pp. (1964 Dover reprint consulted.) Best, L. B. 1974. Blue Racers prey on Field Sparrow nests. Auk 91: 168-169. Best, L. B. 1978. Field Sparrow reproductive success and nesting ecology. Auk 95: 9-22. Cink, C. L. 1977. Snake predation on Bell’s Vireo nest- lings. Wilson Bulletin 89: 349-350. Ervin, S. and C. Rose. 1973. Gopher Snake predation on NOTES ial the Common Bushtit. Auk 90: 682-683. Gregory, P. T. 1975. Arboreal mating behavior in the Red- sided Garter Snake. Canadian Field-Naturalist 89: 461- 462. Martin, K. 1973. Breeding density and reproductive success of robins in relation to habitat structure on logged areas of Vancouver Island, British Columbia. M.Sc. thesis, University of Alberta, Edmonton. 89 pp. Received 19 May 1978 Accepted II] August 1978 Distribution and Movements of Selkirk Caribou, 1972-1974 DAVID J. FREDDY Wilderness Research Center, University of Idaho, Moscow, Idaho 83843 Present Address: Colorado Division of Wildlife, Box 481, Kremmling, Colorado 80459 Freddy, David J. 1979. Distribution and movements of Selkirk Caribou, 1972-1974. Canadian Field-Naturalist 93(1): 71-74. Distribution and movements of Caribou inhabiting the Selkirk Mountains of northeastern Washington, northern Idaho, and southern British Columbia were determined through aerial and ground surveys from 1972 to 1974. During most of the year, Caribou were located in British Columbia at elevations above 1430 m, in close association with the spruce-fir forest. Significant seasonal shifts in elevations frequented by these animals were not detected. Caribou consistently utilized specific travel routes between and within drainages. Their only travel route to the USA was through Kootenay Pass. Continued viability of the population probably depends on safeguarding their movement routes and spruce-fir forest winter ranges. Key Words: Woodland Caribou, Selkirk Mountains, distribution, movements, ranges. Woodland Caribou, Rangifer tarandus caribou (Banfield 1961, p. 70), inhabiting the Selkirk Moun- tains of northern Idaho, northeastern Washington, and southern British Columbia (BC), Canada, may constitute the only population of Caribou frequenting the contiguous United States (USA). Caribou popula- tions formerly existing in Maine and Minnesota were extirpated during the early part of this century (Palmer 1938; Nelson 1947). Populations in north- western Montana and other portions of northern Idaho have apparently been severely reduced in number or extirpated (Evans 1960). Historical reviews pertaining to Selkirk Caribou were assembled by Evans (1960) and Layser (1974). These accounts, based primarily on interviews, sum- marize observations and records of these Caribou from 1800 to 1971. Upon interpretation, these reviews elucidate probable distribution and general move- ments of this herd (Freddy 1974). Estimates of the historic size of this population have been based on inadequate data. Flinn (1959) felt there were at least 100 Caribou in the late 1950s while Layser (1974) estimated that 15-20 animals existed in the early 1970s. Recent work indicates that 25-30 animals form the population (D. Freddy. 1974. Report to inter- national Caribou steering committee. Unpublished report, B.C. Forest Service, Nelson. 9 pp.). Whether this population has recently declined or is currently increasing, decreasing, or stable in size is unknown. In 1971 the University of Idaho Wilderness Re- search Center and several organizations initiated a cooperative study of Selkirk Caribou to document the distribution and movements of this population. The investigation was prompted by ongoing and potential human activities that appeared to be threatening the continued existence of these Caribou. Study Area The study was conducted in the Selkirk Mountains between Priest Lake, Idaho and Kootenay Lake, BC. (Figure 1). Elevations range from 700 m to 2100 m. Maritime air from the Pacific Ocean continually affects the area resulting in frequent cloudiness and precipitation. Lower elevations receive about 75 cm annually and higher elevations, over 250 cm of precipitation, with 70% of this moisture occurring as V2 THE CANADIAN FIELD-NATURALIST snow between October and March (USDA 1972). Snow begins to accumulate at higher elevations during October and by December all elevations are snow covered. In late February or March the snowpack begins to settle and harden as warmer temperatures become more frequent. During April, snow at lower elevations begins to melt and at higher elevations becomes thoroughly settled with depths of 3m common at elevations above 1500 m. By late June, snow at lower elevations has melted but at higher elevations snow will persist in sheltered areas until early July. Western Red-Cedar (Thuja plicata) — Western Hemlock (T7suga heterophylla) and Engelmann Spruce (Picea engelmanii) — Subalpine Fir (Abies lasiocarpa) forest types dominate the area. The spruce-fir forest predominates at elevations above 1500 m, spruce-fir and cedar-hemlock codominate at 1400-1500 m, while the cedar-hemlock forest domin- ates at elevations below 1400 m. A more complete description of these vegetative associations is pro- vided by Daubenmire and Daubenmire (1968). Primary land use has been and continues to be timber production. British Columbia Highway 3 bisects the area and logging roads penetrate nearly every drainage. Methods Distribution and movements of Caribou were determined from aerial and ground observations of Caribou or their tracks. Seventeen aerial surveys were conducted from March 1972 through April 1973 using a Cessna 182 while four surveys were conducted from March to July 1974 with a Bell 206 helicopter. Flights were conducted on days when visibility and contrast were good and a snowpack was present. To avoid duplicate observations of tracks, flights during winter were made after recent snowfalls and during spring after the snowpack had melted sufficiently to remove tracks seen from previous flights. Fixed-winged aircraft were flown at 135-140 km/h at altitudes 150-300 m above the ground, depending on terrain. Helicopter surveys were generally flown at lower altitudes and slower speeds. There was one observer per flight. Aerial surveys followed fixed flight paths to standardize data collection (Freddy 1974). These Surveys concentrated on elevations above 1500 m because Caribou and/or tracks could usually be seen only on the snowpack within the relatively open- crowned spruce-fir forest above this elevation. Mis- identification of tracks during aerial surveys was considered to be minimal. Moose (A/ces alces) and Mountain Goat (Oreamnos americanus) were the only other large ungulates present and the restricted distributions of these two species minimized problems in identifying tracks. Ground surveys were conducted Vol. 93 from February 1972 through August 1974. These surveys were used to supplement aerial surveys especially in areas below 1500 m elevation. Areas selected for search were judged probably to be utilized by Caribou. Searches were made afoot and by snowmobile. Elevations where Caribou or tracks were located were determined to the nearest 30 m from topo- graphic maps and/or the altimeter of survey aircraft. A one-way analysis of variance was used to test for differences in monthly elevations of Caribou loca- tions. Specific travel routes were determined by tracking Caribou on foot or from aircraft. Results and Discussion From February 1972 through August 1974, 238 observations of Caribou or tracks were recorded. Observations occurred during all months except December and January of each year. Caribou frequented areas within BC, Washington, and Idaho. Areas of concentrated Caribou activity occurred in BC with drainages immediately north of Kootenay Pass receiving heavy utilization year round. Areas in the USA most commonly frequented by Caribou were adjacent to Snowy Top and Conti- nental mountains (Figure |). Caribou were found in BC and the USA during both winter and summer but most of the population remained in BC year round. This observed distribution of Caribou (Figure 1) resembled the historic distribution of the population (Freddy 1974). Current limits of distribution were similar to historic bounds and areas adjacent to Snowy Top and Continental mountains and Koot- enay Pass continued as focal points of Caribou activity. Areas in the USA within the southern portion of the population’s range, however, apparently were not used as intensely as in the past. This could have been a function of the years of the study, rather than an avoidance of areas in the USA. Skoog (1968, p. 437) and Kelsall (1968, p. 108) describe distributional shifts of Caribou which can result in temporary absence from areas. Caribou were usually located year round at eleva- tions above 1430 m with the mean elevation of Caribou locations being 1657 + 227 (SD) m. There were no significant differences in monthly elevation means (P < 0.05) which ranged from 1591 + 204 (sD) m in October to 1734 + 235 (Sp) m in September (Table 1). The absence of distinct shifts in seasonal elevations frequented by Selkirk Caribou differs from movements reported for Caribou in Wells Gray Provincial Park (WGPP) in central BC (Edwards and Ritcey 1959). Edwards and Ritcey (1959) concluded that the elevational distribution of WGPP Caribou shifted markedly in response to accumulating or receding snowpacks. During fall, Caribou in WGPP 1979 he @ MTN. COMMON 4 1 t] DISTRIBUTION \ i] \ PRIEST OCCASIONAL LAKE INFREQUENT MOVEMENT ROUTE HIGHWAY FIGURE 1. Distribution, frequency of occurrence, and known movement routes of Selkirk Caribou, 1972-1974. moved down into the cedar-hemlock forest to avoid deepening snow at higher elevations and to forage on understory vegetation. As snow accumulated, Cari- bou ascended to the spruce-fir forest to forage on arboreal lichens, and then again descended to lower elevations in the spring. Although movements of Selkirk Caribou were apparently more variable, both populations frequented lower elevations in fall and associated with the spruce-fir forest in mid-winter. Food habits of Selkirk and WGPP Caribou (Freddy TABLE |—Elevations at which Caribou or their tracks were observed, 1972-1974 Number of Elevation (m), Month * observations Mean +SD Range February 22 1601 + 234 976-1891 March 24 1712 + 229 976-1952 April 19 1674 + 169 1281-1952 May 56 1677 + 185 946-1952 June 27 1665 + 154 1342-1891 July D2 1693 + 210 1312-2074 August 21 1638 + 176 1281-2044 September 14 1734 + 235 1281-2013 October 20 1591 + 204 1281-1952 November 13 1645 + 181 1342-1952 Totals 238 OS ae DLT) 946-2074 a . Surveys were not conducted in December or January. NOTES 73 1974: Edwards and Ritcey 1960) also suggest similar responses to changing snow conditions and availa- bility of forage. Caribou consistently followed specific travel routes between and within drainages (Figure 1). Routes commonly incorporated natural passes along ridges, frequently followed stream bottoms, invariably pro- ceeded through forested areas, and generally con- nected feeding and resting areas used by Caribou. Most routes were utilized during all seasons. Critical to international movement of animals was the travel route through Kootenay Pass where Caribou had to cross a major highway (Figure 1). Kootenay Pass was the only known route used by Caribou to enter the USA, and historical accounts (Spry 1968, p. 480; Layser 1974) support the impor- tance and long-term use of this route by Caribou. Elevations frequented by Selkirk Caribou indicate that these animals utilize the cedar-hemlock forest and especially the spruce-fir forest year round. Forested areas above 1430 m must be considered vitally important to this population. The close association of Selkirk Caribou to the spruce-fir forest during winter reflects their use of arboreal lichens (Alectoria spp.) as a primary winter food (Freddy 1974). Edwards and Ritcey (1960) also found arboreal lichens to be a principal winter food for Caribou in WGPP. Edwards et al. (1960) indicated that mature stands of spruce-fir, as opposed to cedar-hemlock, produced the greater amount of arboreal-lichen biomass available for consumption by Caribou. The association of A/ectoria spp. with mature stands of spruce-fir places this forage in conflict with extensive clearcut logging (Edwards et al. 1960; T. Ahti. 1962. Ecological investigations on lichens in Wells Gray Provincial Park, with special reference to their importance to mountain caribou. Unpublished re- port, Department of Botany, University of Helsinki, Finland. 63 pp.). Consistent use by Caribou of specific travel routes implies that routes should remain undisturbed to allow established patterns of distribution and range use to continue. Klein (1971) reported that Reindeer (R. t. tarandus) have strong traditions for specific migratory routes and that realignment of such routes is difficult. Proposed expansion of BC Highway 3 to four lanes at Kootenay Pass could easily jeopardize future movements of Caribou into the USA. Klein (1971) reported highways and railroads have ob- structed movements of wild reindeer in Norway. The continued viability of the Selkirk Caribou population depends on cooperative international management. Although Caribou continue to frequent the northwestern USA, in all likelihood perpetuation of mature spruce-fir forests frequented by Caribou in BC is essential to the occurrence of these animals in 74 THE CANADIAN FIELD-NATURALIST the USA. Timber harvest programs and other human developments in both BC and the USA must be coordinated to maintain known movement routes and spruce-fir winter ranges. Acknowledgments Financial support for the work was received from the British Columbia Fish and Wildlife Branch, British Columbia Forest Service, Idaho Department of Fish and Game, Idaho Wildlife Federation— District 1, Inland Empire Big Game Council, National Rifle Association, National Wildlife Federation, Priest Lake Sportsmen’s Association, Sierra Club Foundation, US Forest Service—Region |, Washing- ton Department of Game, Washington State Sports- men’s Council, West Kootenay Outdoorsmen, and the University of Idaho. A. W. Erickson and D.R. Johnson are gratefully acknowledged for initiating the study and obtaining financial support. G. Koehler was invaluable as a field assistant during winter. L. H. Carpenter, R. B. Gill, D.R. Johnson, and D.R. Miller read the manuscript. Literature Cited Banfield, A. F. 1961. A revision of the Reindeer and Caribou, genus Rangifer. National Museum of Canada Bulletin 177. 137 pp. Daubenmire, R. and J. Daubenmire. 1968. Forest vegeta- tion of eastern Washington and northern Idaho. Washing- ton Agricultural Experiment Station Technical Bulletin 60. 104 pp. Edwards, R. Y. and R.W. Ritcey. 1959. Migrations of Caribou in a mountainous area in Wells Gray Park, British Columbia. Canadian Field-Naturalist 73(1): 21-25. Edwards, R. Y. and R. W. Ritcey. 1960. Foods of Caribou Vol. 93 in Wells Gray Park, British Columbia. Canadian Field- Naturalist 74(1): 3-7. Edwards, R.Y., J. Soos, and R.W. Ritcey. 1960. Quantitative observations on epidendric lichens used as food by Caribou. Ecology 41(3): 425-431. Evans, H. F. 1960. A preliminary investigation of Caribou in northwestern United States. M.Sc. thesis, Montana State University, Bozeman. 145 pp. Flinn, P. 1959. The Caribou of northern Idaho. Idaho Wildlife Review 11(5): 10-11. Freddy, D. J. 1974. Status and management of the Selkirk Caribou herd, 1973. M.Sc. thesis, University of Idaho, Moscow. 132 pp. Kelsall, J. P. 1968. The migratory Barren-ground Caribou of Canada. Canadian Wildlife Service Monograph Number 3. 340 pp. Klein, D. R. 1971. Reactions of Reindeer to obstructions and disturbances. Science 173: 393-398. Layser, E. F. 1974. A review of the mountain Caribou of northeastern Washington and adjacent northern Idaho. Journal of the Idaho Academy of Science, Special Research Issue 3. 63 pp. Nelson, E. C. 1947. The Woodland Caribou in Minnesota. Journal of Wildlife Management 11(3): 283-284. Palmer, R.S. 1938. Late records of Caribou in Maine. Journal of Mammalogy 19(1): 37-43. Skoog, R.O. 1968. Ecology of the Caribou (Rangifer tarandus granti) in Alaska. Ph.D. thesis, University of California, Berkeley. 699 pp. Spry, I. M. (Editor). 1968. The papers of the Palliser expedition 1857-1860. Champlain Society, Toronto. 694 pp. U.S. Department of Agriculture. 1972. Kaniksu National Forest multiple use plan. Kaniksu National Forest, Sandpoint, Idaho. 50 pp. Received 9 January 1978 Accepted 10 August 1978 Effects of Fire on the Location of a Sharp-tailed Grouse Arena DONALD A. SEXTON! and MURRAY M. GILLESPIE2 'Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 *Department of Renewable Resources and Transportation Services, Box 14, 1495 St. James Street, Winnipeg, Manitoba R3H 0W9 Sexton, Donald A. and Murray M. Gillespie. 1979. Effects of fire on the location of a Sharp-tailed Grouse arena. Canadian Field-Naturalist 93(1): 74-76. Fire is known to be an important factor in creating and maintaining Sharp-tailed Grouse (Pedioecetes Phasianellus) habitat (Amman 1957; Kirsch and Kruse 1972). Controlled burning has been used to improve Sharp-tailed Grouse habitat in several areas (Amman 1957, 1963; Baumgartner 1939; Grange 1948: Kirsch and Kruse 1972; Miller 1963). Few observations are available, however, on the im- mediate effects of fire on Sharp-tailed Grouse behavior. Amman (1957) suggests that fire does not affect arena use by males. Anderson (1969) reports male Greater Prairie Chicken ( 7ympanuchus cupido) 1979 continuing to use an arena the day after it had been burned. During the spring of 1975 we observed the effects of a fire on a Sharp-tailed Grouse arena near Chatfield, Manitoba (50°47’'N, 97°34’W). Since 1970, spring surveys of arenas have been conducted in the Chatfield area. Known arenas were visited in early morning and numbers of male and female grouse recorded. The counts were conducted three or more times a week from early April until mid- June. Arena A was used by males in 1971 but not in 1972 or 1973. Surveys were not conducted in 1974 owing to flooding of much of the area. During the spring of 1975, a new arena (B) was located 480 m north of the now abandoned arena A, in the same area of open grassland (Figure |). Vegetation in the grasslands (Gr) consisted of native grasses, chiefly Needle Grass (Stipa spartea), June Grass (Koeleria cristata), and wheat grasses (Agropyron spp.) as well as a variety of forbs and scattered Saskatoon (Amelanchier alni- folia) and Snowberry (Symphoricarpos occidentalis) shrubs. The shrub area (Sh) contained clumps of Dwarf Birch ( Betula glandulosa) and Saskatoon. The forested portion (Fo) was dominated by Trembling Aspen (Populus tremuloides). When arena B was discovered on7 May, 10 birds of unknown sex were present. Counts on subsequent mornings showed between 7 and 10 males displayed at this arena prior to 23 May. During this period birds were never observed displaying at arena A. On 23 and 24 May, a fire burned the southwestern portion of the grassland area and eliminated all residual grass and forbs but did not greatly affect shrub or tree cover in the area burned. Arena A was completely burned over whereas arena B was unaffected (Figure 1). On 25 May two non-displaying Sharp-tailed Grouse were present on arena A and eight males were displaying on arena B. Between 26 May and | June, one or two birds were displaying on arena A, and four to seven on arena B each morning. By 16 June only one bird remained on arena B while arena A had five to eight males actively displaying. Other neighboring arenas showed a decrease in the number of males during June, as is typical in late spring (Hamerstrom 1939). On 17 June no birds were on arena B but five birds were still attending arena A. After this date birds were not present on either arena. The difference in vegetative cover between arenas A and B prior to the fire was not obvious. After the fire arena B was still covered by rank residual grass and forbs in addition to the current season’s growth. Arena A had only new growth which was sparse until several weeks after the fire. Several authors have shown that most Sharp-tailed Grouse arenas are in areas of sparse ground cover (Amman 1957; Kobridger 1965; Sisson 1975). Ac- NOTES 15 FIGURE 1. (++) on Chatfield study area. Note grassland (Gr), shrub (Sh), and forest (Fo). Location of arenas and extent of burned area cording to Amman (1957), males may abandon sites with tall or dense ground cover. The shift we observed at Chatfield may be explained by birds moving back to the traditional arena (A) after its vegetation cover was reduced by fire to a more acceptable height. The lack of tall cover would permit displaying males to see and be seen, which are two requirements of arena sites (Anderson 1969). In 1976 and 1977, displaying males used arena A regularly but were never seen where arena B had been in 1975. These observations were made during a study of population, movements, and habitat use of Sharp- tailed Grouse conducted by the Manitoba Depart- ment of Renewable Resources and Transportation Services. We thank G. W. Pepper, S. G. Sealy, and M. W. Shoesmith for comments on the manuscript. 76 THE CANADIAN FIELD-NATURALIST Literature Cited Amman, G. A. 1957. The prairie grouse of Michigan. Michigan Department of Conservation Technical Bul- letin, Lansing. 200 pp. Amman, G.A. 1963. Status and management of Sharp- tailed Grouse in Michigan. Journal of Wildlife Manage- ment 27: 802-809. Anderson, R. K. 1969. Prairie chicken responses to chang- ing booming-ground cover type and height. Journal of Wildlife Management 33: 636-643. Baumgartner, F.M. 1939. Studies in distribution and habits of the Sharp-tailed Grouse in Michigan. Trans- actions of the North American Wildlife Conference 4: 485-490. Grange, W. B. 1948. Wisconsin grouse problems. Wiscon- sin Conservation Department Publication 328. 318 pp. Vol. 93 Hamerstrom, F. N., Jr. 1939. A study of Wisconsin Prairie Chicken and Sharp-tailed Grouse. Wilson Bulletin 51: 105-120. Kirsch, L. M.and A. D. Kruse. 1972. Prairie fires and wild- life. Proceedings of the Tall Timbers Fire Ecology Con- ference 12: 289-303. Kobridger, G. D. 1965. Status, movements, habitats and food of Prairie Grouse on a Sandhills Refuge. Journal of Wildlife Management 29: 788-800. Miller, H. A. 1963. Use of fire in wildlife management. Proceedings of the Tall Timbers Fire Ecology Conference 2: 127-143. Sisson, L. 1975. The Sharp-tailed Grouse in Nebraska. Nebraska Game and Parks Commission. 88 pp. Received 13 February 1978 Accepted 6 August 1978 Brewer’s Blackbird Breeding in the Northwest Territories PHILIP H. R. STEPNEY Ornithology Department, Provincial Museum of Alberta, Edmonton, Alberta T5N 0M6 Stepney, Philip H.R. 1979. Brewer’s Blackbird breeding in the Northwest Territories. Canadian Field-Naturalist 93(1): 76-77. The occurrence of Brewer’s Blackbird breeding in the Northwest Territories was verified at Fort Simpson, 27 June 1977. Additional courting individuals were observed at Hay River on two occasions in the same year. This breeding record represents an appreciable northward expansion of the range of this species and is seemingly related to increased modification of previously forested areas. Key Words: Brewer’s Blackbird, breeding, range extension, Northwest Territories. Brewer’s Blackbird (Euphagus cyanocephalus) has increased both its breeding range (Stepney and Power 1973) and winter range (Stepney 1975) inthis century. Typically distributed in western North America, (A.O.U. 1957), habitat modification resulting from agricultural and transportation route development has seemingly aided the expansion of this species (Roberts 1932). Previously, the Meikle River formed the northern limit of the breeding range of Brewer’s Blackbird in the Peace River region of Alberta (Godfrey 1966). In adjacent northeastern British Columbia, a “family” group had been recorded in 1967 at Fort Nelson, where the birds were observed again in 1974 (Erskine and Davidson 1976). Previous authors had not recorded this species in the Fort Nelson region. There are apparently three earlier records of non-breeding Brewer’s Blackbirds occurring in the territories: Fort Simpson, District of Mackenzie, on 15 June 1958, Baker Lake, District of Keewatin, in November 1923 (Godfrey 1966), and north of Old Crow, Yukon Territory, (three birds) in 1971 (Schweinsburg 1974). In 1977 the author observed Brewer’s Blackbirds in two locations in the District of Mackenzie. At Hay River, on 28 May, two pairs of mated birds were observed along the railroad right-of-way adjacent to the airfield. The males were observed courting and guarding the females and engaging in pursuit flights, behavior typical of breeding Brewers Blackbirds (Williams 1952). Time did not permit a search for nests. On 8 September 1977 Brewer’s Blackbirds were again recorded in Hay River. Four mature males were observed within the town, suggesting that a small colony was present in Hay River that summer. At Fort Simpson, on 27 June, a single pair of adults with five fledglings were observed in the cleared area adjacent to the gravel airstrip within the town. The young were flying among the willow shrubs paralleling the airstrip; the adults mobbed me while I searched unsuccessfully for the nest-site, which probably was located on the grass-covered bank of the airstrip drainage ditch. The occurrence of Brewer’s Blackbirds at Fort Simpson is approximately 360 km northeast of their 1979 previous northern limit in British Columbia. This breeding expansion is appreciable but not unexpected in view of the highway and rail-line developments in this originally forested area. It seems likely that the modified habitats along these rights-of-way provided dispersal routes for Brewer’s Blackbird. This species has been observed penetrating forested areas along cleared rights-of-way in areas of Alberta (Smith 1975), Saskatchewan and British Columbia (A. J. Erskine, personal communication). I acknowledge the assistance of Renewable Re- sources Consulting Services Ltd., with whom I was employed at the time of these observations. Literature Cited American Ornithologists’ Union. 1957. Check-list of North American birds. Fifth edition. American Ornithologists’ Union. Erskine, Anthony J. and Gary S. Davidson. 1976. Birds in the Fort Nelson lowlands of northeastern British Colum- bia. Syesis 9: 1-11. NOTES 77 Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada, Bulletin Number 203, Biological Series Number 73. 428 pp. Roberts, T. S. 1932. The birds of Minnesota: Volume 2. University of Minnesota Press, Minneapolis. Schweinsburg, R. E. 1974. An ornithological study of pro- posed gas pipeline routes in Alaska, Yukon Territory and the Northwest Territories, 1971. Arctic Gas Biological Report Series 10: 1-157. Smith, H. C. 1975. The birds north of Lesser Slave Lake, Alberta. Blue Jay 33: 232-239. Stepney, Philip H.R. 1975. Wintering distribution of Brewer’s Blackbird: historical aspect, recent changes, and fluctuations. Bird-Banding 46: 106-125. Stepney, P. H.R. and Dennis M. Power. 1973. Analysis of the eastward breeding expansion of Brewer’s Black- bird plus general aspects of avian expansions. Wilson Bulletin 85: 452-464. Williams, L. 1952. Breeding behavior of the Brewer Black- bird. Condor 54: 3-47. Received 6 April 1978 Accepted 21 September 1978 A Trap to Measure Populus and Salix Seedfall JOHN C. ZASADA! and ROSEANN DENSMORE2 'USDA Forest Service, Institute of Northern Forestry, Fairbanks, Alaska 99801 2Botany Department, Duke University, Durham, North Carolina 27706 John C. Zasada and Roseann Densmore. 1979. A trap to measure Populus and Salix seedfall. Canadian Field- Naturalist 93(1): 77-79. A seed trap, a wooden frame(1 X 1 m X 5 cm) witha plastic sheeting on the bottom to hold moistened potting soil was used in Alaska to determine seedfall for Populus spp. (poplars) and Salix spp. (willows) through germination of seedlings. This seed trap takes advantage of these species’ ability to germinate rapidly under ambient temperatures and optimum moisture conditions. Key Words: Seed dispersal, natural regeneration, seed trap, Salicaceae, Alaska. The types of seed traps used to measure seedfall vary greatly. Forest tree seedfall is usually measured with rectangular or circular traps which vary in surface area and height of collecting surface above ground (Sarvas 1962; Zasada and Gregory 1972). Werner (1975) described a trap that contained a sticky surface to which seeds adhered. Ryvarden (1971) used a water-filled trap to study seed dispersal in alpine areas of Norway. Although these traps could be used for studying Populus (poplar) and Salix (willow) seedfall in Alaska, two factors make them less than perfect. First, within any part of Alaska, there may be more than one species from each genus contributing to the seedfall. Although these species disperse seed at different times, there can be significant overlap. Thus it is desirable to identify the contribution of each species. Seeds of the two genera can be separated on the basis of color (i.e., Populus seed is tan to white in color, Salix seed is green). Further, it is possible after some experience to separate P. tremuloides (Trembling Aspen) from P. balsamifera (Balsam Poplar). Willow seeds, however, cannot be separated easily on the basis of features identifiable in the field. Second, because of the small size of these seeds it is laborious and time-consuming to separate seeds from other debris that lands in seed traps and then to germinate them. We designed, and have been using for 2 yr, a seed trap that takes advantage of the rapid germination of 78 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE |. Seed trap with aspen seedlings in early July. Toothpicks mark seedling locations. Populus and summer-dispersing Salix under opti- mum conditions of temperature and moisture (Zasada and Viereck 1975). Seeds.of these two genera must germinate shortly after landing on the seedbed or they die (Schreiner 1974; Brinkman 1974). Our trap is a I- X I-m wooden frame (5 cm deep) covered on the underside with one layer of 4-mil plastic sheeting (Figure 1). The trap 1s filled with a standard potting soil, such as Jiffy-Mix,! that is kept wet. If flooding from heavy rains is a problem, holes should be cut in the frame just below soil level. So that soil stays wet, holes should not be punched in the sheeting. It is important that the commercial potting mixes not have an inhibitory effect on germination of these species. We have not observed any effect from Jiffy- Mix! or Jiffy-Mix Plus! in growth chamber studies. Traps should be placed in the field prior to seed dispersal. Germinants are counted periodically and left to develop until they can be identified. Seed collected from identified willows and poplars and germinated on trap soils can be used as an aid to identify unknown germinants. We have used the traps on upland and floodplain sites in interior Alaska. Ona birch (4.9 ha) and aspen (2.8 ha) clearcut we used four traps in each area to measure seedfall. Aspen seedfall was estimated to be 383 seeds/ m2 in the aspen clearcut and 376 seeds/ m2 in the birch clearcuts (coefficients of variation 25% 'The use of trade, firm, or corporation names does not constitute endorsement by the U.S. Department of Agri- culture. and 19%, respectively). The first seed dispersal was observed on 10 June, the first germinants between 12 and 16 June, and the last germinants on 14 July (Table 1). Only aspen seedlings were observed in these traps. Figure | shows traps in early July. Four traps placed on a floodplain produced seedlings of a number of species (Table 2). The traps were set out after dispersal of Salix alaxensis. They were within an 8-yr-old stand (traps | and 2) or onan open sandbar adjacent to it (traps 3 and 4). The main species in this stand were S. interior, S. alaxensis, and P. balsamifera;, less common species were S. novae- anglicae and S. monticola. Seedlings of species that are rare or absent on these sites (e.g., S. bebbiana, P. tremuloides) were produced as well as those of species occurring in the stand. TABLE |—Number of aspen germinants and percent of total number germinating on a given date in four seed traps located in clearcuts on upland sites in 1977 Aspen Birch Number of germinants Mean + sD 383 +99 370 22 7/2) (range) (277-S 16) (353-436) Percent germinating 16 June 3p).3) 46.8 23 June 26.2 36.7 3 July 7.8 dil 7 July 9.6 7.5 14 July let Hes) 79 NOTES 79 TABLE 2—Species composition and number of germinants observed in four seedtraps located on floodplain sites adjacent to the Tanana River, Alaska Traps within 8-yr-old stand Traps on open sandbar Species | 2 3 4 Salix lasiandra' 12 5 2 S. bebbiana 33 50 5 12 S. arbusculoides 3 2 = = S. monticola 5 0 = — S. interior 45 281 7 26 S. novae-anglicae 6 2 | I S. planifolia I 2B l | Populus balsamifera 4 2 5 5 P. tremuloides 41 49 48 3 Unknown Salix sp. 14 4 9 3 'Terminology follows Argus (1973). Our observations demonstrate that the seed trap is feasible, but several drawbacks exist. First, depen- ding on the weather conditions, the traps may have to be watered as frequently as every day. Second, the variable measured with these traps is the number of germinants, which is not necessarily the total number of potentially viable seeds reaching the trap. We believe that by keeping the traps saturated, the number of germinants is a good estimate of the number of viable seeds; but this point needs further examination. Literature Cited Argus, George W. 1973. The genus Sa/ix in Alaska and the Yukon. National Museum of Natural Sciences, National Museum of Canada, Publications in Botany Number 2. 279 pp. Brinkman, Kenneth A. 1974. Salix L. In Seeds of woody plants in the United States. USDA Agriculture Hand- book Number 450. pp. 746-750. Ryvarden, L. 1971. Studies in seed dispersal. I. Trapping of diaspores in the alpine zone of the Finsé, Norway. Norwegian Journal of Botany 18: 215-226. Sarvas, Risto. 1962. Investigations on the flowering and seed crop of Pinus silvestris. Finnish Forest Research Institute Publication 53.4. 198 pp. Schreiner, Ernst J. 1974. Populus L. In Seeds of woody plants in the United States. USDA Agriculture Handbook Number 450. pp. 645-655. Werner, P. A. 1975. A seed trap for determining patterns of seed deposition in terrestrial plants. Canadian Journal of Botany 53: 810-813. Zasada, John C. and Robert A. Gregory. 1972. Paper birch seed production in the Tanana Valley, Alaska. USDA Forestry Service Research Note PNW-177. Pacific North- west Forest and Range Experiment Station, Portland Oregon. 7 pp. Zasada, John C. and Leslie A. Viereck. 1975. Effect of temperature and stratification on germination of Alaskan Salicaceae. Canadian Journal of Forestry Research 5: 333-337. Received 7 April 1978 Accepted 19 August 1978 Examination of Overwintering Adult Carabid Beetles for Associated Mites J. E. OLYNYK and R. FREITAG Department of Biology, Lakehead University, Thunder Bay, Ontario P7B S5El Olynyk, J. E. and R. Freitag. 1979. Examination of overwintering carabid beetles for associated mites. Canadian Field- Naturalist 93(1): 79-81. Mites are commonly associated with adult carabids in northwestern Ontario from spring to fall but are not normally found on the beetles during winter. This suggests that the mites are phoretic on the carabids, using them for transport during times of the year when the carabids are active. Key Words: northwestern Ontario, mites, beetles, Carabidae, Acari, ectoparasites. 80 THE CANADIAN FIELD-NATURALIST In North America, collections of carabids during winter have been rare and no previous work has been done on mite populations associated with hibernating carabids. Aitchison (1976), using pitfall traps, studied numerous invertebrate groups in subnivean environ- ments in southern Manitoba. She collected very few carabids in December and March and none in January or February. Olynyk and Freitag (1977) did not capture any carabids in pitfall traps near Thunder Bay, Ontario, between 9 November and 6 April, though Olynyk (1978) found an average of approxi- mately eight mites per captured carabid between May and October. Larochelle (1972a, b, 1973, 1974) made several collections of hibernating adult carabids in southern Quebec but did not record the presence of mites on the beetles (personal communication). The purpose of this study was to determine winter associations of ground beetles and mites in the context of annual relationships. Materials and Methods Carabids were collected between 6 and 26 April 1977 in 200 pitfall traps near Thunder Bay, Ontario (Olynyk and Freitag 1977). Three hand collections were made in the same location in 1977 on 19 November and 3 and 23 December. Mounds of earth, rotten stumps and logs, and surrounding areas on, in, and under the leaf litter were inspected. Larochelle (1974, 1975) noted that such locations were some of the most productive sites for collecting carabids in winter. On 22 April 1978, hand collections were made beside Ontario Highway 588 near the junction of Highways 11/17 and 588 in northwestern Ontario. Areas under loose materials including logs, metal, tires, boards, and cardboard were inspected. All carabids collected were placed in individual vials of 70% ethanol. External surfaces and areas under elytra were examined on each beetle collected under a Wild MS dissecting microscope. Alcohol from vials containing carabids was filtered through Kim- wipe tissue and the tissue then examined for mites. Carabids were identified to species using identifica- tion keys by Lindroth (1961-1969). Results and Discussion Table | contains a list of the carabids collected. On the 53 carabids collected, representing five genera and Six species, we found no mites on external surfaces or | under elytra. Carabids collected in November and December were found beneath rotten stumps and were torpid. Approximately 80% of the carabids collected on 22 April were found beneath a sheet of metal, the rest beneath a nearby sheet of cardboard. Burrows were present, the carabids were congregated and many were torpid. The ground was frozen. Vol. 93 TABLE 1—Carabids collected in early spring and winter of 1977 and 1978 in pitfall traps and by hand near Thunder Bay, Ontario Sex Date Carabid species on 9 26 April 1977 Prerostichus adstrictus l 3 P. pensylvanicus 4 7 19 November 1977 Sphaeroderus nitidicollis I 2 Calosoma frigidum l P. pensylvanicus 2 Agonum decentis ] 3 December 1977 S. nitidicollis l 23 December 1977 S. nitidicollis I 22 April 1978 P. pensylvanicus | 3 Calathus ingratus 1] 14 Greene (1975) found acarid hypopi appearing on carabids in August in Washington, with many hundreds commonly present on the carabids by October. No data were recorded for hibernating carabids, but few if any mites were present on carabids in spring, which suggests that the mites did not overwinter on the carabids. W.M. Graham of Lakehead University studied ants and associated mites populations near Thunder Bay in 1969 and 1970 (March 1978, personal communication). Average mite numbers per ant were lowest in spring and fall, suggesting that the mites did not overwinter on the hibernating ants. Olynyk (1978) noted that the numbers and frequency of occurrence of mites on carabids were lower in spring and fall. As none of the 53 hibernating adult carabids examined carried mites, it can be assumed that the mites do not overwinter on the carabids. Aitchison (1976) found that carabids in southern Manitoba were inactive at temperatures below about -2°C. She collected large numbers of mites in mid- winter and concluded that their activity seemed to be unrestricted by temperatures encountered in her study. Mites phoretic on carabids, using them solely as a transport mechanism, would gain no such benefit from hibernating carabids. It follows that mites phoretic on carabids in other regions likewise are not found on hibernating carabids. Acknowledgments We are indebted to J. P. Walas for his aid in collecting specimens and to the National Research Council of Canada for financial support (Grant No. A4888). IDG) Literature Cited Aitchison, C. W. 1979. The activity of subnivean inverte- brates in Southern Manitoba. M.Sc. thesis, University of Manitoba, Winnipeg, Manitoba. 197 pp. Greene, A. 1975. Biology of five species of Cychrini (Coleoptera: Carabidae) in the steppe region of South- eastern Washington. Melanderia 19: 1-43. Larochelle, A. 1972a. Collecting hibernating ground beetles in stumps. (Coleoptera: Carabidae). Coleopterists’ Bulletin 26(1): 30. Larochelle, A. 1972b. Collecting hibernating Carabidae under snow. Proceedings of the Entomological Society of Washington 74(4): 473. Larochelle, A. 1973. Collecting hibernating ground beetles under snow. Entomological News 84: 82. Larochelle, A. 1974. Winter habits of carabid beetles NOTES 81 (Coleoptera: Carabidae). Great Lakes Entomologist 7(4): 143-145. Larochelle, A. 1975. La chasse aux Carabidae en hiver. Cordulia 1(4): 117-118. Lindroth, C. H. 1961-1969. The ground-beetles of Canada and Alaska. Opuscula Entomologica Supplementum. 1192 pp. Olynyk, J. E. 1978. Carabidae (Coleoptera) and associated Acari in Northwestern Ontario. M.Sc. thesis, Lakehead University, Thunder Bay, Ontario. 73 pp. Olynyk, J. and R. Freitag. 1977. Collections of spiders beneath snow. Canadian Field-Naturalist 91(4): 401-402. Received 15 May 1978 Accepted 29 August 1978 Ring Counts in Salix arctica from Northern Ellesmere Island D. B. O. SAVILE Biosystematics Research Institute, Central Experimental Farm, Ottawa, Ontario KIA 0C6 Savile, D. B. O. 1979. Ring counts in Salix arctica from northern Ellesmere Island. Canadian Field-Naturalist 93(1): 81-82. In 1962 I collected two large specimens of Salix arctica Pall., which had been eroded out of steep- sided postglacial benches roughly 100 m above the present level of Lake Hazen, Northwest Territories, at about 81°50’N, 71°15’W. They were to have been picked up by R. E. Beschel, but, when he failed to do so, they lay forgotten in a drawer for years. I have now cut sections from the main stem and a major branch of each specimen. The sections were polished, cleared with lacquer, and ring counts were made under the dissecting microscope. The short cool growing season in the high Arctic makes some rings very narrow or incomplete; and some such rings may not be definitely distinguishable even under the microscope. The larger plant (Savile 4841A) yielded counts of 84, 84, and 85 (at least partial) rings on a radius of about 36 mm from the markedly eccentric origin. These are minimum counts, because some of the early rings are thought to have included some with almost zero growth. The outermost rings are relatively wide, indicating vigorous growth in the final years of the life of the plant, A large branch yielded 36 rings without any wide outer ones. Possibly death of this branch diverted nutrients into the main stem. R. E. Beschel and D. Webb (1964. Axel Heiberg Island Preliminary Report. Edited by F. Muller. McGill University, Montreal. pp. 189-198. Processed) indicate that S. arctica periodically loses branches. The second specimen (Savile 484/ B), with maxi- mum wood radius of 28 mm, yielded a minimum of 43,44, 44, and doubtfully 48 rings in separate sections, and again most of the last few rings were relatively wide. Sections from both specimens have been deposited in DAO. I suspect that both plants were killed by erosion of the bench on which they grew, because vigorous growth in their last years does not suggest the senescence inferred by Warren Wilson (1964) for plants of this species at Resolute (74°41’N), where little or no growth seemed to occur after about SO yr. Although the Hazen Camp specimens are from about 81°50’N, they are better grown than any at Resolute, largely because Hazen Valley, being ringed by mountains, is subject to dynamically warmed winds and has relatively high summer temperatures; but partly, perhaps, because sites such as the raised branches have good drainage and a deep active layer. The difference is reflected in the flora: about 115 species of vascular plants in Hazen Valley and 70 at 82 THE CANADIAN FIELD-NATURALIST | Resolute. For comparison of these and some other arctic sites see Savile (1972, p. 11). It should be noted that on some sites, notably depressions flooded by Lake Hazen in midsummer, Salix arctica makes very poor growth. It seems probable that, barring soil movement or other accidents, plants on favorable sites in Hazen Valley have either an indefinite life or a much higher life limit than at Resolute. It is worth noting that, of the 12 plants counted from Axel Heiberg Island (all about 79°25’ to 79°30’N) by Beschel and Webb, one showed 87 and all the others between 18 and 64 rings. My specimen, 484/ A, is perhaps the oldest recorded north of 80°N. A few large and vigorous plants up to about 3-m spread were seen near Hazen Camp, but I did not feel justified in sacrificing any of them for the Vol. 93 sake of a ring count. I am grateful to a reviewer of this manuscript for drawing my attention to the mention by Raup (1965) of specimens of Salix arctica at about 73°N in Northeast Greenland of up to 210 and 236 yr old. Literature Cited Raup, H. M. 1965. The structure and development of turf hummocks in the Mesters Vig district, Northeast Green- land. Meddelelser om Gronland 165(3): 1-112. Savile, D. B. O. 1972. Arctic adaptations in plants. Canada Department of Agriculture, Research Monograph 6. Warren Wilson, J. 1964. Annual growth of Salix arctica in the high Arctic. Annals of Botany (London) 28: 71-76. Received 4 July 1978 Accepted 5 September 1978 Giant Cow Parsnip (Heracleum mantegazzianum) on Vancouver Island, British Columbia NEIL K. DAWE! and ERIc R. WHITE? 'Canadian Wildlife Service, R.R. 1, Qualicum Beach, British Columbia VOR 2T0 °5312 Ewart Street, Burnaby, British Columbia V5J 2W4 Dawe, Neil K. and Eric R. White. 1979. Giant Cow Parsnip ( Heracleum mantegazzianum) on Vancouver Island, British Columbia. Canadian Field-Naturalist 93(1): 82-83. The known distribution of the Giant Cow Parsnip ( Heracleum mantegazzianum) on Vancouver Island, British Columbia is reported. A discussion of the time span the plant has taken to become established is included along with notes on the dispersal of the plant from one site of introduction. Key Words: distribution, Heracleum mantegazzianum, Vancouver Island, weeds. The distribution of the Giant Cow Parsnip (Her- acleum mantegazzianum) in Canada was recently described by Morton (1978) as occurring primarily in southern Ontario. He also cites a record from Vancouver on the west coast (Kamermans 1977): however, there is no mention of the plant’s occur- rence on Vancouver Island. Taylor and MacBryde (1977) list but one species of Heracleum for British Columbia: H. sphondylium (= H. lanatum). This note documents the occurrence of H. mantegazzianum on Vancouver Island and will alert naturalists to the possible occurrence of the species in other areas of the province. During summer field work of 1978 we discovered a large number of H. mantegazzianum plants growing along the banks of French Creek, near the creek mouth, approximately 6 km NW of Parksville, British Columbia (49°21’N, 124°22’W). Subsequently two other groups of plants of that species were brought to our attention. Jennifer McGown (personal com- munication) told us of the first group. That group consisted of a small cluster of plants in a vacant Fifth Street lot at Nanaimo, British Columbia, approxi- mately 35 km SE of the French Creek site. The cluster was growing alongside a dry ditch edge amongst a large patch of Rubus discolor. All the plants were taller than 3 m, and all had gone to seed. The second group was discovered near a vacant lot at Victoria, British Columbia in circumstances similar to those of the Nanaimo specimens (Harold Hosford, personal communication). Specimens from these sites were verified (T. C. Brayshaw, personal communication) 1979 and are now onfile at the British Columbia Provincial Museum, Victoria. After the lodging of our H. mantegazzianum species with the Provincial Muse- um, Brayshaw informed us that another specimen was located in the museum herbarium. That specimen, collected in 1973 from the French Creek site, was incorrectly labelled H. lanatum. At the French Creek location the plants were found in large clusters on both sides of the creek. One collected specimen measured 395 cm from the base to the top of the terminal umbel, with a terminal umbel width of 70 cm. The stem was 9 cm thick at the base. The lower 26 cm of the stem was a deep purple while the rest of the stem was green covered with purple spots and blotches decreasing in extent toward the umbel. Clear hairs protruded from most of the spots. The fruit was elliptic and ranged from 12.0 to 13.2 mm in length. The vittae were swollen near the base and at that point averaged slightly over | mm in width. Further investigation revealed large numbers of H. mantegazzianum upstream in suitable locations and subsequently they were found growing along a tributary of French Creek where it crosses Swayne Road, approximately 2km S of Coombs, British Columbia. The plants grew along the banks of the creek, south to a point just past the intersection of the tributary and Winchester Road. Here the local distribution of the plant abruptly ended. French Creek and another of its tributaries crosses Win- chester Road a further 1500 m and 600m north, respectively; however, an inspection of both sites did not reveal any specimens of H. mantegazzianum. Because of that, we feel the area of introduction of the plant lies in the general location of the southernmost tributary of French Creek near its crossing of Winchester Road. We talked with local residents, and in particular to Frances Kroot (personal communication) who lives near the presumed site of introduction, to try to determine the time span H. mantegazzianum has taken to establish itself. Apparently the plant was not known to occur in the area in 1937; however, by 1944 it appeared sporadically where the forest edged away from the creek, and beside bridges where openings were created. Here the plant’s history becomes nebulous, but by 1967, when Frances Kroot came to the area, the plant was well established. By 1975, and likely years before, it was known to be well estab- lished near the mouth of French Creek, some 12 km downstream. As Morton (1978) reports that the principle mode of dispersal is by water, this further supports our conclusions regarding the site of introduction. All the residents we talked with mentioned the stinging effect and the blisters which occur upon coming in contact with H. mantegazzianum, as NOTES 83 described by Morton (1978), and all requested information on methods of eradicating the plant. Bert Topliffe (personal communication) described his attempts to rid them from his yard. He made repeated applications of the herbicide 2,4,5-T and this seemed to be effective shortly after application. The following year, however, they reappeared as numerous as before. He has since found that only by continually mowing the plants does he have any success. Topliffe also mentioned observing a number of people each fall collecting the large umbels. He recalled one occasion when a couple loaded the back of their pick-up truck with the dried stalks and umbels. This, undoubtedly, could disperse the plant over large distances and may account for the Nanaimo specimens. The discovery of H. mantegazzianum on Van- couver Island appears to warrant the concern ex- pressed by Morton (1978). Hazards of coming into direct contact with the plant have already been mentioned. Perhaps more important, however, Is its ability to become well established within a relatively short time, likely at the expense of some of our native species. On Vancouver Island H. mantegazzianum established itself within 30 yr along suitable areas of French Creek and one of its tributaries, moving a distance of approximately 12 km within a period of less than 40 yr. It is well established along virtually the entire waterway from the site of introduction to the French Creek mouth. In some locations the plant grows 80 to 100 m away from the creek edge. The plant’s potential for spreading even further is in- creased through the dispersal of its seeds by humans in their quest of dried umbels, presumably for home decorations. Coupled with those factors are the difficulties encountered in trying to eradicate the plant once it is established. As Morton (1978) points out, we need more information as to the distribution, life history aspects, and control methods of H. mantegazzianum in Canada. Naturalists should also be alert to the possibility that the piant occurs within their area and has to date been passed off as an aberrant form of a locally common species. Literature Cited Kamermans, J. K. 1977. A hairy horror in my Huron haunts. Wood Duck 31: 49-50. Morton, J. K. 1978. Distribution of Giant Cow Parsnip (Heracleum mantegazzianum) in Canada. Canadian Field-Naturalist 92: 182-185. Taylor, R. L. and B. MacBryde. 1977. Vascular plants of British Columbia. University of British Columbia Press, Technical Bulletin Number 4. Received 7 September 1978 Accepted 4 October 1978 84 THE CANADIAN FIELD-NATURALIST Vol. 93 Nesting of Horned Puffins in British Columbia R. WAYNE CAMPBELL,! HARRY R. CARTER,2 and SPENCER G. SEALY? '\British Columbia Provincial Museum, Victoria, British Columbia V8V 1X4 2905 Deal Street, Victoria, British Columbia V85 5G4 3Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2 Campbell, R. Wayne, Harry R. Carter, and Spencer G. Sealy. 1979. Nesting of Horned Puffins in British Columbia. Canadian Field-Naturalist 93(1): 84-86. Summer observations of Horned Puffins (Fratercula corniculata) from 1972 through 1977 suggest that this species is expanding its range along the British Columbia coast. Nesting was confirmed at one new site and probably occurred at three others. Key Words: Horned Puffins, Fratercula corniculata, British Columbia, breeding range. The status of the Horned Puffin (Fratercula corniculata) on the west coast of North America south of its breeding range in southeastern Alaska (A.O.U. 1957) was reviewed recently by Sealy and Nelson (1973) and Hoffman et al. (1975). These authors considered it a late spring and early summer (and occasional winter) visitor to this area but did not report evidence of breeding. Forrester Island, Alaska, was heretofore the southernmost known breeding site in the eastern Pacific (Heath 1915: Willett 1915). Censuses of, and other work in, seabird colonies along the British Columbia coast from 1972 through 1977 indicate (Table 1) that the Horned Puffin is now more widely distributed along the west coast of the Queen Charlotte Islands and Vancouver Island than previously known. It also has been recorded on the central mainland coast. Possible Nesting Sites Marble Island (52° 12’N, 132° 39’W). This is a fairly large, wooded island, about 150 m high, with dense undergrowth above the precipitous cliffs. On 19 July 1977 two Horned Puffins scrambled from a natural rock crevice on a large grassy slope among a colony (hundreds) of Tufted Puffins (Lunda cirrhata). Later, five Horned Puffins were seen at one time circling off the island with Tufted Puffins. During the half hour the Horned Puffins were watched, an adult several times flew to, and almost landed near, its presumed nest site. The following day two Horned Puffins were again seen flying with Tufted Puffins off the island. Unnamed Islet (52°06'N, 131° 14’W). This islet, off the northwest coast of Anthony Island, is about 30 m high, is composed of bare rock with deep cracks and fissures, and has grassy areas along its north tip. Six Horned Puffins were seen flying with Tufted Puffins close to the islet on 2 July 1977. Two days later, a Horned Puffin flew from a crevice near the top of a steep rocky bluff: later we discovered a recently cracked egg in the crevice. Later the same day we watched up to five Horned Puffins, for an hour or so, fly very close to the islet, but none landed or entered a natural crevice. All other possible nest sites were inaccessible without climbing equipment. On 5 August, four Horned Puffins were flushed from rocky vantage points above where they had been seen in early July. Cape St. James (51°56, 131°01’W). This landform consists of the St. James Islands, which are saddle- shaped, bare, and grassy with vertical cliffs about 30 m and the Kerouard Islands, which are mostly grassy, unforested, and up to 76m high. Horned Puffins were recorded near Cape St. James during the summers of 1972 through 1974 and also in 1977. H D. Fisher (letter to SGS, 26 September 1974) was not aware “that the occurrence of Horned Puffins at the Cape (May to August) and carrying fish at that, would be valuable news. The common puffin in the Kerouards is the Tufted Puffin. But among these we always saw a few Horned Puffins. Whenever we were enroute to and from the sea-lion rookeries, they [Horned Puffins] seem to nest on the cliff directly below the weather station on the northernmost Kerouard, a very inaccessible place.” Triangle Island (50°52’N, 129°05’W). This, the most western of the Scott Islands, about 210 m high, is very precipitous with no trees. Horned Puffins were first recorded here in 1972 by C. J. Guiguet and have been observed nearly every summer since by biologists visiting or conducting research (Table 1; Vermeer et al. 1976). Although positive evidence is lacking, up to four pairs may breed on Triangle Island, all at the western end and near “Murre” rock. Discussion The observations presented here and those of Sealy and Nelson (1973) suggest that the Horned Puffin is more common in British Columbia than previously known. If this trend continues, opportunity will exist NOTES 85 TABLE !|—Summer observations of Horned Puffins in British Columbia, 1972-1977 1979 Locality! Date Queen Charlotte Islands Cox Island (Langara I.) 31 July 1977 Darwin Sound May 1977 Marble Island 19 July 1977 Marble Island 20 July 1977 Island NW Anthony I. 2 July 1977 Island NW Anthony I. 4 July 1977 Island NW Anthony I. 5 August 1977 Island E Anthony I. 4 July 1977 Cape St. James May-August 1972 Cape St. James May-—August 1973 Cape St. James May-August 1974 Cape St. James 5 July 1975 Cape St. James 27 July 1975 Cape St. James 3 July 1977 Cape St. James 4 July 1977 Mainland Coast Sinnett Island 27 June 1976 Joseph Island 28 June 1976 Vancouver Island Triangle Island 1 July 1972 Triangle Island 16 June 1974 Triangle Island 2 July 1974 Triangle Island Triangle Island 21 August 1974 June-August 1975 Triangle Island 27 July 1977 Triangle Island 28 July 1977 Triangle Island 1 August 1977 Solander Island 27 June 1975 Barrier Islands 26 June 1975 Wichaninnish Bay 20 June 1975 Number? Remarks | Flying with Tufted Puffins 4 In a flock flying south 5 Suspected nesting 2+ Flying with Tufted Puffins 6 Suspected nesting 5 Suspected nesting 4 Suspected nesting l Flying alone 1 or 2 With Tufted Puffins 1 or 2 With Tufted Puffins 1 or 2 With Tufted Puffins 7 With Tufted Puffins 2 With Tufted Puffins | With Tufted Puffins 2 With Tufted Puffins l Alone off island on water | Flying alone 2 Flying with Tufted Puffins I+ On water near island 2+ On water near island | On nesting cliffs 1-6+ One carrying fish | On rocks on island 3 On water off island 8 On rocks 3 Flying with Tufted Puffins On water off island ! On water ‘Listed from north to south. 2All observations of birds in apparent definitive plumage. to monitor it. Therefore, it is important to interpret carefully our observations so that its status in British Columbia up to 1977 is clear. The finding of a Horned Puffin nest containing an egg, On an unnamed islet northwest of Anthony Island in 1977, provides positive proof of breeding. The observations (Table 1) of birds carrying fish in their bills is evidence short only of finding an egg or chick. Where the above evidence is lacking, observations must be viewed with caution. Myrberget (1959) found that the varying proportion of non-breeding Common Puffins (F. arctica) present in a colony may equal the number of breeding birds there, and after 4 yr of age many individuals prospect for and even “own” burrows but seldom breed (Petersen 1976a). Many more 5-yr-old birds breed but maximum reproductive output does not occur until the birds are 10-11 yr old. In Alaska, many occupied Horned Puffin bur- rows during the breeding season never reach the egg stage, possibly because their “owners” are immatures (D. H. S. Wehle, personal communication). Such im- matures in the Common Puffin are recognizable only on the basis of bill shape and the number and depth of » bill furrows (Petersen 1976b). The presence of a Horned Puffin on a boulder or even in a burrow therefore does not imply nesting. Also, as noted by Lockley (1953), and confirmed by Petersen (1976a), Common Puffins that circle the periphery of a colony are generally immature. The breeders that are feeding young generally fly straight out to sea on leaving the burrows and straight back (carrying fish) when returning. In fact, Brun (1971) considered only those puffins carrying fish in their bills to be breeders when he censused colonies in northern Norway. Hence the presence of Horned Puffins of unknown age at best provides suggestive evidence for breeding. The finding of additional nests and additional observations of birds carrying fish will be required to confirm that this species is continuing to change its status in British Columbia. 86 THE CANADIAN FIELD-NATURALIST Another puffin, the Rhinoceros Auklet (Cero- rhinca monocerata), is actively expanding its breeding range (see Scott et al. 1974) and is recolonizing islands from which it had been exterminated (see Ainley and Lewis 1974). Acknowledgments We appreciate the assistance of R. Billings, T. Carson, H D. Fisher, J. B. Foster, H. M. Garrioch, S. M. Guiguet, M. A. Paul, M. C. Lee, M.S. Rod- way, M. G. Shepard, and K. R. Summers with field aspects of this work. C. J. Guiguet and J. B. Foster kindly read the manuscript. Supporting funds were received from the Ecological Reserve Unit, Depart- ment of Environment, Victoria, British Columbia. Literature Cited Ainley, D. G. and T. J. Lewis. 1974. The history of Fara- llon Island marine bird populations. Condor 76: 432- 446. American Ornithologist’s Union.1957. Checklist of North American birds. Sth edition. Lord Baltimore Press, Baltimore, Maryland. Brun, E. 1971. Census of puffins (Fratercula arctica) on Nord-Fugloy, Troms. Astarte 4: 41-45. Vol. 93 Heath, H. 1915. Birds observed on Forrester Island, Alaska, during the summer of 1913. Condor 17: 20-41. Hoffman, W., W.P. Elliott, and J. M. Scott. 1975. The occurrence and status of the Horned Puffin in the western United States. Western Birds 6: 87-94. Lockley, R. M. 1953. Puffins. J. M. Dent, London. 186 pp. Myrberget, S. 1959. Vekslinger 1 antall lundefgul inne ved kolonian. Sterna 3: 239-248. Petersen, A. 1976a. Age of first breeding in puffin, Fra- tercula arctica L. Astarte 9: 43-50. Petersen, A. 1976b. Size variables in puffins Fratercula arctica from Iceland, and bill features as criteria of age. Ornis Scandinavica 7: 185-192. Sealy, S. G. and R. W. Nelson. 1973. The occurrence and status of the Horned Puffin in British Columbia. Syesis 6: 51-55. Scott, J. M., W. Hoffman, D. Ainley, and C. F. Zeille- maker. 1974. Range expansion and activity patterns in Rhinoceros Auklets. Western Birds 5: 13-20. Vermeer, K., K. R. Summers, and D.S. Bingham. 1976. Birds observed at Triangle Island, British Columbia, 1974 and 1975. Murrelet 57: 35-42. Willett, G. 1915. Summer birds of Forrester Island, Alaska. Auk 32: 295-305. Received 5 April 1978 Accepted I] August 1978 News and Comment Special Appreciation Expressed to R. Emerson Whiting Weare particularly grateful to R. Emerson Whiting for his careful and thorough preparation of the Index for The Canadian Field- Naturalist from 1975 to 1978 (Volumes 89, 90, 91, and 92). His particular concern for accuracy and his attention to details is not only appreciated by the journal but more so by the users of the Index. We sincerely thank Mr. Whiting for his time and effort because the Index is such an important component of each volume. We are pleased to announce that W. Harvey Beck has agreed to take on the task of indexing Volume 93. LORRAINE C. SMITH Editor Second International Congress of Systematic and Evolutionary Biology The Second International Congress of Systematic and Evolutionary Biology (ICSEB-II) will be held at the University of British Columbia, Vancouver, Canada on 17-24 July 1980. The provisional list of symposia topics include these: 1. Arctic refugia and the evolution of arctic biota. Origins and evolution of the north Pacific marine biota. Evolution of reproductive strategies. Evolutionary epigenetics. Evolution of community structure. Green algae and land plant origins. Macromolecular mechanisms in evolution. Se stacey Books Available from IUCN (International Union for Conservation of Nature and Natural Red Data Book — Birds The first part of the completely revised edition of the Red Data Book Volume IT Aves is now available from IUCN. A second part of similar size will be published next year. This new edition has been prepared by Warren King on behalf of the International Council for Bird Preservation (ICBP) and the Survival Service Com- mission of IUCN. King has drawn on the expertise of hundreds of ornithologists throughout the world, including ICBP’s National Sections and Working Groups. The volume covers 199 bird taxa and includes birds that are threatened throughout their range. It does not deal with taxa that may be a threatened category within the boundaries of one nation but are relatively abundant elsewhere. Price (includes binder and dividers) is $30 (US). 87 8. 9. 10. ite Allozymes and evolution. Coevolution and foraging strategy. Evolution of colonizing species. Rare species and the maintenance of gene pools. 12. Paleobiology of the Pacific rim. Sessions for contributed papers and for papers in specialized fields, taxonomic as well as methodo- logical, will also be organized. Those interested in receiving an information circu- lar in the spring of 1979, should write to Dr. G.G.F. Scudder, Department of Zoology, the University of British Columbia, 2075 Wesbrook Mall, Vancouver, British Columbia V6T 1WS. Resources) World Directory of Protected Areas The second instalment of the World Directory of National Parks and Other Protected Areas has been published. Covering a further 43 countries it brings the total number represented in the Directory (the first instalment was published in 1975) to 60. This second instalment also contains additional sheets for five countries previously covered: Australia, Canada, Congo, Iran, UK. Now greatly enlarged the Directory is divided into two volumes: countries A-K, 317 pages; countries L-Z, 330 pages. The 397 new sheets of the second instalment plus the cover for Volume Two may be obtained direct from IUCN, price $60 (US) post free. For the complete two-volume set comprising covers, alphabetical dividers and 647 sheets, the price is $85 (US). 88 THE CANADIAN FIELD-NATURALIST Societies and Amateur Ornithologists The following are taken from the recommendations of the National (USA) plan for Ornithology submitted to the National Science Foundation and the Council of the American Ornithologists’ Union (AOU). 10. Professional societies (preferably the AOU or an intersociety consortium) should form a committee charged with (a) developing projects of national ornithological importance for amateurs, (b) develop- ing centers of responsibility for particular projects, and (c) providing professional assistance in securing and maintaining financial support for the projects. 11. The ornithological societies should undertake the development of a network of involvement by local bird clubs and nature centers. The initial step should be the preparation of a computerized listing of all clubs and contacts. The ultimate goal should be a Federation of American Bird Clubs or an ATO (American Trust for Ornithology) modelled on the British counterpart. 12. The AOU should take the initiative in improv- ing formal coordination in the activities of the ornithological societies and in the work of the profes- sion in general. An obvious option for this, perhaps only as a developmental stage, would be the formation of an intersociety coordinating council whose role would be to consider or initiate long-range policy of IUCN Views on Whale Management The International Union for Conservation of Nature and Natural Resources (IUCN) is committed to the sustainable use of natural resources, including the living resources of the seas. It believes that it is possible to manage whaling ona sustainable basis and is concerned, therefore, that after three years the International Whaling Commission’s (IWC’s) New Management Policy has not been able to achieve this goal. Although the IWC and its Scientific Committee have made commendable efforts to improve man- agement procedures, the outcome has been generally unsatisfactory and there have been some extra- ordinary lapses; for example, the manner in which the quota for North Pacific Sperm Whales was sharply raised in December last year on the basis of questionable data and an inadequate model. The IUCN is also deeply concerned that certain member states so far have not met the IWC’s requirement that full and accurate data and analyses be furnished to the Commission. Those few states complying find their data subjected to intense scrutiny Vol. 93 national or general professional scope, as well as to assist in the integration of routine activities. 13. The ornithological societies should take greater interest in the work of ornithological centers such as organized bird observatories (e.g., Manomet, Point Reyes Bird Observatory) and aid in their development and promotion. 14. The ornithological societies should establish training programs in basic ornithology and research techniques for amateurs. Amateur-oriented work- shops supervised by professionals should become a regular part of annual meetings. 15. Annual meetings of the ornithological societies should be better structured to promote contacts and communication between amateurs and professionals and between junior and senior mem- bers. Part of this goal can be reached through Recommendation 14, but this will not be sufficient. Explicit procedures should be devised, such as scheduled round-table discussions, that will overcome diffidence or reticence and automatically put ama- teurs and professionals and junior and senior mem- bers in personal contact in an arena of common subject-matter interest. From ASC Newsletter 6(5): 53-54, 1978. (Association of Systematics Collections) while those not complying may be awarded quotas in line with past custom rather than with the present conditions of the stocks. The work of the IWC is undermined by member states when they do not provide requested data and when they encourage whaling activities of non-members and report less than their total catch. IUCN suggests to the !WC that, to end such practices and to allay suspicion, it requires its members to provide original catch records and it re-opens discussion of the possibility of placing truly international observers, accredited by the IWC, to all whaling operations. IUCN urges the IWC and member states — to ensure that the quotas they set this year are conservative, particularly with respect to those stocks about whose status there 1s considerable doubt and those stocks which are part of multi- species fisheries; —to ensure that current management policy is thoroughly revised in time for greatly improved procedures to be applied in 1979; —to make good their commitment to gather 1979 adequate data on whales and the ecosystems of which they are part by implementing without further delay the International Decade of Cetacean Research; — to accelerate the revision of the Convention and ensure that its conservation provisions are considerably strengthened and the needs of whale conservation adequately catered for in related instruments for the management of marine resources. Failure to take these measures will confirm IUCN and the rest of the international community in their view — expressed repeatedly since 1972 — that there is no rational alternative to a moratorium on com- mercial whaling. With the support of the United Nations Environ- ment Programme and the World Wildlife Fund, “To Know Ourselves” The history of Canadian science and technology 1s still being so badly neglected by our universities that “Canadian students know virtually nothing about their scientific heritage,” says Thomas Symons, chair- man of the Commission on Canadian Studies. As a result, Symons said in a recent interview, “the technologists and scientists of the country are often underutilized and underestimated by their fellow citizens. A great deal of major scientific and techno- logical work is commissioned outside the country because there just isn’t the realization that we have the capability to do it here.” Symons is the author of the Commission’s 1975 report, To Know Ourselves, one chapter of which was devoted to science, technology, and Canadian studies. The Science Council of Canada is preparing an “issues paper” on science and education in the Canadian context. A recent Council-sponsored sem- inar concluded that there is, at present, “no formal requirement and little interest in putting Canadian content in science courses.” In anticipation of a possible full-scale study in the future, the issues paper will attempt to define and to focus on a number of the serious science-and-education problems highlighted in the Symons report. Symons said he is encouraged by the fact that halfa dozen universities have discussed with him the possi- bilities of either introducing or extending programs in the history of science. A major conference on the history of science was planned for Queen’s University in November 1978. He also said there is some evidence that the attitude of Canadian scientists has been shifting in favor of the idea of Canadian content in science teaching. This shift has, perhaps not sur- NEWS AND COMMENT 89 IUCN has embarked on an expanded program for marine conservation. Three of the objectives of this program are to launch an international system of cetacean sanctuaries, promote the conservation of the living resources of the southern ocean, and develop improved principles and procedures for the man- agement of whaling. IUCN welcomes the participation of the IWC in the operation of the Convention on International Trade in Endangered Species of Wild Fauna and Flora, and hopes that through this and other joint endeavors by concerned governments and international bodies the current problems of the conservation of whales will be satisfactorily resolved. From IUCN Bulletin. New Series 9 (7/8), July/ August 1978. prisingly, been most pronounced in the environmental sciences. In his report, Symons acknowledged that science is universal and international, but he also argued strongly that it nevertheless has a “national and cultural dimension . . . Science is a key ingredient in the cultural fabric of our society.” He suggests that there are “Canadian perspectives, Canadian applica- tions, Canadian motivations (and) Canadian ap- proaches to science that could be described accurately as Canadian studies.” His report in fact contains a litany of scientific subjects that could easily, and without rationaliza- tion, be classified within the ambit of Canadian studies — subjects which the Commission felt were being woefully neglected. For example, it urged that more attention be paid in science education to the’ implications of Canada’s weather and climate; to marine sciences concerned with the continental shelf and offshore conditions; to the geology and geo- graphy of Canada’s land masses; to Canadian mammals; and to Canada’s forests. The report particularly emphasized the lack of research in and about the North and the need for a more sustained and indigenous scientific effort there. It strongly recommended that a university of the North, or at least its nucleus be established immed- lately. The Commission also argued that non-science students in university must be given some training in science and the history of Canadian science. This is needed “to equip non-science students with a better knowledge and understanding of the important part played by science in our society . . . As matters now 90 THE CANADIAN FIELD-NATURALIST stand, universities in Canada are producing each year thousands of graduates whose knowledge and under- standing of the role played by science in our total culture is minimal to zero.” Conversely, science students must be allowed to take courses in the arts and humanities that will give them a basic knowledge of the society in which they live and the social institutions which will inevitably affect scientific research. Achieving this goal is obviously not an overnight kind of problem, but Symons said that a “modest beginning” has been made; a number of universities have indicated to him that they are taking a “fresh Vol. 93 look” at the problem. On the whole, in fact, Symons conveys a generally optimistic message. Although much remains to be done, he feels the Commission has been successful in prodding the scientific community to think about these issues. He also believes that the fact that he is not a scientist, and therefore could not be accused of self- pleading, helped in getting a fair number of the recommendations acted upon. From Agenda, August, 1978. (Bulletin of the Science Council of Canada) Second Annual International Wildlife Film Festival To encourage the production of high quality wildlife films, the University of Montana Student Chapter of the Wildlife Society sponsored the First Annual International Wildlife Film Festival last year. It was a great success and the chapter is again hosting this event. A panel of highly qualified film makers and biologists will judge films pertaining to wildlife and present awards and recognition to the winning films. The deadline for submission of applications and films is 7 February 1979. All entries must have a predominantly wildlife theme and have been pro- duced or released in calendar year 1978. Judging will be held on 10 and I! February and the winning films will be shown on 3 and 4 March at the University Center. Information, rules of eligibility, and application forms can be obtained by writing: Wildlife Film Festival, Wildlife Biology Program University of Montana Missoula, Montana 59812 (Telephone (406) 243-5272). Book Reviews ZOOLOGY A Guide to Bird Finding East of the Mississippi By Olin Sewall Pettingill, Jr., with illustrations by George Miksch Sutton. 2nd Edition: 1977. Oxford University Press, New York. xxvii + 689 pp. U.S. $15.95. When I first began watching birds almost 20 years ago, one of the first books I acquired was O. S. Pettingill’s A Guide to Bird Finding West of the Mississippi, published in 1953. This book and its eastern counterpart, published in 1951, have been constant and valued companions during my travels throughout the United States. Now a new and extensively revised version of the eastern guide has been issued, and a welcome event indeed this is for birdwatchers. Pettingill is presently revising the western guide, with a tentative publication date set for 1980. The format of the book is the same as in the first edition. Following a brief introduction, it is divided into 26 chapters, one for each American state lying entirely east of the Mississippi River (except that Connecticut and Rhode Island are dealt with to- gether, and the New York City region rates its own chapter). Canada is not covered, although several Canadian localities near the U.S. border (e.g., Point Pelee, Ontario and Grand Manan Island, New Brunswick) are discussed under the nearest state. Chapter length varies from 16 pages (Vermont) to 48 pages (Florida). Each chapter consists of two parts: a general account of the state’s birdlife, and a list of specific bird-finding localities, which are discussed under the nearest sizable city or town. Each locality account includes detailed directions for reaching the area and a partial list of birds to be expected, with emphasis on uncommon or spectacular species or noteworthy bird concentrations. Both the general state accounts and specific locality accounts, es- pecially the latter, have been revised substantially from the first edition. As Pettingill notes, the changes in 25 years have been profound, many bird-finding localities have been “developed,” and the distribution of the birds themselves has changed in many cases. Thus some localities have been dropped from the new edition, but many others have been added; in Michigan, for example, 29 localities are described (under 18 cities and towns) in the new edition, versus 16 in the old edition. Locality accounts are generally briefer in the new edition, but for most states, the net result is better coverage overall. The introductory sections in each state chapter are especially illuminating. Pettingill begins with a 91 description of the physiographic regions of the state and their associated vegetation and land-use types. Then come lists of characteristic breeding species for major habitats, with notes on those species that are restricted to certain parts of the state. Next follows a description of bird migration patterns (spatial and temporal) in the state, and finally a brief account of winter birdlife. Having thus “set the stage,” Pettingill then fills in the details in the individual locality accounts which follow. The book’s index is designed for easy location of information on a particular species. Suppose you want to find Bachman’s Sparrows in Alabama. A glance in the index under “Bachman’s Sparrow” tells you that the species is mentioned in Alabama locality accounts on pages 9 and 20; thus you need not wade through the entire Alabama chapter. For English names of birds, the book follows the A.B.A. Checklist, published by the American Birding Association in 1975. Many of these names have not yet been adopted by the American Ornithologists’ Union, and are not yet in general use; however, as the A.O.U. will likely adopt most of them soon, their use by Pettingill may help to prevent the book from becoming rapidly out-of-date. (The first edition includes some names now long obsolete, such as “Holboell’s Grebe” and “Duck Hawk.”) The route directions in most cases are necessarily brief. But I have not experienced the difficulties encountered by one reviewer (J. A. Tucker, 1977, Birding 9: 226-231) in following them. The birder would certainly be advised to have a good road map: handy, especially when trying to find localities in and near large cities. Given the frequent changes in road locations and numbering systems, however, directions for reaching some areas are bound to become out-of- date rather quickly. In looking closely at the chapter on Mississippi (the state I know best), I noted a couple of typographic errors in place-names, and several errors of fact, mostly concerning bird distribution; however, such errors are few and of minor importance. If I do have any serious criticism of the book, it is that, for some states at least, the author has relied too heavily on old information. (For instance, of 14 persons cited as contributing information for the Alabama and Mis- sissippi chapters, only two were not cited in the 1951 edition.) I am not suggesting that some of the areas included are no longer good for birds—only that local 92 THE CANADIAN FIELD-NATURALIST observers now know of better and more accessible areas than some of those included. But I hasten toadd that the selection of localities to include in a book like this is difficult and arbitrary, and no such selection will please all observers. | have now had a chance to use the new “Pettingill” in seven states, and I unreservedly recommend it to anyone looking for birds in the eastern U.S. Although there are now many good bird-finding guides for individual states, Pettingill’s is the only one covering the entire eastern part of the country, and | doubt that The Birds of New Brunswick By W. Austin Squires. 1976. New Brunswick Museum, Saint John, New Brunswick. Second Edition. v + 220 pp., illus. No price given. Ce livre est une nouvelle édition complétement revisée de celle de 1952 épuisée depuis plusieurs années. L’auteur y présente, en 221 pages, la situation ornithologique du Nouveau-Brunswick. L’ouvrage est agrémenté de dix photographies en couleur et autant en noir et blanc; le choix est judicieux et intéressant. J’al apprécié particulierement celles du Martinet ramoneur, de la Sitelle a poitrine rousse, du Grand Puffin et du Pica dos noir. La bibliographie est complete pour la période séparant les deux éditions; il est regrettable que l'on ne retrouve pas dans cette nouvelle édition, toutes les références bibliographi- ques utilisées dans la premiere. Deux index, un regroupant les noms anglais et scientifiques suivi d'un second pour les noms frangais, indiquent le caractere biculturel de la province. Grosso-modo, le livre peut étre divisé en deux sections: l'une traite de différents aspects de lornithologie dans la province et la seconde, constituant la majeure partie de l’ouvrage, est une liste annotée des oiseaux. Dans la premiére section, l’auteur, en plus d’in- troduire la liste annotée, fournit quelques courts chapitres sur des sujets aussi divers que Ihistoire, la biologie, l’écologie et la protection des oiseaux tout en limitant sa discussion a la province. Le chapitre sur Phistoire de lornithologie est lélément le plus original. L’auteur cerne trés bien le développement de lornithologie de 1613 a 1952; cependant, la période de 1952 a nos jours, période ot lornithologie-amateur contribue de fagon importante au développement des connaissances sur la distribution des oiseaux, est déficiente. Ainsi, 1] aurait été intéressant d’avoir des renseignements sur les groupes d’amateurs et de professionnels oeuvrant dans le domaine. A certains endroits de ce chapitre, 11 est difficile de distinguer entre les dates de visites des personnages et celles du Vol. 93 his approach will ever be significantly improved upon. Pettingill is to be congratulated on producing a well done and much needed revision of a book which is already considered a classic. My only lament is — when will someone write a bird-finding guide of similar comprehensiveness and authority for Canada? WAYNE C. WEBER Department of Biological Sciences, Mississippi State Uni- versity, Mississippi State, Mississippi 39762 systeme de références bibliographiques; une typo- graphie différente faciliterait la consultation. Lors de la révision de l’édition originale, on aurait dQ apporter une attention particuliére a la conversion des mesures dans le systéme international (SI). Une carte de la province représente cing régions topo-climatiques, les comtés provinciaux et quelques repéres géographi- ques importants. Un lecteur, non-résident du Nou- veau-Brunswick, apprécierait une carte avec une toponymie plus complete en relation avec la liste annotée. Ainsi, je n’ai pu localiser Pointe Lépreau, localité fréquemment mentionnée dans louvrage; de méme que l’ile Nantucket, homonyme de la célébre ile au large du Massachusetts. Les autres chapitres de cette section sont intéressants: l’introduction et les chapitres sur la migration, la collection et la ter- minologie de la liste annotée doivent étre consultés avant la lecture de la seconde partie de Pouvrage. Cette seconde section fournit une liste annotée de 345 espéces d’oiseaux. Dans la présentation de ce genre de listes, deux approches peuvent étre utilisées; Pune consiste a décrire l’espéce, son habitat et quelques traits de sa biologie, suivi de son statut provincial ou: régional. C’est approche que lon retrouve dans des ouvrages concernant d’autres provinces canadiennes (Salt, W.R. et J. R. Salt, 1976, The birds of Alberta; Tufts, R. W., 1961, The birds of Nova Scotia). L’autre présentation consiste a se limiter au statut de l’espéce, en le précisant par une sélection de mentions. C’est cette derni¢re approche qu utilise Squires et, a mon avis c’est celle quiatteint le mieux les objectifs que ce genre de livre devrait avoir. Dans la présentation de chaque espéce, les noms anglais, frangais et scientifiques sont fournis suivis dun bref résumé du statut et de l’'abondance selon des termes préalablement définis. Pour la majorité des espéces, une bréve description de lhabitat et une sélection de mentions typiques viennent préciser Vabondance. Deux autres paragraphes décrivent 1979 respectivement la distribution générale et provinciale. Enfin, les dates de ponte pour les nicheurs et la période de résidence de lespéce au Nouveau-Brunswick terminent le traitement de chaque espeéce. Ce genre de présentation agrémente la consultation de cette liste. L’auteur a fouillé en profondeur la littérature con- cernant les mentions pouvant se rapporter au ter- ritoire considéré. Cependant, le traitement d’espeéces marginales, quoique précis, souffre d’une déficience bibliographique. Ainsi, auteur aurait pu faire le rapprochement des invasions de Vanneau huppé en 1927 et 1966 avec celles des provinces voisines; les mentions de Fauvettes azurées pour le Québec sont basées sur des spécimens (Ouellet, H., 1967, The distribution of the Cerulean Warbler in the province of Quebec, Canada, Auk 84: 272-274) et non uniquement sur des observations visuelles. Le traite- ment des espéces occasionnelles et accidentelles donne BOOK REVIEWS 93 souvent prise a la critique dans ce genre d’ouvrage; cependant, l’auteur les aborde cas par cas avec un jugement critique. Cette seconde €dition est présentée sur papier de qualité avec une typographie claire et dégagée. Je n’y ai relevé que quelques erreurs typographiques; la seule importante est la référence bibliographique de Tufts (1961) concernant le spécimen d’Ibis blanc. Enfin, je consideére que cet ouvrage est indispensable a ceux qui sintéressent a la distribution des oiseaux dans le nord- est de Amérique du Nord et particuliérement au Canada. PIERRE LAPORTE Service canadien de la faune, Région du Québec, 2700 boulevard Laurier, CP 10100, Edifice “A”, Ste-Foy, Québec GIV 4H5 Mountain Monarchs: wild sheep and goats of the Himalaya By George B. Schaller. 1977. University of Chicago Press, Chicago. 425 pp., illus. US$25. According to the author, Mountain Monarchs is“a companion volume to The Deer and the Tiger in which I treated the ecology and behavior of some large mammals of peninsular India.” For Mountain Mon- archs, Schaller restricts himself, geographically, “to the Great Himalaya of Nepal and India, and to the ranges of Pakistan,” and to the subfamily Caprinae. “Rather than concentrating on one kind of animal in one locality for years, in the accepted manner of current ungulate studies, | moved from area to area, spending one month with one species, then the next one with a different species.” This approach, coupled with a fairly extensive literature review, allows Schaller to discuss a variety of caprine species from a comparative standpoint. It is this comparative treatment of the species that makes the book enjoyable reading. In reviewing the Caprinae, Schaller comments on taxonomy, distribution, physical attributes, popula- tion dynamics, herd dynamics, predators, aggressive behavior, courtship behavior, mother—young rela- tions, and social behavior. Chapters I found particu- larly interesting were those dealing with “Herd Dynamics,” “Aggressive Behavior,” and “Courtship.” “The basic unit in caprid society consists of a female with her offspring. All other social systems represent permutations of this basic theme.” “Most Caprinae herds are open, in that animals join and part, often when population density is great and not as often when it is sparse.” Neither of these comments alone is particularly profound; however, they are significant in that they provide confirmation of caprine behavior not previously studied. Chapter 9, dealing with “Aggressive Behavior,” is probably the strongest and most thorough chapter in the book. It represents a significant review of aggressive behavior and provides comparative data for species inhabiting the Himalayan region. Chapter 10, dealing with “Courtship Behavior,” is not as well organized, but is equally thorough. By comparison chapter I1, “Mother—Young Relations,” is far from thorough. It is probably the weakest section in Mountain Monarchs. The last chapter is somewhat of asummary chapter. ° “When one looks for correlations two environmental variables obtrude: (1) species live longer in a habitat with a predictable food supply, and (2) species dwelling on and around cliffs live longer than those on flat to rolling terrain.” Schaller’s work is intended to provide some basic data on the status of unknown big- game populations in Asia. It is his hope that conservation efforts to preserve these species will be soon forthcoming. “In writing about this vanishing mountain fauna I feel a special urgency. All too often in history the last of a species has disappeared into the belly of a hungry hunter, its epitaph a belch.” PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario POT ITO 94 THE CANADIAN FIELD-NATURALIST Vol. 93 Précis de Zoologie: Vertébrés. 3 — Reproduction, biologie, évolution et systématique. Oiseaux et mammifeéres Par P. P. Grassé. 1977. Masson, Paris. 2e édition. 395 pp. $35.00. Ce volume fait partiellement suite au Précis de Zoologie, Vertébrés, de la méme collection, qui réunissait Anatomie comparée, Biologie et Systéma- tique. Afin de présenter les faits avec plus d’ampleur et d’offrir un apercu plus fidéle de l’état actuel de nos connaissances, l’ouvrage a été divisé en trois volumes indépendants et sensiblement augmentes. C’est un livre de format pratique (16 x 24 x 2cm; | kg), trés bien relié et d’une solidité qui devrait résister a l'usage le plus soutenu. En feuilletant le livre, j'ai été favorablement impressionné par la qualité du papier glacé et la clarté du texte imprimé. II y a plusieurs dessins a l’encre de chine, quelques photographies noir et blanc un peu floues ainsi qu'un croquis en couleur qui n’apporte toutefois rien a la compreé- hension du texte. Le présent livre traite de la reproduction, de Pévolution, des particularités physiologiques, de léthologie et de la systématique des classes des oiseaux et des mammiferes. La partie qui concerne les oiseaux occupe le tiers du livre, celle qui traite des mammifeéres occupe le reste. On y décrit en détail les particularités anatomiques de ces animaux. Par exemple, l’auteur fait une excellente description des adaptations morphologiques qui permettent aux oiseaux de voler. Une section moins importante est consacrée a la systématique. On y présente les Ways of the Six-Footed By Anna Botsford Comstock. 1977. Cornell University Press. Ithaca. x11 + 152 pp. U.S. $5.95. This reissue of the 1903 edition is subtitled on the dust jacket: “A delightful introduction to the world of insects” — so it is! The author’s poetic yet intricate description of the lives of a few common insect species provides the reader with new insight into the lives of insects. The ten chapters cover such subjects as sound communication, the Maple-leaf Cutter, mimicry, socialism, the Mud Dauber and Mason Wasp, the Carpenter Bee, the Basswood Leaf-roller, the Lace- wing, the Seine Maker (Hydropsyche), and the Seventeen Year Cicada. There are 47 illustrations, many of them the author's, for, as stated by Edward H. Smith in the new foreword, “she became one of the foremost wood engravers of her time... .” Not only did she illustrate books for her husband, caractéristiques anatomiques ainsi que quelques aspects des moeurs des principaux genres. Malheureusement, les descriptions sont peu détaillées et le lecteur ne peut pas tellement simaginer lallure des animaux décrits. Enfin, une section est consacrée au comportement. Cette partie de louvrage est tres décevante; le choix des sujets a été fait de fagon arbitraire et chacun des aspects comportementaux s€lectionnés est traité sommairement. Le volume contient aussi quelques hors-textes: un avant-propos, une table des mati€res et un index alphabétique des sujets. Cet index est trés bien fait et permet de trouver rapidement les informations désirées. L’auteur n’a cependant pas incorporé de bibliographie a son ouvrage, ce qui constitue une lacune importante puisque ce traité ne couvre que superficiellement la majorité des sujets. En conclusion, malgré mes remarques critiques, je crois quils’agit tout de méme d’un ouvrage didactique de qualité, du moins ence quiconcerne l’enseignement de la morphologie et de la systématique des oiseaux et des mammiféres. Quant aux moeurs de ces animaux, le lecteur y trouvera une foule dinformations qui Yaideront a s’en faire une bonne idée. JEAN Luc DESGRANGES Service canadien de la faune, Région du Québec, 2700 boulevard Laurier, C.P. 10100, Ste-Foy, Québec G1V 4H5 John Henry Comstock, but co-authored several, as well as publishing some of her own. Her purpose in writing the stories in the present volume was to illustrate the truth that “wherever there is life there are problems confronting it; and that the way of solving these problems has been the way to success in the evolution of a species.” The author has dedicated her book “to all those who have been my good comrades and fellow loiterers in nature’s byways” and it is to these people, or, in other words, all naturalists, that | would recommend this book. WILLIAM B. PRESTON Manitoba Museum of Man and Nature, 190 Rupert Avenue, Winnipeg, Manitoba R3B 0N2 1979 BOOK REVIEWS 95 Birds of Southeastern Michigan and Southwestern Ontario By Alice H. Kelley. 1978. Cranbrook Institute of Science, Bloomfield Hills, Michigan. vi + 99 pp., illus. Paperback US $2.95. Alice Kelley’s book is the latest regionally specific reference on birds in Canada and adjacent United States. Records of birds are taken from the area bounded by Kettle Point on the southeast end of Lake Huron, Rondeau Provincial Park, Toledo, and Flint. The author includes a brief four-page description of the topography, drainage, soils, status of forest and marshland, and areas of particular ecological interest. Although short on illustrations, consisting solely of a map of the region surveyed, the book is long on detailed bird records, providing data on 338 species for the period 1945 to 1974. Each species is listed with abundance rating, residency, record arrival and departure dates, average arrival and departure dates, exceptional sightings and abundance records, breed- ing status, and locations and sites historically favored to viewing the birds. One of the main objectives of Alice Kelley in writing the book is to document the historic changes in bird occurrence in order to monitor effects of rapid environmental alterations. Therefore, the book is written primarily for the seasoned birder/ ornitholo- gist who has advanced beyond basic identification to the point of determining geographic or ecological significance of particular bird sightings at specific locations. Watching Sea Birds By Richard Perry. 1975. Taplinger Publishing Company, New York (Canadian distributer Burns and MacEachern, Toronto). 230 pp., illus. $13.25. There were few books written on the natural history of seabirds of the British Isles in the 1940s that were better, or contributed more to the subject area, than Richard Perry’s Lundy, Isle of Puffins (1940) and Shetland Sanctuary (1948). The information con- tained in these publications was extensive and important at the time; included were precise docu- mentations of the breeding activities of colonial- nesting seabirds at two major sites and numerous unique interpretations of their biological significance. Thus, the republication of these long out-of-print classics under the title Watching Sea Birds is a most welcome event and will allow general readers and students of seabird biology an opportunity to gain easy access to otherwise scarce sources of very useful material. Two disappointing features of the book are these: (1) the I5- x 23-cm format which is unsuitable for most pocket sizes and therefore relatively incon- venient to carry in the field as a complement to most field identification guides; (2) as the author suggests, the general qualitative statements regarding abundance and status have not been rigidly defined in quantitative terms. In 1954, the New York State Federation of Bird Clubs outlined numerical standards for quali- tative terminology, and these could have been incorporated into the report as extensive quanti- tative data were available. This arbitrary system of abundance rating would invalidate most comparisons with those reported in other regional summary accounts using the established stan- dards (e.g., History of the birds of Kingston, Ontario by Helen Quilliam). Disregarding these minor detractions, Birds of southeastern Michigan and southwestern Ontario is a first-rate, comprehensive, regional documentation of birds recommended for those wishing to apply greater significance to their bird sightings and possibly contribute meaningful records for that region. BRENT BEAM Beak Consultants Limited, 6870 Goreway Drive, Missis- sauga, Ontario L4V I1L9 The volume under review is a straightforward reprint of the earlier works containing the identical | text with only minor editorial changes, usually most evident by the removal of out-dated or technically unacceptable sentences and paragraphs. The first half presents the main chapters from the Lundy book on spring occupancy, Atlantic Puffins, Black-legged Kittiwakes, Razorbills, and Common Murres, while the remainder comprises those from the Shetland book on Great Skuas, Arctic Skuas, Northern Gannets, and northern Common Murres. These accounts are the results of intensive studies conducted by the author from 20 March to 7 August 1939 on Lundy Isle in southwestern England and on the Isle of Noss, in the Shetland Islands, from 2 April to 18 September 1945. They provide a detailed and accurate description of the breeding biology of the various species, with particular emphasis placed on behavior, and most of the material covered is as significant and 96 THE CANADIAN FIELD-NATURALIST informative today as it was when first produced. Together, they present a vivid and authoritative account of a community of seabirds made by anastute field observer and go far in capturing and translating the very special atmosphere associated with the study of colonies of seabirds and their islands. There are, however, a few things wrong with this new version; for the most part these are errors in editorial judgment. The flagrant attempt by the author and/or publisher to disguise the reprint as a “new” book by not pointing out the existence of the original works or even the years in which the studies themselves were made (nowhere are the years given) was unwise and totally unwarranted. Although this decision was undoubtedly based on marketing con- siderations, it is difficult to understand how it came about; if anything, the potential for sales should have been enhanced by announcing the volume as a reprint of the earlier books, especially since copies of the originals are difficult and costly to obtain. The omission of all the photographs used to illustrate the earlier books is unfortunate, especially those in the Lundy book, which were carefully integrated with the text to show precise features of certain breeding activities. The substitute black-and-white ink sketches of birds in flight and on land by Richard Richardson do not fill this gap and are, in general, uninspiring and often inaccurate, adding very little to the book. Also, the decision not to reprint the summaries of the breeding schedules for individual species or the appendices (these contain statistical data on nesting and timing, etc.) is hard to accept. A judicious selection for inclusion should have been possible and Bird Hazards to Aircraft: problem and prevention By Hans Blokpoel. 1977. Clarke Irwin, Toronto, in associ- ation with the Canadian Wildlife Service, Evironment Canada and the Publishing Centre, Department of Supply and Services, Ottawa. 236 pp., illus. $9.50. That both birds and aircraft may helplessly tumble to earth after colliding in mid-air has been known since 1912 when the first such fatality occurred. Since that time there have been at least 77 reported bird- aircraft crashes with over 100 lives lost. The total financial loss resulting from these crashes is in the order of $100,000,000. In spite of these staggering financial and personal losses, and the certainty of future bird-aircraft crashes to feed the fears of the world’s millions of flight phobic individuals (30% of the flying population is flight phobic for one reason or another), there has never been a comprehensive guide or manual on the Vol. 93 would have ensured a still wider audience by increasing the value of the new version to both the seabird specialist and serious naturalist. As it now stands, these readers must of necessity go back to the originals for this information. The major shortcoming, however, is the absence of an epilogue by the author to relay his assessment and personal feelings to the reader concerning the signifi- cance and implications of the large reductions in alcid populations that have taken place at his study colonies over the last 30 years. For example, the total number of puffins, razorbills, and murres at Lundy has decreased from about 33,000 breeding pairs in 1939 (from author’s census) to less than 2300 pairs at the present time, with the puffin population virtually extinct (about 40 pairs in 1969-1970). Perry has unfortunately lost a great opportunity to add to the biological record of the populations that he first identified and so carefully appraised. This oversight can only be regretted by those of us concerned about the future welfare of this most threatened group of specialized seabirds and of the richness and diversity of marine ecosystems. Overall, this new book will serve as a reliable and accurate source of the descriptions of seabirds and their breeding activities, and it will be found to be informative and useful to the general reader. D. N. NETTLESHIP Seabird Research Unit, Canadian Wildlife Service, Bedford Institute of Oceanography, P.O. Box 1006, Dartmouth, Nova Scotia B2Y 4A2 of bird/aircraft collision problems of bird hazards to aircraft. Recognizing this fact and drawing on first-hand experience gathered in more than 10 years of scientific study on two continents, Hans Blokpoel presents us with a volume designed to “. . . deal with all aspects of the problem.” This may sound like an ambitious goal but readers will find that Blokpoel’s objective is met very methodically in Bird Hazards to Aircraft by (1) examining the biology of birds as it applies to the problem; (2) examining bird strike statistics to determine the characteristics of the problem; (3) examining engine designs and airports and their surroundings to see how present and future problems may be eliminated or alleviated; and (4) producing references and organizations for further consider- ation. Bird Hazards to Aircraft is written and intended for those directly concerned with the IQ7Y problem, regardless of their speciality. It is very readable. Bird strikes generally occur to the engine of an airplane, at night (based on strike rate), during the spring or autumn migration, at heights under 3000 feet, during take off or landing (1.e., near airports), at aircraft speeds of 80-160 knots and, as in the 1912 crash, the species most often involved is a gull. Birds, however, have struck nearly all leading edges of aircraft, at any time of the day and year, at heights of up to 23000 feet, during all flight phases, at speeds of up to 260 knots, and strikes have involved birds as small as a Ruby-crowned Kinglet or as large as a Sandhill Crane. From here Blokpoel discusses ways of bird- proofing aircraft and on-board means of diverting them from the path of aircraft. Another approach to alleviating the bird hazard to aircraft is to reduce the attraction provided by airports. The most common attractants at airports are food and a safe place to nest or rest. Proper habitat manipulation can reduce both of these. Blokpoel says, “The airport should be made as much as possible into a monoculture, thus supporting only a few bird species against which effective scaring techniques can be developed.” Away from airports the main source of bird problems are from local and migratory flights. It is often possible to shift local bird flights by removing the birds’ food sources or altering their roosting sites. Bears — their biology and management Edited by M.R. Pelton, J. W. Lentfer, and G. E. Folk. 1976. IUCN Publication, New Series Number 40. Mor- ges, Switzerland. 467 pp. US $12. Bears — their biology and management is a selection of papers from the third International Conference on Bear Research and Management. The text contains 45 papers with the papers grouped into 5 sections. The 5 sections cover bear behavior, bears in national parks, management of bears and techniques of management, status of bears, and biology of bears. The authors of the presented papers are international but approach common themes and problems. The eight papers included in section | are all the result of observational studies on bear behavior. Bacon and Burghardt’s paper, “Learning and Color Discrimination in the American Black Bear,” points out that “black bears appear to use their eyesight during ingestive behaviors much more than pre- viously supposed.” Luque and Stokes’ paper, “Fishing Behavior of Alaska Brown Bears,” examines “brown BOOK REVIEWS 97 Migratory movements of birds are much more fixed and Blokpoel discusses bird forecasting systems that have been devised to cope with large numbers of migrating geese and swans. The concluding chapter and bibliography are excellent reference material. Listed are committees and individual names and addresses from 33 countries where the bird strike problem is being studied. The bibliography contains over 400 references and cita- tions to other bird hazard/radar ornithology bibli- ographies. The shortcomings of the book are few and only ofa minor nature. I found that Blokpoel’s occasional anthropomorphic reference made the book more readable; others may not agree. Some pre-1973 species spellings are still employed (e.g., Widgeon). The occasional technique is described without a reference given (e.g., p. 103). The Appendix, Table I- 3, on ground speeds of birds could obviously have been more complete (no gull species are given), especially where the original covered several journal pages. Those few shortcomings, however, are ex- tremely minor and this excellent book should appeal to (and be required reading for) all those, “. . . directly ” involved in or concerned with bird strikes .. .”. D. V. WESELOH Canadian Wildlife Service, Canada Centre for Inland Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6. bear fishing behavior, its development, and _ its relation to environmental and social factors.” Other papers of this section deal with social behavior, — breeding behavior, and threat behavior. Section 2 contains seven papers, all concerned with bears in national parks. Papers of this section discuss people--bear conflicts, movement patterns of bears, territoriality, and bears and garbage. Beeman and Pelton’s paper, “Homing of Black Bears in the Great Smoky Mountains National Park,” demonstrates that “the probability of a bear becoming a nuisance other than in its established home range is quite low.” Although the papers of section 2 have strong management overtones, it is section 3 which contains papers classed as those dealing with management. This section contains 10 papers all of which are interesting. Two of the papers report on bear population control. Greer’s paper, “Managing Mon- tana’s Grizzlies for the Grizzlies,’ suggests that hunting is not a big factor affecting grizzly popula- 98 THE CANADIAN FIELD-NATURALIST tions, but were hunting eliminated the grizzly popu- lation could become a problem. Kemp’s paper, “The Dynamics and Regulation of Black Bear Ursus americanus Populations in Northern Alberta,” dem- onstrates that the adult males were affecting “a regulatory influence on the bear population.” Section 4, the Status of bears, contains 10 papers, mainly Eurasian in content and concerned with declining brown bear populations. Typical of papers contained in this section is Elgmork’s paper, “A Remnant Brown Bear Population in Southern Nor- way and Problems of its Conservation.” This paper points to one of the problems. “The most reasonable explanation for the relatively rapid decline starting in the second half of the 1950s is therefore not hunting and insufficient reproduction but the deterioration of the habitat caused by increasing human activity.” Vol. 93 The biology of bears is the topic of the 10 papers contained in section 5. Some of these papers are strongly physiological in slant. Rogers and Rogers’ paper, “Parasites of Bears: A Review,” I found to be the most interesting paper of this section. On a general level, Bears — Their biology and management contains a wealth of information. Those persons familiar with the 28 papers contained in the 1970 proceedings (IUCN Publication New Series Number 23) will note that these 1974 proceedings appear as a continuation of the earlier proceedings. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario POT 1TO The Earthworms (Lumbricidae and Sparganophilidae) of Ontario By John W. Reynolds. 1977. Life Sciences Miscellaneous Publications. Royal Ontario Museum, Toronto. 1x + 141 pp. $8.00. It is with genuine pleasure that I welcome this guide to some of the lower forms of life, and at a reasonable price. It is regrettable that the title does not include all of Canada; however, the book does purport to be useful in southeastern Canada and the adjacent United States. I suspect that with care this publica- tion would be useful in western Canada as well, forall but one of the nineteen species covered are intro- duced species. Further, fourteen of these species have been recorded from British Columbia. John Reynolds has authored numerous technical and popular articles on earthworms in a number of publications. The foreword (from which I quote later) is by lanR. Ball, Assistant Curator of Invertebrate Zoology, Royal Ontario Museum. Included in the book are an introduction to the general biology of earthworms, methods of study, an illustrated glossary, a key to species, species accounts, and a discussion of distribu- tion and ecology of earthworms. The appendix includes a provincial description. The book is well documented, as indicated by fifteen pages of refer- ences. I have followed the author's instructions on the preservation of earthworms and have found the key workable. Several pages are devoted to each species account, which is illustrated with line drawings and a map of the distribution in Ontario. For each species a synonymy, diagnosis, biology, range (in the world), North American distribution, and Ontario distribu- tion are given. Common names are given in Frenchas well as English. I do not hesitate to recommend this book to anyone with an interest in the lower forms of life, especially to he who “finds his happiness unearthing worms.” WILLIAM B. PRESTON Manitoba Museum of Man and Nature, 190 Rupert Avenue, Winnipeg, Manitoba R3B ON2. The Passenger Pigeon — its natural history and extinction By A. W. Schorger. 1955 (reprinted 1973). University of Oklahoma Press, Norman (Canadian distributor Burns and MacEachern, Toronto). xii + 424 pp. $7.50. The University of Oklahoma Press has done the biological community a great service in reproducing this book. The Passenger Pigeon was an incredibly abundant species, perhaps the most abundant bird that ever lived. Its extinction is the classic disaster story of ecological and conservation literature. Any species as abundant as the Passenger Pigeon (Ectopistes migratorius) is bound to invoke all sorts of legends. The decline of the Passenger Pigeon was close 1979 to the point of no return when scientific investiga- tions of bird behavior and ecology were in their infancy, and most ornithological journals were just beginning or not yet in existence. Thus, most information on the species was a jumble of fact and fiction scattered through old rare books, newspapers, and obscure journals. Schorger did an excellent job of compiling all this scattered data, and then sifting fact from myth. Myths are not simply dismissed, but usually explained as misinterpretation of facts: For example a habit of alighting on the ends of limbs of oak and beech to eat loose nuts that had not yet fallen caused the birds to winnow their wings back and forth to maintain balance. This resulted in a legend that they detached the nuts with blows of their wings. The first chapter is appropriately devoted to early accounts. Seekers of Canadian content will be delighted to learn that the first known record of this bird comes from Prince Edward Island where Cartier saw an “infinite number” on | July 1574. The fifteen remaining chapters document behavior, food, use by humans, anatomy, distribution, and extinction. The last of these chapters is a critique of various drawings and paintings of the birds, based primarily on anatomical facts. In general, the book is well written, easy to read, and remarkably free of technical errors. The caption and legend for Figure | lack any reference to the large nesting area shown on the map. The lengthy original quotes will be somewhat tedious to some readers, but a delight to others. Those of us interested in the history of science will find the 1685 quote on the physiology of “pigeon milk” in Columba livia fascin- ating, but it does seem a little out of place in this book. That the Passenger Pigeon existed in enormous The Bird Finder’s 3-Year Note Book By Paul S. Eriksson. 1976. Paul S. Eriksson Inc., New York. pages unnumbered. US $7.95. This 6- x 9-inch spiral-bound notebook is billed as a natural companion to a field guide for anyone wanting to maintain a permanent record of observa- tions. Each day of the year is given one page. On this page, space is allotted for permanently recording observations for three years. The book provides an organized and orderly way to record year-to-year comparisons of migratory dates, behavioral charac- teristics, unusual observations, identification details, and other important notes that often become mis- placed after the return from a field trip. Since the maintenance of field notes can be extremely im- BOOK REVIEWS 99 numbers cannot be doubted, but the author’s unhesi- tating acceptance that this was the most abundant bird ever known ignores similar estimates for the Red- billed Dioch ( Quelea quelea) of Africa. The use of old and obscure sources with old place names resulted in some errors in distribution, at least in the prairie provinces. These have been corrected by C.S. Houston (Blue Jay (1972) 30: 221-222). One of the most useful features of the book for the serious biologist is the detailed documentation of sources. Some of these lack full publishing details. To these minor criticisms, | would add only that the book is now over 20 years old. Serious researchers will need to consult recent literature for an update of information. Three more recent Canadian references that come to mind are those by Steele (Canadian Field-Naturalist (1967) 81: 172-174) for Ontario, Houston (Blue Jay (1972) 30: 77-83) for Manitoba and Saskatchewan, and Smith and Kidd (Canadian Field- Naturalist (1971) 85: 259) for Alberta. Schorger’s book is an excellent compendium of information that would not have been available without his decades of hard work. I recommend it highly to all ornithologists. To conservationists it will serve as an excellent example of the susceptibility of nature to human thoughtlessness. The fact that there exists only one known specimen of a nestling of what may well have been the most abundant bird ever known is shocking testimony to the lack of foresight by man in regard to this incredible bird. MARTIN K. MCNICHOLL Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta T2E 6M7 portant in future years and is often ignored by even the keenest field observer, any item that will encourage an observer to record his many worthwhile observations is important. Remsen (1977, On taking field notes, American Birds 31: 946-953) recently presented a thorough and enlightening discussion on why one should take field notes, how they should be written, and how they can be used. At the bottom of each page is the inclusion of a quotation by a famous writer, an observation by Eriksson, or space for additional notes. Among the writers quoted are Peterson, Burroughs, Sutton, and Pettingill. A Peterson quote summarizes the im- portance of recording detailed field observations, “If exact numbers of each species are kept, a year-to-year 100 comparison gives a hint of increases or declines. Redstart — common does not mean much, but redstart — 58 does.” A special feature at the back of the book isa 10-page life list of more than 700 species listed alphabetically by their common and scientific names with space provided after each name for writing. Although this feature is an excellent idea, the list contains several errors that may confuse the user. Names are based upon the latest update (32nd supplement) of the A.O.U. Check-list, but usage is not consistent. Among the confusing names discovered in this section are Trudeau’s Tern, a South American species (type specimen taken by Audubon in New Jersey); White Ibis, also listed as Spanish Curlew; Common Golden- eye, also listed as Whistling Duck; scientific name for BOTANY THE CANADIAN FIELD-NATURALIST Vol. 93 Snow Goose given as Chen caerulescens hyperborea; Dipper listed under Ouzel, and Gray Hawk also listed as Mexican Goshawk. The treatment of the junco group is also very confusing. Misspellings spotted in the life list or diary sections are Gadwal for Gadwall, Anus for Anas, Widgeon for Wigeon, and albifrans for albifrons. In summary, this type of diary may benefit many of us who find it difficult “to start from scratch” and to continue the endeavor of maintaining an organized record of field observations after a time. NOEL J. CUTRIGHT Wisconsin Electric Power Company, Milwaukee, Wis- consin 5320] The Rare Vascular Plants of Ontario/Les Plantes Vasculaires Rares de Ontario By George W. Argus and David J. White. 1977. Syllogeus Series Number 14. National Museum of Natural Sciences, Botanv Division, Ottawa. 64 pp. English + 67 pp. French. Paper free. The systematic conservation of our native flora requires current, accurate information concerning the distribution, abundance, and autecology of each taxon. Accordingly, taxa that are found to be rare and/or threatened can receive priority for conserva- tion. Following this strategy, the Systematics and Phytogeography Section of the Canadian Botanical Association established a Rare and Endangered Plants Committee in 1973 to enumerate the rare and potentially endangered native plants in Canada. In concert with these efforts, the Botany Division of the National Museum of Natural Sciences initiated a project to compile a data base on rare and endangered Canadian plants. In the authors words, The Rare Vascular Plants of Ontario is “the first in a proposed series dealing with the rare vascular plants of the provinces and territories of Canada.” (At the time of review similar lists in the Syllogeus Series were also available for Alberta and Nova Scotia.) This publication presents a neatly abbreviated synthesis of information about the rare, native vascular plants of Ontario drawn from many sources: plant distribution maps, literature, IBP/CT check- sheets, the examination of collections in the herbaria at the National Museum of Natural Sciences (CAN) and the Department of Agriculture (DAO), and the considerable input of botanists and naturalists knowl- edgable about the flora of Ontario, who were requested to comment on preliminary lists compiled from the preceding sources. Specifically, the contents of this publication include an introductory section, an annotated list of rare plants, a bibliography, and three appendices. The introductory section briefly explains the history of the project, and gives definitions of rare, endangered, and threatened plants, the organization of the list, future plans for the continuation of this work, potential application of the list for the conservation of the Ontario flora, and notes about the distribution patterns of rare plants in Ontario. The considerable acknowledgments support the author's tenet “that no one person or group has sufficient knowledge of the flora of Canada to make a project of this magnitude a success without the support of all botanists and naturalists.” The present work is a tribute to the authors’ ability to marshal these human resources. Although the authors supply definitions for ‘rare, ‘endangered,’ and ‘threatened, the present work, as suggested by the title, is clearly an enumeration of native vascular plants considered by the authors to be rare. ‘Plant’ is defined “to indicate collectively all the individuals referable to a plant name” including species, subspecies, and varieties. A ‘rare plant’ is defined as one that is restricted to “a small geographic area,” or one that occurs “sparsely over a wide area.” The authors have been cautious to avoid premature 1979 designations of ‘threatened’ or ‘endangered’ in ad- vance of objective information to defend such a designation, noting that for some plants “even information on rarity was scarce.” Pages 11-49 present an annotated list of 611 plants arranged in alphabetical order by scientific name. Where information exists brief notes are supplied for each plant including “synonyms, references docu- menting its status as a rare or endangered plant, its distribution in Canada and Alaska, its distribution in Ontario, references to published or unpublished distribution maps and comments on its status in Canada, special protection and other comments.” The regimented format of this list suggests a computer application for processing future lists. The reader is sure to be alarmed by the length of this list, which represents approximately 19 percent of the entire, native vascular flora of Canada. For many of these plants the center of distribution in Canada, or the entire Canadian distribution is restricted to southern Ontario where the pressures for habitat destruction are severe. For example, in a Canadian context, many Carolinian plants can be considered ‘endemic’ to Ontario. It may be appealing to suggest that the massive landscape conversion in southern Ontario since white settlement is a major reason for this high incidence of rare plants, but the essential historical documentation to advance this postulate is very inadequate. Pages 50-63, inclusive, contain a bibliography in excess of 300 references and three appendices. Appendix | lists 36 plants reported to occur in Ontario whose status requires verification. Appendix II lists 21 alien or doubtfully native plants. Appendix III presents the list of rare plants in taxonomic order following Gray’s Manual of Botany, edition 8. Fungi: Delight of Curiosity By Harold J. Brodie. 1978. University of Toronto Press, Toronto. 131 pp., illus. $10.00. Here is a book of many charms, but how does one classify it? It could scarcely be further removed from the textbook category, although written by a mycol- Ogist. It is certainly not science fiction, though the rank beginner or the skeptic might be excused for thinking so. The author states his aim frankly, to share his personal feelings of curiosity, excitement and delight when confronted with the complexity and perfection of devices in operation among the fungi as they compete for survival and propagation of their kind. Written not for students of the fungi, nor even for those with some background in biology, the book is addressed to the mycologically uninformed public. BooK REVIEWS 101 The publication measures approximately 21.7 x 28.1 cm (8 1/2” X 11”) and is neatly bound in soft cover with a stiff spine. The text is clean, free of typo- graphical error, and organized into two columns to facilitate reading. Although the publication includes both English and French translations, the latter appears unjustified except for the Introduction on pages I-11. The bibliography and appendices are identical duplicates in the two translations while only very elementary English or French translation skills are necessary to comprehend either translation of the annotated rare plant list. Although traditional views concerning the protec- tion of endangered species have focused on wildlife in a quite narrow sense, this publication is evidence of a broadening concern for the stewardship of our biological heritage. As the initial statement in a projected series, The Rare Vascular Plants of Ontario promises great potential to monitor and to update periodically a red list of the Canadian flora. In addition to being an incentive for botanists and naturalists to continue exploratory work towards a more comprehensive understanding of endangered plant life, this report serves as a catalyst to further the appreciation and the protection of our native flora. This significant work deserves the serious attention of all government agencies, private sector organizations and individuals concerned with biological conserva- tion and landuse planning in Ontario. T. J. BEECHEY Provincial Parks Branch, Ontario Ministry of Natural Resources, Whitney Block, Queen’s Park, Toronto, Ontario M7A IW3 It avoids technical jargon, and describes phenomena of the fungus world in simple terms intelligible to non- biologists. Between Preamble and Postcript are eleven un- related chapters, each complete in itself, describing strange fungi or curious features of fungus life. The chapters are bound together very loosely by the fable of the blind men and the elephant, illustrating the author’s contention that most of us, limited by viewpoint and individual powers of perception, fail to perceive and appreciate many aspects of the natural world around us. Some indication of the book’s content may be gathered from reference to the topic of chapter one: Gunnery in the Fungus World. The common mold Pilobolus, a marvel of biological ingenuity, is an 102 example in nature of an automatically controlled missile. By mysterious manipulations of its chemistry, and making use of well-known principles of physics, the fungus has perfected a complex and efficiently designed apparatus for hurling its projectile (a pellet of spores) several thousand times its own diameter, actually directing the aim of the missile with over 80% accuracy. Equally intriguing is the subject matter of the other ten chapters. Not the least of the delights of this book is the author’s facility with English. With a few strokes of the pen, he evokes lively mental images of the unseen fungus world. It is a rare scientist, too, that can align his thinking with that of ordinary folk, to describe a fungus as “suggesting a broad vase filled with bananas,” or again, to depict the fungus-gardening ant carrying big leaf fragments aloft “like a man carrying a sheet of plywood.” The author’s first THE CANADIAN FIELD-NATURALIST Vol. 93 finding of the Coral Fungus is described in a most beautiful piece of writing, revealing a fine-tuned sensitivity to sound, scent, color, and form. Through- out the book, an unexpected sprinkling of phil- osophy and folklore, wisdom and humor, make for enjoyable reading quite apart from the scientific information imparted. The 22 black-and-white photographs of fungi have magnifications ranging from 1/2 x to 10000 x. Also included are a simple glossary, a selected reference to books on related topics, and an index. It is difficult to pigeon-hole this work, but there is no difficulty in recommending it as a delightful and unusual addition to one’s natural history library. SHEILA C. THOMSON 2066 Rideau River Drive, Ottawa, Ontario KIS 1V3 Pollen Flora of Argentina: modern spore and pollen types of Pteridophyta, Gymnospermae, and Angiospermae By Vera Markgraf and Hector L. D’Antoni. 1978. Uni- versity of Arizona Press, Tuscon, Arizona. 208 pp. US $9.50. This publication provides photomicrographs, mor- phologic descriptions, and keys for 374 pollen types. It divided keys into four plant geographic regions to facilitate palynologic work in other similar areas. This text also includes indexes to plant families, plant species, and common names. This much needed text will be very useful to people working in palynology and other disciplines in botany, not only in South America, but in North America, Europe, Africa, Asia and Australia. I. JOHN BASSETT Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 Floristics and Environmental Planning in Western New York and Adjacent Ontario — distribution of legally protected plants and plant sanctuaries By Richard H. Zander. 1976. Occasional Paper Number 1, Buffalo Society of National Sciences, Buffalo, New York. 47 pp., illus. Paper US$2.50 + 35¢ handling. The paper deals with a number of plants that have been given some protection by New York State law. All protected species, except for ferns, orchids, and club mosses are mapped on an individual basis. Many species are discussed in detail with emphasis on geographic distribution and preferred habitat. This publication was obviously written by a botanist who is concerned about the long-term protection of rare plants and their habitats. This reviewer and most readers of The Canadian Field- Naturalist would agree with this idea. The paper’s strength lies in its botanical com- ponents. It is weak inits planning elements. The title is too broad for the material covered in the small publication. In reality the paper deals with rare plants and areas worthy of protection rather than floristics and environmental planning. Zander mentions that Section 193.3 of the New York State Penal Code, “protects certain nature wildflowers, ferns and shrubs in New York State and provides for a small fine for picking, transplanting or removing any of them. However, it affords no protection to these plants from the owners of the land upon which they grow.” Later he mentions that “the recently enacted state Freshwater Wetlands Law... 1979 would give some protection from encroachment to wetlands over 12.4 acres in extent.” These statements tantalize. It would have been worthwhile if the relevant sections of the law had been discussed in detail. Those who are working towards legislature changes to protect natural areas, elsewhere, would have appreciated such information. The document is liberally illustrated with distribu- tion maps. Each has a typical dot format which indicates the presence or absence of a species in a minor political district (townships, town, or city). It would have been better if each station of the uncommon plant had been indicated by a small dot at the appropriate site location, as was done in Fox and Soper’s well read papers on Canada’s Carolinean flora. Unfortunately, several of the maps have no legend and no reference back to the text page where some explanation does occur. Now a few small points. Is Helleborine, Epipactis helleborine, actually so rare as to bea protected plant in New York State? In my experience in southern Ontario it is ubiquitous. In the Regional Munici- pality of Niagara there are important nature reserves that do not appear on the list. The Short Hills Nature Sanctuary, near Forthill, is owned and managed by the Hamilton Naturalists’ Club. The large, new Short Hills Provincial Park has extensive blocks of impor- tant natural areas most of which are protected in the Master Plan. Ideally, what would a planner or other decision- maker need in order to fulfill the expressed desires of this book? Let’s list a few: Atlas of Airborne Pollen Grains and Spores in By Siwert Nilsson, Joseph Praglowski, and Lennart Nilsson. 1977. Ljungforetagen, Orebro, Sweden. 159 pp. US $37.50: Sw Crs 170. This is an excellent introductory text on airborne pollen grains and spores of northern Europe. The atlas contains descriptions and illustrations of more than 70 species utilizing light, scanning and trans- mission electron microscopy. The descriptions are accompanied by distribution maps and flowering period for each species. Many of the species described also occur in Canada and the northern United States. Freshly collected pollen grains and spores were generally used. In some cases pollen was sampled from dried herbarium specimens located in the Botanical Section of the Swedish Museum of Natural History. The pollen grains and spores were embedded in fuschin-stained glycerol jelly and the cover slips on BOOK REVIEWS 103 (1) A law allowing him to practice environmental planning in his area of jurisdiction. (2) The written judgment of reputable botanists on which plants are worthy of protection and why they would be protected. (3) Detailed maps showing the important botanical areas that need to be protected. (4) Published data that supports the boundary delineations of the areas. (5) Political (public) support when a land-use con- flict threatens a delineated natural area. This publication attempts to fulfill several of these points, most specifically 2 and 3. Each of these efforts listed above should be in place at a time well before any on-the-ground activity takes place. For example, most subdivisions in Ontario usually take 5 years from when the first approach is made toa municipality until construction starts. The later an environmentalist enters the planning flow the less likely is his possibility of success. It is the melding of biological data and philos- ophies into the land-use planning regimes of our society that 1s now just getting off the ground, or should I say on the ground. Publications such as this one are efforts in that direction. PAUL F. J. EAGLES Faculty of Environmental Studies, University of Waterloo, Waterloo, Ontario N2L 3GI Northern Europe the slides sealed with paraffin wax. Preparations for scanning, transmission electron microscopy are clear- ly explained in the text. It is unfortunate that there was no description of the equipment used to collect airborne pollen and spores. The amounts of airborne pollen and spores can be summarized more specifically by using one of the collecting devices now used in several countries. | doubt, for example, that the very minute airborne pollen grains from dandelion, Taraxacum spp. or from goldenrod, Solidago spp., would be caught in very large amounts. I. JOHN BASSETT Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 104 Orchid Biology: reviews and perspectives, I Edited by Joseph Arditti. 1977. Cornell University Press, Ithaca. 310 pp. US $29.50. Biology is a wide field of study, even if all the organisms studied belong to a single plant family. Even so, its meaning has to be stretched to cover the range of topics in this book. One third of the text is devoted to “A manual for clonal propagation of orchids by tissue culture.” In this chapter, J. Arditti has gathered together all the techniques brought to his attention by the cut-off date: December 1974. Not only does he cite the original papers but also provides comments based on his own experience. As an annotated review of the literature it should be of some value to people in the area. The other articles seem of less value, either because they cover little new ground or because there is so little good information available. The account of the fossil history of the Orchidaceae appears to fall in the first category. To people like myself who have not read the Schmids’ previous articles on the topic, published in 1973 and 1974 and cited here, their article is interesting and provides a useful warning on the perils of using ENVIRONMENT Nature Quizzes for Canadians By V. and B. McMillan. 1976. Douglas, Vancouver. 144 pp., illus. Paper $3.95. The impetus behind this book evolved from a desire to assist Canadians in learning about their country’s natural environment. I must admit, there are some interesting and informative chapters; however, the text is poorly organized and filled with errors. The numbering systems employed to differentiate the questions are very poor. In many cases the numbers used to label the diagrams are also designed to double as question numbers. On page 89 the first seven questions are labelled alphabetically froma to g, while the remaining questions in the chapter read as numerics (8--20). Very confusing. In the chapter examining the reader’s knowledge of ‘Immature Insects,’ pictures of the adult forms, which the reader is asked to match up with the immature stages, are completely missing. On page 90 (question 11) the authors make reference to a diagram that does not exist. I also found a question (#5, page 15) where the THE CANADIAN FIELD-NATURALIST Vol. 93 secondhand sources of information. I am doubtful, however, of its value to students of orchids. The article on anthocyanins of the Orchidaceae falls into the second class: not enough information is available to justify the length of this section. Two other articles fall outside the bounds of orchid biology to my mind: an account of his involvement with orchids by R. E. Holttum and a discussion of Rumphius’ contribution to orchidology by H.C. D. de Wit. I enjoyed them both as general reading, although that on Rumphius is, in my judgment, rather long. This article and the others would benefit from a more severe editorial hand. As it stands, the section on tissue culture techniques should earn this book a place on the shelves of orchid breeders and the title will recom- mend the book to ardent orchid devotees. It is perhaps only fair to admit that I myself am not one. M. BARKWORTH Biosystematics Research Institute, Ottawa, Ontario KIA 0C6 wrong answer, followed by a correct explanation, was recorded, thus causing a contradiction, which to a novice might prove confusing. On page 41 (question 20) the reader is asked to give the name of a northern freshwater fish belonging to the Gadidae. Directly below the question, not in the answer section where it belongs, is the answer, a labelled diagram of a burbot. There is no consistency in the usage of nomen- clature. I found three incorrect spellings: the red admiral butterfly (Vanessa atlanta) is not Vanessa atalenta (page 38); the moose (A/ces alces) is not Alcea alcea (page 40); the muskrat (Ondatra zibethicus) is not Ondatra zibethica (page 49). I feel that the concept is sound, and that the text has possibilities, but until the book is properly organized and the errors in composition are corrected, the book is not worth buying. PAUL A. GRAY Ontario Ministry of Natural Resources, Box 89, Cochenour, Ontario POV ILO 1979 Book REVIEWS 105 The Shetland Way of Oil. Reactions of a small community to big business Edited by John Button. 1976. Thuleprint Ltd., Sandwick, Shetland. 134 pp. £2.40. This is an admirable little book. Although it deals with events in a faraway place, under a jurisdiction and an administrative system that differ from those in Canada, it offers much sensible advice, as well as stimulating examples of success, to all who care for more in the world they live in than just their own financial prosperity. The most powerful reason for the recent setbacks suffered by the “environmental movement” has been the sustained demand for oil, despite the massive price increases imposed by the OPEC countries. Part of the western reaction was to intensify the search for other supplies. Their most substantial early success was the discovery of large quantities of oil and gas in the northern North Sea and the development of tech- nology to extract oil from under the seabed at unprecedented depths. The nearest deep-water harbor to some of the most promising oil fields was found at Sullom Voe, off the Mainland of Shetland. So there the oil industry descended, with plans for the development of a storage facility for crude oil brought ashore by pipeline and later establishment of a refining plant too. The fourteen chapters of this book, each by a different author, half of them local men, deal chiefly with the responses of the local communities to the potent and unexpected combination of opportunities and hazards. By great good fortune, the expertise and the affluence of the oil companies and their supporters in distant arenas of government did not altogether overwhelm the local talent. The Zetland County Council (now the Shetland Islands Council) quickly devised an Interim Development Plan. It succeeded in obtaining legislation (the Zetland County Council Act of 1974) which gave it powers exceptional for a local authority. The council now acts as the port and harbor authority for Sullom Voe; can acquire land for oil-related development within the designated area; issues licences to dredge (and may refuse to do so) and licences construction work within the three-mile territorial limit; may take shares in commercial undertakings; and may establish a Reserve Fund. With so strong a jurisdictional base the local people have been well placed to exercise control over undesirable development and have succeeded in doing so to a remarkable extent. For naturalists the chapters by Peter Kinnear on “Birds and Oil” and by Brian John on “The Milford Haven Experience” are of greatest interest. Kinnear writes “The value that is placed on birds and other wildlife, which do not provide some immediate financial return, is to a large extent subjective. For many people the value might seem sufficient, but it cuts little ice with either Government or developers, who require a more qualified scientific appraisal. . . . One way of assessing the importance of Shetland’s birdlife is to consider the number of different species and to see what proportion of the total British stocks they constitute.” Fortunately for his chosen approach, those numbers are known and for several species do make up an impressive fraction of the British stock: 90% of the Great Skuas (Stercorarius skua) (or 50% of all those in the northern hemisphere) and 70% of Arctic Skuas (S. parasiticus); 90% of the Whimbrel (Numenius phaeopus), 1/2 to 2/3 of the Red-necked Phalaropes (Phalaropus lobatus), 1/3 to 1/2 of the Red-throated Divers (Gavia stellata) and so on, with nationally significant stocks of at least 24 species. Most of those species are marine and so directly at risk from oil pollution, from pipeline leaks, spills on shore, bilge discharge and tank washing by tankers and, eventually no doubt, tanker collisions or groundings. To look at the probability Kinnear turns to the detailed experience gained over the last fifteen years or so in and around Milford Haven, in west Wales, a major reception area for large tankers and “widely regarded as the cleanest oil port in the world.” Milford Haven handled 53.1 million tons of oilin 1973. Sullom Voe is expected to handle 50 to 150 million tons annually over the next twenty years or so. In the eleven years 1963-1973 Milford experienced an average of 53 spills a year. Sullom can thus expect a similar or larger number, amounting to between 400 and 1200 tons a year, excluding any major accident. But, whereas Milford Haven is subjected to massive tidal flushing, Sullom Voe has only one eighth of Milford Haven’s tidal velocity and only a fraction of its water is changed each tidal cycle. Building up of oil and other pollutants seems inevitable; and a major tanker accident is likely too, because manoeuvring - very large tankers through Yell Sound “will always be a hazardous affair, particularly during fog and severe weather.” Kinnear, however, sees the greatest danger to birds as likely to result from the continuing practice of discharging oily ballast water at sea (in order to save time at the terminal), an activity which it is often hard to detect and prove. He complains that British regulations governing pollution control are in dire need of change. “Although various laws restrict the discharge and dumping of oil in the sea, once there it is incumbent on no-one to do anything about it.” What lessons can naturalists in Canada learn from this book? First, in many areas now being explored for oil we still lack the basic inventories to establish numerically what birds are at risk, although great progress has been made along the eastern Arctic and 106 Atlantic coasts and offshore in the last decade. That task must be completed and the results made known to governments, the oil industry and the public, whether or not they want to hear about them. Second, the best hope for effective actions on behalf of wildlife lies in power being given to local people and in their vigilant and responsible use of those powers. (The Inuit seem to understand this more clearly than many people elsewhere.) Third, wildlife interests are most unlikely to be sufficient to prevent major economic developments, the equivalent of North Sea oil, from taking place; but they can and should make an important contribution to keeping the activities of exploiters cleaner than the latter would like, though not as clean as the birds require. So longas oil is taken THE CANADIAN FIELD-NATURALIST Vol. 93 from under the sea or carried across it, birds will be the losers. “So we return to the moral question and the personal value each one of us places on birds and wildlife. While you decide how you are going to play your part in changing things, spare a thought for the vast numbers of birds which migrate at night and each year die in thousands, attracted by and consumed in the giant flames burning from the rigs across the North Sea” (page 99). HuGH BoyD Canadian Wildlife Service, Environment Canada, Ottawa, Ontario KIA 0E7 EZAIM: écologie de la zone de l’aéroport international de Montréal — Le Cadre d’une recherche écologique interdisciplinaire Par P. Dansereau. 1976. Les Presses de l’Université de Montréal, Montréal. Polycopié. 343 pp. Comme a déja dit Pun des chefs d’€quipe impliqueé dans cette étude écologique de la zone de l’a€roport devenu Mirabel, étude a priori multidisciplinaire: “on a limpression qu'il n’y a que Dansereau qui fait du multidisciplinaire.” Ce commentaire souleéve a lui seul Yampleur du probleme de la communication entre des spécialistes de disciplines aussi variées que la psy- chologie, la geomorphologie, la zoologie. Ce méme probleme cede cependant le pas a la science elle-méme a lintérieur d'une méme spécialiteé. Ayant participé a cette étude, j'ai pu me rendre compte que le travail d’équipe fonctionnait au niveau de chaque équipe dans leur discipline propre, mais que le travail d’intégration multidisciplinaire ne semblait quétre le fruit d'une démarche personnelle, celle de Dansereau. Ce dernier se veut cependant plus positif quant a la possibilité et la réalisation d’un projet multidisciplinaire et complexe a souhait. L’étude EZAIM a été entreprise apres le début des travaux de construction et d’ameénagement de l’aéro- port, ce qui souléve peut-é€tre son caractere secondaire aux yeux des intéréts €conomico-politiques, probleme soulevé par Dansereau lui-méme en fin d’ouvrage d’ailleurs. Le plan de recherche envisagé a été calqué sur ’écosysteme lui-méme, a savoir les différents niveaux trophiques entre lesquels ont lieu des interactions. Les €quipes étaient donc divisées en géomorphologie, écologie végétale, écologie animale (mammiferes, olseaux), géographie humaine (utilisation des terres, industrie et urbanisation), psychiatrie sociale, sans compter les échelons nécessaires de coordination et d’administration. Dansereau insiste sur le fait que cette recherche se veut une expérience, multidisciplinaire soit, mais expérience intégrée par l’auteur dans le contexte d’une “philosophie écologique” quil prone. Le chapitre 2 est peut-étre a ce point de vue le plus important, car Yauteur y expose les définitions de sa démarche qui doit assurer la cohérence a la recherche (ou a toute recherche) qui s’attaque finalement a des €cosystemes tres variés. La définition élargie qu'il donne a l’écosystéme est la suivante: “Un espace limité ot le cyclage des ressources a travers un ou plusieurs niveaux tro- phiques est effectué par des agents plus ou moins fixés et nombreux, utilisant simultanément et successive- ment des processus mutuellement compatibles qui engendrent des produits utilisables 4 courte ou longue échéance.” Se basant sur cette hypothése de travail, et par une description des relations qui interviennent entre les six niveaux trophiques (minéro-, phyto-, zootrophie (herbivorie, carnivorie), investissement et controle) dans des écosystemes types, c’est-a-dire par une analyse écosystématique des paysages, il parvient a définir tout paysage d’un point de vue écologique. Cette analyse consiste en un inventaire des ressources présentes a chaque niveau trophique et des agents et processus ou forces motrices ayant court dans la transformation d’un niveau a l'autre ou d’un paysage a l'autre. Elle implique également la compréhension de l'aspect dynamique du cyclage dans chaque paysage pour mieux situer la stratégie régionale d’un point de 1979 vue écologique, et mener ainsi a la construction de matrices d’impact qui devraient guider la planifi- cation. Mais non content de l’analyse €cosystématique de la zone de Mirabel, Dansereau débouche naturellement sur Vuniversel, soit lorsquwil présente les régions phytotrophiques, dont celles applicables a Mirabel, soit lorsquil présente son nouveau systeme de classification des terres basé sur l’escalade de impact humain. Et ce sont justement ces visées universalistes qui font de cet ouvrage de Dansereau un outil de plus pour les études a caracteére multidisciplinaire. Toute cette philosophie écologique peut se résumer graphiquement par un “boule-de-fléches” (sphére qui contient les circuits de ressources, agents, processus et produits) qui illustre sa démarche. Cette démarche n’a pas été comprise par tous les participants du projet EZAIM, ou selon lexpression de Dansereau “a souvent été perdue de vue.” Du point de vue strictement de la présentation, les erreurs typographiques sont rares. Certaines figures ou tableaux sont incomplets: la légende de la figure 23 nest pas rapportée enti¢rement au tableau XIII tel que mentionneé. Le style est simple et clair, sice n’est le vocabulaire scientifique propre au domaine du botaniste qui transpire tout au long de l’ouvrage. BooK REVIEWS 107 On y trouvera aussi une autocritique, sirement pas assez vive a mes yeux et a ceux de certains autres participants, du projet EZAIM tel qu'il avait été con¢u par l’auteur du présent ouvrage et tel qu’ila été réalisé. Globalement, Vouvrage rend davantage compte de ce que devrait étre une recherche €colo- gique interdisciplinaire a partie d'un modéle, plutdt que d’étre une synthese des résultats écologiques recueillis par chaque équipe. Ce qui retiendra notre attention sera ce modéle €cosystématique de Dan- sereau, sa philosophie qui permettrait d’analyser tous les types de paysage selon un point de vue €cologique, y compris les paysages humains et il faut, je crois rendre hommage a Dansereau pour sa vision syn- thétique du cadre qu'il propose. On ne saurait trop recommander la lecture de cet ouvrage aux aménagistes et aux spécialistes a tous les niveaux trophiques, tant naturels qu’artificiels ou humains, tout au moins pour les changements dans les valeurs sociales qu'il préne pour les développements d’ordre socio-économiques a venir. ANDRE CYR Département de Biologie, Université de Sherbrooke, Sher- brooke, Québec JIK 2RI Outdoors Canada. A unique and practical guide to our wilderness and wildlife Edited by Douglas R. Long. 1977. Reader’s Digest Canada Ltd. and the Canadian Automobile Association, Mon- treal. 383 pp., illus. $21.77. Outdoors Canada is intended to assist Canadians to enjoy their out-of-doors. The book not only tells of the out-of-doors (mammals, birds, plants), but it also explains how to cope in the out-of-doors (camping, canoeing, backpacking). Photographs demonstrate what can be seen, while numerous drawings illustrate and explain the book’s “how to” information. Not all of the information in the section on wildlife is entirely accurate. Some notable inaccuracies would include these statements: “many fur bearing animals are near extinction” (p. 15); “most great whales are endangered species” (p. 13); “the distribution of porcupine is south of the treeline” (should read mainly within the predominantly hardwood forest areas). The section is far from complete but does present an interesting capsulized summary of most groups of animals found in Canada. As noted within the text, birds are animals and are included within the animal section. The treatment given to birds is less complete than that given to mammals. Again some of the book’s statements are questionable. One such statement to which some persons might take exception would be, “the whoop- ing crane is North America’s rarest bird.” The section. on birds will not serve as a field guide; however, it is enjoyable reading. Whereas for the section on Animals the photo- graphs tend to supplement the text, the reverse treatment is given to the section on Plants: photo- graphs are supplemented by the text. Also, the Plant section tends predominantly to focus on flowering plants and generally those that are more common. Again this section will not replace a field-guide treatment as is popularly used by naturalists. Unlike the Animal section, this section appears disorganized. The section on Camping covers the whole gambit of outdoor travel. This section is generally well or- ganized, well presented, and contains much good . advice. One comment I would caution the reader about is this: “to keep your feet dry in leaky boots orin shoes that are not waterproof, wear plastic bags over 108 socks.” If the weather is cold, perspiration within the plastic bags could result in very cold feet (possibly frozen). As the book points out, “experience is probably the surest guide.” As surely as rules are made, there are exceptions. The section, “Outdoors: Just for Fun,” suggests it is a good idea to stay with your canoe after it is capsized. But common sense should prevail and when swamped or capsized in white water, sometimes it is best to leave the canoe. On occasion, people have been known to get pinned between their canoe and rocks, resulting in serious injuries. The last 49 pages of Outdoors Canadais a series of regional (Canadian) maps. These pinpoint the loca- tions of various outdoor areas in Canada. A brief note THE CANADIAN FIELD-NATURALIST Vol. 93 explains the features of specific sites. This section will be of assistance to those persons planning a Canadian out-of-doors vacation. Generally speaking, Outdoors Canada is very much a Reader’s Digest-type book. It is a brief summary of material rather than a complete treatment. The book covers the whole spectrum of the outdoors and admirably does not get bogged down in any one section. For the inexperienced outdoors-person, it 1s an excellent buy. PETER CROSKERY Ontario Ministry of Natural Resources, Ignace, Ontario POT ITO Why Big Fierce Animals Are Rare. An ecologist’s perspective By P. Colinvaux. 1978. Princeton University Press, Prince- ton, New Jersey. viii + 256 pp. $9.50. In an informative and logical fashion Colinvaux presents the rudiments of ecology, and endeavors ...“to trace the status of the ecologist’s quest” ina thorough examination of the numerous riddles of nature. Without obscuring the text with highly technical terms that might discourage the novice, he reveals concerns of modern ecologists, concerns nurtured by pioneers like Charles Darwin, G. F. Gause, and Charles Elton. In 18 chapters he outlines the niche, biogeography, aquatic-terrestrial-atmospheric cycles, succession, co- existence, behavior, species diversity, environmental stability, and the evolution of man’s role in nature. Page by page the ecological story unfolds as the author describes the efforts of men like Alphonse de Candolle, the last man singly to attempt to describe all of the known plants in the world; Vladimir Koppen, the climatologist who mapped world weather using vegetation zones; C. Hart Merriam, the explorer who devised the life zone concept; and Tansley, the English botanist who coined the word ‘ecosystem.’ The subject matter is uniquely accented with a number of interesting anecdotes. For example, in chapter 3, entitled “Why Big Fierce Animals Are Rare,” Colinvaux exposes the myth of 7yrannosau- urus rex amidst a discussion of food-chain dy- namics. He also dispels the fantasies of hope that evolved from the green revolution in a few short sentences: “It is a myth that is probably as impossible to eradicate as the myth that Tyrannosaurus rex was a ferociously active predator. But myth it is. Algae are not more productive than other plants.” I was particularly impressed with the final chapter, “The People’s Place,” in which the author recon- structs the story of man’s climb to environmental dominance. But more than that, Colinvaux stresses the importance of man’s dynamic role in the complex ecosystems of the earth, a role that beginning ecologists must understand in order to develop an appreciation for the relations among other living organisms and their environment. The text is greatly enhanced by a section on ecological reading, which lists many excellent papers on the numerous subjects of the science. A useful index is also included. Unfortunately, the title of the book, Why Big Fierce Animals Are Rare, is mis- leading; in my opinion the title suggests that the book is about predator ecology, not a general overview of ecology. Nonetheless, it is well written, and is as complete a book as one would expect at the introductory level. PAUL A. GRAY Ministry of Natural Resources, Box 89, Cochenour, Ontario POV ILO 1979 OTHER Northern Vagabond. The life and career of J. B. By Alex Inglis. 1978. McClelland and Stewart, Toronto. 256 pp., illus. $14.95. The author begins this interesting book by des- cribing Joseph Burr Tyrrell’s famous 1893 3000-mile trip through the central Barren Lands of Canada — one of the last great feats of exploration in Canada’s history. When Tyrrell’s party of eight set out by canoe from Fort Chippewyan they were not certain whether the Dubawnt and Thelon rivers, along which they planned to paddle, would carry them to Hudson Bay or to the Arctic Ocean; if the latter, their chances of returning alive were slim. Nor were they sure they would encounter game, largely Caribou, to feed them. Their inland exploration ended successfully at Chesterfield Inlet on Hudson Bay, but their trip was by no means over. They had yet to make their way through blizzards and ice along the coast for 400 miles to Fort Churchill, and from there walk and travel by dog sled the 900 miles to Winnipeg and the railway. Tyrrell undertook this voyage while he was a geologist with the Geological Survey of Canada, a position he held from 1881 to 1898 although he had little formal training as a scientist. He spent his summers exploring the wilds of western and northern Canada, his winters in Ottawa writing reports on his geological and other findings. He enjoyed his field work especially (in one week in Alberta he discovered the largest deposit of prehistoric bones and the greatest deposit of workable coal to be found in Canada). He is probably best known scientifically for his postulation, based on rock striations, that during NEW TITLES Zoology Animal facts and feats. 1977. By Gerald L. Wood. Revised edition. Sterling, New York. 256 pp., illus. US $14.95. Animals in peril. A guide to the endangered species of Canada and the United States. 1978. By David C. Grainger. Macmillan, Toronto. 192 pp., illus. $14.95 The beetles of North America. 1977. By Richard Head- strom. Barnes, Cranbury, New Jersey. 448 pp., illus. US $17.50. The California quail. 1978. By A. S. Leopold. University of California Press, Berkeley. xx + 281 pp. US $14.95. Book REVIEWS 109 Tyrrell the Ice Age Canada was not covered by a single massive ice field which advanced and retired once ona long front, but by several glaciers which originated from separate centers and had advanced and retreated on several occasions. During his lifetime, besides his geological reports, he also wrote dozens of books and articles on such topics as Samuel Hearne, David Thompson, Canadian history, and mining in Canada. In 1899 Tyrrell left the Survey because he had been denied promotion, and moved to the Klondike as a Mining Consultant. He lived there in the midst of the gold rush until 1905, learning so much about gold that the rest of his career was centered on this mineral, in part as a goldmine manager near Kirkland Lake, Ontario. When he retired a wealthy man, he turned to managing an apple orchard on the Rouge River, where many of us who lived in Toronto at the time were privileged to meet him. He died in 1957 at the age of 99. This book isa timely one, dealing as it does with one of our first native-born natural scientists. It not only follows Tyrrell’s life in detail, but it gives us a glimpse of the milieu in which an early government scientist worked (The Ottawa Field-Naturalists’ Club meet- ings for example offered Tyrrell and his friends a welcome diversion from their work). We are indebted to Inglis for bringing the life of this remarkable Canadian to our attention. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 A changing world for wildlife. 1977. By G. S. Fichter. Golden/ Western, New York. 141 pp., illus. US $8.95. Coyotes: biology, behavior and management. 1978. Edited by M. Bekoff. Academic, New York. 400 pp. US $34.50. +Endangered birds. Management techniques for preserving threatened species. 1978. Edited by Stanley A. Temple. University of Wisconsin Press, Madison. xxiii + 466 pp., illus. US $9.50. +L’entomologiste amateur. 1977. Par A. Villiers. Savoir en histoire naturelle xxvii. Lechevalier, Paris. 248 pp. illus. 90F. Great Canadian animal stories. 1978. Edited by Muriel 110 Whitaker. Hurtig, Edmonton. 232 pp., illus. $12.95. *Mammals of the palaearctic region: a taxonomic review. 1978. By G. B. Corbet. British Museum (Natural History). Cornell University Press, Ithaca. 314 pp., illus. US $38.50. *Nesting ecology of Canada geese in the Hudson Bay Lowlands of Ontario: evolution and population regulation. 1977. By Dennis G. Raveling and Harry G. Lumsden. Fish and Wildlife Report No. 98. Ontario Government Book- store, Toronto. v + 77 pp. Paper $3.50. The response of peregrine falcons (Falco peregrinus) to aircraft and human disturbance. 1977. By Jim Windsor. Canadian Wildlife Service, Ottawa. 87 pp. The Running Press book of turtles. 1977. By Richard E. Nicholls. Running Press, Philadelphia. 150 pp., illus. Paper US $4.95. The view from the oak: the private worlds of other creatures. 1977. By Judith and Herbert Kohl. Sierra Club/ Scribners, San Francisco. 112 pp., illus. Cloth US $8.95; paper US $4.95. Water birds of California. 1977. By H. L. Cogswell and G. Christman. University of California Press, Berkeley. 399 pp. Paper US $5.75. Wild birds of the Americas. 1977. By Terence Michael Shortt. Houghton Mifflin, Boston. 272 pp., illus. US $14.95. Botany +Ferns of the Ottawa district. 1978. By William J. Cody. Agriculture Canada Publication 974. Supply and Services Canada, Hull. 112 pp., illus. $3.25 in Canada; $3.90 elsewhere. A field guide to edible wild plants of eastern and central North America. 1978. By L. Peterson. Houghton Mifflin, Boston. 384 pp. US $8.95. Grass: the everything, everywhere plant. 1977. By Augusta Goldin. Nelson, New York. 176 pp., illus. US $7.95. Mushrooms of North America. 1977. By O. K. Miller, Jr. Chanticleer/Dutton, New York. 368 pp., illus. Paper US $8.95. +Native trees and shrubs of Newfoundland and Labrador. 1978. By A. Glenn Ryan. Newfoundland Department of Tourism, St. John’s. 117 pp., illus. Free. tNorth American forest lands at latitudes north of 60 degrees. 1978. By various authors. Proceedings of a symposium, University of Alaska, 19-22 September, 1977. School of Agriculture and Land Resources Management, University of Alaska, Fairbanks. 332 pp., illus. Free. Syrup trees. 1978. By Bruce Thompson. Walnut Press, Fountain Hills, Arizona. US $6.95. Wild food plants of Indiana and adjacent states. 1977. By Alan and Sue McPherson. Indiana University Press, Bloomington. viii + 215 pp., illus. Paper US $4.95; cloth US $12.50. Environment Biological data in water pollution assessment: quantitative and statistical analyses. 1978. Edited by K. L. Dickson, THE CANADIAN FIELD-NATURALIST Vol. 93 John Cairns, Jr., and R. J. Livingston. Special Technical Publication 652. American Society for Testing and Materials, Philadelphia. 193 pp. US $17.50 (less 20% to ASTM members). Biological nomenclature. 1978. By C. Jeffrey. 2nd edition. Crame, Russak, New York. viii + 72 pp. US $11.50. The breakdown and restoration of ecosystems. 1978. Edited by M. W. Holdgate and M. J. Woodham. NATO Conference Series I: Ecology, Volume 3. Plenum, New York. 506 pp. US $36. Concepts of applied ecology. 1978. By R.S. DeSanto. Springer-Verlag, New York. 320 pp. Paper US $9.80. +Environmental role of nitrogen-fixing blue-green algae and asymbiotic bacteria. 1978. Edited by U. Granhall. Ecologi- cal Bulletins NFR 26. Swedish National Science Research Council, Stockholm. 400 pp., illus. 100SwCr. Estimating the hazard of chemical substances to aquatic life. 1978. Edited by John Cairns, Jr., K. L. Dickson, and A. W. Maki. American Society for Testing and Materials, Philadel- phia. 283 pp. US $19.50 (less 20% to ASTM members). Evolutionary ecology. 1978. By E. R. Pianka. Harper and Row, New York. 384 pp. US $16.95. The green world: an introduction to plants and people. 1978. By Richard M. Klein. Harper and Row, New York. 512 pp. US $14.95. The management of estuarine resources in Canada. 1978. By Irving K. Fox and J. P. Nowlan. Report No. 6, Canadian Environmental Advisory Council, Ottawa. 51 pp. Free. Mexican wilderness and wildlife. 1978. By Ben Tinker. University of Texas Press, Austin. xii + 132 pp., illus. US $9.95. Reports of the first and second meetings of publie interest groups with the Canadian Environmenta! Advisory Coun- cil. 1978. By anonymous. Report No. 7, Canadian En- vironmental Advisory Council, Ottawa. 124 pp. Free. Resources, environment and economics. 1978. By R. U. Ayres. Wiley-Interscience, Somerset, New Jersey. 240 pp. US $24.95. Sourcebook on the environment. A guide to the literature. 1978. Edited by K. A. Hammond, G. Macinko, and W. B. Fairchild. University of Chicago Press, Chicago. x + 614 pp. US $22. Water and society: conflicts in development. Part 1. The social and ecological effects of water development in developing countries. 1978. Edited by Carl Widstrand. Pergamon, New York. 134 pp., illus. US $20. Miscellaneous Carbon dioxide, climate and society. 1978. Edited by Jill Williams. Proceedings of an IIASA Workshop, 21-24 February, 1978. Pergamon, New York. 304 pp. US $30. Earth observation systems for resource management and environmental control. 1978. Edited by Donald J. Clough and L. W. Morley. Plenum, New York. 475 pp. US $42.50. 72) Journey through the universe. An introduction to as- tronomy. 1978. By T.L. Swihart. Houghton Mifflin, Boston. xvi + 366 pp., illus. US $15.95. The mountains of Canada. 1978. By Randy Morse. Hurtig, Edmonton. 144 pp., illus. $29.95. Quaternary geology: a stratigraphic framework for multi- disciplinary work. 1978. By D. Q. Bowen. Pergamon, New York. 224 pp., illus. Cloth US $30; paper US $12.50. Manual on water. 1978. Edited by C. E. Hamilton. Special Technical Publication 442A. American Society for Testing and Materials, Philadephia. 471 pp. US $28.50 (less 20% to ASTM members). BoOK REVIEWS 111 Correction The listings of “Zoology New Titles,” Canadian Field- Naturalist 92(3): 316, 1978, should read as follows: Biology of bats, volume 3. 1977. Edited by W. A. Wimsatt. Academic, New York. 651 pp., illus. US $59. Biology of bryozoans. 1977. Edited by R. M. Woolacott and R. L. Zimmer. Academic, New York. 556 pp., illus. US $35. tavailable for review *assigned for review Instructions to Contributors Content The Canadian Field-Naturalist is a medium for the publication of scientific papers by amateur and professional naturalists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All readers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. As the journal has a flexible publication policy, items not covered in the traditional sections (Articles, Notes, Letters, News and Comment, and Book Reviews) can be given a special place provided they are judged suitable. Readers are encouraged to support regional, provincial, and local natural history publications as well by submitting to them their reports of more restricted signifi- cance. Manuscripts Please submit, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. Also authors are expected to have complied with all pertinent legislation regarding the study, disturbance, or collection of animals, plants, or minerals. Type the manuscript on standard-size paper, if possible use paper with numbered lines, double-space throughout, leave generous margins to allow for copy marking, and number each page. For Articles, provide a running head, a bibliographic strip, an abstract, and a list of key words. These items are optional for Notes. Generally words should not be abbreviated but use SI symbols for units of measure. Underline only words meant to appear in italics. 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The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. ae G: pe Teneaia” i _ Pre Gass ae? Siete San pe =P eS ( et he peers ot Hale CON 4 att A : joe Se a i ¥ es ber - ‘ cabs r eee een So DG ees wes TABLE OF CONTENTS (concluded) News and Comments 87 Book Reviews Zoology: A guide to bird finding east of the Mississippi — The birds of New Brunswick — Mountain 91 monarchs: wild sheep and goats of the Himalaya — Précis de zoologie: vertébrés. 1: reproduc- tion, biologie, évolution et systématique. Oiseaux et mammiferes — Ways of the six-footed — Birds of southeastern Michigan and southwestern Ontario — Watching sea birds — Bird hazards to aircraft: Problem and prevention of bird/ aircraft collision — Bears: their biology and man- agement — The earthworms (Lumbricidae and Sparanophilidae) of Ontario — The Passenger Pigeon: its natural history and extinction — The bird finder’s 3-year note book. Botany: The rare vascular plants of Ontario/Les plantes vasculaires rares de Ontario — Fungi: 100 delight of curiosity — Pollen flora of Argentina: modern spore and pollen types of Pteridophyta, Gymnospermae, and Angiospermae — Floristics and environmental planning in western New York and adjacent Ontario: distribution of legally protected plants and plant sanctuaries — Atlas of airborne pollen grains and spores in northern Europe — Orchid biology: reviews and perspectives, I. Environment: Nature quizzes for Canadians — The Shetland way of oil: reactions of a small com- 104 munity to big business — EZAIM: écologie de la zone de l’aéroport international de Montréal: le cadre d’une recherche écologique interdisciplinaire — Outdoors Canada: a unique and prac- tical guide to our wilderness and wildlife — Why big fierce animals are rare: an ecologist’s perspective. Other: Northern vagabond: the life and career of J. B. Tyrrell 109 New Titles 109 Mailing date of previous issue 8 January 1979 1978 Council — The Ottawa Field-Naturalists’ Club President: R. A. Foxall E. Beaubien C. Gruchy Vice-President: R. Taylor C. Beddoe P. Hall Treasurer: B. Henson W. J. Cody V. Hume : Recording Secretary: D. R. Laubitz Hones BS aaa Corresponding Secretary: A. Armstrong A. Dugal J. K. Strang C. Gilliatt EVE) Dy iiodd Those wishing to communicate with the Club should address correspondence to: The Ottawa-Field Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. For information on Club activities telephone (613) 722-3050. THE CANADIAN FIELD-NATURALIST Volume 93, Number 1 1979 President’s Message One hundred years in perspective — the changing roles and objectives of The Ottawa Field-Naturalists’ Club ROGER A. FOXALL Guest Editorial Biology — the unknown science? YORKE EDWARDS 6 Editorial The Canadian Field- Naturalist — the status quo or a new direction? LORRAINE C. SMITH 10 Articles Demographic and dietary responses of Red-tailed Hawks during a Snowshoe Hare fluctuation ROBERT S. ADAMCIK, ARLEN W. TODD, and LLOYD B. KEITH 16 The Oriskany sandstone outcrop and associated natural features, a unique occurrence in Canada DIANNE FAHSELT, PAUL MAYCOCK, GORDON WINDER and CRAIG CAMPBELL 28 Breeding areas and overnight roosting locations in the northern range of the Monarch Butterfly (Danaus plexippus plexippus) with a summary of associated migratory routes FRED A. URQUHART and NORAH R. URQUHART 41 Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta DAVID B. SCHOWALTER and JOHN R. GUNSON 48 Wild Mallard stocking in a large marsh habitat ROBERT O. BAILEY 5) Notes Recent collections of the Black Redhorse, Moxostoma duquesnei, from Ontario EDWARD KOTT, ROBERT E. JENKINS, and GREGORY HUMPHREYS 63 Xanthochroism in the Evening Grosbeak CHRISTOPHER W. HELLEINER 66 Interaction between a Long-tailed Weasel and a Snowy Owl PETER) @ Boxe 67 Black-necked Stilts nesting near Edmonton, Alberta ; DICK DEKKER, ROBERT LISTER, TERRY W. THORMIN, D. V. WESELOH, and LINDA M. WESELOH 68 Common Garter Snake predation on Robin nestlings KATHY MARTIN __70 Distribution and movements of Selkirk Caribou, 1972-1974 DAVID J. FREDDY 71 Effects of fire on the location of a Sharp-tailed Grouse arena DONALD A. SEXTON and MURRAY M. GILLESPIE 74 Brewer’s Blackbird breeding in the Northwest Territories PHILIP H. R. STEPNEY 76 A trap to measure Populus and Salix seedfall JOHN C. ZASADA and ROSEANN DENSMORE 1 Examination of overwintering adult carabid beetles for associated mites J. E. OLYNYK and R. FREITAG 79 Ring counts in Salix arctica from northern Ellesmere Island D. B. O. SAVILE 81 Giant Cow Parsnip (Heracleum mantegazzianum) on Vancouver Island, British Columbia NEIL K. DAWE and ERIC R. WHITE 82 Nesting of Horned Puffins in British Columbia R. WAYNE CAMPBELL, HARRY R. CARTER, and SPENCER G. SEALY 84 concluded on inside back cover ISSN 0008-3550 The CANADIAN FIELD-NATURALIST Published b y THE OTTA WA FIELD-NATURALISTS’ CLUB, O , Ottawa, Canada VOL. LY, No. 8 THE HEIDNATUT NATURALIST. published by the ottawa Field-Naturalists’ Club. CONTEN TS. : Past 4, Title Page tor Vol 1X, (TAWA NATUR ALIST oc ee aie hes nee 1 3, Patron, Gouneil, Committees and Leaders of The Ottawa Field ‘Naturalists’ Club for 1895-1806, 3 3, Last of Ordinary and Corresponding Members 3! Urn: Pine (nee ts 4, Calourless Biood in Amimals—Prot, Bf, B, Prince, pa, FLS T & The Rensselaer Grit Plateau RW. Ells, LL.D.) PRS... 7 oe Sale a 8, The Relation of the Atmosphere to Agriculture—F- T. Shutt, MA 5 pa We Beer 12 7. Announcement, Royal Society of Canada ee EE 4 g, Annual Report of Council for 1AGL-1895 1 g, Treasurers Statement -- eae ee aie an ae w 4p, Notes, Review? and ‘Comments, Geategy, Conehalegys Ornitholag > 20 2A yn. Geological Society of Americt 2-7 AR. Faitorial Note 2 oS i ere, OTTAWA, CANADA. PRINTED BY PAYNTER & ApsoTt, 48 RipEad SrReEr. Enteral cad the Oltaea Bost Office os secund last matter 1S. Entered at the ee NOVEMBER 30th, 1941 wa Post Office as cera cl -class matter Centennial Year Volume 93, Number 2 NOVEMBER, 1941 April-J une 1979 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things. The Members of Council are listed on the inside back cover. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors C3D» Bird A. J. Erskine David P. Scott E. L. Bousfield Charles Jonkel Stephen M. Smith Francis R. Cook Charles J. Krebs Robert E. Wrigley George H. La Roi Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang Production Manager: Pauline A. Smith Business Manager: W. J. Cody Subscriptions and Membership Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The Canadian Field-Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879- 1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KILY 4J5 Book Review Editor: Dr. J. Wilson Eedy, R.R. 1, Moffat, Ontario LOP IJO Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon, USA 97731 (address valid until August 1979). Address manuscripts on birds to the Associate Editor for Ornithology: Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0 All other material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0 Urgent telephone calls may be made to the Editor’s office (613-996-5840), the office of the Assistant to the Editor (613-231- 4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager’s office (613-995-9461). Cover: Since The Ottawa Field-Naturalists’ Club was founded in 1879, it has published a scientific journal. Two covers are illustrated. Left, The Ottawa Naturalist, published from 1887 to 1919. Right, The Canadian Field- Naturalist, published from 1919 to the present; it bore this cover design from 1919 to 1944. The Canadian Field-Naturalist Volume 93, Number 2. April-June 1979 Our Responsibilities as Field-Naturalists and Biologists The important message in the guest editorial by Yorke Edwards (Biology — the unknown science? Canadian Field-Naturalist 93(1): 6-9, 1979) is that the world needs more generalists in science and that workers in biology must communicate better. My comments here are offered not only as a reinforcement of his views but also to point out the responsibilities of field-naturalists and biologists regarding the quality, accuracy, and comprehensiveness of their studies. Although for years biologists have been isolated from social and political changes, increasing diaiogue between scientists and the general public and governments 1s taking place. Not all of this is productive. Nevertheless society is becoming more aware that many of the world’s troubles are concerned with changes in normal biological processes and ecosystems and that the nature of environmental problems is complex. In order to formulate policies, decision-makers (normally situated at the government level) must assess the opinions, often conflicting ones, of scientists. Many scientists find it difficult to interpret science to the layman. Moreover, with increasing specialization some scientists can convey information only on limited topics in their own field of interest and are often unable to comprehend the total picture. Therefore, we need more competent generalists to integrate and interpret scientific data and to communicate science to society. As Yorke Edwards wrote, “We need a new morality in the sciences, resulting in major efforts to make science and its influence on Earth understandable.” Because the role of scientists in society is becoming more visible, I think it is germane to point out some of the responsibilities that we should be aware of, but often are not. In particular we should recognize our limitations, strive for accuracy, and maintain credibility. It is important, however, not only that we recognize our own scientific limitations but that we do not exceed them. Unfortunately some biologists extend their studies, interpretations, and conclusions beyond their level of expertise and competence. Sometimes their conclusions are based on results obtained using faulty methods and are thus incorrectly attributed to environmental variation. Sometimes their opinions, whether right or wrong, have been accepted as facts. How does the decision-maker know whose opinion to respect? This is a major problem which has become even more complicated by the mushrooming number of biological consulting firms, some excellent but others abysmally bad. How often has the perhaps glossy, long-winded, eye-appealing final report of the contractor been valued rather than whether the work on which it is based was scientifically sound? How often will firms or individuals take on work that is beyond their qualifications? We know that some environmental consulting firms at least try to make up for their deficiencies in qualified and competent staff by contacting and “picking the brains” of acknowledged experts (often without any financial or other compensation or acknowledgment to the experts for their time and knowledge) whereas others make no attempt to improve the quality of their studies. Some contractors do not have the ability or inclination to do a proper literature search whereas others may do a search, then write a research paper under the pretence that their report contains original work. Biologists or field-naturalists should not be considered specialists outside their own specialty. Environmental problems in particular are very complex, and what may appear on the surface to be simple almost always has complicated interrelated ecological aspects. The tendency to look at one particular aspect of a specific problem without regard for the broader view has sometimes resulted in 113 114 THE CANADIAN FIELD-NATURALIST Vol. 93 disastrous decisions. It is imperative, therefore, that the total involved system be considered if wise decisions are to be made. But so often this doesn’t happen. Too few are able to see, let alone comprehend, the total aspect of biological problems and the complicated interrelationships that exist in nature. Generalists who approach interdisciplinary studies with open eyes and an open mind are needed to understand and to interpret the basic workings of complex ecosystems, 1.e., to put it all together. But they too must know their limitations: they cannot be masters of all disciplines; therefore, they will need to consult specialists on specific aspects of particular problems. Although the popular interdisciplinary method of teamwork is commendable, often it does not fill the need fora broad approach. D. B. O. Savile pointed out some years ago (Unity from diversity in biological research. Transactions of the Royal Society of Canada 4(4): 245-251, 1966) that often each team member sees only his or her aspect of the problem. He elaborated further as follows: “For the clearest and fullest picture of any biological phenomenon, it is helpful if the interdisciplinary approach of the team is complemented by the multidisciplinary approach of the individual.” We have a responsibility to strive for accuracy. Sometimes incorrect descriptions, identifications, computations, etc., appear in reports. It is easier to perpetuate an error than to correct one. Studies should be carried out as thoroughly and competently as possible with careful regard to repeatability, the basis of the scientific method. Moreover, if statements can be misinterpreted, are ambiguous, or are otherwise unclear, they should be rewritten. Often there is considerable pressure to do a study and to prepare the report quickly without particular regard for accuracy and dependability, and consequently too much may be read from limited data. Researchers should be obliged to state the limitations of their methods, data, and conclusions so that there will be less distrust and fewer set-backs but rather an advancement of knowledge. We must endeavor to maintain our credibility and should encourage other biologists and field- naturalists to do so too. This is particularly important when the problems and issues that arise are emotional ones. Interpretations, recommendations, and conclusions should, as far as possible, be based on good sound scientific data. Speculation should be limited to what can be reasonably well supported. We mustn’t compare apples and oranges. Thus the results obtained on one species cannot be directly extrapolated to another species. Our credibility is dependent on the accuracy of our statements, on recognizing the limitations of the data, and on the limitations of our expertise. Obviously if errors are discovered in one part of our work, the results from an entire study will be suspect. For example at a recent naturalists’ meeting, a petition for signatures was presented. Although this petition had an admirable conservation measure as an objective (it was against the legalization of the taking of certain birds of prey), alas the reasoning accompanying it was false. It was alleged that these particular raptors should be saved because as rodent-eaters they are beneficial to farmers. When it was pointed out that these particular birds prey mainly on birds, not rodents, the petition was not withdrawn for this error to be corrected. In such cases, where we make false statements, opponents will dwell on them, and a good cause may be lost. A rather copious volume of “gray” scientific “literature” or pseudoliterature has appeared in recent years. Some of this has resulted from the many contracts let for baseline surveys and preparation of environmental impact statements. The greatest shortcoming of the duly written final reports is that they are not generally subjected to peer review as are research papers published in the primary literature. Frequently errors of commission and omission are put into such records and are perpetuated. Considerable time and effort are required to correct these. Indeed some are never corrected. The solution to this problem is not easy. Because they are not subjected to rigorous (or any in some cases) peer review, there is a tremendous variability in the reports in the gray liter- ature: some reports are very superficial but others are excellent in-depth studies. Better evaluation of these reports is needed. Floral and faunal surveys of many areas, especially national and provincial parks and areas designated by the International Biological Programme, are indeed valuable. But how can a judgment be made on whether the contractors are really trained and reliable? One partial answer is 1979 SMITH: EDITORIAL WS) to be sure voucher specimens are kept, because these at least can be checked for correct identification. We know of at least one government contract where the employees of a consulting firm had misidentified and reported incorrectly ona large proportion of the specimens. It is cases like this that when discovered cause credibility to be lost. But how many go undetected? Many biologists and field-naturalists are concerned about these quick and superficial surveys, with the overconfidence of some investigators, and in the final analysis with credibility. Other scientists are also worried about these problems. In his editorial in the first issue of a new journal Marine Environmental Research (July 1978) Eric Cowell wrote: “Nowhere has departure from scientific integrity been more evident than in the field of environmental research. .. . There is much to learn about the function of our ecosystem and man’s role in the management or mismanage- ment of it. We need clearly defined objectives, impeccable data and analyses and interpretation conducted with integrity.” My plea to field-naturalists and biologists is to follow the old adage “if it’s worth doing, it is worth doing well.” Of course it takes time and effort to do good research, whether it is in the field, in the laboratory or museum, or in the library. Check for accuracy, maintain scientific integrity, exercise care in the interpretation of data, and recognize the limitations. Seek the comments of specialist colleagues but look to generalists to give the total picture. Only if we recognize and carry out these responsibilities can we hope to maintain quality and standards and to contribute positively to society as responsible citizens. LORRAINE C. SMITH Editor Natural Fires as an Index of Paleoclimate J. TERASMAE! and N. C. WEEKS? 'Department of Geological Sciences 2Department of Biological Sciences, Brock University, St. Catharines, Ontario L2S 3A] Terasmae J. and N. C. Weeks. 1979. Natural fires as an index of paleoclimate. Canadian Field-Naturalist 93(2): 116-125. The charcoal abundance and frequency of occurrence varies stratigraphically in postglacial lake sediments, and this information can be used together with palynological and sedimentological data for a reconstruction of paleoclimatic conditions. The gelatin-coated slides method can be used for continuous sampling of lake sediment cores to determine the presence of charcoal particles. Changes in the charcoal profile have been related to changes in the sequence of fossil pollen, specifically that of pine pollen, and from the observed relationship a climatic control of forest fires is inferred. Forest fires have occurred naturally in the Great Lakes — St. Lawrence and Boreal forest regions during the past 9000 yr, and the fire frequency almost doubled during the Pine Pollen Zone about 7000 to 4000 yr ago in the southern Boreal forest. The mean fire frequency in the Lac Louis area was one fire every 95 to 100 yr, but during the period characterized by a pine pollen maximum, 7280 + 240 yr BP (GSC-1481) to 4260 + 240 yr BP (GSC-1491), the fire frequency increased to one fire every 48 to 56 yr. Key Words: forest fires, palynology, paleolimnology, paleoclimate. Natural fires, as distinct from man-induced ones, have been characteristic of terrestrial ecosystems probably for many millions of years. The prerequisites are (1) a readily combustible ‘fuel’ source (vegetation) and (2) an igniting mechanism. The first of these is_ satisfied whenever and wherever a supply of dry vege- tation is available, and both lightning and volcanism have been present as igniting mecha- nisms throughout geological time; however, the frequency of coincident fuel, ignition, and fire has varied through time. The fact that numerous and very extensive forest fires, as well as grassland and also tundra fires, coincide with general or seasonal dry weather episodes (drought) appears to be com- monly accepted (Rowe et al. 1974, 1975). Meteorological conditions leading to such epi- sodes of drought are caused by lack of precipi- tation, relatively high summer temperature, extended periods of sunny weather, and medium- to high-velocity surface wind. A combination of these meteorological variables will intensify the development of drought conditions and, hence, increase the susceptibility of vegetation to fire. The linkage between climatic conditions and the frequency and areal extent of natural fires in vegetation seems to be well established. There- fore we assume that if the frequency of natural fires can be determined by studies of sediment records preserved in lakes and peat bogs, it will 116 be possible to interpret this information in terms of past climatic conditions and to use the time frequency of natural fires as an indicator or index of paleoclimate. There has been also an increasing interest in. the place of natural fire in the environment over the past few years, paralleled by a growing perception of fire as a natural and necessary environmental factor rather than an unnatural “disturbance” that must be controlled and eliminated whenever and wherever possible. The presence of charcoal fragments is the primary evidence that natural fires leave in the sedimentary record, as indicated already by other studies (e.g., Heinselman 1971, 1973; Swain 1973: Rowe and Scotter 1973). The purpose of this study is to investigate the methods used for extracting fire frequency records from lake sediments, and the criteria for selecting appropriate sites for this kind of study. Lake sediments constitute a natural deposi- tory of environmental information that accumu- lates year by year throughout the life-span of a lake. These sediments contain inorganic (sand, silt, clay) particles and organic detritus (pollen, spores, diatoms, cladocera, various other algae and fungi, identifiable plant tissue fragments, and charcoal), as well as chemical residues, precipitates, break-down products, and stable compounds. All these components collectively STS reflect the environmental conditions within the lake as well as in the surrounding area. A stratigraphic study of these sediment com- ponents can establish a time series of ecological events from which paleoecological and environ- mental changes can be inferred on the basis of modern analogues (studies of recent limno- logical-environmental relationships). The study was designed to (1) explore the stratigraphic occurrence of charcoal in sedi- ments of selected lakes for the purpose of establishing a charcoal frequency record for postglacial (Holocene) time, (2) infer from this record the frequency of natural fires (forest fires), and (3) relate this information to the paleoenvironmental changes (including climate and vegetation) that have been established for the same time interval on the basis of palyno- logical studies of the same sediment cores. It was assumed that charcoal particles, origin- ating from forest fires, are transported to the lake by atmospheric turbulence (wind), possibly by surface water runoff, and direct falling of charred plant matter into the lake. Charcoal influx, contributed by all means of transport, would be further dispersed in the lake by surface currents and eventually settles to the bottom where it is incorporated in the accumulating sediment as an identifiable record of the fire. Location and Description of the Study Sites The study sites were four lakes (Figure 1) selected because of availability of background, palynological studies, radiocarbon dating, map- ping of surficial deposits, and the knowledge of postglacial history of the region. Furthermore, palynological studies of lake sediments and peat deposits made at numerous sites throughout Ontario by the senior author over a period of some 20 years (Terasmae 1973) had demon- strated common occurrence of charcoal particles in many palynological preparations. There seemed to be an apparent lack of published information from southern Ontario on charcoal occurrence in lake sediments and its relation- ship to the postglacial history of natural fires. Three lakes are in the Great Lakes — St. Lawrence Forest region (Found Lake, 45°30'65’N, 78°30'80”W; Perch Lake, 46°02’N, 77°21’W; and Boulter Lake, 46°09’15’N, 79°02’W) whereas Lac Louis (47°15’15’N, TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE Ly) 79°07’W) is in the southern part of the Boreal Forest region. The Great Lakes — St. Lawrence Forest is characterized by a mixture of deciduous hardwood tree species and some coniferous tree species, but coniferous tree species attain dominance in the Boreal Forest (Rowe 1959). Some descriptive data on the four lakes and their sediments are summarized in Table 1. All four lakes are different in terms of their geological characteristics. Found Lake is a bedrock basin, surrounded by steep rock slopes, and characterized by only minor inflow of surface and ground water. It is oligotrophic and has a low sedimentation rate (0.41 mm per year). Perch Lake is a depression in surficial deposits (mostly sand) and only a minor part of the shore is bordered by bedrock. It receives some ground- water and surface-water inflow through sur- rounding swamp and bog vegetation that filters out particulate matter, and a small creek flows out of Perch Lake to Ottawa River. It is about 8-10 km from the large sand dune area of the Petawawa delta. The lake is eutrophic and has an average sedimentation rate of 0.61 mm per year. Boulter Lake occupies an ice block depression (a “kettle hole”) in glacial outwash and esker deposits. It has no bedrock shoreline and probably receives some inflow of groundwater. There are no streams flowing into or out of the lake, and it is eutrophic; average sedimentation rate is 0.66 mm per year. Lac Louis (elevation 300 m) is located just below the highest shoreline (305 m) of glacial Lake Barlow-Ojibway in a sand and gravel deposit of an esker complex (Vincent 1973), and it is probably a “kettle” lake. It receives some inflow of groundwater and a small stream flows out of it. It is surrounded by Black Spruce ( Picea mariana) forest and muskeg, with some Trem- bling Aspen (Populus tremuloides) and White Birch (Betula papyrifera). The lake is eutrophic; average sedimentation rate 1s 0.59 mm per year. The lakes share a common characteristic, being essentially closed basins, without any major inflowing streams and with only minor outflow. Furthermore, all lakes are surrounded (except where bedrock forms the shore) by wetland vegetation (bog and swamp) that effec- tively prevents input of particulate matter by 118 THE CANADIAN FIELD-NATURALIST Vol. 93 1 Found Lake 2 Lac Louis 3 Perch Lake 4 Boulter Lake I | | | | | | Ville Marie 2 @ 4 ONTARIO QiViEBEG Qttowa Rive, eet @ Re Fo&smill \ o ALGONQUIN \ 3X PROVINCIAL \ \ PARK _~ —- [P 1 Georgian \ ) et Bay NaN \ BY \ balk River e SS Pembroke OTTAWA 100 kilometres FIGURE |. Index map, showing the location of study sites. Found Lake (site 1), Perch Lake (site 3), and Boulter Lake (site 4) are in the Great Lakes — St. Lawrence forest region, and Lac Louis (site 2) is in the southern part of the Boreal forest region. TABLE |—Some measurements and comparative data on the four lakes studies Found Lake Perch Lake Boulter Lake Lac Louis Elevation (m) 460 156 389 300 Maximum depth of water (m) 34 355) 8.5 7.6 Thickness of organic sediment cored (cm) 430 600 780 540 Surface area (km2) 0.32 0.45 0.25 0.10 Age of oldest organic 10400 + 300 9830 + 250 11800 = 400 9090 + 240 sediment (years BP) (I-7782) (GSC-1516) (GSC-1363) (GSC-1432) Average sedimentation rate (mm per year) 0.41 0.61 0.66 0.59 1979 surface water runoff. It is assumed, therefore, that charcoal particles found in the lake sedi- ment have been transported to the lakes pri- marily by atmospheric dispersal. The only exception would be charcoal input by water runoff from the small area of bedrock slopes adjacent to the lake. This assumption is sup- ported by the observation that many charcoal particles in the lake sediment are in the same size range as pollen (about 20 to 150 um) that is transported to the lakes by atmospheric disper- sal. All four lakes are also characterized by very small watersheds, effectively restricted to the surrounding slopes extending only 100-200 m away from the lake. Only Perch Lake has a larger catchment area that is occupied by wetland vegetation. Methods The lake sediment cores were taken with piston corers; the Brown sampler was used for very soft sediment at and near the water- sediment interface, and the Livingstone sampler for the remainder of the organic and inorganic sediments cored (Mott 1966), after a recon- naissance of lake basin morphometry (depth sounding) and the surrounding landscape had been made. Palynological subsampling of sedi- ment cores, laboratory preparations, and micro- scopic examination of slides followed well established procedures (Faegri and Iversen 1975; Kummel and Raup 1965). The study of charcoal presence and abun- dance was carried out in two different ways. In one case the palynological slides were used for counting of charcoal particles. The principal problem in this case is that samples for a palynological study (0.5 to | cm3) are normally taken at 10-cm (or sometimes 5-cm) intervals from the sediment core, and each sample covers sediment deposition during 10-20 yr, depending on sedimentation rate. This method of sampling obviously does not provide any information for the time intervals (about 100-200 yr) repre- sented by sediment deposition between the sampled intervals; therefore, it is possible that a significant part of the record of forest fires may be missed. To overcome this problem in the charcoal study, Weeks developed another preparation method, based on a technique used for sedi- TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE 119 mentary grain-size analysis by Perrie and Peach (1973) and Peach and Perrie (1975). The method involves two principal steps: preparation of gelatin-coated slides, and transfer of sediment onto the slide. The slides were prepared by first dissolving 1.5 teaspoons “Knox Gelatin” in 300 mL distilled water (the water-gelatin mixture was warmed in a water bath). Clean glass slides were then dipped in the gelatin coating solution (except the frosted ends, used for labelling). The excess gelatin was allowed to run off the slides, which were then placed in a dust-free cabinet to dry for a maximum of 6 h before storage in a clean slide box. The second step began with preparation of the sediment core for sampling. The sediment surface was carefully scraped clean of possible contamination with a spatula, and this surface was made smooth so that the full gelatin-coated area of the sampling slide came into contact with the sediment when the slide was touched or pressed against the core. It should be noted that the sediment cores, after they had been extruded from the coring tubes, were wrapped in plastic and aluminum foil and stored ina temperature-controlled room at 3-5°C to retain the natural moisture content, a normal practice for palynological studies. The sampling was carried out by pressing the gelatin-coated slide lightly to the sediment core: a thin layer of moist sediment sticks to the dry gelatin surface and a “fingerprint” of particles is. lifted off. The amount of pressure on the slide, required for producing a good “fingerprint,” depends on sediment characteristics and mois- ° ture content and this is determined experi- mentally by examining the slides under a micro- scope to make sure that the optimum amount of sediment is picked up by the slide. Too much sediment on the slide makes counting of char- coal particles difficult; too little sediment re- quires the counting of these particles on several slides, which is rather time-consuming. It is expected that further experimentation will lead to a more specific description of this aspect of the method. This sampling method covers the total length of the core without any gaps between samples. Cover slips were placed on the slides and a small amount of xylene was added to the edge of 120 the cover slip. The xylene dispersed throughout the preparation and rendered most of the organic detritus matrix translucent, whereas charcoal particles, pollen grains, and diatoms remained clearly visible. The ‘clearing effect’ of xylene made the identification and counting of charcoal particles much easier (Figure 2). Charcoal particles were identified with the aid of samples collected from sites of recent forest fires and prepared according to methods used in this study. Normally these particles are readily identifiable because some larger (commonly wood) fragments are only partially charred and others still retain characteristics of recognizable plant tissue structure. Some smaller charcoal particles are opaque (black) and differ from mineral grains with respect to fracture, refractive index, and surface characteristics. The common mineral grains are, furthermore, mostly trans- parent and lack cellular structure. The problem of small pyrite grains that occur in some lake sediments (even inside of pollen grains), how- ever, requires further investigation. The charcoal particles were counted in three different ways. In the first case, charcoal was counted together with pollen on slides prepared for palynological study. This technique provides a direct comparison of charcoal abundance with palynological data in the same preparation. In the second case the charcoal particles were counted on the gelatin-coated slides. The pur- pose of this technique was to relate charcoal abundance to sediment stratigraphy because each slide covers about 5cm of sediment deposition. In the third case, an attempt was made to use the Quantimet 720 (a TV image-analyzing computer) for counting of charcoal particles on the gelatin-coated slides. The advantage of this technique is that the instrument can be set to count particles in predetermined size classes, as described by Peach and Perrie (1975) in their study of varved glacial lake sediments. There was some difficulty, however, in having the Quantimet distinguish charcoal particles from other similar particles, and further experiments will have to be made to resolve this problem. For sediment cores from Lac Louis, Found Lake, and Boulter Lake the charcoal occurrence and relative abundance were tabulated on the basis of data obtained from continuous THE CANADIAN FIELD-NATURALIST Vol. 93 sampling by the gelatin-coated slides method. For Perch Lake the charcoal data were obtained by examination of the palynological slides and palynological data were adopted from available pollen diagrams (Boyko-Diakonow and Terasmae 1975). The charcoal data for the three other lakes (Lac Louis, Boulter Lake, and Found Lake) were obtained from the same sediment cores that had been used for palyno- logical study (Vincent 1973; R. J. Mott, personal communication; M. Boyko-Diakonow, personal communication), and a direct com- parison of both kinds of data was again possible. In Figure 3 the abundance of charcoal was calculated in the same manner as the relative pollen percentages of the different tree species; 1.e., the total number of pine pollen grains, for example, is expressed in the pollen diagram as a percentage of the total tree pollen count in a particular preparation (sample). The determination of sedimentation rate was considered an important aspect of the study of stratigraphic dispersal of charcoal in lacustrine deposits because the estimated (or assumed) sedimentation rate provides the time scale for chronostratigraphy of charcoal occurrence and abundance, unless some other means of time calibration is available (for example, when the sediment is annually laminated, or varved). Radiocarbon dates were available for all sedi- ment cores studied and these dates were used to calculate the approximate sedimentation rates that are shown in Table | for each of the lakes investigated. All tabulated data used as the basis of this paper are included in an unpublished report by Weeks (B.Sc. thesis, available from Department of Biological Sciences, Brock University). Results and Discussion Our observations indicate that charcoal does not occur in distinct layers with well defined boundaries in the lake sediments studied. Al- though the abundance of charcoal particles changes stratigraphically, the boundaries of each occurrence of these particles are diffuse, implying that at least some mixing of sediment has taken place during deposition and that charcoal dispersal (in decreasing amounts) may have occurred during several years after the fires that are recorded in the lake sediment. 1979 TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE 12] FIGURE 2. Photomicrographs of gelatin-coated slides from Boulter Lake sediment core: (A) untreated, (B) the same slide after clearing with xylene; and Found Lake: (C) untreated, (D) cleared with xylene. Arrows — charcoal particles; black triangle — pine pollen grain. 122 THE CANADIAN FIELD-NATURALIST Vol. 93 CUPRESSINE AE DEPTH (cm) aa a a Sa a I pr arene fener rel emcee fee pon fa IDO OR ZORS SONG 10 20 30 40 50 60 70 80 0 lOR ZO SOLOREOIO Om co 98307250 (GSC !516) POLLEN PERCENTAGES W@ CHARCOAL POLLEN a a >) Ee WwW {ea) DEPTH (em) GSC 1481 7280+250 POLLEN PERCENTAGES FIGURE 3. Abbreviated pollen diagrams from Perch Lake (A) and Lac Louis (B). The black areas indicate occurrence and abundance of charcoal particles, based on study of palynological slides (Perch Lake) and gelatin-coated slides (continuous sampling) from the Lac Louis sediment core. 1979 The diffuse nature of the charcoal layers in lake sediments has not been explained adequately by this study and the subject of modern charcoal dispersal and deposition in lakes requires more investigation. In the Perch Lake study all palynological slides contained some charcoal, but in view of the large sampling interval (about 10 cm) as compared with the annual sedimentation rate (0.61 mm) it is not possible to be certain of the actual stratigraphic frequency of charcoal oc- currences. Quite certainly many _ charcoal occurrences are not recorded by this type of study and the question of how much information was missed in the Perch Lake study will be investigated further. The Found Lake sediment core contained relatively little charcoal, both in terms of stratigraphic occurrence and abundance. On the other hand, the part of Boulter Lake sediment core (640-750 cm) that was examined in this study (because palynological data were available only for the basal 2 m of this core) contained abundant charcoal at frequent intervals. The Lac Louis sediment core showed a rather high frequency of charcoal occurrence with a non-uniform stratigraphic distribution and varying abundance. A pollen diagram for Lac Louis has been published by Vincent (1973). The sparseness of charcoal in the Found Lake sediment core can be attributed to at least two possible causes. One is the site susceptibility to fire, because the forest of the Found Lake area is dominated by hardwoods, mainly maple (Acer), beech (Fagus), and birch ( Betula). The second is related to the seasonal frequency distribution of thunderstorm activity. According to Chapman (1952) some thunderstorms in this area coincide with the period of fire susceptibility in the hardwood forest, but the greatest thunderstorm activity occurs in June, July, and August when the trees are in full leaf. Autumn fire hazard is reduced by the increased precipitation during that season, and Chapman refers to this north- ern hardwood forest as being the most nearly immune to fire of all forest types. These two causes may explain the paucity of charcoal in the Found Lake sediments during the last 7000 to 8000 yr but this reasoning becomes much weaker for the earlier postglacial time, when, on the basis of palynological TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE 8 evidence, the forest in this area contained more pine and fewer hardwood species some 8000 to 10 000 yr ago, and the forest was dominated by spruce prior to 10000 yr ago. Therefore, it is unlikely that vegetation alone was the factor in fire resistance in early postglacial time, although it may be now. Probably other factors, including topography, microrelief, and soil moisture are also important, and Found Lake may not be a good site for carrying out forest fire frequency studies. Charcoal particles were present throughout the Lac Louis sediment core, and the charcoal profile as shown in Figure 3 was based on the continuous sampling procedure, using gelatin- coated slides. Unfortunately the top and bottom 4 m of sediment core was not available for this study, and hence no charcoal profile is shown for these parts of the core in Figure 3. Interpolation of the radiocarbon dates for Lac Louis (reported by Vincent 1973) gives an age of about 8100 yr BP at the 480-cm depth of sediment. Within the 480 cm, there are 84 charcoal occurrences. If we assume that each is related to a separate fire event, there is one fire every 95 to 100 yr. This is probably an inaccurate estimate, however, because the distribution of charcoal is stratigraphically non-uniform, and a simple expression of mean frequency fails to express the fluctuations of fire frequency throughout the sediment core. For example, fire frequency in the pine zone (Pollen Zones 3 and 4) of the Lac Louis sediment core (Figure 3) is one fire every 48 to 56 yr — at least double the mean fire frequency for the whole core. Although only a part of the Boulter Lake © sediment core was examined for charcoal (640-750 cm), this part of the core contains the pine pollen zone (R. J. Mott, personal com- munication) and both a high frequency and abundance of charcoal. It seems that in Lac Louis, and probably in Boulter Lake, the high frequency and abundance of charcoal coincides with high percentages of pine pollen (the Pine Pollen Zone). For Perch Lake, the fire frequency, based on examination of pollen slides, is calculated to be one fire every 140 to 150 yr. This isa much lower fire frequency than one would expect, given the long history of fire which is evident from palynological data and an environment char- 124 acterized by sand dunes of the Petawawa delta and pine-dominated vegetation. This value also conflicts with the findings of Cwynar (1975) ata similar site, about 50 km to the west where the mean fire frequency in Greenleaf Lake area over the past 1200 yr is one fire every 83 yr. It seems reasonable to conclude that a number of charcoal occurrences have been missed owing to the distance between !0-cm sampling intervals. A study of Lake of the Clouds, Minnesota, relates forest vegetation succession over the past 10 000 yr to climatic change, and charcoal data to the changes in forest vegetation (Swain 1973). Swain reports an increase in fire frequency after 9500 yr BP, which coincides with a pine pollen peak and a climatic warming trend (the hypsi- thermal) which reached an optimum about 7000-6000 yr BP. Consistent high charcoal abundance occurred from 9000 to 6000 yr BP. Then a period of climatic cooling began, spruce and alder increased, White Pine ( Pinus strobus) decreased, and charcoal abundance became irregular from 6000 to 3000 yr BP. From 3000 to 1200 yr BP, a decrease in charcoal occurrence is associated with a similar decline in White Pine. The similarity between the pattern of climate, vegetation, and fire occurrence described by Swain at Lake of the Clouds and that reported by Vincent (1973) and this study at Lac Louis is rather striking. The same stratigraphic pattern of charcoal distribution seems to occur at both sites, but the Lac Louis dates are somewhat younger, possibly because of the difference in the time of deglaciation. The Lake of the Clouds area was deglaciated about 14 500 yr BP, but Lac Louis did not become ice free until about 10 500 yr ago. This implies that vegetation migrated northward with the receding ice and, therefore, radiocarbon dates from Lac Louis are about 7280 + 240 yr BP (GSC-1481) for the beginning of Pollen Zone 3 and 4260+240 yr BP (GSC-1491) for the end of this zone (the Pine Pollen Zone), compared to 9000 and 6000 yr BP for the Minnesota site. Both Lac Louis and Lake of the Clouds lie within the Great Lakes -St. Lawrence and Boreal Forest transition zone. There seems to be good potential for paleoclimatic reconstruction on the basis of charcoal occurrence in, and pollen analysis of lake sediment cores within, this zone. THE CANADIAN FIELD-NATURALIST Vol. 93 The gelatin-coated slide technique for charcoal analysis could be useful in such regional paleo- climatic studies. The results of the present study suggest that it will be possible to work out a postglacial forest fire frequency sequence for at least some regions, such as the deciduous forest and the boreal forest when study sites are selected with appropriate care. Geological site characteristics (topography, types of rocks and surficial de- posits, surface and ground water regime, glacial history and landscape features such as deltas, outwash plains, eskers, etc.) should be con- sidered with respect to fire susceptibility of the vegetation that grows on these different sites. For example, the Found Lake area seems to be rather resistant to forest fires: the Perch Lake area, in contrast, appears to have been especially susceptible to forest fires because of geological characteristics of the landscape that at least locally control the composition of forest. There- fore, these areas are less suitable for forest fire frequency studies than the Lac Louis area, or the Lake of the Clouds area in Minnesota. It appears possible that a fire frequency index can be worked out for Holocene lake sediment sequences and related to paleoclimatic condi- tions that have been inferred, in part, from other kinds of evidence such as palynology and the physical and chemical sediment characteristics. Acknowledgments We gratefully acknowledge the assistance of R. J. Mott of the Geological Survey of Canada who generously provided some sediment core material and some palynological data. We thank J. Simpson (superintendant), D. Brunton, and D. Strickland of the Algonquin Provincial Park for their help with the Found Lake study and for permission to use data from their files, including a pollen diagram compiled for Found Lake by J.H. McAndrews of the Royal Ontario Museum. Special thanks are extended to George H. La Roi for helpful criticism and many suggestions that were invaluable for the improvement of this paper. Partial financial assistance for this study was provided by the National Research Council of Canada (operating grant A5545 to J. Terasmae). 1979 Literature Cited Boyko-Diakonow, M. and J. Terasmae. 1975. Palynology of Holocene sediments in Perch Lake, Chalk River, Ontario. /n Hydrological studies on a small basin on the Canadian Shield. Edited by P. J. Barry. Atomic Energy of Canada, Ltd., Report AECL-5041-I, Chalk River, Ontario. pp. 189-220. Chapman, H.H. 1952. The place of fire in the ecology of pines. Bartonia 26: 39-44. Cwynar, L. 1975. The fire history of Barron Township, Algonquin Park. M.Sc. thesis, Department of Botany, University of Toronto, Toronto, Ontario. Faegri, K. and J. Iversen. 1975. Textbook of pollen analysis. 3rd edition. Blackwell Scientific Publications, Oxford. 295 pp. Heinselman, M. L. 1971. The natural role of fire in North- ern Conifer Forests. /n Proceedings Fire in the northern environment —a symposium. Edited by C. W. Slaughter, R. J. Barney, and G.M. Hansen. Pacific Northwest Forest and Range Experiment Station, Portland, Oregon. pp. 61-72. Heinselman, M. L. 1973. Fire in the virgin forests of the boundary waters Canoe Area, Minnesota. Quaternary Research 3: 329-382. Kummel, B. and D. M. Raup. (Editors). 1965. Handbook of paleontological techniques. W. H. Freeman and Com- pany, San Francisco. 852 pp. Mott, R. J. 1966. Quaternary palynological sampling tech- niques of the Geological Survey of Canada. Geological Survey of Canada, Paper 66-41, Ottawa. 24 pp. Peach, P. A.and L. A. Perrie. 1975. Grain-size distribution within glacial varves. Geology 3(1): 43-46. TERASMAE AND WEEKS: FIRES AS AN INDEX OF PALEOCLIMATE WZ) Perrie, L. A.and P. A. Peach. 1973. Gelatin-coated micro- scope slides in sedimentary size analysis. Journal of Sedimentary Petrology 43(4): 1174-1175. Rowe, J.S. 1959. Forest regions of Canada. Department of Northern Affairs and National Resources,. Forestry Branch, Bulletin 123. 71 pp. Rowe, J.S. and G. W. Scotter. 1973. Fire in the boreal forest. Quaternary Research 3: 444—464. Rowe, J.S., J. L. Bergsteinsson, G. A. Padbury, and R. Hermesh. 1974. Fire studies inthe Mackenzie Valley. Department of Indian Affairs and Northern Develop- ment, Northern Economic Development Branch, ALUR Program, Report ALUR 73-74-61. 123 pp. Rowe, J. S., D. Spittlehouse, E. Johnson, and M. Jasieniuk. 1975. Fire studies in the Upper Mackenzie Valley and adjacent Precambrian Uplands. Department of Indian Affairs and Northern Development, Northern Economic Development Branch, ALUR Program, Report ALUR 74-75-61. 128 pp. Swain, A. M. 1973. A history of fire and vegetation in N.E. Minnesota as recorded in lake sediments. Quaternary Research 3: 383-396. Terasmae, J. 1973. Notes on late Wisconsin and early Holocene history of vegetation in Canada. Arctic and Alpine Research 5(3) Part 1: 201-222. Vincent, J. S. 1973. A palynological study for the Little Clay Belt, northwestern Quebec. Naturaliste Canadien 100: 59-70. Received 10 November 1976 Revised and resubmitted 27 June 1978 Accepted 31 October 1978 Virgin Douglas Fir Forest on Saturna Island, British Columbia THOMAS P. SULLIVAN! Department of Botany, University of British Columbia, Vancouver, British Columbia V6T 1W5 'Present Address: Institute of Animal Resource Ecology, University of British Columbia, Vancouver, British Columbia V6T 1W5 Sullivan, T. P. 1979. Virgin Douglas Fir forest on Saturna Island, British Columbia. Canadian Field-Naturalist 93(2): 126-131. A survey of the Saturna Island Ecological Reserve provided a description of its virgin Douglas Fir ( Psewdotsuga menziesii) forest. The vegetation was quantitatively analyzed with respect to structure and composition. The tree stratum dominated the structure of this reserve, followed secondarily by the low-shrub and bryophyte strata. Psewdotsuga menziesii was the most important tree species with Tsuga heterophylla (Western Hemlock) and Thuja plicata (Western Red Cedar) probablylimited by low precipitation and soil texture. Gaultheria shallon (Salal) was the dominant understory species with Eurhynchium oreganum and Hylocomium splendens dominating the bryophyte stratum. Key Words: British Columbia, coniferous forests, ecological reserve, forest structure, old-growth forests, Pseudotsuga menziesii. Ecological reserves have been established in British Columbia primarily for scientific research and related educational purposes per- taining to our natural environment (Krajina 1973). Included within this proclamation (Eco- logical Reserves Act 1971) is the conservation of natural communities over a long period of time to prevent their alteration by man. The pre- servation of rare or endangered native plants and animals in their natural habitat is of paramount importance in this concept of con- servation. The selection of reserves has been based largely on the previous ecological classification of the province into biogeoclimatic zones. Saturna Island and its ecological reserve, composed of virgin coastal Douglas Fir forest, are located in the dry subzone of the Coastal Douglas Fir (CDF) zone (Krayjina 1965, 1969). An IBP (International Biological Programme) survey of the area was completed in 1968 by T.M.C. Taylor and T.C. Brayshaw (summary sheet of Region |, Area 4 in La Roiet al. 1976). The objective of this study was to provide a detailed ecological description of the virgin Douglas Fir forest on this reserve. This type of description is essential to the understanding and management of ecological reserves. Study Area Saturna Island, comprising approximately 28 km2, is the most southern Canadian island of 126 the Gulf Islands in the Strait of Georgia. It hes between 48°46’-48°49’N and 123°2’-123° 13’W. The Saturna Island Ecological Reserve is composed of two quartersections and encom- passes 1.3 km2. It has a range of elevation of from 150 to 320 m. Figure | shows the interior of this Douglas Fir forest. Weather data for this region are lacking. The study area is characterized by mild temperatures with prolonged cloudy periods, especially in winter, and a small range of temperature seasonally. Fairly wet but mild winters and warm dry summers are prevalent with a long frost-free season. Annual total precipitation is less than 90 cm because of the rain shadow effect of the Olympic Mountains and higher elevations on Vancouver Island. Methods Vegetation sampling was done during July and August 1972. The vegetation was analyzed by strata. The tree, shrub, and high-herb strata were sampled (restricted random sampling scheme) by 21 plots, 10 X 20 m each. The low- herb-dwarf-shrub stratum and bryophyte-lichen stratum were sampled by 210 subplots, each 1X 1m; 10 were placed systematically within each large plot. Data were obtained for diameters of all trees over 10 cm dbh (diameter at breast height) in the plots by species; heights of representative trees in plots using a Blume-Leiss altimeter; density of SULLIVAN: DOUGLAS FIR FOREST, B.C. 27) FiGuRE |. Interior of stand of trees representative of the virgin Douglas Fir forest on the Saturna Island Ecological Reserve. tree saplings (individuals <10cm dbh) by species; density of tree seedlings in subplots by species; percent cover estimates for all plant species in their respective plots and subplots using the coverage scale of Daubenmire (1959, 1968). A complete plant species list for the Saturna Island Ecological Reserve is available from Depository of Unpublished Data, CISTI, National Research Council of Canada, Ottawa, Canada KIA O0S2. Vascular plant nomenclature follows Hitchcock and Cronquist (1973), that of bryophytes follows Lawton (1971) and Stotler and Crandall-Stotler (1977), and that of lichens follows Hale and Culberson (1970). Results Importance values, absolute measurements, and diameter size classes for the tree stratum are presented in Table 1. The importance value is a summation of percentages of relative density + relative basal area + relative frequency (after Curtis and McIntosh 1951; Bray and Curtis 1957; Curtis 1959). Pseudotsuga menziesii has the highest importance value at 190 and the most consistent distribution of size classes throughout the stand. The average percent cover values for trees and saplings are presented in Figure 2. Douglas Fir trees are located on every plot and exhibit consistently high cover values through- out the reserve. This dominant species has a mean cover value of 52%. Unfortunately, it was not possible to obtain core samples from the protected trees on this reserve (regulations prohibit any disturbance of vegetation on ecological reserves), and hence there are no estimates of stand age. Average percent cover values for the three species in the low-shrub stratum and the two fern species in the high-herb stratum are in Figure 2. Gaultheria shallon (Salal) is distributed throughout most of the reserve and completely dominates the low-shrub layer. The average percent cover values for the most dominant species in the low-herb — dwarf- shrub and bryophyte strata are presented in 128 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE I—Importance values, absolute measurements, and diameter size classes.for tree species in the Saturna Island Ecological Reserve, based on measurements in 21 plots, 10 X 20 m each > 10cm dbh — size class distribution (cm) density-ha ! Importance Absolute Absolute Absolute Max value* density basal area frequency <10 cm ht. Species (0-300) (ha7!) (m2?.ha"!) (%) Seedlings dbh 10- 15- 20- 30- 41- l= 61- (m) Pseudotsuga menziesii 189.8 619 39.3 100 3.100 469 162 110 157 100 64 21 5 43.3 Tsuga heterophylla 63.5 154 8.7 62 6,600 429 38 40 33 21 17 5 0 35.1 Thuja plicata 46.7 123 6.6 43 900 374 50 19 38 2 7 5) 2 442 Total all species 300.0 896 54.6 — 10,600 1272 250 169 228 123 88 31 7 — *Importance value = Relative density + relative basal area + relative frequency (after Curtis and McIntosh 1951: Bray and Curtis 1957: Curtis 1959). Figure 3. Eurhynchium oreganum completely dominates the bryophyte layer with a mean coverage of 23%. Hylocomium splendens is the second most important species with a mean coverage of 5%. Discussion Pseudotsuga menziesii is the dominant tree species, constituting the uppermost layer of the forest canopy. This species is suitably adapted to all possible moisture and nutrient levels con- trolled by the soil and topography on this reserve. The highest productivity of the species Tree and High Shrub Stratum Pseudotsuga menziesii Isuga_ heterophylla Thuja olicata Low Shrub Stratum Gaultheria, shalion Berberis nervosa Rosa gymnocarpa High Herb Stratum Pteridium, aquilinum. Polystichum munitum in this dry subzone of the CDF zone is a site index of 45 to 48 mat 100 yr. Trees near the base of the slope on this reserve will probably attain this size class at 100 yr. Because this dry subzone is the least productive forest area of all the coastal parts of the mesothermal climatic region in British Columbia (Krajina 1969), Pseudot- suga menziesii does not achieve the growth reached by this species in more favorable regions. Tsuga heterophylla (Western Hemlock) is in low abundance and, in general. trees of this species are smaller than those of Pseudotsuga 30 40 . +50... GCiaae Average percent cover FIGURE 2. Average percent cover values for tree, low-shrub, and high-herb strata in the Saturna Island Ecological Reserve, based on estimates in plots and subplots. SS) Low Herb-Dwarf Shrub Stratum Festuca occidentalis Festuca sSubulata’ Galium aparine Elymus glaucus Trientalis latifolia Fragaria vesca Listera cordata Bryophyte Stratum Eurhynchium oreganum Hylocomium splendens Dicranum fuscescens Scapania nemorgsa Isothecium _stoloniferum Rhytidiadelphus triquetrus Hypnum circinale Lepidgzia reptans Rhizomnium glabrescens Lophocolea bidentata O NO SULLIVAN: DOUGLAS FIR FOREST, B.C. 129 aS 6 Average percent cover FIGURE 3. Average percent cover values for low-herb — dwarf-shrub and bryophyte strata in the Saturna Island Ecological Reserve, based on estimates in subplots. menziesii. Most Western Hemlock seedlings and saplings are growing on rotting wood compared with the soil or litter habitat characteristic of Douglas Fir seedlings. This low density and growth on decaying wood is typical of Tsuga heterophylla in the dry subzone of the CDF zone, which generally presents limiting factors for the growth of this species. It has a very high shade tolerance and prefers soils which supply low amounts of fairly well balanced nutrients, and these conditions exist only on podzolized soils frequently moistened by rainfall (Krajina 1969). Adequate precipitation (at least 165 cm annually) and a true podzol soil are not present on the ecological reserve, thereby limiting the success of Tsuga heterophylla. Thuja plicata is found downslope in the northern half of the reserve in association with Tsuga heterophylla and the dominant Pseudot- suga menziesii. It also occurs sporadically in clumps in local topographic depressions where some surface water is present in all seasons except summer. This species has smaller trees © than Pseudotsuga menziesii but does grow very well at lower elevations. The shade tolerance of Thuja plicata is as high as that of Tsuga heterophylla but its other ecological charac- teristics are quite different (Krajina 1969). Precipitation and soil texture, as they affect availability of water and nutrients, are probably limiting factors for Thuja plicata and Tsuga heterophylla in the Douglas Fir forest com- prising this ecological reserve. Pseudotsuga menziesii 1s well adapted to subhumid or even dry climates and if its moderate nutritional requirements are met, it tends to be very successful in both dominance and abundance, relative to other species in this study area. 130 The associations of the Coastal Douglas Fir zone have been studied and described by Mueller-Dombois (1959) with reference to the Nanaimo Lakes region and adjacent valleys. But there have been no plant community studies on this biogeoclimatic zone on the Gulf Islands. Three of the seven recognized associations of the Coastal Douglas Fir zone (dry subzone) are present on this ecological reserve. The Salal, moss, and Sword Fern (Polystichum munitum) associations are all represented. The Salal association (ca. 65% relative area) is most widespread and occurs with all three tree species. Herbs and bryophytes are suppressed owing to the presence of Gaultheria shallon. The moss association (ca. 25% relative area) alternates with Salal in a patchy distribution throughout the reserve. Eurhynchium oreganum and Hylo- comium splendens best represent this com- munity in their appearance as mats on the forest floor. The Sword Fern association (ca. 10% relative area) is probably best represented on this reserve at lower elevations. Here Pseudotsuga menziesii achieves its best growth in both height and diameter. Thuja plicata also follows this trend but is still the subdominant species. Polystichum munitum is not abundant in the forest but exhibits its highest cover value in this part of the reserve. A brief discussion of the probable history and future successional status of this forest is of importance to the understanding and manage- ment of this ecological reserve. There are a few isolated individuals of Pseudotsuga menziesii scattered throughout the forest, which are much larger and older than the majority of trees. Most of these trees, dead and alive, show evidence of fire damage. Also, traces of burnt logs and wood are distributed over the reserve. Therefore, at least part of the area has experienced past fire, which probably affected the growth of trees in the present-day forest. It is possible that the very large trees that survived the fire provided a seed source for regrowth of the Douglas Fir forest. Logging in the past is evident on the extreme eastern part of the reserve but is definitely absent from all other areas. The future forest will continue to be dom- inated by Pseudotsuga menziesii. Its seedlings are the most evenly distributed of all species and THE CANADIAN FIELD-NATURALIST Vol. 93 exhibit shade tolerance on all available habitats. The high density of seedlings and saplings of Tsuga heterophylla and Thuja plicata is due to clumping in restricted habitats, which contrasts with the widespread occurrence of Pseudotsuga menziesii. Both these species will remain restricted to specific sites in correlation with water and nutrient requirements. In the understory, the pattern of Salal, mosses, and open areas should continue to predominate, with suppressed herb and reduced shrub layers due to the shade cast by Gaultheria shallon. It is possible that Salal may expand and shade out regions of the moss association if soil moisture and topographic variation do not continue to support this Salal-moss distribution. Finally, the browsing of vegetation by Columbian Black-tailed Deer (Odocoileus hemionus columbianus) may become important if the population density of these animals increases. My sightings of deer during fieldwork indicated that these animals were very abundant throughout the reserve and on Saturna Island in general. The understory species, Gaultheria shallon, Rosa gymnocarpa, Pteridium aqui- lium, and = Polystichum munitum exhibited some degree of damage from browsing. Effects on tree seedlings were not noticeable, but according to J. Revel (1963 unpublished report, Faculty of Forestry, University of British Columbia) and Cowan and Guiget (1970), this species of deer prefers seedlings of Western Red Cedar and Douglas Fir. Western Hemlock is usually exempt from browsing. Therefore, the future successional status of this forest could be controlled by the effects of seedling browsing as well as the ecological tolerances already dis- cussed for the three tree species. In addition, deer trails on some downslope regions of the reserve have disturbed the ground cover of mosses exposed to the passage of these animals. Acknowledgments I thank Charles J. Krebs for reviewing the manuscript and Charles E. Beil and the Ecolo- gical Reserves Committee through V. J. Krajina for financial support. Many thanks are extended to Walter R. Kaiser and family for help with the fieldwork. I am most grateful to Nancy Turner and William Zales for identification of vascular plants and bryophytes. Ig Literature Cited Bray, J. R. and J.T. Curtis. 1957. An ordination of the upland forest communities of southern Wisconsin. Ecolo- gical Monographs 27: 325-349. Cowan, McT. I. and C. J. Guiguet. 1970. The mammals of British Columbia. Queen’s Printer, Victoria. pp. 366-369. Curtis, J.T. 1959. The vegetation of Wisconsin: an ordination of plant communities. University of Wisconsin Press, Madison, Wisconsin. pp. 63-83. Curtis, J. T. and R. P. McIntosh. 1951. An upland forest continuum in the prairie-forest border region of Wis- consin. Ecology 32: 476-496. Daubenmire, R. 1959. A canopy-coverage method of vegetational analysis. Northwest Science 33: 43-64. Daubenmire, R. 1968. Plant communities. Harper and Row, New York. pp. 37-96. Ecological Reserves Act. 1971. Statutes of the Province of British Columbia. Queen’s Printer, Victoria. 3 pp. Hale, M. E. and W. L. Culberson. 1970. A fourth checklist of the lichens of the continental United States and Canada. Bryologist 73: 499-543. Hitchcock, C.L. and A. Cronquist. 1973. Flora of the Pacific Northwest. University of Washington Press, Seattle. 736 pp. SULEIVAN SS DOUGIEAS FUR] ORESIi= BiG: 131 Krajina, V. J. 1965. Biogeoclimatic zones and classification of British Columbia. Ecology of Western North America 1: 1-17. Krajina, V.J. 1969. Ecology of forest trees in British Columbia. Ecology of Western North America 2: 1-147. Krajina, V. J. 1973. The conservation of natural ecosystems in British Columbia. Syesis 6: 17-31. La Roi, G. H., T. A. Babb, and B. J. Perley. 1976. Cana- dian National Directory of IBP areas. University of Alberta Press, Edmonton. Lawton, E. 1971. Moss flora of the Pacific Northwest. Hattori Botanical Laboratory. Nichinan, Miyazaki, Japan. Mueller-Dombois, D. 1959. The Douglas Fir associations on Vancouver Island in their initial stages of secondary succession. Volume, I, 2. Ph. D. thesis, University of British Columbia, Vancouver. Stotler, R. and B. Crandall-Stotler. 1977. A checklist of liverworts and hornworts of North America. Bryologist 80: 405-428. Received 27 June 1978 Accepted 2 November 1978 Colonial-nesting Herring Gulls and Common Terns in northeastern Saskatchewan HARRY A. STELFOX! and GREGG J. BREWSTER? 'Wildlife Research Division, Saskatchewan Department of Tourism and Renewable Resources, 2602 8th Street East, Saskatoon, Saskatchewan S7H 0V7 2Habitat Protection and Development Division, Saskatchewan Department of Tourism and Renewable Resources, 2602 8th Street East, Saskatoon, Saskatchewan S7H 0V7 Stelfox, Harry A. and Gregg J. Brewster. 1979. Colonial-nesting Herring Gulls and Common Terns in northeastern Saskatchewan. Canadian Field-Naturalist 93(2): 132-138. Observations of colonial-nesting birds were conducted during the summers of 1973 and 1974 as part of the Wildlife investigations of the Churchill River Study in northeastern Saskatchewan. Herring Gulls (Larus argentatus) and Common Terns (Sterna hirundo) were the only species found to breed in abundance within the surveyed area. All of the Herring Gull colonies and the majority of the Common Tern colonies were located on Reindeer Lake, rather than on the smaller lakes and river channels along the Churchill and Reindeer rivers. The mean size of nesting colonies was relatively small.compared to those of more southerly latitudes. Favored nesting sites were small rocky islands which were very susceptible to flooding. Common Terns appeared to be better adapted to utilizing the more ephemeral nesting habitat along the river systems than Herring Gulls. ~ Key Words: Common Terns, Herring Gulls, nesting colonies, habitat, behavior, Saskatchewan. The breeding range of the Common Tern (Sterna hirundo) extends throughout Saskat- chewan and that of the Herring Gull (Larus argentatus) covers most of the northern half of the province (Gollop 1969). This paper presents gull and tern nesting observations made during the summers of 1973 and 1974 while we were conducting wildlife studies in northeastern Saskatchewan for the Churchill River Study (CRS). The CRS was undertaken by the governments of Canada, Manitoba, and Saskat- chewan to assess the potential environmental impact of a proposed hydroelectric dam on the Churchill River 65 km west of the Manitoba- Saskatchewan border (Barber et al. 1975). The study area lies 300 km northeast of Prince Albert, Saskatchewan, and is characterized by Precambrian rock and boreal forest. The area investigated includes the Churchill River from Drinking Falls to the Island Falls Dam, Manawan Lake, all of the Reindeer River and most of Reindeer Lake (Figure 1). The primary objective of our gull and tern investigations was the location of nesting colonies to determine population levels and distribution. A secondary objective was an assessment of the nesting habitat. Methods High-level black-and-white air photos were examined to locate small isolated islands and shoals representing potential nesting sites. In late June 1973 an aerial survey was flown over most of Reindeer Lake to check these locations and numerous others for signs of nesting activity. Nest sites on the Churchill and Reindeer rivers were located during the course of other wildlife aerial and ground surveys. Once during late June or early July of each year a sample of nesting islands was examined on the ground to determine numbers of nests, eggs, young, and adults, as well as nesting habitat characteristics. Colonies on Reindeer Lake were examined between 19 and 23 June in both years. In 1974 elevations of islands and individual nests above water level were surveyed at 10 selected colony sites using a Cooke survey transit and stadia rod. Nest distribution and substrate characteristics were also noted. Results Common Terns Twenty-seven colonial nesting sites were located, 17 on Reindeer Lake, 4 on the Churchill River, 4 on the Reindeer River, and 2 on 132 1979 STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 133 @ Common Tern Nesting Colony A Herring Gull Nesting Colony ® Common Tern and Herring Gull Nesting Colonies on the same island Water bodies surveyed for colonial nesting birds wwe ose Ug h\s 3 SASKATCHEWAN Prince albert OSaskatoon ORegina VEOLINVW 6 Southend Horriot R. Ww? NMUMIHILVASYS FIGURE 1. Location of Common Tern and Herring Gull nesting colonies within the study area. 134 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE |—Number and average size of Common Tern and Herring Gull nesting colonies Number of colonies located (and sampled) Mean number (and range) of Mean number (and range) of adults per sampled colony nests per sampled colony Location 1973 1974 1973 1974 1973 1974 Common Terns Reindeer Lake 13(5) 15(6)* 23(10-45) 25(6—5S0) 12(4-26) 21(5-38) Reindeer River 2(2) 3(3) =" 34(18-—50) 55(35-66) 19(9-28) 25(11-33) Churchill River 4(3) flooded 28(8-45) = 14(9-22) = Manawan Lake 2(2) not surveyed 43(40-45) — 29(17-40) — Means 29 35 16 23 Herring Gulls Reindeer Lake 9(5) NG) 19(10-S5) 18(6—45) 13(2-31) 13(3-22) *Includes five colonies not sampled in 1973. **Includes two colonies not sampled in 1973. ***Includes four colonies not sampled in 1973. Manawan Lake (Figure 1). Not all colony sites were used in both years. In 1974, colony sites on the Churchill River were completely flooded asa result of unseasonally high water levels. Mean colony size was small, ranging from 16 nests in 1973 to 23 in 1974 (Table 1). The two colonies on Manawan Lake were relatively large, averaging 29 nests in 1973. Colonies examined on Reindeer Lake averaged only 12 nests in 1973 and 21 nests in 1974. By 21 June, most colonies had nests with eggs, with mean clutch sizes ranging from | to 2.8. On 20 June 1973, however, two colonies on Rein- deer Lake had several adult birds and nest depressions with no eggs, whereas a colony on the Reindeer River had the first young hatching on 23 June 1973. Reliable productivity data were not obtained because most sampled colonies were visited only once during the season. Some loss of eggs from flooding was observed, with low near-shoreline nests being washed over during storms and strong winds. Heavy rains which resulted in sharply rising water levels also took their toll. One colony on Reindeer Lake examined on 23 June 1974 had recently been destroyed by humans: 10 adult terns remained on the shoal but no re-nesting had been initiated. Herring Gulls Eleven different colony sites were located, all on Reindeer Lake. Four of these were shared with Common Terns (Figure 1). In addition, a single Herring Gull nest with eggs was at a Common Tern colony of Steephill Lake (Rein- deer River) in both 1973 and 1974, and lone Herring Gull nests were observed at four locations on Reindeer Lake, three of these being associated with Common Tern nesting sites. Colonies averaged 13 nests in 1973 and 1974 (Table 1). In 1973 most nests contained eggs, while in 1974 a large percentage of the nests attended by adults contained no eggs. Nest Island Characteristics All Herring Gull and Common Tern nesting islands were very low (<3 m above water level) and small (<1000 m’), except for one larger Herring Gull nesting island (Colony #10, see Table 2). Generally less than 50% of the surface of nest islands supported vegetation, primarily grasses, forbs, mosses and lichens, with an occasional shrub or tree. But Colony #10 in Table 2 and a Common Tern colony on the Reindeer River (Figure 2) had 70-80% of their surface area covered by shrubs and trees, whereas several islands had no growing vegeta- tion (Figure 3). The vegetation and small amount of associated soil occurred on the higher portions of nesting islands where there was no extensive erosion by ice and wave action. The substrate of nesting islands varied from large fragmented blocks to smoother solid rock and scattered boulders and stones. Occasionally some finer sand and pebble deposits were present near the water’s edge. Driftwood was strewn over the lower portions of some islands and completely covered one (Colony #4, Table )). ~ 1979 STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 135 FIGURE 2. Common Tern nesting colony on Steephill Lake (Reindeer River). EMSs SN Ds a \ ee SS FiGuRE 3. Herring Gull colony. 136 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 2—Island size and nest distribution for 10 Herring Gull and Common Tern nesting islands on Reindeer Lake, 1974 Nest height above water (m)** Island Island Colony height (m) area (m2) Species l 1.6 155 Common Tern 2 Dl 349 Common Tern Herring Gull 3 1.0 697 Common Tern Herring Gull 4 0.8 387 Common Tern Herring Gull 5 3.1 960 Common Tern Herring Gull 6 2.4 968 Common Tern Herring Gull 7 2.4 465 Herring Gull 8 2S) 426 Herring Gull 9 Dal 426 Herring Gull 10 2.0 3988 Herring Gull Means 1.8 586 Common Tern Dal 1083 Herring Gull Number of nests* Mean Range 5) 0.4 0.3-0.4 19 0.6 0.4-0.8 l 1.3 38 0.4 0.2-0.5 | 1.0 16 0.2 0.1-0.4 3 0.5 0.2-0.8 36 1.2 0.2-2.0 I 0.2 14 0.9 0.8-1.1 2D 0.8 0.5-1.1 22 0.5 0.1-0.8 6 0.5 0.3-0.8 10 0.8 0.5-1.4 13 0.5 0.2-1.1 0.7 0.7 *All recognizable nests, with or without eggs, were included in the measurements. ** All measurements were taken from 21 to 23 June 1974 when the water level on Reindeer Lake was 331.1 m (1103.8 ft.) above sea-level. Distribution of Nest Sites Common Tern nests were usually located near the lower edge of the vegetated portions of islands among scattered grass tussocks and moss-lichen mats. Nests were sometimes nothing more than a shallow depression in a thin layer of soil, or a crevice in the rock. Herring Gulls constructed nests of grasses and small twigs, frequently on exposed rocks or pieces of driftwood, affording a good vantage point. The mean height of both tern and gull nests above late June water levels in 1974 was 0.7 m (Table 2). No nests were located more than 2.0 m above water level. The mean nearest- neighbor distance between the edges of Common Tern nests was 0.9 m, with the closest nests being only 0.3 m apart. The mean distance between adjacent Herring Gull nests was 3.3 m, while the shortest distance was 2.1 m. Discussion and Conclusions Herring Gulls and Common Terns were the only species of Laridae found to breed in abundance within the study area. All of the Herring Gull colonies and most of the Common Terns were found on Reindeer Lake rather than the smaller lakes and river channels to the south (cf., Vermeer 1975). This distribution is thought to reflect the more stable nesting conditions on Reindeer Lake, where water levels are not subject to rapid large-scale fluctuations such as those that occur on the rivers. Common Tern colony locations and sizes on the Churchill and Reindeer rivers changed from 1973 to 1974 because of late spring flooding in 1974. McNicholl (1975) points out that in unstable habitats larids generally demonstrate reduced nest site tenacity and increased group adherence, thus aiding the rapid pioneering of alternate nesting habitat. The shifting nest site locations and relatively larger average size of Common Tern colonies that we observed in 1974 on the river system support this theory. We observed no evidence of Common Terns nesting singly, while we found five such cases in Herring Gulls. Several other cases of Herring Gulls nesting as single pairs have been reported in the literature (Harper 1953; Nero 1963; Vermeer 1975). In general our data would suggest that nest site tenacity is stronger in the Herring Gulls while group adherence is stronger in the Common Terns, thus enabling the terns more successfully to colonize riverine habitats. A noteworthy feature of our tern and gull nesting colonies is their small mean size. This appears to be characteristic for these more 1979 northern latitudes as indicated by other workers (Mowat and Lawrie 1955; Scotter 1961; Nero 1963, 1967: Weber 1976). No colonies of over 100 nests were reported for either species and most were considerably smaller. By contrast, Common Tern colonies in southern Saskat- chewan are characteristically large, ranging in size from 100 to 1000 nests (Ferry 1910; Lahrman 1957; Sanderson 1966; C. S. Houston, personal communication). In the Great Lakes, Common Tern colonies may number over 2500 birds, and Herring Gull colonies usually exceed 100 birds (Morris and Hunter 1976). Vermeer (1975) studied one Herring Gull colony of 161 nests at Kawinaw Lake in southern Manitoba. Several factors may account for such variation. Differential biological productivity and food availability is likely a primary factor (cf., Vermeer 1970). The waters in our study area are relatively nutrient-poor (oligotrophic) and have a cover of ice until mid- to late May. The size and distribution of nest islands, plus predation factors, may be important as well. Favored nesting sites of both gulls and terns were very low, small, rocky islands with restricted vegetation growth. Wind, wave and ice action are believed to be important elements in maintaining the exposed mineral substrates and sparse vegetation cover utilized for nesting. Nest sites thus occurred within a few feet of sur- rounding water levels and were very susceptible to flooding from both natural and artificial water level increases. A hydroelectric dam and reservoir as pro- posed by Saskatchewan Power Corporation at Wintego Rapids* on the Churchill River would result in water levels rising 28.5 m at Wintego Lake, 5.3 m at Steephill Lake, and 2.3 mat Keg Lake. All eight of the islands used for nesting by Common Terns on the Churchill and Reindeer rivers would be flooded; however, at least 11 new potential nesting islands would be created by the flooding. Reindeer Lake, which would be in- cluded within the proposed reservoir, would not exceed its historical levels, resulting inno change in numbers and locations of nesting islands. Once filled and operational, reservoir water levels would steadily rise a full 1.2 m from 1 May to | November to replace the winter drawdown. Much of this rise would come in June and July when runoff is heaviest (Hofer 1975). Our data STELFOX AND BREWSTER: COLONIAL BIRDS, SASKATCHEWAN 137) on the distribution of gull and tern nests (Table 2) suggests that on suitable nesting islands within the proposed Wintego Reservoir a 0).7-m in- crease in water levels during June and July would flood more than 50% of the gull and tern nests. Further studies are needed at these northern latitudes to understand more fully the factors influencing population distributions and pro- ductivity. It is necessary to have a better understanding of behavioral responses to changing environmental conditions, as well as food and nesting habitat requirements, before one can reasonably predict the possible impacts of future northern developments on colonial- nesting gulls and terns. Acknowledgments This study was funded by, and carried out for, the Churchill River Study under the direction of the Wildlife Research Division, Department of Tourism and Renewable Resources, Govern- ment of Saskatchewan. Valuable field assistance was provided by D. G. Hjertaas, D. G. McInnes, Re Wo Melinchuk: J. Es Polson. vand | Dsair Sherratt. We thank C.S. Houston for many helpful suggestions and critical review of the manuscript. *On 18 September 1978 the Government of Saskatchewan, following the recommendation of the Churchill River Board of Inquiry, announced that it would not permit construction of the proposed hydroelectric installation at Wintego Rapids. Literature Cited Barber, S. R., H. A. Stelfox, and J. D. Boden. 1975. Wild- life (Saskatchewan). Churchill River Study, Final Report 28. Saskatoon, Saskatchewan. 272 pp. Ferry, J. F. 1910. Birds observed in Saskatchewan during the summer of 1909. Auk 27: 185-204. Gollop, J.B. 1969. Birds in Saskatchewan. Jn Atlas of Saskatchewan. Edited by J.H. Richards and K. I. Fung. Modern Press, Saskatoon. pp. 85-90. Harper, F. 1953. Birds of Nueltin Lake Expedition. Ameri- can Midland Naturalist 49: 1-116. Hofer, R. D. 1975. Hydrology (Saskatchewan). Churchill River Study, Final Report 2. Saskatoon, Saskat- chewan. 61 pp. Lahrman, F. W. 1957. Birds of the Isle of Bays. Blue Jay 15: 106-110. McNicholl, M.K. 1975. Larid site tenacity and group adherence in relation to habitat. Auk 92: 98-104. 138 Morris, R. D. and R.A. Hunter. 1976. Factors influenc- ing desertion of colony sites by Common Terns (Sterna hirundo). Canadian Field-Naturalist 90: 137-143. Mowat, F. M. and A. H. Lawrie. 1955. Bird observations from southern Keewatin and the interior of northern Manitoba. Canadian Field-Naturalist 69: 93-116. Nero, R. W. 1963. Birds of the Lake Athabasca region, Saskatchewan. Saskatchewan Natural History Society, Special Publication Number 5. 143 pp. Nero, R.W. 1967. The birds of northeastern Saskat- chewan. Saskatchewan Natural History Society, Special Publication Number 6. 96 pp. Sanderson, R. M. 1966. The colonial birds at Suggi Lake, Saskatchewan in 1966. Blue Jay 24: 121-123. THE CANADIAN FIELD-NATURALIST Vol. 93 Scotter, G. W. 1961. Summer observations of birds in northern Saskatchewan, 1960. Blue Jay 19: 70-74. Vermeer, K. 1970. Breeding biology of California and Ring-billed Gulls. Canadian Wildlife Service, Report Series Number 12. 37 pp. Vermeer, K. 1975. Food habits and breeding range of Herring Gulls in the Canadian prairie provinces. Condor 75: 478-480. Weber, W. C. 1976. Birds of north-central Manitoba, 1973. Blue Jay 34: 84-94. Received 10 August 1978 Accepted 23 November 1978 Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario P. BARCLAY-ESTRUP! and RICHARD A. SIMS? 'Department of Biology, Lakehead University, Thunder Bay, Ontario P7B 5El 2Canadian Forestry Service, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario P6A 5M7 Barclay-Estrup, P. and R. A. Sims. 1978. Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario. Canadian Field-Naturalist 93(2): 139-143. White Elm (U/mus americana) stands near Thunder Bay, Ontario were investigated for epiphyte cover. Lichens dominated the cover in upper quadrats and mosses dominated in quadrats near the ground. In both upper and lower quadrats, mean cover was about 10%. Total cover was greatest on the north sides of the trees and tended to increase as the distance from the city increased. The stand located close to a kraft pulp and paper mill had the lowest number of species. Three species of liverworts, 12 mosses, and 17 lichens were recorded. Key Words: northern Ontario, White Elm, epiphytes, pollution. White Elm, U/mus americana, is rapidly disappearing from much of its former range in Canada and the United States owing to large- scale mortality caused by the introduced Dutch Elm Disease. This unfortunate circumstance has resulted in increased interest in studies relating to White Elm, such as that of Mahoney (1973) 1n northern Michigan and Newberry (1974) in Wisconsin. A further point of interest is that the groves in our region, the northwest shore of Lake Superior, form part of the northern boundary of White Elm’s range. The purpose of this study was to record the epiphytes on White Elm near Thunder Bay before Dutch Elm Disease removes the elm population, and to note epiphyte variability in relation to distance from the city of Thunder Bay. Methods During September and October of 1973, 50 trees were examined in five natural stands (10 trees per stand) located at various distances from the city of Thunder Bay. These stands were as follows: 1. Great Lakes Paper. On the south bank of the Kaministiquia River directly south of the Great Lakes Paper kraft pulp and paper mill. This small stand is on the south end of Thunder Bay city and is considered to be at 0 km (48°21’W, 89°19’N, 194 m above sea- level (asl)). 2. Pointe de Meuron. This relatively large stand of elm is on the north bank of the Kaminis- tiquia River adjacent to the reconstructed Old Fort William and is 3 km west of the Great Lakes Paper site (48°21’W, 89°21’/N, 195 m asl). 3. Slate River. This medium-sized stand is on the west bank of the Slate River just north of Highway 608 and is 16 km southwest of Great Lakes Paper (48°16’W, 89°29’N, 224 m asl). 4. Kakabeka Gorge. This is a small stand located on the east bank of the Kaminis- tiquia River about 300 m downriver from Kakabeka Falls (36 m high) and is 23.4 km west-north-west of Great Lakes Paper (48°24’W, 89°37'N, 229 m asl). 5. Blind Line Road. This small stand is on the south bank of a small creek just north of Blind Line Road and 250 m west of High- way 11-17. This site is 25.6 km west-north- west of Great Lakes Paper (48°25’W, 89°38’N, 284 m asl). All sites have Balsam Poplar (Populus bal- samifera), Black Ash (Fraxinus nigra), Moun- tain Maple (Acer spicatum), and Red-osier Dogwood (Cornus stolonifera). Also generally present are Eastern White Cedar (Thuja occi- —— dentalis), White Birch (Betula papyrifera), Speckled Alder (A/nus rogosa), and Red Rasp- berry (Rubus idaeus). Table | shows additional characteristics of the sites and of the trees in the stands. 139 140 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Mean soil and tree characteristics. DBH = Diameter at breast height (range in parentheses) Site Soil pH % Carbon Trees, DBH (cm) Bark pH Great Lakes Paper 5.9 11.0 44.9 (30.5-71.1) 4.2 Pointe de Meuron 5.9 13.1 37.7 (19.7-55.9) 6.2 Slate River 5.8 14.4 38.0 (22.9-49.5) 6.4 Kakabeka Gorge 6.8 5.3 24.8 (17.2-45.7) 6.5 Blind Line Road 6.4 18.8 21.3 (15.9-31.8) 6.6 Mean 6.2 IDES 381.3} OS 9=7 11 1b) 6.0 TABLE 2—Presence (X) and frequency of epiphytes on White Elm at the five study sites. Frequency figures are out of a possible total of 50 trees. ¢ = found only in top quadrats: b = found only in bottom quadrats: no notation = found in both top and bottom quadrats. GLP = Great Lakes Paper: PM = Pointe de Meuron: SR = Slate River: KG = Kakabeka Gorge: BL = Blind Line Road Presence Frequency Top Bottom Whole Species GLP PM SR KG BL quadrats quadrats tree Frullania eboracensis Xx Xx xX xX 28 15 31 Frullania bolanderi x 5 | 5 Porella platyphylla xe xe 0 2 D Anomodon minor x? x xX x 6 20 20 Orthotrichum speciosum x x! x 15 Fi 19 Brachythecium salebrosum x x xX 8 13 15 Pylaisiella selwynii Xx X X 5 7 1] Pylaisiella polvantha x! x 10 10 11 Brachythecium reflexum x xe x 3 3 3 Platydicta subtile xX | 5 5 Plagiothecium denticulatum Xe, 0) 2 2 Amblystegium juratzkanum x? 0 l l Orthotrichum obtusifolium x! | 0 l Dicranum montanum x 0 I | Mnium cuspidatum xe 0 I l Physconia grisea x xX xX x xX 30 23 33 Candelaria concolor xe Xx xX x! 28 10 29 Parmelia sulcata xX xX x! xX 21 9 21 Physcia orbicularis xX xX xX x 18 6 19 Physcia adscendens x x! x’ 11 3 12 Physcia aipolia xX x! Xx 7 3 9 Bacidia chlorococca* X 4 7 8 Lepraria membranacea xX 3 4 5 Lecanora impudens x X xX 3 3 4 Lobaria pulmonaria Xx x! Xx 4 l 4 Xanthoria polycarpa x! x! 4 0 4 Cetraria ciliaris x! 3 0 3 Lecanora conizaea xX 2 2 3 Evernia mesomorpha x! 2 0 2 Lepraria neglecta X | l 2 Parmelia saxatilis x! 2 0 2 No. of species, top 4 18 [2 12 ae) No. of species, bottom 6 16 9 10 13 XG=aMil No. of species per tree 6 20 12 15 IG SNe * Bacidia chlorococca, while not noted in the quadrats. was also present on the elms at both Slate River and Blind Line Road. 1979 Each of the 10 trees ineach stand was analyzed for epiphyte presence using the method em- ployed by Rao and Le Blanc (1967). In this method, two circular quadrats were used per tree. One quadrat (bottom) was from the tree base at 0 cm to 30 cm above the base; the second quadrat (top) was also 30cm deep and was located at 120 cm to 150 cm above the tree base. All mosses, liverworts, and lichens were recorded. Cover was also estimated using the Domin Scale for this purpose. Conversion of the Domin Scale data to percent cover was based on the following equivalents: 10 = 100%, 9 = 75%, 8 = 60%, 7=40%, 6= 30%, 5 = 20%, 4=5%, 3=3%, 2=1%, 1=0.5% and K =0.25%. The directional quarter of greatest cover for each tree was noted. Sorensen’s Coefficient of Similarity was used to compare quadrats and trees and also as a basis for a Bray and Curtis (1957) ordination in which each tree (top and bottom combined) was used as one quadrat. All field studies were done by the second author. Determinations were also done by the second author with assistance from C. Garton (bryophytes) and the first author (macro- lichens). All crustose lichens were verified by J.W. Thompson. Voucher specimens were SITES Great Lakes Point de Meuron Slate River Kakabeka Falls Blind Line Road | @ Uy ZO Wi@ 37% 3 A ‘ 4a 5 # FIGURE |. An ordination of whole trees from five sites using the Sorensen Coefficient of Similarity method of Bray and Curtis (1957). BARCLAY-ESTRUP AND SIMS: WHITE ELM EPIPHYTES 14] deposited in the Lakehead University Her- barium. Results The species of lichens and bryophytes present at each site, the number of species per tree, the number of species in top quadrats, the number of species in bottom quadrats, and the number of species per site in all quadrats are shown in Table 2. The total number of species for all sites is 32, made up of 3 hepatics, 12 mosses, and 17 lichens. The coefficient of similarity ordination of whole trees is illustrated in Figure | to show relationships between sites. Percent cover values by site are shown in Figure 2. When all five sites are considered 40 30 20 % COVER FIGURE 2. Cover relationships of bryophytes and lichens showing values for top quadrats (above line) and bottom quadrats (below line) at the five study sites. The sites are arranged in order of increasing dis- tance from Thunder Bay. L, lichens: B, bryophytes: T, total of both lichens and bryophytes: GLP, Great Lakes Paper site: PM, Pointe de Meuron site: SR, Slate River site: KG, Kakabeka Gorge site: BR, Blind Line Road site. 142 together, lichens contribute a mean cover of 5.9% in the top quadrats and 2.1% inthe bottom quadrats. Byrophytes contribute 3.6% in the top quadrats and 8.0% in the bottom quadrats. Overall mean epiphyte cover was 9.5% in the top quadrats and 10.2% in the bottom quadrats. In 68% of the trees the greatest cover 1s on the north quarter of the trees, 14% on the west, 10% on the east, 2% on the south, and in 6% two or more quarters were equal. Discussion The most obvious observation from the data obtained in this study is that the Great Lakes Paper site is quite different from the other four areas. The number of species of epiphytes in each of the three groups, and the total number of species for all groups, are significantly lower (P< 0.01) in the Great Lakes Paper site than in any of the other areas. This is perhaps even more significant when related to the mean DBH (diameter at breast height) of the stands, as Great Lakes Paper has the highest mean DBH of any of the areas, indicating greater age which often results in high epiphytic cover. The bark pH of 4.2 of the Great Lakes Paper site was much lower than on the other sites, which had a mean bark pH of 6.4. The emissions from the nearby pulp mill are probably responsible for the singular character of this site when compared to the other four sites. The mean bark pH of 6.0 is, incidentally, comparable to that found by Skye (1968) for U/mus sp. in the Stockholm region, and also for White Elm in Central Wisconsin (Newberry 1974). When all sites are considered together as representative of the Thunder Bay region, the number of species of lichens compares favorably with the number noted by Mahoney (1973) on White Elm in northern Michigan, where 15 species were found in one area and 8 species in another for a total of 17. Other publications that have included data on White Elm epiphytes are Hale (1952, 1955), Culberson (1955), Le Blane and De Sloover (1970), and Newberry (1974). For various reasons (e.g., data for elm are combined with that of other species: whole trees were investi- gated: trees were of urban environments) com- parison with all but Newberry (1974) are perhaps not very useful, except for the obvious con- THE CANADIAN FIELD-NATURALIST Vol. 93 clusion that generally the more trees and the larger the area of the tree included, the greater the number of epiphyte species that will be encountered. Newberry’s (1974) study, however, does give separate White Elm data and is related to sulphur dioxide emissions from a kraft paper mill. In Newberry’s study a species list is not included but 18 species and three varieties of lichens are common on White Elm and eight more lichen species are classified as rare (six more fruticose lichens are present but the phorophyte is not specified). Candelaria con- color, Parmelia sulcata, Physcia orbicularis, s.1., and Physconia grisea are apparently common in Thunder Bay as well as in northern Michigan and central Wisconsin. Physcia setosa, the most common lichen in the Michigan study, is rare in Thunder Bay and was not noted on the elms studied. Physcia elaeina, a ubiquitous species in Wisconsin, has not been reported in the Thunder Bay region. Parmelia rudecta and P. caperata were present in both the American studies, but although present in Thunder Bay, were not recorded on the elms investigated. One of the most common epiphytes in the Thunder Bay area 1s Hypogymnia physodes. This species, however, was not noted on the White Elms of this study or by Mahoney (1973) in Michigan and was reported as rare in Wisconsin, indicat- ing if not phorophyte specificity at least phoro- phyte preference. Although the bryo-flora was not considered by Mahoney (1973) or Newberry (1974) some general comments are perhaps warranted. None of the three liverworts found are rare. But, while Frullania eboracensis, the most common epi- phytic hepatic in this study, is also one of the most common in the Thunder Bay region the other two species are scarce. In the mosses, moreover, Anomodon minor, the most common moss on elm, is also scarce, and Hypnum pallescens, usually a very common epiphytic moss, was not recorded on any of the 50 elms studied. Table 2, besides showing the species present, clearly indicates that liverworts and lichens are more common in the upper quadrats and that the mosses are more common in the lower quadrats. The ordination (Figure |) based on presence- absence data also shows the Great Lakes Paper 1979 site to be rather different from the other four sites. The Kakabeka Gorge site is set off as well, but not to the same extent and the ordination indicates relationships among the Pointe de Meuron, Slate River, and Blind Line Road sites. A second ordination using the reciprocal averag- ing technique (Hill 1973) gave essentially the same results except that the Kakabeka Gorge stand is quite separate from the other groups. The cover relationships between groups are similar to the species numbers relationships, with bryophytes being greater in the lower quadrats and lichens greater in the upper quadrats. Cover relationships between sites, however, are quite different from species num- bers relationships. In Figure 2 the sites are arranged according to increasing distance from Thunder Bay City. Coincidentally, this is in decreasing order of tree DBH which probably represents decreasing mean age of stands as well. Increasing distance from Thunder Bay results in increasing cover, attributable at least in part to decreasing aerial poijlutants. The pattern is not as straightforward as this, however, as the very high bryophyte cover at Kakabeka Gorge must be related to the higher humidity levels resulting from spray from nearby Kakabeka Falls. In any event the Great Lakes Paper site has the lowest cover values and Blind Line Road, although it 1s apparently the youngest stand, has cover values equivalent to those of the larger-diameter trees at both Pointe de Meuron and Slate River. In conclusion it can be stated that the epiphyte flora of White Elms near Thunder Bay 1s relatively comparable to the floras of elms in Michigan and Wisconsin. Also, if Dutch Elm Disease, which was first reported near Thunder Bay in 1976 (Anonymous 1976), should become severe in these isolated stands of elms in this northern part of the tree’s range, some record is now available of the present epiphyte flora. BARCLAY-ESTRUP AND SIMS: WHITE ELM EPIPHYTES 143 Acknowledgments We thank W. M. Graham of the Lakehead Biology Department for useful discussions: J. W. Thompson, Department of Botany, Uni- versity of Wisconsin for determining a number of lichens: and the National Research Council of Canada for support under Grant Number A- 6121. Literature Cited Anonymous. 1976. Tree diseases. Forestry Bulletin, June— July. Great Lakes Forest Research Station, Sault Ste. Marie, Ontario. Bray, J. R. and J. T. Curtis. 1957. An ordination of the upland forest communities of southern Wisconsin. Eco- logical Monographs 27: 325-349. Culberson, W. L. 1955. The corticolous communities of lichens and bryophytes in the upland forests of northern Wisconsin. Ecological Monographs 25: 215-231. Hale, M.E., Jr. 1952. Vertical distribution of crypto- gams in a virgin forest in Wisconsin. Ecology 33: 398-406. Hale, M. E., Jr. 1955. Phytosociology of corticolous cryp- togams in the upland forests of southern Wisconsin. Ecology 36: 45-63. Hill, M.O. 1973. Reciprocal averaging: an eigenvector method of ordination. Journal of Ecology 61: 237-249. Le Blanc, F. and J. De Sloover. 1970. Relation between industrialization and the distribution and growth of epiphytic lichens and mosses in Montreal. Canadian Journal of Botany 48: 1485-1496. Mahoney, R. 1973. The lichen flora on Ul/mus americana in northern Michigan. Michigan Botanist 12: 163-166. Newberry, G. 1974. The influence of a sulfate-process paper mill on corticolous lichens. Bryologist 77: 561-576. Rao, D.N. and F. Le Blanc. 1967. Influence of an iron- sintering plant on corticolous epiphytes in Wawa, On- tario. Bryologist 70: 141-157. Skye, E. 1968. Lichens and air pollution, a study of cryptogamic epiphytes and environment in the Stockholm region. Acta Phytogeographica Suecica 52, Uppsala. ' 123 pp. Received 14 February 1975 Resubmitted 16 October 1978 Accepted 16 November 1978 Habitat Utilization and Population Densities of the Amphibians of Northeastern Alberta WAYNE ROBERTS and VICTOR LEWIN University of Alberta Museum of Zoology, Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 Roberts, W. and V. Lewin. 1979. Habitat utilization and population densities of the amphibians of northeastern Alberta. Canadian Field-Naturalist 93(2): 144-154. During 1976 and 1977 a survey of the herpetofauna of boreal forest habitats of northeastern Alberta was undertaken. The Wood Frog (Rana sylvatica) was the most abundant and widespread species and was collected at 24 of 25 sites examined. Boreal Chorus Frogs ( Pseudacris triseriata maculata) and Canadian Toads (Bufo hemiophrys) were common but found at less than half of the study sites. All three species had spawned prior to 12 June. Natural ponds and borrow pits were the most frequently used spawning sites. Canadian Toads also spawned in flowing water and lake margins. All species had metamorphosed by early August. Wood Frogs metamorphosed at a mean snout—vent length of 17.2 mm, Canadian Toads at 12.4 mm, and Boreal Chorus Frogs at 13.7 mm. Spawning for each of these species probably does not occur until individuals are in their third summer of life. Population densities of Wood Frogs, Boreal Chorus Frogs, and Canadian Toads reached estimated maxima of 19.6, 2.3, and 12 per 1000 m? during 1977. Maximum densities of all species were found within 50 m of the nearest water body in moist habitat vegetated by sedges, grasses, horsetails, willows, and poplar. Lower densities were found in upland mixed woods and no anurans were found in dry areas with sandy substrate and Jack Pine forest. Other amphibian and reptile species were not encountered. Key Words: habitat, population densities, boreal forest, Alberta, Wood Frog (Rana sylvatica), Boreal Chorus Frog ( Pseudracis triseriata maculata), Canadian Toad (Bufo hemiophrys). The herpetofauna of northeastern Alberta is poorly known. Only a few locality records for four species of amphibians and a single reptilian species exist for northeastern Alberta (Logier and Toner 1961). The Canadian Toad (Bufo hemiophrys), the Boreal Chorus Frog ( Pseuda- cris triseriata maculata), and the Wood Frog (Rana sylvatica) are known from the Fort McMurray area (University of Alberta Museum of Zoology = UAMZ collections). Little has been published regarding the abundance and life histories of these species within Alberta, especi- ally in the northern half of the province. Harper (1931) reports the occurrence of the Leopard Frog (Rana pipiens) and Red-sided Garter Snake (Thamnophis sirtalis parietalis) in the extreme northeastern corner of Alberta. The present study was undertaken to determine reproductive phenology, habitat utilization, and population densities of the three amphibian species commonly found in the boreal forest region of northeastern Alberta. Most amphibians and reptiles appear to have well defined home ranges (Porter 1972). Quanti- tative studies of home ranges have been con- ducted for the Wood Frog by Bellis (1965), the Chorus Frog by Kramer (1973, 1974), and the Canadian Toad by Breckenridge and Tester (1961). The tendency for individuals to spend extended periods of time at a given locality permits population estimates to be made by “total” counts, that is, the sum of individuals marked (counted) within a given area. Such counts may be higher than the actual number present within an area at a given time. This overestimate is the result of (1) counting all (or nearly all) true residents plus a number of transients; (2) counting individuals whose home ranges do not lie completely within the study plots; (3) mortality not being taken into account. The studies by Breckenridge and Tester (1961), Kramer (1973, 1974), and Bellis (1965) were conducted in relatively small areas and thus provide a limited amount of comparative infor- mation with respect to habitat preferences. Information on habitat utilization can be ob- tained by conducting regular periodic censuses of a large number of sample sites selected to include a wide range of habitat types. Study Area Twenty-four potential spawning sites (Figure 1) were selected in 1976 and 1977 within the boreal forest region of northestern Alberta. These sites consisted of 12 streams, five borrow pits (excavations resulting from the removal of 144 1979 ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS. ALBERTA Peace River Lake oo Athabasca Claire Wood Buffalo National Athabasca River Edmonton, 520 miles 20 30 km Birch sete: Mountains Ells River Gregoire \ \ J SS t “4 6 8 9 loose Ss. oe + rks ¢/ HZ FIGURE |. Location of amphibian study sites in northeastern Alberta. 145 146 road building materials), four natural ponds, portions of the shores of two lakes, and a bog. During 1977, 16 of these sites plus a dry Jack Pine (Pinus banksiana) ridge were studied to determine habitat utilization and population density. Methods Each site was visited during the day at least eight times during June, July, and August and the amphibian species present, and their abun- dance, were recorded in the following manner. The visits were equally distributed during morn- ings and afternoons and as well between cool and warm days for each site. During each visit the observer walked completely around the margins of ponds and borrow pits or fora fixed distance (approximately 400 m) along the mar- gins of lakes or flowing waters. Amphibians . observed were identified to species and the numbers of individuals of each were recorded. Individuals were assigned to developmental stages (if larvae) or size classes (if terrestrial) in order to describe their development, time of metamorphosis, and growth during their first year of life. The following arbitrary classes were employed, with equivalent stages of Gosner (1960) given in brackets: (1) eggs (1-25); (2) tadpoles without limbs (26-31); (3) tadpoles with hind limbs present (32-40); (4) tadpoles with four limbs present (41-42); (5) transforming tadpoles (semi-aquatic or terrestrial with tail still evident) (43-45); (6) young-of-the-year with tail completely resorbed (46); (7) yearlings (the smallest size class of subadults or adults); (8) adults (large subadults and adults). The smaller number of classes or stages distinguished by relatively gross differences facilitates the des- cribing of large numbers of larvae in the field. The head-body lengths (= length) of post- metamorphic frogs and toads were measured to the nearest 0.1 mm according to Conant (1975). Small subadults were similarly measured, up to and including the time of metamorphosis of young-of-the-year. Young-of-the-year were measured again in late August. The data should provide objective criteria for distinguishing underyearling (class 6), yearling (class 7), and older (class 8) individuals, as preliminary obser- vations here indicate that these groups occur in mutually exclusive length-frequency groups. THE CANADIAN FIELD-NATURALIST Vol. 93 Subsamples of aggregations of larvae were collected and preserved in 10% formalin. These were later identified to genus using keys in Blair et al. (1968). Specimens of each species and their developmental stages from each site were col- lected throughout the study and deposited inthe University of Alberta Museum of Zoology (Accession number 76-31). At each habitat sampling site three plots, 10 X 100 m, were established at right angles to the water body (if present). Where more than one habitat type was present the plots were established so that all types were sampled. Where a water body was present the following information was recorded: basin morphology, submergent and emergent vegetation present. Owing to the large area encompassed by the sample plots only the dominant vegetation types were mapped for each plot. A census of each of the sample plots was taken 10 times during the summer of 1977. During each visit the observer noted weather conditions, walked the plot, traversing its width 20 times going away from the pond edge and 20 times on the return trip to the pond ina crisscross pattern. During each visit the species present, age class (as determined by length), location on the plot, and habitat type with which each individual was associated was recorded. Subadults and adults were marked by removal of the phalanges distal to the web on one digit of the left hind foot: thus they were not individually marked. The popula- tion of adults on each plot was regarded as the total number of individuals so marked. The sample plots were subdivided by orange surveyor’s tape placed at 20-m intervals. Indi- viduals found between the water margin and the 20-m marker were assigned the mean value of 10 m, and individuals found between 20- and 40- m markers were assigned the mean value of 30 m and so on for purposes of calculating the mean distance that each species was found from the water margin during each month of the study. Results and Discussion During the 1976 survey 478 adult and sub- adult amphibians were counted of which 225 were Wood Frogs, 207 Canadian Toads, and 16 Boreal Chorus Frogs. No spawning pairs or egg masses were found during the present study. IY) Tadpoles were present in all of the known spawning sites during mid-June. Abundance, Spawning, and Development Wood Frog The Wood Frog was present at all sites and spawned in 14 of the 24 sites studied. This species spawned in natural ponds, borrow pits, and a backwater on a stream. Egg masses of this species are attached to aquatic vegetation (Wright and Wright 1949): thus Wood Frogs are most likely to spawn in standing water where suitable vegetation may be found. Spawning had apparently been completed before the 1976 study was initiated in mid-June. This species spawns as early as Marchin the southern portion of its range but as late as July in the Northwest Territories (Wright and Wright 1949). Within the study area metamorphosis occurred as early as 30 June and was complete by early August. The period during which metamorphosis occur- red within the study area was exceptionally short. Metamorphosis elsewhere may occur from late May to mid-September (Wright and Wright 1949) and overwintering of tadpoles may occur at northern latitudes (Bleakney 1954). Wood Frogs metamorphose at a size of 14-22 mm_ head-body length (Wright and Wright 1949). Postmetamorphic individuals in the present study ranged from 14.4 to 18.7 mm CsI pe NE= 918) and trom: se) ton24-0, mim (x = 20.5, N=12) by late August. Yearlings ranged from 20.4 to 31.0 mm (x = 25.4, N = 13) in late June and thus can be readily dis- tinguished from young-of-the-year. Yearlings apparently do not spawn, as frogs of this size were not found in breeding aggregations. The length of spawning individuals ranges from 29 to 50 mm for males and 34 to 56 for females throughout the range of this species (Wright and Wright 1949). Individuals of this size within the study area are almost certainly two or more years old. Schueler (1973) also found that frogs of this species did not breed until they were two or more years old. Dispersal of post-spawning adults away from the spawning sites occurs but some adults may be encountered near the margins of water bodies throughout the summer. Young-of-the-year also leave the spawning sites shortly after meta- morphosis and only small numbers were en- ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 147 countered at these sites during late August. Wood Frogs are terrestrial and are occasionally found far from water in moist wooded areas. They are known to overwinter under leaf litter (Wright and Wright 1949; Hodge 1976) and under logs or stumps. Boreal Chorus Frog The Boreal Chorus Frog was seen at seven of the study sites and spawned at three of these. This species spawned in two of the borrow pits studied and in one natural pond. The eggs are laid in small loose clusters attached to vege- tation, frequently in temporary ponds (Wright and Wright 1949). Spawning within the study area had been completed prior to 12 June 1976. Spawning is known to occur as early as late March in Indiana (Wright and Wright 1949), and as early as late April at Lake Wabamun, Alberta. Calling individuals have been heard as early as 3 April (1976) along the Red Deer River near Innisfail (52°N) (personal observations W.R.). Spawning was observed in the north- western quarter of Alberta in late May of 1976 and likely occurred within the study area at that time. Metamorphosis occurred as early as 24 July and was completed shortly after 3 August. As with the Wood Frog, metamorphosis of this species occurred within a short period. Boreal Chorus Frogs throughout their range meta- morphose at lengths of 7.5-13.0 mm (Wright and Wright 1949). Postmetamorphic individuals measured during the present study were much larger, 12.6-14.8 mm (x = 13.7, N = 14). Other northern Alberta specimens (N = 18, UAMZ collection) averaged 12.7 mmat, or shortly after, | metamorphosis. The relatively large size of Boreal Chorus Frogs at the time of metamor- phosis may be the result of development at low temperatures as is true for many poikilotherms (Ray 1961). The size at which young-of-the-year overwinter may be a critical factor affecting their ability to persist during periods of stress. Larger individuals may be more successful in this regard. Yearlings in a number of northern Alberta collections ranged from 15.9 to 17.3 mm during May. Spawning adults, taken from breeding aggregations, in Northern Alberta ranged from 19.3 to 28.2 mm (Xx = 23.0, N = 28). The size of spawning individuals throughout the range of this species ranges from 21 to 32 mm for 148 males and 20.0 to 37.5 mm for females (Wright and Wright 1949) indicating that individuals within the study area probably do not spawn until they are two or more years old. It is of interest that Matthews (1971) reports that marked individuals in a montane population do not breed until their fourth summer. Boreal Chorus Frogs are known to overwinter in Alberta by burrowing under decaying stumps and into anthills (personal observations W. R.). Canadian Toad The Canadian Toad was present at nine of the study sites during 1976 and spawned at seven of these. This species utilized the widest variety of spawning habitats including natural ponds, borrow pits, streams, and lake margins. The eggs of Canadian Toads are laid in long strings and may or may not be associated with vegetation (personal observations W. R.). The long strings of eggs may be less subject to displacement by moving water than the attached spherical or subspherical egg masses of frogs and thus facilitate spawning in waters with a slight current, wave action, and no vegetation. This increases the variety of spawning habitats available to the species; however, egg and tadpole mortality may be higher owing to predation and displacement by running water. The fecundity of Canadian Toads (3354-5842 eggs, N =3 (V. L. unpublished data)) is higher than that of Wood Frogs (x = 777 eggs (Herreid and Kinney 1966)) and Chorus Frogs (x = 597 eggs (Whitaker 1971)), which may compensate for the mortality arising in the wide variety of spawning habitats utilized. These strings are also adaptive as some eggs will usually remain submerged if the water level recedes. Spawning within the study area had apparently been completed prior to 12 June. Spawning com- menced during May along the Red Deer Riverin 1976 and had occurred into mid-June at the Same site in previous years. Within the study area metamorphosis occurred as early as 25 June and continued until early July. The develop- mental time for this species appears to be unknown. Postmetamorphic individuals of this Species range from 9.0 to 13.5 mm in head-body length (Wright and Wright 1949). Individuals sampled during the present study ranged from IS*to 13.2 mm (x = 1274- N= 11) and had at- tained head-body lengths of 19.3-28.0 mm THE CANADIAN FIELD-NATURALIST Vol. 93 (xX = 24.8, N = 10) by late August. The smallest yearling found in late June was 22.0 mm long; thus yearlings are readily distinguishable from young-of-the-year. Spawning individuals of this species range from 58 to 68 mm for males and 56 to 80 mm for females (Wright and Wright 1949): it is unlikely that individuals of this species within the study area spawn until they are two or more years old. Dispersal of adults from spawning sites occurs soon after spawning but individuals may be found along the margins of these sites through- out the summer. Young-of-the-year toads are often found in high numbers along the margins of spawning areas throughout the summer. Lillywhite and Wassersug (1974) regard the aggregating behavior of young-of-the-year Bor- eal Toads (Bufo boreas) as a retained larval characteristic. Young-of-the-year Canadian Toads were abundant along the Red Deer River (a spawning site) in early September of 1976 but were absent after mid-September, probably having buried themselves in suitable substrate for overwintering. Habitats and Population Densities Nine habitat types were identified within 17 study sites. These are ranked in order of decreasing wetness as follows: (1) water, (2) sedge (Carex spp.) fen, (3) grass (Graminae) meadows, (4) willow (Salix spp.) bog, (5) Trembling Aspen (Populus tremuloides) forest, (6) Black Spruce (Picea mariana), (7) White Spruce (Pica glauca), (8) upland mixed wood (Populus tremuloides, Picea mariana, _ P. glauca), (9) Jack Pine (Pinus banksiana). Because visits to each site were equally divided between mornings and afternoons and pro- portionately between cool and warm days, any biases in the number of amphibians observed resulting from these factors were minimized. In fact 28.6% of the amphibians were counted on cool mornings (23.7% of the visits) and 14.4% were seen on cool afternoons (14.8%): 34.7% were seen on warm mornings (33.1%), and 22.3% were seen on warm afternoons (28.4%). Wood Frog The extent to which each habitat type was utilized by each age class of the Wood Frog during May—August of 1977 is shown in Figure 2. Wood Frogs were most frequently found in 1979 KEY HABITAT TYPES: | Water 2 Sedge Fen 3 Grass Meadow 4 Willow-bog 5 Aspen Poplar 6 Black Spruce 7 White Spruce 8 Upland Mixed Wood 9 Jack Pine Dry SYMBOLS} 15 | | Adults (N=15) 10 Ne Yearlings (N=10) Z Young-of-year (N=5) rn) Ww a = 5) 2 z = 2 = a 2 < 5 O ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 149 WOOD FROG FIGURE 2. Habitat utilization by Wood Frogs in northeastern Alberta during May—August 1977. grass meadows, willow bogs, and aspen poplar habitat, which are the most moist terrestrial habitats available. Heatewole (1961) regards the Wood Frog as a forest-dwelling species and suggests that the shade provided by trees is an important factor in determining habitat selec- tion. He demonstrated experimentally that substrate structure and substrate moisture are important factors governing the selection of microhabitat. Bellis (1962) states that Wood Frogs are restricted to forested areas, particu- larly lowland bogs. Shelford (1913) demon- strated that Wood Frogs prefer an environment with moist air. Marshall and Buell (1955) found that in exposed areas with sparse Wood Frog populations the vapor pressure deficit was higher than in shaded tamarack (Larix /arcinia) habitat with a dense Wood Frog population. It thus appears that moisture content of both the substrate and air are important factors in determining the suitability of habitat types for Wood Frogs. Tall grasses, willows, and aspen poplars growing in moist soil in low-lying areas provide a favorable environment for Wood Frogs within the study area, while much of the forest habitat, especially Jack Pine area, is too dry with respect to substrate moisture and perhaps the humidity of the air. Heatwole (1961) suggests that the Wood Frog is less terrestrial than previously thought (see Wright and Wright 1949) and points out that its association with woods and damp leaf litter occurs after woodland ponds dry up. Our study sites were associated with more permanent bodies of water. Wood Frogs were rarely found at distances greater than 100 m from a water body and the mean distance for all age classes during each month was less than 50 m. Bellis 150 (1962) regards the Wood Frog as intermediate between aquatic and terrestrial anurans in its ability to withstand body water loss. Only large Wood Frogs are found in dry habitats and it is probably the large individuals that emigrate and colonize new areas (Bellis 1962). We also noted that larger Wood Frogs tended to be further from water, perhaps owing to their greater ability to tolerate body water loss. During 1977, 646 Wood Frogs were counted. Population densities of this species ranged from 0.0 to 19.6 individuals (yearlings or older) per 1000 m2 (Table 1). The highest population density for this species (19.6 adults per 1000 m2?) was found in an area with a mixture of grasses, willow, and aspen poplar. These plant communi- ties are frequently found individually, mixed or intergrading with one another adjacent to water bodies. Bellis (1961) found maximum densities of about 17 individuals per 1000 m2 in Tamarack and Calla Lily (Calla palustris) habitat. Boreal Chorus Frog Boreal Chorus Frogs were most frequently found near the water margin or in shallow water amongst vegetation (Figure 3). Too few year- lings or adults were found to determine what, if any, habitat type is preferred. Whitaker (1971) regards the Chorus Frog as a woodland species that occurs most frequently near water, while Stebbins (1954) notes that in the west this species seems to be primarily an inhabitant of open moist grasslands. Kramer (1974) found that the TABLE |—Densities of amphibians in selected habitat types within the study area. Numbers in parentheses indicate maximum estimates Density per 1000 m2 Boreal Habitat Wood Chorus Canadian types Frog Frog Toad Water Sedge Willow (2.3) 0.45 Grass (19.6) 2.83 (12) 0.83 Horsetail Poplar White Spruce Black Spruce 1.08 0.07 0.16 Upland mixed j wood Jack Pine 0 0 0 THE CANADIAN FIELD-NATURALIST Vol. 93 home ranges of Chorus Frogs contained breed- ing pools and that most frogs remained within 100 m of these pools throughout the summer. In Alberta the Boreal Chorus Frog is known from the prairies, aspen parkland, and boreal forest regions (UAMZ collections), but always near water. It is the most widespread amphibian species within Alberta and persists in a wide variety of moist habitat types providing that at least temporary ponds are available for spawn- ing and for development of the tadpoles. Boreal Chorus Frogs were found closer to water on the average than the other anuran species present. Adults and young-of-the-year were found at a mean distance of less than 30 m from water during all months. During June of 1977 nonbreeding individuals were found at a mean distance of 54 m from the water margin. This may be the result of their being found close to their overwintering sites. Unlike breeding individuals they probably do not move immedi- ately to water bodies after emergence in the spring. During July and August most indi- viduals of all age classes were found within 20 m of the water margins. Kramer (1973) notes that while a few individuals may be found over 200 m from breeding pools, most are found within 100 m. Individuals may be found buried under leaf litter during the day (Kramer 1973). This may serve as protection against diurnal preda- tors and desiccation. These observations may indicate that this small species is more prone to desiccation than are the larger species. During 1977, 148 Boreal Chorus Frogs were counted. Insufficient data were obtained to determine population densities on most study plots. Two sites yielded adult population densi- ties of 2.3 individuals per 1000 m2 (Table 1). Kramer (1974) obtained densities of 0.55 adults per 1000 m2 from a 33-acre (13.4-ha) study area in Indiana. His data probably include most of the adults present; estimates from the present study, on the other hand, are probably con- servative owing to the secretive nature of the Boreal Chorus Frog. Larger numbers of the Boreal Chorus Frog are assumed to be present because of the numbers heard calling (but seldom seen) during May and June. Canadian Toad The Canadian Toad was found most fre- quently in grass meadows and willow bogs near 1979 BOREAL CHORUS KEY HABITAT TYPES: | Water Wet 2 Sedge Fen 3 Grass Meadow 4 Willow- bog 5 Aspen Poplar 6 Black Spruce 7 White Spruce 8 Upland Mixed Wood 9 Jack Pine Dry SYMBOLS: 15 - | | Adults (N=15) uy b:] Yearlings (N=10) 6 Z Young -of—year (N=5) ” a Ww a = =) Zz z = es) ce a = ) z z =< = x a = <= 2308 6 7B © M AY Vol. 93 CANADIAN TOAD Bernd DOQQ iS 73 © AUGUST HABITAT TYPES FIGURE 4. Habitat utilization by Canadian Toads in northeastern Alberta during May—August 1977. however; individuals were seldom found suf- ficiently far upland that they were in particularly dry habitat. During 1977, 130 Canadian Toads were counted. Population estimates for this species reached a maximum of 12 adults per 1000 m2 (Table 1), only slightly over half of the maximum density for Wood Frogs. Kelleher and Tester (1969) found Canadian Toads in much greater abundance. This was probably owing to an abundance of excellent overwintering habitat combined with a lack of competition from other anurans along the margins of a prairie lake. General Discussion The relative abundance of the three species found during both years of the study may be misleading. The Wood Frog is certainly wide- spread and abundant within the study area. This species is the most widely distributed amphibian in North America and occurs further north than any other species. Its adaptations to life in cold regions are summarized by Hodge (1976). Canadian Toads, however, may be much less abundant than the Wood Frog, as 182 of the 207 adult and subadult Canadian Toads seen during 1976 were young-of-the-year. These were found in postmetamorphic aggregations along the mar- gins of spawning sites. In spite of its ability to use the widest variety of spawning habitats (among the three species found here) this species spawn- ed in only 7 of the 24 sites studied compared with 14 sites utilized by the Wood Frog. The Boreal Chorus Frog was heard at or near all of the study sites but spawned in only three of these. Its utilization of temporary ponds and standing bodies of water for spawning sites may reduce the number of potential sites from 24 to 10 for this species. The small size, cryptic coloration, and secretive behavior of Boreal Chorus Frogs 1979 ROBERTS AND LEWIN: AMPHIBIAN POPULATIONS, ALBERTA 153 CANADIAN TOAD 2 1000m 9 ine) ::Subadults: 2222) ° Individuals / (eo) Wiilow Sedge, Grass = < ud ac | ae w” ac are) = ao = fe) ac ac fe) rr) fay z Oo a. Horsetail Cattail 60 CHORUS FROG WOOD FROG Upland Jack Pine Mixed Sand Ridge DISTANCE FROM WATERIN METRES FIGURE 5. Schematic distribution and population density of adult and subadult amphibians in northeastern Alberta in relation to vegetation and water. make them difficult to see and collect even when they are present in large numbers at spawning SiLeSee SINCE teas) sae stenhesthial species. and frequently hides under logs and stones, it is the most difficult to see of the three species present. This species 1s probably more abundant than our data indicate. Leopard Frogs are apparently absent from our study area although they occur further north, probably as glacial relicts. The Red-sided Garter Snake was also not encountered; how- ever, it may occur sporadically, as we have a recent and reliable sight record of one at Point de Roche on the west shore of Lake Claire (J. Kristensen, Department of Zoology, University of Alberta, personal communication). Thus the known herpetofauna of northeastern Alberta consists of four amphibian species: the Leopard Frog, Wood Frog, Boreal Chorus Frog, and Canadian Toad, and a single reptile, the Red-sided Garter Snake. The Wood Frog is the most abundant and widespread species. The three common amphibian species share a num- ber of life history features. They overwinter in terrestrial sites and spawn in the spring. Larval development is rapid and metamorphosis occurs during the first summer of life. Borrow pits as well as natural ponds appear to be favored as spawning sites over streams and lakes, although lakes are used by toads. Subadults and adults are terrestrial; however, most are found within 50 m of a water body. Highest densities of all species are found in moist habitats, near water vegetated by sedges, grasses, horsetail, willow, and aspen. The habitat utilization, population density, and distribution of age classes for each species is schematically summarized in Figure 5. The vegetation profile is based on the frequency of occurrence of dominant species within each 20-m segment of the 48 sample plots adjacent to 154 THE CANADIAN FIELD-NATURALIST water. The density values for each amphibian species are based on totals of subadults and adults for all sites and thus represent mean densities. Wood Frogs may be found over 100 m from water; however, most are found within 50 m. Small individuals tend to be found closer to water than large ones. Chorus Frogs are uncommon at distances greater than 20 m from water and none were found further than 100 m - from water. Canadian Toads gradually decrease in abundance at distances greater than 40 m from water. No amphibians were found in dry sandy areas with Jack Pine forest habitat. Acknowledgments Financial support for this study was provided by Alberta Oil Sands Environmental Research Program (AOSERP Project TF 5.1). L. Brusnyk provided excellent field assistance and, as well, helped with data analysis. Both L. Brusnyk and B. Herbert prepared the figures. J. O. Murie provided useful suggestions during the initial phase of this study. Literature Cited Bellis, E. D. 1961. Cover value and escape habits of the wood frog in a Minnesota bog. Herpetologica 17: 228- 231. Bellis, E.D. 1962. The influence of humidity on wood frog activity. American Midland Naturalist 68: 139-148. Bellis, E. D. 1965. Home range and movements of the wood frog in a northern bog. Ecology 46: 90-98. Blair, W.F., A. P. Blair, P. Brodkorb, F. R. Cagle, and G. A. Moore. 1968. Vertebrates of the United States. 2nd edition. McGraw-Hill Book Co., New York. 616 pp. Bleakney, J. S. 1954. Range extensions of amphibians in eastern Canada. Canadian Field-Naturalist 66: 165-171. Breckenridge, W. J. and J. R. Tester. 1961. Growth, local movements and hibernation of the Manitoba toad, Bufo hemiophrys. Ecology 42: 637-646. Conant, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. 2nd edition. Houghton Mifflin Co., Boston. 429 pp. Gosner, K. L. 1960. A simplified table for staging anuran embryos and larvae with notes on identification. Herpe- tologica 16: 183-190. Harper, F. 1931. Amphibians and reptiles of the Athabasca and Great Slave lakes region. Canadian Field-Naturalist 45: 68-70. Heatewole, H. 1961. Habitat selection and activity of the wood frog, Rana sylvatica Le Conte. American Midland Naturalist 66: 301-313. Henrich, T. W. 1968. Morphological evidence of secon- dary intergradation between Bufo hemiophrys Cope and Bufo americanus Holbrook in eastern South Dakota. Herpetologica 24: 1-13. Vol. 93 Herreid, C. F. and S. Kinney. 1966. Survival of Alaskan Woodfrog (Rana sylvatica) larvae. Ecology 47: 1039- 1041. Hodge, R. P. 1976. Amphibians and reptiles in Alaska, the Yukon and Northwest Territories. Alaska Northwest Publishing Co., Anchorage, Alaska. 89 pp. Kelleher, K. E. and J. R. Tester. 1969. Homing and sur- vival in the Manitoba toad, Bufo hemiophrys, in Min- nesota. Ecology 50: 1040-1048. Kramer, D. C. 1973. Movements of western chorus frogs Pseudacris triseriata triseriata tagged with Co. Journal of Herpetology 7: 231-235. Kramer, D. C. 1974. Home range of the western chorus frog Pseudacris triseriata triseriata. Journal of Herpe- tology 8: 245-246. Lillywhite, H. B. and J. Wassersug. 1974. Comments on a postmetamorphic aggregate of Bufo boreas. Copeia 1974: 984-986. Logier, E.B.S. and G.C. Toner. 1961. Check list of amphibians and reptiles of Canada and Alaska, a revision of contribution No. 41. Royal Ontario Museum, Contribution Number 53. 92 pp. Marshall, W.H. and M. F. Buell. 1955. A study of the occurrence of amphibians in relation to a bog suc- cession, Itasca State Park, Minnesota. Ecology 36: 381— 387. Matthews, T. C. 1971. Genetic changes in a population of boreal chorus frogs ( Pseudacris triseriata) polymorphic for color. American Midland Naturalist 85: 208-221. Porter, K.R. 1972. Herpetology. W.B. Saunders Co., Toronto. 523 pp. Ray, C. 1961. The application of Bergmann’s and Allen’s rules to the poikilotherms. Journal of Morphology 106: 85-108. Schueler, F. W. 1973. Frogs of the Ontario coast of Hudson Bay and James Bay. Canadian Field-Naturalist 87: 409-418. Shelford, V. E. 1913. The reactions of certain animals to gradients of evaporating power of air. A study of experimental ecology. Biological Bulletin (Woods Hole) 25: 79-120. Soper, J.D. 1964. The mammals of Alberta. Hamly Press Ltd., Edmonton. 402 pp. Stebbins, R. C. 1954. Amphibians and reptiles of western North America. McGraw-Hill Book Co., Toronto. 536 pp. Tamsitt, J. R. 1962. Notes ona population of the Manitoba toad (Bufo hemiophrys) in the Delta marsh region of Lake Manitoba, Canada. Ecology 43: 147-150. Underhill, J. C. 1961. Intraspecific variation in the Dakota toad, Bufo hemiophrys, from northeastern South Dakota. Herpetologica 17: 220-227. Whitaker, J.O. 1971. A study of the western chorus frog, Pseudacris triseriata, in Virgo County, Indiana. Journal of Herpetology 5: 127-150. Wright, A. H. and A. A. Wright. 1949. Handbook of frogs and toads of the United States and Canada. Comstock Publishing Co., Ithaca, New York. 640 pp. Received 8 August 1978 Accepted 27 November 1978 Caribou Distribution and Group Composition Associated with Construction of the Trans-Alaska Pipeline RAYMOND D. CAMERON,! KENNETH R. WHITTEN, ! WALTER T. SMITH,2 and DANIEL D. ROBY: ‘Alaska Department of Fish and Game, 1300 College Road, Fairbanks, Alaska 99701 2Department of Biology, Idaho State University, Pocatello, Idaho 83209 3Alaska Cooperative Wildlife Research Unit, University of Alaska, Fairbanks, Alaska 99701 Cameron, Raymond D., Kenneth R. Whitten, Walter T. Smith, and Daniel D. Roby. 1979. Caribou distribution and group composition associated with construction of the Trans-Alaska Pipeline. Canadian Field-Naturalist 93(2): 155-162. Caribou surveys were conducted periodically along the Trans-Alaska Pipeline haul road on the central Arctic Slope between June and November 1975. Mean calf percentage observed in summer was approximately one-third lower than that obtained from concurrent aerial surveys of both the pipeline corridor and adjacent areas; however, fall means were identical. In both summer and early fall, mean latitudes calculated for groups with and without calves along the haul road were more southerly than for the corresponding group types observed through aerial survey. A more detailed regional comparison of survey data revealed corridor-related abnormalities in Caribou distribution and group composition. No Caribou, or relatively low numbers, were observed in the northernmost segment of the pipeline corridor near Prudhoe Bay, and calf percentages in summer were consistently lower for each of four arbitrarily established regions of the haul road than expected on the basis of aerial survey results; fall calf percentages did not differ appreciably. Mean group size was generally lower along the haul road than for comparable areas to the east and west. Responses of Caribou to the pipeline corridor in general, and of cows and calves in particular, are discussed in relation to inherent avoidance tendencies as modified seasonally by terrain, group dominance, and human activity. Key Words: Caribou, Rangifer, pipeline, disturbance, group composition, Prudhoe Bay, ecological distribution. The discovery of vast petroleum reserves at tion-related human activity. Consequently, a Prudhoe Bay in 1968 resulted in construction of | major study was undertaken to evaluate the the Trans-Alaska Pipeline, designed totransport reactions of Caribou to the Trans-Alaska Pipe- crude oil to Valdez on the Gulf of Alaska. North line on a seasonal basis and to document any of the Yukon River, the pipeline and its associated shifts in annual movement patterns associated haul road were to traverse the ranges and range occupancy. To date, this investigation of two major subpopulations of Barren-ground _ has been largely restricted to the Arctic Slope. Caribou (Rangifer tarandus granti), the Porcu- Based on summer and fall surveys in this pine and Western Arctic herds (Skoog 1968; region between 1969 and 1972, A. Gavin (1973, Hemming 1971), and concern was focused onthe Report by Atlantic Richfield Company) re- potential restriction of Caribou movements. ported a decline in Caribou numbers from 26000 In 1971 and 1972 K. N. Child (1973, Comple- to 2500, and he thought these Caribou to be tion Report, Alaska Cooperative Wildlife Re- “offshoots” from the Porcupine and Western search Unit, University of Alaska), using Arctic herds. Cameron and Whitten (1979), simulated pipeline and pipeline crossing struc- however, have recently identified a third sub- tures at Prudhoe Bay, attempted to assess the population of about 5000 Caribou whose range effectiveness of various above-ground designsin is centered on the pipeline route along the permitting free movements of Caribou. The Sagavanirktok River (Figure |). Seasonal move- majority of approaching Caribou avoided the ments are primarily north-south, between simulation and, of the successful crossings, calving grounds near the coast and winter range about 3 times as many were by use oframpsthan in the northern foothills of the Brooks Range. by passage under the structure. This detailed The identity of this “Central Arctic herd” has study provided valuable preliminary data but apparently been confounded by periodic influx was limited to observations on summer range. from one or both of the larger adjacent herds, Thus, little was known of the seasonality of although such outside influence was thought to Caribou responses to pipelines and construc- be minimal during the course of the present 155 THE CANADIAN FIELD-NATURALIST Vol. 93 156 OLIKTOK | ry 4 STATION 1 DEADHORSEC}/ » / i” Y = MITIIN LI le! le\ FRANKLIN s) BLUFFS PUMP STATION 2 KEY PUMP. STATION 3 ee soe —— Above-ground Pipeline ecoeeeBuried Pipeline = Survey Coverage GALBRAITH-+ LAKE t PUMP STATION 4 FIGURE |. Survey coverage along the Trans-Alaska Pipeline corridor and regional boundaries established for comparison of haul road and aerial survey results. IST) study. This report deals with changes in lati- tudinal position, group size, and composition of Caribou along the Trans-Alaska Pipeline haul road during summer and fall 1975. These data are compared with similar values obtained through concurrent aerial survey of alargerarea extending east and west of the corridor. The haul road between the Yukon River and Prudhoe Bay was completed in fall 1974, and pipeline construction efforts were near maxi- mum during the major portion of this investiga- tion. The pipeline is 122 cm in diameter and is alternately buried and above ground depending on soil stability; ground clearances of elevated sections range from less than | m toa maximum of about 5 m. Methods Systematic surveys were conducted from the Trans-Alaska Pipeline haul road between June and November 1975. Generally these were scheduled twice-monthly, each survey consisting of two complete trips over the 263-km distance between Pump Station 4 and Prudhoe Bay (Figure 1). As only half of the route could be covered in | d, most surveys required 4 d for completion. Owing to inclement weather, survey components were not necessarily on consecutive CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE 157 days, but an attempt was made to minimize the time interval for each series. Inclusive dates are given in Table 1. A pickup truck, generally with one driver/ observer, was used for all road surveys. Speeds were adjusted to 40-65 km/h, depending on visibility, to provide reasonably constant per- ceptibility of Caribou for anestimated 1.5 kmon either side of the haul road. Only Caribou observed initially with the naked eye were recorded, but binoculars and spotting scope were used as required to obtain additional information. Pertinent data recorded for each Caribou sighting included the following: Location — road distance from a known point: Number of Caribou per group ( defined arbi- trarily as a single Caribou, or two or more Caribou separated by less than an estimated 300 m); Group composition — calf — less than | yr old, yearling — more than | yr old but less than 2 yr old, cow — female more than 2yr of age, bull — male more than 2 yr of age, adult — more than | yr old, sex unknown, unknown — unclassified as to sex or age. TABLE I—Changes in Caribou numbers and mean group composition along the Trans-Alaska Pipeline haul road, June—November 1975 Groups without Total classified Groups with calves calves Total number Number Number Number Survey of caribou of Bulls, Calves, of Bulls, Calves, of Bulls, dates observed caribou % % caribou” % % caribou” % June 11-18 91 79 66 0 0 0 0 79 66 June 24-July 2 361 342 86 3 19 8 21 323 92 July 10-17 677 677 52 32 499 35 43 178 100 July 24-August 2 136 136 98 l 5 60 20 131 100 August 7-13 273 273 TH 7 Sy) 2 32 216 97 August 20-28 156 149 92 l 11 19 18 138 98 September 3-6 202 200 83 3 23 30 26 177 90 Summer mean*® 65 13(21) 31 40(35) 94 September 20-28 602 416 44 17 333 30 23 83 99 October 24-28 54 31 36 24 25 26 30 6 80 November 5-10 176 37 49 10 19 25 19 18 74 November 19-25 92 36 43 11 25 30 17 11 73 Fall mean® 43 17(17) 29 23(21) 92 “Total caribou in groups with no “unknowns” (see Methods). *Total caribou in groups with one or more calves present. “Total caribou in groups with no calves. “Aggregate percentages; those in parentheses were determined from aerial surveys (Cameron and Whitten 1979). 158 Road locations were converted to their lati- tudinal equivalents using 1:63 360 scale maps of USGS Topography Series. For each survey series a mean latitudinal position of Caribou was calculated as described by Cameron and Whitten (1979). Results and Discussion Caribou Numbers and Group Composition Numbers and group composition of Caribou observed from the haul road during each survey period are given in Table |. Groups with one or more “unknowns” (see Methods) were excluded from composition calculations. Ninety-eight percent of the total Caribou sighted during summer were successfully classified. During fall, however, bright sun and snow frequently created distortion which precluded classification at distances greater than about 400 m, and entire groups were recorded as unknowns; the pro- portion classified decreased to 56%. Cows and yearlings were often difficult to distinguish, and many such individuals were classified as “adults” (see Methods) when more specific identification was impossible. Calves, however, were relatively easy to recognize and reported percentages are thought to be very reliable. Similarly, iden- tification of bulls was rarely in question. During summer 13% of total Caribou ob- served from the haul road were calves, com- pared with a corresponding value of 21% obtained by systematic aerial survey of a larger area roughly centered on the pipeline route (Table 1). In contrast, a mean of 17% calves was observed in fall during both road and aerial surveys, indicating that a representative portion of the herd was present along the corridor. Within each season, calf percentages for groups with calves were similar for road and aerial observations (Table 1). It can be calculated from the summer road data in Table |, however, that the number of Caribou observed in groups with calves averaged 33% of the total classified, whereas 60% of Caribou classified by air were observed in groups with calves (see Cameron and Whitten 1979): respective mean values for fall were similar at 77 and 81%. In addition, observations from the haul road indicate that bulls were present in low numbers in groups with calves but consistently predominated in groups without calves, and the percentages did not THE CANADIAN FIELD-NATURALIST Vol. 93 cb) 79) 3 = re a) —® 0 Total Caribou Observed —-—8 Caribou in Groups w/Calves ta D. ye 4 Caribou in Groups w/o Calves \ co) 1S) (ep) ey, S ro) _ (©) Cc ro) ® = Jun Jul Aug Sep Oct Nov Month FIGURE 2. Seasonal changes in mean latitudinal position and mean group size of Caribou along the haul road and a comparison with corresponding results from aerial surveys. differ appreciably with season for either group type (Table 1). Clearly then, the low proportion of calves recorded during haul road surveys in summer was due primarily toa local reduction in the relative number and/or size (see below) of groups with calves rather than a lower per- centage of calves in such groups. Latitudinal Movements Changes in mean latitudinal position of Caribou along the haul road are depicted in Figure 2a. Throughout summer, groups with calves remained farther north than groups without calves. A similar trend was obtained from concurrent aerial surveys (Cameron and Whitten 1979), and these results are plotted for comparison. Thus, the more northerly location of the mean shown for aggregate sightings by air ISA) is partially attributable to a greater proportion of groups with calves (see previous section) because they tend to occupy higher latitudes. Positions determined by aerial survey for groups both with and without calves, however, were also farther north than those calculated from road survey observations. Corresponding means did not differ appreciably in September, but the same general trend was again apparent in November (Figure 2a), despite the fact that no overall differences in calf percentage were observed (Table 1). Thus, dissimilar composi- tion does not fully account for the more southerly distribution of Caribou along the pipeline corridor. Rather, local abnormalities are indicated. Group Size Two distinct peaks in group size are shown in Figure 2b for haul road observations, one in mid-July during post-calving aggregation and another in September just prior to the rut. Except for late July and early September, the mean size of groups with calves was greater than that for groups without calves. A further comparison of these means with those deter- mined from aerial surveys indicates that in June, August, and September the size of correspond- ing group types was higher for aerial than for haul road surveys. A valid comparison is not possible for November because of the extremely small sample size (Table |). Because the same | June O Rr oeias > | August Percent of Total Caribou Observed CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE Bik 159 criteria for group identification were used in both survey procedures (see Methods), these data suggest avoidance of the area by larger groups, group fragmentation, and/or a de- creased tendency for group coalescence near the pipeline corridor. Differences in Regional Distribution and Group Composition To permit a more accurate assessment of latitudinal distribution and group composition of Caribou, four regions were delineated along the pipeline corridor for direct comparison of aerial and road survey observations. These regions were centered on the corridor, and each was bounded arbitrarily by '4 degree of latitude and 2 degrees of longitude (Figure 1). The regional distribution of total Caribou obser- vations for both aerial and road surveys is shown in Figure 3, and applicable calf percentages are presented in Table 2. It was assumed that aerial survey results reflected the true distribution of Caribou among the four regions and provided representative data on composition within each region. Corresponding data from the haul road, obtained concurrently, are compared with these “expected” values. As sample sizes were fre- quently low, statistical analysis of differences in group composition was not feasible, but some overall and regional trends are apparent. Figure 3 demonstrates a distinct north-south gradient in regional density of Caribou for both November [] Air September “abl sf hi. a ZvOrenG [PREZ aac, Region FiGurE 3. Regional distribution of Caribou (see Figure 1) determined from corresponding aerial and haul road surveys. 160 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 2—Regional comparison of calf percentages determined from corresponding aerial and haul road surveys, June-November 1975 Region | Region 2 Region 3 Region 4 Number Number Number Number Survey of Calves, of Calves of Calves, of Calves, Month* method* caribou” % caribou” % caribou” % caribou” % June Air 4 0 65 2 159 10 231 27 Road 0 121 0 195 4 26 12 August Air 0 — 49 0 122 9 200 34 Road 4 0 37 0 183 3 49 27 September Air 32 0 174 21 18 DD 99 20 Road 59 6 125 9 wy) 25 0 = November Air 238 7 256 17 101 27 57 44 Road 36 11 0 — 0 — 0 = “Aerial surveys (Cameron and Whitten 1979) or haul road surveys within the regions shown in Figure 1. "Number of caribou classified, i.e., total caribou in groups with no “unknowns” (see Methods). June and August, with aerial sightings in region 4 representing 50 and 55% of the total, respec- tively. In contrast, the majority of haul road observations during summer were in region 3, and relatively few Caribou were observed near the coast. Calf percentages for regions 3 and 4 of the haul road were appreciably lower than expected, while few calves were noted in regions 1 and 2 from either road or air (Table 2). For all regions combined, calf percentages determined from aerial and haul road surveys, respectively, were 17 and 3 in June, and 20 and 7 in August. Thus, the more southerly distribution of Cari- bou within the pipeline corridor in summer was due in part to a general avoidance of northern sections near Prudhoe Bay. In addition, aggre- gate and regional calf percentages were con- sistently lower than the expected values, indi- cating a disproportionate avoidance of the pipeline corridor by groups with calves. The resulting preponderance of bulls (i.e., groups without calves), by virtue of their latitudinal preference (Figure 2a), contributed further to a southward displacement of Caribou along the haul road. By late September a southward movement of Caribou had occurred. Approximately half of the total sightings from both air and haul road were in region 2, but no Caribou were observed along the corridor in region 4 where nearly one- third of the total was observed during aerial surveys (Figure 3). During both aerial and road surveys the majority of Caribou were found in groups with calves (Table 1), and mean group size increased (Figure 2b) as pre-rut aggrega- tions formed. Although aerial observations indicated that groups without calves remained substantially farther south (Figure 2a), such differences are of little quantitative importance as these Caribou represented less than 10% of total sightings. Regional differences in calf percentage demonstrate no consistent trend (Table 2), but combining data for regions 1, 2, and 3 results in an estimated 17% calves for both air and road surveys. Thus, in September a strong avoidance of the Prudhoe Bay area is indicated, but sightings of Caribou elsewhere along the haul road appear to reflect overall herd composition, and disproportionate avoidance by groups with calves was not apparent. In November aerial surveys established that fewer Caribou were occupying coastal areas. Most Caribou within the corridor were in region 1, and none were observed in regions 3 and 4 (Figure 3). Total haul road sightings in Novem- ber were among the lowest recorded (Table 1), and any conclusions regarding differences in calf composition would be equivocal because of small sample size. Nevertheless, combined data for fall again indicate that a more representative portion of the herd approached and was ob- served from the haul road, although continued avoidance of northern areas is indicated. Relatively low numbers of Caribou near the haul road in region 4 during summer and fall may partially explain the small group size observed within the pipeline corridor (Figure 2b). Groups with calves observed along the coast 9 by air were generally larger than comparable inland groups, and their avoidance of the haul road area would tend to depress the calculated mean irrespective of other influences on group size. Groups without calves, however, exhibited a similar decrease in summer and a corridor- related reduction in group size remains suspect. Avoidance Behavior of Caribou Avoidance of the Prudhoe Bay area was noted throughout the course of this study. The coastal region near Prudhoe Bay, recently the site of increased construction and exploration activity, was previously a portion of the calving grounds for the Central Arctic herd. As late as 1972 K. N. Child (1973, op. cit.) and A. Gavin (1973, op. cit.) reported calving within or immediately adjacent to the Prudhoe Bay complex. With facilities expansion and continued human activi- ty over the past 3 or 4 yr, local occupancy by Caribou has generally decreased, and in 1975 we observed no neo-natal calves from this northern- most section of the haul road. The area was also previously invaded by Caribou during annual post-calving movements along the coast (Cam- eron and Whitten 1979) and during oscillatory movements to and from the coast in response to changing insect density (Child 1973, op. cit.; White et al. 1975). Evidence for these occur- rences 1s still visible as Caribou trail systems and, although movements within the Prudhoe com- plex are still detectable, they are now mere remnants of past activities. Post-calving and insect-induced movements, however, still occur elsewhere along the coast between the Colville and Canning rivers, indicating avoidance of a specific portion of summer range with continued occupancy of adjacent regions. Disturbance- related abandonment of range is thought to be a gradual process, occurring with increasing avoidance of adverse stimuli (Klein 1971; Calef 1974), and the recent history of changing Caribou occupancy near Prudhoe Bay appears to reflect this pattern. In summer, avoidance of the pipeline corridor was primarily by cows and calves. Cows are most sensitive to unusual stimuli just before par- turition or during the early stages of labor (Lent 1966), but the present results suggest that heightened sensitivity extends through the first 2 or 3 months post-partum. Other reports indicate CAMERON ET AL.: CARIBOU DISTRIBUTION NEAR PIPELINE 16] that cows and young calves are more easily alarmed by, and more likely to flee from, a potential threat than are male Caribou, barren cows, or cows with older calves (de Vos 1960: Lent 1966; Bergerud 1974). Loud noises, unless associated with moving objects, do not generally alarm Caribou which perceive and identify adverse stimuli through visual and olfactory means (Bergerud 1974). Cows with young calves, however, consistently take flight from distant stimuli, using visual analysis without verifica- tion by scent (Bergerud 1974). Bergerud (1974) and Lent (1966) reported that alert posture assumed by individuals within a group did not generally induce group flight unless cows and neo-natal calves were present. Also, we noted that cow-calf pairs on the calving grounds were difficult to approach on foot to within 400 m. Because of the importance of visual stimuh, local terrain relief may influence the degree of avoidance of the pipeline corridor. North of Pump Station 2 the haul raod traverses more than 100 km of continuous coastal plain (see Figure 1) which is preferred summer range for the Central Arctic herd. Construction ac- tivity 1s frequently visible for 20 km or more, and the greater inherent sensitivity of cows and calves together with greater visibility provided by flat terrain may account for the observed avoidance of the haul road during summer. Bulls do not appear to be as sensitive to local human activity; however, since the mean latitude of groups without calves (1.e., principally bulls) was also displaced southward some evidence exists for low-level avoidance of northern areas. Avoidance of the Prudhoe Bay complex continued during fall, and most Caribou were observed farther inland. In September and October groups along the haul road were well mixed sexually, and cow-calf avoidance of the pipeline corridor appears to have diminished. This may be partially a result of a change in group dominance associated with breeding; rutting bulls clearly influence group activity (Bergerud 1973; Cameron and Whitten, un- published observations; Roby 1978) and may alter the “normal” avoidance behavior of cow- calf pairs. Perhaps more importantly, the ad- vanced age of calves may have reduced the sensitivity of maternal cows to human dis- turbance. 162 THE CANADIAN FIELD-NATURALIST Continued study is necessary to assess the long-term responses of Caribou to oil develop- ment in the Arctic. Sustained exposure of Caribou to these stimuli may increase the degree of avoidance, stabilize it at the current level, or ultimately result in a reversal with subsequent accommodation. Further, it is not known whether this avoidance response, by virtue of its probable influence on free movement, will seriously alter herd productivity. Such are matters for conjecture, and should be addressed by future investigations. Acknowledgments This study was funded jointly by the Alaska Department of Fish and Game through the Joint State/ Federal Fish and Wildlife Advisory Team and by Federal Aid in Wildlife Restoration Project W-17-8. Logistic support by Alyeska Pipeline Service Company is gratefully acknowledged. Literature Cited Bergerud, A. T. 1973. Movement and rutting behavior of caribou (Rangifer tarandus) at Mount Albert, Quebec. Canadian Field-Naturalist 87: 357-369. Bergerud, A. T. 1974. The role of the environment in the aggregation, movement, and disturbance behavior of caribou. Jn The behaviour of ungulates and its relation to management. Volume 2. Edited by V. Geist and F. Walther. International Union for Conservation of Nature and Natural Resources Publications, New Series Number 24. Vol. 93 Calef, G. W. 1974. The predicted effect of the Canadian Arctic Gas pipeline project on the Porcupine caribou herd. Jn Research reports, Volume IV, Environmental impact assessment of the portion of the Mackenzie gas pipeline from Alaska to Alberta. Environmental Protec- tion Board, Winnipeg, Manitoba. pp. 101-120. Cameron, R. D. and K. R. Whitten. 1979. Seasonal move- ments and sexual segregation of caribou determined by aerial survey. Journal of Wildlife Management (in press). De Vos, A. 1960. Behavior of barren-ground caribou on their calving grounds. Journal of Wildlife Management 24: 250-258. Hemming, J. E. 1971. The distribution and movement patterns of caribou in Alaska. Alaska Department of Fish and Game, Wildlife Technical Bulletin Number 1. 60 pp. : Klein, D. R. 1971. Reaction of reindeer to obstructions and disturbances. Science 173: 393-398. Lent, P. C. 1966. Calving and related social behavior in the barren-ground caribou. Zeitschrift fur Tierpsycholo- gie 6: 701-756. Roby, D. D. 1978. Behavioral patterns of barren-ground caribou of the Central Arctic herd adjacent to the Trans- Alaska oil pipeline. M.Sc. thesis, University of Alaska. 200 pp. Skoog, R. O. 1968. Ecology of caribou ( Rangifer tarandus granti) in Alaska. Ph.D. thesis, University of California, Berkeley, California. 699 pp. 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. In Eco- logical investigations of the tundra biome in the Prudhoe Bay region, Alaska. Edited by J. Brown. Biological Papers of the University of Alaska, Special Report Number 2. pp. 150-201. Received 4 October 1977 Accepted 27 December 1978 Interesting Vascular Plants from Southeastern Yukon Territory GEORGE W. SCOTTER! and WILLIAM J. CODY? 'Canadian Wildlife Service, Room 1000, 9942 — 108 Street, Edmonton, Alberta T5K 2J5 2Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 Scotter, George W. and William J. Cody. 1979. Interesting vascular plants from southeastern Yukon Territory. Canadian Field-Naturalist 93(2): 163-170. Recent vascular plant collections in extreme southeastern Yukon have added considerable data on the flora of the Yukon Territory. This paper reports 19 taxa as newto the Yukon (Borrychium virginianum var. europaeum, Dryopteris spinulosa, Matteuccia struthiopteris var. pensylvanica, Agrostis exarata, Cinna latifolia, Muhlenbergia mexicana, Oryzopsis asperifolia, Poa ammophila, Sphenopholis intermedia, Parietaria pensylvanica, Vicia americana, Viola rugulosa, Circaea alpina, Aralia nudicaulis, Osmorhiza depauperata, Castilleja miniata, Anaphalis margaritacea var. subalpina, Aster ciliolatus, and Bidens cernua) and 43 taxa as extensions of known ranges into the southeastern Yukon. Key Words: vascular plants, Yukon Territory, flora, range extensions. During July of 1977, the senior author, in connection with other work, had the oppor- tunity to collect vascular plants from three sites in extreme southeastern Yukon. Although 214 taxa of vascular plants were collected, the purpose of this paper is to report only new records and major extensions of known ranges for the Yukon Territory. Field studies were made at one cool spring and two hot springs. The cool spring site, the most intensively collected, was near two terraced tufa deposits at the base of a limestone ridge on the east side of the Coal River. Scalloped terrace- faces on the west side of the pools were covered with a variety of bryophytes and vascular plants (Figure 1). Specimens from poorly drained communities below the two tufa deposits and in surrounding Black Spruce (Picea mariana), White Spruce (P. glauca), Lodgepole Pine (Pinus contorta var. latifolia), and Balsam Poplar (Populus balsamifera) stands were also col- lected. The locations (Figure 2) and elevation of the collection sites, date of collection, collectors’ names and collection numbers follow: 1. Cool springs and adjacent areas near the Coal River, 60°09’N, 127°26’W, elevation 650-750 m, 16-19 July 1977, Scotter and T. Ahti 24304-24598. 2. Hot springs and adjacent areas near Larsen Creek, 60°12’N, 125°32’W, elevation 760 m, 20-22 July 1977, Scotter 24599-24756. 3. Hot springs and adjacent areas near the Beaver River, 60°23’N, 125°34’W, elevation 450 m, 22 July 1977, Scotter 24757-25775. The southeastern Yukon is mostly forested with open stands of Black Spruce, White Spruce, and Lodgepole Pine. Small portions of the area extend above treeline, which occurs at approxi- mately 1200-1350 m asl. The area lies within the Liard and Hyland plateaus and the whole area was covered by at least one advance of Cordil- leran ice. For greater details on vegetation and geology of the region readers are referred to Douglas and Norris (1959), Bostock (1948), Gabrielse and Blusson (1969), Rowe (1972), and Oswald and Senyk (1977). Voucher specimens have been deposited at the Agriculture Canada herbarium (DAO). Unless otherwise indicated, all specimens were deter-. mined by Cody. A complete species list of the plants collected is available and may be obtained by contacting the authors. In the annotated list, voucher numbers follow the species; site numbers are in brackets. Species not previously reported from the Yukon are noted with an asterisk (*). Nomenclature follows Porsild and Cody (1979). The ecology of springs in northwestern Cana- da has been little studied except for Liard Hot Springs, British Columbia (Porsild and Crum 1961) and some near Hole-in-the-Wall Lake, Nahanni National Park, Northwest Territories (Arnold 1961). The three springs studied in the Yukon are very different from each other in structure and associated vegetation so that 163 Vol. 93 THE CANADIAN FIELD-NATURALIST 164 istic «tg land ali Phe AMOR SRS dal Are et ” ia) FIGURE |. The most common plants at the cool spring site growing on top of the tufa terraces were tall White Bog Orchid (Hahenaria dilatata), Marsh Grass-of-Parnassus ( Parnassia palustris var. neogaea), Meadow Fireweed (Epi/obium glandulosum var. adencaulon), Yellow Monkey Flower (Minulus guttatus), Pink Fleabane ( Erigeron philadelphicus), Yellow Ragwort (Senecio pauperculus). and several species of grasses and sedges. Mats of velvety-green bryophytes covered the sides of the moist tufa terraces. 1979 kilometres Yukon Territory RR eS Coal River springs SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 165 Nahanni National Northwest Territory SAGAS ai) PRA r ( Beaver River \ springs Larsen Creek \ springs FIGURE 2. Map of the southeastern Yukon and adjacent regions showing the three collection sites in relation to Liard Hot Springs and Nahanni National Park. generalization on ecology is difficult. The Liard Hot Springs and the springs near the Coal River. however, have a close affinity in both structure and associated vegetation. *Botrychium virginianum var. europaeum (Rattle- snake Fern). 24475, 24478 (1); 24712, 24673 (2). Previously known from northern British Columbia (Hultén 1968; Porsild 1951) and several sites in southwestern District of Mackenzie (Cody 1963: Scotter and Cody 1974). Athyrium filix-femina ssp. cyclosorum (Lady Fern). 24473 (1). This appears to be the second report of this species from the Yukon Territory. Porsild (1951) found A. filix-femina ssp. cyclosorum at a hot spring site in the Mayo District. In the southwestern District of Mackenzie it is known from only three localities (Jeffrey 1961; Porsild 1961; Scotter and Cody 1974), two of which are hot spring sites. *Drvopteris spinulosa 24468, 24472, 24482 (1). (Spinulose Wood Fern). 166 Rare in the northwestern part of its range. A site in Nahanni National Park, the second site in the District of Mackenzie, is soon to be reported by Cody et al.’ * Matteuccia struthiopteris var. pensylvanica (Pteritis nodulosa) (Ostrich Fern). 24764 (3). According to Hultén (1968), this fern is restricted to small isolated areas in Alaska. In northern British Columbia, Porsild and Crum (1961) reported that it formed dense colonies near and below the springs at Liard Hot Springs. In the southwestern District of Mackenzie it occurs on the flood plain of the Liard River nearly as far downstream as Nahanni Butte (Cody 1963) and onthe floodplains of the lower South Nahanni River (Scotter and Cody 1974). Equisetum hyemale var. affine (Scouring Rush). 24743 (2). According to the map in Hultén (1968), this collection extends the known range to the south- eastern Yukon Territory. Selaginella selaginoides (Spikemoss). 24527 (1); 24639 (2). These collections of this species in the southeastern Yukon Territory change the position of the delimiting line in Hultén (1968) to include all of the southern Yukon Territory. x Agrohordeum macounti (Elymus macounii). 24617 (2). This hybrid of Agropyron trachycaulum and Hordeum jubatum was known inthe Yukon Territory by Porsild (1951) from only three collections. In the herbarium of the Department of Agriculture at Ottawa (DAO), there are now an additional eight collections. The hybrid is thus of much more frequent occurrence than previously suspected, and may occur wherever the two parents are found growing in close proximity. * Agrostis exarata. 24345, 24351, 24358 (1); 24622B (2). Cody and Porsild (1968) extended the known range of this species from the Pacific Coast to the District of Mackenzie at Tungsten. Scotter and Cody (1974) have since reported additional sites at both hot and cool springs in Nahanni National Park. Bromus ciliatus (Fringed Brome). 24320, 24332 (1); 24610, 24611 (2). Hultén (1968) shows only three widely separated Yukon sites on his map. The species is not common in 3Cody, W. J., G. W. Scotter, and S. S. Talbot. Additions to the vascular plant flora of Nahanni National Park, Northwest Territories (in preparation). THE CANADIAN FIELD-NATURALIST Vol. 93 southwestern District of Mackenzie, but Porsild (Porsild and Crum 1961) has collected it at Liard Hot Springs. * Cinna latifolia (Wood Reedgrass). 24412 (1). Cody (1963) reported a number of stations along the lower Liard River, and Porsild and Crum (1961) found it to be occasional in rich mixed aspen and spruce woods at Liard Hot Springs. Danthonia intermedia (Wild Oat-Grass). 24770 (3). This collection helps fill in the known distribution of this species between sites in southern Yukon Territory (Porsild 1951, 1966) shown on Hultén’s (1968) map, and sites reported by Scotter and Cody (1974) and Cody et al.3 in Nahanni National Park. Glyceria striata var. stricta (Fowl Manna Grass). 24342, 24416, 24523, 24555 (1); 24625 (2). Porsild (1951) surmised that this grass would be found in the Yukon Territory because !t was already known from central Alaska and the Liard Hot Springs. Hultén (1968) depicted a collection from near Watson Lake (Calder & Kukkonen 27617 and Calder & Gillett 26544 [DAO]). The specimens cited above thus represent the second report of this species for the Yukon Territory. * Muhlenbergia mexicana. 24629 (2). (det. C.G. Reeder). Porsild (Porsild and Crum 1961) and Welsh (1974) reported this grass from the Liard Hot Springs. * Oryzopsis asperifolia (Mountain-Rice). 24435 (1). In northern British Columbia, O. asperifolia is known from Mile 578 Alaska Highway, 59°38’N, 127°29’W (Calder & Gillett 25653 [DAO]). In the District of Mackenzie it has recently been reported from Nahann: National Park (Cody et al.3), an extension of range to the northwest from the Salt Plains west of Fort Smith (Raup 1936). Phalaris arundinacea (Reed Canary Grass). 24408 (1). The map in Hultén (1968) depicts collections from Whitehorse (Porsild 1951) and near Teslin Lake. Additional Yukon Territory specimens at DAO are from Albert Creek, Mile 643.5 Alaska Highway (Calder & Gillett 26545, Harms 17199). * Poa ammophila. 24596 (1). (det. M. Barkworth). This is an extension of the known range of some 900 km to the south from the type region at the mouth of the Mackenzie River on the arctic coast of the District of Mackenzie. It 1s not known to occur between those collections. Schizachne purpurascens. 24434 (1). Previous collections in the Yukon Territory were from along the Canol Road (Porsild 1951; Hultén 1968). 1979 * Sphenopholis intermedia. 24314 (1); 24602, 24612, 24740 (2). Porsild (1951) and Hultén (1968) reported this grass from Liard Hot Springs and Tanana (Manley) Hot Springs in Alaska. Scotter and Cody (1974) reported it from Nahanni National Park and commented on its presence elsewhere in the District of Mackenzie. Carex atherodes. 24510, 24511 (1). These collections will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern section of the Yukon Territory. Carex concinna. 24452 (1). This collection will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern section of the Yukon Territory. Juncus balticus var. littoralis (J. arcticus spp. ater, sensu Hultén 1968) (Wire Rush). 24540 (1). This collection will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern section of the Yukon Territory. Allium schoenoprasum var. sibiricum (Wild Chives). 24563 (1). This collection will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern section of the Yukon Territory. Smilacina trifolia (Three-leaved Solomon’s-seal). MTS 3) (De Apparently rare in southeastern Yukon Territory. The single dot onthe map in Hultén (1968) is probably based on specimens gathered at Watson Lake by Raup and Correll (Porsild 1974). It was also collected there by Mitchell 9] and Gillett 3618 (DAO). Another unreported collection is from the Cassiar Mountains, 60°18’N, 130°18’W, Poole 27 (DAO). Goodyera repens (Rattlesnake Plantain). 24437 (1). According to the map in Hultén (1968), previous collections from the Yukon Territory have ali been made west of 130° W longitude. Salix longistylis (S. alaxensis var. longisiylis). 24584 (1): 24611, 24644, 24662 (2). These collections will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern section of the Yukon Territory. The species is, as well, based on collections at DAO, much more frequent in the Mackenzie Mountains in the western District of Mackenzie, than indicated on that map. Salix padophylla. 24518, 24576 (1); 24660 (2). These collections will move the delimiting line on the map in Hultén (1968) eastwards to include the extreme southeastern section of the Yukon Territory. To the east it has been recorded from several sites in Nahanni National Park (Scotter and Cody 1974). SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 167 * Parietaria pensylvanica (Pellitory). 24725 (2). The distribution of this species in Canada is quite spotty (Bassett et al. 1974) because of its apparently restricted habitat requirement of cool, moist, shady, rocky situations. Porsild (1961) reported the ex- tension of the known range from central British Columbia north to Liard Hot Springs, and discussed the distribution. Urtica gracilis (Nettle). 24462 (1); 24732 (2). Apparently rare in the Yukon Territory. Hultén (1968) indicates collections from the Dawson area, and near the British Columbia border. Polygonum amphibium var. stipulaceum (Water Smartweed). 24413, 24513 (1). These collections will move the delimiting line on the map in Hultén (1968) eastwards to include the extreme southeastern section of the Yukon Territory. Minuartia dawsonensis. 24365, 24372, 24592B (1); 24604, 24641 (2). These collections will move the delimiting line on the map in Hultén (1968) eastwards to include the extreme southeastern part of the Yukon Territory. It is also known from Nahanni National Park (Scotter and Cody 1974). Silene menziesii. 24483 (1). On the basis of the distribution map in Hultén (1968), this species appears to have a broken distribution in the northern part of its range. This collection will move the delimiting line on the map in Hultén (1968) to the east to include the extreme southeastern part of the Yukon Territory. Ranunculus abortivus (Kidneyleaf Buttercup). 24605 (2). This is apparently a rare species in the Yukon Territory. Hultén (1968) shows only four dots from the south central part of the Territory. Thalictrum sparsiflorum var. richardsonii (Meadow- Rue). 24406, 24551 (1). These collections, from the extreme southeastern part of the Yukon Territory, are intermediate between the sites on the map in Hultén (1968) and the Liard River sites in southwestern District of Mackenzie reported by Jeffrey (1961). Amelanchier alnifolia (Saskatoon). 24422, 24594 (1); 24682 (2). These collections will move the delimiting line on the map in Hultén (1968) eastwards to include the extreme southeastern part of the Yukon Territory. Dryas drummondii (Yellow Dryad). 24590 (1): 24626 (2). The collections, from the extreme southeastern part of the Yukon Territory, are from an area inter- 168 mediate between two areas of distribution outlined by Hultén (1968). Geum aleppicum var. strictum (Avens). 24618, 24731 (2). Hultén (1968) shows only a single disjunct locality in southwestern Yukon Territory. The main range extends across Canada from the east coast and reaches Nahanni National Park in southwestern District of Mackenzie. Sanguisorba sitchensis (S. stipulata, sensu Hultén (1968)) (Burnet). 24415 (1). This collection, from the extreme southeastern part of the Yukon Territory, will move the delimiting line on the map in Hultén (1968) to the east and thus include all of the southern Yukon. Astragalus americanus. 24564 (1). According to Hultén (1968), previous collections from the Yukon Territory have all come from west of longitude 132°W. Oxytropis deflexa var. foliolosa. 24646 (2). This collection helps complete our knowledge of the distribution of var. foliolosa in the Yukon Territory by extending the delimiting line around the southern population to the extreme southeastern Yukon. The map in Hultén (1968) depicts widely separated northern and southern populations in the Yukon Territory. *Vicia americana (Wild Vetch). 24745 (2). This species occurs as far north in British Columbia as Liard Hot Springs (Porsild and Crum 1961) while in southwestern District of Mackenzie it is found along the Liard River (Jeffrey 1961) and in Nahanni National Park (Scotter and Cody 1974). Geranium richardsonti. 24747 (2). Hultén (1968) shows only a few collection sites in central Yukon Territory and the hot springs at Tungsten in the headwaters of the Flat River in the District of Mackenzie. It is also known from Hole-in- the-Wall Lake in Nahanni National Park (Porsild 1961) in the District of Mackenzie. Viola langsdorffti. 24638, 24726 (2). Hultén (1968) indicates two inland sites in central and southern Yukon Territory which are remote from the mainly coastal range of the species. The collec- tions cited here represent another site more remote from the main range. *Viola rugulosa (Western Canada Violet). 24735 (2). This is a westward extension of range from sites in the Liard and South Nahanni valleys in southwestern District of Mackenzie reported by Jeffrey (1961) and Scotter and Cody (1974). THE CANADIAN FIELD-NATURALIST Vol. 93 *Circaea alpina (Enchanter’s Nightshade). 24758 (3). This circumpolar species is known from northern British Columbia at Liard Hot Springs (Porsild and Crum 1961), and in the adjacent District of Mac- kenzie in the headwaters of the Flat River and along the Liard River (Scotter and Cody 1974). * Aralia nudicaulis (Wild Sarsaparilla). 24427 (1). Porsild and Crum (1961) found this species at Liard Hot Springs, while Jeffrey (1961) and Scotter and Cody (1974) reported it from several sites in the southwestern District of Mackenzie. Cicuta maculata var. angustifolia (Water Hemlock). 24739 (2). Only one other collection is known from the Yukon Territory by the authors: Tatchun River, Mile 117.5 Mayo Highway, Taylor 4080 (DAO). * Osmorhiza depauperata (O. obtusa) (Sweet Cicely). 24479 (1); 24687 (2). Recorded from adjacent northern British Columbia at Liard Hot Springs by Porsild and Crum (1961), and from the Liard River Valley in southwestern District of Mackenzie by Cody (1963). Collomia linearis. 24730, 24742 (2). This is a native North American species which Hultén (1968) considers to be introduced inthe Yukon Territory. It seems doubtful that it was introduced to this remote and pristine site. Previous collections, according to Hultén’s map, are all west of longitude 135°W. Scutellaria galericulata var. pubescens (Skullcap). 2ATTLA (3). Previously known from the Yukon Territory only from the vicinity of Mayo (Porsild 1951, 1974; Hultén 1968). Castilleja caudata. 2456S (1). This collection extends the known range of this species to the southeast from western and northern Yukon Territory. * Castilleja miniata (Red Indian Paint-brush). 24683, 24728 (2). This is a northward extension of range from north central British Columbia. Mimulus guttatus (Yellow Monkey-flower). 24306, 24339 (1); 24678 (2). According to Hultén (1968) and Porsild (1974) this species barely enters the southern part of the Yukon Territory near Whitehorse and Watson Lake. It has been reported from mineral springs in Nahanni National Park (Scotter and Cody 1974). The plant was very abundant at the Coal River springs. 1979 Galium triflorum (Sweet-scented Bedstraw). 24516 (1); 24675 (2). This species, according to the map in Hultén (1968), barely enters the southern part of the Yukon Territory west of 133°W longitude. Our collections will move the delimiting line eastward to include the extreme southeastern Yukon Territory. In the adjacent Dis- trict of Mackenzie it is known along the Liard River (Cody 1961: Jeffrey 1961) and in Nahanni National Park (Scotter and Cody 1974); in northern British Columbia it has been found at Liard Hot Springs (Porsild and Crum 1961). * Anaphalis margaritacea var. subalpina (Pearly Ever- lasting). 24727 (2). The site at Hole-in-the-Wall Lake, Nahanni Na- tional Park, reported by Porsild (1961) was widely disjunct from the Alaska Panhandle sites shown on the map in Hultén (1968). The specimen reported here helps fill the gap between these widely separated areas. Arnica cordifolia. 24387 (1). This collection comes from an area intermediate between the sites shown on the map in Hultén (1968), and an apparently isolated site on the lower Liard River reported by Raup (1947). Aster alpinus ssp. vierhapperi. 24783 (1). This collection from the extreme southeastern part of the Yukon Territory will move the delimiting line in Hultén (1968) to the east, thus bringing it close to the populations in southwestern District of Mackenzie cited by Scotter and Cody (1974). * Aster ciliolatus. 24738 (2): 24775 (3). These collections extend the known distribution of A. ciliolatus to the west from Fort Liard on the Liard River in southwestern District of Mackenzie (Jeffrey 1961). Aster modestus. 24399 (1): 24356, 24648, 24784 (2). Hultén (1968) indicated a collection from near Dawson and Porsild and Crum (1961) reported this aster at Liard Hot Springs in northern British Columbia. * Bidens cernua (Nodding Beggar-ticks). 24767, 24774 (3). In central Alaska this species is known from the Yukon River drainage (Hultén 1968) while in the District of Mackenzie it is known from Nahanni National Park, Great Slave Lake, and the Slave River. Erigeron philadelphicus (Pink Fleabane). 24336, 24340, 24350 (1); 24608, 24647, 24676 (2). Hultén (1968) indicates a single Yukon Territory locality near Old Crow, from whence it has also been reported by Wein et al. (1974). Porsild and Crum (1961) reported it at Liard Hot Springs. SCOTTER AND CODY: VASCULAR PLANTS, YUKON TERRITORY 169 Hieracium scabriusculum. 24757, 24759B (3). Hultén (1968) indicates two sites for this species, one near Watson Lake and another near Dawson. Solidago canadensis var. salebrosa (Canadian Gol- denrod). 2474] (2). Hultén (1968) indicates collections from the vicinity of Dawson and Watson Lake. Literature Cited Arnold, E. W. 1961. Plant communities of a hot spring in the Mackenzie Mountains, Northwest Territories. M.Sc. thesis, University of Michigan, Ann Arbor. 34 pp. Bassett, I. J.. C. W. Crompton, and D. W. Woodland. 1974. The family Urticaceae in Canada. Canadian Journal of Botany 52: 503-516. Bostock, H.S. 1948. Physiography of the Canadian Cor- dillera, with special reference to the area north of the fifty-fifth parallel. Geological Survey Canada, Memoir 247. 106 pp. Cody, W. J. 1961. New plant records from the upper Mackenzie River Valley, Mackenzie District, Northwest Territories. Canadian Field-Naturalist 75: 55-69. Cody, W. J. 1963. A contribution to the knowledge of the flora of southwestern Mackenzie District. Canadian Field-Naturalist 77: 108-123. Cody, W. J.and A. E. Porsild. 1968. Additions to the flora of continental Northwest Territories, Canada. Canadian Field-Naturalist 82: 263-275. Douglas, R. J.W. and D. K. Norris. 1959. Fort Liard and La Biche map-areas, Northwest Territories and Yukon. Geological Survey Canada, Paper 59-6. 23 pp. Gabrielse, H. and S. L. Blusson. 1969. Geology of Coal River map-area, Yukon Territory and District of Mac- kenzie. Geological Survey Canada, Paper 68-38. 22 pp. Hultén, E. 1968. Flora of Alaska and neighboring terri- tories. Stanford University Press, Stanford. 1008 pp. Jeffrey, W. W. 1961. Notes on plant occurrence along lower Liard River, Northwest Territories. National Mu- seum of Canada Bulletin 171: 32-115. Oswald, E. T. and J. P. Senyk. 1977. Ecoregions of Yukon Territory. Department of Fisheries and Environment, — Canadian Forestry Service Report BC-X-164. 115 pp. Porsild, A. E. 1951. Botany of southeastern Yukon adja- cent to the Canol Road. National Museum of Canada Bulletin 121: 1-400. Porsild, A. E. 1961. The vascular flora of an alpine valley in the Mackenzie Mountains. National Museum of Canada Bulletin 171: 116-130. Porsild, A. E. 1966. Contributions to the flora of south- western Yukon Territory. National Museum of Canada Bulletin 216: 1-86. Porsild, A. E. 1974. Materials for a flora of central Yukon Territory. National Museum of Canada, Publications in Botany, Number 4: 1-77. Porsild, A. E. and H. Crum. 1961. The vascular flora of Liard Hot Springs, B.C., with notes on some bryophytes. National Museum of Canada Bulletin 171: 131-197. Porsild, A. E. and W. J. Cody. 1979. Vascular plants of continental Northwest Territories. National Museums of Canada, Publications in Botany, Number 8. (in press.) 170 THE CANADIAN FIELD-NATURALIST Raup, H. M. 1936. Phytogeographic studies in the Atha- baska-Great Slave Lake region. I, Catalogue of the vascular plants. Journal of the Arnold Arboretum 17: 180-315. Raup, H.M. 1947. The botany of southwestern Mac- kenzie. Sargentia 6: 1-275. Rowe, J. S. 1972. Forest regions of Canada. Department of the Environment, Canadian Forest Service Publication Number 1300. 172 pp. Scotter, G. W. and W. J. Cody. 1974. Vascular plants of Nahanni National Park and vicinity, Northwest Terri- tories. Naturaliste Canadien 101: 861-891. Vol. 93 Wein, R. W., L. R. Hettinger, A. J. Janz, and W. J. Cody. 1974. Vascular plant range extensions in the northern Yukon Territory and northwestern Mackenzie District. Canada. Canadian Field-Naturalist 88: 57-66. Welsh, S. L. 1974. Anderson’s flora of Alaska and adjacent parts of Canada. Brigham Young University Press, Provo, Utah. 724 pp. Received 15 August 1978 Accepted 22 January 1979 Notes Fall Foods of Common Snipe on the Copper River Delta, Alaska STANLEY E. SENNER and P. G. MICKELSON Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99701 Senner, Stanley E. and P. G. Mickelson. 1979. Fall foods of Common Snipe on the Copper River Delta, Alaska. Canadian Field-Naturalist 93(2): 171-172. Stomachs from 22 Common Snipe (Capella gallinago delicata) collected at three sites on the Copper River Delta, Alaska, were examined to determine food habits. Cranefly larvae, Prionocera sp.(1 ipulidae) were the most important food items and occurred in 75 to 86% of the stomachs. These and beetle larvae were the major foods, as previously shown for other breeding and staging areas in Canada. Key Words: Common Snipe, food habits, Copper River Delta, Alaska. Common Snipe (Capella gallinago delicata) breed primarily across boreal North America (Tuck 1972), although some are found breeding in arctic regions (e.g., Colville River, Alaska (Kessel and Cade 1958)). Most literature on snipe in Alaska concerns their distribution (e.g., Gabrielson and Lincoln 1959). There are no previous studies on food habits of snipe in Alaska, although Tuck (1972) described food habits of snipe at breeding and fall concentration sites in Canada. During fall the Copper River Delta in south- central Alaska may host several 10 000s of snipe (isleib and Kessel 1973). The majority of these migrate south, presumably to wintering grounds from British Columbia to California (Burleigh 1955; Tuck 1972), although snipe rarely have been recorded in mid- winter on the western Copper River Delta (Isleib and Kessel 1973). Study Area Snipe were collected at three sites on the Copper River Delta. Nine snipe were from Mile 20 of the Copper River Highway (60°27’N, 145°16’W). Carex spp., Eleocharis spp., Equisetum spp., and Hippuris vulgaris dominated this meadow habitat. Five snipe were collected in mixed sedge-grass-forb habitat dominated by Carex lyngbyaei, Deschampsia sp., Festuca rubra, and Hedysarum alpinum along lower Alaganik Slough (60°26’N, 145°17’W). Eight snipe were from the east Copper River Delta (60°17’N, 144°56’W) in habitat dominated by A/nus crispa, Myrica gale, and Carex lyngbyaei. Methods All snipe were taken between | September and 10 October in 1976 (10 specimens) and 1977 (12 specimens). Contents of the snipe gizzard and proventriculus, and sometimes the esophagus, were removed and placed in Whirl-paks with hexamethy- lenetetramine-buffered formalin (Mueller 1972). Time between collection of the bird and removal of stomach contents varied up to several hours. Because of differential digestion rates and post-mortem digestion (e.g., Tuck 1972), only the number of stomachs with recognizable items are reported here. Tuck (1972) suggested that frequency of occurrence (Hartley 1948) might be more meaningful than volume or weight for presenting food habits data. Our results are expressed in frequency of occurrence because volume or weight assigns great importance to large or difficult-to-digest items of food and underemphasizes the importance of small or easily digested food items. Vegetable debris, seeds, and mineral grit were not considered as food (Tuck 1972). White and Harris (1966) and Tuck (1972) found that plant fibers and seeds remained relatively unchanged by any digestive process and were finally regurgitated. Results and Discussion Nineteen of 22 snipe collected on the Copper River Delta contained food items (Table |). The most important food items, represented by up to 43 | individuals in a single stomach, were Cranefly larvae, Prionocera sp. (Tipulidae), which were present in 75 to 86% of the stomachs from all three sites (Table 1). The mean length of 26 intact larvae was 24.4 + 5.82 mm. Beetles, especially larval Dytiscidae (probably Agabus sp.) were the second most important item in the diet of snipe during fall (Table 1). These occurred in 14 to 75% of the stomachs from snipe collected from the three sites. Both cestodes and nematodes were found in snipe stomachs at Mile 20 (Copper River Highway) and Alaganik Slough, but neither was found in snipe taken on the east Copper River Delta. The nematodes did not appear to be parasitic on the snipe. The results of this study agree with those sum- marized by Tuck (1972) in that insects, particularly 171 172 TABLE 1—Frequency of occurrence of food items in the stomachs of 19 Common Snipe collected at three sites on the Copper River Delta, Alaska Percent frequency of occurrence East Copper Alaganik River Food Mile 20 Slough Delta categories (N=8) (N=4) (N=7) Animal taxa Cestoda IDES 50.0 — Nematoda 25.0 25.0 — Lumbricidae 12.5 -— — Gastropoda, undetermined 12.5 _ — Pelecypoda, undetermined 12.5 _ _— Insecta, undetermined — — 14.3 Diptera (total) 75.0 100.0 85.7 Tipulidae Prionocera sp. larvae 75.0 75.0 85.7 Pedicia sp. larvae a 25.0 — Chironomidae larvae = 25.0 28.6 Coleoptera (total) 50.0 75.0 14.3 Undetermined adult — 25.0 14.3 Dytiscidae larvae (largely Agabus sp.? and Hydroponidae) 375 75.0 14.3 Carabidae adult 12.5 — --- Staphylinidae adult — — 14.3 Araneida, undetermined a 25.0 — Non-food items! Vegetable debris 87.5 75.0 100.0 Seeds 87.5 75.0 85.7 Mineral grit 100.0 75.0 71.4 1See Tuck (1972). Diptera and Coleoptera larvae, are taken frequently. Tuck (1972) mentions that Tipulidae are especially significant, primarily on snipe breeding grounds, but also on fall staging areas in Canada. In northern latitudes Tipulidae are of great importance to other scolopacid shorebirds (see Chernov 1967; Holmes and Pitelka 1968). Our study has confirmed their value on THE CANADIAN FIELD-NATURALIST Vol. 93 an important staging and breeding ground for Common Snipe along the north Gulf of Alaska coast. Acknowledgments We thank Daniel D. Roby, William A. Lehn- hausen, John W. Smith, Brian E. Lawhead, and Stephen Murphy for assistance in collecting snipe. George J. Mueller, Stephen F. MacLean, Jr., and Jacqueline D. LaPerriére helped identify invertebrate specimens. Stephen F. MacLean, Jr., James C. Bartonek, Leslie Tuck, and D. Keppie reviewed the manuscript. Literature Cited Burleigh, T. D. 1955. Investigations of woodcock, snipe and rails in 1954. U.S. Fish and Wildlife Service Special Scientific Report, Wildlife 28. 62 pp. Chernov, Y.I. 1967. [Trophic relationships of birds to insects in the tundra zone.] Ornitologi’ya 8: 133-150. (Translated from Russian by S. F. Maclean, Jr.) Gabrielson, I. N. and F. C. Lincoln. 1959. Birds of Alaska. Wildlife Management Institute, Washington, D.C. 922 pp. Hartley, P. H. 1948. The assessment of the food of birds. Ibis 90: 361-381. Holmes, R.T. and F.A. Pitelka. 1968. Food overlap among coexisting sandpipers on northern Alaskan tundra. Systematic Zoology 17: 305-318. Isleib, M. E. “Pete” and B. Kessel. 1973. Birds of the North Gulf Coast-Prince William Sound Region, Alaska. Biological Papers of the University of Alaska Number 14. 149 pp. Kessel, B. and T. J. Cade. 1953. Birds of the Colville River, northern Alaska. Biological Papers of the University of Alaska Number 2. 83 pp. Mueller, G. J. 1972. Field preparation of marine specimens. University of Alaska Museum, Fairbanks. 44 pp. Tuck, L. M. 1972. The snipes. Canadian Wildlife Service Monograph Number 5. 428 pp. White, M. and S. W. Harris. 1966. Winter occurrence, foods, and habitat use of snipe in northwest California. Journal of Wildlife Management 30(1): 23-34. Received 5 April 1978 Accepted 23 November 1978 1979 NOTES 73} Ivory Gull Colonies in Southeastern Ellesmere Island, Arctic Canada T. FRISCH and W. C. MORGAN Geological Survey of Canada, 588 Booth Street, Ottawa, Ontario KIA 0E4 Frisch, T. and W. C. Morgan. 1979. Ivory Gull colonies in southeastern Ellesmere Island, Arctic Canada. Canadian Field-Naturalist 93(2): 173-174. Five Ivory Gull colonies, ranging in population from about 12 to 60 birds, have been discovered in the upland icefields of southeastern Ellesmere Island. The colonies occur on cliff-faces of precipitous nunataks as much as 26 km from the nearest sea coast. One colony is established as a nesting site; the others are probable breeding places. Key Words: Ivory Gull, Ellesmere Island, Arctic, colonies, nunatak nesting site. Only one active breeding colony of the Ivory Gull (Pagophila eburnea) has heretofore been known from the North American Arctic: Seymour Island, a small low island north of Bathurst Island (MacDonald 1976). In the summer of 1977, while engaged in a helicopter-supported geological survey, the authors found five colonies of Ivory Gulls in the highlands of southeastern Ellesmere Island, an environment very different from that of Seymour Island. This note records the newly-discovered localities and our observations. The area of Ellesmere Island that contains the colonies is bounded by latitude 78°15’N and longi- tude 81° W and traversed by a major fiord, Makinson Inlet (Figure 1). Most of the land area comprises upland icefields and has a maximum relief of 1500 m; rock exposures are found mainly in coastal cliffs and nunataks. The terrain shown in Figure 2 lies north of Makinson Inlet between colonies 4 and 5 (Figure 1) and is representative of much of the area, which is underlain largely by metamorphic and granitic rocks of the Canadian Shield. The only permanent human settlement in the area is the small Inuit community of Grise Fiord, on the south coast of Ellesmere Island. Positions of the five Ivory Gull colonies discovered are plotted on Figure | and more accurate locations in the Universal Transverse Mercator Grid system on National Topographic System 1:250 000 maps are given below. All sites are on rock ledges near the tops of steep cliff-faces of nunataks several hundred metres high at elevations of 450 to 750 m above sea-level. Only site 2, which was located purely by chance after the helicopter had landed and shut down, was observed at relatively close range. Owing to their inaccessibility, and lack of time and/or the threat of excessive disturbance by the helicopter, the other sites could not be examined closely. Numbers of gulls seen are very approximate and are based chiefly on estimates made during a single fly-past. Information on each colony is listed in the following order: NTS 1:250 000 map sheet; approxi- mate location on map; UTM easting and northing in Zone 17; approximate elevation above sea-level; distance to nearest sea coast; date of sighting; observer. Other details follow. Site 1. Craig Harbour 49A; NE corner; 508600, 8532300; 750 m; 23 km; 11 August; TF. Sixty birds flying around cliff ima “80 u TI ee jee Bae ~~ = q eS = Adogan Inlet as Ss Lagi ny! BS tie eC nN 184 4, Ne ee ae Talbot Inlet SS } y ww ‘eas a 4 i Ge ? BAF FIN BaeAw 114 \, (¢ YW )Clarence ( Head \ i Bes) ee 5 ey We = ¢ aS ( X ff ie i pa ire we ) ( ; 25km i J 1 1 FIGURE |. Sketch-map of southeastern Ellesmere Island showing five sites of Ivory Gullcolonies. Broken lines mark the edges of icefields and glaciers but nunataks are not shown. 174 THE CANADIAN FIELD-NATURALIST Vol93 FIGURE 2. Typical terrain north of site 4. and resting on ledges. A flat area on top of the nunatak was free of snow but not occupied by birds. Site 2. Clarence Head 39B; NW corner, 526000, 8537400; 450 m; 14km; 12 August; WCM. Twelve to 15 birds seen near top of cliff. Four large young birds visible by eye in nests on excrement-stained ledges. No unhatched eggs were visible through binoculars. Observations were made from an icefield looking down on the site across a gully. Site 3. Talbot Inlet 39C and part of 39D; Thorndike Peaks; 541200, 8565000; 600 m; 6 km; 12 August; WCM. Fifty birds in the air, resting on ledges, and taking off from ledges. Site 4. Talbot Inlet 39C and part of 39D; Inglefield Mountains; 543600, 8597600; 600 m; 12 km; 29 June; WCM. At least 50 birds suddenly encountered when helicopter flew around corner of nunatak. Site 5. Ekblaw Glacier 39F and 39E; head of Cadogan Glacier; 549900, 8679000; between 600 and 750 m; 26 km; 15 August; TF. Thirty birds on ledges and flying around nunatak. Site 2, southwest of Smith Bay (Figure 1), is thus established as a new breeding place of the Ivory Gull. Although we have no direct evidence of breeding at the other localities, we consider it more than probable that these are also nesting sites. Nesting sites of the Ivory Gull similarly situated on nunataks well inland from the coast have been reported from southern Vestspitsbergen by Birken- majer and Skreslet (1963) (see also Lovenskiold 1964). No doubt sites of this type are used because, among other factors, they provide a haven safe from predators. Furthermore, Polar Bear (Ursus mari- timus) kills provide a major source of food for the Ivory Gull (MacDonald 1967, p. 6) and all the Ellesmere Island sites are relatively near the east coast, which, according to our observations, supports a sizable bear population, particularly in Smith Bay. Our chiefly aerial coverage of southeastern Elles- mere Island was by no means complete and, judged from essentially chance discoveries of five colonies and a number of sightings of individual Ivory Gulls, it seems certain that more colonies exist in the upland icefields. Also, our sightings lend credence to uncon- firmed reports from the 19th century. of a breeding place on the east coast of Ellesmere Island, north of the area described here (MacDonald and Macpherson 1962). In any event, southeastern Ellesmere Island is now established as an important breeding ground of the Ivory Gull. Acknowledgments We thank S. D. MacDonald, D. N. Nettleship, and H. Ouellet for reviews of this note and the Polar Continental Shelf Project for support in the field. Literature Cited Birkenmajer, K. and S. Skreslet. 1963. Breeding colony of ivory gulls in Torell Land, Vestspitsbergen. Norsk Polar- institutt Arbok 1962: 120-126. Lovenskiold, H. L. 1964. Avifauna Svalbardensis. Norsk Polarinstitutt Skrifter 129. MacDonald, S. D. 1976. Phantoms of the polar pack ice. Audubon 78(3): 2-19. MacDonald, S. D. and A. H. Macpherson. 1962. Breeding places of the Ivory Gull in Arctic Canada. National Museum of Canada Bulletin 183. pp. 111-117. Received 23 August 1978 Accepted 17 November 1978 79 NOTES 175 Wind-caused Death of Great Cormorant MARTIN K. MCNICHOLL! and GEOFFREY G. HOGAN Department of Biological Sciences, Brock University, St. Catharines, Ontario L2S 3A1 'Present Address: Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta T2E 6M7 MeNicholl, Martin K. and Geoffrey G. Hogan. 1979. Wind-caused death of Great Cormorant. Canadian Field- Naturalist 93(2): 175. Key Words: Accidental death, Great Cormorant, Prince Edward Island, weather, wind. At 18:32 on 10 April 1977, McNicholl watched an adult Great Cormorant (Phalacrocorax carbo) fly from the open sea to ledges near the top of a cormorant colony cliff at Cape Tryon on the northern shore of Prince Edward Island. Strong north winds, rated as 5 or more on the Beaufort scale ( = 29-38+ km/h), were blowing at the time, and visibility was reduced by blowing snow, although the snow also heightened light levels. The bird tried three times to land, but each time gusts caused it to circle back towards the sea. On the fourth approach, the bird was gliding into a position from which it could land when a sudden gust of wind threw it breast-first into the cliff face. The bird flapped one wing twice, then ceased all motion. A Common Raven (Corvus corax) attempted to land near the corpse 2 min later, but was unable to do so. The next day a raven was eating the dead bird. On 4 April, another day of strong north winds with sudden gusts, McNicholl observed another adult Great Cormorant to be blown about 30 m along the cliff-face on attempting to land. This bird was able to land briefly at the new locale along the cliff, but was driven away by other birds there. These observations were made during regular checks of the colony from a cleft in the cliff which hid the observer from birds not actually perched on ledges on the cliff. This cleft was over 200 m from the spot where the bird was killed. McNicholl visited the colony from 22 March to 22 April during which period cormorant activity declined markedly on days with gusty winds, but birds were able to land readily in high winds if these were steady. Byrd and Tobish (1978) attributed the deaths of 48 chick and two adult kittiwakes ( Rissa brevirostris and R. tridactyla) to wind during a violent storm in Alaska, but did not observe the precise manner of deaths. There are many reports in the literature of deaths of birds during storms, and several reports of deaths by collison with man-made objects (see review by Weir 1976), but we are not aware of other reports of deaths caused by collisions at familiar colony sites in winds not associated with violent storms. The above observations suggest that strong gusty winds could pose a serious problem to large seabirds at cliff colonies. Stonehouse (1964) mentioned that frigate- birds (Fregatidae) have difficulty in landing during strong winds, but did not attribute deaths to this cause. These observations were made during a study by Hogan under the supervision of R.D. Morris. Financial support was provided by the Prince Edward Island Fish and Wildlife Division of the Department of the Environment, a National Research Council of Canada grant to R.D. Morris, and an Ontario Graduate Scholarship to Hogan. The manuscript was improved on the basis of comments by R. K. Ross and an anonymous reviewer. Literature Cited Byrd, G. V. and T. G. Tobish. 1978. Wind-caused mortali- ty in a kittiwake colony at Buldir Island, Alaska.: Murrelet 59: 37. Stonehouse, B. 1964. Frigatebird. Jn A new dictionary of birds. Edited by A. L. Thomson. McGraw-Hill, New York. pp. 325-326. Weir, R. D. 1976. Annotated bibliography of bird kills at man-made obstacles: a review of the state of the art and solutions. Canadian Wildlife Service, Ontario Region, Ottawa. Received 18 September 1978 Accepted 17 November 1978 176 THE CANADIAN FIELD-NATURALIST Vol. 93 Habitat Selection by Wintering Snowy Owls (Nyctea scandiaca) M. Ross LEIN and G. A. WEBBER Department of Biology, University of Calgary, Calgary, Alberta T2N 1N4 Lein, M. Ross and G. A. Webber. 1979. Habitat selection by wintering Snowy Owls (Nyctea scandiaca). Canadian Field- Naturalist 93: 176-178. Habitat selection by Snowy Owls (Nyctea scandiaca) wintering near Calgary, Alberta, was studied in 1973-1976. Habitats were classified as being residential, fallow, stubble, hayfield, or pasture, and areas of each were determined. Compared to the relative areas of the various habitats, owls were seen more frequently in stubble and hayfields, and less frequently in the other three habitat categories. This selection is believed to be related to the varying densities and accessibility of small mammals in the different habitats. Key Words: Snowy Owls, habitat selection, winter, prairie farmland, prey availability. Studies of the behavioral ecology of the Snowy Owl (Nyctea scandiaca) in winter are sporadic and incomplete. Gross (1927, 1931, 1944, 1947) docu- mented several irruptions between 1926 and 1946 but did not obtain comprehensive information on aspects of the owls’ ecology other than food habits. Most subsequent publications deal with unusual locality records or numbers of birds, or with food habits, although a few (Keith 1960, 1964: Quilliam 1965: Weir 1973) consider other behavioral or ecological aspects. The virtual lack of intensive or long-term investigations of the biology of the species in areas where it regularly winters is especially Surprising since some owls may actually spend more time in the south than on their arctic breeding grounds (Bird 1972). The senior author began a long-term study of wintering Snowy Owls near Calgary, Alberta in 1973. Snowy Owls have been recorded in numbers in this area every winter since 1964-65 (Bird 1972: personal observations). This note reports some initial findings of this investigation on the habitats utilized by owls on their winter range. Methods Each winter from 1973-74 to 1975-76, 8-12 survey routes were censused by members of the Calgary Field Naturalists’ Society, using a modification of Gollop’s (1965) procedure. Each 64-km (40-mi) route was surveyed every 2-4 wk by two volunteers who drove the route at a speed of 30-50 km/h, stopping every 3.2 km (2.0 mi) to scan the surrounding area with binoculars. Each owl’s location was plotted on a topographic map and pertinent standardized data, including a habitat description, were recorded. This analysis is limited to one route in 1974—75 and three routes in 1975-76 because of the effort involved in calculating the habitat composition of routes and the availability of recent aerial photography. The routes chosen were the most productive in terms of total owls recorded, and were located in mixed agricultural land, just east of Calgary. A habitat map was prepared for each route, including an area extending 1.6 km (1.0 mi) on each side of the survey route and 1.6 km before the beginning and after the ending of the route. The nature of each field or other habitat unit was identified from aerial photos or by direct inspection. All habitat determinations were made after cultivation ceased in the fall and prior to its resumption in the spring. Areas of each habitat on each route were estimated using the standard dot grid method. We recognize five habitat types on the study area: 1. Residential: farmyards, rural residences, feedlots, hamlets, and other areas of high human activity. 2. Summerfallow: cultivated fields where either no crop was planted during the previous growing season or the stubble was ploughed up after harvest. These areas are virtually bare earth with only traces of plant material on the surface. 3. Stubblefields: areas where cereal crops have been grown and harvested, leaving only the basal 10-20 cm of the stems. Fields usually persist in this form until ploughing and seeding in the following spring. 4. Hayfields: areas planted to non-native grasses and cut yearly for livestock fodder. They are rarely used as pasture and are ploughed and reseeded at intervals of longer than one year. 5. Pastures and Sloughs: areas not subjected to annual cultivation for the raising of cereal crops or hay, often marginal land which may be used as pasture for livestock. Sloughs fill with spring runoff but are typically dry throughout much of the summer, autumn, and winter. This is a highly heterogeneous category which is not easily subdivided. We tallied the survey results by the habitat in which -each owl was located when sighted. When an owl was perched on a fenceline that represented a boundary IA) between two different habitats, a value of 0.5 owl was assigned to each of the two categories. Results The distribution of habitat classes on the survey routes is presented in Table 1. The major category is stubblefields, comprising almost 60% of the area. Residential areas are a very minor category. The distribution of owl sightings across these habitat classes is also shown in Table |, along with an expected distribution of sightings based on a null hypothesis of no habitat selection. The observed distribution is significantly different from that expec- ted (chi-square goodness of fit test, P< 0.01), demon- strating a degree of habitat selection. Owls are recorded more frequently than expected in stubble and hayfields, and less frequently than expected in the other three habitat categories. Discussion The overall preference for open habitats by winter- ing Snowy Owls is well-known (Bent 1938: Gross 1947: Keith 1960). In the Calgary area, most owls are found in agricultural land east of the city: few birds are observed in the more wooded aspen parklands or foothills to the west (Bird 1972; M.R. Lein, un- published data). A similar avoidance of wooded regions is noted elsewhere (Gross 1947: Nagell and Frycklund 1965); however, the present study appears to be the first attempt to demonstrate habitat selection by Snowy Owls at a finer level. We suggest that the preference for stubblefields and hayfields is related to the availability of small- mammal prey in such areas, as would be predicted by optimal foraging theory (MacArthur and Pianka 1966; Pyke et al. 1977). Summerfallow fields are relatively barren and offer little to rodents either in cover or food. Pastures are usually heavily grazed and offer little cover for small mammals. Non-grazed sloughs and roadside ditches offer excellent cover for rodents, which are relatively abundant in such habitats (P. C. Boxall, unpublished data), but rank vegetation and deep snow cover may sharply limit NOTES 177 their availability to owls. Low numbers of owls around residential areas may be a direct response to human disturbance. Thus, the habitat distribution of the owls suggests that they may be choosing areas (stubblefields and hayfields) in which rodents are both relatively numerous (P. C. Boxall, unpublished data) and available to predators. It is perhaps surprising that a more pronounced degree of habitat selection is not shown. Several reasons may be suggested. The first is the relatively crude level of the analysis. Owls certainly do not categorize habitats in the same manner as do human investigators, and an analysis of distribution using different habitat classes might show more striking differences. Second, and perhaps more important, is the implicit assumption of the present study that the owls recorded in a habitat were actually choosing it as a hunting area. Although hunting attempts have been observed in all habitats throughout the day, it is possible that owls may choose different areas for roosting and “loafing” than those in which they hunt. We know very little about the hunting behavior or activity cycles of wintering Snowy Owls. Owls tend to use elevated perches such as trees or utility poles early in the morning and toward sunset (Keith 1964: M. R. Lein, unpublished data) and to perch on or near the ground during mid-day. It has been suggested that these high perches are hunting perches and that this represents a diurnal pattern of hunting behavior. Unfortunately, most of the data used in this study were gathered during the middle of the day and are unsuitable for a test of such an hypothesis. If it were possible to identify owls likely to be actively hunting, and to carry out the analysis on only those owls, stronger habitat selection might be expected. Acknowledgments We thank those members of the Calgary Field Naturalists’ Society who conducted owl surveys: this study would not have been possible without their efforts. We gratefully acknowledge P. C. Boxall for providing us with preliminary results of his investiga- tion on Snowy Owls, and for commenting on the TABLE |—Distribution of habitats, and of owl sightings, on the survey routes analyzed. Habitat category Pasture Residential Fallow Stubble Hayfield + slough Total Percentage of total area 1.81 Da 58.48 6.55 18.04 100.00 Number of owls observed 2.5 20.5 116.0 18.0 18.0 175.0 Number of owls expected Ball) 26.46 102.34 11.46 SiESy/ 175.00 178 THE CANADIAN FIELD-NATURALIST manuscript. T. E. Dickinson also provided helpful suggestions. Financial support was received in the form of operating grants from the University of Calgary and the National Research Council of Canada to M. R. Lein. Literature Cited Bent, A. C. 1938. Life histories of North American birds of prey. Part 2. United States National Museum Bulletin Number 170. Bird, C. D. 1972. The Snowy Owl in the Calgary area, 1964-1972. Calgary Field Naturalist 3: 106-107. Gollop, J. B. 1965. Snowy Owls and Gray Partridge near Saskatoon in the winter of 1963-1964. Blue Jay 23: 28-31. Gross, A. O. 1927. The Snowy Owl migration of 1926-27. Auk 44: 479-493. Gross, A. O. 1931. Snowy Owl migration — 1930-31. Auk 48: SOI-S11. Gross, A. O. 1944. Food of the Snowy Owl. Auk 61: I-18. Gross, A. O. 1947. Cyclic invasions of the Snowy Owl and the migration of 1945-46. Auk 64: 584-601. Vol. 93 Keith, L. B. 1960. Observations on Snowy Owls at Delta, Manitoba. Canadian Field-Naturalist 74: 106-112. Keith, L. B. 1964. Territoriality among wintering Snowy Owls. Canadian Field-Naturalist 78: 17-24. MacArthur, R. H. and E. R. Pianka. 1966. On optimal use of a patchy environment. American Naturalist 100: 603- 609. Nagell, B. and I. Frycklund. 1965. Invasionen av fjalluggla (Nyctea scandiaca) i sodra Skandinavien vintrarna 1960- 1963 samt nagot om artens beteende pa Overwintring- slokalerna. Var Fagelvarld 24: 26-55. Pyke, G.H., H.R. Pulliam, and E. L. Charnoy. 1977. Optimal foraging: a selective review of theory and tests. Quarterly Review of Biology 52: 137-154. Quilliam, H.R. 1965. Winter study of Snowy Owls on Wolfe Island, 1965. Ontario Field Biologist 19: 1-8. Weir, R. D. 1973. Snowy Owl invasion on Wolfe Island, winter 1971-72. Ontario Field Biologist 27: 3-17. Received 21 March 1978 Accepted 17 November 1978 Two-lined Salamander, Eurycea bislineata, in Labrador FRANCIS R. COOK! AND JOHN PRESTON2 'Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8 2Environment Canada, 814 Hudson Drive, Labrador City, Newfoundland A20O I1M5 Cook, Francis R. and John Preston. 1979. Two-lined Salamander, Eurvcea bislineata, in Labrador. Canadian Field- Naturalist 93(2): 178-179. In mid-August 1972 a single Two-lined Sala- mander, Eurycea bislineata, was collected by J. P. approximately 16 km (10 mi) NE of Labrador City on the west side of a small brook that drains into Moose Head Lake, about 0.6 km (1 mi) upstream from the lake (52°58’N, 66°40’W). The specimen is a recently transformed juvenile 63mm in total length and 29.5mm snout to anterior margin of vent. It is catalogued as National Museum of Natural Sciences (Herpetology Section) 17948. This is the first example of the species collected in Labrador and is a 290-km (180-mi) range extension north of the nearest records on the north shore of the Gulf of St. Lawrence (Bleakney 1958, map 85: Logier and Toner 1961, map 19; Power 1965). Based on two collections which have not been previously docu- mented, Conant (1975, map 240) shows that in interior central Quebec the range of the Two-lined Salamander extends north to near the latitude of southern Labrador. These collections are a single specimen from the Lac Charon area 32 km (20 mi) from Chibougamau taken by Jean-Guy Rossignol 3 July 1967 (NMNS 10287), and five specimens col- lected along the Rupert River, 48 km (30 mi) west of its source at Lake Mistassini approximately 20 July 1967 by Daniel Roy (American Museum of Natural History 88025-29; transmitted by Roger Conant). Bleakney (1968, map 82) plotted a record of the Red- backed Salamander, Plethodon cinereus, in the Lake Melville area of central Labrador. This was based ona report from W. J. Smith who in the summer of 1955 had the local occurrence of a small salamander mentioned to him on several occasions (Bleakney 1958, p. 15). Because of gross similarities between Plethodon and Eurycea, however, we suggest that this record may be more logically assigned to the latter and 1979 suggest a wider Labrador range for it than our single specimen establishes. The presence of Eurycea in Labrador and central Quebec is a further indication of the need for a northward revision of Bleakney’s Herpetofaunal Section 5 to include his previously disjunct Section 5A as has already been proposed by Cook and Folinsbee (1975) on the basis of recent records for the Blue-spotted Salamander, Amby- stoma laterale, the only other salamander verified for Labrador to date. Literature Cited Bleakney, J. Sherman. 1958. A zoogeographical study of the amphibians and reptiles of eastern Canada. National Museum of Canada Bulletin 155: 1-119. NOTES 179 Conant, Roger. 1975. A field guide to reptiles and am- phibians of eastern and central North America. Houghton Mifflin Company, Boston. xviii + 429 pp. Cook, Francis R. and John Folinsbee. 1975. Second record of the Blue-spotted Salamander from Labrador. Canadian Field-Naturalist 89(3): 314-315. Logier, E. B.S. and G. C. Toner. 1961. Check list of the amphibians and reptiles of Canada and Alaska. Royal Ontario Museum, Toronto, Life Sciences Division Con- tribution 53: 1-92. Power, G. 1965. Notes on the cold-blooded vertebrates of the Nabisipi River region, County Duplessis, Quebec. Canadian Field-Naturalist 79(1): 49-61. Received 9 November 1978 Accepted 4 December 1978 First Canadian Record of a Flathead Catfish E. J. CROSSMAN! and J. H. LEACH? ‘Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 2Ontario Ministry of Natural Resources, Fisheries Research Station, RR #2, Wheatley, Ontario NOP 2P0 Crossman, E. J. and J. H. Leach. 179-180. 1979. First Canadian record of a Flathead Catfish. Canadian Field-Naturalist 93(2): Pylodictis olivaris was captured in the Canadian waters of Lake Erie on 22 August 1978. This is a 30-km range extension north and is the first Canadian record. Key Words: Flathead Catfish, Pv/odictis olivaris, Canadian record, Lake Erie. A live specimen of Pylodictis olivaris, the Flathead Catfish (Ictaluridae), was netted in Lake Erie and constitutes the first time the species has been captured in Canadian waters. The fish was taken by Rudy Krause, of K. W. Krause and Sons Fisheries, Leamington, Ontario, on 22 August 1978. It was caught ina commercial trapnet west of Point Pelee, 4.8 km north of the tip of the point (41°56’50”N, 82”31’40”W). The bottom of the net site was silty clay and the water temperature at the time of capture was 24.3°C. The catfish was 52 cm in total length and weighed 1.79 kg. Trautman (1957) indicated that the species has been known to reach 134.6 cm length and 37.2 kg weight. The Canadian specimen had a noticeable deformity of the middle region of the caudal verte- brae, but otherwise agreed with the morphological characteristics of the species as given by Trautman (1957) and Hubbs and Lagler (1964). The most striking feature other than head shape, which separ- ates this species from other local catfishes, is the strong backward extensions of the premaxillary tooth-patches. These were very obvious in this specimen. The species has long been known in the United States waters of Lake Erie, and Trautman (1957) indicated records north to the international boun- dary. This latest record from the northern side of Lake Erie itself represents a range extension of 30 km north of previous records and is somewhat unusual because this species is more abundant in the Ohio River and its tributaries than in the Lake Erie portion of Ohio. Trautman listed only five specimens taken in the Lake Erie waters of Ohio from 1938 to the date of writing (1956?). He suggested that “there appears to be a small population . . . in the Huron River, where they are taken yearly.” The mouth of the Huron River in Ohio is due south of Point Pelee. In describing the total range of the species, Hubbs and Lagler (1964) called it rare in Lake Erie and mentioned that it is usually a large-river form. This catfish had not been previously reported from 180 adjacent Canadian waters, in spite of the intensive commercial fishery in that area. It was the only specimen seen so that it is impossible to say now whether it is a stray or whether a small population has already been established in the Point Pelee area. The shallow waters from western Lake Erie to Lake St. Clair have contributed most of the limited number of records of freshwater fishes that have moved north into Canadian waters over the past 25 yr. Catches in those waters should be carefully scrutinized and all unusual items reported. The specimen has been added to the reference collection of the Royal Ontario Museum (ROM Cat. No. 34561). THE CANADIAN FIELD-NATURALIST Vol. 93 We thank Rudy Krause for his care in recognizing that the fish was different, and for taking the trouble to turn it over to the Ontario Ministry of Natural Resources. Literature Cited Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great Lakes Region. University of Michigan Press, Ann Arbor, Michigan. 213 pp. Trautman, M.B. 1957. The fishes of Ohio. Ohio State University Press, Columbus, Ohio. 683 pp. Received 2 October 1978 Accepted 4 December 1978 Earthworm Cocoons as a Drift Component in a Southern Ontario Stream DONALD P. SCHWERT! and KENNETH W. DANCE? 'Geology Department, Stevens Hall, North Dakota State University, Fargo, North Dakota 58105 2Ecologistics Limited, 309 Lancaster Street West, Kitchener, Ontario N2H 4V4 Schwert, D. P. and K. W. Dance. 1979. Earthworm cocoons as a drift component in a southern Ontario stream. Canadian Field-Naturalist 93(2): 180-183. Cocoons representing at least six species of Lumbricidae were isolated from drift subsamples of a stream in Waterloo County, Ontario in the wpring of 1976. Of the 308 cocoons, 92% were viable. We present arguments to support cocoon drift as a potentially important mechanism in lumbricid dispersal. Key Words: cocoons, Lumbricidae, Oligochaeta, biological drift, dispersion, Ontario, geographical distribution. In southern Ontario, as in many other regions of North America, large populations of lumbricid earthworms often occur in moist lowlands adjacent to streams and lakes. When temperature, moisture, and light conditions permit, a number of the lumbricid species in these areas are active on or just below the soil surface. Significant activity combined with surface runoff from rainstorms and snowmelt can result in large numbers of earthworms and their cocoons becoming trapped in waterways. Their subsequent downstream drift might be expected to be a significant factor in lumbricid dispersal, but this mechanism has remained poorly studied. Bouché (1972) hypothesized that stream drift had an impor- tant influence on the distribution of some hygrophilic lumbricid species in France, and Ward (1976) applied a similar hypothesis to explain the recolonization of a riffle area by the lumbricid Evseniella tetraedra in a Colorado stream. No published observations of the drift of lumbricid cocoons exist, however, probably because these might be easily confused with plant seeds during sorting. During the latter 5 mo of a 13-mo stream drift study in southern Ontario (Dance and Hynes, in press), sorting of earthworm cocoons from freshly collected drift samples was initiated. We present here data on this component and suggest its possible significance with respect to lumbricid dispersal. The Study Area All of the examined cocoons were obtained from drift samples of two headwater branches of Cana- gagigue Creek in a predominantly agricultural region immediately north of Floradale in Waterloo County, Ontario (43°39’N, 80°35’W). Stations | and 2 were located on the permanent, spring-fed east branch of ITY the creek. Stations 3 and 4 were located on the west branch, an intermittent stream which flowed continu- ously during the five months of this study. Station | was a gravelly riffle, in a treeless pasture 350 m downstream from a White Cedar ( Thuja occidentalis) woods. Station 2 was a cobble and boulder riffle at the downstream margin of an extensive American Beech (Fagus grandifolia) and Sugar Maple (Acer sac- charum) stand. Station 3 was at the downstream edge of a short, silty riffle in a treeless pasture. The streambank at Station 4 was lined with herbs and Salix spp., and the bed consisted of boulders and cobbles resting on silts. Methods Drift was subsampled continuously using a special- ly designed apparatus of 253 um Nitrex cloth net mounted over an aluminum and steel frame (Dance et al., in press). Water flowed without wave interference through a vertical opening | cm wide and 100 cm high into a long sample bag. The contents of the trap were placed weekly into a plastic bag and frozen until sorting. Under a binocular microscope, cocoons were sorted from thawed samples which had been collected between February and June 1976 and placed into labelled vials of 70% ethanol. Attempts at placing generic or specific names on the cocoons were inhibited by distortions in the cocoon shape resulting from the sample procedures and by the frequent absence of the diagnostic end “tufts” which had apparently broken off during the turbulence of drift. Names could, therefore, be placed on only a small number of the cocoons through comparison with a cocoon reference collection and with the cocoon descriptions in Evans and Guild (1947) and Gerard (1964). Mean monthly discharge volumes during the study period for each of the sample stations were calculated from data on the creek provided by the Canada Water NOTES 18] Survey and the School of Engineering at the Uni- versity of Guelph. Utilizing this information and the calculated average component of the total discharge continuously subsampled by each trap (for detailed procedure, see Dance et al., in press), a multiplication factor was derived estimating the total monthly number of cocoons passing each sample point. Results A summary of the mean monthly discharge values for the four Canagagigue Creek sample stations is presented in Table 1. Discharge volumes fluctuated widely for each station during the study period but for all four were maximal in March and minimal in June. Monthly summaries of the actual cocoon content of the drift samples and of the estimated total cocoons flowing past each station are presented in Table 2. Cocoons were absent from all February samples, but maximum numbers were recorded in March at the east branch stations and in April at the west branch stations; the west branch stations were inoperable during most of March because of flooding. Except for a sharp increase in June numbers at Station 2, the incidence of cocoons generally decreased into the early summer. TABLE |—Estimated mean 1976 monthly discharge (L/s) for four Canagagigue Creek drift sample sites (Waterloo County, Ontario) Station Month | 2 3 4 Feb. 202 455 3174 3398 Mar. 901 1126 3066 6089 Apr. AY || 410 394 446 May 151 275 220 298 June 148 24] 47 87 TABLE 2—Summary of the 1976 monthly totals of earthworm cocoons per drift trap (and estimated total cocoons flowing past each sample station) for four Canagagigue Creek drift sample sites in Waterloo County, Ontario, and their percent viability Number of cocoons (estimated total) Station Numbers of viable Cocoon cocoons (% of Month | 2 3 4 total monthly total) Feb. 0 (0) 0 (0) 0 (0) 0 (0) 0 = (=) Mar. 52 (17731) 42(22 105) Flood Flood 94 88 ( 93.6) Apr. 34 ( 5964) 33 ( 9167) 10 (1149) 52 (11 556) 129 110 ( 85.3) May 22 ( 4000) 6 ( 1875) 1( 88) 8 ( 2963) 37 36 ( 97.3) June 18 ( 3529) 29 (13 809) 0( 0) 1( 345) 48 48 (100.0) Total 308 282 ( 91.8) 182 Nearly 92% of the cocoons were found to contain sperm and albumen, or some stage in the develop- ment of the embryonic mass, and these were termed “viable,” or potentially capable of hatching. All cocoons resembled in general form those of the Lumbricidae. A qualitative examination of the undis- torted component yielded cocoons resembling the following taxa in the family: Aporrectodea spp., Dendrodrilus rubidus, Eisenia fetida, Eiseniella tet- raedra, Lumbricus terrestris, Octolasion teyrtaeum. We have collected adults of each of the above species from sites in Waterloo County, and all of the species have been recorded from southern Ontario by Reynolds (1977). One adult Eisenia fetida and one adult Eiseniella tetraedra were also collected from the drift samples. Discussion Fluctuations in the monthly totals of the observed cocoons undoubtedly reflect both the physical state of the environment and the cocoon production of the local fauna. The absence of cocoons from the February samples would be expected because of the frozen condition of the soil, but the March and April cocoon peaks are probably the result of both the high rate of surface runoff from snowmelt and rain and a high degree of surface activity and cocoon pro- duction by the earthworm fauna. The precise dis- tances which any of the cocoons drifted to the trap sites remains unknown, although presumably none of the cocoons were derived from sources greater than 2-3 km upstream, the area of the upper Canagagigue watershed. The small size (< 6mm) and tough, spheroidal outer walls of lumbricid cocoons are ideally suited for long and rigorous transport in streams. Individuals of several lumbricid species have been shown to be capable of prolonged submersion (Roots 1956; Edwards and Lofty 1972), and the successful hatching and growth while submerged of the lumbricid Allolobophora chlorotica has been demonstrated by Roots (1956). From these studies and from the remarkably high viability of the drift cocoons of the present study, successful hatching of cocoons could be expected in areas of a stream where the cocoons had been deposited near the margin or in the bottom sediments of pools. A major consequence, therefore, would be the establishment of the transported species in lowland areas along the stream. Because cocoons have subsequently been obtained from random drift samples in other streams of southern Ontario and of Newfoundland, the phe- nomenon of cocoon drift appears to be widespread. We have also collected large immature and adult Lumbricidae from other streams in Ontario, as wellas in Newfoundland, New York, and Pennsylvania. THE CANADIAN FIELD-NATURALIST Vole9s Although these worms survived submergence in rigorous environments, the battered nature of many of the specimens indicates that these large, soft-bodied stages have difficulty in physically surviving pro- longed transport. In addition, this stage is particularly susceptible to predation by fish; the palatability of cocoons to fish is unknown, but presumably low. The successful drift of cocoons may be of con- siderable importance in explaining the modern dis- tribution of many lumbricid species in North Ameri- ca. All of the species identified from Canagagigue Creek are of taxa hypothesized by Gates (1970) and Reynolds (1974) as having been introduced into North America from Europe. The widespread distribution of many of these species across Canada and the United States is attributed by Gates (1976) as being primarily due to transport by human activity. While man has undoubtedly influenced earthworm distribution, he cannot, as Ball (1975) argues, logically claim respon- sibility for the entire distribution of lumbricid earthworms across the continent. Transport of co- coons, however, from one point of intensive lumbricid establishment upstream to other points downstream could result in the subsequent colonization of the lower watershed by these species. Although further investigation is needed to support this mechanism, stream drift may be as important to the dispersal of earthworms as it is to many other groups of invertebrates. Acknowledgments We are grateful to those property owners near Floradale, Ontario, who granted us access to Cana- gagigue Creek for sampling. We also appreciate the useful suggestions of A. D. Harrison of the University of Waterloo regarding the manuscript. Funding for publication of this paper was provided by Ecologistics Limited and by North Dakota State University. Literature Cited Ball, I. R. 1975. Nature and formulation of biogeographi- cal hypotheses. Systematic Zoology 24(4): 407-430. Bouché, M. B. 1972. Lombriciens de France, écologie et systématique. Institut National de la Recherche Agrono- mique, Paris. 617 pp. Dance, K. W. and H. B. N. Hynes. /n press. A continuous study of the drift in adjacent intermittent and permanent streams. Archiv fur Hydrobiologie. Dance, K. W., H. B.N. Hynes, and N. K. Kaushik. /n press. Seasonal drift of solid organic matter in two adjacent streams. Archiv fur Hydrobiologie. Edwards, C. A. and J. R. Lofty. 1972. Biology of earth- worms. Chapman and Hall, Ltd., London. 283 pp. Evans, A.C. and W.J. McL. Guild. 1947. Cocoons of some British Lumbricidae. Annals and Magazine of Natural History, Series I1, 14: 714-719. Gates, G. E. 1970. Miscellanea megadrilogica VII. Mega- drilogica 1(2): 1-14. 1979 Gates, G. E. 1976. More on earthworm distribution in North America. Proceedings of the Biological Society of Washington 89(40): 467-476. Gerard, B. M. 1964. Synopses of the British fauna. Number 6. Lumbricidae (Annelida) with keys and descriptions. Linnaean Society of London. 58 pp. Reynolds, J. W. 1974. Are oligochaetes really hermaphro- ditic organisms? Biologist 56(2): 90-99. Reynolds, J. W. 1977. The earthworms (Lumbricidae and Sparganophilidae) of Ontario. Life Sciences Miscel- laneous Publications, Royal Ontario Museum. x + 141 PP. NOTES 183 Roots, B. I. 1956. The water relations of earthworms. II. Resistance to desiccation and immersion, and behaviour when submerged and when allowed a choice of environ- ment. Journal of Experimental Biology 33: 29-44. Ward, J. V. 1976. Lumbricid earthworm populations in a Colorado mountain stream. Southwestern Naturalist AME Wns. Received 29 September 1978 Accepted 27 December 1978 Flowering Plant Phenology at Sheep Mountain, Southwest Yukon Territory MANEFRED HOEFS Yukon Game Branch, P.O. Box 2703, Whitehorse, Yukon Territory YIA 2C6 Hoefs, M. 93(2): 183-187. 1979. Flowering plant phenology at Sheep Mountain, southwest Yukon Territory. Canadian Field-Naturalist The dates of initiation of flowering of 60 vascular plant species in the Kluane Lake area, Yukon Territory, are presented for the 1970, 1971, and 1972 seasons. The altitudinal advance of plant development is determined, using Carex filifolia as the indicator species. Key Words: phenology, flowering plants, subarctic botany, Carex filifolia, Kluane Lake, Yukon Territory. The data presented here are part of a larger investigation carried out on Dall Sheep (Ovis dalli dalli) and their range in the Kluane Lake area of the Yukon Territory since 1969. The specific study area is “Sheep Mountain,” an important winter range of a Dall Sheep population located at the southeast shore of Kluane Lake near the mouth of the Slims River (61°00’N, 138°30’E). For details on the vegetation, climate, geology and soil of the area, the reader 1s referred to Hoefs et al. (1975) and for details on the sheep population, their range use patterns, and forage selection, to Hoefs (1975). During this investigation it became clear that a number of activities of sheep, for instance forage selection, use of various plant communities, and vertical migration, were closely linked to plant phenological phenomena. A number of these were investigated. My studies (Hoefs 1974) have demonstrated that the flowers of certain plant species are preferred forage items for Dall Sheep. This paper deals with the dates of initiation of flowering of 60 vascular plants on Sheep Mountain. A number of factors have been used to explain the vertical seasonal movements of ungulates; these include snow conditions and other weather factors, avoidance of blood-sucking insects, protection of winter ranges, and advantages with respect to forage (Dixon 1938; Murie 1944: Blood 1963; Egorov 1967; Hebert 1972). Some indications of the vertical march of phenology can be obtained from the dates of first flowering of plant species at different altitudes. A more accurate quantification, however, is possible by - observation of the performance of the same species of plant at various altitudes. This study determined altitudinal advance using Carex filifolia as the indicator species. Carex filifolia was selected because it is (a) a fairly abundant plant in all dry grassland associations in the boreal, subalpine, and alpine biogeoclimatic zones of the study area (Hoefs et al. 1975); (b) one of the most preferred Dall Sheep forage plants (Hoefs 1975); (c) an “early bird” in a pheno- logical sense, flowering earlier and reaching the annual maximum of growth before any of the other important forage plants (Figure 1). Methods Sixty species of flowering plants, known to be used by sheep (Hoefs 1975), were mapped and marked by wooden stakes on Sheep Mountain in 1969. The dates 184 THE CANADIAN FIELD-NATURALIST Species April May June July 20 30 10 20 30 10 20 30 10 20 Pulsatilla patens mm | | ee Carex filifolta ee meee | | ee Townsendta hookertr | fees Rae | Arctostaphylos rubra i ie a ae eee Ee Anemone parviflora eee | eae ee ee ee Potentilla hookertana eng | ee Saxtfraga opposttifolta Rat (eee eae) Oxytropts visetda oe | Tees | S| | ea Lupinus arcticus | me ee Arctostaphylos uva-urst a aaa Linum perenne = Ce ed Erigeron composttus ae SS Lr aa) Plantago canescens aS an ae ea ee ee a Smilacina racemosa Sees | | pat le) Carex aquatilis mT Se een re Hedyearum alpen). Tas Fee | ee ae | Amevanchversaint porta) (ee Se ee | Mertensta paniculata a er eee ee OCNSEC TECTED ESS ae. |. 2 le wnemoneimelte peda BN 2 | Anemone drummondtr ee] ee eee eee eee Dryas drummondit [| | fT Saxtfraga tricusptdata aaa ae Dryas integrifolia [| oot TT Casstope tetragona ea ae eS Taraxacum officinale =r SS pilopium latifoltum ———==rS Astragalus umbellatus a aE ee See Lappula myosotts an ee ee eee ee Potentilla fruttcosa Pe ee Myosotts alpestris eee cee Aster stbtrtcus |. eel Pentstemon gormant an ae ee ee ee eee Erigeron purpuratus [ee ee ee Aster alptnus i a Oxytropts huddeTsonit i as ee ee Sedum rosea PI ao Rosa actcularts LO a ee ee tlene acaults esac Sipe ae. a as Artemtsta hyperborea ae Se haar area Arntea alptna Vieburnum edule Aehtllea borealts Artemista alaskana Itnnaea borealts PeSaeS h O| nz oe Cie) pi lobiummuanguetd folie 0 | ee Calamagrostts purpurascens Poe eee Vaeetntum vttts-tdaea Aster yukonense Ledum palustre Poa glauca gadenus elegans oltdago multrtradiata Crepte nana | || dl lo DEC CNIS UCI feta GC CLC a | oe Ol | i | Erigeron caespitosus ecese| Ge ss | |e 5| ele | altum boreale (ce a ae a a ay ESTs | Hordeum jubatum Pe ee ie ae ein a a Artemisia rupestris aie a eS ee Lz 2 Agropyron yukonense FIGURE |. Dates of first flowering of Sheep Mountain plants for 1970, 1971, and 1972. Vol. 93 179 at which these plants first developed fully opened flowers were recorded during 1970, 1971, and 1972 seasons. Ten clumps of Carex filifolia were selected at each of five altitudinal sites (935, 1000, 1330, 1590, and 1910 m) and measurements of growth, flowering, fruiting, and initiation of dormancy were carried out at weekly intervals from early May to late October 1970. The selected clumps were protected from grazing by exclosure fences. Nomenclature of plant species follows Hultén (1968) with the exception of Artemisia hyperborea, which is described in Porsild (1966). Results and Discussion Dates of First Flowering Figure | lists 60 flowering plants that grow on the range of the Dall Sheep population under study, and gives their first flowering dates arranged approxi- mately in chronological sequence. Because observa- tions were carried out over three seasons, with considerable differences in weather patterns, the first flowering dates are ranges of 10d to 2 wk and not individual days. Great variations were observed particularly between the spring of 1971 compared to 1970. Some flowers came into bloom as much as 12 d earlier in 1971. The data of Figure | can therefore be interpreted as follows: beginning of range is initial flowering date in 1971, end of range is initial flowering date for 1970. The date for 1972 is about half-way between those two extremes. The data presented here, particularly for June when many plants came into bloom, must be considered as approximations since it was not physically possible to check every marked species every day. It is unlikely, however, that any of the dates are in error by more than 4d. Figure | includes a number of species which grow in various altitudes and in various plant associations (aspects) at the same altitude. Even though only specimens growing in the most “advanced” sites in a phenological sense, which were usually south-facing slopes in the boreal zone, were marked and are considered in Figure I, records were kept on the delays observed if such species also grew on north- facing slopes and at higher elevations. Detailed information on the floristic composition of the various plant associations on Sheep Mountain, and their distribution with respect to aspect and altitude are given in Hoefs et al. (1975) and need not be repeated here. In general it was found that there was a 10- to 14-d delay in plant development between north- and south-facing slopes at the same altitude, and a delay of about | d for each 35 m of altitude at the same aspect. An example may demonstrate this point. Pulsatilla patens is the first plant, with the exception of willows, to have open flowers. On sunny, south- NOTES 185 facing protected slopes with grassland vegetation in the boreal zone (1000 m), its flowers may in favorable springs (1971) be seen as early as 22 April. This species grows also in alpine elevation (1667 m), where it did not come into bloom before 17 May (1971). Figure | reveals that Pulsatilla patens is the only species that may come into bloom in April, about 26 species begin to bloom during May in an average year, 19 species in June, and 15 species in July. Although there are many exceptions to this rule it is in general true that members of the families Ranunculaceae and Legu- minosae are among the first to come into bloom while members of the Compositae and Gramineae families are among the last. Altitudinal Advance of Plant Development Figure 2 shows the growth curves of Carex filifolia at five sites with increasing altitudes. Lines connecting the points at which 50% and 100% of the annual growth was achieved give an indication of the vertical phenological march. For reasons as yet unexplained, the plants at an elevation of 1000 m were pheno- logically more advanced than those at an altitude of 935 m. One reason may have been that the plants at the lower altitude were more often subject to shade cast by surrounding trees during the course of a day. Because the lines connecting 50% and 100% growth points are parallel it will suffice to discuss one of them. At an altitude of 1000 m Carex filifolia reached its maximum annual growth around 18 June, at 1330 m around 26 June, at 1590 m around 5 July, and near the peak of Sheep Mountain, at 1910 m, it took till 15 July, 1970. The speed of vertical advance of plant development was therefore 910 m per month or about 315 m per 10 d. It is not possible to quantify within similar terms the sequence of dry-up, since this is influenced not only by temperature or altitude but also by moisture deficiency. This speed of vertical advance agrees with Hopkins’ (1920) bioclimatic law, which states that . there is a delay in vegetation development of about 3 to 4d for each 100- to 130-m increase in altitude, as well as with Stoddart and Smith’s (1955) remark that “Most observers agree that the (altitudinal) difference approaches one day for each 100 feet of altitude.” My studies documented a very close relationship between vertical migrations of Dall Sheep and plant phenology, particularly in spring. During May and June sheep leave their winter ranges and move up high, making use of plant associations with new growth and avoiding others that are still dormant. This pattern of range use continues on summer range, where advanced plant associations on south- and west-facing slopes were primarily used in June and early July while plant associations on east and northern aspects were used in late July and August. Selection with respect to phenological stage was also 186 THE CANADIAN FIELD-NATURALIST 100 50 100 50 100 50 100 50 % "green'! matter in total biomass 100 50 50 % of annual _—[— Vol. 93 growth completed Maximum of annual growth completed FIGURE 2. Growth of Carex filifolia at various altitudes in 1970, demonstrating vertical phenological advance. observed when feeding took place within one plant association. New growth was preferred to the mature and dormant stage of a species. Special attention was paid to flowers, particularly large and showy ones. Pulsatilla patens and Oxytropis viscida flowers are important forage items in early spring when the sheep are still on winter range, while Pedicularis spp., Dryas integrifolia, Hedysarum alpinum, Epilobium lati- folium, and Oxytropis huddelsonii flowers are important on alpine summer range. Fall migration from alpine summer ranges commences in late August and early September. Only ewes and lambs, however, appear to arrive at lower elevations in time to utilize certain plant species that are still “green.” Heavy use is made at this time of the various willow species in the subalpine shrub zone (1200 to 1600 m), when all the 1979 vegetation of the alpine zone is already dormant. Forage selection and range use patterns are dealt with in detail in Hoefs (1975). Literature Cited Blood, D. A. 1963. Some aspects of behaviour of a bighorn herd. Canadian Field-Naturalist 77(2): 77-94. Egorov, O. V. 1967. Wild ungulates of Yakutia. Israel program for scientific translations, Jerusalem. United States Department of Commerce. Dixon, J. S. 1938. Birdsand mammals of Mount McKinley National Park. United States Department of Interior, National Park Service, Fauna Series Number 3. Hebert, D. 1972. Differences between years and nutrient cycles. Alberta Fish and Wildlife Division, Northern Wild Sheep Council, Symposium Proceedings 1972: 15— DD, Hoefs, M. 1974. Food selection of Dall sheep. Jn The behaviour of ungulates and its relation to management. Volume 2. Edited by V. Geist and F. Walter. Inter- national Union for Conservation of Nature and Natural Resources. pp. 759-786. NOTES 187 Hoefs, M. 1975. Ecological investigation of Dall sheep and their habitat. Ph.D. thesis, University of British Columbia, Vancouver. ; Hoefs, M., I. McT. Cowan, and V.J. Krajina. 1975. Phytosociological analysis and synthesis of Sheep Moun- tain, southwest Yukon Territory, Canada. Syesis 8 (Supplement 1): 125-228. Hopkins, A.D. 1920. The bioclimatic law. Journal of Washington Academy of Science 10: 34-40. Hultén, E. 1968. Flora of Alaska and neighboring ter- ritories. Stanford University Press, Stanford, California. Murie, A. 1944. The wolves of Mount McKinley. Fauna of the National Parks of the United States, Fauna Series Number 5. United States Department of Interior. Porsild, A. E. 1966. Contributions to the flora of south- western Yukon Territory. National Museum of Canada, Contributions to Botany IV, Bulletin 216. Stoddart, L. A. and A. D. Smith. 1955. Range manage- ment. McGraw-Hill Book Company, New York. 433 pp. Received 8 September 1978 Accepted 5 December 1978 Eggshell Thickness in American Shorebirds before and since DDT MICHAEL L. MORRISON and LLOYD F. KIFF Western Foundation of Vertebrate Zoology, 1100 Glendon Avenue, Los Angeles, California 90024 Morrison, Michael L. and Lloyd F. Kiff. 1979. Eggshell thickness in American shorebirds before and since DDT. Canadian Field-Naturalist 93(2): 187-190. Thickness indices of eggshells collected before and since use of DDT began were obtained for 31 species and subspecies of American shorebirds. Little change in thickness was found between early and recent samples, the maximum being -4.7% in “American” Oystercatcher (Haematopus ostralegus palliatus) eggshells from Texas and the Atlantic coast of the United States. Key Words: American shorebirds, eggshell thickness, DDT, bioaccumulation, pesticides, chemical pollutants. Significant eggshell thinning has been documented in many species of wild birds 1n recent years, nearly all of it attributable to the effects of p,p’-DDE, a breakdown metabolite of DDT (Cooke 1973: Stickel 1975: Peakall 1975). Thinning has been most severe in the eggshells of bird- and fish-eating species (Ratcliffe 1970: Anderson and Hickey 1972), but there have been few investigations of eggshell thickness in middle trophic-level birds, including shorebirds of the sub- order Charadrii. Minor shell thickness changes were found for Pluvialis apricaria and Tringa nebularia by Ratcliffe (1970), and for Philohela minor by Dilworth et al. (1972) and Kreitzer (1972). Studies of organochlorine residues in shorebirds or their eggs are more numerous. Keith and Gruchy (1972) summarized data from several sources, and additional figures for body and egg residue levels have been reported for various species by Enderson and Berger (1968), Flickinger and King (1972), White et al. (1973), Peakall (1976), and Walker (1977). Owing to its importance as a game species, several studies are available on residue levels in American Woodcocks ( Philohela minor) (Pearce 1971: McLane et al. 1971, 1973; Dilworth et al. 1972, 1974; Clark and McLane 1974). Most of these studies indicated that organochlorine residues, including DDE, rarely exceeded 4 ug/g wet weight; however, significantly higher levels of DDE were found in the bodies of Numenius americanus (14.0 ug/g wet weight) in Alberta (Peakall 1976), Philohela minor (13.0 ug/g wet weight) in a heavily sprayed area in New Brunswick (Pearce 1971), Pluvialis dominica (62.0 ug/g lipid), and Numenius phaeopus (39.0 ug/g lipid) in Alaska (Walker 1977), Vol. 93 LD-NATURALIST THE CANADIAN FY: 188 1891-1 “1000 > d xxx “10'0 >dxx S00 >dx ‘sisayjuaied ul UdAId pojUdsaidds sayoinjo Jo Jaquinn,, 8I- Kr0o00+9S0 POOOFLSO (8)67 (0@)6L OL6I~ OS6l SPr6l ~ I1€61 egouury ‘sndojupuny piupjppdospy + 7000+ 850 7000+ 850 (€7)88 (SIIZ S961-1S6I ZLr6l - II161 BSR] “DUIdjD °) 07- 6000F0S0 €000F1S0 (5)91 (81)L9 £961 ~0961 Sr6I ~ ZZ6I BYSRIY ‘Ipuing “) CT «=—£000F PFO SOOOFSPO (ZIEh (ZI)hh €S61 ~ 8h6l Srol — ZE6l BQOMURIAL “D//IINUILU *>) «1 C- €000F 9F0 £000+F LPO (OZZL (O@)8L $961 ~- IS6l be6l — PZ6l BYSPLY “UNL +) + 7000+ 9¢'0 cOO0+9b0 (OZLL (97/66 1961 - I1S6l Lb6l — rI6l BYSeLY “MypIsnd slapd) 61- rooo+ ESO CO0O0F FSO (OZ)S8ZL (OZ)SL £961 IS6l 9b6Il ~ 7761 BYSEIY “SMLIDIIN “df *L€ 7000+ ¢S0 PrO00+F PSO (OZ)8ZL (S1)OL €961~ IS6l Sb6l ~ I€61 BYSELY “SMIDGO] ‘d 8c 0100+ 020 900°0 + CL'0 (8)0€ (0@)6L CL6I1~ IS6l 9bP6l ~— E£6I BIUJOJIR) “0/001 sndospjpoyd TI- 9000+ 080 SOO0+ 180 (91)¢9 (81/69 €961~ I1S6l Sb6l ~ PZ6l BYSeLY “D/DYdawOUD[ALU *y xL7+ 80004520 SOO0+FELO (91)h9 (LI)89 $961 ~ IS6l Lh6l ~ 8761 BYSPLY “SatdsaIUul DIADUAA ~ $0004 690 LOO0+F 790 (LIDL9 (81)89 9961 OS6l Le6l ~ O16] BIULOJIPR) DMD/NIDU SIMI 81 6000 + 011 6000+ ZI'l (S)6l (OWLL 8961 ~ TZS6l IPél — Z88I sexo] ‘smipuyndiiuas snsoydosidoin J 91+ TlI0O0FS90 PO0O0F P90 (Pr)9I (CI8rh L961 ~ £961 9b61 — 9761 BLOG “DLIDIIOS DBULATL vI- rlodoO+erl 1100+ SrI (El)0S (IZ)LL 8661 - 8h6l Lr6l — vI6l URI] SMUDILIALUD “A x8 © + 1100+ 601 900°0 + S01 (L)LT (0Z)08 €S61~ IS6l Sr6l — £61 egowuery ‘sndoapyd SMIUAUIAN 80 LOOOF ETI TlOOF HT CUdDEr (LI)S9 6961 OS6I Lr6l ~ 6261 BLOG|Y “popes DsOwUur] yI+ 8000+ £20 S000 + 7L0 (L)IT (02)09 1961 ~ 8r6l IPr6l ~ 9761 BIUIOFITR ) SHUT PUDXolD wD x67 LOOOF6L0 LOOO+F ISO (HIDLP (0Z)08 LL61~ 8r6l IP6l — O161 PIUIO PPR) “SM4Af1ION * vyo+ 800045680 L000 + £80 (6)L7 (8)pc 8961 1961 St6l ~ 9061 BpllO|{ “DIUOS}IM °) *C1~ L0O00+ £80 L000 + 680 (L)0Z (91)9h S961 ~ ZS6I IP6l ~ 0061 SBX9] “DIUOS[IM “) *91— 8004190 9000 + 7290 (8)9Z (CI)8S €S6Il -8r6l SPr6l —- I€6l eqoMUPY, ‘SMIDLUyDdiLUas SNLApoADYD x07+ LOOO0+ 001 7000+ 860 (57)96 (ZZ)L8 S961 8r6l Lr6l — HZ6l BYySeIYy “vjospionbs -q 900'0 + €8'0 9000+ 780 (SI) 85 (L1)89 8961 6r6l SP6l ~ 0061 BYSBLY “DIIUILUOP SIPDINA]d FE + PIOO- SII 80004111 (Q)ZE (OWLL TL61-—8r6l +br6l - 8I6l BIUIOJIPD ‘DUDILALUD DAISOMAANIIY 61 Sl00+ P01 8000+ 901 (9)p7 (Z1)89 6S6l OL6I — 8681 yriy) “smuDoixal sndojupwuipy wxxE€- LIOOFLYI PIOOFCSI (ElPE (0Z)ZS 9961 -6r6l LOI - Z68I ISBOD Doe "Sf “MUDWUY.IDg ‘0 “H xaxl¥- O1O0F ZHI PIOOF6rI (OZ)PS (OZ)IS 6961 -SS61 961 - 881 IsvO OUR “S*f] FY sexo ‘smpyjvd snsajp.uso sndoiwappy €l+ poo0o0+LL0 90004920 (77)88 (€1)9S $961 ~ 8h6l £761 — 968I OXI] “F ‘vsouids punlvr osuey) u d u a (aU) (q) pajdwies uoisay ‘saisads % aS F Xopul uray 8339 JO “ON poriod polad JUDY Alley SBIIV URILIIWY YWON SNOLIVA WOIJ SS9UYIIYI [[IYSdsa PIlgoIOYS — ] AIAV]E 1979 and in the eggs of Philohela minor (14.9 ug/g wet weight) in New Brunswick (Dilworth et al. 1972). Ina DDT-sprayed area around Churchill, Manitoba, the invertebrate prey of shorebirds contained DDE residues of 0.3 and 0.4 ug/g wet weight, and shore- birds there accummulated DDE residues of up to an average of 39.4 ug/g wet weight (Brown and Brown 1970). The DDE residue levels reported in these latter studies were at least as high as those associated with serious eggshell thinning in certain raptors and fish- eating birds (Blus et al. 1974; Peakall 1976; Kiff et al., in press), yet there have been no reports of such changes in the eggshells of these or other shorebirds. This study was undertaken to determine whether significant changes in eggshell thickness of American shorebirds have occurred since the introduction of DDT in the mid-1940s. Methods Empty dry eggshells of charadriine shorebirds in the collection of the Western Foundation of Verte- brate Zoology were weighed to the nearest 0.001 g on a Mettler P120 balance, and their length and breadth were measured to the nearest 0.01 mm with Helios dial calipers. A shell thickness index (shell weight X 100/length X breadth) was calculated for each eggshell; such an index is correlated with actual eggshell thickness (Anderson and Hickey 1972). Eggs that were broken, that had blowholes greater than 3mm in diameter, or that were collected in an advanced stage of incubation, were excluded from the analyses. The nomenclature and species sequence used here follows Morony et al. (1975). Results The mean thickness indices for pre-1947 (before- DDT) and post-1947 (since-DDT) North American shorebird eggshells are shown in Table |. Seven species had slightly thicker eggshells in the recent samples, four showed no change, and 16 species had thinner eggshells in the post-1947 samples. Eggs of NOTES 189 another species, Charadrius wilsonia, were thicker in Florida but thinner in Texas than pre-1947 indices. The maximum amount of difference between the pre- and post-1947 samples were -4.7% in the “American” Oystercatcher (Haematopus ostralegus palliatus) in Texas and along the southern Atlantic coast of the United States. Recent eggshells of the “black” Oystercatcher (H. o. bachmani) in California and Oregon were 3.3% thinner than the mean thickness of the pre-1947 sample. Many shorebirds included in this study migrate and winter in South American countries where DDT is still intensively used. For comparative purposes, we also measured eggshells of three resident species of Chilean shorebirds of three different families (Table 2). As with the North American species, only minor changes were noted. The only statistically significant difference, -4.1%, was found in a plover, Vanellus chilensis. Discussion Eggshell thinning exceeding 20% has generally resulted in reproductive failure and population declines in the species involved (Keith and Gruchy 1972: Stickel 1975), but the biological significance of thinning less than 10% is not well understood (Faber and Hickey 1973). In this study, the maximum increase (+3.8%) was similar to the maximum decrease (4.7%) in thickness index. Although statistically significant, these minor index changes are probably due to sampling artifacts (e.g., observer error, insufficient sample size, geographical variation), rather than pesticide effects. We know of no biological phenomena which could readily explain an increasing shell thickness within the span of time these eggs were collected. Disruptions in eggshell ultrastructure and chemical composition that reduced egg hatchability in a population of Common Terns (Sterna hirundo) were - attributed to DDE contamination even in the absence of marked shell thinning (Fox 1976). Thus, the lack of substantial thinning of American shorebird eggshells TABLE 2 — Eggshell thickness of Chilean shorebirds Early Recent period period No. of eggs® Mean index + SE % Species (E) (R) E R E R Change Nycticryphes semicollaris 1934 — 1938 28(15) 0.93 + 0.011 1962 — 1969 19(11) OLS ae OOS = 3 Vanellus chilensis 1934 — 194] 24(8) 1.21 0.017 1960 — 1968 33(10) 1.16+0.011 4.1* Gallinago paraguaiae 1933 — 1940 40(20) 0.87 + 0.008 1957 — 1969 22(12) 0.88 £ 0.013 + 1.1 “Number of clutches given in parentheses. *P< 0.05. 190 in recent years does not prove that these species are free of pesticide-induced reproductive problems. It is possible that shorebirds have a lower sensitivity to DDE-induced eggshell thinning than many higher trophic-level species, in addition to usually possessing lower residue burdens. Peakall (1975) categorized the charadriiforms as being “moderately sensitive” to DDE, based on data on Herring Gulls (Larus argentatus) presented in Hickey and Anderson (1968). Although DDE residues in these species have evidently not reached a level at which they cause eggshell thinning, migrant shorebirds may still repre- sent the most important source of DDE contamina- tion for Arctic raptors, including Peregrine Falcons (Falco peregrinus) and Gyrfalcons (F. rusticolus). Several studies have indicated that migratory shore- birds possess the highest organochlorine residues of any prey item taken by these falcons (Cade et al. 1968: Enderson and Berger 1968: White et al. 1973: Walker 1977). Acknowledgments We appreciate the support of Ed N. Harrison and the Western Foundation of Vertebrate Zoology. Critical review by A. J. Erskine, J. J. Hickey, and D. B. Peakall enhanced the quality of the paper. Julie Kiff and Dana Gardner assisted in preparation of the manuscript. Literature Cited Anderson, D. W. and J. J. Hickey. 1972. Eggshell changes in certain North American birds. Proceedings of the XV International Ornithological Congress. pp. 514-540. Blus, L.J., B.S. Neely, Jr.. A. A. Belisle, and R.M. Prouty. 1974. Organochlorine residues in Brown Pelican eggs: relation to reproductive success. Environmental Pollution 7: 81-91. Brown, J. J. and A. W. A. Brown. 1970. Biological fate of DDT in a sub-arctic environment. Journal of Wildlife Management 34: 929-940. Cade, T. J., C. M. White, and J. R. Haugh. 1968. Pere- grines and pesticides in Alaska. Condor 70: 170-178. Clark, D.R., Jr. and M. A.R. McLane. 1974. Chlori- nated hydrocarbon and mercury residues in woodcock in the United States, 1970-1971. Pesticides Monitoring Journal 8: 15-22. Cooke, A. S. 1973. Shell thinning in avian eggs by environ- mental pollutants. Environmental Pollution 4: 85-52. Dilworth, T.G., J. A. Keith, P. A. Pearce, and L.M. Reynolds. 1972. DDE and eggshell thickness in New Brunswick woodcock. Journal of Wildlife Management 36: 1186-1193. Dilworth, T.G., P. A. Pearce, and J. V. Dobell. 1974. DDT in New Brunswick woodcocks. Journal of Wildlife Management 38: 331-337. Enderson, J.H. and D.D. Berger. 1968. Chlorinated hydrocarbon residues in peregrines and their prey species from northern Canada. Condor. 70: 149-153. THE CANADIAN FIELD-NATURALIST Vol. 93 Faber, R.A. and J.J. Hickey. 1973. Eggshell thinning, chlorinated hydrocarbons, and mercury in inland aquatic bird eggs, 1969 and 1970. Pesticides Monitoring Journal 7: 27-36. Flickinger, E.L. and K. A. King. 1972. Some effects of aldrin-treated rice on Gulf Coast wildlife. Journal of Wildlife Management 36: 706-727. Fox, G. A. 1976. Eggshell quality: Its ecological and physiological significance in a DDE-contaminated Com- mon Tern population. Wilson Bulletin 88: 459-477. Hickey, J.J. and D.W. Anderson. 1968. Chlorinated hydrocarbons and eggshell changes in raptorial and fish- eating birds. Science 162: 271-273. Keith, J. A. and I. M. Gruchy. 1972. Residue levels of chemical pollutants in North American birdlife. Pro- ceedings of the XV International Ornithological Congress. pp. 437-454. Kiff, L. F., D. B. Peakall, and S. R. Wilbur. Recent changes in California Condor eggshells. Condor. /n press. Kreitzer, J. F. 1972. Thickness of the American Woodcock eggshell, 1971. Bulletin of Environmental Contamination and Toxicology 9: 281-286. McLane, M.A.R., L.F. Stickel, and J.D. Newsom. 1971. Organochlorine pesticide residues in woodcock, soils, and earthworms in Louisiana, 1965. Pesticides Monitoring Journal 5: 248-250. McLane, M. A.R., L. F. Stickel, E. R. Clark, and D.L. Hughes. 1973. Organochlorine residues in woodcock wings, Il states — 1970-71. Pesticides Monitoring Journal 7: 100-103. Morony, J.J., Jr., W.J. Bock, and J. Farrand, Jr. 1975. Reference list of birds of the world. American Museum of Natural History, New York. 207 pp. Peakall, D. B. 1975. Physiological effects of chlorinated hydrocarbons on avian species. /m Environmental dynamics of pesticides. Edited by R. Haque and V. H. Freed. Plenum Publishing Company, New York. pp. 343-360. Peakall, D.B. 1976. The peregrine falcon (Falco pere- grinus) and pesticides. Canadian Field-Naturalist 90: 301-307. Pearce, P. A. 1971. Side effects of forest spraying in New Brunswick. Transactions of the Thirty-sixth North Ameri- can Wildlife and Natural Resources Conference. pp. 163-170. Ratcliffe, D. A. 1970. Changes attributable to pesticides in egg breakage frequency and eggshell thickness in some British birds. Journal of Applied Ecology 7: 67-115. Stickel, W. H. 1975. Some effects of pollutants in terrestial ecosystems. /n Ecological toxicology research. Edited by A.D. McIntyre and C.F. Mills. Plenum Publishing Company, New York. pp. 25-74. Walker, Wayman. 1977. Chlorinated hydrocarbon pol- lutants in Alaskan Gryfalcons and their prey. Auk 94: 442 447. White, C.M., W.B. Emison, and F.S.L. Williamson. 1973. DDE in a resident Aleutian Island peregrine population. Condor 75: 306-311. Received 2 October 1978 Accepted 2 January 1979 1979 NOTES 19] Response of Wintering Moose to Mechanical Habitat Rehabilitation in Alaska M. SIGMAN Alaska Cooperative Wildlife Research Unit, University of Alaska, Fairbanks, Alaska 99701 Present address: Ester, Alaska 99725 Sigman, M. 1979. Response.of wintering Moose to mechanical habitat rehabilitation in Alaska. Canadian Field-Naturalist 93(2): 191-193. Moose (Alces alces gigas) formed winter concentrations following mechanical crushing of vegetation in a 28-yr-old burn. Compared to a control area, the density of Moose and calf survival in the rehabilitated area were consistently higher in January—March 1975. Persistent large aggregations, although dynamic in nature, were related to the distribution of crushed mature hardwood stands because Moose fed on aspen bark. Observed differences between the two areas resulted from the provision of a concentrated, available, and nutritious food supply in combination with less severe snow conditions. Key Words: Moose, Alces alces gigas, habitat, rehabilitation, aggregation, behavior. In 1975, as part of efforts to manage habitat to stabilize a declining Moose population, Kenai Nation- al Moose Range (KNMR) staff used LeTourneau tree crushers to rehabilitate a portion of an area burned in 1947. An important result of the winter crushing of mature and regrowth vegetation was the immediate presence of large concentrations of Moose in the crushed area. My objectives in observing these concentrations were to compare the use of the area to that of a control area and to observe behavioral responses of Moose to the habitat disturbance. Study Area The Willow Lakes Rehabilitation Area (WLRA) is located in the KNMR in the northwestern portion of the Kenai Peninsula, Alaska, and is part of a large plain composed of flats, low ridges, hillocks, muskeg, and numerous small lakes. The particular area selected to be crushed could be easily reached froma road and the vegetation was considered representative of the 1947 burn. A 460-ha (1137-acre) doughnut- shaped area was crushed while the “doughnut hole” was scheduled for a later controlled burn. Two areas were surveyed by use of fixed-wing aircraft: a 2330-ha (5750-acre) area which included crushed and uncrushed portions and a control area of similar size approximately 6 km away in the 1947 burn. Both areas included many lakes; the actual area used by Moose for feeding and bedding was roughly equivalent. J. L. Oldemeyer, Denver Wildlife Research Center (unpublished data) has provided detailed information on the vegetation in the project area before and after crushing. In general, before crushing, islands of remnant mature White Spruce (Picea glauca)/ Trembling Aspen (Populus tremuloides)/ Paper Birch (Betula papyrifera) stands were interspersed with regrowth birch-spruce (P. glauca and P. mariana) stands. Methods Semi-monthly surveys of the WLRA and control area were flown from 2 December 1974 until 2 May 1975. I used information from these surveys to select suitable areas for viewing large concentrations of Moose. I conducted long (4-8 h) observations of large aggregations in the crushed area and noted all Moose in the general area on my way to and from specific off- road observation sites of aggregations. In addition, I conducted a daily road survey along the southern boundary of the project area. Owing to lack of access- ibility, however, I made no attempt to observe Moose in the control area. Ground observations continued, weather permitting, from 14 February until 6 April ISHS. During long observation periods, I observed aggre- gations from natural blinds (e.g., a clump of birch trees on a nearby hill) approximately 200 m from the Moose. After allowing a period for the animals to resume undisturbed behavior after my arrival at the blind, I recorded data at 15-min intervals, noting the number of Moose visible from my vantage point, the number of Moose in aggregations, the general behavior of individual Moose (e.g., feeding vs. lying), and the presence of collared Moose (collared previ- ously by the Alaska Department of Fish and Game). I also sketched the relative locations of individuals. Between these observations, I scanned the area frequently for movements, arrivals and departures in aggregations, and interactions. Results The WLRA received much more use than the control area. Aerial observations documented 1374 d of Moose use in the WLRA compared to 466 d in the control area. Observed densities ranged from 2.5 to 9.8 Moose per km? in the WLRA compared to 1.3 to 3.1 Moose per km? in the control area (Table |). The higher densities in the crushed area built up between 192 TABLE 1—Comparison of Moose densities observed in the Willow Lakes Rehabilitation Area (WLRA) and the control area Moose/ km2 Date of survey WLRA Control 23 Dec./74 25) 24 Dec./74 3 10 Jan./75 6.3 ral 20 Jan./75 7.0 De SEkeby 5 6.3 2.0 20 Feb./75 9.8 2.6 6 Mar./75 VD 1.6 14 Mar./75 6.2 DM 1 Apr./75 7.8 2.4 18 Apr./75 4.2 3 early and mid-winter, then decreased in late winter, while the densities in the control area remained low throughout the winter. Calf survival was much higher in the WLRA (Table 2). Cow-calf ratios ranged from 26 to 62 calves per 100 adults, compared with those in the control area, which decreased steadily from 36 calves per 100 adults to none by 18 April. Ground observations in the WLRA documented an additional 324 d of use by Moose. Although group- ings of one, two, or three Moose were commonly observed travelling or feeding in any portion of the area, all larger groups were observed in crushed mature hardwood stands. The animals initially con- sumed branches and twigs of birch and aspen trees. Concentrations persisted in these area for several days apparently to consume aspen bark from the downed mature trees. Based on a vegetation type map of the TABLE 2—Comparison of calves per 100 adults observed in the Willow Lakes Rehabilitation Area (WLRA) and the control area Calves/ 100 adults Date of survey WLRA Control 23 Dec./74 39 24 Dec./74 36 10 Jan./75 2 23 20 Jan./75 31 28 5 Feb./75 34 13 20 Feb./75 28 13 6 Mar./75 ; 37/ 24 14 Mar./75 27 II 1 Apr./75 31 4 18 Apr./75 62 0 THE CANADIAN FIELD-NATURALIST Vol. 93 area, maximum densities within these crushed stands varied from 0.25 to 30 Moose per km3 (n = 34). Concentrations persisted but the aggregations themselves were extremely dynamic during the long observation periods. Repeated observations of collar- ed Moose indicate that the individual length of stay was variable, ranging from | to 112 d, which suggests that considerable turnover occurred throughout the winter. With the exception of cow-calf pairs, Moose rarely left or entered the stands in groups. Individuals rarely interacted even at high densities. Only three brief interactions occurred during 48.1 h of observation involving 405 Moose. Discussion The comparison of Moose densities within the WLRA and those in the control area demonstrates the rapid build-up of Moose following crushing and a prolonged use of the crushed area by many Moose. The observed mid-winter aggregations, although not notably larger than those observed in late-winter situations (Timofeeva 1967; Peek et al. 1974; Rose- neau and Stern 1974), were extremely dense within the crushed mature hardwood stands. The lowland winter range of the 1947 burn is generally considered poor winter range (Oldemeyer et al. 1978). Thus, it is likely that the major factor responsible for the aggregations and concentrations of Moose in the WLRA was a food supply that was highly nutritious (J. L. Oldemeyer, personal com- munication), concentrated, and available. Snow conditions were also variable between the two areas. Movement of the large crushers to various sites compacted the snow into trails and the removal of vegetation resulted in wind compaction of newly- fallen snow. Sigman (1977) noted that the increasing snow depths in typical 1947 burn habitats, in the nearby Moose Research Center, appeared related to the timing of calf deaths. In the WLRA, the reduction of energy expenditure necessary to move through deep snow and to locate food was combined with an increased energy intake from the food supply. This combination may have acted to reverse some of the stressful effects of typical winter conditions and seemed to have benefitted calves in particular. The immediate and prolonged use of the rehabili- tated area demonstrates flexible aspects of Moose behavior patterns. Use of the disturbed area confirms the ability of individual Moose opportunistically to locate a small area where winter conditions are more favorable for survival and the ability of wintering Moose to tolerate high densities. Acknowledgments Funding for this study was provided through Pittman-Robertson funds, administered by the 1979 Alaska Department of Fish and Game (ADFG) and the Alaska Cooperative Wildlife Research Unit. Logistical support was provided by the staff of the Kenai National Moose Range and the Moose Re- search Center, Kenai, Alaska. In particular, Robert Ritchey, KNMR, provided aerial sightings and Paul Arneson, ADFG, provided some ground sightings. John Oldemeyer, KNMR, and Lyman Nichols, ADFG, reviewed the manuscript and provided help- ful suggestions. Literature Cited Oldemeyer, J. L., A. W. Franzmann, A. L. Brundage, P. D. Arneson, and A. Flynn. 1978. Browse quality and the Kenai Moose population. Journal of Wildlife Manage- ment 41(3): 533-542. NOTES 193 Peek, J. M., R. E. LeResche, and D. R. Stevens. 1974. Dy- namics of Moose aggregations in Alaska, Minnesota, and Montana. Journal of Mammalogy 55(1): 126-137. Roseneau, D. G. and P.M. Stern. 1974. Distribution of Moose, Muskox, and Sheep in northeastern Alaska, 1972. In Distribution of moose, sheep, muskox, and fur- bearing mammals in Northeastern Alaska. Edited by R. D. Jakimchuk. Canadian Arctic Gas Study Limited. Biological Report Series, Volume VI, Chapter I. Sigman, M. J. 1977. The importance of the cow-calf bond to overwinter moose calf survival. M.Sc. thesis, University of Alaska, Fairbanks, Alaska. 185 pp. Timofeeva, E. K. 1967. Onthe behavior of moose, based on observations made in the northeastern Leningrad oblast. Vestnik Leningradskogo Universiteta, Number 15. (Translation). 11 pp. Received 2 October 1978 Accepted 3 January 1979 New Localities for the Northern Spring Salamander and the Four-toed Salamander in Southwestern Quebec DAVID M. GORDON Box 180, Macdonald College Post Office, Quebec H9X 1C0 Gordon, David M. 1979. New localities for the Northern Spring Salamander and the Four-toed Salamander in south- western Quebec. Canadian Field-Naturalist 93(2): 193-195. New localities for the salamanders Gyrinophilus porphyriticus porphyriticus and Hemidactylium scutatum are documented and the habitat at the collecting sites described. This is the first record for Gyrinophilus west of the Richelieu River and only the second record outside the Appalachian Mountain region. Hemidactylium is reported for the second time south of the St. Lawrence River in Quebec. Key Words: salamanders, Gyrinophilus, Hemidactylium, geographic distribution, Quebec. The geographic distribution of salamanders in Quebec is poorly known (see Figure 1). The secretive nature and seasonally variable habits of these animals and, until recently, the lack of systematic field work have contributed to this situation. The recent efforts of the author in conjunction with the National Museum of Natural Sciences, and of Weller (1977) have expanded our knowledge of the nature of the distributions of. two of the species of salamanders in Quebec. This note documents two new localities for the Northern Spring Salamander and one new locality for the Four-toed Salamander. Gyrinophilus porphyriticus porphyriticus During the course of field work (15 August 1973; 29 April and 19 May 1975) two larval and 10 meta- morphosed Northern Spring Salamanders (Gyrino- Philus p. porphvriticus) were discovered in the headwaters of Riviere aux Outardes-Est, near Frank-. lin Centre, Huntingdon County (45°01’N, 73°54’W),. Riviére aux Outardes-Est originates at a swamp and small lake at the top of Covey Hill at an elevation of 210-225 m. At the collecting site the stream cuts through a stand of Eastern Hemlock (7suga cana- densis) and mixed hardwoods of Sugar Maple (Acer saccharum), American Beech (Fagus grandifolia), and Yellow Birch (Betula alleghaniensis). The stream varies in width from 4 to 10 m, with steep banks ranging from 10 to 80cm in height. The stream bottom consists of gravel and sand with some detritus; rocks of all shapes and sizes provide cover in the stream and along the banks. An extensive network of passages among the rocks occurs where the stream has eroded the bank. The Northern Dusky Salamander ( Desmognathus fuscus fuscus) occurs in the seepage areas along the 194 banks and the Northern Two-lined Salamander (Eurycea bislineata bislineata) is abundant through- out the stream. The Spotted Salamander (Amby- stoma maculatum), the Blue-spotted Salamander (Ambystoma laterale), and the Red-backed Sala- mander (Plethodon cinereus cinereus) occur in the woods adjacent to the stream. This is the first record of the Spring Salamander west of the Richelieu River, and only the second record of the species from outside of the Appalachian Mountain region of Quebec. Weller (1977) reported Spring Salamanders from Yamaska Mountain of the Monteregian Hills on the St. Lawrence Lowlands. More recently, he has taken four metamorphosed Gyrinophilus from one locality on Shefford Moun- tain (Figure 1) at the edge of the Appalachian Mountain range. The Dusky Salamander and the Two-lined Salamander were also found at this locality. These specimens and those I collected have 76° 75° THE CANADIAN FIELD-NATURALIST Vol. 93 been deposited in the herpetological collection of the National Museum of Natural Sciences as NMC 17816, and NMC 16379, 16767, 16791 respectively. Hemidactylium scutatum The Four-toed Salamander (Hemidactyvlium scu- tatum) has been reported from four scattered locali- ties in Quebec of which only one is south of the St. Lawrence River (Figure 1). On 29 April 1976, two metamorphosed Hemidactylium were taken from a site 11 km E of Covey Hill, Huntingdon County, Quebec (45°01’N, 73°38’W). The collecting area was a woodlot of Eastern Hemlock, Sugar Maple, American Beech, and Yellow Birch. The salamanders were discovered in a pile of bark at the base of a dead tree, close to a small woodland pond. Moss covering the rocks and logs, and bark and leaf litter provided cover. The ground was extremely wet with many scattered pools of FIGURE |. Solid circles depict localities for Gyrinophilus p. porphyriticus in Canada, triangles the known localities of Hemidactvlium scutatum in Quebec. Locality 1, Franklin Centre Gyrinophilus record, 2, Shefford Mountain Gyrinophilus record; 3, the new record for Hemidactylium. The inset maps illustrate the North American ranges according to Conant (1975): A, Gyrinophilus porphyriticus and B, Hemidactylium scutatum. 1979 temporary water. Sphagnum moss was present only in small widely scattered patches. This habitat is virtually identical to an area on Ile Perrot, Quebec where I have collected Four-toed Salamanders in past years, and to another Ile Perrot locality reported by McCoy and Durden (1965). Denman(1961, 1965) and Gorham (1955) found Hemidactylium in similar forest types. The Red-back Salamander and the Blue- spotted Salamander occurred in the immediate vicinity, and the Red-spotted Newt (Notophthalmus viridescens) was abundant in the nearby woodland pond. This is the second record of Hemidactylium south of the St. Lawrence River in Quebec. The specimens are catalogued as NMC 17539. My thanks go to Barbara L. Brown for her assistance in the field, to Wayne F. Weller who provided me with a copy of his manuscript on stream salamanders in advance of publication, and allowed me to publish his Gyrinophilus record, and to Francis R. Cook for his advice and encouragement regarding the field work and the manuscript. Portions of this study were financed by the National Museum of NOTES 195 Natural Sciences, as part of a herpetofaunal survey in Quebec in 1975. Literature Cited Conant, R. 1975. A field guide to reptiles and amphibians of eastern and central North America. 2nd edition. Houghton Mifflin Company, Boston. xvii + 429 pp. Denman, N.S. 1961. A range extension of the Four-toed Salamander in eastern Canada. Canadian Field-Natur- alist 75: 110. Denman, N.S. 1965. Further records of the Four-toed Salamander with remarks on its habitat in Quebec province. Canadian Field-Naturalist 79: 76-77. Gorham, S. W. 1955. Notes on the Four-toed Salamander in the province of Quebec. Canadian Field-Naturalist 69: 167. McCoy, C.J. and C.J. Durden. 1965. New distribution records of amphibians and reptiles in eastern Canada. Canadian Field-Naturalist 79: 156-157. Weller, W. F. 1977. Distribution of stream salamanders in southwestern Quebec. Canadian Field-Naturalist 91; 299- 303. Received 8 April 1978 Accepted I] January 1979 First Record of the Long-tailed Shrew (Sorex dispar) in New Brunswick GORDON L. KIRKLAND, JR.,! DAVID F. SCHMIDT,2 and CAROL J. KIRKLAND! 'Vertebrate Museum, Shippensburg State College, Shippensburg, Pennsylvania 17257 ?Division of Mammals, National Museum of Natural History, Washington, D.C. 20560 Kirkland, Gordon L., Jr., David F. Schmidt, and Carol J. Kirkland. 1979. First record of the Long-tailed Shrew (Sorex dispar) in New Brunswick. Canadian Field-Naturalist 93(2): 195-198. One subadult male Long-tailed Shrew (Sorex dispar) was collected in Albert County, New Brunswick, on 25 July 1978. This is the first New Brunswick record for this species and the fifth Canadian specimen collected. The New Brunswick specimen was compared to Sorex gaspensis and New England S. dispar to ascertain its specific identity. Key Words: Sorex dispar, Long-tailed Shrew, New Brunswick, first record, Sorex gaspensis. The Long-tailed Shrew (Sorex dispar) is known from Canada on the basis of four specimens (Ameri- can Museum of Natural History 17434447) collected at two localities in Quebec near the United States border: 16 km (10 mi) S of Armstrong, near Lac du Portage; and south of Cartierville, just a few yards north of the New Hampshire border (Peterson 1966) (Figure 1). With the exception of these specimens which at present cannot be located for study, S. dispar is confined to the United States in the Appalachians and adjacent mountains in a narrow belt extending from Maine to North Carolina (Kirkland and Van Deusen 1979). The closely related Gaspé Shrew (Sorex gaspensis) is an endemic Canadian species which until recently was thought to be restricted to the Gaspé region of Quebec. But it has recently been collected in such disjunct locations as Mount Car- leton, New Brunswick (Peterson and Symansky 1963) and Cape Breton Island, Nova Scotia (Roscoe and Majka 1976) (Figure 1). In an attempt to determine whether the hiatus in the distribution of S. gaspensis is real or the product of inadequate sampling, field work was conducted in southeastern New Brunswick and in the Cobequid Mountains and on North Mountain in 196 THE CANADIAN FIELD-NATURALIST GASP E fo} NEW BRYN SW I FIGURE |. Vol. 93 PEON SOLA g fr sS PRINCE EDWARD ISAM CAPE BRETON ISLAND Kilometers Location of capture sites of Sorex gaspensis (open circles), Maine Sorex dispar (dots), SSC 8393 (dot within circle), and previously recorded Quebec S. dispar (triangles), based on Godin (1977), Kirkland and Van Deusen (1979), and Peterson (1966). One mark may represent more than one locality if several specimens have been collected from different localities in the same region. Nova Scotia during July 1978. In 2057 trapnights (TN) of sampling effort at six localities in Nova Scotia and New Brunswick, no S. gaspensis were collected; however, on 25 July 1978, a single subadult male S. dispar was trapped 5.3 km N, 3.5 km W of Riverside-Albert, Albert County, New Brunswick. This locality is approximately 305 km ESE of the nearest previous locality for this species at Beaver Creek, Aroostook County, Maine (Godin 1977). The New Brunswick specimen (Shippensburg State College 8393) was captured in a Museum Special trap baited with rolled oats and set in rocks, 30cm below the surface, on a rocky east-facing hillside in a deciduous-coniferous forest (elevation 180 m). The capture site was dominated by Yellow Birch (Betula alleghaniensis) with Mountain Maple (Acer spicatum) and Balsam Fir (Abies balsamea) as subdominants. Living ground cover at the site averaged 51-75% and consisted of mosses, ferns, and seedlings of Mountain Maple. In 306 TN of sampling effort, 23 small mammals representing six species were captured in the trapline that yielded the single S. dispar: seven Smoky Shrews (Sorex fumeus), four Short-tailed Shrews ( Blarina brevicauda), one North- ern Flying Squirrel (Glaucomys sabrinus), eight Southern Red-backed Voles (Clethrionomys gap- peri), and two Woodland Jumping Mice (Napaeo- zapus insignis). The trap in which the S. dispar was captured had yielded a male S. fumeus the previous morning (24 July). The standard external measure- ments and 16 selected cranial and mandibular characters for specimen SSC 8393 are presented in Table |. Because the New Brunswick specimen was captured near the center of the disjunction in the range of S. gaspensis and over 300 km from the nearest S. dispar 1979 TABLE !—Measurements for six external and 16 cranial/ mandibular characters of New Brunswick specimen SSC 8393 and the ranges of measurements for these 22 characters in 23 S. gaspensis and 56 New England S. dispar (All linear measurements in millimetres; weight in grams) Character SSC Range Does S. gaspensis S. dispar Total length 123 95-127 103-136.5 Tail length 56 45- 55* 46- 61 Hind foot length 13 10.5 -12.5* 12- 15 Ear length 8 5- 9 5- 9 Body length 67 45-77 48- 79 Weight 4.2 2.2 - 4.3 4.0 - 4.9 Greatest length [ASS ISAS les* svleS a 18e3, Condylobasal length 16.8 15.35-16.35* 16.45-17.7 Interorbital breadth 3.15 2.8 — 3.5 3.1 — 3.6 Cranial breadth 8.25 Toll = Sa 7.5 — 8.3 Molariform tooth TOW 3.9 Bre 3-69%80 3/5 — 43 Cheek tooth row 4.55 4.05-— 4.4* 4.1 - 49 Total tooth row 7.05 6.35- 7.25 6.8 — 7.6 Incisor width 1.15 1.0 - 1.3 1.1 - 1.4 Canine width 1.65 ES lRS Sth elE4S eS: Molar width 3.8 3.2 — 3.8 3.65— 4.15 Nasal length 6.25 5.4 — 6.3 5.6 — 7.25 Palatal length V2 6.0 — 6.75* 6.6 — 7.5 Post-palatal length 8.2 6.9 — 7.6* 7.3 — 8.35 Mandible length I 10.15 9.1 -— 9.9* 9.75-10.75 Mandible length II —:11.00 9.7 -10.8* 10.1 -11.35 Mandible height 32 2.95— 3.95 3.05— 4.0 *SSC 8393 exceeds range for character. locality, it was important to ascertain its specific identity correctly. Was it a S. dispar, a S. gaspensis, or was it intermediate between these two taxa? The existence of an intergrade specimen at this New Brunswick locality would call into question the specific status of S. gaspensis (Kirkland and Van Deusen 1979). Univariate and multivariate statistical analyses were performed to compare the New Brunswick specimen with 23 S. gaspensis and 56 New England S. dispar. For localities of these specimens and descriptions of the measurements see Kirkland and Van Deusen (1979). All skulls were measured by the senior author with Helios dial micrometers (calibra- tion 0.05 mm) under a dissecting microscope. Comparisons of the New Brunswick specimen with S. gaspensis and New England S. dispar revealed that the New Brunswick specimen exceeded the size range of S. gaspensis for 12 of 22 characters and equalled the largest known specimen for one other (Table 1). The New Brunswick specimen fell within the size range of New England S. dispar for all 22 characters (Table 1). Data from 247 S. dispar with locality latitudes ranging from from 35.6° N (North Carolina) to NOTES OF 45.9° N (Maine) were analyzed with the SPSS Computer Package (Nie et al. 1975) to produce regression lines of size versus latitude for each of 22 characters. These regression equations were used to predict the characteristics of a S. dispar from the latitude of the New Brunswick specimen (45.8° N). The New Brunswick specimen fell within the 95% confidence interval of the predicted value (see Zar 1974) for 21 of 22 characters (Table 2). In addition, discriminant function analyses (BMD 07M, Dixon 1967) were performed on the New Brunswick speci- men, 14 S. gaspensis, and 25 New England S. dispar using 18 characters. The New Brunswick specimen was assigned by the computer analysis to S. dispar both when assigned to S. gaspensis ( P = 0.753) and to New England S. dispar (P= 1.000). The respective discriminant scores (first two canonical variables) for SSC 8393, the 14 S. gaspensis, and 25 S. dispar were = 0358s = L010 x = 2 965. \(range —45I 9 ato =12395)), Yi = —0:000) (range: = —1-956n tom 24 Nl): X= 1779 (range 01169) to 431479), Yo= 0.000 @Gange™ —1-499) to 12788), when) SSG 8393 was assigned a priori to S§. gaspensis, and X = +3.415, Y=-0.010; X= -4.402 (range -5.944 TABLE 2—Comparisons of the observed values of 22 morphological variables in New Brunswick Long-tailed Shrew (Sorex dispar) specimen (SSC 8393) from 45.8°N with the predicted values and 95% confidence intervals of a Sorex dispar from that latitude based on regression analysis of 247 S. dispar collected at latitudes 35.6°-45.9°N 95% Character SSC _ Predicted Confidence 8393 value interval Total length 123 115.92 103.76-128.08 Tail length 56 54.68 49.02- 60.34 Hind foot length 13 13.06 11.51- 14.61 Ear length 8 7.54 4.51- 10.57 © Body length 67 61.35 5056= 72514 Weight 4.2 4.37 DM= O03} Greatest length 17.85 18.02 17.34— 18.70 Condylobasal length 16.8 16.97 16.23- 17.71 Interorbital breadth 3.15 3.47 3.13— 3.81 Cranial breadth 8.25 7.87 V3= B37 Molariform tooth row 3.90 4.07 3.87— 4.27 Cheek tooth row 4.55 4.46 4.22— 4.70 Total tooth row 7.05 7.30 6.98=— 7262 Incisor width 1.15 1.36 1.22— - 1.50 Canine width 1.65 1.51 les38= 1k69 Molar width 3.80 3.89 36l= 4217 Nasal length 6.25 6.20 5.62— 6.78 Palatal length V2) 7.04 6.64— 7.44 Post-palatal length 8.20 7.94 7.46— 8.42 Mandible length I 10.15 10.08 9.66— 10.50 Mandible length II 11.00 11.01 10.49— 11.53 Mandible height 3.20 3.50 3.12— 3.88 198 to —2.916), Y =-0.000 (range -1.708 to +1.595); XK = +2.371 (range -0.633 to +4.744), Y = -0.000 (range -2.226 to +1.901) when SSC 8393 was assigned a priori to S. dispar. The results of these three analyses suggest that the New Brunswick specimen is a Sorex dispar and resembles S. dispar from New England. Acknowledgments The field work in Nova Scotia and New Brunswick was supported by a grant from The Explorers Club. We thank Henry W. Setzer and personnel of the New Brunswick Department of Natural Resources and the Nova Scotia Department of Lands and Forests for their assistance. We acknowledge Don E. Wilson and Michael A. Bogan for critically reviewing this manuscript. Literature Cited Dixon, W.J. 1967. BMD Biomedical Computer Pro- grams. University of California Press, University of California Publications in Automatic Computation, Number 2. 600 pp. Godin, A. J. 1977. Wild mammals of New England. Johns THE CANADIAN FIELD-NATURALIST Vol. 93 Hopkins University Press, Baltimore. 304 pp. Kirkland, G. L., Jr. and H. M. Van Deusen. 1979. The shrews of the Sorex dispar group: Sorex dispar Batchelder and Sorex gaspensis Anthony and Goodwin. American Museum of Natural History Novitates. Jn press. Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and D.H. Bent. 1975. Statistical package for the social sciences. 2nd edition. McGraw-Hill Book Company, New York. 675 pp. Peterson, R. L. 1966. The mammals of eastern Canada. Oxford University Press, Toronto. 465 pp. Peterson, R.S.and A. Symansky. 1963. First record of the Gaspé Shrew from New Brunswick. Journal of Mam- malogy 44: 278-279. Roscoe, B. and C. Majka. 1976. First records of the Rock Vole (Microtus chrotorrhinus) and the Gaspé Shrew (Sorex gaspensis) from Nova Scotia and a second record of the Thompson’s Pygmy Shrew ( Microsorex thompsoni) from Cape Breton Island. Canadian Field-Naturalist 90: 497-498. Zar, J. H. 1974. Biostatistical analysis. Prentice-Hall, Inc., Englewood Cliffs, New Jersey. 620 pp. Received 20 November 1978 Accepted 11 January 1979 Intraspecific Food Theft by the American Kestrel PETER M. FETTEROLF Department of Zoology, University of Toronto, Toronto, Ontario M5S 1A1 Fetterolf, P. M. 1979. Intraspecific food theft by the American Kestrel. Canadian Field-Naturalist 93(2): 198. One male American Kestrel, Falco sparverius, was observed in an unsuccessful attempt to steal prey from another male American Kestrel. Key Words: American Kestrel, prey theft, Falco sparverius. On 24 March 1978, I watched two male American Kestrels, Falco sparverius, at the southern edge of the Toronto Island Airport, Toronto, Ontario. The weather was cold (+3°C) and clear. The birds, perched approximately 5 m apart in 15- to 20-m willows, Salix rigida, faced the open grassland to the north. At 12:02 EST, one kestrel plunged from its perch and captured a rodent, probably a vole, Microtus sp. The bird did not mantle its prey, whereupon the second male swooped down from its perch, grabbed the vole, and rapidly flew north low across the airport runways. The victim took flight immediately, climbed above the robber and dove upon it three times, shrieking the “klee” call just prior to contact each time. At each contact, the pursurer apparently dug its talons into the fleeing bird. On the third assault, the robber dropped the rodent and departed. The other kestrel pounced on the prey at once, assumed an exaggerated mantling posture with the wingtips nearly touching, and ate the rodent. The theft, chase, and successful recapture of the prey took less than 90 s and covered a distance of about 40 m. No reports of prey theft by American Kestrels were found but a European Kestrel, F. tinnuculus, robbed prey from a Short-eared Owl, Asio flammeus (Boyle, G. L. 1974. Kestrel taking prey from Short-eared Owl. British Birds 67: 474-475). Prey-robbing by kestrels seems to be rare, but it is not surprising that the observed encounter occurred at a time of the year when food supplies were likely to be low. Special thanks go to Keith Bildstein for comment- ing on an earlier draft. Received 3 November 1978 Accepted 14 January 1979 1979 NOTES 199 First Record of the Northern Brook Lamprey, Ichthyomyzon fossor, in the Nelson River Drainage, Manitoba J. JYRKKANEN!? and D. G. WRIGHT3 ‘Manitoba Department of Renewable Resources, Hadashville, Manitoba *Present address: Box 396, Garibaldi Highlands, Squamish, British Columbia VON ITO ‘Fisheries and Environment Canada, Fisheries and Oceans, Freshwater Institute, 501 University Crescent, Winnipeg, Manitoba R3T 2N6 Jyrkkanen, J. and D. G. Wright. 1979. First record of the Northern Brook Lamprey, /chthyomyzon fossor, in the Nelson River drainage, Manitoba. Canadian Field-Naturalist 93(2): 199-200. Fourteen specimens of Northern Brook Lamprey, /chthyomyzon fossor, were collected from the Birch River, a tributary to the Winnipeg River in southeastern Manitoba on 8 and 13 May 1977. This represents the first record of this species in the Nelson River drainage and is a significant extension of the presently known range. Key Words: Ichthyomyzon fossor, range extension, Nelson River drainage, Manitoba, new records, geographical distribution. Fourteen specimens of Northern Brook Lamprey, Ichthyomyzon fossor, were captured by hand from the Birch River, upstream of the town of Prawda, Manitoba (49° 39’N, 95°48’W) on 8 and 13 May 1977. The Birch River is a tributary of the Winnipeg River, in the Nelson River system, flowing into Hudson Bay. This represents the first record of this species in the Nelson River drainage system and Is significant in that it is a range extension of over 500 km from the nearest known population. Identification of the specimens was confirmed by E. J. Crossman of the Royal Ontario Museum. Morphological characteristics on which species iden- tification is based are as follows: lateral teeth all uni- cuspid; the supraoral lamina a single bicuspid tooth; infraoral lamina cusps blunt; diameter of the sucking disc less than one half the length of the branchial region; myotomes in the trunk, between the last gill opening and the anus, number between 49 and 54; total length of mature individuals between 100 and 255 mm. The specimens have been deposited in the collection of the Royal Ontario Museum, Toronto and are catalogued under the number ROM 34264. Of the 14 specimens collected, 10 were males, averaging 116.5 mm (SE =3.6 mm), and four were females, averaging 136.5 mm (SE = 9.6 mm) in total length. All specimens were sexually mature with females having free ova within the body cavity. It is assumed that they were spawning at the time of collection. Ichthyomyzon fossor is currently known to exist over a very limited range. Hubbs and Trautman (1937) report that the species is abundant in all of the Great Lakes drainages of the State of Michigan (Erie, Huron, Michigan, and Superior) and from Scott Creek in the Mississippi River drainage of Wiscon- sin. Leach (1940) reports the occurrence of the species in the Tippecanoe River (Mississippi River drainage) in northern Indiana. Dymond (1947) indicates that J. fossor was resident in the Thames River system, while Vladykov (1949) reports that specimens were collect- ed in the Yamaska and St. Francis rivers of southern Quebec. Scott and Crossman (1973) add that the distribution of the species includes Georgian Bay, Lake Nipissing, and the north shore tributaries of Lake Superior. The Birch River is a small, fairly slow-moving river with an estimated maximum flow of 5.7-8.5 m3/s (200-300 ft3/s) and a low flow of less than 0.15 m3/s (5 ft3/s). Winter flows may be augmented by overflow from the Shoal Lake viaduct entering the upper reaches of the Birch River at East Braintree. The substrate of the Birch River is highly varied with silty sediments in the quieter reaches of the stream, gravel and cobble riffles, bedrock outcroppings, and several small waterfalls. A thick mat of algae covers the . shallow, rocky substrates in the summer. Hubbs and Trautman (1937) state that /. fossor lives in creeks and small rivers, apparently avoiding both small brooks and large rivers. It has never been recorded in lakes, either small or large. This would seem to preclude the hypothesis that the Birch River population has been derived from either Lake Superior or Wisconsin populations. Vladykov (1949) reports that ammocoetes of /. fossor are sold as bait for sport fishing in Quebec. This statement tends to support a hypothesis that the Birch River population may have been introduced by an angler who dis- carded his remaining bait after a fishing trip. Because the species 1s not parasitic, the presence of I. fossor in the Nelson River drainage basin should not present any possibility of endangering or destroying native fishes. 200 Acknowledgments The authors acknowledge the assistance of D. P. Scott, Fisheries and Oceans, Winnipeg, C. C. Lind- sey, Department of Zoology, University of Manitoba, Winnipeg, and E. J. Crossman, Curator of Ichthy- ology and Herpetology, Royal Ontario Museum, Toronto, in confirming the identification of the specimens, and thank W. Coder and J. Johnson, Manitoba Department of Renewable Resources, Hadashville, for their assistance in the field. Literature Cited Dymond, J.R. 1947. A list of freshwater fish of Canada east of the Rocky Mountains, with keys. Royal Ontario Museum of Zoology, Miscellaneous Publication Number 1. 55 pp. THE CANADIAN FIELD-NATURALIST Vol. 93 Hubbs, C. L. and M. B. Trautman. 1937. A revision of the lamprey genus /chthvomyzon. Museum of Zoology, Uni- versity of Michigan, Miscellaneous Publication Number 35. 109 pp. Leach, W. J. 1940. Occurrence and life history of the northern brook lamprey, /chthvomyzon fossor, in Indi- ana. Copeia 1940 (1): 21-34. Scott, W. B. and E. J. Crossman. 1973. Freshwater fishes of Canada. Bulletin of the Fisheries Research Board of Canada 184: 49-51. Vladykov, V. D. 1949. Quebec lampreys ( Petromyzonidae). 1. List of species and their economic importance. Depart- ment of Fisheries, Province cf Quebec, Contribution Number 26. 67 pp. Received 6 November 1978 Accepted 12 January 1979 Blue Grouse Brood Hen — Black Bear Confrontation M. G. SULLIVAN Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 Sullivan, M. G. 1979. Blue Grouse brood hen — Black Bear confrontation. Canadian Field-Naturalist 93(2): 200. On 14 July 1977, while engaged in a census of Blue Grouse (Dendragapus obscurus) near Courtenay, Vancouver Island, British Columbia, I heard a banded female grouse clucking loudly (indicative of brood disturbance) as I approached the area. From about 75 m away I observed the grouse flying at a Black Bear (Ursus americanus). Flaring about a metre from the bear, the hen then landed on the ground about 20 m away, clucking and rushing about. The bear ran towards the grouse, whereupon she flew at the bear and again landed some distance away. Through these maneuvers, the hen seemed to lead the bear out of a marshy valley onto a rocky ridge. At this point, the bear appeared to catch my scent and ambled out of sight. The hen then returned to the marshy area. Although no chicks were found, I later discovered that this hen had a brood at that time. Her behavior was similar to that displayed towards humans when chicks are present. Searching the area, I found several stumps that had been freshly torn apart by a bear searching for insects. Ripe Huckleberries (Vaccinium ovalifolium) were plentiful and are an important food for bears in this area (unpublished data). Thus, a food shortage was unlikely at the time. An interesting addition to this incident is that on31 May 1977, I had found a leg band from an adult male Blue Grouse in a month-old bear scat in the same general area. This bird was banded as an adult on 16 June 1975, about 2.4 km from where the band was found and may have been eaten as carrion. Received 14 February 1978 Accepted 7 November 1978 News and Comment Editor’s Report for 1978 The Canadian Field-Naturalist received 149 manu- scripts for consideration in 1978. This number is consistent with the 147 submitted in 1976 and the 137 uel OMT In 1978, 77 papers (38 Articles and 39 Notes) originally submitted from 1975 to 1978 were publish- ed in Volume 92 of The Canadian Field-Naturalist. The breakdown according to subject matter is as follows: birds, 25; mammals, 23; plants, 17; inverte- brates, 5; amphibians and reptiles, 4; fishes, 2; and other, 1. Comparison with the number of manuscripts published in the two previous years, 124 (34 Articles and 90 Notes) in 1976 and 101 (30 Articles and 71 Notes) in 1977, shows that the number of Notes published in 1978 was considerably lower. No explanation is apparent for this; certainly there have been no particular delays in the publication process nor any changes in acceptance criteria. Occasionally there is a long lapse after a manuscript has been returned to its authors for revision and before it is resubmitted and accepted for publication. In fact, this interval has even exceeded three years! Therefore, the proportions of accepted papers with respect to the years in which they were submitted (number accepted/number received) have increased over the earlier published figures. The revised propor- tions are as follows: 1974 — 116/152; 1975 — 122/ 167; 1976 — 90/147; and so far for 1977 — 86/137. The financial position of The Canadian Field- Naturalist is currently good because several authors have paid page charges for all published pages rather than just the obligatory charges for pages over six. Therefore, for the first time in several years, it was not necessary to submit an application for a grant to the Natural Sciences and Engineering Research Council (formerly the granting body was the National Re- search Council of Canada). Furthermore, our printer, M.O.M. Printing of Ottawa, has tried to keep our printing costs down while at the same time continuing to put out a quality product. The financial outlook then for 1979 is conditionally bright. LORRAINE C. SMITH Editor Proposals Invited from Field Research Investigators The Center for Field Research, a non-profit organization established to raise private funds for field research and to encourage public understanding of science, is currently accepting proposals for 1980. Projects are considered on the basis of scientific merit and their ability to utilize the assistance of motivated lay volunteers in the field. In the past seven years, The Center for Field Research and its affiliate, Earthwatch, have raised over $1.5 million from interested members of the public who have also contributed their time and skills to worthy research in a wide variety of disciplines. This year, it expects to grant another $500,000 in support of 66 projects, including the observation of sea cow distribution and behavior, western Australia; a study of the ecology and social behavior of the spotted hyena, Kenya; an archaeological and archi- 201 tectural investigation of Repton Anglo-Saxon church, England; an anthropological study of return- ing migrants, Newfoundland; a paleontological sur- vey and fossil collection of Devonian rocks, Idaho; and an archaeological study of early Iron Age settlement and economic systems, West Germany. Proposal deadlines for 1980 research are: | June 1979 (for work beginning after | December 1979): | October 1979 (for work beginning after | June 1980): and 15 January 1980 (for work beginning after | September 1980). Scholars of all nationalities and from all disciplines are invited to apply. For appli- cation guidelines write Nancy Bell Scott, The Center for Field Research, 10 Juniper Road, Box 127-Q, Belmont, Massachusetts 02178 (Phone 617- 489-3032). 202 Inland Bird Banding This journal, formerly Inland Bird Banding News, published by the Inland Bird Banding Association and printed by Allen Press, has been drastically changed by its new Editor. It is now a quarterly refereed journal and will no longer contain the news and other columns that it once had. These items will continue in some form in a newsletter. The Editor is actively soliciting manuscripts that deal with bird-banding techniques and the results of bird-banding studies, especially studies dealing with birds from middle North America (Canada to the Gulf Coast). Interested ornithologists and students of birds who have suitable material for publication (preferably manuscripts of 10 pages or less in length) should THE CANADIAN FIELD-NATURALIST Vol. 93 contact the Editor, Jerome A. Jackson, Department of Biological Sciences, Mississippi State University, Mississippi State, Mississippi 39762 (phone 601- 325-5722). Bluebird Society A new society, the Bluebird Society of North America, has been formed. Its address is as fol- lows: Box 6295, Silver Springs, Maryland 20906. We thank Colleen Hislop for correcting the page proofs for this issue. Book Reviews ZOOLOGY Where to Find Birds in British Columbia By David M. Mark. 1978. Kestrel Press, New Westminster. 72 pp., illus. Paper $3.00. (Available by mail from Kestrel Press, P.O. Box 2054, New Westminster, British Columbia V3L 5A3.) This small book (13.5 X 21 cm) gives bird-finding information for 49 areas (“sites”) in British Columbia (hereafter B.C.), which are described, region by region, for eight geographic regions of the province. A sketch map shows the general locations of birding areas in each region, but the reader will need a good road map to use with the book. The introduction (six pages) includes a helpful discussion of bird dis- tribution in B.C. as related to the Biotic Areas of Munro and Cowan (B.C. Provincial Museum Special Publication 2, 1947). Following this, eight pages are occupied by a photocopy of the recent (1977), com- prehensive checklist of B.C. birds issued by the Provincial Museum, which identifies those species known to breed in B.C. and those of accidental occurrence. The main part of the book (pages 19 to 63) is devoted to area accounts, which vary in length from five lines to two pages. These are generally up-to-date and informative, with specific and accurate route directions, including distances (the author is a geographer!), and usually with lists of particular bird species to expect. Finally, there is a seven-page section on “sought-after species” (those most often sought by birders), with references to area accounts and some- times additional comments as well. Besides the author, 15 other individuals or groups contributed information for the book. My only major criticism of this guide is that coverage is poor for some parts of B.C. (As I was one of the contributors, | am perhaps indicting myself for not providing more information!) For example, Vancouver Island (and the Victoria area in par- ticular) receives rather scanty treatment considering its diverse avifauna and the heavy visitation it receives from out-of-province birders. None of the five national parks in B.C. is mentioned. On the other hand, coverage is quite good for the Vancouver area and most of the southern interior, and the accounts included for those areas can scarcely be faulted. | sincerely hope the gaps in coverage will be filled ina future edition. All in all, this is a very good bird-finding guide, at least for a pioneering effort in a province previously lacking one. It will prove very helpful both to B.C. birders and to those visiting the province from elsewhere. WAYNE C. WEBER Department of Biological Sciences, Mississippi State Uni- versity, Mississippi State, Mississippi 39762, USA The Moths of America North of Mexico, Including Greenland. Fascicle 22.2, Noctuoidea (in part): Lymantriidae By D.C. Ferguson. 1978. Classey and the Wedge Entomo- logical Research Foundation (distributed in North America by Entomological Reprint Specialists, Los Angeles). US $48 ($40 by subscription). This fascicle, often advertized as soon appearing, finally did. It was eagerly awaited by all who will have to use it. We have to admit that the Lymantriidae area very difficult family and many things had to be elucidated to make publication useful: names, distri- bution, and general considerations. This begins with the family name, and everybody concerned is glad that there were no changes in this as well as in often used generic names. These have been not quite stable in the past because of international faunal and taxonomic complications, like Gynae- phora, Dasychira, Lymantria, Leucoma, Euproctis. Appreciable conservatism 1s also shown in specific names, e.g., Euproctis chrysorrhoea. One may hope that this final nomenclature finds easy access into our collections and papers. One might question whether it was necessary to fill the literature with all the new “subspecific” names. More biochemical and larval taxonomic research would probably have obliterated some of the “sub- species” and shown them to be species in their own right. Here is certainly a field open for further research. The keys are in general good, short, useful and working. The larval key to the last instar larvae of Orgvia, however, leaves a question open: how can there be a description ofa larva of a “subspecies” when thee somscalled@ sty pels 1OlethisiessubSpeciesia mS) as questionable as can be (“/eucostigma plagiata’) and the larvae in question just do not yield adults like this “type”? Otherwise this key is helpful and correct. Certainly the most difficult genus is Dasychira. LS) 204 THE CANADIAN FIELD-NATURALIST Here Ferguson, who was especially involved for decades with this genus, can proudly show the fruits of long labor. The explications about the generic name now make Dasychira finally definite for our species, and we have to congratulate Ferguson for his success in re-finding D. tephra. From this point of departure everything else falls into place. We are also thankful that he was able to confirm and give new meaning to the names and taxa so familiar to all of us but so often doubted by overzealous workers over many years. I mean obliquata, cinnamomea, atrivenosa, meridion- alis, basiflava, leucophaea, dorsipennata, plagiata, grisefacta. Other names like aridensis, parallela, lemmeri, pini disappeared after having been the reason for much misunderstanding in the past. The other difficult genus in the family is Orgyia. Regrettably one cannot say what we just said about Dasychira in this case. There are many needless mistakes in Ferguson’s treatment. There are also good points, and these are the following: (a) the confirmation of the fact that all North American species belong to one only well-defined genus, Orgyia. (6) the confirmation of O. cana as species in its own right. This, however, is not a “revised status,” as it was already treated as such in Entomologische Zeitschrift 83: 12, 1.e., in 1973, a publication which is left out of the literature quoted. The problem with the associa- tion of the larvae in this group would have been completely and easily solved if Ferguson had used the preserved and associated specimens of the McFarland rearings in the Los Angeles County Museum of Natural History. It has to be pointed out again that MONA authors seem to rely too exclusively on the collections of the United States National Museum, as already Rindge objected in Journal of the Lepi- dopteran Society 28: 4; (c) giving taxonomic status to these puzzling very large California Orgyia (magna). Further research 1s here very much necessary: (d@) the detection of the old Boisduval specimen which Guérin-Méneville figured and used for the name of O. detrita, in the collection of the United States National Museum. It should be remarked that “Degens Bd. Am. B.” is not “apparently... an unpublished name” but is Latin for “coming from Boisduval America Borealis” (the Latin verb is dego, -i, -ere); (e) the additional knowledge about O. falcata and its larva. On the negative side there have to be mentioned the following points: (a) the laconic statement “the female genitalia have not been studied.” Besides the fact that they have been very extensively studied in several issues of Ento- mogische Zeitschrift and are especially in the genus Orgvia of decisive taxonomic importance, in a Vol. 93 “definite” presentation of North American moths, this is not quite understandable; (b) the omission of all observations about pupae is likewise not quite understandable, the more so as the form of the pupa is also species characteristic; (c) the omission of any study of the eggs by means of the scanning electron microscope (SEM) and electro- phoresis and also of allimportant larval characters, as mandibles, ocelli, setae on thoracic legs, headcap- sules, surface of exterior cuticle; (d) the claiming of a “new synonymy” for O. definita kendelli on p. 75 while this synonymy was long established in Entomologische Zeitschrift 83: 14 (Definita kendelli = leucographa), 1.e., in 1973; (e) the establishing of “new status” in the rindgei/ leuschneri complex while Chua et al. had already established and published in Journal of Research on the Lepidoptera 15: 4 (1976) specific status for O. rindgei. This has been also repeated by Riotte in Entomologische Zeitschrift 87: 3 (1977). Some special words have to be said on the “enfant terrible” of the whole fascicle: O. /euwcostigma. Looked at as Ferguson presents it, it well makes sense. The fact, however, is that the realities are quite other: (a) The aedoeagus which is said “may be at once distinguished from all other species in that it is apically tapered to a point” is blunt and not tapered to a point as SEM micrographs show. (6) The (not studied) female genitalia would have helped to an other and correct classification. (c) Use of the earlier published results of egg electrophoresis would have shown that indeed /eucostigma is sympatric to the extreme with wardi which in no way 1s a “subspecies” or “synonym” of anything. (d) Larval structures, if they had been used, would have shown the same (they also were previously published). (e) To use the so called “Walker type” of Acypha plagiata, abdomen- less as it is and without any locality label, as type fora “subspecies” of /ewcostigma in Nova Scotia, replacing wardi, 1s at best absurd. Insect pins may be convincing sometimes; in a case like this, certainly not. The Walker specimen could be used for the real /euco- stigma in Nova Scotia but I can see no need for this. Competent workers in the British Museum came toa quite other evaluation of the Walker specimen and placed it together with /eucostigma from Wisconsin as best match. Therefore, one should list the Walker type as “incertae sedis.” One good thing at least should be mentioned, however: with Abbot’s larval painting of leucostigma declared as lectotype of the species, we have now a good basis for it. We also think that the male figure of Abbot’s /ewcostigma is really definita (many of Abbot’s plates are mixed with non- conspecific creatures). Very good also is Ferguson’s final clearing up of the /ewcographa Geyer problem: we have nothing against his treatment here. 1979 The plates in this volume are of the expected quality and with the richness of depicted specimens very useful. The only thing we miss Is a single specimen of Orgvia wardi from the type locality, Prospect Road, N.S., perhaps best together with a /eucostigma from the same locality to show the difference. The Nova Scotia specimens figured are all /eucostigma. In the literature quoted we miss the important paper by Chua et al., 1976, Investigation of selected species of the genus Orgyvia (Lymantriidae) using isoelectrofocusing in thin layer polyacrylamide gel (Journal of Research on the Lepidoptera 15(4): 215— 224) as well as Riotte, 1973, Uber Orgvia (O.) gulosa und Orgyvia (O.) cana (Lep.: Lymantriidae) (Ento- mologische Zeitschrift 83(12): 129-140) and by the Hawks, Falcons and Falconry By Frank L. Beebe. 1976. Hancock House, Saanichton, British Columbia. 320 pp., illus. $12.95. Any author attempting to write an original book on birds of prey or falconry today, faces a difficult task, but Beebe certainly does offer something new in this most recent book. Perhaps the book should be retitled “Hawks, Falcons, Falconry and Human Attitudes”; through the entire book, there pervades an undertone of bitterness towards an anti-falconry movement, which culminates in the final chapter, “The endangering of the Peregrine: A study in environmental strategy.” A reader cannot help but be impressed by the collection of facts which, according to the author, led to classifying the Peregrine Falcon as an endangered species. After seven years of intense involvement with birds of prey and their human admirers, be they scientists, bird-watchers, falconers, photographers, or others, I find it difficult to believe that these facts constitute a case for the premeditated abolition of falconry. Biologists will be disappointed by the rather brief treatment of general raptor biology but Beebe demonstrates a good grasp of raptor behavior. His observations and theory on the adaptive significance of a reversible hallux, prone sleeping position, and very deep sleep associated with Arctic-nesting Gyr- falcons (p. 167) is interesting and certainly warrants further investigation. On p. 180, the author provides some fascinating insight into the influence of geo- graphical landforms on hunting strategies of river- nesting Peregrine Falcons. The descriptions for each species covered in the book are fairly standard, but tend to lean toward the western areas of Canada with which the author is clearly most familiar. Much, if not all, of this material has been reproduced from his earlier published work BOOK REVIEWS 205 same author, 1977, Abschliessende Bemerkungen zu den Studien uber nordamerikanische Arten der Gattung Orgvia (Lepidoptera: Lymantriidae) (Ento- mologische Zeitschrift 87(3): 9-12, concerning defi- nita, rindgei, and leucographa = detrita). The criticisms, however, do not interfere with the great value of the fascicle otherwise. We certainly welcome its final appearance and wish it good and longlasting success. URC2 EMR TOnmeE Department of Entomology, B.P. Bishop Museum, P.O. Box 6037, Honolulu, Hawai, USA 96818 and Royal Ontario Museum, Toronto, Ontario “Field studies of the Falconiformes of British Columbia,” No. 17 Occasional Paper Series, British Columbia Provincial Museum. The author is some- what inconsistent in his descriptions of the various subspecies, providing them for some species, but not for others. The section on falconry techniques should prove extremely useful to falconers, as Beebe has chosen to present detailed accounts of certain aspects of this art rather than give a shallow description of falconry as a whole. For example, he has given a thorough description of how to fly a bird to the lure (p. 239). Having conducted research on kestrel reproduction, I must dispute the author’s statements regarding the timing of the breeding for this species (p. 132). Beebe claims that “kestrels arrive on their breeding terri- tories in April” and that “eggs are produced, depending on latitude, from late May to mid-June.” In southern Quebec at least, kestrels generally arrive as early as the latter half of March and lay eggs during the last two weeks of April and the first two weeks of May, somewhat dependent on weather conditions from year to year. Furthermore, my studies and those of others (Bent, A. C. 1938. Life histories of North American birds of prey. Part 2: Orders Falconiformes and Strigiformes. Dover Publications, New York. 482 pp.; Balgooyen, T. G. 1976. Behavior and ecology of the American Kestrel (Falco sparverius) in the Sierra Nevada of California. University of California Pub- lications in Zoology 103: 1-83), give an average egg size of kestrels as approximately 35 X 28 mm rather than the 39 X 29 mm Beebe reported. The first-year mortality rate of 90% quoted in this book for birds of prey on p. 207 is surely the highest ever reported in the literature. Also, this reviewer was 206 not aware that the seal hunt had been closed (see p. 298). There is much unnecessary repetition in photo- graphs and an overabundance of head studies, some of very poor quality, e.g., Peale’s Falcon, p. 170. The ink drawings, with some exceptions, are quite well done, as well as functional. The flight silhouettes throughout the book are excellent. One illustration, that of a falconer’s knot, is apparently missing on p. 193, and on p. 239 the labels on the two diagrams apparently have been assigned incorrectly. These latter two errors exemplify the poor editing found throughout the book. More than 50 typo- graphical and spelling errors were located in the text and there was inconsistent use of names, i.e., kestrel versus sparrowhawk, and even spelling, 1.e., grey versus gray. References were not presented in a consistent manner and sometimes are incorrect, e.g., Morlan W. Nelson (1965), p. 152. Much space is Fishes of the World By J.S. Nelson. 1976. Wiley, New York. xii+416 pp., illus. US $26.00. The author wrote this book “to present a modern introductory systematic treatment of all major fish groups.” It is intended to serve as a reference for professionals and as a text for fish systematics courses. This book, therefore, was doomed to failure before it was written, and ensured of success before it was off the press. Fish systematics is ina state of rapid change and the healthy diversity of opinion on various aspects of fish systematics assures that some of the included material is outdated, but the need for any recent synthesis is obvious to anyone trying to teach the subject. Following the prefatory remarks, the text includes an Introduction which comments on numbers of fishes, their diversity, distribution, and classificatory schemes. Then follows a classification of the phylum Chordata in which fishes are treated to at least the familial level. Appendix I is a checklist of extant classes, orders, suborders, and families. Appendix 2 is a series of 45 distribution maps of families of fishes. There is an extensive bibliography and the Index includes the genera mentioned in the text. Nelson considers six classes of fishes (four extant) comprising 46 orders and 450 families. An outline drawing of a “typical” representative of each of most families is included. Characteristics for each included family or higher taxon are usually included, although they are not always complete. Recent revisionary papers on the orders are mentioned and comparisons THE CANADIAN FIELD-NATURALIST Vol. 93 occupied by overly emphatic titles in the first nine sections where only brief treatment of the subjects follows. There is no doubt that Beebe has had much experience with both wild and captive birds of prey and for this reason, I am more inclined to recommend his other book, “Field studies of the Falconiformes of British Columbia” for raptor en- thusiasts not especially interested in falconry. Fal- coners from coast to coast, however, will likely applaud Beebe’s efforts in this book to speak on their behalf and it is to this audience that I heartily recommend this book. DAVID M. BIRD Raptor Research, Department of Renewable Resources, Macdonald Campus of McGill University, Ste. Anne de Bellevue, Quebec H9X 3MI with previous classificatory syntheses are included. The number of genera and approximate number of species in each order and family are given, and insome cases these taxa are mentioned. There are, of course, the inevitable minor errors: e.g., there is only one species of Zanclus, not two; Terapon is misspelled Therapon: and a few of more consequence, e.g., the retention of the families Rosauridae and Kasidorodae. For a volume of its size and complexity of nomenclature, there are re- markably few typographical errors and the references to literature through 1973 are quite complete. In- evitably this reference will be compared to the recent book by Lindberg which, unfortunately, in the English translation has the same title. The two books complement one another: Nelson’s classification is more modern, has some distribution maps, and costs less than Lindberg’s, which has better illustrations, a more extensive list of references and, of course, keys to families. By and large Nelson has achieved what he set out to do, and despite differences of opinion one may have with the classification he uses or the philosophy by which he arrived at it, this is a most useful text. I recommend it highly to all serious students of fishes. C. GRUCHY National Museum of Natural Sciences, Ottawa, Ontario KIA OM8 1979 BOOK REVIEWS 207 Nesting Ecology of Canada Geese in the Hudson Bay Lowlands of Ontario: evolution and population regulation By Dennis G. Raveling and Harry G. Lumsden. 1977. Ontario Fish and Wildlife Research Report Number 98. Ontario Government Bookstore, Toronto. v + 77 pp. $3.50. The report is divided into two major sections, nesting ecology and population limitations. Objec- tives for the study of the nesting ecology of Canada Geese of the Mississippi Valley Population were to examine apparent optimum and accomplished repro- ductive rates, habitat preferences, and density and spacing of nests in relation to the regulation of population size. The primary objective for the second portion of this report was to examine nesting ecology to provide insight into the factors limiting goose numbers. These objectives have certainly been ful- filled. The study has direct management implications and extends our understanding of the evolution of waterfowl adaptations to their environment. The 159-square-mile study area located west of James Bay was examined intensively, by walking and helicopter searches, in 1967-1969 for nesting geese. The researchers were fortunate in having a “late” year, an “early” year, and an “average” year, making for some interesting year-to-year comparisons. Among the many factors measured that relate to nesting ecology were habitat preferences, nest site location, nest phenology, nest density and spacing, clutch size, nesting success, predator influences, fertility, and goose behavior. An especially interesting facet of the study involved the development of a model of body weight dynamics of adult females in spring that calculated the caloric cost of migration, maintenance, Birds in Boreal Canada By Anthony J. Erskine. 1977. Canadian Wildlife Service, Report Series 41. Supply and Services Canada, Ottawa. 71 pp., illus. $5 in Canada; $6 elsewhere. This book presents “an overall review of the boreal avifauna, its composition, evolution, and prospects for survival.” For eight summers, Erskine made quantitative studies of boreal birds, from New Brunswick to British Columbia, utilizing 40-hectare (16-acre) plots, each of which he visited three or more times. Merriam’s concept of a boreal life zone is seen to be a misleading oversimplification. Erskine shows how bird species composition varies greatly between black spruce, tamarack, balsam fir, and aspen. Further subcategories such as ‘shrubs after burns,’ ‘farms,’ and ‘balsam poplar of flood-plain forest,’ allow even better understanding of bird distribution and abundance. egg laying, and incubation and predicted weights and control of clutch size. Energy stores accumulated during spring migration were shown to be essential to flight to nesting range, waiting for nest sites to become snow free, egg formation, and maintenance through incubation. Clutches were largest in an “early” year and in early nests within a year. The mechanism of cessation of egg laying appears to be the exhaustion of accumulated reserves to a “winter” weight level. Winter mortality was clearly the major limiting factor for Mississippi Valley Population geese, and this mortality was overwhelmingly accounted for by hunting. The report contains 54 tables and 34 figures, all of which are neat and clear. Several black-and-white aerial photographs illustrate the various habitat types in the study area. The text is lucid and very readable. In fact, the only negative aspect of the entire report is its slightly awkward size, being 8 X 11".”' rather than 8", X 11”. Anyone interested in avian nesting ecology and in the importance of well-designed and thorough field studies will find this report to be an excellent source. NOEL J. CUTRIGHT Wisconsin Electric Power Co., 231 W. Michigan, Milwaukee WI 53201 'Book Review Editors Note: 20 cm X 30cm as result of Ontario Government Metrification program. Although the American Robin and the White- throated Sparrow occupy no less than ten of these habitat niches, the Black-and-White Warbler is listed for only one (poplar-birch forest). The Tennessee Warbler is listed for broad-leaved understory of spruce, for jack pine, for alder understory of tamarack bog forest, for young aspen, and openings (western), but is rare in pure spruce. In stands where balsam fir predominates, one may expect to hear the Winter Wren, Blackburnian and Black-throated Green War- blers, with Bay-breasted Warblers in the more mature stands, and Magnolia Warblers in the younger growth, while the Ruffed Grouse and Blue Jay replace the Spruce Grouse and Gray Jay. One should look for Nashville Warblers in tamarack bog forest. Inevitably others will disagree with Erskine on some minor points. I would not agree that the Red- 208 tailed Hawk is more characteristic of conifer than of broad-leaved forests. I would add Sharp-tailed Grouse, Black-billed Magpie, and House Sparrow to the list of birds characteristic of northern farms, the Great Crested Flycatcher, Cedar Waxwing, and Connecticut Warbler to the list for aspen forest, especially in parkland areas, and the Vesper Sparrow to the list for more southerly jack pine forests. I do not agree that Least Sandpipers breed in boreal fens at this latitude. Erskine contends that birds of the boreal forest have attracted less study than those of any other part of Canada. Nevertheless, his is a personal project that fails to review many published studies from the THE CANADIAN FIELD-NATURALIST Vol. 93 region; Flack’s review of aspen parkland birds is mentioned but some of Flack’s important conclusions are ignored. This is a landmark publication. It heralds a new recognition by the Canadian Wildlife Service of its responsibility for “dickey birds” and their ecology. It contains a wealth of ecologic and distributional data, brilliantly analyzed and synthesized. It demands careful and repeated perusal by everyone interested in boreal birds. C. STUART HOUSTON 863 University Drive, Saskatoon, Saskatchewan S7N 0J8 Lexique anglais-francais, Termes techniques 4 I Usage des Biologistes By Jean Vaillancourt. 1978. Editions de ’Université d’Ot- tawa, Ottawa. 427 pp. Paperback $12.00. Biologists, like the animals they study, must look beyond linguistic and national boundaries, to be aware of developments in their field. This lexicon, of a high degree of completeness, will help the biologist reap the full benefits of discoveries in two of the world’s most important languages. The lexicon consists of two principal parts. The first and largest is an English-French lexicon with the entries listed alphabetically in English and numbered consecutively. The emphasis is thus on providing the equivalent word in the other language rather than a definition. The second section is an alphabetical index of French terms keyed by number to the first section. This approach permitted condensation of the text and reduction in cost without loss in utility. The English-French lexicon is laid out in the following manner: the English word in boldface type, which permits the word to be easily picked out, followed by one or more classificatory abbreviations such as Anat., Biol., Zool., Océanogr., Méd., Bot., Ichtyol., Ornith., Micr., etc.; the French synonym or synonyms; the plural form (only when irregular); and the gender. The only other even roughly comparable sources the reviewer is aware of are the following: Glossaire de biologie animale by Roger Husson (1970, Gauthier- Villars, Paris), which is unilingual and gives about 2500 definitions; the Dictionnaire frangais-anglais, anglais-frangais des termes médicaux et biologiques by Pierre Lépine and Philip R. Peacock (1974, Flammarion, Paris), having some 6500 words; and the Dictionary of biology by Gunther Haensch and Giselin Haberkamp de Anton (1976, Elsevier Sci- entific Publishing Company, Amsterdam), with syn- onyms in four languages for about 9800 terms. (There are also a number of unilingual English biological dictionaries.) None of these give comparable cover- age. To check the thoroughness of coverage the first 25 biological terms found in the preceding diction- aries were checked to see whether they were included in Vaillancourt’s lexicon. Of the 25 terms in all three, about 90% were found in Vaillancourt. The reviewer has a rather specialized manuscript dictionary of ichthyology. Even in this case about half these specialized terms were found in Vaillancourt. The coverage 1s clearly very good. Over 10 400 terms are included. The disciplines covered include bacteriology, bot- any, zoology, marine biology, anatomy, systematics, evolution, ecology, limnology, oceanography, genet- ics, paleontology, pharmacology, and physiology as well as related terms in geology and geography. In a discipline as broad as biology it would be impractical to include every term. Only a few omissions were noted — neotype (néotype), meristic (méristique), and ray in the sense of fin ray (rayon), although ray in the sense of a kind of fish was included. Ichthyologist and fish(eries) biologist are equated, although in general use the first is usually restricted to the theoretical student of systematics, anatomy, evolution, ecology, and zoogeography of fishes, as opposed to the applied management of fishes; however, these are minor points. I wholeheartedly recommend the addition of this book to your biological library. There have been many times that I wished I had a comparable authoritative source book. Vaillancourt is to be congratulated for his breadth of coverage which will make it invaluable for the student and professional. Don E. MCALLISTER National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8 IO79 BOOK REVIEWS 209 An Introduction to the Aquatic Insects of North America Edited by R.W. Merritt and K.W. Cummins. 1978. Kendal/ Hunt, Dubuque, Iowa. 441 pp., illus. US $18.95. Nearly twenty years have passed since a new comprehensive reference work on aquatic insects has been published. Although the publication of this book dealing with aquatic entomology will not render any of the existing texts obsolete, it is a long awaited up- to-date synthesis of the taxonomic status of the various groups of aquatic insects. The greatest improvement over existing books of this kind is the shift in emphasis from the traditionally taxonomic theme to one that includes extensive ecological information. The taxonomic treatment of the insect orders presented in the text has been restricted for the most part to the familial level. Only a few of the smaller orders such as the Megaloptera and the Neuroptera, and a few families of aquatic Diptera, notably the Tipulidae,Culicidae, and Simuliidae, are dealt with at the generic level in the keys. The opinion of the editors was that a single reference work treating all major life stages to the generic level, as well as including the ecological data, would be of prohibitive size. Pre- sumably the inclusion of the extensive ecological information for the orders has been done at the expense of the taxonomic sections to provide a more encompassing review. This decision by the editors is commendable for it has resulted in a wide range of information available in a single text. In view of the frequent changes that occur at the generic and specific levels, the restriction of the taxonomic information to the family level by the editors should also result in a book less subject to the need for major revisions. In addition each section contains a listing of the major references to more detailed taxonomic studies. Thus the lack of generic keys has little impact on the usefulness of the book. Although the taxonomic sections generally deal with the organisms at the family level, most of the genera are mentioned in the ecological treatment of each order. The ecological data has been compiled into tables and the editors have maintained a consistency throughout the book while including as much ecological information as possible in a readily obtainable form. This is especially significant. con- sidering the diversity of contributors involved in the preparation of the text. The reader need only find the genus of interest and read across the page to determine its habitat, behavioral characteristics, trophic rela- tionships, and distribution in North America. As well, references to specific ecological studies are also included if more information is required. Additional topics pertaining to aquatic entomology serve as introductory to the systematic and ecological sections on the insect orders. The section dealing with sampling methods and equipment deserves special mention. The full range of aquatic habitat types is presented in tabular form with a listing of the sampling methods and equipment considered most suitable for each habitat type. The references to pertinent literature that are contained in this section appear to have been carefully selected to include both proponents and critics of the various methods outlined in the tables. The most commonly used equipment is well illustrated by diagrams, and a list of suppliers, complete with addresses is provided in the text. Canadian readers, however, will be disappointed that this list contains only suppliers in the United States and does not mention Canadian distributors. The chapter dealing with the general morphology of the aquatic insects is, like the rest of the book, well illustrated by diagrams. Unfortunately, only the stoneflies are illustrated in the immature stage, which in a book geared towards students, provides little guidance for those attempting to work with highly specialized forms suchas the larvae of aquatic diptera. A generalized treatment of phylogenetic relation- ships and evolutionary adaptations of aquatic insects completes the non-systematic sections of the book. The book appears to be aimed towards the amateur collector and the beginning student of aquatic entomology, but its use is by no means restricted to this group. Although the generalized treatment of the topics contained in the text clearly demonstrates this bias, the work is also an excellent source book that the professional can draw from. The editors have defi- nitely succeeded in producing a work that will be of use to all levels of interest and expertise in aquatic entomology for many years. REIN JAAGUMAGI Department of Entomology and Invertebrate Zoology, Royal Ontario Museum, Toronto, Ontario MS5S 2C6 210 Nichoirs d’ oiseaux Par Raymond Cayouette, illustrations de Jean-Luc Grondin. 1978. La Société zoologique de Québec, Charlesbourg, Québec. 36 pp. $4. Cette plaquette brochée de 36 pages a des dimensions qui sortent de l’ordinaire (25,5 x 20,0 cm), mais qui contribuent a mettre en relief la valeur générale de Tlouvrage. La composition et la présentation graphiques sont trés agréables et de bon gout. Le papier est de bonne qualité et limpression est soignée. La reproduction des planches en couleurs compte parmi les meilleures que j’aie eu Yoccasion d’examiner au cours des derni€res années et rend justice au talent de l’artiste. L’ouvrage comprend cinq parties principales qui varient en importance. L’introduction et les généralités occupent les huit premieres pages et on trouve en neuviéme page une liste des espéces qui utilisent des nichoirs. Les auteurs traitent en détail de vingt espéces d’oiseaux nicheurs dans les vingt pages suivantes, et des modéles de nichoirs accompagnés de dimensions apparaissent a la fin de ’ouvrage. Ce qui frappe surtout dans cette plaquette c’est la finesse des illustrations, c’est du Jean-Luc Grondin a son meilleur. Grace a ce travail Grondin s’affirme parmi les meilleurs artistes-naturalistes du pays. Son style est sir et il maitrise trés bien sa technique. I] concilie a la fois le souci du détail et un grand sens artistique et démontre qu'il est aussia l’'aise avec les illustrations en couleurs qu’avec le dessin au trait qu'il a si heureusement utilisé antérieurement. Le texte Raymond Cayouette est clair, précis et bien rédigé. On y trouve une quantité de détails intéressants et généralement précis sur la nidification des oiseaux en général et sur les 20 espéces qui sont traitées dans le texte. Ces derniéres sont accompagnées dillustrations. En page 5, l’auteur mentionne que l’Hirondelle pourprée “a diminué de Birds in Peril: a guide to the endangered birds By John P. S. Mackenzie. 1977. Pagurian Press, Toronto. 191 pp., illus. Hardback $14.95, paper $7.95. I have often felt that an author who attempted to write a topical book on an environmental issue did so at considerable risk, for new information Is constantly being revealed that dates the author’s effort or contradicts his work. Endangered species has especially been the topic of considerable verbiage, much of it ill-informed. Despite these obstacles, John Mackenzie has made a worthwhile effort in bringing to the public’s attention the plight of some endangered or threatened birds. THE CANADIAN FIELD-NATURALIST Vol. 93 facon évidente la quantité de moustiques” dans une région des Etats-Unis. Or il s’est avéré, a la suite d’études récentes, que lHirondelle pourprée con- somme peu de moustiques! On a quelquefois con- fondu “couvée” avec “ponte.” Ponte désigne toujours l'ensemble des oeufs pondus par un oliseau. J’ai constamment noté que les longueurs des oiseaux different appréciablement, dans certains cas, des longueurs reconnues pour ces espeéces. Les périodes de couvaison données sous le vocable “Durée de l’incubation” ne sont pas toujours en accord avec celles reconnues dans des études de ces espéces. De méme le réle des adultes dans la couvaison, tel qu’indiqué dans le texte, n’est pas toujours conforme aux données fournies ailleurs! Les aires de nidification au Québec sont générales, mais précises sauf dans le cas de ’Etourneau sansonnet que lon fait nicher a Fort-Chimo! Les dimensions recommandées des_ nichoirs apparaissent dans un tableau ingénieux en page 33 et dérogent peu des dimensions recommandées par plusieurs auteurs. Les modeéles de nichoirs suggérés en pages 34, 35 et 36 ne sont pas innovateurs, mais suffisent pour qui veut se donner la peine de les construire pour le plus grand bénéfice des oiseaux qui les utiliseront. En résumé, cette petite brochure, malgré quelques imprécisions de texte, est un excellent achat, méme a $4.00, a cause de la beauté des illustrations et des renseignements qu’elle renferme. C’est sans réserve que j’en recommande lachat a tous ceux qui s’intéressent aux Oiseaux et qui veulent leur offrir des sites de nidification supplémentaires. HENRI OQUELLET Musée national des Sciences naturelles, Musées nationaux du Canada, Ottawa, Ontario. KIA 0M8 of Canada and the United States The book relates the history of 20 species or sub- species of birds in North America, how they became threatened, and what steps are being taken to help prevent their extinction. Refreshingly, stories have been included to illustrate that intelligent wildlife research and management can help bring a species back to a reasonable population. Terry Shortt produced twenty color paintings for the book. They have been delicately reproduced and capture the spirit of his art. Additional line drawings are throughout. As usual, the work of Terry Shortt is superb. Having just visited an exhibit of his work that 1979 included the original color paintings for the book, I recommend the book for Shortt’s paintings alone. The introductory pages are clearly but simply presented with no new insights or information offered. A discussion on the complex factors that usually work together to cause a species to become threatened would have been helpful. For example, should threatened wildlife be managed on a species, sub-species, or population level? Or, should an animal be considered endangered when it is merely occurring on the fringe of its range? The initial discussion was inadequate. Each discussion is broken into sub-headings: history, decline, life cycle, aspects of biology and behavior, what is being done and the future. This is a good feature, for it provides a wide range of information that can easily be found for each species. Much more could have been added and the reader should not view the text as definitive accounts. Occasionally the writing is unclear. On page 12, for example, the comment, “commercial interests such as mining, oil, gas . . . support environmental studies” “financed a multitude of environmental studies on all aspects.” True, corporations have undertaken many studies but we cannot lightly dismiss corporate BOTANY Manual of the Vascular Plants of Wyoming By Robert D. Dorn. 1977. Garland Publishing Inc., New York and London. 1498 pp. (2 volumes). US $95.00. These two volumes, if they had been typeset rather than typewritten double-spaced, and often with large blank spaces on the pages, could have been condensed into one easily handled volume. The author has deliberately not included infra- specific taxa, some annual grainfield weeds, species growing only under irrigation conditions suchas lawn weeds, and species to be expected but not yet found in the state. One cannot argue about a decision to produce a conservative treatment, and as well not to include extraneous taxa. It does seem unfortunate, however, that a large part of the introduced flora has been ignored. This is particularly so, because students who will test the keys will probably do so on plants gathered from about townsites and college campuses, and thus be frustrated because the correct answers are not always forthcoming. The order of the flora is strictly alphabetical within the three large groupings of Pteridophytes, Gymno- sperms, and Angiosperms. Descriptions are relatively short: habitat information is adequate; distributions BOOK REVIEWS PAU responsibility for on page 22, the author mentions the hazards of oil tanker traffic, which will be greatly increased when the pipeline is complete. As far as lam concerned, the jury is still out on both government and corporate responsibility towards the environ- ment. Wildlife is just not perceived by decision- makers as a competing resource. The inclusion of success stories in the management of endangered species is a welcome addition. Dis- cussions on the Trumpeter Swan, Hudson Godwit and others is helpful. I found too much emphasis placed on the work being done by the United States Fish and Wildlife Service, although this may reflect the most ambitious program. I had the feeling at times that the “what is being done” section was overkilled and tended to be repetitive. One can quarrel about many “nits” especially by a reviewer who works in the wildlife management field, but on balance, the book is well put together and provides a good source of information for the layreader. GERALD B. MCKEATING 14 DeSavery Crescent, Toronto, Ontario M4S 2L2 are given only for the state of Wyoming and sometimes only by the name of a county; synonomy is minimal; the few line drawings scattered throughout are by Jane L. Dorn. An appendix near the end of the second volume includes a page on geography and climate, a map of the state depicting counties, sections on historical vegetation, present vegetation, natural vegetation types, selected references, rare and endangered spe- cies, early collectors in Wyoming, and a summary giving a breakdown by family of the 605 genera and 2144 species treated. This might perhaps better have been placed with the introductory material in the first volume. The high cost, coupled with the bulkiness of this work, will undoubtedly mean that few students, for whom it was written, will put it on their bookshelves. WILLIAM J. CODY Research Institute, Agriculture Canada. KIA 0C6 Biosystematics Ottawa, Ontario D2 THE CANADIAN FIELD-NATURALIST Vol. 93 A Provisional Checklist of Species for Flora North America (Revised) Edited by S.G. Shetler and L. E. Skog. 1978. Missouri Botanical Garden Monographs in Systematic Botany Volume |: 1-199. Paper US $6.50. This revised checklist which includes 16 274 species, 2350 genera and 233 families is a tangible result of the efforts of many individuals, and particularly of Shetler, towards what was eventually to have pro- duced a Flora of North America. It 1s indeed a step towards such a flora, and is a document from which a general review and revision of the flora can begin. The area covered is that part of North America north of the Mexican border, including Greenland. Included are names of taxa, authorities, coded information of four categories of plant character- istics, range given by 16 regions, and the source of the information. There is no synonymy. The computer printout has been photoreduced in order to incor- porate more information on each page, and this has made legibility somewhat more difficult. WILLIAM J. CODY Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 Garden Spice and Wild Pot-herbs. An American Herbal By W.C. Muenscher and M.A. Rice. 1978. Cornell Uni- versity Press, Ithaca, New York. 213 pp., illus. US $16.95. This book contains accounts, botanical, culinary, and horticultural, of approximately 70 plants, pri- marily cultivated plants, used as herbs and spices. An illustration of each plant is provided, showing the stage at which the particular plant should be harvested. The illustrations themselves have been prepared from the original wood-cuts by Elfriede Abbe. Included in the botanical aspects of the book are short accounts of the plant families represented as well as brief descriptions of each of the species included. Scientific names are given, with some frequently used synonyms being listed in the appen- dix, as well as common English, French, and Italian names. The culinary notes are rather general but include information on the usual uses of various portions of the plants. I suspect that cooking has a “common-name problem” similar to that which led to the establishment of scientific nomenclature: horse- radish sauce is described as “an alcoholic infusion” to my great surprise. I can only suspect that Muenscher and I know different sauces by the same name. Medicinal and nutritional notes are also included, the former I suspect being primarily of the folk- medicine genre. It is unfortunate that the glossary is restricted to botanical terms, for many of those used in the medicinal portion are probably unfamiliar to the majority of readers. What does a carminative drink do? For those with gardens, but particularly those with gardens in southern Canada, the horticultural in- formation will be of interest. In this regard, the hardiness table, showing survival rates at Ithaca, New York, for the winter of 1942-43, offers more concrete information than is usually available. My own reaction to the type-face used, Goudy Kennedy Bold and Italic, was negative: it seemed hard on the eyes. On the other hand I think it did help to induce a relaxed, browsing mood and this is, | would say, a book for browsers rather than readers. It contains a lot of fascinating information and I anticipate dipping into it fairly frequently both when | am planning a meal and when asked a question by a home gardener interested in growing herbs and spices. It is a reprint of the original 1955 edition. MARY E. BARKWORTH Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 How to Identify Grasses and Grasslike Plants (Sedges and Rushes) By H. D. Harrington. 1977. Swallow Press Inc., Chicago, Illinois. 142 pp., illus. US $3.95. Grasses, sedges, and rushes are often not even considered “proper” flowering plants by amateurs and some beginning botanists. This erroneous impression is reinforced by non-technical identification manuals which totally ignore the three families. The absence of an attractive corolla can apparently only be accepted in trees, which, of course, do not have ffowers as in “How to identify the flowers of ABC.” Undoubtedly, learning to identify and eventually to recognize the grasses, sedges, and rushes is not 179 straightforward. The structures involved are some- what different and generally rather small. Neverthe- less so much of our vascular plant flora belongs to these three groups that the botanist, whether amateur or professional, who ignores them or lumps them into a few genera ignores a substantial proportion of the plants in the area. Harrington’s book is not an identification manual. It is, however, an illustrated dictionary and advice manual that anyone tackling the three families for the first time will find an invaluable reference. Terms are not only explained in words but are also illustrated with line drawings. Moreover beginners will be parti- cularly pleased to find that words such as ‘anther’ are illustrated as well as ‘aphyllopodic.’ Most of the book is concerned with grasses (six Flora of Alberta — a checklist By D. F. Brunton. 1977. Alberta Department of Recrea- tion, Parks and Wildlife, Edmonton. 43 pp. Free. This small booklet represents a checklist of the vascular plants of Alberta that has been compiled by the author from E.H. Moss’s (1959) Flora of Alberta, J. C. Packer and M. G. Dumais’ (1972) “Additions to the Flora of Alberta” (Canadian Field-Naturalist 86: 269-274), and J. C. Packer’s (1974) A Supplement to E. H. Moss’s Flora of Alberta. The listed species’ names and very abbreviated annotations concerning their generalized distributions in Alberta (e.g., SW & N) or rarity status (e.g., R) have been abstracted entirely from these earlier sources. Thus, this booklet does not represent either a nomenclatorial updating or a reconsideration of the taxa composing the Alberta flora. For most purposes, readers would seem better advised to use the original sources, plus B. Boivin’s (1968-72) Flora of the Prairie Provinces. For more updated information on the status of Alberta’s rare plants, readers should see G. W. Argus and D. J. White’s (1977) “The Rare Vascular Plants of Alberta” (Syllogeus, Number 17). Despite their obvious limitations, I believe such provincial, state or regional checklists of the flora may serve useful purposes. Although not true in this case, Freshwater Wetlands: ecological processes and Edited by Ralph E. Good, Dennis F. Wigham, Robert L. Simpson, and Crawford G. Jackson, Jr. 1978. Aca- demic Press, New York. 378 pp., illus. This book developed from a conference entitled “Freshwater Marshes: Present Status, Future Needs.” BOOK REVIEWS 713) chapters) with one chapter each for sedges and rushes. In addition, there is a chapter on collecting and pressing and another listing identification manuals that were available at the time of writing. Harrington is consistently encouraging, straightforward, and realistic. He states quite clearly that identification of the three families is not easy but not impossible for the non-professional taxonomist. In reading the book it is quite evident that he has helped numerous students in the past. He will continue to do so. MARY E. BARKWORTH Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 checklists often provide the only coverage of the total known flora of a region and represent an important preliminary stage towards the ultimate production of a floral manual with keys and descriptions. A checklist provides a brief, easily consulted outline of the flora and can be conveniently used to check off the plants encountered in more local species’ inventories. Space for just such a purpose has been provided in the present booklet. It may prove to be most useful for environmental consultants who are involved with baseline inventory studies of particular areas for impact assessments preceding planned developments. Personally, I find such a checklist to be a convenient source for quickly checking the spellings and author- ships of species’ scientific names. I believe that the practicality of the present checklist of the flora of Alberta could have been improved by the inclusion of at least the more important synonyms, especially - those needed to correlate the species’ names used in the floral manuals of Moss, Boivin, and others. V. L. HARMS Fraser Herbarium, University of Saskatchewan, Saskatoon STN 0WO management potential The objective of both the book and the conference was to consider all ecological aspects of wetland eco- systems in addition to the production ecology of marshes. This objective, of course, entailed an expansion of the object of study from marshes to the 214 total spectrum of “wetlands.” Wetlands are generally accepted to be the broadest classification category comprising all lands inundated by standing water fora sufficient portion of growing seasons to establish a hydrophytic vegetation, or lands with a high water table and an accumulation (generally accepted to be greater than 30cm) of raw organic matter which essentially provides the rooting medium for vege- tation. This development from conference to book eluci- dates the genesis of the book’s title Freshwater Wetlands but it in no way facilitates this reviewer’s acceptance of the term. In the study of marshes, freshwater and saline marshes are convenient and practical subdivisions of the field of study. In the study of wetlands, however, the acceptance of a category of freshwater wetlands involves a flagrant discrimination against a small group of saline wet- lands. If generally adopted, this concept could inhibit the investigation of the practical criteria which may be used to distinguish the freshwater and saline sub- divisions since workers would be reluctant to ap- proach the ‘no man’s land’ between these two, assumed, compartments. An erroneous conclusion has been drawn from the developmental sequence which proceeded from a study of marshes to a study of the total wetlands spectrum. This conclusion is — given the direction of this sequence — marsh must be the ‘whole’ into which all the pieces are to be fitted. Hence we find such anomalies as “bog marshes” (which are fens!), “marsh exploitation by peat mining,” “meadow” (blanket bog). An analysis of the chapters of this book which deal _ specifically with wetland ecosystems indicates that there has been only a modest attainment of the goal of expanding the scope of the field of study. About 75% of the content of these chapters is devoted to a consideration of marshes, 15% considers open fens, 5% considers a combination of open fens and meadows, and 5% considers the overall category of wetlands. Hence, the consideration of marshes pre- ponderates, and swamps, open and treed bogs, and treed fens, all substantial portions of the wetlands spectrum, have not been treated in this volume. Because primary production processes form the first section of this book it may be assumed that the editors assign first priority to this aspect of wetland ecosystems. On lands which provide essential agri- cultural and forest crops, emphasis on production 1s warranted and the tedious, time-intensive activity of quantitative production evaluations is a worthy endeavor. On wetlands, however, the emphasis on THE CANADIAN FIELD-NATURALIST Vole93 production processes is not justified. If the case for the conservation of wetlands is to be convincingly presented to legislators and to the public, conserva- tionists and scientists should give highest priority to understanding the role of wetlands in serving as groundwater recharge areas, as stream-flow regu- lators (roles which, at best, achieve no more than brief references in this book), in serving as wildlife habitats and as an important component of an aesthetically- pleasing, varied landscape. Accepting this book’s emphasis on marshes, I must observe I have not experienced any competition in my annual harvests of Typha latifolia pollen and a significant harvest of marsh emergents for cattle fodder is unlikely since it would seriously conflict with other marsh uses and because, as indicated in this book, the bulk of the mineral nutrients of marshes are contained in the substrata. I shall temper my predominately negative re- marks with a few concluding, positive observations. The Summary and Recommendations sections, which conclude each of the four parts of the book, provide good summary statements on the stage of develop- ment of the art and the high priority projects for future work on the four ecosystem aspects — primary production processes, development processes, nutri- ent dynamics, and management potential. The contribution of Richardson et al. (“Nutrient Dynamics of Northern Wetland Ecosystems”) utili- zes a rational wetlands classification scheme so that some of the important nutrient and environmental criteria which characterize the different wetland types begin to emerge. The contribution of Sloey et al. (“Management of Wetlands for Nutrient Assimilation”) is an excellent dissertation on a topic which ts likely soon to be an environmental concern. His cautionary statement is worth quoting for The Canadian Field- Naturalist readers: “In the past we caused deterioration of the quality of our surface waters by using them to treat our wastes. Lest we make the same mistakes in handling our valuable and diminishing wetlands, it is mandatory that we carry out long-term, carefully- monitored experiments at a severely-limited number of sites. It is important that those conducting these experiments document changes in the natural system that could signal future problems.” A. N. BOISSONNEAU Ontario Ministry of Natural Resources, Ontario Centre for Remote Sensing, 880 Bay Street, Toronto, Ontario MS5S 1Z8 1979 ENVIRONMENT Boreal Ecology By William O. Pruitt. 1978. Studies in Biology No. 91. Arnold (Canadian distributor MacMillan, Toronto). 73 pp., illus. Paper $4.80; cloth $10.20. Boreal ecology is a book of definition and hence one of introduction. Like most other books in this series it is designed to keep teachers and students alike informed of recent developments in biology. It is a book about snow, and to this end Pruitt maintains that the . . . “ecological effects of snow cover are so multi-faceted and all prevading that one might accurately state that boreal ecology is the study of the ecology of snow.” The book deals in generalities and briefly covers the following aspects of boreal ecology: radiant energy and light; water, soil and permafrost; bioclimate: characteristics of boreal vegetation; boreal animals, complete with a discussion of some of their adapta- tions to the boreal environment, ecosystems, and food webs: human utilization of boreal regions: research methods and procedures peculiar to boreal regions. Behind Pruitt’s exacting and scientific description of the boreal forest regions, there is a message. In essence, it centers upon his belief that continued misuse of this fragile environment means inevitable BOOK REVIEWS D5) degradation and eventual destruction. Disruption of the faunal contingent of the boreal forest is well illustrated in the case of the Alaska Tundra Vole (Microtus oeconomus). Populations of this vole have extended their range from normal riverine locations into the crevices formed from tractor trains, which through repeated use have exposed the perma- frost and subjected it to severe melting. Says Pruitt, “|. . one can only speculate on the possible genetic results of having a population attenuated to one or two Microtus wide and perhaps thousands of Micro- tus long.” It seems to me that we should begin to stop, look, and listen before unknowingly stepping into trouble. The text is well organized, easy to follow and supported with maps, diagrams, labelled photographs and tables. The reference list is extensive enough to serve as the beginning point of a literature search. It is an excellent piece of work and I recommend it to interested beginners and professionals. PAUL A. GRAY Ontario Ministry of Natural Resources, Box 89, Cochenour, Ontario POV ILO The Environmental Impact of Outdoor Recreation By G. Wall and C. Wright. 1977. Department of Geog- raphy Publication Series No. 11, University of Waterloo, Waterloo. x + 69 pp., illus. Paper $5. This book evolved out of *...a need to synthesize the research on environmental impact of recreation, to draw planning and management implications from the research, and to identify areas of further research.” The text is organized into nine sections: the introduction, which outlines the problem, the study, and the methodology in environmental impact re- search: geology; soil; vegetation; water quality: wildlife: air; implications and conclusions, complete with planning and site management recommenda- tions; and references. The impact of outdoor recreation on geology and air, determined by the authors not to be a major problem, is given little space in the analysis. Soil is considered at length; however, most of the literature comes from the United States and England, and consequently, in the Canadian context much of the research has little meaning. Compaction, organic matter, soil nutrients, temperature, run-off and drainage changes, erosion, and soil organisms are considered. The authors suggest that more research is required on temperature, soil organisms, bacteria,’ and nutrients. Vegetation, which is usually first to be affected by physical disturbance, is reportedly well covered in the literature. It is reviewed in terms of change in percentage cover and species diversity, plant growth and age structure, and habitat diversity and mechan- ical damage. The authors believe that more work is required because the research to date has“... yielded conflicting results, especially in terms of compaction, soil moisture, nutrient levels and soil organisms, and their associated affects on vegetation species and growth rates.” A considerable amount of research into water quality has been conducted in Canada, especially in Ontario. This work has been completed primarily out of concern for the detrimental effects that cottaging, camping, boating, fishing and other related activities 216 might have on the aquatic ecosystems. Wall and Wright concentrate their discussion on pathogens, nutrients and plant growth, dissolved oxygen, and pollutants. They suggest that research must continue because “... there is a lack of quantitative analyses of change in water quality brought about by different types and intensities of recreational use.” The wildlife section consists of a discussion on disturbance, loss and gain of habitats, alteration of population, and changes in species composition. In my opinion the authors have unsuccessfully at- tempted to summarize the impact of recreation on wildlife in flowchart fashion. They list the immediate results of intrusion into a wildlife habitat as killing, disturbance and alteration, which results in adapta- tion, or migration, and a change in reproduction levels. The final result is a change in the population and a different species composition. It goes without saying that upon intrusion into a wildlife habitat the first three events occur; however, I believe that the primary effect of an intrusion is an alteration of habitat, which in turn causes disturbance L’inventaire du Capital Nature: méthode de territoire (3éme approximation) By M. Jurdant, J. L. Bélair, V. Gerardin, and J. P. Ducruc. 1977. Péches et Environnement Canada, Québec. Ap- provisionnements et services Canada, Ottawa. 202 pp., illus. $7.00, $8.40 (other countries) French. The purpose of this volume, aimed at those individuals involved in land management, is to introduce a method for land classification and ecological survey. It has taken ten years of field verification and re-evaluation to perfect this method and present it in a workable document. Initially the development of the method was started in 1967 asa pilot project in the Saguenay/Lac-St.-Jean region of Quebec; its workability was verified when Jurdant’s team undertook the ecological survey of 350 000 km? in the James Bay area. Chapter | outlines the goal and objectives of the ecological inventory. The goal is to produce carto- graphic documents that are characterized by the most permanent bio-physical parameters of the environ- ment and hence provide an ecological basis for planning the integrated management of the resources of a given territory. They achieve this goal by producing an ecological map that summarizes the bio- physical parameters investigated. In Chapters 2 and 3, the authors discuss the basic concepts and criteria used in ecological classification. THE CANADIAN FIELD-NATURALIST Vol. 93 resulting in productivity alteration, mortality, emi- gration or immigration, or adaptation. A change in the population and the species composition then occurs. The book is reasonably well organized and comes complete with graphs, flowcharts, and a reference list. The composition is weak in a few areas and the authors have a tendency toward wordiness; for example, “Even without the intervention of man, the environment would not be unchanging, but would be in a perpetual state of flux.” They might have said, ‘Even without the intervention of man, the environ- ment is naturally dynamic.’ I recommend this book to those unfamiliar with this subject and interested in developing a basic under- standing of environmental impact. PAUL A. GRAY Ontario Ministry of Natural Resources, Box 89, Cochenour, Ontario POV ILO classification et de cartographie écologique du The basic taxonomic unit for the inventory is the ecological phase which is defined by the combination of certain vegetation and soil characteristics (this uses the concept that vegetation and soil are the best integrators of variation in biosystems and geosystems respectively). For the method to work the classifica- tion criteria must indicate the intrinsic properties and the actual state of the ecosystem. The criteria must be measurable and should contribute in some way to vegetation evolution. Twenty-one criteria are listed for both terrestrial and aquatic ecosystems. Chapters 4 to 9 describe in very precise terms the levels of ecological detail that are observable at different cartographic scales, the methods of classifi- cation and cartography, and the methods for analyz- ing vegetation. An example of what and how to measure, how to record data at all ecological reference sites, and how the cartography is interpreted and applied for management purposes is given. The maps and annexes demonstrate the workability of the ecological survey method. As I have indicated to many of my colleagues, this book will be an invaluable asset to ecologists, planners, and managers at all levels of government, and to private industries and especially to those involved with the preparation of environmental impact statements. 1979 The book is replete with original drawings and caricatures by L. Renaud, that add to the readability of the prose. In addition, the authors have included several flow charts depicting the basic concepts of the ecological survey and outline how a multidisciplinary team should be organized to carry out a successful BOOK REVIEWS 2A) ecological inventory. GILLES ROBITAILLE Environment Canada, Laurentian Forest Research Centre, Box 3800, Ste-Foy, Québec GIV 4C7 Applications de la Télédétection a Etude de la Biosphére By C. M. and M. C. Girard. 1975. Masson, Paris. 186 pp., illus. Paper in French. 96F. Remote sensing techniques (both instrumentation and application) have developed very rapidly since 1970. This text, published in 1975, is very scientific in its approach, is extremely well documented, is clear and to the point, is logically structured, and is very particular to details in methodology and definition. The work examines a wide spectrum of applications and the in-depth analysis of examples will certainly be appreciated by the readers. The Girards have or- ganized this work into four parts, each complete in itself. In part one, ‘Remote Sensing,’ the authors present a discussion of the principles involved. They describe the characteristics and the energy sources of the electromagnetic spectrum, the influences of the atmosphere, and explain the mechanisms involved for the peculiar behavior of vegetation, soil, water, ice and snow in different spectral ranges such as the visible and near infrared, middle and far infrared, and the very high frequencies. Both photo and non- photographic instrumentation is described. Data treatment analysis, i.e., visualization, increasing con- trast, and information selection, as well as factors that influence the utilization of data are given. Part Two is entitled ‘Methods’ with the emphasis being placed on methods of photo and image interpretation. Sixty percent of the book (Part Three) is devoted to ‘Interpretation.’ It is a detailed account applied to vegetation, pedology, agronomy, and to anthropo- genic influences. Text examples are often related to actual photographs. The interpretation of vegetation is looked at from the point of view of species identification, recognition of vegetation assemblages, edaphic conditions, and succession of plant com- munities. The pedology section looks at the ident- ification of soil characters that are both readily and not readily discernable. Examples are also given. The interpretation of agricultural lands is also treated. The authors give information on crop identification, soil modification, and touch on the problem of agronomic cartography. They also expand on the utilization of remote sensing for pest control. Anthropogenic influences on the environment are increasing. The artificial interferences by man that lead to urban- ization are not treated. What the authors do look at is the interpretation of influences on the natural environment and rural modifications. In Part Four an example of how high-altitude data (35 km high from balloons) are utilized in inter- pretation and thematic cartography (..e., geology, soil utilization) is given. This part concludes with a flow chart on data treatment by computers. During the three years since publication, many innovations have occurred in the field of remote sensing. This text remains a valuable source of material for those interested in acquiring a basic applications reference. It is recommended for use at the university level and by current practitioners. GILLES ROBITAILLE Environment Canada, Laurentian Forest Research Centre, Box 3800, Ste-Foy, Québec GIV 4C7 Wildness Is All Around Us: notes of an urban naturalist By Eugene Kinkead. 1978. Dutton, (Canadian distributor Clarke Irwin, Toronto). 178 pp., illus. $12.50. Wildness Is All Around Us is an informative and entertaining book that offers a variety of insights into some items of natural history, plus many entrancing digressions. For example, the essay on the coyote (pronounced ky oat) describes not only its dis- tribution, paleontological and recent history, appear- ance, behavior, relationship with man, and repro- duction with other canids, but also includes lengthy asides on endangered species (of which the coyote 1s certainly not one), on sheep and sheep-ranching, on 218 wolves, and on the principles of heredity. But there is almost nothing about coyotes in cities, despite the subtitle of the book. Kinkead also includes a long essay on the House Sparrow, a fascinating history of this species’ intro- duction to North America (Manhattan) in and after 1848 to reduce the measuring-worm population, (even though anti-sparrow organizations existed in England as early as 1744), and its subsequent conquest by 1880 of much of the continent. Until that time these widely-acclaimed birds were shipped freely about the country to eradicate noxious insect pests. After that time, however, the sparrow was seen rather as “an evil of monstrous proportions.” Sparrows molested over 70 species of native birds; ate fruits, vegetables, grain crops, and urban plantings; and around dwellings were dirty, noisy, and blatantly promiscuous. The New York Times predicted in 1887 that once Americans realized how tasty sparrows roasted with bacon were, their numbers would be decimated by cooks, but this did not happen. Instead sparrow populations continued to expand until about 1910, when the increase in the automobile and consequent decrease of horses and the grain they ate undercut the sparrows’ food supply, especially in winter and in cities. Since then sparrow populations, although still numerous, have stayed within more reasonable bounds. The other chapters in this book are really vignettes rather than essays. The first describes the pande- monium which took place one spring when a Steller’s OTHER THE CANADIAN FIELD-NATURALIST Woln9s Eider from Siberia was spotted off the Boston South Shore. The area was soon swamped with birders wanting to increase their Life List, alerted by a message taped for the “Voice of Audubon” Boston telephone number. The second is about a rare birch tree, Betula uber, rediscovered with the help of the author after 60 years beside a creek in the Appala- chians of Virginia. The geological history and the wildflowers of Central Park in New York City are the subjects of the other two chapters. Although Eugene Kinkead is primarily a writer rather than a biologist, he includes in this book a wealth of biological information. He is not correct, however, in saying that no person in North America has been attacked by a healthy wolf. In northern Ontario in 1942 a wolf knocked a railman off his handear, and, while canoeing in 1893 through the barren lands of Canada, J.B. Tyrrell’s party was beseiged by wolves. Similar attacks have occurred in Eurasia. I think also few would agree with the dust- cover statement that the fox is threatened with extinction. This well-written book with its few, evocative sketches uses a format which could profitably be a model for many would-be authors writing about nature. | recommend it highly to both laypeople and professional biologists. ANNE INNIS DAGG Box 747, Waterloo, Ontario N2J 4C2 Field Photography: beginning and advanced techniques By Alfred A. Blaker. 1976. Freeman, San Francisco. (Canadian distributor Oxford University Press, Toronto). 450 pp., 55 black and white plates, 12 color plates, plus 41 pp. field book. $19.95. This book is a very comprehensive treatise on the subject of field photography. It follows the same format and style as the author’s previous publication, Photography for Scientific Publication: a handbook (Freeman, San Francisco, 1965), which presented a synthesis of laboratory photographic technique. The purpose of this publication is instructional; the author states in the preface that, “this book will represent a self-teaching aid to help interested persons improve the quality of photographs they may need.” The text accomplishes this purpose. The book is organized into three sections: Back- ground, Basic Photography, and Field Photography Techniques. There are also three appendices contain- ing film speed data as well as development and copying information. A supplemental pamphlet.sum- marizes mathematical tables and formulae in a form suitable for field use. The background section introduces the reader to scientific field photography. It discusses merits of photography, photographic training, and offers ad- vice on photographic operations. This information is valuable to persons currently involved in professional photographic capacities. The author continues this section with a discussion on the tools of photography. He accurately and extensively describes cameras, film, 1979 development materials, and essential accessories used in modern photography. This discussion has two minor, unavoidable problems: personal bias and some outdated information. The second part of the text, basic photography, is most informative to the incipient nature photog- rapher. An initial short chapter describes considera- tions related to composition of photographs, oras the author puts it, “seeing a picture.” This section is very informative, and is highly recommended for both inexperienced and veteran photographers. Blaker next presents explanations concerning basic topics of photography such as exposure, photo- graphic filters, and darkroom procedures. The data given here are detailed and it is recommended that the reader be concerned only with problems of his/her particular interest. Although the writing style here is clear enough, there is some difficulty in grasping the extensive reference material in one quick reading. If a simplistic discussion of these fundamental topics is required one should seek elsewhere. The final portion of the text deals specifically with field techniques. Blaker commences this section with an appeal to field workers to maintain ethical ecological considerations while endeavoring to cap- ture a picture. This is sound advice that all nature photographers should attend. A short section on climatic problems in photography and proper equip- ment preparation for field work, rounds out this interesting division of text. The remaining chapters of this section contain information on essential topics of nature photog- raphy: close-up, long-range (telephoto), and wide- angle photography. The author begins with close-up (macro) imaging. He explains equipment, back- ground theory (of focal length changes, etc.) and comments on special problem areas. This information NEW TITLES Zoology Avian breeding cycles. 1977. By R. K. Murton and N. J. Westwood. Clarendon (Oxford University Press, New York). xiv + 594 pp., illus. US$48. The behavior of fish and other aquatic animals. 1978. Edited by D. I. Mostofsky. Academic, New York. xiv + 394 pp., illus. US$27.50. *Behavioural ecology: an evolutionary approach. 1978. Edited by John R. Krebs and Nicholas B. Davies. Black (US and Canadian distributor Sinaver, Sunderland, Maryland). 512 pp., illus. Cloth US$34; paper US$17.50. BOOK REVIEWS AIS) is very useful, especially the data on close-up photography with flash equipment. The chapter on telephoto work is unfortunately not as helpful. Some advice here is questionable; for example, the point is made that teleconverters give as good (or better) results for a portion of the cost of telephoto lenses. A pertinent point overlooked is that the image one obtains 1s directly related to the quality of the lens with which it 1s captured. Blaker does not explain that teleconverters of sufficient optical quality are not inexpensive. A telephoto lens with its added versatility may on occasion be a better choice of equipment. The next chapter gives a brief description of wide- angle photography. Although little emphasis is placed on landscape photography, the remainder of the chapter is complete. The last two chapters deal with stereo photography, which is not an essential element in most field work. But this section presents interesting information that may invoke novel methodology in future field photography. Field Photography 1s a book that offers something for every nature photographer. The more experienced will want this text for use of its inexhaustive reference data and extensive bibliography. Beginners will find the material on basic photographic procedures very worthwhile, and advice on problem areas invaluable. All will appreciate its wide scope, logical presentation, and picture quality. A word of caution: this publica- tion is designed as a reference text and is written accordingly. ALAN J. KENNEDY 11416-50 Avenue, Edmonton, Alberta T6H 0J3 Bird flight. 1978. By Georg Ruppell. Translated from the German edition (Munich, 1975). Van Nostrand Reinhold, New York. 192 pp., illus. US$18.95. Birds as builders. 1977. By P. Goodfellow. Arco, New York. 168 pp., illus. US$15.95. *Birds of man’s world. 1978. By D. Goodwin. Cornell University Press, Ithaca. vi + 183 pp., illus. US$10.45. A birdwatcher’s adventures in tropical America. 1977. By A. F. Skutch. University of Texas Press, Austin. xii + 327 pp., illus. US$13.95. 220 *A bird watcher’s guide to the eastern United States. 1978. By Alice M. Geffen. Barron’s Education Series (Canadian distributor Burns and MacEachern, Toronto). 384 pp., illus. US$15.95. t+Ducks, geese and swans of the world. 1978. By Paul A. Johnsgard. University of Nebraska Press, Lincoln. xxl + 404 pp., illus. US$35. East African mammals. An atlas of evolution in Africa. Volume 3, part A (Carnivores). 1977. By J. Kingdon. Academic, New York. vill + 476 pp., illus. US$74.25. Eels: a natural and unnatural history. 1978. By C. Moriarty. Universe, New York. 192 pp., illus. US$1S. *The Gannet. 1978. By Bryan Nelson. Butes, Vermillion, South Dakota. 336 pp., illus. US$25. Granivorous birds in ecosystems. 1977. Edited by J. Pinowski and S. C. Kendeigh. Cambridge University Press, New York. 451 pp., illus. US$47.50. A guide to fishes of the temperate Atlantic coast. 1977. By M. J. Ursin. Dutton, New York. xx + 268 pp., illus. Cloth US$10.95; paper US$5.95. Insect life. 1977. By M. Tweedie. Collins, London. 192 pp.. illus. US$8.95. {Migratory game bird hunters and hunting in Canada. 1978. Edited by H. Boyd and G. H. Finney. Canadian Wildlife Service Report Series Number 43. Supply and Services Canada, Hull. 125 pp., illus. $7.50 in Canada; $9 elsewhere. Octopus. Physiology and behaviour of an advanced inverte- brate. 1978. By M. J. Wells. Chapman and Hall, London and Halsted (Wiley), New York. xiv + 418 pp., illus. US$42.50. +The order Microsauria. 1978. By R.L. Carroll and P. Gaskill. Memoirs Volume 126. American Philosophical Society, Philadelphia. 211 pp., illus. Paper US$15. Parental behavior in birds. 1977. Edited by R. Silver. Dowden, Hutchinsonand Ross, Stroudsburg, Pennsylvania. xvill + 436 pp., illus. US$24.50. Quantitative ethology. 1978. Edited by P. W. Colgan. Wiley- Interscience, New York. xvi + 364 pp., illus. US$25. Readings in sociobiology. 1978. Edited by T. H. Clutton- Brock and P. H. Harvey. Freeman, San Francisco. x + 394 pp., illus. Cloth US$18; paper US$9. Songbirds of the eastern and central states. 1977. By T. L. Rising. Tundra (Scribner’s), Plattsburgh, New York. viii + 48 pp. US$3.95. Studer’s popular ornithology: the birds of North America. 1977. Edited by J. H. Studer. Original publication 1881. THE CANADIAN FIELD-NATURALIST Vol. 93 Harrison House/ Barre (Crown), New York. x + 301 pp., illus. US$39.95. Sulidae. Gannets and boobies. 1978. By J.B. Nelson. Aberdeen University Study Series, Number 154. Oxford University Press, New York. xii + 1012 pp., illus. US$98. Transactions of the XIII International Congress of Game Biologists. 1977. Proceedings of a conference in Atlanta, Georgia 11-15 March, 1977. Wildlife Society, Washington. 538 pp. Paper US$10. *Vanishing birds. Their natural history and conservation. 1978. By Tim Halliday. Holt, Reinhart, and Winston, Toronto. US$16.95. Ways of wildlife. 1977. Edited by E. Horwitz. Wildlife Society, Washington. 157 pp. Paper US$2.95. Botany Dangerous plants. 1977. By John Tampion. David and Charles (Canadian distributor Douglas, David and Charles, Vancouver). 176 pp., illus. $12.50. Darwin and his flowers: the key to natural selection. 1977. By M. Allan. Taplinger, New York. 318 pp., illus. US$14.50. Fenland. Its ancient past and uncertain future. 1978. By H. Godwin. Cambridge University Press, New York. 203 pp.., illus. US$18.95. The flora of Canada: part 1 — general survey; part 2 — Pteridophyta, Gymnospermae, Monocotyledoneae; part 3 — Dicotyledoneae (Saururaceae to Violaceae). 1978. By H. J. Scoggan. Publications in Botany Numbers 7-1, 7-2, and 7-3. National Museums of Canada, Ottawa. 1115 pp. $89 for 3-volume series in Canada: $106.80 elsewhere. An illustrated history of herbals. 1977. By F. J. Anderson. Columbia University Press, New York. xvi + 270 pp., illus. US$16.95. Important forest trees of the United States. 1978. By U.S. Forestry Service. Public Documents Distribution Center, Pueblo, Colorado. 70 pp., illus. US$2.10 plus 25% foreign’ handling. Introduction to world vegetation. 1978. By A. S. Collinson. Allen and Unwin, Winchester, Massachusetts. 192 pp., illus. Paper US$8.75; cloth US$16.95. Manual of woody landscape plants: their identification, ornamental characteristics, culture, propagation and uses. 1978. Stripes, Champaign, Illinois. 536 pp., illus. $15. The moss flora of Britain and Ireland. 1978. By A. J. E. Smith. Cambridge University Press, New York. C 900 pp.., illus. US$59.50. 1979 Orchid biology: reviews and perspectives, I. 1977. Edited by J. Arditti. Comstock (Cornell University Press), Ithaca. 312 pp., illus. US$29.50. +Physiology and biochemistry of seeds in relation to germination. Volume 1, development, germination and growth. 1978. By J. D. Bewley and M. Black. Springer- Verlag, New York. xi + 306 pp., illus. US$45. River plants. 1978. By S. M. Haslam. Illustrated by P. M. Wolseley. Cambridge University Press, New York. C 350 pp., illus. Cloth US$62.50; paper US$14.95. {Tropical trees and forests. An architectural analysis. 1978. By F. Hallé, R. A. A. Oldeman, and P. B. Tomlinson. Springer-Verlag, New York. xvi + 441 pp., illus. US$62.50. Tropical trees as living systems. 1978. Edited by P. B. Tomlinson and M. H. Zimmerman. Cambridge University Press, New York. C 624 pp., illus. US$49.95. *Vascular plant families. 1977. By James Payne Smith, Jr. Mad River Press, Eureka, California. 320 pp. US$7.85. + Wild coffee and tea substitutes of Canada. 1978. By Adam Szezawinksi and Nancy Turner. Edible Wild Plants of Canada, Number 2. National Museum of Natural Sciences, Ottawa. I11 pp., illus. $6.95. Environment Ecology and environmental planning. 1978. By J. M. and M. A. Edington. Chapman and Hall, London and Halsted (Wiley), New York. viii + 246 pp., illus. US$22.50. Ecology field glossary: a naturalist’s vocabulary. 1977. By W. H. Lewis. Greenwood, Westport, Connecticut. x11 + 152 pp.. illus. US$1S. *The ecology of North America. 1978. By V. E. Shelford. First paperback edition (original publication 1963). Uni- versity of Illinois Press, Urbana. 610 pp., illus. No price given. Enjoying nature with your family. 1977. By M. Chinery. Crown, New York. 192 pp., illus. US$12.95. Environmental assessment law in Canada. 1978. By Paul D. Edmund. Edmund-Montgomery, Toronto. xx + 380 pp. $30. tEnvironmental chemistry and cycling processes. 1978. Edited by D. C. Adriano and |. L. Brisbin, Jr. Proceedings of a symposium, Augusta, Georgia, April 28-May 1, 1976. U.S. Department of Energy Symposium Series Number 45. National Technical Information Service, Springfield, Vir- ginia. 943 pp., illus. Paper US$I5. Environmental issues. 1978. Edited by M. W. Holdgate and G. F. White. SCOPE 10. Wiley, Toronto. 242 pp. Paper $23. BOOK REVIEWS Up) Environment and society: an introductory analysis. 1977. By B. Harvey and J.D. Hallett. MIT Press, Cambridge. x + 163 pp., illus. Paper US$6.95. The limits of altruism: an ecologist’s view of survival. 1977. By Garrett Hardin. Indiana University Press, Bloomington. vi + 154 pp., illus. US$10. Nature/science annual, 1978 edition. 1977. By the editors of Time-Life Books. Time-Life, Alexandria, Virginia. 192 pp., illus. US$7.95. Nature’s economy: the roots of ecology. 1977. By D. Worster. Sierra Club, San Francisco. xii + 404 pp. US$15. Pacific seashores: a guide to intertidal ecology. 1978. By Thomas Carefoot. University of Washington Press, Seattle. 208 pp., illus. Paper US$12.95. Principles of pollination ecology. 1978. By K. Faegriand L. van der Pil. Third revised edition. Pergamon, New York. 242 pp., illus. Cloth US$25; paper US$15. Pyramids of life: illuminations of nature’s fearful symmetry. 1977. by J. Reader and H. Croze. Lippincott, Philadelphia. 222 pp., illus. US$12.95. Risk assessment of environmental hazard. 1978. By R. W. Kates. SCOPE 8. Wiley, Toronto. 160 pp. Paper $13.35. A Sierra Club naturalist’s guide to the deserts of the southwest. 1977. By P. and L. Larson. Sierra Club, San Francisco (Canadian distributor Wiley, Toronto). 288 pp.., illus. Cloth US$9.59; paper US$5.95 (Cdn. $7.95 paper). The thin edge: coast and man in crisis. 1978. By A. W. Simon. Harper and Row, New York. xii + 180 pp. US$10. The underwater wilderness: life round the great reefs. 1977. By C. Roessler. Chanticleer (Dutton), New York. 319 pp., illus. US$35. *Vermilion Lakes Banff National Park: an introductory © study. 1978. By Bow Valley Naturalists, Banff, Alberta. iv + 68 pp., illus. Paper $3. Miscellaneous Astronomy now. 1978. By J.M. Pasarhoff. Saunders, Philadelphia. xvi + 400 pp., illus. Paper US$1I1.95. Astronomy, the cosmic journey. 1978. By W.K. Hart- mann. Wadsworth, Belmont, California. x11 + 536 pp., illus. US$16.95. Bibliography of published and unpublished literature on the Hudson Bay Lowland. 1978. By S. E. Haworth, D. W. Cowell, and R. A. Sims. Report O-X-273. Great Lakes Forest Research Centre, Sault Ste. Marie. 270 pp. Free. Bio-babel. Can we survive the new biology? 1978. Edited by Allen R. Utke. John Knox Press, Atlanta. 248 pp. US$11.95. 22D Biométrie et écologie. 1978. By J.M. Legay and R. Tomassone. Societé frangaise de Biométrie, Jouy-en-Josas, France. xi + 388 pp., illus. US$8. The faces of the Great Lakes. 1977. By J. Ela. Sierra Club, San Francisco. 192 pp., illus. US$24.50. Hunger on planet earth. 1977. By J. Archer. Crowell, New York. vill + 216 pp., illus. US$7.95. Land/water classification. 1978. By D.M. Welch. Eco- logical Land Classification Series Number 5. Supply and Services Canada, Hull. x +54 pp., illus. Paper $4.50 in Canada; $5.40 elsewhere. + Last of the naturalists: the career of C. Hart Merriam. 1977. By K.B. Stirling. Revised edition. Natural Sciences in America Collection. Arno, New York. 472 pp., illus. US$23. *Lexique anglais-francais, termes techniques a lusage des biologistes. 1978. By Jean Vaillancourt. Editions de ?Uni- versité d’Ottawa, Ottawa. 427 pp. $12. THE CANADIAN FIELD-NATURALIST Vol. 93 Nature to be commanded. Earth-science maps applied to land and water management. 1978. By U.S. Geological Survey. Public Documents Distribution Centre, Pueblo, Colorado. 96 pp., illus. US$6.25 plus 25% foreign handling. Simulation modelling of environmental problems. 1978. Edited by F. N. Frenkiel, D. W. Taylor, and D. W. Goodall. SCOPE 9. Wiley, Toronto. 128 pp. Paper $13.35. Sun/earth. How to use solar and climatic energies. 1978. By Richard C. Crowther. Scribner, New York. vil + 232 pp., illus. Paper US$8.95. The world of Roger Tory Peterson: an authorized biog- raphy. 1977. By J. C. Devlin and G. Naismith. Illustrations and paintings by Roger Tory Peterson. Quadrangle (New York Times), New York. xxii + 266 pp., illus. US$14.95. +Available for review *Assigned for review The Ottawa Field-Naturalists’ Club Minutes of the Ninety-Ninth Annual Business Meeting of The Ottawa Field-Naturalists’ Club The 99th Annual Business Meeting of The Ottawa Field-Naturalists’ Club was held in the auditorium of the National Museum of Natural Sciences, on 10 January 1978. The President, R. A. Foxall, called the meeting to order at 8:17 p.m., with a quorum of 26 persons present (final total, 27). The Recording Secretary read the minutes of the 98th Annual Meeting, which were approved on motion (by K. Strang, 2nd, H. Thomson). R. Foxall referred to business arising from the minutes. The Council had directed the Publications Committee to consider the status of The Shrike as a Club publication; the Committee has recommended that The Shrike be considered a publication of The Ottawa Field-Naturalists’ Club, a recommendation accepted by the Council. As recommended at the last Annual Meeting, the Conservation Committee had considered the matter of the necessity for approval by Council of all reports and briefs; and had reeommend- ed that all significant submissions, containing policy statements or contentious issues, should be cleared by the President or his designate. A General Meeting of the Club, on Centennial planning, had been held in June 1977, to inform the membership of the projects being undertaken, and to get ideas of the interests of the members. In the absence of the Treasurer, the Club’s accountant, M. Brigham, was called upon to present the 1977 financial report. Unfortunately, this had not been audited, owing in part to the current prevalence of the ‘flu’. R. Taylor, the Chairman of the Finance Committee, explained that the change of the financial year, to end at the end of September, was to relieve the pressure on the Treasurer’s Assistant and the Auditors to produce the statement in time for the Annual Business Meeting; closing the books in December had always been difficult, particularly with membership dues still coming in at that time. For this reason, this year’s statement covers only nine months. Taylor also explained that the apparent large profit to the Club was due to the lack of allocation of funds to The Canadian Field- Naturalist. Excluding ‘windfall’ items such as life memberships, which cannot be budgeted, the Club had actually operated at a slight loss, justifying the increase in membership dues for 1978. The Canadian Field- Naturalist profit was due mainly to the success of the raptor issue; the grant request to the National Research Council for 1978 has been reduced accordingly. R. Foxall raised the point that the Auditors, approved at the last Annual Meeting, were G. J. Wasteneys and D. Potter; since that time, D. Potter has changed his job and will not have time to do the audit before the end of February. Wasteneys is still available, but the Constitution requires two auditors. C. Gruchy moved (2nd, E. Dickson) that the responsibility for finding a second auditor for the financial year ending September 30, 1977 be delegated by the Annual Business Meeting to the President of The Ottawa Field-Naturalists’ Club; this was approv- ed unanimously. Gruchy then questioned the need for the Club to have two auditors: M. Brigham agreed that one person could do the job. It was recommended that this matter should be examined by the Council to see whether a change to the Constitution may be in order. R. Taylor moved (2nd, C. Gruchy) adoption of the interim, unaudited, financial statement covering | January 1977 to 30 September 1977: the motion was approved. The Annual Report of Council, to be published in The Canadian Field-Naturalist, was read by R. Foxall. A. Erskine pointed out that there were deficiencies in, rather than lack of, knowledge of our area’s natural history: and Todd suggested that the Pink Road feeder should in some way be identified as the Lucerne feeder mentioned in the previous report. Acceptance of the report, as amended, was moved by S. Hamill, 2nd, C. Gilliatt, and approved. M. Brigham moved a vote of thanks (2nd, J. Harrison) to all the volunteers who look after the Club’s bird feeders: George McGee and Bill Holland, Mr. and Mrs. Hugh Munro and W. Earl Godfrey, Jean Hastie, and John Dubois. Motion approved unanimously. Foxall then called on C. Gruchy, as Chairman of | the Nominating Committee, to present the slate. Gruchy apologized for not having published a request for nominations, but said that despite this, one nomination had been received from the membership at large. He then presented the slate: President: R. A. Foxall; Vice-President: R. Taylor; Treasurer: B. Henson; Recording Secretary: D. Laubitz; Corresponding Secretary: S. Armstrong: additional members of Council: E. Beaubien, C. Beddoe, W. Cody, J. Diceman, E. Dickson, A. Dugal, C. Gilliatt, C. Gruchy, P. Hall, J. Harrison, V. Hume, H. MacKenzie, J. Murray, M. Ney, G. Patenaude, K. Strang, S. Teeple, E. Todd. On motion (by C. Gruchy, 2nd, M. Ney) all of those nominated were declared elected. R. Foxall, on behalf of the Club, gave thanks to those Council members who were retiring: Tony Erskine, Gavin Nicholson, Gerry Oyen, Pamela Sims, and Stan van Zyll de Jong. 223 224 The Auditors for 1977-1978, G. J. Wasteneys and M. Brigham, were approved on motion(by R. Taylor, 2nd, E. Todd). Foxall then reported that plans for some of the Centennial projects are proceeding well, but that despite many appeals there have been very few volunteers, and some of the planned activities may have to be dropped. It is felt that the Club deserves a good Centennial celebration, but perhaps our ideas have been too ambitious. We will also need many volunteers to do small jobs during 1978. E. Dickson asked whether the list of people who have offered their skills to help the Club had been consulted by the Centennial Planning Committee; D. Laubitz answer- ed that it had been consulted when specific skills had been needed. M. Brigham raised the point that some of the proposed Centennial projects concern saleable items: that the cost of these items may be about equal to the amount for which they can be sold; and queried whether consideration had been given to the need for advertising to promote sales. R. Foxall replied that THE CANADIAN FIELD-NATURALIST Vol. 93 the costing of items for sale is indeed a difficult task: and that advertising was being considered an impor- tant adjunct to selling Centennial items. He emphasiz- ed that 1978 will be a very busy year for the Council and the Club. J. Harrison moved (2nd, A. Hanes and E. Dickson) a motion of thanks to Harry Thomson for his long and dedicated service to the Club, as Business Manager of Trail & Landscape, from which position he is now retiring after I] years. The motion was warmly approved by all present. E. Todd thanked the President, on behalf of the Club, for carrying on the business of the Club during 1977, a job which Todd knew by experience to be a frequently thankless one. Adjournment of the meeting (on motion by B. Henson, 2nd, K. Strang) was at 9:35 p.m. Following the refreshments, color slides were shown by J. Harrison, M. Ney, and D. Black. D. R. LAuBITZ, Recording Secretary Report of Council of The Ottawa Field-Naturalists’ Club The consideration and support of Centennial projects has taken up a large proportion of Council’s time during 1978. One result of the Centennial program is that we have been reminded that our members may have in their possession valuable archival materials referring to The Ottawa Field- Naturalists’ Club. Club members are therefore re- minded that any relevant materials for which they no longer have any use will be gladly received by the Public Archives. For the Centennial Steering Committee,1978 was a period of major activity, leading to Council approval of 14 of the 33 Centennial projects that had been proposed. The Club pin, designed by W. Rath, will go on sale at the Centennial Banquet, to be held on May 19. An exhibition, ‘100 years of looking at nature,’ will open on May 18 as the preliminary to a weekend of seminars, workshops, and field trips on natural history topics. In September, we shall combine a day of field trips with a Club picnic. Hasty-notes, designed by E. Dickson, and the Centennial Calendar, featur- ing works by Club artists, have already been produc- ed; being prepared for publication are a reissue of John Macoun’s Autobiography; ‘Orchids in the Ottawa District’ by A. and J. Reddoch; Cumulative Indices to Transactions of The Ottawa Field-Natural- ists’ Club and The Ottawa Naturalist compiled by J. Gillett; and a revised ‘Checklist of Birds of the Ottawa Area’ by B. Barrett. A stereo recording of * Nature Sounds in the Ottawa Area’ by F. M. Brigham is close to completion. Two projects still in the early stages of preparation are ‘Birds in the Ottawa Area 1979, a record of bird sightings and abundance in Centennial year to be published in 1980; and the Macoun Club’s natural history ‘Trail and Guide, which was held up owing to a forced change of location. (Hue MacKenzie) The Centennial Ways & Means Committee was set up to assess Centennial projects’ costs; to determine the availability of The Ottawa Field-Naturalists’ Club funds to meet these costs: and to recommend ways of raising additional funds should they be required. Total project costs are now estimated to be approxim- ately $40,500, with recovery through sales of about $18,400, giving an estimated net outlay of $22,100. Moneys available through Club funds are about $50,000: no action to obtain additional funds from outside the Club was considered necessary. (C. Gilhatt) Finance Committee: The 1977-1978 budget was adopted by Council early in the year. With the financial year ending in September, the books were presented to the auditors on time, and the 1978-1979 budget was presented to and approved by Council at its December meeting. Ivy Hewis resigned as Assistant 1979 to the Treasurer early in 1978: the services of Lois Cody were engaged for this position temporarily, and the position was advertised to the Club membership. Of the three applicants, Mrs. Cody was judged to be the most suitable, and Council approved her appoint- ment on a continuing basis. The committee recom- . mends that the constitutional requirement of two auditors be changed to one; it also recommends that one person be appointed to be responsible for the inventory of sales of all items excluding back issues of Club journals. (R. Taylor) The Membership Committee reported that the total Club membership now stands at 1150 compared with 1163 in 1977; the membership structure is given in the table below. The Club lost a well known naturalist and Honorary Member in January 1978, when Hoyes Lloyd died. At the Annual Dinner in April, W. K. W. Baldwin was presented with an Honorary Member- ship. Marsh Ney, Chairman of the committee, resigned in August when he moved to Victoria, B.C. (F. E. Goodspeed) The Publications Committee reported that, since the last report, The Canadian Field-Naturalist was published in four issues, Volume 91(4) and Volume 92(1, 2, 3), with a total of 430 pages containing 37 articles, 46 notes, 37 book reviews, and a listing of 266 new natural history book titles. The publication was supported in part by a grant of $2200 from the National Research Council, which is very gratefully acknowledged. Trail & Landscape was published in five issues for a total of 152 pages of articles and information of interest to local members. Marc Forget, the Business Manager, has had to resign; we thank him for his contribution to Trail & Landscape. The Shrike, our local bird-watchers newsletter, was published in eight issues containing 55 pages. The committee recommended amendments to By-laws 6 and 9; submitted a grant application for $8000 to the Natural Sciences and Engineering Research Council to assist the publication of the indices to the THE OTTAWA FIELD-NATURALISTS’ CLUB 225 Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist. An analysis of the authorship of articles published in The Canadian Field-Naturalist was carried out at the request of Council. Based on a sample of issues published between 1950 and 1977, our study indicated that there had been a steady decline in the number of papers published by authors not identified with an institu- tional address. We emphasize The Canadian Field- Naturalist’s status as a major national, Canadian publication open to all workers in the field of natural history; advice and guidance are available to any author requiring assistance in documenting research results for publication. (J. K. Strang) During 1978 the Excursions and Lectures Commit- tee organized 10 monthly meetings, with an average attendance of 30, in the auditorium of the National Museum of Natural Sciences; three photographic workshops; the Annual Dinner, with Clarence Tillen- lus, artist and conservationist, as speaker: and excursions covering general topics (14), birds (17), and flowers, butterflies, geology, fish, mushrooms, and microscopic life (one of each). The bus excursions are still popular and will be continued: and the use of younger members as assistant field trip leaders has been successful both as help to leaders and for training future leaders. Jeff Harrison, Chairman of the committee, resigned in May when he moved to Newfoundland. (Jeanette Dean) The Conservation Committee spent most of its time in 1978 studying and responding to Conservation Lands in Ottawa-Carleton and the subsequent re- visions to this study. Proposed changes to the original (1974) Ottawa-Carleton Regional Official Plan in- cluded numerous boundary changes and six deletions ~ of Conservation Areas, due in part to pressure from affected landowners. Asa result of three major reports prepared by the committee, some of the revised boundaries were expanded, and two of the deleted Canadian Canadian Membership (local) (other) USA Foreign Totals Individual 454 (438) 321 (342) 102 (106) 3 (5) 880 (891) Family 198 (207) 26 (22) 1 (2) 2 (1) 227 (232) Sustaining 9 (10) nil (3) nil nil 9 (13) Life 13 (9) 7 (6) 3} (1) 2 (2) 25 (18) Honorary 5 (5) 4 (4) nil nil 9 (9) Totals 679 (669) 358 (377) 106 (109) 7 (8) 1150 (1163) Changes +10 -19 -3 —] —13 Figures in brackets represent 1977. 226 areas were proposed for reinstatement. During the year it became apparent that, despite The Ottawa Field-Naturalists’ Club representation on advisory committees and direct communication with planning staff and politicians, without input by individual members of the Club the main viewpoint represented in response to the planning report would be that of those who were vehemently opposed to conservation zoning within the Region, as expressed at public meetings. Letters giving background to the issues were therefore sent to all local members, requesting them to write to their local politicians on the issues; there was a 10% response, which was much appreciated. Eleven field trips, to collect data, were held during the year, and it is clear that continued activity to try to influence planners and politicians will be needed for many months to come. How the committee’s work will affect the status of conservation in the region is not yet known. (Ewen C. D. Todd) The Macoun Field Club Committee reported that there are 65 members in the club in this their 30th year. During 1978 consideration was given to publicizing the club, but it was decided that, rather than advertise, information about the club would be distributed to organizations where it would be readily available to those seeking such a club; local schools were also informed of the club’s activities, and the members of the Senior group offered to speak to schools, and to lead Club activities, etc. A donation received in memory of John Bird, from Mrs. Furness Thompson, of Pennsylvania was used to buy a pair of binoculars for the club. (S. Gray) The Education and Publicity Committee was responsible for the acquisition and installation of a THE CANADIAN FIELD-NATURALIST Vols Club phone in 1978, with the number to be listed in the local directory. Other activities included providing judges for the Ottawa Science Fair, and $100 in prizes; involvement in the mail campaign for conservation support; requests for volunteers for weekday activi- ties: and some advertising of monthly meetings in the local papers. A special field trip and scavenger hunt was organized in September for Club members and their children; it is difficult to tell whether the idea was well received as it was raining and no one came. (Elisabeth Beaubien) The Council has also been concerned with im- proving relationships both inside and outside the Club. There has been considerable discussion of the sort of relationship The Ottawa Field-Naturalists’ Club should have with the Federation of Ontario Naturalists, and it was concluded that The Ottawa Field-Naturalists’ Club should have Federated Club status; the election of Courtney Gilliatt to the Federation of Ontario Naturalists’ Board of Directors was welcomed. Long-term discussions on the desired relationship between The Canadian Field-Naturalist and The Ottawa Field-Naturalists’ Club have led to the recommendation that an ad hoc committee of Council be set up to study the roles of our publications and their relationship to the Club: and to produce a discussion paper on journal policies and editorial guidelines, to be brought before Council. As a result of a desire to strengthen the relationship between Council and the membership, a Council reporter was appointed, whose articles on Council activities now appear regularly in Trail & Landscape. Compiled from committee reports and Council minutes by D. R. LAUBITZ, Recording Secretary Auditor’s Report To: Members of The Ottawa Field-Naturalists’ Club We have examined the balance sheet of The Ottawa Field-Naturalists’ Club as at September 30, 1978 and related statements of Profit and Loss. Our examination included a general review of the accounting procedures and such tests of the records and supporting vouchers as were considered necessary in the circumstances. In our opinion these financial statements present fairly the financial position of the organization as at September 30, 1978 and the result of its operations for the period in accordance with generally accepted accounting principles. December 2, 1978 (Signed) F. M. BRIGHAM G.J. WASTENEYS 1979 THE OTTAWA FIELD-NATURALISTS’ CLUB pi} The Ottawa Field-Naturalists’ Club Balance Sheet as at September 30, 1978 Assets Current Gashmancdhvermude posits she sacks gies we ceases $ 38,240.19 ACCOUMS TECOVADIG hea ook dbaoode ood net owed b ocoroo Ol 2 SDI ACCTUGG WASTES sey eo cree ee Ae eo ee Nach ree Greer an 10,822.26 Rrepaldeexpenses;—_centemmMialll nya... ee oot eis S32 2242339 Sie Investments Canada Savings Bonds due October 31, 1978 ........... 10,700.00 Fixed at cost FSU OITA TO taper Mak ARN cess tae Weeae Guede dad Qlci ead aapntiees a) Sees 52950 Wesseaccummlatedsdepreciation ase a. c-oee eees oe e 435.12 94.38 Total assets 64752012 Liabilities and Surplus Current liabilities AGCOWRIS DEVAS eine cere eh en ee oe ener $ 1,506.54 IDetierral: Tine aeons ee alga ean teenie oe e ee ene 7,675.83 $ 9,182.37 Surplus Balancer @ ctobe wall wlO Te tar ses eieceec te ave cyaneisow stele cies enc 43,934.07 AGGE MCOMS Over Qos 5 socewsssoaouo gubleo do ue he @ttawa Ficld=Naturalists: @lub)..... 4... - $ 4,155.60 hier Canadian-hicldsNatunalist.s esc la. 7,480.08 11,635.68 DATES SSvicnlysas) WIUS soocoesduceesedasccdn so ooos 35,5095) Total lalbiiGes Aine SWRI Se de coeuoceaeoonceoamdsoud: $ 64,752.12 (Signed) Geoffrey Wasteneys, Auditor F. Montgomery Brigham, Auditor B. C. Henson, Treasurer 228 THE CANADIAN FIELD-NATURALIST Vol. 93 The Ottawa Field-Naturalists’ Club Statement of Income and Expenditure for the year ended September 30, 1978 Income Apportionment of membership fees PACTinier call Wrens eae ete ect cee, azar heed Wn cd asa, vad ehh ee $ 6,642.72 [STIS a es ie aa en Oe Ces se oe cess Beye ee ae ars nce oh ens Aa 720.00 3.7). 362072 iinailiscleandscape:—— SUDSCHIPtIONS) rae ess sie ee ee - 212.00 = Back mUmMbens sence as cccrtakee clone 35.00 247.00 ING® NOE SNA BCG Ore oO ODIO CRE COED EER ARC eae 442.78 Salles — decals, GmMleMMS oocccocvcccccuccgoooosouCb OOO 70.90 8,123.40 ISSUES 85 Gece aetin esl outches OIERES eno > US ERTIES RL TENS, oe 2,051.99 $ 10,175.39 Expenditure Trail & Landscape 1? {Lo} WAS RTA Speer So, oer a ete ao RNa ene ee a Ba Deh) Gigculatrone ryan eer ocr niet emis ae tien hea 119.61 Edit peancdrothices cats marches c.g setulae ace ees 54.09 [mI@ NIXON ANE, 5G oliaie Suck aco RR Se Buns Pec choke cheno ee cere nr oe ___ 440.00 3,191.40 Committee activities — net EXCursiOnseanG MleChunes mum ern cene et ie ees oe eee (146.50) NECN DEns Mp rte mmencccnestae conte sieesea ee ea aon se sacasia ae aus Ql 22 Miaic ou G hulp th arta, bears cms eek tener nian ae ae Lee 136.62 G@OnSenvanvOnbete eeepc rae eared ote tet ci eepers RTA ee 175.83 Gente rian alee eae vate eh yc shes Uy ge tht a 27.00 Bind@liced ercwanct ake cattery keke ER art Ree ee 590.11 ORC hVGRS UVC ar cnr wectaecicgan ac eee a eo Ee 40.83 1,781.11 BaldiwimyScholarships, ssc kns cca ee eee ak ek oe scutes 250.00 SPEClARACKIVITIES urs ahs ete nce tcstiarche eRe tan ae oi nests a aictac’ 114.91 GOuMCIIREPENSES pry er cans oe roe eae Oe cc ehererc kote 120.00 OHHICERS UD PICS teres ae ey he oe ae mee cette Mieka Tone 276.37 FLO MOVANT eas ee nee eee ena eG iene ata aes otioaets 200.00 MS Celeane@ USire tts se dete eek Ach ah ee enue hae LI 86.00 1,047.28 6,019.79 Excess of income over expenditure ...................... $ 4,155.60 1979 THE OTTAWA FIELD-NATURALISTS’ CLUB The Ottawa Field-Naturalists’ Club 229 Statement of Income and Expenditure of The Canadian Field-Naturalist for the year ended September 30, 1978 Income Apportionment of The Ottawa Field-Naturalists’ Club Membership Fees PNGNINU gee eet tae re eee eee ac eh SNe MR ose aa vane ohe ls $ 4,428.48 LUG so. gigs oNERSRe ne Een eres Ne te PER cere 480.00 4,908.48 SUMOSCHMIUTOMS teensy ei ccaus ayes tune wks S-oreye eieeaiet © sisieye Seis) werets 14,737.94 ublicationi—sREphINtS vcpusee ee ae 4 fem s oe oe yt «sete 5,761.00 = lalates andutalbssettines quer aryacine sicily 2,025.50 SpE XGA YP ARES sols eieueysueroney=fed-lepsrepaiahsiavora svoie 7,897.00 SB ACK SMUIMIDEES ints cscr-cereveseicics hee sicnstel onererene 1,803.73 Grants — National Research Council of Canada ........ 2,900.00 — Canadian National Sportsmen’s Show ........ 125.00 OMe TION CTES ita. cece sertnele se crete se tee nisin wie ses elsiasies 2,214.11 See MAINS Chaput stectens cccie venole e's aleuctic vena lenase, estates 962.13 Expenditure Peatitbo SAVIN Merete ep cree ce ree cuesin stsnosecl ina tet aye aapcysGeteforeleltoren ot =ne 23,079.80 RGD TANN USHA SP chic arith ayicye {nv yrensyrays lasaysay stray SeeKoer olel Ble lasevalio'ss Se Sse = 3,073.09 GincwlatirOnyseryeetia ates whee cei sualere whale Glew ote siahsusisyore GS se ws diinmeg ade xpeNnSeSiseysews cise aveach che avetebeiemeuey oie eo as choi shes OihicemaSSISCAMt i) sas seers a Sestabdetas a testes tava slisieieleos Sarena tens FAOSUAG Cemeteries ceersecnere | cei a Re Sisie ste etetetlehd. Sieitie iene aVolekesape CHMGS. SUYD/TNSS serene lS a Aue en tenn Chenin gies ete eae Mtoe AOI Nec inl emcee alec suSt wet cus rcita estes le okt cone cue ees sci age foga7eh sce, ueMORs IMDS celle OU Shara ce ctspots cae steyey acess sunt caeeeiarcne cos soe eaetsncueneia es Excess of income over expenditure .................0.65, $ 19,646.42 17,487.23 3,025.00 3,176.24 26,152.89 3,627.09 I TSe23 1,891.50 896.05 570.45 1,502.00 33.60 $ 43,334.89 35,854.81 $ 7,480.08 Instructions to Contributors Content The Canadian Field-Naturalist is a medium for the publication of scientific papers by amateur and professional naturalists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All readers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. As the journal has a flexible publication policy, items not covered in the traditional sections (Articles, Notes, Letters, News and Comment, and Book Reviews) can be given a special place provided they are judged suitable. Readers are encouraged to support regional, provincial, and local natural history publications.as well by submitting to them their reports of more restricted signifi- cance. Manuscripts Please submit, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. Also authors are expected to have complied with all pertinent legislation regarding the study, disturbance, or collection of animals, plants, or minerals. Type the manuscript on standard-size paper, if possible use paper with numbered lines, double-space throughout, leave generous margins to allow for copy marking, and number each page. For Articles, provide a running head, a bibliographic strip, an abstract, and a list of key words. These items are optional for Notes. Generally words should not be abbreviated but use SI symbols for units of measure. Underline only words meant to appear in italics. The names of authors of scientific names should be omitted except in taxonomic manuscripts or other papers involving nomen- clatural problems. Authors are encouraged to use “proper” common names (with initial letters capitalized) as long as each species is identified by its scientific name once. Although we prefer the names of journals in the Literature Cited to be written out in full, these may be abbreviated following the Bibliographic Guide For Editors & Authors, The American Chemical Society, Washington, D.C. (1974). Unpublished reports should not be cited here. Next list the captions for figures (numbered in arabic numerals and typed together on a separate page) and present the tables (each Reviewing Policy of The Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor (who reviews it himself or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, chosen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific titled, numbered consecutively in arabic numerals, and placed on a separate page). Mark in the margin of the text the places for the figures and tables. Extensive tabular or other supplementary material not essential to the text, typed neatly and headed by the title of the paper and the author’s name and address, should be submitted in duplicate on letter-size paper for the Editor to place in the Depository of Unpublished Data, CISTI, National Research Council of Canada, Ottawa, Canada KIA 0S2. A notation in the published text should state that the material is available, at a nominal charge, from the Depository. The Council of Biology Editors Style Manual, 4th edition (1978) available from the American Institute of Biological Sciences, is recommended as a guide to contributors. Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Tllustrations—Photographs should have a glossy finish and show sharp contrasts. Photographic repreduction of line drawings, no larger than a standard page, are preferable to large originals. 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The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. i 7 a TABLE OF CONTENTS (concluded) News and Comment 201 Book Reviews Zoology: Where to find birds in British Columbia — The moths of America north of Mexico, including Greenland: Fascicle 22.2, Noctuoidea (in part): Lymantriidae — Hawks, falcons and falconry — Fishes of the world — Nesting ecology of Canada Geese in the Hudson Bay Lowlands of Ontario: evolution and population regulation — Birds in boreal Canada — Lexique anglais-frangais, termes techniques a usage des biologistes — An introduction to the aquatic insects of North America — Nichoirs d’oiseaux — Birds in peril: a guide to the endangered birds of Canada and the United States No S eS) Botany: Manual of the vascular plants of Wyoming — A provisional checklist of species DAT for Flora North America (revised) — Garden spice and wild pot-herbs: an American herbal — How to identify grasses and grasslike plants (sedges and rushes) — Flora of Alberta: a checklist — Freshwater wetlands: ecological processes and management potential Environment: Boreal ecology — The environmental impact of outdoor recreation — L’inventaire ZAltS du capital nature: méthode de classification et de cartographie écologique du territoire (3e€me approximation) — Applications de la télédétection a l’étude de la biosphere — Wildness is all around us: notes of an urban naturalist Other: Field photography: beginning and advanced techniques 218 New Titles 219 The Ottawa Field-Naturalists’ Club D8 Mailing date of previous issue 5 February 1979 1979 Council — The Ottawa Field-Naturalists’ Club President: Roger Taylor Elisabeth Beaubien Fran Goodspeed i : " pat Charlie Beddoe Peter Hall Vice-President: Courtney Gilliatt aml Rekha Fe Nine conic Treasurer: Barry Henson Frank Bell Frank Pope 2 } : Bill Cody lan Sneddon Recording Secretary: Diana Laubitz Janel Dicemean Ken Strang Corresponding Secretary: Valerie Hume Ellaine Dickson Ken Taylor Loney Dickson Ewen Todd Past President: Roger Foxall Marc Forget Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. For information on Club activities telephone (613) 722-3050. THE CANADIAN FIELD-NATURALIST Volume 93, Number 2 1979 Editorial Our responsibilities as field-naturalists and biologists LORRAINE C. SMITH Articles Natural fires as an index of paleoclimate J. TERASMAE and N. C. WEEKS Virgin Douglas Fir forest on Saturna Island, British Columbia THOMAS P. SULLIVAN Colonial-nesting Herring Gulls and Common Terns in northeastern Saskatchewan HARRY A. STELFOX and GREGG J. BREWSTER Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario P. BARCLAY-ESTRUP and RICHARD A. SIMS Habitat utilization and population densities of the amphibians of northeastern Alberta WAYNE ROBERTS and VICTOR LEWIN Caribou distribution and group composition associated with construction of the Trans-Alaska pipeline RAYMOND D. CAMERON, KENNETH R. WHITTEN, WALTER T. SMITH, and DANIEL D. ROBY Interesting vascular plants from southeastern Yukon Territory GEORGE W. SCOTTER and WILLIAM J. CODY Notes Fall foods of Common Snipe on the Copper River Delta, Alaska STANLEY E. SENNER and P. G. MICKELSON Ivory Gull colonies in southeastern Ellesmere Island, arctic Canada F. FRISCH and W. C. MORGAN Wind-caused death of Great Cormorant MARTIN K. MCNICHOLL and GEOFFREY G. HOGAN Habitat selection by wintering Snowy Owls (Nyctea scandiaca) M. Ross LEIN and G. A. WEBBER Two-lined Salamander, Eurycea bislineata, in Labrador FRANCIS R. COOK and JOHN PRESTON First Canadian record of a Flathead Catfish E. J. CROSSMAN and J. H. LEACH Earthworm cocoons as a drift component in a southern Ontario stream DONALD P. SCHWERT and KENNETH W. DANCE Flowering plant phenology at Sheep Mountain, southwest Yukon Territory MANFRED HOEFS Eggshell thickness in American shorebirds before and since DDT MICHAEL L. MORRISON and LLOYD F. KIFF Response of wintering Moose to mechanical habitat rehabilitation in Alaska M. SIGMAN New localities for the Northern Spring Salamander and the Four-toed Salamander in southwestern Quebec DAVID M. GORDON First record of the Long-tailed Shrew (Sorex dispar) in New Brunswick GORDON L. KIRKLAND, JR., DAVID F. SCHMIDT. and CAROL J. KIRKLAND Intraspecific food theft by the American Kestrel PETER M. FETTEROLF First record of the Northern Brook Lamprey, /chthvomyzon fossor, in the Nelson River drainage, Manitoba J. JYRKKANEN and D. G. WRIGHT Blue Grouse brood hen ~ Black Bear confrontation M. G. SULLIVAN 113 116 126 132, 139 144 155 163 195 198 199 200 concluded on inside back cover ISSN 0008-3550 The CANADIAN FIELD-NATURALIST Published b y THE OTTAWA F IELD-NATURA LISTS’ CLUB, O , Ottawa, Canada The CANADIAN FIELD-NATURALIST ublished by THE Z o) Fe FURALISTS A wa ana Pp E OTTAWA FIELD-NATURA CEL Ott d B awa, C a THE movers SS ee Published by The Ottawa Fjeld-Naturalists’ Club Editor: ARTHUR GIBSON, mntomological Branch, Devs! OLTAWA- ent of A wricullure, Associate Editors: FIARLAN I. SMITH. w. H. PIARRINGTON, Pp. A. TAVERNER, Anthropology: Entomology. Ornithology. Mi. Q. MALTE, ph.D.. M. ¥. WILLIAM, PHD. dus RE- LAMBE, ¥.G.S.. Botany - Geology- Palaeontology’ por, JonNMACOUN, MA. arto KLoTZ, TAs: Cc, GoRDON Hewits, BD Sc., Conchology- Metgorology- Zoalogy- ee CONTENTS: —_—— On Cheneosaurus tolmanensis, @ new genus and species of trackodont dinosaur from. the Edmonton cretaceous of Alberta, BY Lawrence 4 117 sR By John M. Coop 123 « feeding habits ; = M. Sternberg 129 Concer ario Cra} ON Noa ee 130 Bird Notes. By L. Mcl. Verrill Age The padson-Merrill press, Limited. dedicat a Special Tssue qssumD MARCH 9TH, 1 tye : Entered at Ottawa Post Office Se See Ob mailer 2 lO raptors—especially the Pereg 5 E : grine Falcon Volume 90, Number 3 July-September 1976 Centennial Year Volume 93, Number 3 July-September 1979 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Mrs. Edward Schreyer The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things. The Members of Council are listed on the inside back cover. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors @) DBind A. J. Erskine George H. La Roi E. L. Bousfield Charles Jonkel David P. Scott Francis R. Cook Charles J. Krebs Stephen M. Smith W. O. Pruitt, Jr. Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang Production Manager: Pauline A. Smith Business Manager: W. J. Cody Subscriptions and Membership Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The Canadian Field-Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879- 1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 45 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, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, Oregon, USA 97731 (address valid until August 1979). Address manuscripts on birds to the Associate Editor for Ornithology: Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0 All other material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0 Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231- 4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461). Cover: Since The Ottawa Field-Naturalists’ Club was founded in 1879, it has published a scientific journal. Two covers are illustrated. Left, The Ottawa Naturalist, published from 1887 to 1919. Right, The Canadian Field-Naturalist, published from 1919 to the present; the format with a photograph on the cover was introduced in 1970. The Canadian Field-Naturalist Volume 93, Number 3 July-September 1979 To a Bigot For years you have been telling us about the supreme importance of your laboratory studies in biology and we have been convinced that your research is making contributions to scientific knowledge. But we have also listened, sometimes with considerable repressed anger (because we know it is useless trying to argue with you: “Don’t confuse me with facts, my mind is made up!”) as you denigrate any studies dealing with natural history. Unfortunately, despite our vehement, logical protests, you refuse to hear the other side of the story but remain adamant, being dogmatically convinced only of the correctness of your own beliefs and opinions. You think further that investigations on whole animals or plants are merely diversions from what you consider to be worthwhile science. But then you are not alone because there are many others like you. Why am I writing this editorial? Well, I hope to warn others about your bigotry, especially because you are often in a position to influence or make decisions regarding the research and careers of field-biologists. Indeed, I have often heard you dismiss the merits of scientists’ publications as of “little scientific value” because they were concerned with ecology, life history, taxonomy, or distribution of animals and plants, and were published in “just naturalists’ journals.” You downgrade the worth of field-biologists but are firmly and smugly convinced of your own worth. In your narrow-mindedness you automatically equate the total amount of a research grant with the scientific worth of the researcher. We think you are wrong. Everyone recognizes that some types of research are very costly but these studies are not necessarily, and often are not, the most valuable and significant. If a researcher can carry out his studies with a grant of x dollars, why should he ask for 3x or 30x dollars other than to gain more prestige in the eyes of people like yourself? Some projects do not require grants of thousands and thousands of dollars for very expensive materials and equipment. Fortunately some research, and it is often excellent research, can be done by dedicated naturalists and field-biologists with a minimum of funds or even none at all. Logically, value judgments should be made on the work accomplished and not on the size of the grants. If you look at history, you'll find that some outstanding scientists carried out first-rate science both in the laboratory and field with a minimum of technical help and materials. “Oh yes,” you'll say, “but times have changed.” Molecular biology, and studies of the physiology and biochemistry of small bits of living protoplasm or ultrastructures are the “in” fields but for these research projects (like yours) is good funding absolutely essential, even at the expense of other biological research? How often have persuasive and persistent people like you convinced granting bodies and governments to fund your research at enormous expense and cared little for those who ask for a small share. In fact, when you think certain mission-oriented or applied problems are more acceptable, haven’t you slanted your research grant proposal towards some aspect of cancer, energy, food production, or the environment in order to increase your chances in the competition for available funds? Some of us have been trained and have worked in your discipline as well as in field biology and consequently we have an understanding of both these branches of biology. You, however, know only one. Although we doubt you'll ever listen, you should be aware that knowledgeable and broad-minded biologists recognize the importance of all good biological research. Of course we understand that you value your studies highly but we cannot, and will not, accept your narrow-minded view that your researches are inherently a cut above others, especially natural history and field studies. There is no doubt in our minds that investigations of plants, animals, and whole communities are of the utmost importance. The more we learn about ecosystems and the more we discover how plants and animals have adapted to their changing environments, the better we will be able to conserve and manage our renewable natural resources, and to make predictions for the future. On the other hand, as you learn in your research more and more about less and less, how can you even begin to understand most of the natural world? No doubt you won’t read this editorial because The Canadian Field-Naturalist to you is “just another naturalists’ journal,” not a laboratory-oriented one and in your eyes doesn’t rate. But I hope other field- biologists and naturalists will add their voices to mine so that in the end we will be able to counteract and destroy the false premises on which your bigotry is built. LORRAINE C. SMITH Editor 231 Seasonal Growth, Food, and Feeding Habits of Young-of-the-year Black Crappie in the Ottawa River JOHN MARK HANSON and S. U. QADRI Department of Biology, University of Ottawa, Ottawa, Ontario KIN 6N5 Hanson, John Mark and S. U. Qadri. 1979. Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie in the Ottawa River. Canadian Field-Naturalist 93(3): 232-238. In the Ottawa River, during the first growing season, young-of-the-year Black Crappies (Pomoxis nigromaculatus) attained an average total length of 68.4 mmanda mean weight of 4.03 g. The length-weight relationship was log WT = -4.84 + 2.97 log TL. Young-of-the-year Black Crappies fed mostly on copepods and cladocerans from June to September but switched to amphipods in October. They are diurnal in habit and feed mostly on organisms in the mid-waters of the littoral zone. Key Words: growth, food organisms, feeding behaviour, Black Crappie, Pomoxis nigromaculatus, juveniles, copepods, amphipods, seasonal variations, Gatineau, Quebec, Ottawa River. Black Crappies, Pomoxis nigromaculatus, in the Ottawa River are at the northern limit for the species (Scott and Crossman 1973); the fish are restricted to downstream of the Chaudiére Falls (McAllister and Coad 1974). Throughout its range in the United States little information is available on growth and feeding of young-of-the-year. In Canada, the feeding biology of the adult fish has been studied only by Keast and Webb (1966) and Keast (1968). The purpose of the present study was to investigate the seasonal growth, food, and feeding habits of young-of-the-year Black Crappies in the Ottawa River. 30 20 ° ® _ = je) = ® Qa ‘= a s '0 is) = June July Methods and Materials The study site was a section of the Ottawa River, Kettle Island Bay, a long narrow bay, 5 ha in area (75°39’0’N, 45°28’8”W). The average depth of the area studied was 3 m and the maximum was 6 m. The bay is unique in that the sand is overlain with silt and decaying detritus. The river current near the bay mouth has an average velocity of 0.63 m/s (R. G. Warnock, University of Ottawa, unpublished data) but the current inside the bay depends on wind direction and velocity. The water temperature at the surface for May to October is given in Figure |. Water hardness was 25-50 mg/L, oxygen 6-9 mg/L, carbon Aug. Sept. FiGuRE 1. Water temperatures for Kettle Island Bay in 1977. M3y) 1979 dioxide 8-15 mg/L, and pH 7.1 + 0.3; suspended solids vary greatly. In the bay, emergent vegetation consists of Sagit- taria latifolia, Scirpus sp., and Polygonum sp.; dense stands of Elodea sp. and Ceratophyllum sp. with scattered Potamogeton spp. occur from 0.2 to about 1.5m; Potamogeton spp. and Nymphaea sp., with Elodea sp. and Ceratophyllum sp. growing under- neath, occur on the two humps (at 1.5 and 2.0 m) and from the sandbar to the south shore on the eastern end of the bay. This sandbar is flanked by thick beds of Vallisneria sp. Sampling for Black Crappies was done by towing a green 6.35-mm mesh seine, 1.83 by 30.5 m. One end was fixed on shore as the other was towed out from shore and back by a boat with a 20-HP outboard motor. From the 2000 or more young captured on each sample date, only 30-35 were kept; these were sampled for lengths (mm), weight (g), scales, and stomach contents. Sampling was done at 2-wk intervals from May to October of 1977 and about 300 young were randomly selected for detailed examina- tion. Organisms from the stomachs of fish captured in the field were identified (mostly at order level) and the number of organisms per fish, percent frequency, and percent volume were compiled on a monthly basis. Total length (mm) was used throughout. For com- parison to other studies, conversion factors for fork length and standard length were obtained as follows: HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE Fork length (cm) psy NS 233 SL = 0.8302 TL — 0.3206 (based on 71 individual measurements) and FL = 0.94 TL + 0.0135 (Figure 2). Wet weights were taken to the nearest 0.01 g. The anterior scale radius was measured with an ocular micrometer (40 X) and reduced to actual size in millimetres. Fulton’s condition factor was calculated for each fish. The significance of the difference of sample means was determined by a /-test. Food and feeding habits of young Black Crappies were observed in the field and in the aquarium. From late June to early August, 10 young-of-the-year Black Crappies were maintained in an aerated all-glass aquarium, 61 X 30.5 X 30.5 cm. City of Ottawa water (pH 6.9-7.2,; hardness 40-50 mg/L) was kept at 22+1°C. Substrate from Kettle Island Bay was planted with Vallisneria sp., Potamogeton sp., and Ceratophyllum sp. Lighting was from fluorescent lights and sunlight. In order to determine what foods could be eaten, food preferences, and feeding chronology, the fish in the aquarium were fed the following: Brine Shrimp (Artemia salinas) nauplii, cladocerans, copepods, amphipods (Hyallela azteca), free-swimming nema- todes, dipteran (Chironomus sp.) larvae, white worms (Enchytraeus albidus), vestigial winged fruit flies (Drosophila melanogaster), newborn Guppies ( Poeci- lia reticulatus), and shiner (Notemigonus crysoleucas and Notropis spp.) fry. The young crappies were observed for | h after the food was introduced. | 2 3 4 5 6 7 8 9 Total length (cm) FIGURE 2. Relationships of fork length to total length of young-of-the-year Black Crappies in 1977. FL = 0.9443 TL + 0.0135. Averages of lengths with 95% confidence limits are shown. 234 Results Large numbers of young-of-the-year Black Crappie were captured near the Elodea and Ceratophyllum beds until October when the weeds disintegrated and the young moved into deeper waters. During the study period all sizes of Yellow Perch (Perca flavescens), Northern Pike (Esox Jucius), Largemouth Bass (Micropterus salmoides), Pumpkinseed (Lepomis gibbosus), Rockbass (Ambloplites rupestris), Brown Bullhead (Jctalurus nebulosus), Golden Shiner ( Note- migonus crysoleucas), Emerald Shiner (Notropis atherinoides), and Silvery Minnow (Hybognathus nuchalis) were captured consistently, but in varying proportions, with the Black Crappies. The first batch of young Black Crappie was captured 20 June 1977. The mean length of the first sample of young Black Crappie was 25.5 mm and rapid growth occurred until mid-August when growth in length slowed to a value of 68.4 mm (Figure 3). The length-weight relationship (Figure 4) is exponential with a slope of 2.98. Rapid growth in weight occurred from July to late September (Figure 5) and slowed to a mean value of 4.03 g. The mean condition factor was 1.30 (Figure 6) except for early August when the value of 1.01 is Total length (cm) Ts) S50) 15 3| June July THE CANADIAN FIELD-NATURALIST Vol. 93 significantly (P< 0.005) lower than adjacent values. Scales appeared on fish at about 25 mm in length, weighing an average of 0.23 g. The relationship between length and the anterior scale radius (Figure 7) is given by Scale Radius = 0.1094 TL —-0.1023, show- ing direct proportionality for young-of-the-year Black Crappies. Table 1 summarizes, by month, the food items taken by young Black Crappies in Kettle Island Bay. The percentage of empty stomachs was always less than 9 and the stomachs containing food were usually at least half full. Copepods occupied 48 to 90% of stomach volumes from June to September but were only 13% of the volume in October. Cladocerans occurred to a maximum of 50% of stomach volume in September but only 4% by volume in August. Amphipods occurred maximally at 57% of the volume in October. Under conditions of high winds and/or roiled water, young Black Crappies could not be observed feeding. On very bright days, very few young Black Crappies were observed feeding within 30 cm of the surface, except in direct shade. Otherwise there was no observed preference for position in the water column. Sept. FIGURE 3. The increase in length of young-of-the-year Black Crappies during the summer and fall of 1977. Averages and 95% confidence limits are presented. Curve fitted by eye. 1979 LOgjo weight 0.1 02 03 04 05 06 07 08 09 10 LOgio total length FIGURE 4. Length-weight relationship of young-of-the-year Black Crappies in 1977. log WT = -4.84 + 2.965 log TL. Length in millimetres, weight in grams. During the feeding studies in the aquarium it was noted that after sunset the young crappies remained motionless near the bottom either in a corner or among the vegetation. Only nematodes were entirely ignored, although dipterans that reached the bottom and amphipods that clung to weeds usually escaped. Brine Shrimp nauplii were most readily consumed. The agility of newborn Guppies made them difficult to capture. All other foods offered were eaten within the hour of observation. Discussion The mean total length of 68 mm for the first growing season for Black Crappie in the Ottawa River is comparable to values of 61 to 69 mm in Minnesota (J. H. Keuhn, unpublished data; Moyle and Burrows 1954) and 68 mm in South Dakota (J. T. Shields, HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE 235 Weight (g) June July Aug. Sept. Oct. FIGURE 5. The increase in weight of young-of-the-year Black Crappies during the summer and fall of 1977. Averages and 95% confidence limits are presented. Curve fitted by eye. unpublished data; Vanderpuye and Carlander 1971) but are far smaller than those fish in southern ponds (130-200 mm, Tucker 1973) and lakes (47-91 mm, Erikson 1952). The northern growing season is shorter and food is less abundant than in the more eutrophic southern waters. The length-weight relationship has a slope of almost 3.0 and it does not vary significantly from this value throughout its range (Table 2). The fairly consistent condition factor (Figure 6) supports the argument that body proportions of the young crappies were constant during the study period. The significant decrease in condition in early August occurred after a 2'4-wk period of unusually cold, wet, windy weather during which the young Black Crap- pies were inactive. Contrary to our findings, Neal (1963) found that temperature and water levels did not affect the growth of Black Crappies in Clear Lake, Iowa. The shorter growing season in the Ottawa River would tend to exaggerate any condition affecting growth. Table 3 shows that young-of-the-year Black Crappies have essentially the same condition factor throughout the range. 236 THE CANADIAN FIELD-NATURALIST Vol. 93 1.4 he $030 (24) 1.3 (35) i ‘6 ~b01 fe, ten (i) Condition factor [So eso lom esl 157 rs 15a SO anova June July Aug. Sept. Oct. FIGURE 6. Condition factor of young-of-the-year Black Crappies during the summer and fall of 1977. Averages and 95% confidence limits are presented. Sample size is in parentheses. TABLE |—Percent volume (above), percent occurrence (center), and number of prey items per fish (below) of the stomach contents of young-of-the-year Black Crappie in 1977 July August 9.86 September 9.86 6.97 October 1979 Anterior scale radius (mm) Total length (cm) FIGURE 7. The relationship of total length to the anterior scale radius for young-of-the-year Black Crappies in 1977. Scale radius = 0.1094 TL -0.1203. Averages and 95% confidence limits are shown. HANSON AND QADRI: YOUNG-OF-THE-YEAR BLACK CRAPPIE M37) The body length to scale radius relationship was directly proportional as was found for young crappies in South Dakota (Vanderpuye and Carlander 1971) and in Iowa (Erikson 1952). Later year classes may or may not have direct proportionality (Carlander 1977). The food eaten by young Black Crappies does not seem to vary much throughout its distribution. Pearse (1919) and Ewers (1934) reported that young Black Crappies ate copepods and cladocerans in early stages, with a progression to insects. Amphipods were not an important food item in either of these studies but the number of fish examined was small and some of the fish may have been yearlings. The destruction of the weedbeds, notably Elodea sp., in October seems to account for the increased availability of amphipods as a food item. This was also reflected in the increased numbers of odonatans taken at the same time. The relative unimportance of dipterans in the diet can be attributed to their benthic habits and mostly noctur- nal movements and the diurnal feeding habits of young Black Crappies. Mathur and Robbins (1971) found that young White Crappies (Pomoxis an- nularis) are diurnal and eat much the same food items as young Black Crappies. Although Reid (1949) reported that Black Crappies in Florida do not eat fish until they are about 80 mm long, Burris (1956) found a 25-mm individual that had eaten a fish fry. In this study, a small percentage of Black Crappies from 25 to TABLE 2—Length-weight relationships of young-of-the-year Black Crappies from various sources Length-weight relationship Source Location Total length (mm) Alabama Y oung-of-the-year Oklahoma 15-56 South Dakota South Dakota Y oung-of-the-year Y oung-of-the-year Ottawa River 25-68 log WT = -4.710 + 2.914 log TL log WT = -5.659 + 3.351 log TL log WT = -5.252 + 3.198 log TL log WT = -5.019 + 3.075 log TL log WT = -4.835 + 2.97 log TL Tucker (1973) Burris (1956) Nelson (1974) Vanderpuye & Carlander (1971) Present study TABLE 3—Condition factor of young-of-the-year Black Crappies from various sources Location Total length (mm) Alabama 50-155 Alabama Y oung-of-the-year Oklahoma 15-56 (hatchery fish) Minnesota (standard) — Ottawa River 25-68 Condition factor Source 1.21 Swingle (1965) 1.00-1.45 Tucker (1973) 0.79 Burris (1956) < 1.05 (poor) 1.22-1.50 (average) > 1.88 (excellent) Carlander (1944)* 1.30 Present study *Unpublished data with Minnesota Bureau of Fisheries Resource Investigations Report 41, revised, 40 pp., typewritten (quoted in Carlander 1977). 238 33 mm long ate fry. Both shiner fry and newborn Guppies were eaten in the aquarium but the agility of the Guppies suggests that shiner fry, too, may soon become too difficult for young Black Crappies to capture under natural conditions. Literature Cited Burris, W. E. 1956. Studies of the age, growth, and food of known age young-of-the-year black crappies and of stunted and fast growing black and white crappies of some Oklahoma lakes. Ph.D. thesis, Oklahoma A & M College, Miami. 88 pp. (quoted by Carlander 1977). Carlander, K. D. 1977. Handbook of freshwater fishery biology. Volume 2. Iowa State University Press, Ames, Iowa. 431 pp. Erikson, J. G. 1952. Age and growth of the black and white crappies, Pomoxis nigromaculatus (Le Sueur) and Pomo- xis annularis Rafinesque, in Clear Lake, Iowa. Iowa State Journal of Science 26: 491-505. Ewers, L. A. 1934. Summary report of crustaceans used as food by the fishes of the western end of Lake Erie. Transactions of the American Fisheries Society 64: 379-390. Keast, A. 1968. Feeding biology of black crappie, Pomox- is nigromaculatus. Journal of the Fisheries Research Board of Canada 25: 285-297. Keast, A. and D. Webb. 1966. Mouth and body form relative to feeding ecology in the fish fauna of a small lake, Lake Opinicon, Ontario. Journal of the Fisheries Re- search Board of Canada 23: 1848-1867. Mathur, D. and T. W. Robbins. 1971. Food habits and feeding chronology of young white crappie, Pomoxis annularis Rafinesque, in Conewingo Reservoir. Transac- tions of the American Fisheries Society 100: 307-311. McAllister, D.E. and B.W. Coad. 1974. Fishes of Canada’s National Capital Region. Fisheries and Marine Service, Miscellaneous Special Publication 24. 200 pp. THE CANADIAN FIELD-NATURALIST Vol. 93 Moyle, J. B. and C. R. Burrows. 1954. Manual of instruc- tions for lake survey. Minnesota Bureau of Fisheries Research Unit Special Publication 1. 70 pp. (quoted by Carlander 1977). Neal, R. A. 1963. Black and white crappies in Clear lake, 1950-1961. Iowa State Journal of Science 37: 425-445. Nelson, W. R. 1974. Age, growth, and maturity of thirteen species of fish from Lake Oahe during the early years of impoundment, 1963-1968. U.S. Fish and Wildlife Service Technical Paper 77: 1-29. (quoted by Carlander 1977). Pearse, A. S. 1919. Habits of the black crappie in inland lakes of Wisconsin. U.S. Bureau of Fisheries Document 867: 5-16. Reid, G. R. 1949. Food of the black crappie, Pomoxis nigromaculatus, in Orange Lake, Florida. Transactions of the American Fisheries Society 79: 145-154. Scott, W. W. and E. J. Crossman. 1973. Freshwater fishes of Canada. Fisheries Research Board of Canada, Bulletin 184. 966 pp. Swingle, W. E. 1965. Length-weight relationships of Ala- bama fishes. Auburn University Agricultural Experi- mental Zoology Series in Fisheries 3. 87 pp. (qouted by Carlander 1977). Tucker, W.H. 1973. Food habits, growth, and length- weight relationship of young-of-the-year black crappie and largemouth bass in ponds. Jn Proceedings of the 26th Annual Conference of South Eastern Game and Fish Commissioners. Edited by A. L. Mitchell. pp. 565-577. Vanderpuye, C.J. and K.D. Carlander. 1971. Age, growth, and condition of black crappie, Pomoxis nigro- maculatus (Le Sueur), in Lewis and Clark Lake, South Dakota, 1954-1967. Iowa State Journal of Science 45: 541-555. Received 25 November 1978 Accepted 22 January 1979 Nesting and Food-Storage Behavior of Peromyscus maniculatus gracilis and P. leucopus noveboracensis C. CRAIG TADLOCK and HAROLD G. KLEIN Department of Biological Sciences, State University of New York, Plattsburgh, New York 12901 Tadlock, C. Craig and Harold G. Klein. 1979. Nesting and food-storage behavior of Peromyscus maniculatus gracilis and P. leucopus noveboracensis. Canadian Field-Naturalist 93(3): 239-242. In a laboratory study we compared the nesting and food-storage behavior of Peromyscus maniculatus gracilis and P. leucopus noveboracensis (Rodentia: Muridae) given a choice between upper and lower nest-boxes. Peromyscus |. noveboracensis preferred upper nesting locations while P. m. gracilis showed no significant preference between upper and lower nest-boxes, although more lower nest-boxes were used. These results agree with those of a previous field study on the ecological distribution of these subspecies where both inhabited an upland forest habitat providing both tree- and ground-holes. Peromyscus |. noveboracensis stored significantly more food and tended to build heavier nests than P. m. gracilis. This may be related to the higher preferred ambient temperature for P. I. noveboracensis and its more southern geographic distribution as compared to P. m. gracilis. Where these subspecies occur in the same habitat, competition between them may be minimized because of differences in height-preference for nesting and food-storage. Key Words: Peromyscus, nesting, food-storage, height-preference, interspecific competition, sympatry, behavior, ecological, geographical, physiological. Previous studies have shown that the Deer Mouse, Peromyscus maniculatus gracilis, and the White- footed Mouse, P. leucopus noveboracensis, are occasionally found living within the same woodland habitats of northeastern United States and south- eastern Canada (Klein 1960; Smith and Speller 1970). Smith and Speller (1970) found both taxa cohabiting an upland forest habitat in southeastern Ontario while an adjacent mixed forest and a cedar forest were occupied only by P. m. gracilis. They observed that a significantly greater number of P. /. noveboracensis ran up trees when released from traps, compared to P. m. gracilis, which generally ran to ground refuges. Because there were probably more tree-holes in the upland forest than in the other forest habitats, one hypothesis presented by these authors was that P. /. noveboracensis occurred in the upland forest because it could find refuge in the tree-holes present. The lack of tree-holes in the mixed and cedar forests prevented inhabitation by P./. noveboracensis because of competition by P. m. gracilis for ground-holes. The objective of this study was to determine whether a difference in nest-height preference occurs between these subspecies under laboratory condi- tions. Differences in food-storing and nest-building behavior, which might contribute to ecological sep- aration of these taxa, were also investigated. Material and Methods Seven adult male Peromyscus m. gracilis and 10 adult male P./. noveboracensis were live-trapped within 40 km of Plattsburgh, Clinton County, New York, and were individually housed in plastic cages” (25 X 10 X 15 cm) for 2-5 mo prior to testing. The animals were housed and tested in a windowless, air- conditioned room maintained at 20.6 + 2°C and with a 13 light:11 dark cycle. A 7.5-W light provided dim illumination during the dark phase. Food pellets (Agway Charles River Rat and Mouse Formula) and water were provided ad libitum. The test apparatus was a modified version of one used by Klein and Layne (1978). It consisted of a pressed-board (Masonite) box 60 X 60 X 122 cm. Eight removable nest-boxes (16 X 14 X 14cm) with removable tops were attached to the outside of the box, with four nest-boxes at each of two levels. Circular entrance holes (5-cm diameter) to the upper and lower nest-boxes were located 98 and 4 cm, respectively, from the floor of the apparatus. Access’ to each upper nest-box was provided by a transversely- grooved wooden stick which extended obliquely within the apparatus from its floor to the nest-box entrance hole. The apparatus had a removable pressed-board floor and a wire mesh (7.5 mm) cover. The floor of the apparatus was covered witha 7.5-mm layer of a clay-based cat-box absorbent litter (Poise). A food-dish and water-bottle were centrally located. Two types of nesting material were provided. Pieces of paper-towel material (13 X 2 cm) and pieces of non- surgical absorbent cotton (2 + 0.5 g) were scattered on the apparatus floor at the beginning of each test period. Each mouse was tested for a 7-night period. The 10 P. |. noveboracensis were tested consecutively prior to testing the seven P. m. gracilis. Between tests of the two subspecies, the entire apparatus was washed with U9) 240 a mild ammonia solution and allowed to air-dry fora period of 5 wk. Between tests of individual mice, the apparatus was cleaned with warm water and the locations of the nest-boxes were randomized. We believe that these procedures eliminated any attrac- tion or avoidance responses by the test animals to specific nest-boxes or nest-box levels. The results were inspected for any evidence of selection of previously used nest-boxes by the test animals; none was found. On Day |, at approximately 6 h into the light phase of the daily cycle, each animal was put into the apparatus through a lower-level entrance hole. On Days 2 through 7, at about 6 h into the light phase, the following data were recorded: location of mouse, location of nest(s), location and number of food pellets, and location and number of feces. On Day 8, at 4h into the light phase, the animal was removed from the apparatus. In addition to the above data, the weights of the nests and stored food were recorded. The apparatus was cleaned and a new mouse was introduced. Results Every mouse visited each of the eight nest-boxes, as determined by the presence of feces in the nest-boxes. Maximum-minimum thermometer data indicated no difference in temperature between upper and lower nesting levels, eliminating this factor as a possible explanation for the nesting patterns observed in this study. Peromyscus |. noveboracensis exhibited a clearcut preference for upper nest-boxes but P. m. gracilis individuals were divided in their choice of nest-box level (Table 1). Individually, the mice of both taxa were highly consistent in their nestbox-level selection during the 7-night period. None nested at both levels of the apparatus and each mouse could be classified as either an “upper” or “lower” nester (Table 1). Nest construction was similar in both subspecies. All nests were constructed of shredded cotton; no THE CANADIAN FIELD-NATURALIST Vol. 93 paper-towel material was incorporated into any of the nests. The nests were all built in a spherical form, with a small cavity in the center and with two openings. A comparison of the nest weights of the two subspecies, regardless of level, indicated a tendency for P. 1. noveboracensis nests to weigh more than P. m. gracilis nests (Table 1), but the difference was not significant (Mann-Whitney U test, P= 0.094). This was caused partly by the great variability in nest weights, which ranged from 17.9 to 76.2 g in P. 1. noveboracensis and from 9.5 to 53.2 g in P. m. gracilis. A comparison of the nests built in the upper boxes showed that P. /. noveboracensis nests were significantly heavier than P. m. gracilis nests (Table 1). No significant difference was found between weights of upper and lower nests of P. m. gracilis (Mann-Whitney U test, P= 0.228). Eight of the 10 P. 1. noveboracensis constructed only one nest during the study period and these nests were used in calculating the mean nest weight (Table 1) in order to make it comparable to the mean nest weight for P. m. gracilis, which was calculated from seven single nests. The other P. /. noveboracensis individuals built two and three nests each within the 7- night period, respectively. Each of these nests weighed less than the mean weight of the eight single nests, and their weights were not included in the calculation of the mean nest weight because of this bias. Food stores were usually independent of nests. Only one P. m. gracilis stored food in its nesting-box. Five P. /. noveboracensis stored food in their nesting- boxes; of these, four individuals had additional food stores, from three to six each. Peromyscus |. nove- boracensis stored significantly more food than P. m. gracilis (Table 2). Intrasubspecific comparisons of weights of food stored in upper versus lower boxes showed no significant differences in either subspecies (Table 2). Although there was a tendency for P. /. noveboracensis to store more food in upper than in lower boxes, and for P. m. gracilis to store more food TABLE |—Summary of nesting data for mice of each subspecies nesting at each nest-box level during the 7-night study period ee SS ee a ee Se ee ere a IE Nest Subspecies location P. m. gracilis Upper Lower P. |. noveboracensist Upper Lower Weight of all Number of nests for each mice Nest weight (g) subspecies (g) Spee Mean + SE Mean + SE 4 IBS Se DSP + 3 34.0 + 12.29 mae 2b * . SOO ree 35.6 + 6.7 oooooooooo—owooaovwoaoasw@@w9w9wTw0DaO=oOD®ooo SESE eo *Difference between these weights was significant at P = 0.016; Mann-Whitney U test (Siegel 1956). tOnly eight single nests were used for calculating mean nest weights. See text for explanation. 1979 TADLOCK AND KLEIN: PEROMYSCUS NESTING BEHAVIOR 241 TABLE 2—Summary of food-storage data for mice of each subspecies storing food at each nest-box level during the 7-night study period. N = 7 for P. m. gracilis; N = 10 for P. 1. noveboracensis Weight of Weight of total Mean food stored food stored Food-store number of at each level (g) by subspecies (g) Subspecies location food-stores Mean + SE Wien xe Se i Upper 1.1 9.58 + 5.84** in A P. m. gracilis ee 26 18.55 + 5.65 28.13 + 10.90 ; ; Upper DD 45.46 + 13.94** me cs P. 1. noveboracensis es 19 30.38 + 15.54 75.84 + 17.38 *Difference between these values was significant at P< 0.05; Mann-Whitney U test (Siegel 1956). **Difference between these values was significant at P= 0.018; Mann-Whitney U test (Siegel 1956). in lower boxes, these differences were not significant (Table 2). A comparison of food-storage between the subspecies at each level showed that in the upper boxes P. /. noveboracensis stored significantly greater weights of food than P. m. gracilis (Table 2). In the lower boxes, the weights of food stored by P. /. noveboracensis were not significantly higher than those stored by P. m. gracilis. Discussion The three variables (nest-height preference, nest weight, and amount of food stored) measured in the artificial environment of our study indicated a significantly greater usage of upper nest-boxes by Peromyscus |. noveboracensis compared to P. m. gracilis. With respect to nest-height preference, the choice of upper nest locations by P. /. novebora- censis was unquestionable; a preference for upper- level food storage sites, although not significant, was relatively strong. This agrees with Nicholson’s (1941) finding that a southern Michigan population of P. /. noveboracensis used tree nest-boxes (0.9 to 9.2m above ground) to a much greater extent than ground- level boxes. Peromyscus m. gracilis showed no significant preference for nest-box level in nesting or food storage, although heavier nests were built and more food was stored in lower boxes. The patterns of nest- box utilization by these subspecies in our artificial environment supports the hypothesis of Smith and Speller (1970) that cohabitation of these subspecies in their upland forest habitat was based, in part, on differences in the utilization of above-ground cavities. Implicit in Smith and Speller’s hypothesis was the assumption that P. /. noveboracensis was forced to occupy arboreal situations because of competition for ground holes from P. m. gracilis; however, the con- sistent preference of P. /. noveboracensis for upper boxes for nesting in our study suggests that this may be a behavior typical for this subspecies. Therefore, the observed arboreal tendencies of P. 1. novebora- censis in the Smith and Speller (1970) study may not have been a change in the behavior of this population because of competition from P. m. gracilis. The lack of nest-height preference by P. m. gracilis in our study suggests an ability of this taxon to adjust to various nesting situations. It agrees with the widespread distribution of P. m. gracilis in the Smith and Speller (1970) study and the observation that P.m. gracilis individuals ran up trees on some occasions, although they usually retreated into ground holes. It also suggests that P. m. gracilis would not be a serious competitor of P. /. noveboracensis for arboreal nest sites except in periods of high population density in P. m. gracilis. The low population density of P. m. gracilis in the Smith and Speller (1970) study, coupled with the tendency of released mice to use ground refuges, is in agreement with this inference. Although P. /. noveboracensis stored a significantly greater amount of food than P. m. gracilis, and P. 1. noveboracensis tended to build heavier nests than P. m. gracilis, it is possible that these differences (mainly attributable to the apparent inhibition of P. m. gracilis in its utilization of upper nest-boxes) were not entirely behaviorally-based. The thermo- regulatory function of the nest in Peromyscus has been well-documented (Sealander 1952; Glaser and Lustick 1975). A difference in ambient temperature preference has been experimentally demonstrated between P. /. noveboracensis (32.4°C selected) and P. m. gracilis (29.1°C selected) by Ogilvie and Stinson (1966). The differences between these preferred ambient temperatures and the ambient temperature of our study (20.6°C) were probably sufficient to elicit temperature-induced food-storing and nest-building responses in these animals. A lowered ambient temperature (7° versus 27°C) caused an increase in 242 THE CANADIAN FIELD-NATURALIST food-storing behavior in P. /. noveboracensis (Barry 1976). An ambient temperature of 20°C elicited a maximal paper-shredding, nest-building response in P. |. noveboracensis, compared to the responses at 25° and 30°C (Jaslove and McManus 1972). The dif- ferences in food-storing and nest-building behavior shown by the subspecies in our study may have been differential responses to the low ambient temperature of our study relative to the preferred ambient temperatures of these taxa. The differences in pre- ferred ambient temperature is presumably based ona physiological difference between the subspecies. Riedel (1967) demonstrated that, at low temperatures, P. m. gracilis survived better than P. 1. noveboracensis, whereas the opposite was true at high temperatures. A situation of severe competition for nest sites may exist, however, between P. Jeuwcopus and a closely related species, P. gossypinus, where they are sym- patric in the northern part of the Gulf coastal plain and in the Mississippi Valley of southern United States. McCarley (1963) concluded that in this area P. gossypinus excluded P. leucopus from lowland forest habitats by interspecific competition. Com- petition for arboreal nest-sites may have been a major factor in this interspecific relationship. Ten male P. gossypinus palmarius from Florida showed a very strong preference for upper nest-boxes when tested in an apparatus similar to that of our study (Klein and Layne 1978). A direct comparison (using a similar apparatus) of the nest-height preferences and inter- actions of individuals from sympatric populations of P. leucopus and P. gossypinus might clarify the extent and nature of their interspecific competition. The behavioral and physiological differences be- tween P. m. gracilis and P. |. noveboracensis are probably based on adaptations related to the geo- graphic-ecologic distributions of these subspecies (and their respective species). For example, nesting in an arboreal site would be more appropriate in areas of mild winter climate, whereas in areas of severe winter climate it might cause a serious thermoregulatory problem. Such a species- or subspecies-typical be- havior, if unmodifiable, could be a factor limiting the geographic-ecologic distribution of the taxon. The different nest-height preferences shown by these subspecies may reflect the climatic conditions in their respective environments. Although the ranges of P. /. noveboracensis and P. m. gracilis broadly overlap, P. |. noveboracensis is more southern than P. m. gracilis in its geographic distribution and, where these taxa are sympatric, corresponding ecologic dif- ferences in local distribution generally occur (Hamil- ton 1943; Klein 1960; Miller and Kellogg 1955). Where cohabitation of these subspecies occurs, our study suggests that sufficient differences exist in the height-preference aspect of nesting and food-storage Vol. 93 behavior to result in decreased competition between them. Acknowledgments We thank James N. Layne for the valuable suggestions he made during the preparation of this paper. Literature Cited Barry, W. J. 1976. Environmental effects on food hoarding in deermice (Peromyscus). Journal of Mammalogy 57: 731-746. Glaser, Harriet and Sheldon Lustick. 1975. Energetics and nesting behavior of the northern white-footed mouse, Peromyscus leucopus noveboracensis. Physiological Zoology 48: 105-113. Hamilton, W. J., Jr. 1943. The mammals of eastern United States. Comstock Publishing Co., Ithaca, New York. Jaslove, S. W. and J. J. McManus. 1972. The influence of temperature on feeding, drinking and nest construction by the white-footed mouse, Peromyscus leucopus. Bulle- tin of the New Jersey Academy of Science 17: 1-6. Klein, H. G. 1960. Ecological relationships of Peromyscus leucopus noveboracensis and P. maniculatus gracilis in central New York. Ecological Monographs 30: 387-407. Klein, H. G. and J. N. Layne. 1978. Nesting behavior in four species of mice. Journal of Mammalogy 59: 103-108. McCarley, Howard. 1963. Distributional relationships of sympatric populations of Peromyscus leucopus and P. gossypinus. Ecology 44: 784-788. Miller, G.S., Jr. and Remington Kellogg. 1955. List of North American Recent mammals. U.S. National Museum, Bulletin 205. Nicholson, A. J. 1941. The homes and social habits of the woodmouse (Peromyscus leucopus noveboracensis) in southern Michigan. American Midland Naturalist 25: 196-223. Ogilvie, D.M. and R.H. Stinson. 1966. Temperature selection in Peromyscus and laboratory mice, Mus musculus. Journal of Mammalogy 47: 655-660. Riedel, D. 1967. Effect of environmental temperature on the body temperature, oxygen consumption and survival of two species of Peromyscus (Rodentia: Cricetidae). M.Sc. thesis, Carleton University, Ottawa. (Reported in Smith and Speller 1970.) Sealander, J. A. 1952. The relationship of nest protection and huddling to survival of Peromyscus at low tempera- ture. Ecology 33: 63-71. Siegel, S. 1956. Non-parametric statistics for the be- havioral sciences. McGraw-Hill Book Co., New York. Smith, D. A. and S. W. Speller. 1970. The distribution and behavior of Peromyscus maniculatus gracilis and Pero- myscus leucopus noveboracensis (Rodentia: Cricetidae) in a southeastern Ontario woodlot. Canadian Journal of Zoology 48: 1187-1199. Received 2 November 1978 Accepted 8 February 1979 Life History Characteristics of Little Brown Bats (Myotis lucifugus) in Alberta DAVID B. SCHOWALTER,! JOHN R. GUNSON,! and LAWRENCE D. HARDER? 'Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2 2Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9 Schowalter, D. B., J. R. Gunson, and L. D. Harder. 1979. Life history characteristics of Little Brown Bats (Myotis lucifugus) in Alberta. Canadian Field-Naturalist 93(3): 243-251. Life history data of Little Brown Bats (Myotis lucifugus) were recorded from 1971 to 1978 during the course of a rabies study in Alberta. The study is the northern-most examination of the life history of the species to date. The number of bats that inhabited the 269 maternity colonies examined, varied from less than 15 to approximately 1100 adults. Seventy-five percent of the colonies in central Alberta were in occupied buildings. Differences in the biology of Little Brown Bats from more southern studies included later commencement of parturition, shortened period of parturition, non-breeding of juvenile females, substantial numbers of females at non-maternity shelters during mid-summer, extensive roosting at exposed locations during summer and fall, and aggregations of unknown function in abandoned buildings during fall. Adult females gained weight during and after pregnancy and in late summer, and lost weight at parturition and in late July. Juvenile males entered hibernation weighing less than adult males. Adult males were seldom located in maternity roosts but predominated at a hibernaculum. Key Words: Little Brown Bat, Myotis lucifugus, life history, Alberta. The Little Brown Bat (Myotis lucifugus) is one of the most widely distributed species of North Ameri- can bats (Hall and Kelson 1959); its range encom- passes a wide variety of habitats and climates. Latitudinal gradients in photoperiod and the duration © of the seasons could induce selective pressures that might result in behavioral and physiological variation within this species. The biology of M. /. lucifugus in eastern North America (reviewed by Humphrey and Cope 1976) and of M. 7. occulatus in New Mexico (O’Farrell and Studier 1973, 1975; Studier and O’Farrell 1972; Studier et al. 1973) have been extensively studied, but life history information for more northern populations of Little Brown Bats is necessary to identify and define characteristics that vary with latitude. The initiation of a research program on bat rabies was prompted by an outbreak of rabies in Alberta in 1970 and the first diagnosis of this disease in a bat from the province in the following year. Dorward et al. (1977) reported preliminary conclusions regarding the incidence of rabies in bats from Alberta. This paper presents biological data on the Little Brown Bats collected during the course of the study. Observa- tions were made from 49°N to 57°N, but most information was collected between 52°N and 55°N, the region commonly known as ‘central Alberta.’ This area is 4-6° north of studies in Ontario (Fenton 1969, 1970), approximately 11° north of studies in Indiana and Kentucky (Humphrey and Cope 1976), and approximately 14° north of studies in New Mexico (O’Farrell and Studier 1973, 1975). The Little Brown Bats that inhabit the non- mountainous portions of Alberta have been identified as M. 1. lucifugus (Hall and Kelson 1959; Soper 1964; Banfield 1974); however, southern Alberta is occupied by a separate form, M./. carissima (Smith and Schowalter 1979). Unless otherwise indicated and with the exception of a discussion of the provincial distribution of M. lucifugus, the data presented here deal with M./. lucifugus as it occurs in central Alberta, although some observations may be from areas of intergradation with the southern form. Also discussed are bats from a hibernaculum near Cado- min (see Figure 1) which have been previously classified as M./. pernox (Soper 1964) but are indistinguishable from M./. lucifugus of central’ Alberta (H. C. Smith, Provincial Museum of Alberta, personal communication). Methods Most colonies and some shelters were located through complaints of bats made to government offices by the public. Systematic surveys, responses to newspaper advertisements, and a questionnaire mailed to rural landowners provided additional locations. History sheets of bats sent to the Animal Diseases Research Institute (Western) [ADRI(W)] of Agriculture Canada at Lethbridge, Alberta for rabies testing provided the majority of the locality records for individual bats. Additional individual records were obtained from specimens at the Provincial Museum of Alberta, the Museum of the Department of Zoology at the University of Alberta, as well as 243 244 THE CANADIAN FIELD-NATURALIST Rocky 200 km Individual O Colonies — estimated size Solonetzic Soils 1-25 e (78) 26-100 ®© (99) 101-250 @ (57) 251-500 @ (28) over — 500 (5) Vol. 93 Suntain HouseO FicuRE 1. Distribution and estimated size of Little Brown Bat maternity colonies located in Alberta, 1971-1977. from published data and our own field activities. Most roosts were entered to identify and enumerate the bats present. Bats were periodically collected at maternity roosts for rabies testing and additional bats were captured at maternity roosts known to harbor rabies, a variety of non-maternity shelters, a hiber- naculum, and by mist-netting over ponds and streams. Aggregations of bats in buildings in fall (‘fall aggrega- tions’) were sampled in 1975. The following informa- tion was recorded for each bat: age (juvenile or adult) as determined by the closure of the epiphyses of the finger joints, sex, and weight. Examination of the nipples of adult female bats (Racey 1974) provided an indication of their reproductive history, and the current reproductive condition of a sample of female bats taken prior to the commencement of parturition was determined by uterine examination. The number of fetuses and the horn of implantation were recorded for pregnant females. Eight non-parous unsuckled female bats collected at shelters in mid-June were aged by dental annuli (Schowalter et al. 1978b) to determine the ages of non-breeding females. 1979 Results and Discussion Distribution We examined 269 maternity roosts of Little Brown Bats. Sufficient data to determine the relative abun- dance and distribution of this species are available only from the more heavily settled southern and central areas of the province. Concentrations of colonies were evident in several lake-cottage develop- ments, in certain rural areas of knob and kettle (‘pot- hole’) topography in central Alberta, and in areas of extensive irrigation development near Brooks and Lethbridge. Few colonies were located near Vegre- ville, where the habitat resembled that of areas with many colonies, or north of Brooks and Medicine Hat (Figure 1). Questionnaire responses, interviews with residents, and the pattern of submission of rabies- suspect bats all suggested that few Little Brown Bats are found in these regions. Silver-haired Bats ( Lasion- ycteris noctivagans) (Schowalter et al. 1978a) and Hoary Bats (Lasiurus cinereus) (Schowalter and Dorward 1978) have been submitted to ADRI(W) from these areas with sufficient frequency to suggest that the absence of Little Brown Bats is probably not an artifact of the reporting system. The area of low abundance of Little Brown Bats in Alberta corresponds, in part, to the distribution of solonetzic soils (Figure 1) and the distribution of grasslands (see Hardy 1967). Water bodies in solon- etzic soil zones are known to support different arthropod faunas than those of other areas (Rawson and Moore 1944) and the food supply for bats in solonetzic soil areas may thus be of different quality than in other areas. Grasslands are less heavily settled and have far fewer trees than other agricultural areas and thus offer fewer potential roost sites. These factors do not, however, appear to explain completely the observed distribution of Little Brown Bats. Maternity Colonies Although some complaints of bats in cottages were not investigated and colonies in unoccupied buildings and natural roosts were less likely to be reported, occupied buildings were apparently favored for colony formation in central Alberta. Of the 196 M. /. lucifugus maternity colonies located and classified in this region, 147 were in occupied buildings and, with the exception of one colony in a tree, the remainder were in cottages, abandoned houses, sheds, and barns. Maternity roosts were typically dark and poorly ventilated and almost always subject to heating by the sun, as described by Fenton (1970) and Humphrey and Cope (1976). Most colonies occupied a single building, although bats in larger colonies appeared to use as many as six buildings. The estimated number of adult Little Brown Bats in individual maternity colonies ranged from less than 15 to 1100; between 50 SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA 245 and 300 was usual. From 1972 to 1975 bats returned to the maternity roosts during the first half of May. During 1976 and 1977, years with early and mild springs, bats were reported as early as 24 April (1976) and many roosts contained bats by | May. The earliest date that a juvenile was taken at a non-nursery roost was 9 July (1976), but juveniles and suckled females were regularly observed at such locations only during the last week of July, indicating that nursery break-up was advanced by that time. By the second week of August most maternity roosts had few bats, but juveniles were found at this type of roost as late as 26 August (1975). Juvenile bats outnumbered adult females in our August samples from maternity colonies (69:30), indicating that adults tend to leave nurseries sooner than juveniles, as Humphrey and Cope (1976) have reported. The sexes were equally represented in our collec- tion of fetuses and juveniles from maternity colonies, but adult males were seldom found in these roosts (Table 1). This low occurrence of males agrees with the observation by Humphrey and Cope (1976) that males rarely frequent maternity colonies in the northern portion of the species range compared with regions to the south. TABLE |1—The percentage of males among samples of Little Brown Bats from central Alberta: 1972-1978. Sample size in parentheses Percentage of males Sample type Fetuses Juveniles Adults Maternity roost 53 (61) 49 (395) 0.4 (807) Shelters May 3 (Sl) June 41 (66) July 20 ~=(5) 38@8)) Aug. 31 (45) 24 (49) Sept. 41 (106) 30 (147) Fall aggregation 38 = (8) Li CS) Swarming 67 (30) 80 (81) Hibernation 78 (270) Other species of bats were seldom encountered in Little Brown Bat maternity roosts. Three attics used by Little Brown Bats were also colonized by Big Brown Bats (Eptesicus fuscus). Little Brown Bats appeared to have recently colonized two of these attics after modifications to the buildings and the two species used different parts of the attics. In the third case a single beam separated the roosts of the species, but at the time of observation the majority of the Big Brown Bats were in the walls of the building rather than immediately adjacent to the Little Brown Bats. A 246 Silver-haired Bat was observed in one colony on 17 May (1974). Shelters We recognized two types of non-nursery roosts or shelters used by Little Brown Bats. Some bats persistently used concealed locations in barns, attics, log buildings, piles of lumber, and behind shutters, tree bark, and a piece of tin wrapped around a tree trunk. One banded male was observed on each of six inspections of a roost over three summers. Shelters in exposed situations, typically darkened recesses of brick buildings between 2 and 7 m above the ground, were used daily by | to 22 bats. There was evidently a high turnover of individuals at these locations: some property owners destroyed roosting bats daily, seemingly without reducing the number of bats using the roost. Bats banded at this type of shelter have not been recaptured at the original banding site. More bats were collected at exposed roosts at night than during the day. Although bats roosting at shelters during the night usually became torpid, many of those that were not disturbed, aroused and departed prior to sunrise. Bats found during the day were almost always torpid. With the exception of a male collected on 23 May 1978, all bats captured at shelters during May were females (Table 1); many had previously suckled young (Table 2) and may have been migrating to maternity colonies. Male Little Brown Bats were also poorly represented (1/24) in May submissions to ADRI(W) from Alberta and May samples from shelters in Michigan (Miller 1955). These observations probably relate to the tendency of males to depart from the hibernacula later than females (Humphrey and Cope 1976). TABLE 2—The percentage of unsuckled adult female Little Brown Bats among samples from central Alberta: 1972- 1978. Sample size in parentheses Percentage Sample type Period unsuckled Maternity roosts Prior to 15 June! 17 (329) After 1 July? 18 (331) Shelters May 30 (30) June 95 (39) July 88 (8) Aug. 41 (37) Sept. 44 (91) Fall aggregations Aug.—Sept. 18 (17) ‘Pregnant unsuckled bats taken after 15 June could have been identified as suckled. 2Bats which appeared to be unsuckled after | July are not considered likely to give birth that year. THE CANADIAN FIELD-NATURALIST Vol. 93 Female Little Brown Bats were observed at shelters in Alberta proportionately more frequently than has been reported elsewhere (Table 3). The proportion of females appears to increase with latitude (Table 3), but the distance from hibernacula is known to affect the proportion of males in maternity roosts (Humph- rey and Cope 1976) and may affect the sex ratio in shelters. Female Little Brown Bats that use shelters during June have been thought to be non-parous (Miller 1955; Humphrey and Cope 1976). In Alberta the majority of females taken at shelters during June were unsuckled (Table 2). None of eight females captured at shelters, whose uteri were examined was pregnant. Palpation of other female bats at shelters during June produced no evidence of pregnancy. Although sex-ratio data for bats caught at shelters (Table 1) represent observations made at varying intervals with varying effort over 5 yr, significant annual variation is assumed not to have occurred. TABLE 3—Percentage of males among Little Brown Bats taken in shelters during June from various North American locations Latitude Percent Location north males (n) Reference Oklahoma 35 100 (17) Glass and Ward (cited in Humphrey and Cope 1976) Indiana 39 100 (3) Humphrey and Cope 1976 West Virginia 39 New York 42 100 (40) Krutzsch 1961 87 (63) Davis and Hitchcock 1965 Michigan 42 84 (85) Miller 1955 Vermont 42 98 (648) Davis and Hitchcock 1965 Alberta 54 41 (66) This study The use of shelters increased during late July with the appearance of juvenile and suckled adult female bats. The largest numbers of bats were collected at shelters around the end of August and generally declined thereafter. During August and September bats frequented roosts not used earlier in the year. Bats roosting on white buildings were easily seen from roadways and were collected by driving through business districts. Exposed roosting was observed or reported from throughout central Alberta. Bats roosting at such locations may have been migrants passing through unfamiliar areas. An adult male Little Brown Bat banded at such a shelter in Edmonton on 8 September 1977 was recaptured 5 d later near Rocky Mountain House, a movement of 173 km. 1979 Frequent use of exposed roosts by Little Brown Bats has not been reported previously. The roosting is so obvious that it is doubtful that it would be missed by bat researchers if it occurred in their research areas. Efforts to locate bats or bat droppings at potential exposed shelters in Lethbridge and Medicine Hat did not produce evidence of bats, suggesting that the subspecies found in southern Alberta may not use exposed roosts. The behavior and transience of bats at shelters were similar to those of the Cave Bat (Myotis velifer) at ‘transient colonies’ in early spring and late autumn in Kansas (Kunz 1974). This behavior is thought to be important in reducing metabolic expenditure and in migration. In central Alberta we have taken a few Keen’s Bats (Myotis keenii) and Big Brown Bats at shelters used by Little Brown Bats, and Silver-haired Bats are frequently encountered by window washers in exposed situations on higher buildings in Edmonton. As similar observations in other areas are not known to us, exposed roosting in central Alberta may be important in_ thermo- regulation, possibly in response to unidentified climatic factors. Pregnancy Suckled ‘a E © b& co) = £ = C) s > a] ° a SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA 247 Fall Aggregations We have observed concentrations of 500 or more Little Brown Bats in buildings on three occasions during late August and early September. All of the buildings used by these ‘fall aggregations’ were abandoned and unheated, but the bats were not torpid during the day. Local residents reported that one of these buildings had been used by bats during several autumns. The predominance of heavy female bats in two small samples (Table 1, Figure 2) suggests that fall aggregations may be related to migration rather than to breeding. Humphrey and Cope (1976) noted evidence of “autumn dispersal and transient nursery visitation” distinct from swarming which takes place at caves. Fall aggregations in Alberta may be part of the same phenomenon noted by Humphrey and Cope (1976), but few data are available on activity of Little Brown Bats away from caves at this time of year. Weight Dynamics Adult female Little Brown Bats gained weight during pregnancy, after parturition, and prior to hibernation (Figure 2). Weight losses occurred as a 8 Fall Aggregation Lactation MAY JUNE JULY AUGU ST SEPTEMBER FIGURE 2. Seasonal variation in body weight of adult female Little Brown Bats in central Alberta (mean + SE, range), 1971-1977. 248 result of parturition and from unknown causes at the end of July, at least among females in maternity roosts. Data from Ontario (Fenton 1970) are comparable to those presented here, but they do not extend into August, so that it is not known whether the weight loss observed at that time is general for the species. Juvenile Little Brown Bats gain weight rapidly until late July (Figure 3). Stomachs of juveniles examined at that time contained insects only, insects and milk, or milk only, indicating that weaning had begun, Weights are less consistent after that time, suggesting that juveniles may lose some weight after colony break-up. A collection on 20 August (Figure 3) included many juveniles that appeared to be starving and were of low weights. Swarming and Hibernation A preponderance of males was netted at the mouth of Cadomin Cave during September 1977 (Table 1), a finding that is similar to observations elsewhere (Davis and Hitchcock 1965; Hall and Brenner 1968; NURSERY 9 8 : 9 7 > io 6 = ~ 8 AS 5 4 20 NON-NURSERY JUNE JULY AUGUST SEPTEMBER Month FIGURE 3. Seasonal variation in body weight of juvenile Little Brown Bats in central Alberta (mean + SE, range), 1974-1977. THE CANADIAN FIELD-NATURALIST Vol. 93 Fenton 1969; Humphrey and Cope 1976). Swarming may have ended at Cadomin Cave between 20 and 27 September 1977 as only four individuals were captured on the later date. This change was probably not caused by weather as Long-legged Bats (Myotis volans) were captured in greater numbers on 27 than on 20 September. Juvenile bats captured on 20 September 1977 at Cadomin Cave weighed significantly less (t-test, P<0.01) than juveniles at shelters in Edmonton and Edson on 19 and 20 September (Table 4). Weights of adult males in the two samples were nearly identical. Too few adult females were captured at Cadomin Cave to allow comparison. Juveniles may have lost weight in moving to the cave from maternity roosts, whereas the distance moved by adult males from the summer range may have been less, as suggested by the observations of Humphrey and Cope (1976). The mean weight of adult males at Cadomin was signifi- cantly greater (t-test, P<0.01) than that of juvenile males, indicating that juveniles likely enter hiber- nation weighing less than adults. TABLE 4—Mean weights (+ 95% confidence intervals) of Little Brown Bats captured at shelters in Edmonton and Edson and at Cadomin Cave 19-20 September 1977. Differences tested by Student’s t-test Juvenile Juvenile Adult Location females males males Cadomin 8.7+0.4 (7) 8.6+0.4 (17) 9.8+0.2 (44) Shelters 9:9) == O'S) (116) 92 == 056 0) O30 2518) P<0.01 P<0.02 P<0.8 The earliest and latest dates that we have found numerous bats hibernating were 20 September 1977 and 15 May 1975, respectively. Fifteen bats captured on the spring date were all males, which suggests that most or all females may have left the hibernaculum. The sex ratio of hibernating bats favored males (Table 1) and was similar to that reported elsewhere (Humphrey and Cope 1976). The locations of the hibernacula of most Little Brown Bats found in Alberta are unknown despite inspection of mines and caves and continuing liaison with caving organizations. The only suitable areas for hibernacula within the province are in the Rocky Mountains and a large area of karst in the northeast of the province. The numerous coal mines of the central and southern portion of the province, which would likely offer potential hibernacula, have been routinely blasted shut in accordance with provincial law. Three hibernacula were located: Cadomin Cave and a cave near Nordegg, both in the Rocky Mountains, over- 1979 winter between 500 and 1000 bats, and a third cave in Wood Buffalo National Park in the northeast contains a small number of hibernating Little Brown Bats. Many bats would have to travel 450 km from maternity roosts to reach either of these areas. Reproduction Of the 183 adult female Little Brown Bats collected in maternity roosts prior to the onset of parturition (15 June), 91% were pregnant. Twin fetuses were observed twice among 312 pregnancies; 290 of the single fetuses had implanted in the right horn. The incidence of pregnancy among 162 bats that had previously suckled young (95%) was significantly higher (x test, P<0.001) than that among 21 unsuckled females (52%). Pregnant, unsuckled bats generally weighed less than pregnant, suckled females and they carried fetuses that often appeared smaller than those of suckled bats. Three of eight unsuckled bats caught in nurseries during mid-July were resorbing embryos. The earliest neonate was observed on 15 June 1975 and the latest date a pregnant bat was captured was 15 July 1976. Parturition appeared to be highly synchronous both within and between colonies as reflected in the relatively consistent patterns of weight gains of pregnant females and juvenile bats (Figures 2, 3) even though the samples were taken from colonies in a variety of buildings and over a broad geographical area. Based on fetal inspections and observations of neonates, the majority of the bats born in 1972 through 1975 were born during the last week of June. Parturition tended to commence earlier during 1976, possibly owing to the early arrival of adult female bats at the maternity roosts. Greater variation in fetal and juvenile development was noted during that year. Limited observations during 1977 indicated similarity with 1976. Like the females of many temperate-zone vesperti- lionids (Asdell 1964), the majority of female M. /. lucifugus in central Alberta do not bear young as yearlings. Schowalter et al. (1978b) reported that yearling bats comprised a smaller than expected proportion of the 90 adult female bats in a nursery in central Alberta and that all of the yearlings present were non-parous. All eight of the unsuckled, non- parous females from shelters that we have aged by counts of dental annuli were young of the previous year. Non-parous yearling bats probably comprise the majority of the unsuckled bats found both in maternity colonies and at shelters in central Alberta (Table 2). Juvenile Little Brown Bats are thought to be reproductively active in New Mexico (O’Farrell and Studier 1975) and Indiana (Humphrey and Cope 1976) as non-parous females were almost entirely SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA 249 absent. Davis and Hitchcock (1965) found 11 of 21 yearling Little Brown Bats (aged by closure of finger epiphyses) examined in New England to be pregnant, suggesting that nonbreeding by juvenile females may tend to increase with latitude. This may account for the tendency for the increased proportion of females in shelters in June with increased latitude (Table 3). Differing timing of events both within the year and within the life of individual female bats parallels increased latitude. Later commencement of parturi- tion in Little Brown Bats with increased latitude is well documented (Fenton 1970; Humphrey and Cope 1976). O’Farrell and Studier (1975) have also observed that the period of parturition becomes abbreviated in more northern populations. Both these trends are evident in comparisons of data from central Alberta with those available from areas to the south. In southern Alberta, which has a frost-free period that is approximately 20 days longer than in central Alberta (Government and University of Alberta 1969), parturition of M. 1. carissima is estimated to extend from at least as early as mid-June to the first of August, a much longer period than in M. /. lucifugus in central Alberta. Determination of mean parturition date of M. /. carissima was not possible from the limited samples as parturition was highly variable within and between colonies and between years; however, parturition in 1975 and 1976 was later there than in central Alberta. A collection made south of Lethbridge on 12 July 1976 included seven lactating and nine pregnant females, and two volant young. Pregnant females were taken at the same colony 21 July 1975. Observations at this colony in July 1977 indicated that most young were born by at least mid- July of that year. Visits to other colonies in the region also indicated variability of parturition. The longer, later, and more variable period of parturition in southern Alberta may relate to minimal confrontation with cold weather on the maternity range and to the longer warm season which would, in many years, allow late-born bats sufficient time to prepare for hibernation. O’Farrell and Studier (1975) related a brief parturi- tion period of Myotis thysanodes to synchrony of departure from hibernacula. Although it is possible that most female Little Brown Bats found in central Alberta follow this pattern, the geographic extent of the region and the number of bats involved suggest that this is not a complete explanation of the brief parturition period observed. Female bats may return to maternity roosts before food is abundant and temperatures are favorable to take advantage of the earliest arrival of favorable conditions. Low tempera- tures are known to slow fetal development in some bats (Racey 1973; Studier and O’Farrell 1972; Studier et al. 1973) and fetuses may tend to be at a similar 250 stage of early development at the time of arrival of — warm weather, regardless of the date of arousal from hibernation and arrival at the colony of the females. In years with mild early springs fetal development in arriving females would not be slowed and an earlier, more variable parturition period would result, as observed in 1976. Whereas the gestation period of early-arriving females may tend to be extended by cool conditions in early May, gestation periods of late-arriving females may be shortened by the com- paratively high temperatures during long daylight periods that are characteristic of late June. These considerations suggest that the relative abbreviation of the parturition period observed in Alberta, while probably adaptive to bats living at higher latitudes, may largely be a response to environmental condi- tions rather than a specific regional adaptation of bats. The results described here were not obtained through research designed specifically to examine bat life-history. Detailed examination of a single colony or local population rather than examination of many colonies over a large geographical area would clarify details of phenology and growth patterns. Little Brown Bats occur far north of the present study area and study of such boreal populations would identify adaptations to exceedingly long daylight periods. The trends identified here may be more evident in those populations. Acknowledgments The research described here was part of a joint program of the Alberta Fish and Wildlife Division, ADRI(W) and the Veterinary Services Division of the Alberta Department of Agriculture. Funding was largely provided by the latter agency. The continued interest and support of H. Vance and G. Whenham of the Veterinary Services Division is gratefully ac- knowledged. J. Bradley and W. Dorward of ADRI(W) agreed to the use of data from their files and consented to our use of rabies-suspect specimens. D. Bedard, H. Boumans, P. Cole, W. Johnson, R. Masuda, D. Meyer, D. Pipella, N. Previsich, B. Prins, and W. Wynnyk assisted in various ways in field and laboratory. H. Smith of the Provincial Museum of Alberta provided bat bands and showed a continuing interest in our research. Parks Canada and the Alberta Parks Division consented to and assisted with activities in parks. G. Lynch, A Todd, and J. Murie reviewed drafts of this paper. Literature Cited Asdell, S. A. 1964. Patterns of mammalian reproduction. Cornell University Press, Ithaca. 670 pp. Banfield, A. W. F. 1974. The mammals of Canada. Univer- THE CANADIAN FIELD-NATURALIST Vol. 93 sity of Toronto Press, Toronto. 438 pp. Davis, W.H. and H.B. Hitchcock. 1965. Biology and migration of the bat, Myotis lucifugus, in New England. Journal of Mammalogy 46: 296-313. Dorward, W.J., D.B. Schowalter, and J. R. Gunson. 1977. Preliminary studies of bat rabies in Alberta. Canadian Veterinary Journal 18: 341-348. Fenton, M. B. 1969. Summer activity of Myotis lucifugus (Chiroptera: Vespertilionidae) at hibernacula in Ontario and Quebec. Canadian Journal of Zoology 47: 597-602. Fenton, M.B. 1970. Population studies of Myotis luci- fugus (Chiroptera: Vespertilionidae) in Ontario. Royal Ontario Museum, Life Sciences Contribution 77: 1-34. Government of Alberta and The University of Alberta. 1969. Atlas of Alberta. University of Alberta Press. 158 pp. Hardy, W. G. (Editor). 1967. Alberta/A natural history. Hurtig, Edmonton. 343 pp. Hall, E. R.and K. R. Kelson. 1959. The mammals of North America. Volume I. Ronald Press, New York. 546 pp. Hall, J.S.and F. J. Brenner. 1968. Summer netting of bats at a cave in Pennsylvania. Journal of Mammalogy 49: 779-781. Humphrey, S.R. and J.B. Cope. 1976. Population eco- logy of the Little Brown Bat, Myotis lucifugus, in Indiana and north-central Kentucky. American Society of Mammalogists, Special Publication Number 4. 81 pp. Kunz, T.H. 1974. Feeding ecology of a temperate in- sectivorous bat (Myotis velifer). Ecology 55: 693-711. Krutzsch, P.H. 1961. A summer colony of male Little Brown Bats. Journal of Mammalogy 42: 529-530. Miller, J.S. 1955. A study of the roosting habits, and of the environmental factors concurrent with the time of evening flight, of Little Brown Bats (Myotis lucifugus) in northern lower Michigan. Ph.D. thesis, University of Michigan. 86 pp. O'Farrell, M. J. and E. H. Studier. 1973. Reproduction, growth and development in Myotis thysanodes and M. lucifugus (Chiroptera: Vespertilionidae) in north- eastern New Mexico. Ecology 54: 18-30. O’Farrell, M. J. and E. H. Studier. 1975. Population struc- ture and emergency activity patterns in Myotis thysanodes and M. lucifugus (Chiroptera: Vespertilionidae) in north- eastern New Mexico. American Midland Naturalist 93: 368-376. Racey, P. A. 1973. Environmental factors affecting the length of gestation in heterothermic bats. Journal of Reproductive Fertility (Supplement) 19: 175-189. Racey, P. A. 1974. Aging and assessment of reproductive status of Pipistrelle bats ( Pipistrellus pipistrellus). Journal of Zoology (London) 173: 264-271. Rawson, D.S. and J. E. Moore. 1944. The saline lakes of Saskatchewan. Canadian Journals of Research, D 22: 141-201. Schowalter, D. B. and W. J. Dorward. 1978. Some western Canadian bat records. Blue Jay 36: 49-50. Schowalter, D. B., W.J. Dorward, and J. R. Gunson. 1978a. Seasonal occurrence of Silver-haired bats (Lasionycteris noctivagans) in Alberta and British Colum- bia. Canadian Field-Naturalist 92: 288-291. Schowalter, D. B.,L. D. Harder, and B. H. Treichel. 1978b. Age composition of some vespertilionid bats as determin- 1979 ed by dental annuli. Canadian Journal of Zoology 56: 355-358. Smith, H. C. and D. B. Schowalter. 1979. A subspecies of Little Brown Bat new to the province of Alberta. Blue Jay 37: 58-62. Soper, J. D. 1964. The mammals of Alberta. Hamley Press, Edmonton. 402 pp. Studier, E. H.and M. J.O’Farrell. 1972. Biology of Myotis thysanodes and M. lucifugus (Chiroptera: Vespertilion- SCHOWALTER ET AL.: LITTLE BROWN BATS, ALBERTA 251 idae). I. Thermoregulation. Comparative Biochemistry and Physiology 41 A: 567-596. Studier, E. H., V. L. Lysengen, and M. J. O’Farrell. 1973. Biology and Myotis thysanodes and M. lucifugus (Chirop- tera: Vespertilionidae). II. Bioenergetics of pregnancy and lactation. Comparative Biochemistry and Physiology 44A: 467-471. Received 10 November 1978. Accepted 12 February 1979. Winter Denning of the Striped Skunk in Alberta JOHN R. GUNSON and RONALD R. BJORGE Alberta Fish and Wildlife Division, O.S. Longman Building, 6909 116 Street, Edmonton, Alberta T6H 4P2 Gunson, J. R. and R. R. Bjorge. 1979. Winter denning of the Striped Skunk in Alberta. Canadian Field-Naturalist 93(3): 252-258. Winter denning of the Striped Skunk (Mephitis mephitis) was studied in Alberta and adjacent Saskatchewan between 1971 and 1977. Most skunks denned under buildings and for periods up to 150 d. Skunks denned in communes and alone. Communal dens contained a number of female skunks (mean of 5.8 per den) and usually one adult male skunk. Solitary dens were occupied by males only, usually juveniles. Range of numbers of skunks in 61 communal dens was 2-19. Available comparative data suggest commune size increases with latitude, an adaptation to optimize winter survival and reproductive success. Age-specific selection seemed to operate in some communes with juvenile females excluded from certain dens. Juvenile males visited den sites in fall, but only rarely denned with female communes. Key Words: Mephitis mephitis, Striped Skunk, denning, communes, sex, age. The Striped Skunk (Mephitis mephitis) dens for extended periods during winter (Allen and Shapton 1942: Dean 1965; Verts 1967). More recent studies of denned skunks during winter have explored the potentials of disease transmission (Houseknecht 1969), activities (Sunquist 1974), physiological and behavioral adaptations (Mutch and Aleksiuk 1977), and habitat selection (Mutch 1977). We observed winter denning of skunks during 1971-1977 in studies of skunk rabies in Alberta and adjacent Saskatchewan (Gunson et al. 1978) and in an intensive population study of skunks in central Alberta (Bjorge 1977). This paper summarizes observations of den descriptions, numbers, sex and age, periods skunks spent in dens, and of movements to and from dens, and considers the adaptive significance of these aspects of winter denning of skunks near the northern limit of the species distribution. Study Area Most dens were located in the Alberta—Saskat- chewan border area where the bulk of rabies-related work was conducted (Figure 1). The area is aspen parkland (Bird and Bird 1967) in the northern half and prairie (Webb et al. 1967) in the southern half. It was settled between 50 and 100 yr ago and current land use has been in effect for many years. Cattle ranching and dry-land farming predominate in the short-grass prairie and cereal grain production and mixed- farming in the parkland. The climate of the study area is continental, characterized by warm summers and cold winters (Government and University of Alberta 1969). Variations in annual precipitation and number of frost-free days from north to south were = 28/18 cm and = 100/130 d. The 130-km? intensive study area (Tofield), located = at 53°23’N, 112°43’W, appeared typical of the aspen parkland. Eight major forested areas (mostly Trem- bling Aspen, Populus tremuloides), ranging 30-65 ha, occurred as well as numerous smaller areas with trees and brush along fence rows, road allowances, farm- yards, and wetlands. About 150 ponds, many of them seasonal, dotted the area. Predominant crops were cereal grains, forage, livestock, and poultry. Of 112 farmyards on the study area, 40 were abandoned. Methods Dens were located during six winters (1971-72 to 1976-77) primarily by visual examination of likely locations such as old buildings, culverts, and ground burrows. When located, skunks were removed from dens chiefly by carbon-monoxide gassing with two 2- cycle gasoline-operated motors, shooting (22-calibre rifle), and kill-trapping with mostly 220 Conibear traps. Collection of all skunks was often possible through removal of floorboards or occasionally by jacking or digging under foundations. Only dens - examined during November to March, and only those dens where collectors were convinced all skunks were captured, were used in calculations of numbers and of sex and age. Skunks were live-captured at Tofield during the non-winter period by trapping (National, Rudolph Skunker, and wooden and metal box traps were used), night-lighting (Jacobson 1969), and excavation of maternal dens. Live-captured skunks were handled according to the techniques of Jacobson et al. (1970) including netting, ear-tagging, weighing, sexing, and release. Radiotransmitters, similar to those used by Brand et al. (1975) and developed by personnel of the Department of Electrical Engineering, University of Alberta, Edmonton, were fitted to skunks. Signals were received via a truck-mounted non-directional whip antenna or a hand-held directional loop antenna and a portable receiver. Locations were determined by triangulation. DS) 1979 GUNSON AND BJORGE: STRIPED SKUNKS WINTER DENNING U3)3) | ALBERTA N 100 km | 62 mi x Edmonton ofield Parkland = ~ f 7 Calgary \ Wen f \ e \ e eye Pra \ 7 ee “e Ficure |. Locations of 61 communal dens of Striped Skunks in prairie and parkland portions of Alberta and adjacent Saskatchewan. Dental ages of collected skunks were determined by counts of cemental annuli in canines (Rakowski 1972; Casey and Webster 1975) after histological prepar- ation at Matson’s Audiovisual and Microscopic, Milltown, Montana. Live-captured young of the year were classified as juveniles until 31 December on the basis of weight, nipple development, and general appearance. Results and Discussion Location and Description of Dens Our searches for winter dens were directed prima- rily at rural, usually abandoned, buildings; thus the precise proportional use of buildings versus burrows, culverts, and other possible locations during winter was not determined. Of 73 communal ( >1 skunk) dens, 72 were located under buildings; one was in a culvert. Of 44 solitary (1 skunk) dens, 37 were under buildings, five were in culverts, and two were in ground burrows. Five of the six culvert dens were located in the southern prairie region. Both ground burrows were located at Tofield after extensive searching and tracking in snow during early winter 1971-72. The relatively large number of female skunks (see next section) under some buildings and the availability of other buildings without skunks, but used during previous winters, suggested that most female skunks, especially in the parkland, denned under buildings. During a depopulation at Tofield in December 1974, 19 (53%) of 36 female skunks tagged during June to October of that year were removed from six communal dens. Depopulation was incomplete at one den and skunks could not be removed from one other potential communal den because of its concrete structure. The above, and probable losses from death and emigration between summer and winter, support our belief that the majority of female skunks in the parkland den under buildings. Comparable recapture rate at buildings for males was only 11.9%, suggesting that many male skunks den in locations other than buildings. Of the two ground burrow dens located, one was used by a tagged male; the sex of the skunk in the other burrow was not determined. Related research (Gunson et al. 1978) implied relatively heavy skunk use of buildings during winter in the parkland versus prairie. Skunk depopulations by poisoning at buildings effectively reduced skunk populations in the parkland, but not in the prairie. Use of buildings as winter dens of skunks appeared greater in our study area than that recorded in more southerly areas. Allen (1939) and Storm (1972) reported that buildings were used only occasionally by skunks in Michigan and Illinois, respectively, It may 254 be that buildings were more available to skunks in our area than in other areas. Larger commune size and lower densities (unpublished data) of skunks in our area compared to more southerly areas probably favored the selection of building dens. A preference for building dens would result in greater proportions of skunks utilizing such dens in areas or periods of lower density. Building dens offer skunks more space for walking and use of latrines and for breeding during years of longer winters. Skunks made use of the space between ground and floorboards in most building dens. All winter dens that we observed contained dried vegetation, mostly grass. Ground burrows, usually associated with cellars, were present under some buildings. Construc- tion of burrows was likely necessary in some cellars to create a more confined den area. Numbers, Sex, and Age of Skunks in Dens A mean of 6.7 skunks was collected from 61 communal dens where collectors were convinced all skunks were taken (Table 1). Females were more common incommunal dens than males; in fact, no den contained more than one live male. Allen (1939) noted that only one male was present with denned females in his area. We obseved 43 dens of solitary males; denning by solitary females was not observed. Dens of solitary males and females were reported by Allen (1939) and Allen and Shapton (1942) in Michigan. TABLE 1—Composition of 61 communal winter dens of the Striped Skunk in Alberta and adjacent Saskatchewan (dens where all skunks were collected) Number of skunks Sex Total Mean (range)/den Females 353 5.8 (1-18) Males 53 0.9 (O- 1) Both sexes 406 6.7 (2-19) Somewhat more skunks were taken from communal dens in parkland versus prairie (means of 6.8 of 32 dens and 6.4 of 29 dens, respectively (Student’s t-test, P<0.4)). The largest commune was located in Township 57, the most northerly den we observed. Comparable data from other areas, although meagre, suggest commune size increases with latitude. Shirer and Fitch (1970) working in Kansas noted three skunks denned together, but for only short periods during winter. Numbers of skunks in dens was 2 to 11 in 10 dens (mean of 5.1) in Michigan (Allen 1939; Allen and Shapton 1942), two to six per den in Minnesota (Houseknecht 1969; Sunquist 1974), and THE CANADIAN FIELD-NATURALIST Vol-93 eight in one den in Colorado (Yeager and Woloch 1962). Larger communes in more northerly areas are likely an adaptation of the species to more rigorous climates. Mutch and Aleksiuk (1977) demonstrated that den temperatures vary directly with numbers of skunks in the den. It follows that females in larger groups would expend less energy in surviving longer winters, during which they do not feed, and conserve more energy for reproduction. Length of winter importantly influences production of skunks. Litters of young skunks at Tofield were significantly larger (Student’s t-test P<0.05) in 1973 (mean of 5.2) after a relatively mild winter compared to size of litters following the severe winter of 1973-74 (mean of 2.5) (Bjorge 1977). Juvenile (<1 year) females represented 47.7% of 327 aged females (Figure 2). In contrast, juvenile 70 60 50 Solitary Dens Males 40 n=38 0 20 10 50 Communal Dens Males n=51 50 is) PROPORTION (°%o) OF SAMPLE Communal Dens Females n=327 0 O ; : ; | =] = — 0.5 15 2.5 3.5 4.5 5.5 6.5 AGE-GROUP FIGURE 2. Age-structure of Striped Skunks in solitary and communal dens in Alberta and adjacent Saskatchewan. 1979 males comprised only seven (13.7%) of 51 aged males in communal dens. Most of the lone males were young; 26 (68.4%) of 38 aged solitary males were juveniles. No lone male was over 3 yr of age. Comparisons of age structures of skunks in pop- ulations during summer and fall and in communal winter dens at Tofield (Table 2) indicated that the juvenile proportion in the female cohort was some- what similar in dens and free-ranging skunks. Juvenile males, however, although common (> 60%) in male cohorts in populations, were absent in six communal dens during winter. In summary, winter dens of the Striped Skunk in our area included communal denning of females, usually with one adult male, and solitary denning of other males, including most juveniles. TABLE 2—Comparisons of age structures of Striped Skunks in six communal dens and in populations on the Tofield, Alberta study area Proportion (%) juveniles (n) Source Period Male Female Communal dens 1974-75 O0( 5) 47 ( 32) Population! 1974 66 ( 41) 35 ( 34) Population! 1971-74 62 (134) 37 (140) 'Live-trapped during June through October; excludes captures at communal dens in September and October. Den Formation Live-trapping at Tofield during September through October 1971-74, indicated that male skunks visited winter den sites other than the one they would occupy. From two to three male skunks were captured at each of six sites at which communes were forming during fall. Because only one male winters at each commune a male selection process must occur. Much of the selection might be a result of avoidance, probably juveniles tending to avoid more experienced adults. At each of two sites in 1974, adult and juvenile males were captured at the den site in fall; only the adults were present with the female commune in winter. Some competition, involving fighting, probably oc- curs during the fall pre-denning period. Three males with fresh scars on ears and face were observed during that period. Some unsuccessful (solitary) males visited com- munal dens during winter. A farmer poisoned a male skunk at one communal den in southern Alberta in late February. During the first week of March four live skunks, including one male, were collected at the den. We believe it highly probable that the second GUNSON AND BJORGE: STRIPED SKUNKS WINTER DENNING DES male joined the commune after the death of the former male. Tracks of skunks to and from dens were common during that collection period. At each of two other communal dens one live and one dead male skunk were collected. During a thaw in February 1972, at Tofield, a solitary male visited a commune 1 km from his burrow. Data from Tofield suggested that occupation of a specific communal den by females in at least some cases probably depended on age and experience. Five of seven females at one winter den at Tofield were adults, only one of which was a yearling. R.R.B. considered this area to have a higher density of skunks than most of the remainder of the Tofield area. At least four of the eight skunks at that winter den were captured and tagged during 1973 and 1974 at another den site used the previous three winters, but not in 1974-75. All or most of the group moved some 2.5 km to the new winter den during the fall pre-denning period in 1974. An adult male wintered alone at the original den. These observations suggest the com- mune was highly cohesive, and may explain the observed relatively low proportion of juveniles. Other young females may have been forced to den elsewhere. We noted that two other communes changed dens between years. Another communal den of seven females, located in an area R.R.B. considered to be of lowskunk density, consisted of six juveniles and one yearling. We collected some evidence that suggested these were not siblings. First, as discussed later, juvenile females at Tofield were highly mobile, travelling many kilo- metres to communal dens (Bjorge 1977). If such extensive dispersal of juvenile females is common it is unlikely that siblings would den at the same location. Second, of 62 young skunks, including 27 females, tagged in natal dens in 1973 and 1974, none was recaptured in communal dens on the study area even though survival of young was demonstrated at seven . of 15 dens. Survival was not recorded at the other natal dens. Allen (1939) concluded that winter groups of female skunks in his area were not families. Juvenile proportion of females in 56 communes was not bimodally distributed (Figure 3) as one might have expected from the selection process outlined above. Selection against juveniles at some dens could potentially create a situation of most dens with either low or high proportions of juveniles. Thirty-three (59%) of 56 dens contained < 30% or > 60% juveniles. Many of the dens in our sample were located in areas where skunks were being depopulated for rabies control. Because skunk sign was more abundant and more readily observed at communal dens than at solitary dens, skunks were more likely to be discov- ered and poisoned at communal dens. Higher than normal mortality at communal dens may account for 256 W Number of Dens u 0 11621 0«631)S 41—s“O51 4 vi at n | I | | I 10 20 30 40 50 60 70 80 90 100 Juvenile Proportion % FIGURE 3. Juvenile proportion of female Striped Skunks in 56 communal dens in Alberta and adjacent Saskatchewan. the variation between observed and expected distri- butions. Juveniles might be more readily accepted into dens of reduced size. Denning Period Success of live-capture at sites of communal dens at Tofield increased from 69 and 124/ 1000 trap-nights in July and August, respectively, to 275 in September; skunks began to harvest vegetation around den entrances in late September, as observed by Sunquist (1974) in Minnesota. This seemed to represent a pre- denning activity as skunks were still free-ranging during September and October, as evidenced by success of night-lighted captures along roads (Gunson et al. 1978) and by live-capture success at non-denning sites at Tofield. Adult skunks likely began denning before juveniles. Night-lighted captures of free- ranging skunks during late September and October were highly biased towards juveniles (unpublished data), and mean distances of adult females from communal dens at Tofield during September and October were significantly (Student’s t-test P< 0.05) less than for juveniles. Live-capture at communal den sites was most successful in October (306/ 1000 trap- nights). Sunquist (1974) noted that juveniles were more active than adults in November and initiated THE CANADIAN FIELD-NATURALIST Vol. 93 winter denning an average of 7 d later than adults. In 1973 activities of skunks beyond the immediate vicinity of winter dens at Tofield were terminated bya 25-cm snowfall on 10 November. Observations at 10 occupied dens during winter 1971-72 indicated that activity beyond a few metres of the den did not occur during early December to mid- February. A marked increase of activity was evident at six sites following a thaw on 12 February, with tracks leading from or to dens in four instances. Activity during mid-winter thaws was common during the study period, but evidently was mostly confined to males; of 13 road-killed skunks examined during December through February, 12 were males. Dean (1965) also noted the reactivation of winter- denned skunks during above-freezing temperatures. Emergence of skunks from two ground burrows occurred in mid-February and mid-March 1972. The latter emergence occurred in an area of snow accumulation and the emerging skunk had to burrow through approximately 50 cm of snow to leave the den. Movements of skunks from dens were common by mid-March. Males appear to leave dens earlier than females. Two skunks captured away from dens at Tofield in March were males. Preponderance of male skunks in fur harvest records (see Verts 1967) are likely explainable by greater and earlier activities of male skunks during winter and spring as recorded here. Dates of departure of six radio-collared adult female skunks from dens at Tofield were all between 2 and 9 April 1974. Because some of the six skunks may have denned before the 10 November snowfall (day 1), minimum period in winter dens during 1973-74, a winter of unusual severity, was 143 to 150 d. During the winter of 1971-72, a winter of moderate severity, most skunks were dénned by mid-November and skunks in communes were mostly inactive until mid- March, a period of approximately 120 d. These observations indicate that female skunks generally occupy winter dens in the parkland of Alberta between 120 and 150 d, and that males usually den for shorter periods. In Alberta the winter period is shorter in the prairie portion of the province and skunks in that area denned for correspondingly shorter periods. We observed numerous tracks leading from dens in southern areas of the province during March and male-female pairs of skunks were often observed at non-winter sites during that month. Skunks occupied winter dens 79-125 d in Minnesota (Sunquist 1974) and 62-87 d in Illinois (Verts 1967; Storm 1972). Movements to and from Winter Dens Juveniles travelled further than adults to winter dens; mean greatest distances from dens of 47 adult 1979 and 22 juvenile skunks during summer were 2.3 and 3.1 km, respectively. Adults tended to occupy the same general areas during consecutive summers (Bjorge 1977). Four adult females, monitored by telemetry from 46 to 85 d after leaving the den in spring, travelled greatest straight-line distances of 1.6 to 3.4 km. Two other females, monitored from July and August travelled 3.0 and 3.5 km to their winter dens. Movements of juvenile female skunks to winter dens were 13.7, 5.9, and 5.6 km from initial locations in July, September, and October, respectively. One other juvenile female travelled 21.7 km to a farmyard where she was shot before denning. Extensive dispersals of juveniles were recorded in another intensive study area in the prairie region in Alberta (P. Andersen, University of Calgary, personal commun- ication). The observed mobility of juveniles may be related to a search for acceptance into a winter commune. Concluding Discussion Carnivores have evolved a wide range of species- specific winter denning strategies varying from the mostly solitary denning of bears to the highly communal denning of female Striped Skunks as reported here. Social organization of denning would appear to be importantly related to reproduction in carnivores. European Badgers (Meles meles) and American Badgers (Taxidea taxus) apparently den either in pairs or solitarily (Neal 1948; Ewer 1973; Lindzey 1978). Breeding of badgers and bears occurs during summer (see Ewer 1973), and is followed by delayed implantation; thus, communal denning in these species would not enhance reproduction. Because Striped Skunks breed during or immediately after winter denning, reproduction is potentially more successful where numbers of females are available to at least one male. Ewer (1973) noted that Verts observed one male would breed several captive female skunks in short order; such behavior would ensure that all females become pregnant. Pregnancy rates of Striped Skunks in Alberta were high during the study period (unpublished data). Communal winter denning is common in Raccoons (Procyon lotor), although group size is usually small (Ewer 1973), despite one den of 23 individuals (Mech and Turkowski 1966). Family groups of Raccoons den together (Whitney and Underwood 1952). Our observations suggest that family groups of Striped Skunks in Alberta do not often den together. Genetic variation would be maintained by family separation in areas where a significant proportion of breeding occurs during winter denning. In summary, communal denning appears to have adaptive value for winter survival and reproductive GUNSON AND BJORGE: STRIPED SKUNKS WINTER DENNING a3} success of Striped Skunks, especially in more northern latitudes where the period of inactivity and fasting is longer, where energy conservation can be critical, and where breeding may be influenced by winter distribution. Acknowledgments This study was supported by the Veterinary Services Division, Alberta Department of Agriculture and by the Fish and Wildlife Division, Alberta Department of Recreation, Parks and Wildlife. The Saskatchewan Department of Natural Resources (now Tourism and Renewable Resources) kindly provided permission to collect skunks in adjacent Saskatchewan. The capable technical assistance of P. Andersen, P. Cole, W. Etherington, E. Ewaschuk, W. Johnson, L. Pecharsky, D. Pipella, D. Schowalter, and W. Wynnyk is acknowledged. D. Schowalter kindly reviewed earlier drafts of this paper. W. M. Samuel of the University of Alberta supervised much of the Tofield studies during 1973 and 1974. Literature Cited Allen, D. L. 1939. Winter habits of Michigan skunks. Journal of Wildlife Management 3(3): 212-228. Allen, D. L. and W.W. Shapton. 1942. An ecological study of winter dens with special reference to the eastern skunk. Ecology 23(1): 59-68. Bird, C. D. and R. D. Bird. 1967. The Aspen parkland. Jn Alberta, A natural history. Edited by W.G. Hardy. Hurtig Publishers, Edmonton, Alberta. pp 135-149. Bjorge, R.R. 1977. Population dynamics, denning and movements of Striped Skunks in central Alberta. M.Sc. thesis, University of Alberta, Edmonton. 96 pp. Brand, C. J., R. H. Vowels, and L. B. Keith. 1975. Snow- shoe Hare mortality monitored by telemetry. Journal of Wildlife Management 39(4): 741-747. Casey, G.A. and W.A. Webster. 1975. Age and sex. determination of Striped Skunks (Mephitis mephitis) from Ontario, Manitoba and Quebec. Canadian Journal of Zoology: 223-226. Dean, F. C. 1965. Winter and spring habits and density of Maine skunks. Journal of Mammalogy 46(4): 673-675. Ewer, R. F. 1973. The carnivores. Weidenfeld and Nicolson, London. 494 pp. Government and The University of Alberta. 1969. Atlas of Alberta. University of Alberta Press. 158 pp. Gunson, J. R., W. J. Dorward, and D. B. Schowalter. 1978. An evaluation of rabies control in skunks in Alberta. Canadian Veterinary Journal 19: 214-220. Houseknecht, C. R. 1969. Denning habits of the Striped Skunk and the exposure potential for disease. Bulletin of the Wildlife Diseases Association 5: 302-306. Jacobson, J. O. 1969. Application of a nighttime roadside census to Striped Skunk population studies. M.Sc. thesis, North Dakota State University, Fargo, North Dakota. 83 Pp: 258 Jacobson, J. O., E. C. Meslow, and M. F. Andrews. 1970. An improved technique for handling Striped Skunks in disease investigations. Journal of Wildlife Diseases 6: 510-512. Lindzey, F. G. 1978. Movement patterns of badgers in northwestern Utah. Journal of Wildlife Management 42(2): 418-422. Mech, L. D. and F. J. Turkowski. 1966. Twenty-three Rac- coons in one winter den. Journal of Mammalogy 47(3): 529-530. Mutch, G. R. P. 1977. Locations of winter dens utilized by Striped Skunks in Delta Marsh, Manitoba. Canadian Field-Naturalist 91(3): 289-291. Mutch, G. R. P. and M. Aleksiuk. 1977. Ecological aspects of winter dormancy in the Striped Skunk (Mephitis mephitis). Canadian Journal of Zoology 55: 607-615. Neal, E. 1948. The badger. Collins, London. Rakowski, P. W. 1972. Studies on the Striped Skunk in southeastern North Dakota. M.Sc. thesis, North Dakota State University, Fargo. 62 pp. Shirer, H. W. and H.S. Fitch. 1970. Comparison from radio-tracking of movements and denning habits of the THE CANADIAN FIELD-NATURALIST Vol. 93 Raccoon, Striped Skunk and Opossum in northeastern Kansas. Journal of Mammalogy 51(3): 491-503. Storm, G.L. 1972. Daytime retreats and movements of skunks on farmlands in Illinois. Journal of Wildlife Management 36(1): 31-45. Sunquist, M. E. 1974. Winter activity of Striped Skunks (Mephitis mephitis) in east-central Minnesota. American Midland Naturalist 92(2): 434-446. Verts, B. J. 1967. The biology of the Striped Skunk. University of Illinois Press, Urbana. 288 pp. Webb, R., A. Johnston, and J. D. Soper. 1967. The prairie world. Jn Alberta, A natural history. Edited by W.G. Hardy. Hurtig Publishers, Edmonton, Alberta. pp. 93- 115. Whitney, L. F. and A.B. Underwood. 1952. The Rac- coon. Practical Science Publishing Company, Orange, Connecticut. Yeager, L. E. and J. P. Woloch. 1962. Striped Skunk with three legs. Journal of Mammalogy 43(3): 420-421. Received December 1978 Accepted 5 March 1979 Range Extensions of Vascular Plants in Northern Yukon Territory and Northwestern District of Mackenzie JOHN A. NAGY,! 3 ARTHUR M. PEARSON,! 4 BERNARD C. GOSKI,! 5 and WILLIAM J. Coby? 1Canadian Wildlife Service, Environment Canada, Edmonton, Alberta T5K 2J5 2Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 3Present Address: Lethbridge Community College, Lethbridge, Alberta TIK 1L6 _ 4Present Address: Government of the Yukon Territory, Whitehorse, Yukon Y1IA 2C6 5Present Address: Alberta Fish and Wildlife Division, Edson, Alberta TOE OPO Nagy, John A., Arthur M. Pearson, Bernard C. Goski, and William J. Cody. 1979. Range extensions of vascular plants in northern Yukon Territory and northwestern District of Mackenzie. Canadian Field-Naturalist 93(3): 259-265. Between 1970 and 1975 plant communities in the Yukon Territory and the District of Mackenzie north of 67°N were investigated. We record three taxa new to the known flora of the Yukon Territory and one new to the flora of the District of Mackenzie. Among the other vascular plants listed here, nine are new to the range predicted by Hultén within the Yukon Territory, 23 are extensions within predicted ranges and thus corroborate these predictions, and 29 are extensions beyond predicted ranges. Key Words: vascular plants, Yukon Territory, District of Mackenzie, flora. Between 1970 and 1975 over 800 collections of — tics Research Institute, Canada Agriculture, Ottawa, vascular plants, representing 220 species, 53 sub- Ontario. species and seven varieties, were made during Cana- dian Wildlife Service studies in the northern Yukon Collection Sites Territory. Hultén (1968), Welsh and Rigby (1971), Approximate collection site locations are shown in and Wein et al. (1974) have reported extensive Figure 1. Other site data are listed below. collections and recent range extensions for vascular 1) Sam Lake, 68°25’N, 138°35’W, elevation ca. plants from our study area. We record here additional 427 m (1400 ft). Marshy polygonal; dry-to-wet species to the reported flora of the Yukon Territory tundra and hummocky lakeside areas; marshy and northwestern District of Mackenzie. We also note and wet shrubby lakeside areas. species that occur outside of the ranges predicted by 2) Canoe River, 68°37’N, 138°43’W, elevation ca. Hultén (1968), or were collected for the first time in 305 m (1000 ft). Wet sedge meadows west of northern Yukon Territory within ranges predicted. In river. addition, a number of species are recorded as the 3) Ridge 1.6km east of Sam Lake, 68°24’N, northernmost or southernmost collections within 138°33’W, elevation ca. 488 m (1600 ft). Dry ranges from which collections are known. rocky and moist-to-wet tundra on ridge; dry The study area and collection locations are shown south- and north-facing scree slopes; wet snow in Figure 1. The 31 collection sites from which plant bed community near drainage. specimens were obtained ranged in elevation from 122 4) Anker Creek, 68°43’N, 137°33’W, elevation ca. to 152 masl on the Arctic Coastal Plain, 244 to 122 m (400 ft). Alder-willow bluffs on moist-to- 427 masl on the Old Crow Flats, and 152 to wet south-facing slope; dry rocky talus slope. 1067 masl in the Barn, British, and Richardson 5) Trail River, 68°57’N, 138°56’W, elevation ca. mountains. Bostock (1961) and Welsh and Rigby 152 m (500 ft). South-facing river bank. (1971) describe the physiography of the area. 6) Headwaters of Anker Creek, 68°30’N, 138°19’W, Our activities were primarily concentrated in areas elevation ca. 945 m (3100 ft). Alpine slope and that had not received attention from previous investi- tundra. gators. Collectors were A. M. Pearson 1970, 1973, 7) 6.4km northwest of Sam Lake, 68°28’N, 1974, 1975; J. A. Nagy 1974, 1975; M. Dennington 138°40’W, elevation ca. 457 to 762 m (1500 to 1975; B. C. Goski 1973, 1974: D. Pearson 1974; C. B. 2500 ft). Areas of dry, rocky and moist-to-wet Larsen 1973, 1974; J. W. Nolan 1973. The collections tundra; moist-to-wet and swampy areas in and made during 1970 and 1973 were identified by A. E. along edge of White Spruce ( Picea glauca) stand; Porsild, National Museum of Natural Sciences, willows (Salix spp.) in and along edge of White Ottawa, and those collected during 1974 and 1975, by Spruce stand; grassy slopes on upper edge of W. J. Cody and G. A. Mulligan (Draba), Biosystema- White Spruce stand; dry rocky ridge and slope. 259) 260 THE CANADIAN FIELD-NATURALIST Vol. 93 0 kilometres ©O BEAUFORT SEA SS Oo miles 30 Herschel MACKENZIE BAY 68° { «e eg 40° (38° 136° FicurE |. Locations of 31 vascular plant collection sites in the northern Yukon Territory and northwestern District of Mackenzie. 1979 8) 9) 10) 11) 12) 13) 14) 15) 16) 17) 18) 19) 20) 21) 22) 23) 24) 25) 26) 2i)) Ridge 1.6km (1 mi) north of Sam Lake, G8S225UNe 1382350W, elevation’ “cae. 58m (1700 ft). Dry rocky slopes. Dog Creek, 68°25’N, 138°38’W, elevation ca. 427 m (1400 ft). Willows along drainage; and tundra. Headwaters of Boulder Creek, 68°26’N, 138°13’W, elevation ca. 488 m (1600 ft). Moist- to-dry rocky south-facing slopes; stabilized south-facing talus slope; mossy wet stream bed; moist tundra along creek. Ridge above Boulder Creek, 68°25’N, 138°16’W, elevation ca. 914 m (3000 ft). Dry rock south- facing slope. Tributary of Babbage River, 68°38’N, 138°01’W, elevation ca. 427 m (1400 ft). Wet mossy stream bank; wet streamside willow stand; dry south- facing slope. Tributary of Fitton Creek, 68°33’N, 138°10’W, elevation ca. 457 m(1500 ft). Moist drainage; dry rocky ridge. Trail River, 68°58’N, 138°58’W, elevation ca. 396 m(1300 ft). Moist drainage; dry south-facing slope. Spruce Creek, 68°56’N, 138°41’W, elevation ca. 305 m (1000 ft). White Spruce stand on dry south-facing slope. Old Crow Flats, 68°09’N, 139°35’W, elevation ca. 274 m (900 ft). Bog. Old Crow Flats, 68°12’N, 139°49’W, elevation ca. 305 m (1000 ft). White Spruce stand on dry ridge; marshy area. Old Crow Flats, 68°12’N, 139°48’W, elevation ca. 305 m (1000 ft). Sphagnum bog. Old Crow Flats, 68°01’N, 139°48’W, elevation ca. 244 m (800 ft). Sphagnum bog. Headwaters of Babbage River, 68°38’N, 139°32’W, elevation ca. 457 m(1500 ft). Willows along stream bank; tundra. Margaret Lake, 68°50’N, 140°36’W, elevation ca. 457 m (1500 ft). Tundra along Firth River. Babbage Falls, 68°43’N, 139°03’W, elevation ca. 274 m (900 ft). Rock crevices along falls; dry rocky south-facing slope. Bell River, 67°43’N, 136°33’W, elevation ca. 427 to 457 m (1400 to 1500 ft). Scree slope; White Spruce — White Birch (Betula papyrifera) — willow stand; sphagnum bog; moist slope. Firth River, 69°20’N, 139°33’W, elevation ca. 152 m (500 ft). North-facing rock outcrop. Boulder Creek, 68°25’N, 138° 15’W, elevation ca. 457 m (1500 ft). Moist creek-side areas near headwaters. Sam Lake, 68°30’N, 138°35’W, elevation ca. 762 m (2500 ft). Dry hilltop. Mount Sedgwick, 68°58’N, 139°01’W, elevation NAGY ET AL.. NORTHERN VASCULAR PLANTS 261 ca. 305 m (1000 ft). Dry slope near ridge top; moist creek-side area. Old Crow, 67°35’N, 139°50’W, elevation ca. 250 m (820 ft). Roadside near airstrip. Rapid River, 68°16’N, 137°08’W, elevation ca. 914m (3000 ft). Sedge association in thick organic soil on moist south-facing rock slide. Margaret Lake, 68°48’N, 140°36’W, elevation ca. 457 m (1500 ft). Lakeside sphagnum bog. Bear Creek, 67°58’N, 136°12’W, elevation ca. 1067 m (3500 ft). Moist basin. 28) 29) 30) 31) Vascular Plant Range Extensions The 65 taxa reported here are listed with collection sites, habitats, collection years, and collection numbers. Nomenclature follows Hultén (1968) and if names used by Porsild and Cody (1968) differ, they are also given, in parentheses. A complete set of voucher specimens has been deposited in the Cana- dian Wildlife Service Herbarium at Edmonton, with duplicates of the 1974 collections deposited at the Canada Agriculture Herbarium in Ottawa (DAO). Three taxa, Luzula Wahlenbergii ssp. Wahlen- bergii, Potentilla rubricaulis, and Galium Brandegei are reported as new to the flora of the Yukon Territory, and one species, Cerastium maximum, as new to the flora of the Continental Northwest Territories. The new range data provided here on the taxa listed represents a contribution to our knowledge of the phytogeography of these taxa, as well as a contribu- tion towards a flora of Yukon Territory. Lycopodiaceae Lycopodium annotinum ssp. pungens (Stiff Club-moss). Site 1: shrubby lakeshore, 74-58; site 4; alder, 74-189; site 5: river bank, 74-10; site 23: sphagnum bog, 74-683. Helps complete gaps in the predicted distribution between sites in northwestern District of Mackenzie and Alaska, extending the known range northward to the Arctic Coastal Plain (Hultén 1968). Lycopodium complanatum (Ground-Cedar). Site 23: mixed White Spruce — birch — willow, 74-68]. Extends the known range northward from collection localities reported by Hultén (1968) and Wein et al. (1974) along the Yukon River drainage. Polypodiaceae Cystopteris fragilis ssp. fragilis (Fragile Fern). Site 22: crevices along falls, 74-675. Hultén (1968) predicted the occurrence of both ssp. fragilis and ssp. Dickieana in northern Yukon Territory and did plot one collection from Yukon Territory north of 65°N, along the Yukon River drainage near the Alaska border. Welsh and Rigby (1971) reported three specimens from the British Mountains, but did not give their collections a subspecific designation. The specimen reported here is the first record of ssp. fragilis 262 within the predicted range for northern Yukon Territory (Hultén 1968). Dryopteris fragrans (Fragrant Cliff-Fern). Site 5: river bank, 74-13; site 14: south-facing slope, 74-435. Welsh and Rigby (1971) reported the first collection from the northern Yukon Territory. The specimens reported here are the most northern collections, and help complete gaps in the predicted distribution for northern Yukon Territory (Hultén 1968). Sparganiaceae Sparganium minimum (Bur-reed). Site 18: emergent aquatic, 74-556. Extends the known range significantly northward from collection localities in west central Yukon Territory to the Old Crow Flats (Hultén 1968). Potamogetonaceae Potamogeton alpinus ssp. tenuifolius. Site 18: sphagnum bog, 74-550. Extends the known range to the northwest of Welsh and Rigby’s (1971) recent first collection for Yukon Territory north of 65°N. Potamogeton perfoliatus ssp. Richardsonii (P. Richard- sonii) (Red-head Pondweed). Site 16: submerged aquatic, 74-511. First record from the predicted range for the Yukon Territory north of 65° N (Hultén 1968). Helps bridge the gap in the distribution between collection sites in Alaska and northwestern District of Mackenzie. Gramineae Calamagrostis canadensis ssp. Lansdorffii (Blue-joint). Site 18: sphagnum bog, 74-549; site 25: creek bank, 73-/, 73-3. Hultén (1968) predicted the occurrence of both ssp. canadensis and ssp. Langsdorffii in northern Yukon and reported collections of both subspecies adjacent to the Yukon Territory border. Welsh and Rigby (1971) reported a specimen of C. canadensis from the Old Crow Flats, but did not give their collection a subspecific designation. This is the first report of ssp. Langsdorffii from the predicted range for Yukon Territory north of 65°N. Calamagrostis purpurascens. Site 21: tundra, 74-667. Hultén (1968) cited collections from adjacent areas in Alaska and northwestern District of Mackenzie, while Welsh and Rigby (1971) reported the first collections from northern Yukon Territory. The specimen reported here is the most northern collection within the predicted range for Yukon Territory (Hultén 1968). Festuca ovina ssp. alaskana. Site 8: dry rocky ridge, 74-320. Hultén (1968) reported a collection from the Yukon River drainage near the District of Mackenzie border, and Cody has collected it in the Richardson Mountains to the east in District of Mackenzie (Cody and Porsild 1968). The specimen reported here is the most northern collection in the predicted range in Yukon Territory. Cyperaceae Carex glacialis. Site 7: rocky south-facing slope, 74-254. First record within the predicted range for Yukon Territory north of 65°N (Hultén 1968). Carex lapponica (C. canescens var. subloliacea). Site 19: sphagnum bog, 74-560. Extends the known range signifi- cantly northward from central Yukon Territory to the Old Crow Flats (Hultén 1968). THE CANADIAN FIELD-NATURALIST Vol. 93 Carex membranacea. Site 6: alpine tundra, 74-346; Site 7: White Spruce stand, 74-615; site 7: dry rocky ridge, 74-288; site 20: tundra, 74-586a. Hultén (1968) plotted collection localities from adjacent areas in Alaska and District of Mackenzie. The specimens reported here are the first collections from the predicted range for Yukon Territory north of 65°N. Carex Williamsii. Site 16: bog, 74-508. First reported for northern Yukon but is from south of the range predicted by Hultén (1968) for the northern Yukon Territory, and represents only the fourth collection for Yukon Territory (Porsild 1975). Eriophorum angustifolium ssp. triste (E. triste). Site 6: alpine tundra, 74-347; site 7: dry rocky ridge, 74-118. Hultén (1968) indicated a collection from the Firth River drainage in Alaska near the Alaska border. The specimens reported here are the first collections from the predicted range for Yukon Territory north of 65°. Eriophorum russeolum var. albidum. Site 1: marsh, 74-75. Hultén (1968) cited collection sites from adjacent areas in Alaska and northwestern District of Mackenzie. The specimen reported here is the first collection from the range predicted by Hultén (1968) for northern Yukon Territory- Porsild (1951, 1975) has reported it from along the Canol Road and the Ogilvie Mountains. Araceae Calla palustris (Wild Calla). Site 18: sphagnum bog, 74-546; site 28: roadside, 73-8. Hultén (1968) reported a collection from the headwaters of the Yukon River. Our collection extends the known range northwestward to the Old Crow Flats. Lemnaceae Lemna trisulca (Star-Duckweed). Site 18: floating aquatic in sphagnum bog, 74-554. Northernmost collection within the range predicted by Hultén (1968) for Yukon Territory. Juncaceae Juncus articus ssp. alaskanus (J. balticus var. alaskanus). Site 21: riverside tundra, 74-664. Welsh and Rigby (1971) reported the first collection (presumably var. alaskanus) of J. arcticus from northern Yukon Territory, but did not give their specimen a subspecific designation. This is thus only the second record of ssp. alaskanus within the range predicted by Hultén (1968) for Yukon Territory north of 65°N. Luzula arctica (L. nivalis). Site 10: south-facing slope, 74- 170. First report from the predicted range for Yukon Territory north of 65°N (Hultén 1968) and helps bridge the gap between collection localities in Alaska and northwestern District of Mackenzie. Luzula arcuata s. lat. Site 13: drainage, 74-383. First collection from Yukon Territory north of 65° N, and is from near the northern limit as predicted by Hultén (1968). Luzula multiflora s. lat. Site 3: dry rocky hillside, 74-633; site 7: tundra, 74-24]. First collections within the range predicted by Hultén (1968) for Yukon Territory north of 65°N. Luzula parviflora. Site 12: stream bank, 74-2/2. First collection from Yukon Territory north of 66°N, extending the known range northward to the Barn Mountains (Hulten 1968). 1979 Luzula Wahlenbergii ssp. Wahlenbergii. Site 1: polygonal ground, 74-269. Hultén (1968) reported a collection from northwestern District of Mackenzie adjacent to the Yukon Territory border. The specimen reported here is the first collection from the range predicted by Hultén (1968) for Yukon Territory and is new to the Territory. Orchidaceae Spiranthes Romanzoffiana (Hooded Ladies’-Tresses). Site 7: bog, 74-503; sphagnum bog, 74-56]. First collection reported from Yukon Territory north of 65° N, extending the predicted range westward from collection localities in northwestern District of Mackenzie to northern Yukon Territory (Hultén 1968). Salicaceae Salix Barrattiana. Site 14: drainage, 74-463. First specimen collected in northern Yukon Territory, extending the predicted range eastward from collection sites in Alaska (Hultén 1968). Caryophyllaceae Cerastium maximum. Site 31: alpine basin, 73-46. Weinetal. (1964) extended the range of C. maximum eastward to the headwaters of the Yukon River from near the Alaska border (Hultén 1968). The collection reported here extends the range further eastward to the Richardson Mountains of northwestern District of Mackenzie. This species is thus new to the flora of Continental Northwest Territories (Porsild and Cody 1968). Silene acaulis ssp. subacaulescens (Moss Campion). Site 7: alpine tundra, 74-341; site 30: sphagnum bog, 73-5. Hultén (1968) plotted a collection locality along the Firth River drainage near the Alaskan border. The specimens reported here are the first collections from the predicted range for Yukon Territory north of 65°N (Hultén 1968). Ranunculaceae Anemone narcissiflora Ss. lat. Site 29: sedges on rock slide, 73- 1. Hultén (1968) indicates several collection sites of A. narcissiflora ssp. interior from the District of Mackenzie adjacent to the Yukon Territory border. Our collection is the first report from the predicted range for Yukon Territory north of 65°N. Ranunculus Eschscholtzii. Site 25: creek bank, 73-9. Hultén (1968) reported a collection from District of Mackenzie adjacent to Yukon Territory. The specimen reported here is the first collection from Yukon Territory north of 65°N, extending the known range westward from northwestern District of Mackenzie to the Barn Mountains. Papaveraceae Papaver Hultenii. Site 24: rock outcrop, 75-/. First collection within the predicted range for northern Yukon Territory (Hultén 1968). Papaver Macounii (P. Keelei). Site 7: White Spruce stand, 74-618; site 10: south-facing slope, 74-161; site 27: creek bank, 70-22; site 30: sphagnum bog, 73-4. Hultén (1968) indicated collection localities from adjacent areas in Alaska and northwestern District of Mackenzie. These are the first specimens collected in the predicted range for Yukon Territory north of 65°N (Hultén 1968). NAGY ET AL.: NORTHERN VASCULAR PLANTS 263 Cruciferae Cardamine umbellata. Site 1: lakeshore, 74-641. First collection from Yukon Territory north of 65°N, extending the known range significantly northward to the Barn Mountains (Hultén 1968). Draba alpina. Site 7: dry rocky ridge, 74-100. Welsh and Rigby (1971) reported a collection from the Firth River drainage, within the range predicted by Hultén (1968). Our collection extends the known range southeastward from this station. Draba borealis. Site 24: rock outcrop, 75-4. Hultén (1968) plotted an isolated collection in southwestern Yukon Territory from about 137° W. A more recent map is given by Mulligan (1970). The specimen reported here is the first collection from northern Yukon Territory, and extends its known range significantly northward from about 63°N. Draba crassifolia. Site 24: rock outcrop, 75-9. Hulten (1968) reported an isolated collection from the Mackenzie River drainage adjacent to northern Yukon Territory. A more recent map is given by Mulligan (1975). The specimen reported here is the first collection from Yukon Territory north of 65°N, extending the known range westward. Draba incerta. Site 7: dry rocky ridge, 74-98; site 10: south- facing slope, 74-158, 74-164; site 13: dry rocky ridge, 74-404. Hulten (1968) plotted three collections from southern Yukon Territory in close proximity to the British Columbia border. A more extensive map is given by Mulligan (1972). The collections reported here extend the known range signifi- cantly northward to northern Yukon Territory. Droseraceae Drosera rotundifolia (Round-leaved Sundew). Site 18: sphagnum bog, 74-543. Hultén (1968) reported an isolated collection from the Mackenzie River drainage. Our specimen is the first collection from Yukon Territory north of 65°N, extending the known range northward to the Old Crow Flats. Saxifragaceae Saxifraga caespitosa (Tufted Saxifrage). Site 11: dry rocky ridge, 74-183. Extends the known range southeastward from the recent first collections reported from northern Yukon Territory by Welsh and Rigby (1971) and Wein et al. (1974). Saxifraga hieracifolia. Site 21: tundra, 74-587. Hultén (1968) and Wein et al. (1974) reported collections from localities along north coastal Yukon Territory. The collection reported here extends the known range slightly southward along the Firth River drainage. Ribes triste (Red Currant). Site 4: alder stand on river bank, 74-190. Previously reported from the Old Crow Flats and northwestern District of Mackenzie by Hultén (1968). The specimen reported here extends the known range northward to the arctic coastal plain. Rosaceae Dryas octopetala L. ssp. alaskensis (D. alaskensis). Site 7: White Spruce stand, 74-82; site 27: dry ridge, 70-22. Wein et al. (1974) reported the first collection of this species for northern Yukon Territory from the headwaters of the Yukon River drainage. The collections reported here extend the 264 known range northward and bridge the gap between collection localities in Alaska and northwestern District of Mackenzie (Hultén 1968). Potentilla elegans. Site 3: dry rocky slope, 74-36; site 6: alpine tundra, 74-350; site 11: dry rocky ridge, 74-179, site 13: drainage, 74-377. Extends the known range eastward from the recent extensions reported by Welsh and Rigby (1971) and Wein et al. (1974). Potentilla hyparctica. Site 11: dry rocky ridge, 74-184; site 14: south-facing slope, 74-453. First collections reported for Yukon Territory north of 65°N, extending the known range eastward from localities in Alaska (Hultén 1968). Potentilla Hookeriana ssp. Hookeriana (P. nivea ssp. Hookeriana). Site 22: rock crevices, 74-671. Hultén (1968) reported collections of P. Hookeriana from coastal northern Yukon Territory and the Old Crow Flats. The collection reported here extends the known distribution southeastward within the predicted range (Hulten 1968). Potentilla rubricaulis. Site 3: dry rocky ridge, 74-24, 74-237; site 7: dry rocky ridge, 74-103, 74-114, 74-257, 74-289, alpine tundra, 74-334; site 8: dry rocky slope, 74-70; site 10: south-facing slope, 74-154, 74-168. First collections from Yukon Territory, expanding the known range west- ward from northwestern District of Mackenzie (Hultén 1968). Rubus arcticus ssp. acaulis (R. acaulis). Site 16: bog, 74-504. Hultén (1968) reported collections from adjacent areas in Alaska and northwestern District of Mackenzie. The specimen reported here is the first collection from the predicted range for Yukon Territory north of 65°N (Hultén 1968). Sibbaldia procumbens. Site 6: alpine tundra, 74-360. Hultén (1968) reported an isolated collection from northwestern District of Mackenzie. The specimen reported here is the first collection from Yukon Territory north of 65° N, and extends the known range westward to the Barn Mountains from the Richardson Mountains in northwestern District of Mac- kenzie. Leguminosae Oxytropis borealis (O. glutinosa). Site 24: rock outcrop, 75-18. First report from the range predicted for northern Yukon Territory, and helps bridge the gap in its distribution between localities in northwestern District of Mackenzie and Alaska (Hultén 1968). Oxytropis deflexa var. sericea. Site 16: bog, 74-496. First collection from Yukon Territory north of 65°N, extending the known range northward to the Old Crow Flats (Hultén 1968). Oxytropis nigrescens ssp. pygmaea ( O. pygmaea). Site 7: dry rocky ridge, 74-55, 74-109; site 11: dry rocky slope, 74-174; site 26: dry hilltop, 70-20h; site 29: sedge association on rock slide, 73-0. Hultén (1968) reported several collec- tions from the Alaskan coast adjacent to the Yukon Territory border. The collections reported here extend the known range across northern Yukon Territory. Pyrolaceae Moneses uniflora (One-flowered Pyrola). Site 7: White THE CANADIAN FIELD-NATURALIST Vol. 93 Spruce stand, 74-605. Hultén (1968) plotted a collection site along the Yukon River drainage near the Alaska border. Our collection extends the known range northward to the Barn Mountains. Pyrola secunda ssp. obtusata (One-sided Pyrola). Site 7: White Spruce stand, 74-617; site 16: bog, 74-493; site 17: White Spruce stand, 74-525. Hultén (1968) indicated a collection site from the Firth River drainage along the Alaska border. The specimens reported here are the first collections from the predicted range for northern Yukon Territory. Ericaceae Chamaedaphne calyculata (Leather-leaf). Site 19: sphagnum bog, 74-562. Hultén (1968) plotted a collection site along the Yukon River drainage near the Alaska border. The collection reported here extends the known range north- ward, well onto the Old Crow Flats. Ledum palustre ssp. groenlandicum (L. groenlandicum) (Labrador-tea). Site 15: White Spruce stand, 74-474. Hultén (1968) reported a collection from the Yukon River drainage near the village of Old Crow. Our collection extends the known range northward to the Barn Mountains. Boraginaceae Eritrichium splendens. Site 7: dry rocky ridge, 74-261, 74-306. Hultén (1968) and Wein et al. (1974) reported specimens from the Yukon River drainage near the Alaska and District of Mackenzie borders, respectively. The collections cited here extend the known range northward to the Barn Mountains in northern Yukon Territory. Scrophulariaceae Pedicularis lapponica. Site 1: tundra, 74-199, 74-234; site 9: tundra, 74-222; site 26: dry hilltop, 70-20L. Hultén (1968) reported a collection from the Old Crow Flats in northern Yukon Territory. The collections reported here extend the known distribution northward to the Barn Mountains, but still fall within the predicted range (Hultén 1968). Lentibulariaceae Pinguicula villosa. Site 1: polygonal ground, 74-268; site 18: sphagnum bog, 74-552; site 26: dry ridge, 70-20m. Most northern collections within the range predicted by Hulten (1968). He had plotted a collection site on the Yukon River drainage near the southern edge of the Old Crow Flats. Utricularia vulgaris (Bladderwort). Site 18: submerged aquatic, 74-555. First collection from the predicted range for Yukon Territory north of 65°N (Hulten 1968). Rubiaceae Galium Brandegei. Site 19: sphagnum bog, 74-558. First collection from the predicted range for the Yukon Territory and is new to the Territory. Helps bridge the gap in the distribution between collection localities in Alaska and northwestern District of Mackenzie (Hultén 1968). Caprifoliaceae Linnaea borealis ssp. americana (Twinflower). Site 3: dry rocky slope, 74-630. first collection from its predicted range for Yukon Territory north of 65°N. 1979 Compositae Aster alpinus ssp. Vierhapperi. Site 3: dry rocky ridge, 74-283; site 7: dry rocky ridge, 74-260, 74-272, 74-293. Wein et al. (1974) reported a recent range extension from the Yukon River drainage in northern Yukon Territory adjacent to the District of Mackenzie border. The collections reported here extend the known range northward to the Barn Mountains. Antennaria Friesiana ssp. compacta (A. neoalaskana). Site 3: dry rocky ridge, 74-239; site 7: dry rocky ridge, 74-113; site 8: dry rocky ridge, 74-3/1b; site 10: south slope, 74-144; site 13: dry rocky ridge, 74-405, 74-4179. Hultén (1968) reported a collection from the Old Crow Flats. The specimens reported here are the northernmost collections within the range predicted by Hultén (1968) for northern Yukon Territory. Senecio atropurpureus ssp. tomentosus (Kjellm.) Hult. (S. Kjellmannii). Site 11: dry rocky ridge, 74-186. First known collection from its predicted range in Yukon Territory north of 65°N, and helps bridge the gap in the distribution between collection localities in Alaska and District of Mackenzie. Taraxacum phymatocarpum. Site 7: rocky alpine tundra, 74-105, 74-325, 74-338. Hultén (1968) reported a collection from Herschel Island. These collections extend the known distribution within the predicted range, southward to the Barn Mountains. Acknowledgments The research was undertaken as part of a broader study of the ecology of the arctic mountain Grizzly Bear under the auspices of the Canadian Wildlife Service. We thank Anne Gunn, Canadian Wildlife Service, who read the manuscript and provided helpful suggestions. NAGY ET AL.:. NORTHERN VASCULAR PLANTS 265 Literature Cited Bostock, H.S. 1961. Physiography and resources of the northern Yukon. Canadian Geographical Journal 63: 112-119. Cody, W. J.and A. E. Porsild. 1968. Additions to the flora of Continental Northwest Territories, Canada. Canadian Field-Naturalist 82: 263-275. Hultén, E. 1968. Flora of Alaska and neighboring terri- tories. Stanford University Press, Stanford, California. 1008 pp. Mulligan, G. A. 1970. Cytotaxonomic studies of Draba glabella and its close allies in Canada and Alaska. Canadian Journal of Botany 48: 1431-1437. Mulligan, G. A. 1972. Cytotaxomomic studies of Draba species in Canada and Alaska: D. oligosperma and D. incerta. Canadian Journal of Botany 50: 1763-1766. Mulligan, G. A. 1975. Draba crassifolia, D. albertina, D. nemorosa, and D. stenoloba in Canada and Alaska. Canadian Journal of Botany 53: 745-751. Porsild, A. E. 1951. Botany of the southeastern Yukon adjacent to the Canol Road. National Museum of Canada Bulletin 121: 1-400. Porsild, A. E. 1975. Materials for a flora of Central Yukon Territory. National Museums of Canada, Publications in Botany, Number 4. 77 pp. Porsild, A.E. and W.J. Cody. 1968. Checklist of the vascular plants of Continental Northwest Territories, Canada. Plant Research Institute, Canada Department of Agriculture, Ottawa. 102 pp. Wein, R. W., L. R. Hettinger, A. J. Janz, and W. J. Cody. 1974. Vascular plant range extensions in the northern Yukon Territory and northwestern Mackenzie District, Canada. Canadian Field-Naturalist 88: 57-66. Welsh, S. L. and J. K. Rigby. 1971. Botanical and physio- graphic reconnaisance of Northern Yukon. Brigham Young University Science Bulletin, Biological Series, Volume XIV, Number 2. 64 pp. Received 21 September 1977 Accepted 9 March 1979 Food and Feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, Ontario HUGH R. MACCRIMMON and ROBERT W. PUGSLEY Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1] MacCrimmon, H. R. and R. W. Pugsley. 1979. Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, Ontario. Canadian Field-Naturalist 93(3): 266-271. Feeding by adult Rainbow Smelt (Osmerus mordax) is selective and varies with fish size, sex, and season. Intensity of feeding is lowered prior to spawning. Zooplankton is the major food, and we found no evidence of predation on the eggs or young of sport fishes. Fish were found in the stomachs of 11% of the females and 4.3% of the males and were mainly young-of-the-year smelt in summer and lake Emerald Shiners ( Notropis atherinoides) in winter and spring. Fish were eaten only by smelt of age groups I to V (156 to 204 mm in fork length) while crustaceans were the principal food of young-of-the-year smelt. Key Words: seasonal variation, selection, food preference, diet, predation, food organisms, Osmerus mordax, limiting factors, body size, feeding behavior. With the exception of a few lakes in the eastern part of the province (Dymond 1944), the invasion of Rainbow Smelt, Osmerus mordax, into Ontario inland waters is of recent origin. Smelt first appeared in Lake Simcoe (44°23’N, 79°18’W) in 1962, pre- sumably as a result of an unrecorded release about 1960: a substantial naturalized smelt population resulted (MacCrimmon and Skobe 1970). Because of a decline in Lake Trout (Salvelinus namaycush) and Whitefish (Coregonus clupeaformis) fisheries concurrent with the naturalization of smelt in Lake Simcoe, and because of the smelt’s unique dentition (McAllister 1963), it is a popular accusation that the smelt is a serious predator on the eggs and young of commercial and sport fishes. This assump- tion is strengthened by the fact that Lake Simcoe smelt can be readily caught by anglers using hooks baited with fresh or salted Emerald Shiners, Notropis atherinoides, especially in the winter ice fishery. Studies elsewhere, however, have not confirmed that the smelt is an important predator on fishes (Creaser 1925, 1928: Kendall 1927: Greene 1930: Schneberger 1936: Gordon 1961: Ferguson 1965: Rupp 1968: Burbidge 1969: Delisle 1969: Lackey 1969: Anderson and Smith 1971: Selgeby et al. 1978). We examined the stomach contents of smelt taken monthly by gill nets set in the lake and of mature adults collected from a river spawning area. Seasonal patterns of food and feeding of smelt in Lake Simcoe are reported, and the importance of predation on fishes assessed. Materials and Methods The area of intensive study included a 54-km2 section of Lake Simcoe north of Sibbald Park, extending west from Georgina Island to Jackson’s Point and then north to the 22-m depth contour. Monthly samples were taken from this area between August 1970 and August 1971 by means of gill nets set in the late morning for periods not exceeding | h. Specimens were chilled and examined in the labora- tory within 3 h of capture. An additional sample of 336 smelt was collected from the 1971 spawning run in the nearby Pefferlaw River and frozen for later study. The data from fish stomachs taken during the spawning period are dealt with separately from those of other months. Stomach contents were examined under a binocu- lar microscope at 16X using a technique similar to that of Thompson (1959), Ferguson (1965), and Burbidge (1969). The volume of partially digested food was estimated as a percentage of its contribution to the total stomach food volume. The amounts of the individual food items in the remaining volume were expressed as percentage contribution to the total volume of undigested recognizable food items. The frequency of occurrence of specific food items is expressed as a percentage of those stomachs contain- ing recognizable items. Stomachs containing only chyme or fluid were considered to be empty. Results Examination of the stomachs of 1416 adult smelt from Lake Simcoe between August 1970 and 1971 found that 76% contained food and, of these, recognizable items were present in 91.5% (Table 1). Feeding varied seasonally, both in kind and quantity of food eaten. All smelt stomachs collected in early June contained food, as did over 84% of those collected in July to December. Food intake was minimal in March and April before the spawning period but increased during and after spawning among mature fish captured during the Pefferlaw River spawning run. Females resumed feeding more quickly than males after spawning. Stomach fullness normally fluctuated between 47 and 69% of stomach 266 1979 MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE 267 TABLE !—Monthly stomach contents of mature Rainbow Smelt showing the number containing food, fullness, and the occurrences of recognizable food, all expressed in percentages Stomachs Stomachs Stomach containing Sample containing fullness recognizable Month size food by volume food items fe) ? 3 2 fe) 2 fe) 2 Aug. 22 89 100.0 98.1 50.9 46.7 90.9 92.1 Sept. i 18 29 100.0 96.6 51.7 55.9 50.0 Sal Oct. 37 68 100.0 100.0 SPT 50.9 89.2 77.9 Nov. ] 36 — 86.1 — 68.6 — 96.8 Dec. Mp) 78 100.0 98.7 S33 60.0 100.0 98.7 Jan. 72 60 100.0 100.0 57.9 62.6 94.4 98.3 Feb. 48 60 72.9 71.7 36.3 44.9 97.1 88.4 Mar. 54 61 16.7 21.3 55.0 46.5 100.0 100.0 Apr.* 163 62 Die) 38.7 10.3 11.3 100.0 100.0 May 28 68 64.3 82.4 58.1 68.5 100.0 100.0 June 46 58 100.0 ~ 100.0 64.0 67.6 87.0 89.7 July 29 83 86.2 86.8 65.2 48.7 92.0 87.5 Aug. 4] 83 97.6 92.8 70.3 59.0 97.5 96.1 By sex Number 581 835 38] 695 381 695 349 635 Mean 65.6 83.2 S245) 55.0 91.6 91.4 Combined (6+@) Number 1416 1076 1076 984 Mean 76.0 54.0 91.5 Mature fish on Pefferlaw River spawning grounds 336 42.0 45.9 100.0 *Excludes spawning fish. capacity, but the food volume in April was only 10%. Although stomach fullness varied slightly (1.0-16.5%) between males and females in any one monthly sampling period, the yearly average was nearly the same (Table 1). The importance of each food item varied with season and both sexes typically ate the most abundant and/or largest organisms available. In winter (January to March) copepods (Cyclops sp. and Diaptomus oregonensis) were the most important food items eaten by both sexes, followed by clado- cerans (mainly Bosmina sp.) which became prevalent in February. Amphipods (Gammarus) and fish (N. atherinoides, and young-of-the-year O. mordax) made up 28 and 40% respectively, of the food in February and March. In April, prior to spawning, copepods and tendipedids were the most frequent organisms eaten (Table 2). In smelt captured during the spawning run in the Pefferlaw River, fish eggs (O. mordax and Cato- stomus commersoni) accounted for 55.7% of the food volume in those stomachs containing food. No extraneous matter was found in the eggs. Emerald Shiners were found in 31% of these fish, and made up 30% of the total volume of food consumed in the spawning and post-spawning period. The remaining volume of recognizable food items consisted of unidentified fish larvae (2.8%), Gammarus (0.7%), and algae (0.7%). Empty stomachs were found in 58% of fish collected and many of these were enlarged and contained only clear fluid. Smelt collected in the lake in May and June had fed principally on the dipteran larvae of Chaoborus albipes, Tendipes tentans, Pentaneuris monilis, and the ephemeropteran nymph of Hexagenia sp. No bottom detritus was present in stomachs containing these benthic organisms. In July, cladocerans ( Lepto- dora kindtii), tendipedidae pupae, and copepods were the dominant food items. In September to December, fish (mostly preserved Emerald Shiners used to prebait Whitefish fishing grounds) and cladocerans (Bosmina sp. and Daphnia longispina) made up a greater proportion of the food than did copepods (Table 2). Although the seasonal variation in food items eaten was similar in both male and female smelt, slight differences in stomach volume and frequency of occurrence of these items did occur. Male smelt ate 268 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 2—Monthly stomach contents of male and female Rainbow Smelt expressed by percent volume and by frequency (in parenthesis) of occurrence Month Cladocera Copepoda Tendipedidae Aug. 53.6 4.6 32.0 (88.1) (82.2) (47.5) Sept. 49.6 18.9 13.6 (75.0) (78.6) (42.9) Oct. 49.1 5.8 6.8 (65.1) (58.1) (12.8) Nov. 76.0 6.0 0.7 (100.0) (100.0) (6.7) Dec 92.1 dS — (100.0) (100.0) — Jan. 5.4 82.9 0.3 (80.3) (92.1) (3.1) Feb. 36.0 30.7 0.3 (70.8) (79.2) (1.4) Mar. 0.5 55.0 — (4.5) (59.1) = Apr. — 69.9 14.2 — (71.1) (21.3) May _ 8.5 46.5 (25.0) (67.7) June 2.0 19.7 52.0 (12.0) (25.0) (53.3) July 46.5 9.4 IW 3a (83.7) (45.3) (43.0) Aug. 80.3 DP? 11.0 (91.2) (30.1) (31.9) Mean, sexes combined 39.5 23.9 15.8 (64.4) (64.0) (26.3) Mean, by sex 3 36.0 35.9 14.2 (66.2) (70.5) (27.0) S 41.5 17.3 16.6 (63.5) (60.5) (26.0) Food item Fish Chaoborus Hexagenia Gammarus 6.0 3.0 — — (6.9) (3.0) — — 17.1 _ 0.4 _— (21.4) — (3.6) 34.1 _ — 3.0 (37.2) = — (5.8) 11.0 — — 6.3 (16.7) — — (16.7) — — —_— 0.4 — _ _ (1.0) 7.1 0.2 — 0.9 (@A))) (2.4) _ (7.9) 11.0 5.1 _ 16.8 (12.5) (6.9) —_ (19.4) 26.1 3.2 _ 14.1 (36.4) (4.5) — (18.2) 1.7 oo _ 3.3 (1.6) _ — (4.9) 4.4 9.3 24.3 4.3 (4.4) (27.9) (30.9) (5.9) + 2.1 28.0 = aa (6.4) (38.0) — Dell DAES 6.9 = (2.3) (46.5) (8.1) Peo) 3.8 -- — (2.7) (16.8) — — 7.6 33.9) 49 2.6 (8.6) (9.6) (3.0) (3.6) 315) 2.1 48 2.8 (4.3) (8.0) (1.2) (3.4) 9.9 49 5.0 2.4 (11.0) (10.7) (5.4) (3.9) cladocerans and copepods in equal quantities (36% by volume), supplemented by tendipedidae (14.2%) and, to a lesser extent, by Hexagenia, fish, Gammarus, and Chaoborus, respectively. Females fed primarily on Cladocerans (41.5%), followed, in decreasing amounts, by copepods, tendipedidae, fish, Chao- borus, Hexagenia, and Gammarus (Table 2). Larger smelt exhibited a more varied diet than did smaller fish (Table 3). The occurrence of tendipedidae (larvae and pupae), Chaoborus, Hexagenia, and fish (Emerald Shiners and young-of-the-year Rainbow Smelt) each increased with fish length. Fish first appeared in the diet of smelt at Age Group |. Most of the identifiable fish eaten throughout the year were young-of-the-year smelt. During the autumn and winter months, preserved (salted) Emerald Shiners, which are used as bait and for prebaiting Whitefish fishing grounds, were eaten occasionally. In the spring (mostly after the smelt had spawned), live shiners were consumed in substantial quantities (30.8% by volume in the river sample). The high percentage of Emerald Shiners occurring in the October sample (34% by volume) resulted from a collection of smelt made in proximity to an area newly baited for Whitefish. The only other fish found in smelt stomachs were two larval White Suckers (C. commersoni) (15 mm in length) and a single small darter (Etheostoma sp.). Discussion As a result of the broad temperature regime favorable for smelt feeding and growth (Ferguson 1965), the smelt is able to feed throughout the year and at various water depths on a variety of food 1979 MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE 269 organisms, especially zooplankton supplemented sea- sonally by small fish and benthic invertebrates. Similar items of diet are commonly used by smelt in the Great Lakes (Kendall 1927: Schneberger 1936: Baldwin 1948: Hale 1960: Gordon 1961; Anderson and Smith 1971) and in other inland waters (Greene 1930; Reif and Tappa 1966: Rupp 1968; Burbidge 1969: Delisle 1969; Lackey 1969). The diet of landlocked Arctic Smelt (O. mordax dentex) in the Rybinsk reservoir in Russia (Inanova et al. 1970) also resembles that of freshwater Rainbow Smelt in Lake Simcoe. A comparison of the stomach contents of male and female fish indicates that although the food items are similar, the females eat more of the larger organisms (fish and dipteran larvae) and havea more varied diet. Also, recently spent males are slower to resume feeding than are females. There is no mention of these phenomena elsewhere in the published literature. The absence of debris in smelt stomachs that contain eggs in the spring months, and benthic larvae and pupae at most times, indicates that feeding is selective in Lake Simcoe. Adult smelt generally select the largest organisms available and the larger fish have a more complex diet than the smaller fish. Similar feeding behavior was observed among smelt in lakes Superior (Hale 1960), Huron (Baldwin 1948), Champlain (Green 1930), and Gull Lake, Michigan (Burbidge 1969). Lake Simcoe smelt feed extensively on eggs and fish during the spawning and post-spawning season in the rivers. Although over half the volume of food consumed at those times was comprised of smelt and sucker eggs, eggs were not found in smelt stomachs at other times. Creaser (1925) also found small quanti- ties of smelt eggs in the stomachs of smelt captured in the spawning run in Cold Creek, Michigan. Stomachs of 12% of the adult smelt taken on their spawning grounds in the Pefferlaw River contained Emerald Shiners. At other times of the year fish were found in the stomachs of 8.6% of the adult smelt [EOE SENT examined and were primarily young-of-the-year smelt in summer and fall, and salted Emerald Shiners in winter. The selection of fish at certain seasons has been reported in other papers. For example, 97.3% of the stomachs of 147 smelt taken in Crystal Lake, Michigan in September contained Emerald Shiners (Creaser 1925). Larvae of the Cisco (Coregonus artedii) were found in 17% of the stomachs of smelt taken in May from Black Bay of Lake Superior (Selgeby et al. 1978). In Lake Huron, Emerald Shiners were found in 8% of smelt stomachs but only in July and August, and young-of-the-year smelt were found OTN OR OD os in 23% of smelt stomachs but only in September ay yall ae el” sale (Gordon 1961). In New York State, the food of the S22 SSSSefa8 large race of Lake Champlain smelt contained a lower Gammarus 10.0 4 6.7 Fish 4.] 10.3 14.3 26.7 Hexagenia | I 13 50.0 Food item 18.4 19.0 26.7 25.0 Copepoda Tendipedidae Chaoborus 100.0 - 90.0 _- 89.7 65.6 Wee 54.0 26.4 45.2 28.6 33.3 40.0 25.0 50.0 spawning season Cladocera 100.0 100.0 97.4 92.6 93.1 83.3 80.0 75.0 Stomachs containing recognizable food items 15 4 320 Total remains Fish empty 3 5 3 2 3 16 of fish 24 9 79 No. m) (m interval TABLE 3—Percent annual frequency of occurrence of major food items eaten by adult Rainbow Smelt (130 to 209 mm fork length) excluding fish taken during Length SS SS ei ia 270 percentage of fish during June and July, while, in the Saranac Lakes, fewer fish were eaten in summer (Greene 1930). Similarly, in Lake Erie, fish occurred in only 2.3% of smelt stomachs in spring and summer, but increased to 51.2% during the autumn (Price 1963). There is no evidence from any study. including this one. that any segment of the smelt population becomes totally piscivorous. either on a seasonal or permanent basis: or that smelt predation is an obvious factor in the suppression of any sympatric population. The feeding selectivity of the Rainbow Smelt. how- ever. could exert competitive pressure on specific food items required at some critical stage in the develop- ment of other fish species. For example. during the summer Lake Simcoe smelt eat more Leptodora kindtii even though smaller cladocerans are more abundant. Also Reif and Tappa (1966) suggest that the differential predation of stocked smelt on zoo- plankton in Harvey’s Lake, Pennsylvania, has result- ed in the disappearance of L. kindtii and the replacement of the larger Daphnia pulex by the smaller D. dubia. It is possible that competition for copepods between adult smelt and larval lake Cisco has led, in part. to the decline of the Cisco in the waters of western Lake Superior (Anderson and Smith 1971). Thus. in considering possible impacts of an invad- ing fish species on the welfare of indigenous species. such as the relationship between the naturalized smelt and native Lake Trout (Sa/velinus namaycush) and Whitefish in Lake Simcoe, a knowledge of relative diet and food preferences of the species at all stages of life. as well as ecosystem responses. is essential. The importance of the observed cannibalism on eggs and young-of-the-year smelt as a self-regulatory popula- tion control is not known but in view of the increasing abundance of the species is obviously not a serious constraint at the ambient stage of naturalization. There are insufficient data on the food of other fish species in Lake Simcoe to permit consideration of the likelihood of important interspecific competition for food. nor is there evidence of predation by Rainbow Smelt on either the Lake Trout or Whitefish popula- tion. The significance on the Lake Simcoe biosystem of observed variations in the extent and composition of the diet of the Rainbow Smelt relative to season. sex. and size of fish is worthy of more profound investigation. Acknowledgments We are indebted to the staff of the Lake Simcoe Fisheries Assessment Unit of the Ontario Ministry of Natural Resources for their aid and the use of their facilities. Financial assistance for the study was provided by the Canadian National Sportsmen’s Show and the National Research Council of Canada. THE CANADIAN FIELD-NATURALIST Vol. 93 Literature Cited Anderson, E. D. and L. L. Smith, Jr. 1971. A synoptic study of food habits of 30 fish species from western Lake Superior. University of Minnesota Agricultural Experi- mental Station, Technical Bulletin 279. 199 pp. Baldwin, N.S. 1948. The American Smelt, Osmerus mor- dax (Mitchill) of South Bay. Manitoulin Island. Lake Huron. Transactions of the American Fisheries Society 78: 176-180. Burbidge, R. G. 1969. Age. growth, length-weight relation- ship. sex ratio. and food habits of American Smelt. Osmerus mordax (Mitchill), from Gull Lake. Michigan. Transactions of the American Fisheries Society 98: 631— 640. Creaser, C. W. 1925. The establishment of the Atlantic smelt in upper waters of the Great Lakes. Papers of the Michigan Academy of Science. Arts. and Letters 5: 405-424. Creaser, C. W. 1928. The food of yearling smelt from Michigan. Papers of the Michigan Academy of Science. Arts, and Letters. 10: 427-431. Delisle, C. 1969. Ecologie. croissance et comportement de léperlan du Lac Heney. comté de Gatineau ainsi que la répartition en eau douce au Québec. Ph.D. thesis. Department of Biology, University of Ottawa. 189 pp. Dymond, J. R. 1944. Spread of the smelt (Osmerus mor- dax) in the Canadian waters of the Great Lakes. Can- adian Field-Naturalist 58: 12-14. Ferguson, R. G. 1965. Bathymetric distribution of Ameri- can Smelt. Osmerus mordax. in Lake Erie. University of Michigan Great Lakes Research Division Publication. 13: 47-60. Gordon, W.G. 1961. Food of the American Smelt in Saginaw Bay. Lake Huron. Transactions of the American Fisheries Society 90: 439-443. Greene, C. W. 1930. The smelts of Lake Champlain. Jn A biological survey of the Champlain watershed. Supple- ment to the 19th Annual Report. 1929. of the New York Conservation Department. pp. 105-129. Hale, J. G. 1960. Some aspects of the life history of the smelt (Osmerus mordax) in western Lake Superior. Minnesota Fish and Game Investigations. Fish Series 2: 25-41. Inanova, M.N., I. Ye. Permitin, and S.N. Polovkova. 1970. Seasonal distribution features of adult landlocked smelt (snetok). Osmerus eperlanus eperlanus morpha spirinchus Pallas. in the Volga Reach of Rybinsk reservoir. Journal of Ichthyology 10: 678-685. Kendall, W. C. 1927. The smelts. U.S. Bureau of Fisheries. Bulletin 11: 217-375. Lackey, R. T. !969. Seasonal depth distribution of land- locked Atlantic Salmon. Brook Trout. landlocked Ale- wives. and American Smelt in a small lake. Journal of the Fisheries Research Board of Canada 27: 1656-1661. MacCrimmon, H.R. and E. Skobe. 1970. The fisheries of Lake Simcoe. Fish and Wildlife Branch. Department of Lands and Forests (Ontario Ministry of Natural Re- sources). 140 pp. McAllister, D. E. 1963. A revision of the smelt family. Osmeridae. Bulletin of the National Museum of Canada 191, Biological Series 71. 53 pp. 1979 Price, J. W. 1963. A study of the food habits of some Lake Erie fish. Bulletin of the Ohio Biological Survey (New Series). Volume II (No. 1). 89 pp. Reif, C. B. and D. W. Tappa. 1966. Selective predation: smelt and cladocerans in Harvey Lake. Limnology and Oceanography I 1: 437-438. Rupp, R.S. 1968. Life history and ecology of the smelt. Osmerus mordax, in inland waters of Maine. Maine Department of Inland Fisheries and Game. Fisheries Research and Management Division. Final Report. 36 pp. Schneberger, E. 1936. The biology and economic impor- tance of the smelt in Green Bay. Transactions of the American Fisheries Society 66: 139-142. MACCRIMMON AND PUGSLEY: RAINBOW SMELT FOOD, LAKE SIMCOE Ti Selgeby, J. H.. W.R. MacCallum, and D. V. Swedberg. 1978. Predation by Rainbow Smelt (Osmerus mordax) on Lake Herring (Coregonus artedii) in western Lake Superior. Journal of the Fisheries Research Board of Canada 35: 1457-1463. Thompson, R.B. 1959. Food of the squawfish. Prycho- cheilus oregonensis (Richardson). of the lower Columbia River. U.S. Fish and Wildlife Service. Fisheries Bulletin 60 (158): 43-58. Received 15 January 1979 Accepted 16 March 1979 Song Pattern of the Cypress Hills Population of White-crowned Sparrows M. ROSS LEIN Department of Biology, University of Calgary, Calgary, Alberta T2N IN4 Lein, M. R. 1979. Song pattern of the Cypress Hills population of White-crowned Sparrows. Canadian Field-Naturalist 93(3): 272-275. A disjunct population of White-crowned Sparrows (Zonotrichia leucophrys oriantha) in the Cypress Hills of Alberta possesses a distinctive song pattern which differs markedly from that of birds of the same geographic race in the Rocky Mountains of western Alberta. This suggests that this population is truly isolated and does not exchange individuals with populations in the main range of the subspecies. Key Words: White-crowned Sparrow, Zonotrichia leucophrys oriantha, Cypress Hills, Alberta, song dialects. The Cypress Hills of southeastern Alberta and southwestern Saskatchewan support an island of mesic forest habitat surrounded by drier shortgrass prairie. The plateau averages about 450 m higher than the surrounding plains, and this higher elevation results in lower temperatures and higher precipita- tion. Consequently, the region shelters disjunct populations of many plants and animals characteristic of the coniferous forests of the montane or boreal regions of Canada (Halladay 1965). Although it seems probable that many of these disjunct populations, including conifers (Thompson and Kuit 1976) and land snails (Russell 1951), are completely isolated from other conspecific popula- tions because of the problems of dispersal across long distances of inhospitable terrain, the situation is less clear for populations of mobile organisms such as birds. The Cypress Hills Plateau possesses disjunct populations of at least 12 passerine species, separated from other populations in the Rocky Mountains or boreal forest by gaps of 250-300 km. It is likely that conifer forests became established on the Cypress Hills via colonization from migrating belts of con- iferous forest following the receding Laurentide ice sheet at the end of the Wisconsin glaciation, and were isolated by the expansion of the grassland steppe vegetation, beginning about 10 000 yr ago (Thomp- son and Kuijt 1976). The populations of montane or boreal birds may have been isolated in the Cypress Hills at this time, or they may have colonized the area somewhat later. A third possibility is that these populations are not truly isolated, but continuously exchange individuals with conspecific populations in other areas. The disjunct population of White-crowned Spar- rows (Zonotrichia leucophrys) breeding in the Cy- press Hills is assigned to Z. /. oriantha, a black-lored race that also inhabits the Rocky Mountains from southern Alberta and British Columbia south to northern New Mexico, and west to south central Oregon and central eastern California (American Ornithologists’ Union 1957). Although the Cypress Hills population differs in mensural and plumage characters from other populations of oriantha, this variation is of a clinal nature and does not warrant taxonomic separation (Banks 1964). Local popula- tions of White-crowned Sparrows, particularly the Pacific coast races Z. /. nuttalli and Z. |. pugetensis (Marler and Tamura 1962: Baptista 1975, 1977), may be characterized by distinctive song “dialects.” Whether these dialects are an adaptation to reduce gene flow between populations, as suggested by Nottebohm (1969, 1970), is unclear, but they certainly may act as markers for groups of birds with little exchange of individuals (Baptista 1975). During the summer of 1978 I had the opportunity to record songs of White-crowned Sparrows in the Cypress Hills. This paper describes the song pattern of this population and uses this evidence to suggest that this population is indeed isolated. Methods Recordings were made in Cypress Hills Provincial Park, Alberta, 5-8 June 1978. A minimum of 15 individuals were taped along the eastern and northern sides of Reesor Lake and in the valley of Battle Creek. Other birds were heard but not recorded. The sparrows were singing strongly in clumps of bushes, mostly hawthorn (Crataegus spp.), roses (Rosa spp.), and Buckbrush (Symphoricarpos occidentalis), ad- joining patches of open grassland. Songs were recorded with a Nagra 4.2 tape recorder and a Gibson E. P. M. parabolic microphone. Sona- grams were prepared with a Kay Elemetrics 6061B Sona-Graph, using the wide-band filter and high- shape setting. DTZ 1979 Results and Discussion Of the 15 males whose songs were recorded, 14 possessed the black lores characteristic of oriantha. One male at Reesor Lake, however, had white or pale gray lores, a character found in the northern race of the species, Z. /. gambelii. Representative songs of each of the 15 males are shown in Figure |. With one exception, all are similar. They consist of a more or less segmented introductory whistle, a pair of warbled syllables, two buzzes, and a terminal slow trill. Although there is some variation in the form of the warbled syllables among birds, this is minor and all are clearly variants of the same pattern. Approximately 6-10 other males that were heard but not recorded possessed songs of this common pattern. The one exception (Figure |O) had a song in which the terminal trill was absent and the pair of warbled syllables was replaced by a four-note segment with a very different sound quality. The absence of the trill may not be significant because birds frequently sing incomplete songs; however, none of the songs heard from this male over a period of about 20 min had the trill, and all had the four-note segment. The male with the unusual song was not the individual with the gambelii phenotype. The latter bird’s song (Figure I C) was comparable to those of other males of this population. The song pattern of White-crowned Sparrows in the Cypress Hills is markedly different from the “typical” song of oriantha in the Rocky Mountains of southern Alberta, the nearest conspecific population. Figure 2 shows songs of birds from three localities in the Rockies, spanning a distance of approximately 200 km from Waterton Lakes National Park in the south to the Highwood Pass in the north. All have similar songs, with three whistled notes in place of the warbled syllables of the Cypress Hills birds, and a terminal trill consisting of simpler notes than those of Cypress Hills songs. No-.songs resembling those of Cypress Hills birds have been recorded in the Rocky Mountains of Alberta. The sharp differences between the songs of Cypress Hills birds and those of birds from the Rocky Mountains of Alberta suggest that there is little, if any, exchange of birds between these two oriantha populations, and that the Cypress Hills White- crowned Sparrows form a discrete population. Be- cause juvenile birds probably learn their song prior to dispersing from their natal area (Marler 1970), any bird immigrating to the Cypress Hills from another population would be expected to sing a different song pattern. The apparent absence of songs resembling those of montane oriantha suggests that there is little influx of birds from this area. Other disjunct populations of White-crowned Sparrows are found on other montane “islands” in LEIN: CYPRESS HILLS WHITE-CROWNED SPARROWS 23 Montana (Thompson 1978). The song patterns of these populations are unknown. But the fact that White-crowned Sparrows are absent from the Sweet- grass Hills (100 km WSW of the Cypress Hills and 140 km E of the Rocky Mountains) although suit- able habitat is available (Thompson 1978), also suggests a low rate of immigration of birds from the main range into outlying populations. The general similarity of songs of birds over a wide area in the Rocky Mountains (Figure 2) argues that there is considerable exchange of birds between populations in this region. Studies of oriantha populations in California and Colorado (Baker 1975; Orejuela and Morton 1975), however, have demon- strated the existence of dialects in the songs of populations separated by only 16 km. This suggests that population structure, dispersal, and the develop- ment of song dialects may show a considerable amount of variation within this single subspecies. There are several possible explanations for the presence of a white-lored bird in the Cypress Hills. Migrant gambelii have been recorded there (Taverner 1927; Rand 1948), and this individual may represent the result of past introgression. But birds with white lores have been collected on several occasions far to the east of the range of gambelii, or any of the other white-lored races (Rand 1948), and this may be a character that is exhibited occasionally in generally black-lored populations. The one individual with an aberrant song (Figure 10) may have copied the song of another population (Baptista 1974) or it may reflect an error in song- learning (Baptista 1975). Acknowledgments I gratefully acknowledge the aid of Valerie A. Haines in recording songs in the Cypress Hills, and the valuable comments on the manuscript made by Luis F. Baptista. Literature Cited American Ornithologists’ Union. 1957. Check-list of North American birds. Fifth edition. Port City Press, Inc., Baltimore, Maryland. Baker, M.C. 1975. Song dialects and genetic differences in White-crowned Sparrows (Zonotrichia leucophrys). Evo- lution 29: 226-241. Banks, R.C. 1964. Geographic variation in the White- crowned Sparrow Zonotrichia leucophrys. University of California Publications in Zoology 70: 1-123. Baptista, L. F. 1974. The effects of songs of wintering White-crowned Sparrows on song development in seden- tary populations of the species. Zeitschrift fur Tier- psychologie 34: 147-171. Baptista, L. F. 1975. Song dialects and demes in sedentary populations of the White-crowned Sparrow (Zonotrichia leucophrys nuttalli). University of California Publications in Zoology 105: 1-52. 274 THE CANADIAN FIELD-NATURALIST Vol. 93 A aaa MAAK J | \\ \\ ee pe i FIGURE 1. Songs of male White-crowned Sparrows from Cypress Hills Provincial Park, Alberta. 1C is the song of a white- lored bird; all others are songs of black-lored birds. Time scale is in seconds. Each division of ordinate scale is 1 kHz. 1979 HAaALe. Q ama HAAAAAA AS 1.0 2.0 Ss LEIN: CYPRESS HILLS WHITE-CROWNED SPARROWS 2S Seg — F & cae HAAKKaS cast 44444444 FIGURE 2. Representative songs of oriantha White-crowned Sparrows from the Rocky Mountains of Alberta. 2A and 2B are from Waterton Lakes National Park; 2C and 2D are from the Sheep River Valley; and 2E and 2F are from Highwood Pass, Kananaskis Valley. Time scale is in seconds. Baptista, L. F. 1977. Geographic variation in song and dialects of the Puget Sound White-crowned Sparrow. Condor 79: 356-370. Halladay, I. R. 1965. Recent biota of the Cypress Hills Plateau: a general survey of the natural history. Jn Cypress Hills Plateau guidebook, Part 1. Edited by R. C. Zell, 15th Annual Field Conference, Alberta Society of Petroleum Geologists. Pp. 37-54. Marler, P. 1970. A comparative approach to vocal learn- ing: song development in White-crowned Sparrows. Journal of Comparative and Physiological Psychology, Monograph 71: 1-25. : Marler, P. and M. Tamura. 1962. Song “dialects” in three populations of White-crowned Sparrows. Condor 64: 368-377. Nottebohm, F. 1969. The song of the Chingolo, Zono- trichia capensis, in Argentina: description and evaluation of a system of dialects. Condor 71: 399-315. Nottebohm, F. 1970. Ontogeny of bird song. Science 167: 950-956. Orejuela, J. E. and M.L. Morton. 1975. Song dialects in several populations of Mountain White-crowned Spar- rows (Zonotrichia leucophrys oriantha) in the Sierra Nevada. Condor 77: 145-153. Rands, A. L. 1948. Birds of southern Alberta. National Museum of Canada, Bulletin Number 111. Russell, L.S. 1951. Land snails of the Cypress Hills and their significance. Canadian Field-Naturalist 65: 174-175. Taverner, P. A. 1927. Some recent Canadian records. Auk 44: 217-228. Thompson, L.S. 1978. Species abundance and habitat relations of an insular montane avifauna. Condor 80:: 1-14. Thompson, L.S. and J. Kuijt. 1976. Montane and sub- alpine plants of the Sweetgrass Hills, Montana, and their relation to early postglacial environments of the northern Great Plains. Canadian Field-Naturalist 90: 432-448. Received 10 January 1979 Accepted 26 February 1979 Nesting Biology and Development of Young in Ontario Black Terns ERICA H. DUNN Department of Biology, Trent University, Peterborough, Ontario Present address: Long Point Bird Observatory, P.O. Box 160, Port Rowan, Ontario NOE 1MO E. H. Dunn. 1979. Nesting biology and development of young in Ontario Black Terns. Canadian Field-Naturalist 93(3): 276-281. Habitat, nesting substrate, egg characteristics, and growth are described for Black Terns ( Chlidonias niger) nesting on Lake Erie. Black Terns prefer moderate density of emergent vegetation (mainly Typha) in about | m of water, adjacent to open water, and return to the same nest site or area as long as it is suitable. Egg and growth characteristics are similar to those for other temperate-nesting tern species. Minnows form a good portion of the biomass consumed by chicks in this area. Key Words: Black Terns, Chlidonias niger, nesting, growth, egg characteristics. Black Terns (Chlidonias niger) nest widely through- out southern Canada, but their breeding biology is not well known. As the preferred habitat is cattail (Typha spp.) marsh with a metre or more of water, and the young after the first 2 or 3 d leave the nest when disturbed, accessibility for observation is restricted. Only one intensive study of breeding biology has been done (Cuthbert 1954), and this included few data on development of young. This paper fills in some gaps in knowledge of nesting biology and gives the first data on growth of chicks. Methods This study area was an approximately 15-ha marsh on the north side of Long Point, near Port Rowan, Ontario (42°35’N, 80°24’W). A description of the vegetation and bird life is given in Dunn and Nol (1976), and mammals seen in the area were Muskrats (Ondatra zibethicus) and Mink (Mustela vison). A canoe was used to traverse the study area. There were approximately 30-40 Black Tern nests in the area, in small groups. Twenty-three nests in four groups were studied in 1975 and 32 in five groups in 1976. Nests were identified with tagged 2-m stakes pushed into the mud. Eggs were marked with indelible felt-tip marker, and measured with dial calipers to the nearest 0.01 mm (length and width at broadest point), and were weighed several times during incubation to the nearest 0.1 g with a Pesola spring balance. Occasion- ally eggs were floated ina beaker of fresh lake water to determine degree of development (Hays and LeCroy 1971). Fences of hardware cloth were placed around the nests during incubation to prevent the young from leaving the nest site during development. The fences were 0.4—0.5 m in diameter and 0.3 m high. The adults readily incubated and cared for young within the fences, and early placement made it easier to catch adults later in roofed traps of similar size, with entrance holes in the top. Adults and young were banded with U.S. Fishand Wildlife Service aluminum bands. Color bands were used on adults until it became clear that they were almost never visible. Chicks were weighed every | or 2 d to the nearest 0.1 g, and tarsus and ulna were measured with the adjacent bones held at right angles. Measurements of skull were from bill tip to back of the skull, with the ruler held parallel to the bill; and of culmen from bill tip to feather line on the upper mandible. Sheath and feather length were measured for the 9th primary, outer rectrix, and a few body feathers on the anterior parts of dorsal and ventral tracts. On four occasions, observations of nests were made from a blind mounted permanently on an anchored rowboat. Results and Discussion Black Tern nests were formed of dead cattail, most commonly on a pre-existing mat of floating dead TABLE |—Percentage of Black Tern nests built on various substrates: A, Ontario (N = 24)—this study; B, Michi- gan (N = 23)—Cuthbert (1954); C, Iowa (N = 197)—Berg- man et al.; D, California (N = 40)—Gould (1974) A B (Cc D Floating dead vegetation 75 48 11 15 Floating boards or logs 17 22 33 Cattail rootstock 8 53 Muskrat-built structures 22 36 33 Abandoned birds’ nests” 20 Broken-down bulrushes 9 “Muskrat lodges or feeding platforms of freshly cut vegeta- tion. *Platforms of old grebe ( Podiceps) and Forster’s Tern nests. Black Terns have also been found nesting on abandoned American Coot (Fulica americanus) nests (Cuthbert 1954). 276 1979 cattail lodged in emergent vegetation (Table 1). No nests were found on Muskrat houses or feeding platforms, although these were abundant. The majority of substrates were floating, in water 1-1.2 m deep. A few nests in a nearby marsh were found on isolated small patches of soft mud, but these nest sites were otherwise normal (see below). Black Tern nests on dry land are rare (Cuthbert 1954; Richardson 1967). These terns are opportunistic in the choice of supporting structure for their nests, althougn in my study floating vegetation and boards seemed to be preferred over Muskrat houses (Table 1). Previous papers have not commented on the relative avail- ability of other substrates. Nest characteristics were described for 25 nests. Typically, they were about 2-5 cm high and 25 cm across. The underlying mats of dead cattail were lodged in living cattail, usually with heavy growth on at least one side, and averaged 8 m? (ranging from 0 for a nest on a board to 417 m? in area). Open water was, on average, 4 m away (range: 0.5-12 m). Some nests were in large pools, and these were generally on boards or rootstock, depending less on emergent vegetation to support the nest substrate. Moderate cattail growth seemed preferred (standing at least | m above the water but dispersed enough so that a canoe could be forced through). A few nests were in thin new growth, but were rare in dense old stands. Other studies show that the species composition of aquatic vegetation at Black Tern breeding sites varies markedly, but that its density 1s usually moderate. Water is usually | m deep and nests normally are adjacent to open water (Cuthbert 1954; Richardson 1967; Bergman et al. 1970: Gould 1974). Although nests were usually well spaced (only 2 or 3 within a 25-m2 area), a few were within 3 m, with a clear view between them. Groups had up to 10 nests, and probably resulted from chance distribution of good sites. About one quarter of all nests were not near other tern nests. This dispersion of nests is typical (Cuthbert 1954: Gould 1974), although nests may occasionally be much closer together (Hoffman 1926). Because floating vegetation became waterlogged by late July and winter storms altered nest-site charac- TABLE 2—Success of Black Terns nests DUNN: BLACK TERN NESTINGS 27), teristics, and because water levels and cattail growth alter over a period of years, terns must seek new nest sites each year. McNicholl (1975) predicted that under these circumstances, the birds would continue to usea nesting area as long as the habitat is suitable, but when they must move, they would go as a group to a new site. This is in contrast to the site tenacity demon- strated by several gull and tern species with permanent nesting areas. A few re-trap data suggest that Black Terns do return to the same general location of a previous breeding site and will nest there again if possible. Of five birds trapped in 1976, which had been banded previously, one was at nearly the same nest site as in 1975 and another was within the same general locality, probably moving less than 50 m. (Changes in habitat between years made exact mapping difficult.) Two birds banded in 1972 were also found in 1976 within a short distance of the banding site. The fifth bird moved farther, about 75-100 m, but was still within the same patch of marsh vegetation as in 1975. The Black Terns in my study moved in 1977, after winter storms had so altered the habitat that no suitable floating platforms were available for nest substrates. So many Black Terns nest in the Long Point marshes, however, that it would have been impractical to trap other areas to determine whether McNicholl’s (1975) prediction on group adherence held true. The outcome of the 55 nests located during the 2 yr is given in Table 2. The majority of known losses occurred in the egg stage. In most cases, eggs disappeared or did not hatch, and a few nests were destroyed by storms. Undoubtedly, some losses resulted from my activities (eggs cracked during trapping of adults, young in retaining fences eaten by Mink). Nevertheless, Bergman et al. (1970) showed a very similar success rate (29% of 192 nests) in a much larger study with no handling of eggs or birds. There was no correlation between success and nest-site characteristics either in my study or in that of Bergman et al. (1970). There were no significant differences in measure- ments of eggs according to sequence in the clutch 1975 No. % Successful’ 9 39 Unsuccessful 9 39 Undetermined 5) DD, Total 23 anaes otal sali] Opa amen Total No. YG No. % 6 19 15 27 23 72 32 58 3 9 8 15 32 55 “Young hatched, but later fate may have been unknown. Nests in which all chicks were known to have died were classified as unsuccessful. 278 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 3—Mean +SD (N) weights and measurements of Black Tern eggs Egg 1 Egg 2 Egg 3 All eggs Weight (g) 10.63 + 0.96 (8) 11.42 + 0.51 (6) 10.92 + 0.45 (6) 10.95 + 0.76 (20) Length (mm) 33.96 + 0.98 (19) 34.32 + 1.45 (13) 34.51+1.13 (20) 34.344 1.32 (91) Width (mm) 24.88 + 0.63 (19) 25.32 + 0.57 (13) 24.73 £0.45 (20) 24.92 + 0.62 (91) Shape index* 74.04 + 2.67 (15) 74.22 + 2.84 (12) 72.00 + 2.69 (18) 72.69 + 3.72 (74) “100 X width/length (Collins and LeCroy 1972). (Table 3), although third eggs were longer and narrower and had a lower shape index (Collins and LeCroy 1972). There was a tendency for the first egg to be smallest and the second to be largest (Table 4). In contrast, the smallest egg for Common Terns (Sterna hirundo) is usually the third one. As in Black Terns, the third egg is usually the most narrow (Gemperle and Preston 1955; Gochfeld 1977). Fresh egg weight of Black Terns was 17.3% of adult body weight, a slightly lower percentage than shown by the larger Common and Roseate Terns (S. dougallii) (Collins and LeCroy 1972). TABLE 4— Percentage of Black Tern nests in which given eggs in the laying sequence were largest, of middle size, or smallest in the clutch. N= 10 clutches. Volume calculated as per Shott and Preston (1975) Egg | Egg 2 Egg 3 Largest in clutch 10 60 30 Medium sized 30 30 40 Smallest in clutch 60 10 30 Development of embryos could be followed through flotation of eggs. The pattern for Black Terns is very similar to that in Common Terns (Hays and LeCroy 1971), although Black Tern eggs sometimes float a day earlier (Table 5). Not enough eggs were floated to give further details, but within a clutch, the degree of flotation normally indicated the true laying sequence. Incubation begins after laying of the first egg and eggs are usually laid a day apart (Haver- schmidt 1945), so that the eggs are graded in amount of incubation. TABLE 5—Flotation and hatching schedule of Black Tern eggs Days after laying of egg Development 6-7 Vertical, sits on pointed end 8 Some eggs float 10-11 Last eggs float 19 - 20 Eggs pip 20 — 22 Eggs hatch Weights of three freshly hatched chicks averaged 7.25 g, and subsequent growth is detailed in Figures 1-3. The average weight for day 0 (day of hatch) chicks in Figure | includes birds which may have been hatched as much as 10-20 h before being weighed. Weight gain in the chicks shows a pattern very similar to that of other temperate-nesting species. The growth curves of terns can best be fitted to logistic equations, and growth constants calculated for various species are compared in Table 6. Although Black Terns appear to grow more rapidly than the other terns, they 70 4.37634 34 te 60 50 Dh fo} MEAN ADULT WEIGHT = 63.4g (N=27) WEIGHT (GRAMS) ol {o) 20 20 AGE (DAYS) FIGURE |. Weights of Black Tern chicks. Horizontal bar shows mean, heavy vertical gives SD, and thin vertical bar indicates range. The numbers indicate sample size. 1979 DUNN: BLACK TERN NESTINGS 279 TABLE 6—Comparative growth in northern-nesting tern species Species K* ‘10 — 90° Adult wt., g Source Black Tern 0.365 12.1 63 This study Arctic Tern 0.288 isi 104 Ricklefs 1973 Common Tern 0.300 14.6 125 Ricklefs 1973 0.238 18.5 116 LeCroy and Collins 1972 Roseate Tern 0.237 18.5 108 LeCroy and Collins 1972 Sandwich Tern 0.258 237 Ricklefs 1973 17.0 (Sterna sandvicensis) F “A constant relating to the growth curve, representing rate at which asymptotic weight is attained. Calculated according to Ricklefs (1967). *Time in days between achieving 10% and 90% of asymptotic weight (Ricklefs 1967). are also much smaller as adults. Comparisons of growth in relation to asymptotic weight between Black Terns and other semi-precocial gulls, terns, and skuas (Table 6; Ricklefs 1973, p. 188) suggest that the Black Tern has a typical growth rate for its size. Although few chicks were followed through the entire growth period, some clearly lagged behind in weight gain, as is also seen in other tern species (e.g., Langham 1972). In contrast to what LeCroy and Collins (1972) found for Common and Roseate Terns, however, feather growth in laggard Black Tern chicks was also retarded. The weight loss which occurred after attainment of asymptotic weight (close to adult weight) is also common in other terns (LeCroy and Collins 1972; LeCroy and LeCroy 1974). Of the weight and linear measures shown in Figures 1 and 2, no single one proved suitable for aging chicks accurately in the field, although skull length was helpful for younger chicks and culmen length for older ones. Considered all together, these average measure- ments can be used to estimate the age of an unknown chick within a 3- to 4-d range. The tarsus reaches full adult length in the chicks well before fledging, but skull and culmen must both continue to grow after first flight (Figure 2). Growth of the body feathers levelled off by the end of the weight increase period, but both tail and wing feathers were still growing rapidly. The latter in particular can be helpful in aging nestlings (Figure 3), as was also found for Common and Roseate Tern chicks (LeCroy and Collins 1972). The egg tooth in 10 chicks dropped off before day 5, and in one case, on day | or 2. Typically it seems to disappear on day 3 or 4, making this species similar to Common, Forster’s, and Arctic Terns (Sterna para- disaea), but faster in losing the egg tooth than Roseate or Least Terns (S. albifrons) (McNicholl 1971: LeCroy and Collins 1972). On four occasions, observations of nests with single chicks were made froma blind mounted ona rowboat. The watches totalled 7 h for achick 5 d old, 2.25 hfora 15-d-old, and 5.25 h for two 19-d-olds in separate nests. These watches were too few to support general conclusions on behavior, but certain observations are worth noting. The single S-d-old chick was fed more often than the older ones, and was the only one brooded. It was fed 5.4 times per hour, as opposed to 3.6 per hour for the 15-d chick and 1.7 times per hour for each of the two 19-d chicks. Brooding took place for 14% of the observation period. These figures correspond well with those of Cuthbert (1954), who made much lengthier observations of chicks less than 9 d old. There is apparently an increase in feeding rate over the first few days, then a decline after about 10d, when most weight gain is complete. Of 602 feedings observed by Cuthbert (1954), 5% of the items were minnows, 3% dragonflies and, overall, 93.6% were insects. Of the 56 food items seen in this study, 13% were minnows, and 6% were dragonflies. The remainder were usually unidentifiable small objects (therefore probably not minnows) and seemed to include many grubs and larvae. Although often described as primarily insectivorous, Black Terns clearly use fish as a substantial portion of the food fed to chicks. Given the large size and digestibility of the: minnows as compared to insects (Dunn 1973), fish probably provide at least one-third of the protein required by the growing chicks. Begging, feeding, and brooding behaviors corres- ponded closely to those described in detail by Cuthbert (1954) and Baggerman et al. (1956). One parent seemed to bring mainly fish to the nest, while the other brought small items, often in series of closely spaced feedings. Most of the chicks’ non-feeding activity consisted of preening, walking, or resting, and occasionally pick- ing up and immediately dropping bits of vegetation. There was a notable change in behavior by 19 d of age, when the chicks spent a good deal of time flapping their wings and hopping into the air. Five chicks disappeared from the retaining fences on day 19, only one stayed longer; earlier departures were 280 THE CANADIAN FIELD-NATURALIST Vol. 93 TARSUS CULMEN Ae | Sl 8G he | 4 al eos \ ee MEAN ADULT MEAN ADULT TARSUS = 17.3mm CULMEN = 26.5 mm (N= 27) (N= 29) LENGTH (mm) O) oO 0) Je tt als 3 50 + an sales 40 cae 30 MEAN ADULT SKULL = 59.1 mm 20 (N= 29) O 3 lO I5 0) © 5 lO I5 20 AGE (DAYS) FIGURE 2. Measurements of Black Tern chicks. Legend as in Figure 1. 1979 100 80 60 LENGTH (mm) 20 AGE (DAYS) FIGURE 3. Development of feathers in Black Tern chicks (total length, including sheath). Legend as in Figure 1. Upper bars indicate length of 9th primary, lower bars show outer rectrix growth. believed to be caused by predation or faulty fencing. The chick that was underweight throughout the growth period stayed inside the fence the longest. Delayed fledging of underweight Common Terns was noted by LeCroy and LeCroy (1974). I originally thought disappearances of young represented first flights, but the last occurrence took place while the nest was under observation. The chick flapped its wings and barely hopped the fence, then swam away. Exact age of fledging is undetermined, but must be close to 19 d. Baggerman et al. (1956) said flight first occurred 3 wk after hatching, but Cuthbert (1954) felt it was closer to 25 d after hatching. Black Terns are very similar to other temperate- nesting species in both egg and growth characteristics, even though other species studied to date are primarily piscivorous and nest along marine shores rather than in marshes. Acknowledgments This study was supported by the Frank M. Chapman Memorial Fund in 1975, and in 1976 by the National Research Council of Canada. The Long Point Bird Observatory provided facilities, a canoe, and numerous assistants. I particularly thank Erica Nol for extensive help in field work, and Marshall DUNN: BLACK TERN NESTINGS 281 Field for aid in designing traps and giving access to unpublished banding records. Literature Cited Baggerman, B., G. P. Baerends, H. S. Heikens, and J. H. Mook. 1956. Observations on the behaviour of the Black - Tern (Chlidonias n. niger(L.)), in the breeding area. Ardea 44: 1-71. Bergman, R.D., P. Swain, and M. W. Weller. 1970. A comparative study of nesting Forster’s and Black Terns. Wilson Bulletin 82: 435-444. Collins, C. T. and M. LeCroy. 1972. Analysis of measure- ments, weights, and composition of Common and Roseate Tern eggs. Wilson Bulletin 84: 187-192. Cuthbert, N. L. 1954. A nesting study of the Black Tern in Michigan. Auk 71: 36-63. Dunn, E. H. 1973. Energy allocation of nestling Double- crested Cormorants. Ph.D thesis, University of Michigan, Ann Arbor. Dunn, E. H. and E. Nol. 1976. /m Cattail marsh. Fortieth breeding bird census. American Birds 31: 83. Gochfeld, M. 1977. Intraclutch variation: the uniqueness of the Common Tern’s third egg. Bird-Banding 48: 325-332. Gemperle, M.E. and F. W. Preston. 1955. Variation of shape in the eggs of the Common Tern in the clutch- sequence. Auk 72: 184-198. Gould, G.I., Jr. 1974. Breeding success of piscivorous birds at Eagle Lake, California. M.S. thesis, Humboldt State University, California. Haverschmidt, F. 1945. De broedduur van de Zwarte Stern Chlidonias niger (L.). Ardea 33: 237-240. (English summary.) Hays, H. and M. LeCroy. 1971. Field criteria for deter- mining incubation stage in eggs of the Common Tern. Wilson Bulletin 83: 425-429. Hoffman, P. W. 1926. Nesting of the Black Tern in Wisconsin. Auk 43: 86-87. Langham, N.P. FE. 1972. Chick survival in terns (Sterna spp.) with particular reference to the Common Tern. Journal of Animal Ecology 41: 385-395. LeCroy, M. and C. T. Collins. 1972. Growth and survival of Roseate and Common Tern chicks. Auk 89: 595-611... LeCroy, M. and S. LeCroy. 1974. Growth and fledging in the Common Tern (Sterna hirundo). Bird-Banding 45: 326-340. MeNicholl, M. K. 1971. The breeding biology and ecology of Forster’s Tern (Sterna forsteri) at Delta, Manitoba. M.S. thesis, University of Manitoba, Winnipeg. MeNicholl, M.K. 1975. Larid site tenacity and group adherence in relation to habitat. Auk 92: 98-104. Richardson, F. 1967. Black Tern nest and egg moving experiments. Murrelet 48: 52-56. Ricklefs, R. E. 1967. A graphical method of fitting equa- tions to growth curves. Ecology 48: 978-983. Ricklefs, R.E. 1973. Patterns of growth in birds. II. Growth rate and mode of development. Ibis 115: 177-201. Shott, A. R. and F. W. Preston. 1975. The surface area of an egg. Condor 77: 103-104. Received 30 May 1978 Accepted 27 February 1979 Summer, Autumn, and Winter Diets of Elk in Saskatchewan HUGH M. HUNT Wildlife Research Division, Department of Tourism and Renewable Resources, 2602 8th Street East, Saskatoon, Saskatchewan S7H 0V7 Hunt, H. M. 1979. Summer, autumn, and winter diets of Elk in Saskatchewan. Canadian Field-Naturalist 93(3): 282-287. The composition of the contents of 88 Elk (Cervus elaphus) rumens obtained in summer, autumn, and winter from the boreal forest region of east-central Saskatchewan demonstrated that woody browse was the preferred forage of Elk. Relatively constant amounts of forbs (13-17% of rumen contents) and sedges (Carex spp.) and grasses (7—-8%) were present in the rumens in each season. Cultivated crops made up over one-quarter of the forage found in autumn rumens. Summer rumen compositions were characterized by similar high proportions of browse in rumens from each of four areas in east-central Saskatchewan and slight variations from area to area in the minor amounts of sedges and grasses, forbs, and agricultural crops. Significant differences from area to area in the individual component foods in each broad forage class in the rumens demonstrated the variable and adaptable nature of the diet of Elk in Saskatchewan. Key Words: Elk, food, browse, Saskatchewan, Cervus elaphus. Extensive habitat alteration has occurred in the boreal forest of Saskatchewan as a result of increased timber utilization and accelerated land clearing for agriculture along the forest fringe. Knowledge of the seasonal composition of the diet of any wildlife species is basic to understanding its ecology, particularly in terms of how habitat changes may affect food quantity and quality. An investigation of the diet of Elk or Wapiti (Cervus elaphus) in the boreal forest was undertaken because other food studies did not appear to be directly applicable to Saskatchewan Elk. In a review of 48 studies of the foods eaten by Elk in the western United States, Kufeld (1973) determined that grasses were the preferred forage of Elk in all seasons except summer when forb use was high. In the only published study of Elk diet in habitat closely comparable to Saskatchewan’s boreal forest, Riding Mountain National Park in Manitoba, woody browse was the preferred forage with grass and grass-like plants abundant in the diet only in spring and early winter (Blood 1966). Study Areas and Methods From 1973 to 1977, contents of 88 Elk rumens were sampled from the Porcupine Forest, the Squaw Rapids area, the Fort a la Corne Wildlife Manage- ment Unit, and the region surrounding Candle Lake in east-central Saskachewan (Figure 1). The distribution _ of rumen samples by area and season is givenin Table 1. The Porcupine Forest is predominantly a Trem- bling Aspen (Populus tremuloides) forest with some mixedwood (aspen —- White Spruce (Picea glauca)) portions. The Squaw Rapids area consists of aspen and Balsam Poplar (Populus balsamifera) stands, mixedwood stands (aspen — Balsam Poplar — White Spruce, aspen — Jack Pine (Pinus banksiana)) and softwood stands (Black Spruce (Picea mariana)) 282 along the Saskatchewan River, and the sedge (Carex spp.) meadow - willow (Salix spp.) complexes of the Cumberland Delta of the Saskatchewan River. Aspen, aspen-—Jack Pine, and Jack Pine stands characterize the Fort ala Corne Wildlife Management Unit whereas the area around Candle Lake is typical of the mixedwood section of the boreal forest as described by Rowe (1972) with aspen, White Birch (Betula papyrifera), White Spruce, Jack Pine, Black Spruce, and Tamarack (Larix lJaricina) the main tree species. Nomenclature for the vascular plants follows Moss (1959) and Budd and Best (1969); most common names are taken from Budd and Best (1969). The mean daily temperature in January in east- central Saskatchewan is about —20°C; mean July temperature is about 18°C. Annual preciptation averages about 42 cm, approximtely two-thirds of which falls as rain from April to October. Yearly snowfall averages about 130cm and mid- to late winter snow depths average about 45 cm. Causes of death of the sample animals were collisions with automobiles (8) and trains (1), sport- hunting (53), unregulated hunting (7), winter mor- tality (1), and a biological collection program (18). The rumen samples were from 5 male calves, 48 adult males, 3 female calves, 23 adult females, and 9 animals TABLE |—Distribution of rumen samples by area and season Area Porcupine Squaw Fort a la Candle Season Forest Rapids Corne Lake Summer 28 5) 13 4 Autumn 12 — 3} — Winter 13 l l 8 Total 53 6 17 12 1979 Nipawin Saskatchewan Kilometres 0510 20 30 40 A Candle Lake B Squaw Rapids C Fort ala Corne D Porcupine Forest SASKATCHEWAN HUNT: ELK DIETS, 23) Arbortield Hills 3) Wildcat Wilderness Area Posquia —— Preecevi/le FIGURE 1. Approximate limits of Elk range in east-central Saskatchewan from which the Elk rumens were obtained. of unknown sex and age. Most rumen samples were stored by freezing as suggested by Korschgen (1971) but a few were preserved in 10% formalin. After thawing, a 1.1-L (1- qt) aliquot of rumen content was washed through a 5.66-mm mesh screen (U.S. Standard Sieve Series Mesh No. 3'4) following Dirschl’s (1962) recom- mendation. The material remaining on the screen was placed on a white enamel tray, covered with water, and separated into pre-weighed petri dishes using forceps. Rumen components were identified macro- scopically to the lowest taxonomic level possible. Mushrooms were not identified beyond order, and grasses were identified only to family (Gramineae); fine material was sorted by forage class as uniden- tified herbaceous material or unidentified woody material. Dry weights of the components were recorded after they had dried for 24-48 h at 70°C. Data for the identified portions of the samples were grouped according to region and season regardless of sex or age class. The grouped data are presented by the aggregate percentage method of Martin et al. (1946). Seasonal groupings were made primarily according to plant phenological developments described by Stew- art et al. (1977), and correspond approximately to the periods 15 June - 30 September (summer), | Octo- ber — 30 November (autumn), and | December — 30 April (winter). Summer represented the period following new vegetative growth in spring until killing frosts halted annual growth, and autumn was the non- green period following summer before snow depths became a critical factor influencing forage avail- 284 ability and elk mobility. Winter was the period characterized by snow cover and lower temperatures although this period did not end with the disap- pearance of snow but rather with the commencement of new vegetative growth. Results and Discussion Browse was the major component in the diet of the Elk in east-central Saskatchewan, especially in sum- mer and winter (Figure 2). When leaves were available on the woody plants in summer and autumn, Elk consumed both leaves and twigs; twigs predominated in winter diets. Grasses and sedges were of minor importance in each season and forbs were of moderate importance. Agricultural crops were eaten when available in the autumn. Blood (1966) reported similar use of browse, grasses and sedges, and forbs by Elk in Riding Mountain National Park, Manitoba. Elk ate a variety of browse species. Aspen was the most important browse species both in proportion of rumen contents and frequency of occurrence (Table 2). Willows, the second most frequently occurring browse species, were common in rumens from each season. Wild Red Raspberry (Rubus strigosus), rose (Rosa spp.), and High-bush Cranberry (Viburnum trilobum) were common forages in summer although rose was found frequently in rumens from each season. The 10 browse species occurring most SUMMER (n=50) Browse 49.9°% WINTER (n=23) AUTUMN (n=15) FIGURE 2. Composition of the rumen contents from Elk in summer, autumn, and winter in east-central Sas- katchewan. THE CANADIAN FIELD-NATURALIST Vol. 93 frequently in the rumens were aspen (84% occur- rence), willow (74%), rose (63%), Wild Red Rasp- berry (45%), High-bush Cranberry (34%), Red-osier Dogwood (Cornus stolonifera) (30%), Saskatoon (Amelanchier alnifolia) (15%), Jack Pine (15%), Choke Cherry (Prunus virginiana) (13%), and Pin Cherry (Prunus pensylvanica) (13%). The five most frequently occurring species accounted for 82% of the dry weight of the browse identified. Blood (1966) found rose, aspen, willows, and Saskatoon were the most important browse items in rumens from Riding Mountain National Park, making up about 87% of the total browse segregated. Kufeld (1973) listed Saskatoon, aspen, Choke Cherry, and willow as highly valuable Elk forages. The abundance of most browse species in the rumens varied seasonally. Use of leaves of woody species was presumably related to their high levels of crude protein and total digestible nutrients as demon- strated by Stewart et al. (1977). Rose hips and the fruit and leaves of High-bush Cranberry were important foods in late summer. Significant use of fruits by White-tailed Deer (Odocoileus virginianus) has been reported by authors such as Healy (1971), Skinner and Telfer (1974), and Puglisi et al. (1978). Red-osier Dogwood was consumed in quantity only in autumn, and Pin Cherry was common only in summer and autumn rumens. Similar seasonal use of these two species by Moose (Alces alces) in Minnesota was reported by Peek et al. (1976). The frequency of occurrence of rose in the rumen samples decreased in winter probably because snow covered this low- growing shrub. Forbs were eaten mostly in the summer and autumn but were almost as important in winter. In Elk rumens from Riding Mountain National Park, Blood (1966) found forbs comprised a higher proportion in the summer (26%) than in this study (17%), but only one- half the level noted in this study in autumn and winter. The principal forbs in the Saskatchewan rumens were Wild Vetch (Vicia americana), Cream-colored Vetch- ling (Lathyrus ochroleucus), Wild Peavine (Lathyrus venosus), Prickly Sow-thistle (Sonchus asper), Can- ada Thistle (Cirsium arvense), Fireweed (Epilobium angustifolium), and horsetail (Equisetum spp.), whereas the principal forbs in the Riding Mountain rumens were Wild Peavine, Cream-colored Vetchling, Dandelion (Taraxacum officinale), Fireweed, and Coltsfoot (Petasites sagittatus) (Blood 1966). In recent cutovers in Saskatchewan, proliferation of invader or pioneer plant species such as Prickly Sow- thistle and Canada Thistle provided a source of forage; similar forages were probably not available to Riding Mountain Elk as logging is not permitted there. Blood (1966) concluded that certain forbs in Riding Mountain National Park were eaten in excess 1979 HUNT: ELK DIETS, SASKATCHEWAN 285 TABLE 2—Detailed seasonal composition of rumen contents from Elk in east-central Saskatchewan. Forages comprising <1% of the rumen contents are grouped into the “others” categories and the mean proportion of unidentified material in the samples is also included Summer (n=50) Autumn (n=15) Winter (n=23) Food % Dry % Dry % Dry categories weight Frequency weight Frequency weight Frequency Trembling Aspen (Populus tremuloides) 11.9 72.0 20.7 100.0 26.4 100.0 Willow (Salix spp.) ES 70.0 1.5 73.3 15.9 82.6 Raspberry (Rubus strigosus) 11.6 62.0 _ — - 34.8 Rose (Rosa spp.) 12.3 80.0 — 60.0 — = High-bush Cranberry (Viburnum trilobum) 16.3 48.0 1.7 4.0 — = Red-osier Dogwood (Cornus stolonifera) o — 8.3 66.7 2 39.1 Pin Cherry (Prunus pensylvanica) 4.4 18.0 Del 13.3 — = Saskatoon (Amelanchier alnifolia) — — od} 20.0 — — Jack Pine (Pinus banksiana) 1.3 16.0 1.8 13.3 1.9 13.0 Choke Cherry (Prunus virginiana) a= _ — — 2.4 Dllel/ Honeysuckle (Lonicera spp.) = == = = 1.9 13.0 Others 3.4 _ 3.4 = 2.0 — Unidentified browse 5.9 44.0 9.1 U3) 19.1 Slee. Total browse 74.6 49.9 72.8 Sedge (Carex spp.) 2.0 46.0 — 40.0 4.7 39.1 Grass 4.9 68.0 5.8 60.0 82 39.1 Prickly Sow-thistle (Sonchus asper) 1.2 22.0 5.9 33.3 Dey 17.4 Vetchling, Peavine (Lathyrus spp.) 4.2 22.0 — = 1S 13.0 Vetch (Vicia americana) 30) 52.0 — 26.7 = = Mushroom EY 18.0 — — — — Fireweed (Epilobium angustifolium) 1.9 36.0 — _— oe Horsetail (Equisetum spp.) _ — 2.8 13.3 2.2 8.7 Canada Thistle (Cirsium arvense) -- -- — — 4.6 13.0 Alfalfa (Medicago sativa) 19) 2.0 1.6 33) 4.0 4.3 Barley (Hordeum vulgare) — 10.8 40.0 _— Oats (Avena sativa) a _ 5.9 33.3 _ Wheat (Triticum aestivum) — _ 3.1 6.7 2.0 4.3 Cereal stems — _— 6.0 26.7 — Others Del — Spl _ DED os Unidentified herbs _— _ Bal 20.0 _— — Total herbs 25.4 50.1 DY) Unidentified material 37.6 96.0 40.4 9353 Dei 86.9 of their availability while some other abundant forbs made up a small or negligible part of the diet. The same was probably true for east-central Saskatche- wan. Cultivated crops such as Wheat (Triticum aesti- vum), Oats (Avena sativa), Barley (Hordeum vul- gare), and Alfalfa (Medicago sativa) were used extensively in autumn. This may cause significant conflicts along the forest-farmland interface, par- ticularly in years of weather-delayed harvests. Elk use of agricultural crops was not mentioned by Blood (1966), and Alfalfa is the only agricultural crop present in the Saskatchewan rumens included in Kufeld’s (1973) list of Elk forages. Agricultural crops may be particularly important to Elk in autumn for maintenance of good body condition prior to the lengthy period of negative energy balance that Stewart et al. (1977) describe as characteristic for Moose in winter. Mid- to late summer rumen compositions for the four areas consisted predominantly of woody plant material (Figure 3). The minor amounts of sedges and grasses, forbs, and agricultural crops varied slightly from area to area. Examination of a more detailed breakdown of the rumen compositions for the four areas (Table 3) indicated that although the propor- tions of the browse forage class for each area were relatively similar, the individual component foods of the browse and other forage classes differed from area to area. For example, the proportion of aspen in the rumens ranged from over half at Candle Lake to only trace amounts at Squaw Rapids. Willows were 286 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 3—Composition of the rumen contents from Elk in summer in four areas of east-central Saskatchewan. Forages comprising <1% of the rumen contents are grouped into the “others” categories and the mean proportion of unidentified material in the samples is also included Porcupine Squaw Rapids Fort a la Corne Candle Lake (n=28) (n=5) (n=13) (n=4) Food % Dry % Dry % Dry % Dry categories weight Frequency weight Frequency weight Frequency weight Frequency Trembling Aspen (Populus tremuloides) V3 71.4 = = 13.3 84.6 53.7 100.0 Willow (Salix spp.) 2.9 SA BoP) 60.0 Sal 9253 DES 100.0 Rose (Rosa spp.) 14.5 85.7 11.2 80.0 11.6 76.9 — 50.0 Raspberry (Rubus strigosus) 10.7 53.6 27.6 100.0 10.6 76.9 1.4 50.0 High-bush Cranberry (Viburnum trilobum) — 21.2 60.7 41.4 100.0 1.1 15.4 — = Balsam Poplar (Populus balsamifera) 2 17.9 = — — — — = Pin Cherry (Prunus pensylvanica) 3.1 el — — 10.4 53.9 — — Manitoba Maple (Acer negundo) = 3.6 — —_ — _ — — Choke Cherry (Prunus virginiana) — — — _ 1.1 15.4 — — White Birch (Betula papvrifera) 1.0 3.6 — —_— — — — _ Currant, Gooseberry ( Ribes spp.) = = = = = — DT 25.0 Jack Pine (Pinus banksiana) — — — _ 4.2 53.9 2.4 25.0 Others 2.1 ~ 0.4 — 1.1 — 1.0 = Unidentified browse US 53.6 10.7 80.0 2.6 23.1 — —_ Total browse ales) 9355 Tales 83.1 Sedge (Carex spp.) Dod) 53.6 1.5 60.0 — — 2.4 50.0 Grass 8.0 75.0 — _ 1.5 76.9 — — Prickly Sow-thistle (Sonchus asper) 1.4 14.3 2.2 20.0 a _ Bo) 75.0 Vetchling, Peavine (Lathyrus spp.) 6.0 32.1 _ 333 Hell - = Vetch (Vicia americana) 5.3) 57.1 = — 9.2 53.9 — 25.0 Sarsaparilla (Aralia nudicaulis) _ 1.1 20.0 — — = 25.0 Fireweed (Epilobium angustifolium) 3.0 42.9 -— — — 1.3 50.0 Cow-parsnip (Heracleum lanatum) — — — — — = 12 25.0 Mushroom — _ i _ od) 46.2 4.2 25.0 Lichen (Cladonia spp.) — — — — — — 2.9 25.0 Barley (Hordeum vulgare) — _ — — 7.4 Ali — — Others 0.9 _ 1.7 — Dp? _ Dell _ Unidentified herbs 1.4 21.4 — — — = — Total herbs 28.5 6.5 28.9 16.9 Unidentified material 41.2 96.4 Dia 100.0 38.6 100.0 22.4 75.0 common in the rumens from the Candle Lake area and Fort a la Corne but less abundant in the rumens from the Porcupine Forest and the Squaw Rapids region. Pin Cherry was a significant component only in the samples from the Porcupine Forest and Fort a la Corne; Jack Pine was found only in rumens from Fort a la Corne and the Candle Lake area. High-bush Cranberry formed a major part of the food in rumens from the Porcupine Forest and Squaw Rapids area but was less common in rumens from the other areas. Mushrooms formed a large part of the non-woody forage in samples from both Fort a la Corne and the Candle Lake area whereas lichens (Cladonia spp.) appeared in notable quantity only in the rumens from the Candle Lake region. Deer have also been noted to eat mushrooms, but the significance of this is unknown (Healy 1971; Skinner and Telfer 1974). The cosmopolitan nature of the diet of Elk likely is a factor that enables them to thrive in many types of habitat. This study demonstrates the importance of twigs and leaves of woody plants in the summer, autumn, and winter diet of Elk in east-central Saskatchewan in contrast to Kufeld’s (1973) synthesis of Elk diets in the western United States in which grasses are the preferred forage. Land clearing along the forest fringe and subsequent planting of exotic grasses to enhance cattle production will make those areas less attractive to Elk as supplies of many of the preferred forages will be reduced. Clearcut logging in the commercial forest will stimulate production of most of the preferred Elk forages but forest managers must ensure that the proper balance between food and cover is maintained. 1979 PORCUPINE (n=28) SQUAW RAPIDS (n=5) Sedge,Grass 2.4% FORT A LA CORNE (n=13) CANDLE LAKE (n=4) FIGURE 3. Composition of the rumen contents from Elk in summer from four areas in east-central Saskatch- ewan. Acknowledgments This project was part of a study of the ecology of Elk in Saskatchewan conducted by the Wildlife Research Division of the Saskatchewan Department of Tourism and Renewable Resources. I am indebted to R. Tennent, D. Arneson, D. Phillips, and B. Weichel for analysis of the rumen samples. The assistance of many department members in collecting rumen samples and for review of the manuscript is also appreciated. HUNT: ELK DIETS, SASKATCHEWAN 287 Literature Cited Blood, D. A. 1966. Range relationships of Elk and cattle in Riding Mountain National Park, Manitoba. Canadian Wildlife Service Wildlife Management Bulletin, Series 1, Number 19. 62 pp. Budd, A.C. and K.F. Best. 1969. Wild plants of the Canadian prairies. Canada Department of Agriculture Publication 983. 519 pp. Dirschl, H. J. 1962. Sieve mesh size related to analysis of antelope rumen contents. Journal of Wildlife Manage- ment 26(3): 327-328. Healy, W. M. 1971. Forage preferences of tame deer in a northwest Pennsylvania clear-cutting. Journal of Wildlife Management 35(4): 717-723. Korschgen, L. J. 1971. Procedures for food-habits analysis. In Wildlife management techniques. Edited by R.H. Giles, Jr. Wildlife Society, Washington, D.C. pp. 233-250. Kufeld, R. C. 1973. Foods eaten by the Rocky Mountain Elk. Journal of Range Management 26(2): 106-113. Martin, A. C., R. H. Gensch, and C. P. Brown. 1946. Al- ternative methods in upland gamebird food analysis. Journal of Wildlife Management 10(1): 8-12. Moss, E. H. 1959. Flora of Alberta. University of Toronto Press, Toronto, Ontario. 546 pp. Peek, J. M., D. L. Urich, and R. J. Mackie. 1976. Moose habitat selection and relationships to forest management in northeastern Minnesota. Wildlife Monographs, Num- ber 48. 65 pp. Puglisi, M. J.,S. A. Liscinsky, and R. F. Harlow. 1978. An improved methodology of rumen content analysis for White-tailed Deer. Journal of Wildlife Management 42(2): 397-403. Rowe, J.S. 1972. Forest regions of Canada. Canada Department of Environment, Canadian Forestry Service Publication Number 1300. 172 pp. Skinner, W.R. and E.S. Telfer. 1974. Spring, summer, and fall foods of deer in New Brunswick. Journal of Wildlife Management 38(2): 210-214. Stewart, R. R.,R. R. MacLennan, and J. D. Kinnear. 1977. The relationship of plant phenology to Moose. Saskatche- wan Department of Tourism and Renewable Resources Technical Bulletin Number 3. 20 pp. Received 4 November 1978 Accepted 28 March 1979 Taxonomy, Distribution, and Ecology of the Cliff-brake Ferns (Pellaea: Polypodiaceae) in Alberta DANIEL F. BRUNTON Box 59, Canmore, Alberta TOL O0MO Brunton, Daniel F. 1979. Taxonomy, distribution, and ecology of the cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta. Canadian Field-Naturalist 93(3): 288-295. The genus Pellaea in Alberta consists of three species of calcicolous rock ferns. Pellaea atropurpurea, an apogamous triploid, is rare in the province. It is locally abundant in the Bow Valley of western Alberta, preferring hot arid sites on partially-shaded, heavily fractured limestone cliff-faces with a southwestern exposure. Pellaea occidentalis, a sexual diploid, is uncommon in Alberta but is locally abundant in the Bow Valley of western Alberta. It is found on arid south- or southwest-facing, open limestone cliff-faces. Pellaea suksdorfiana, an apogamous tetraploid, is uncommon in western Alberta where it ranges throughout the mountains. It prefers cool shaded, east- to north-facing limestone or calcareous sandstone cliff-faces, usually near water. Pellaea occidentalis and P. suksdorfiana are distinct from the eastern P. glabella, an apogamous tetraploid, on the basis of distinctive characteristics of range, morphology, cytology, and ecology. Characteristics for separating P. suksdorfiana, P. occidentalis, and P. atropurpurea are provided. Forty-five previously undocumented records for Alberta Pellaea are listed. Key Words: Pellaea atropurpurea, Pellaea occidentalis, Pellaea suksdorfiana, taxonomy, distribution, ecology, cytology, Alberta, Polypodiaceae. The cliff-brake ferns ( Pe//aea) have been described as “.. comparatively rare in Canada or at most locally abundant...” (Rigby and Britton 1970). This state- ment still applies to the status of these small calci- colous rock ferns despite substantial additional study in parts of Ontario and Quebec (Lafontaine and Brunton 1972; Brunton and Lafontaine 1974). In western Canada the genus is represented by three species, P. atropurpurea(L.) Link, P. occidentalis (E. Nels.) Butters, and P. suksdorfiana Butters. The latter two are frequently treated as varieties of the eastern P. glabella. This study was initiated to determine whether ecological evidence exists to support specific status for the three Alberta taxa of Pe/laea and to gain a better understanding of its distribution and abundance in the province. Taxonomy The taxa of Pe/laea in Alberta have been described under several names and/or combinations at a species and varietal level. Close morphological similarity has encouraged this, even though there are clear cytolo- gical differences (Tryon 1957). The three Alberta species of Pellaea are as follows: 1) Pellaea atropurpurea (L.) Link 2) Pellaea occidentalis (E. Nels.) Butters = P. glabella Mett. ex Kuhn var. occiden- talis (E. Nels.) Butters P. glabella Mett. ex Kuhn var. nana (Richards.) Cody = P. pumila Rydb. 3) P. suksdorfiana Butters = P. glabella Mett. ex Kuhn var. simplex (E. Nels.) Butters = P. atropurpurea (L.) Link var. simplex (Butters) Morton Few authors today question the validity of P. atropurpurea as a distinct species (Tryon 1957, and others). It is an apogamous triploid (n and 2n = 87). It is also the largest species of Canadian Pellaea. It occurs across the country from British Columbia to Quebec and exhibits a distinctive morphology. The questions arise with P. occidentalis and P. suksdor- fiana, which are often treated as varieties of P. glabella. Pellaea occidentaiis is a sexual diploid (n = 29, 2n = 58) and is a tiny plant of the dry interior of North America. Pellaea suksdorfiana is an apogamous tetraploid (n and 2n = 116) which is a species of the Cordilleran region of western North America. Both exhibit distinctive morphologies and ecology. The species with which they are often combined, P. glabella, is an apogamous tetraploid (n and 2n = 116). Although maintaining an equivalent cytology to that of P. suksdorfiana, it is disjunct from that taxon by over 1300 km. It is quite clearly different in morpho- logy and cytology from P. occidentalis. Without the benefit of known ecological differences and with limited range data available, it is understandable that many authorities have preferred to treat these as one species. A. F. Tryon (Gray Herbarium, Harvard University, personal communication 1977) is in- 288 1979 clined to retain this status for the utility of indicating relationships in the genus. On the basis of more complete range data and the demonstration of ecological preferences of each species which were observed during our study, we feel that there is sufficient evidence to justify each taxa being treated specifically. ’ Banff. Bt AS : Mt Mt i Cory Norquay\ A A Jasper National Park C ) Unpublished and/or new stations A Previously published records National (Rigby and Britton. 1970) Park — ~s kilometres Kananaskis Provincial A Park —— BRUNTON: PELLAEA IN ALBERTA ‘Three A Sisters 289 Distribution and Ecology Pellaea atropurpurea (Purple Cliff-brake) (Figures 1, 2), This fern is known sparingly in Canada from British Columbia to Quebec. It is considered to be the rarest Pellaea in Canada (Rigby and Britton 1970), although recently it has been found to be locally abundant in 4 kilometres ” Ga] CALGARY ye ~~ Bow X ie angina ne © n eT haa hbridge Ue fil ¢ FIGURE |. Distribution of Pe//laea atropurpurea in Alberta. 290 parts of Ontario and western Quebec (White 1977). Only recently noted in the province by the revision of a specimen collected in 1900 at Banff (Rigby and Britton 1970), it is now known from I1 sites in Alberta. These are all in or near the Bow Valley, alow dry valley in the eastern slopes of the Rocky Mountains. The additional stations are listed below. Herbarium acronyms follow Holmgren and Keuken (1974) with the exception of DFB which represents the author’s personal herbarium. FIGURE 2. Robust clump of Pellaea atropupurea growing in wide fissure in southwest-facing limestone cliff-face. Gap Lake, Exshaw, Alberta, 18 February 1978. 4 mi [6.4 km] E of Canmore, Porsild 18147 in 1951 (CAN), Brunton 1289 in 1976 (DFB,DAO, CAN); 1.5 mi [2.4 km] SE of Exshaw, Bird 18500 in 1967 (UAC), Brunton 1307 in 1977 (DFB); 4 mi [6.4 km] SE of summit of Mount Head, Bird 28833 in 1972 (UAC); 2.5 km NE of Exshaw, Brunton and Brunton 1247 in 1976 (DFB,DAO,ALTA,CAN); 2 km NE of Exshaw, Brunton and Grant 1522 in 1977 (DFB,CAN); 2.5 km W of Exshaw, Brunton 1530 in 1978 (DFB); 4.5km W of Exshaw at Gap Lake, THE CANADIAN FIELD-NATURALIST Vol. 93 Brunton 1528 in 1978 (DFB,DAO): 1 km NE of Exshaw, Brunton and Coneybeare 1532 in 1978 (DFB,CAN); 2.5 km S of Kananaskis Environmental Sciences Center, Barrier Lake, Brunton 1538 in 1978 (DFB); 12 mi[19.3 km] S of Seebe, Cody, Jaques and Corbin 23037 in 1977 (DAO): southeast Barrier Lake, Kananaskis Valley, Brunton 1623 in 1979 (DAO). The species is exceptionally abundant at several sites, particularly the Grotto Mountain and Gap Lake sites (Porsild 18147 and Brunton 1289, and Brunton 1528 respectively). Alberta specimens are typically less than 20 cm tall. Only at the above sites (and at Barrier Lake, Cody et al. 23037) were plants noted which approached 30 cm in height, a condition more typical of eastern Canadian plants. This is likely a reflection of the much drier climate of Alberta. We observed that P. atropurpurea in drier-than-normal sites in Quebec were smaller than the typical. Alberta P. atropurpurea is found on dry steep exposed, heavily-fractured limestone rock slopes or cliffs. With one exception (Heart Mountain, Bird 18500), the stations exhibited a southwestern ex- posure. Individual plants were partially shaded by rock ledges and/or scattered woody vegetation, usually Ribes oxycanthoides and/or Potentilla fruti- cosa. The plants grow out from under a sheltering ledge or boulder and are found only in those crevices in which a considerable amount of soil and/or plant debris has accumulated. The site preference was very consistent with other Canadian site descriptions (Brunton and Lafontaine 1974). The maximum elevation noted for any station of P. atropurpurea was 1650 m above sea level. In almost every case, the Woolly Lip-fern (Chei- lanthes feei) was a close (and frequently abundant) associate. These warmer-than-normal sites were also typified by a number of montane species which are very local in distribution in Alberta. They include Douglas Fir (Pseudotsuga menzieii), Limber Pine (Pinus flexilis), and Rocky Mountain Juniper (Juni- perus scopulorum — possibly J. scopulorum X Juni- perus Ssp.). Purple Cliff-brake, being the largest and easiest to collect of Alberta Pe//aea, is probably represented in herbaria in greater than natural frequency vis. other Pellaea. Pellaea suksdorfiana (Smooth Cliff-brake) (Figures 3, 4). This species occurs in Canada only in the Cordil- leran region of British Columbia and Alberta, where it is considered to be uncommon and local (Taylor 1970: Rigby and Britton 1970). Prior to this study, it was known from five locations in Alberta. Twenty additional stations are listed below: L979 Spray Falls, Banff N.P., Warren in 1934 (ALTA): 3 mi [4.8 km] W of the Kananaskis Forest Experi- ment Station, Porsild and Lid 19424 in 1956 (CAN): 5 mi [8.1 km] S of Kananaskis Forest Experiment Station, Porsild and Lid 19424 in 1956 (CAN); E of Corral Creek, De Smet Range, Jasper N.P., Porsild 21285 in 1959 (CAN); Talbot Lake, Jasper N.P., Porsild 22377 in 1960 (CAN); Bearspaw Dam, Calgary, A. Birdin 1965 (UAC), Brunton 1534 in 1978 (DFB); entrance near Hinton, Pegg 2095 in 1965 (CAN,ALTA); southwest side Lac des Arc, Bow BRUNTON: PELLAEA IN ALBERTA 291 Valley, Brunton 1139 in 1976 (DFB); W of Barrier Pass, Jaques 5059 in 1974 (UAC), Brunton 1293 in 1976 (DAO,DFB): E of Jasper, Jasper N.P., Ogilvie in 1960 (UAC); west side Heart Mountain 1.5 mi [2.4 km] SE of Exshaw, Brunton 1308 in 1977 (DFB): 2.5 km W of Exshaw, Brunton and Paton 1337, 1338 in 1977 (DFB), Brunton 1531 in 1978 (DFB); 4.5 km W of Exshaw at Gap Lake, Brunton 1529 in 1978 (DFB); | km NE of Exshaw, Brunton and Coney- beare 1533 in 1978 (DFB); 12 mi[19.3 km] of Seebe, Cody, Jaques and Corbin 23038 in 1976 (DAO). The ‘Three A Sisters Jasper National Park @ Unpublished and/or new stations Banff National Park — A Previously published records (Rigby and Britton. 1970) 100 kilometres Kananaskis Provincial Park 47 CALGARY 5; a Bow AD UF Prato SPE FIGURE 3. Distribution of Pellaea suksdorfiana in Alberta. 292 Sight Records: 6.5 km E of Canmore on Grotto Mountain (Brunton 1977); north side Banff Airfield, Banff N.P. (Brunton 1978); south side Tunnel Mountain, Banff N.P. (Brunton 1978); west side Hwy 40 at Barrier Lake (Brunton 1978); east side Hwy 40 at Barrier Lake (Brunton 1978). Virtually all of these stations are from cliffs of limestone. All but two of the stations near Exshaw (Brunton 1529 and Brunton and Coneybeare 1533) and one by Barrier Lake (Jacques 5059, Brunton 1293) are adjacent to water bodies. These latter stations are on north- or northeast-facing slopes. FIGURE 4. Typical frond of Pellaea suksdorfiana growing out from fissure in southeast-facing limestone cliff-face. Gap Lake, Exshaw, Alberta, 18 February 1978. Pellaea suksdorfiana prefers cool heavily-shaded, less arid sites than the other Alberta Pellaeas, normally selecting waterside sites with an eastern exposure. It often grows in thin fissures in open rock. Rarely exceeding I5cm in height (and usually smaller) it is, nonetheless, typically larger than P. occidentalis. In those sites which are south-facing (e.g., Brunton and Paton 1338) there is heavy shade from surrounding trees. Individual clumps are open and sprawling, with fronds extending well out from the rock-face. THE CANADIAN FIELD-NATURALIST Vol. 93 Pellaea suksdorfiana is uncommon throughout the Rocky Mountains of Alberta. It extends out onto the prairie in two sites (near Cochrane and Calgary). For a morphological comparison of this species with P. occidentalis see Table 1. Pellaea occidentalis (E. Nels.) Butters (Western Cliff- brake) (Figures 5, 6). This is a species of the dry interior plains and low mountains of central North America. It is not found W of Alberta or E of Manitoba in Canada (Rigby and Britton 1970; Taylor 1970; Tryon 1957). Only in Wyoming does it extend across the Continental Divide into areas of moister climate (Hitchcock et al. 1969). It has the most exclusive range of any Canadian Pellaea, overlapping with others only in southwestern Alberta. Prior to this study it was known from three locations in Alberta. Fourteen additional stations are listed below: Southwest slope Sawback Range, Banff N.P., Por- sild and Breitung 15482 in 1946 (CAN); southern slope Mount Cory, Banff N.P., Porsild 17993 in 1951 (CAN); Kootenay Plains W of Nordegg, Dumais in 1976 (ALTA,DFB); Grotto Mountain, 6.5 km E of Canmore, Brunton 1290 in 1976 (DFB,DAO); south- east end Mount Rundle, Canmore, Brunton and Paton 1328 in 1977 (DFB,DAO): north side Barrier Lake, Cody, Jaques and Cordin 23043 in 1976 (DAO): north side Crowsnest Lake, Crowsnest Pass, Brunton 1613 in 1978 (DFB,DAO). Sight Records: 2.5 km NE of Exshaw (Brunton 1976); 2 km NE of Exshaw (Brunton 1977); 4.5 km W of Exshaw at Gap Lake (Brunton 1978); south side Stoney Squaw Mountain, Banff N.P. (Brunton 1978); southwest side Mount Norquay, Banff N.P. (H. F. Coneybeare 1978); north side Banff Airfield, Banff N.P. (Brunton 1978); 2.5 km W of Exshaw (Brunton 1978). Western Cliff-brake is the smallest Canadian Pellaea, rarely exceeding 10 cmin height. The Alberta stations are on exposed hot limestone cliffs and rock slopes with a southern or southwestern exposure. In every station examined, Cheilanthes feei was a close associate. It is typically found on the most exposed cliffs and rock areas. It is locally very abundant in the Bow Valley. The well-known Tunnel Mountain station in Banff National Park supports hundreds of thousands, if not millions, of plants. It is the largest site known to date. This species grows in very dense clumps in a “pin- cushion” shape out of pocks and/or fissures in the rock face. The fronds rarely extend out beyond the surface of the rock face (as a water conservation measure?). Despite the fact that it grows close to the British Columbia border in Banff National Park and 1979 BRUNTON: PELLAEA IN ALBERTA 293 the Crowsnest Pass, there are no records for this species W of the Continental Divide in Canada. The increase in precipitation on the west side of the divide is dramatic and appears to create unsuitable con- ditions for P. occidentalis, even on sites which seem to be otherwise quite suitable. ’ Banff Mt ie ie Cory Norquay\ _ KS Jasper National Park ® Unpublished and/or new stations A Previously published records (Rigby and Britton. 1970) kilometres Kananaskis Provincial ‘Three A Sisters Ona ae ae “ageonogtion (pn a : ont : é & * Sr Outside of the Bow Valley, the Western Cliff-brake is rare in Alberta. Being very inconspicuous, however, it may well have been overlooked. It should be watched for elsewhere in dry valleys along the eastern slopes of the Rocky Mountains and on cliff areas in the prairie (e.g., the Milk River canyon). 4 kilometres 6 8 10 \2, CALGARY & Rive’ i eae Suen uF Feats Whe SL FIGURE 5. Distribution of Pellaea occidentalis in Alberta. 294 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE 6. Typically dense clump of Pellaea occidentalis growing in hair-line fissure in open, south-facing limestone rock face. South (5 km) of Canmore Alberta, | November 1977. TABLE 1—Comparison of characteristics of Pellaea atropurpurea, P. occidentalis, and P. suksdorfiana. Source: 1, Rigby and Britton (1970); 2, Tryon (1957); 3, This study Characteristics Chromosome no. No. of spores per sporangium Frond length Frond displacement* Rachis color* Stipes* Pinnae* Pinnae angle to rachis Site preference* P. occidentalis 2n = 58 64 (small) Maximum |[5 cm Dense, thickly clumped “pin cushions” flush with rock face Golden to dark brown Thin and brittle old stipes inconspicuous Mitten-shaped and sessile; margins strongly reflexed Perpendicular Dry exposed SW-facing limestone P. suksdorfiana 2n = 116 32 (large) Maximum 20 cm Open and spreading, extending out beyond rock face Reddish-brown to brown-purple Thicker and sturdy, rarely sparsely pubescent, old stipes conspicuous and often silvery color Oblong-lanceolate and (at least lower) petioled; margins strongly reflexed Acute Shaded cool, E-or N-facing calcareous cliffs by water P. atropurpurea 2n = 87 32 (large) Maximum 35 cm Erect and spreading, extending up and outward from ledges far beyond rock face Purple to blackish-purple Thick and sturdy, heavily pubescent old stipes abundant, silvery color Lanceolate to oblong- lanceolate and petioled; margins slightly reflexed Broad Dry partially-shaded, SW-facing limestone Source 1 1,2 13 E253 *Feature of particular value for field identification. 1979 Discussion There are many more stations for Pellaeain Alberta than was previously thought. It isa common group on the limestone cliffs and rock faces of the Bow Valley in western Alberta. It is reasonable to expect that more stations will be found in other drier mountain valleys along the eastern slopes of the Rocky Mountains. Pellaea suksdorfiana is the most frequently ob- served Cliff-brake in Alberta, occurring uncommonly in cooler shaded, more moist cliffs. Pellaea atro- Purpurea appears to be locally common on sites similar to that for P. occidentalis, both preferring hot dry open limestone cliffs and rock faces. Pellaea atro- purpurea prefers a more shaded and more heavily fractured rock than does P. occidentalis. With three of the four Canadian species occurring in Alberta, the Pellaea flora of the province is particularly rich. With so many stations growing in close association in the Bow Valley, there is an excellent opportunity to study the ecology of these ferns and to search for (as yet unknown) hybrids. As Tryon (1957) suggests that P. suksdorfiana may have arisen as a fertile hybrid between P. atropurpurea and P. occidentalis, this becomes an interesting possi- bility. Acknowledgments My thanks to A. F. Tryon, M. Dumais, H. F. Coneybeare, W.J. Cody, D.M. Britton, and D. Jaques for information concerning their collections and observations. The curators of CAN, DAO, ALTA, and UAC provided access to these collections. J. M. Gillett provided valuable assistance at CAN, as did W. J. Cody and C. Frankton at DAO. In addition, BRUNTON: PELLAEA IN ALBERTA 295 Cody loaned a number of specimens to me for examination. D. M. Britton, University of Guelph, reviewed an earlier manuscript and provided valuable criticism. B. Wackerle typed several drafts of the manuscript. I was accompanied in the field on several occasions by D. Paton, H. F. Coneybeare, J. D. Lafontaine, and B. Mitton. The late A. E. Porsild provided important collection data early in the work and encouraged this study. A particular vote of thanks is owed my daughter Joni who hovered precariously with me on numerous cliffs without complaint. Literature Cited Brunton, D. F. and J. D. Lafontaine. 1974. The distribu- tion of Pellaea in Quebec and eastern Ontario. Naturaliste Canadien 101: 937-939. Hitchcock, C. L., A. L. Cronquist, and M. Ownbey. 1969. Vascular plants of the Pacific Northwest. Volume 1. University of Washington Press, Seattle. Holmgren, P.K. and W. Keuken. 1974. Index herbar- iorum, Part I. The herbaria of the world, 6th edition. Regnum Vegetabile 92: 1-397. Lafontaine, J.D. and D.F. Brunton. 1972. The Purple Cliff-brake, Pellaea atropurpurea (L.) Link, in western Quebec. Canadian Field-Naturalist 86: 297-298. Rigby, S.J. and D. M. Britton. 1970. The distribution of Pellaea in Canada. Canadian Field-Naturalist 84: 137-144. Taylor, T. M. C. 1970. Pacific Northwest ferns and their allies. University of Toronto Press, Toronto. Tryon, A. F. 1957. A revision of the fern genus Pellaea section Pellaea. Annals of the Missouri Botanical Garden 44: 125-193. White, D. 1977. Rare plant survey: revisions. Trail and Landscape 11: 22-25. Received 29 November 1978 Accepted 31 March 1978 Characteristics of Peregrine Falcons Migrating through Central Alberta, 1969-1978 DICK DEKKER 3819 — 112A Street. Edmonton, Alberta T6J 1K4 Dekker. Dick. 1979. Characteristics of Peregrine Falcons migrating through central Alberta. 1969-1978. Canadian Field- Naturalist 93(3): 296-302. In central Alberta from 1969 to 1978, 541 sightings of migrating Peregrine Falcons (Falco peregrinus) were recorded in spring, and 34 in fall. Exclusion of suspected duplications gave 226 spring migrants and 27 fall migrants. In spring, adults arrived an average of 9 d before the immatures. The great majority (86%) of spring migrants passed through the study area between 4 May and 23 May. Presence of adults peaked from 4 May to 7 May, of immatures from 12 May to 15 May. Fall sightings occurred from 10 September to 4 October. In spring, males comprised 33% of the adult sample and 35% of the immatures identified. Aggression between the sexes is suggested as a reason for the observed imbalance in the sex ratio. The proportion of adults to immatures in spring was | to |. Spring migrants that were seen to leave the study area ina northerly direction did so by soaring and high-altitude sailing. Strong head winds did not appear to deter some adults from migrating. Foraging activity was seen to take place throughout the day and as late as | h after sundown. Key Words: Peregrine Falcon, Falco peregrinus, migrations, Alberta. Fall migration of northern Peregrine Falcons has been studied along the coastal and Great Lakes flyways (e.g., Hunt et al. 1975: Mueller and Berger 1961: Ward and Berry 1972), but very little infor- mation has been recorded about their spring migra- tion. Isolated spring sightings of peregrines have been reported from across the continent, as evidenced by scattered notes in American Birds. During a spring study of migrating raptors along Lake Ontario only four peregrines were seen in 108 d (Haugh and Cade 1966). The paucity of data has led to speculation about the routes, timing, and age structure of the peregrines moving north. The fact that few subadults have been observed on the breeding grounds suggested that mortality of first-year birds is high. Cade (1960) postulated that some immature peregrines may not return to the Arctic and Subarctic until their second year. This paper provides information on the timing of spring and fall migrations of peregrines through Alberta, and the age, sex, and behavior of these migrants. Study Area and Methods The study area is a crescent-shaped strip of land situated on the rolling plains of central Alberta and is roughly 15 X 2 km. Bordered by a marshy lake, it is dominated by grain fields and pastures which hold scattered pockets of Trembling Aspen (Populus tremuloides) and willow (Salix spp.). Shallow depres- sions collect melt-water in spring, attracting ducks and shorebirds in large numbers during that season. The study area was regularly visited from late winter to late autumn to observe the hunting habits of raptorial birds. During the 47-d period 15 April to 31 May, visits to the area averaged 27.9 d in each of the 10 yr. During the 45-d period from 1 September to 15 October visits averaged 14.4 d. Days afield varied, some lasting from sunrise until nightfall and others spanning only 3 or 4h during the afternoon or morning. Actual hours afield in spring were tabulated only in the last 5 yr (Table 1). In searching for peregrines, no set procedures were followed. Vehicular travel in the study area was limited, especially during wet periods. Most points were accessibie only on foot, and 5-25 km were covered by hiking each day. Resting peregrines were detected by scanning the landscape through 10-power binoculars. In calm weather, the falcons sat on posts or stones and were readily found, but strong winds induced them to shelter on the ground. Very rarely were trees used as perches. As a rule, to avoid flushing them, resting peregrines were not approached closely. No attempts at trapping or photography were made. Most falcons were watched until they left of their own accord. The majority of sightings were of flying peregrines. To maximize chances of seeing hunting falcons, I stayed near points where foraging peregrines had previously been observed. Alarm reactions of prey species, especially shorebirds, usually signalled the appearance of a falcon. Often, while I was quietly sitting in the open or in a parked car, the falcons seemed to ignore my presence and approached boldly, providing numerous close-range views. As the various age and sex groups differ in appearance, and some 296 1979 birds show individual differences in plumage and behavior, individuals sometimes could be recognized with certainty: this helped me to minimize duplicate counts. The following procedure was used to reduce possible duplications. When a bird identifiable as to age and sex, for instance an immature female, was observed on several consecutive days, each observa- tion was recorded for the maximum count, but only one sighting of an immature female was included in the minimum count, unless there was evidence of more than one immature female falcon present in the study area. Sightings of unidentified peregrines were included in the maximum count even though sex and age were unknown, but not inthe minimum count if a classified peregrine was known to be present in the study area at the time. Two or more interacting falcons were seen fre- quently in aggressive encounters or in what appeared to be cooperative hunting. Whenever this happened. I could assess their numbers more accurately. The passage of peregrines, like that of other migratory birds, is seldom an evenly-spaced phe- nomenon. There were periods when none were seen in the study area, followed by days or parts of days when several appeared in rapid succession. The rate of arrival and departure influenced my ability to calculate the number of individual peregrines seen passing through the study area. I believe that the true number is considerably less than the maximum count presented but somewhat higher than the minimum figure. Results and Discussion Numbers Sighted Peregrines were sighted from 16 April to 30 May (Figure |) and again from 10 September to 4 October (Figure 2). From 1969 to 1978,541 peregrine sightings were recorded in spring (Table 1), and 34 in fall (Table 2). These include duplicate counts except the most obvious ones. The minimum counts pooled for 10 yr are 226 in spring (Table 1) and 27 in autumn (Table 2). From 1976 to 1978 both maximum and minimum spring counts were much higher than average, which can be attributed only partly to an increase in field time (Table 1). These years were characterized by early springs and low meltwater levels, with few ponds available to ducks and waders, which instead concentrated on the marshy lakeshore bordering the study area. In consequence the falcons hunted over a relatively narrow strip of land and were more readily observed than in wet springs when they foraged over much of the surrounding countryside. Timing of Migrations The earliest dates on which peregrines were sighted in the study area in different years varied from DEKKER: MIGRATING PEREGRINES IN ALBERTA 297 16 April to 9 May. These early arriving falcons were always adults. The mean date of first arrivals was 27 April. First dates for the brown-backed immatures varied from 27 April to 10 May. The mean first arrival date for immatures was 5 May.9 d behind the adults. The last spring dates on which adults were seen in the study area ranged from 11 May to 27 May witha mean last date of 19 May. Comparable dates for immatures varied from 2! May to 30 May witha mean of 25 May, 6 d after the adults. The great majority of peregrines (86%) passed through the area from 4 May to 23 May. Daily sightings showed a peak on 13 and 14 May. The peak of adult sightings occurred between 4 and 7 May, 8 d before the peak of immatures which fell between 12 and 15 May (Figure 1). The limited data from the autumn show sightings from 10 September to 4 October with a peak on 23 September (Figure 2). Subspecies and Plumage Variation White (1968) recognized three subspecies of Pere- grine Falcons in North America. Falco p. anatum formerly nested in nearly all suitable habitat south of the tundra except the northwest Pacific coast where F. p. pealei occurs. Falco p. tundrius inhabits the Arctic and Subarctic. During this study no representatives of F. p. pealei were identified. Most of the migrants seen could not be assigned to any subspecies. A few falcons that appeared typical, in some respects, of F. p. anatum were seen during the early part of each spring season, while typical examples of F. p. tundrius were among the last peregrines to pass through. The _ criterion used for field identification of the two subspecies was dorsal coioration which could best be appraised when the bird was in flight. The blue-gray color of adult F. p. anatum was lightest on the rump and darkened to nearly black on the head, tail, and wingtips. This variegated color pattern of F. p. anatum was also noted by Beebe (1974). When in flight, typical F. p. tundrius adults appeared evenly | blue-gray on the entire dorsal surface. They usually looked smaller and slenderer than typical F. p. anatum birds of the same sex. All adult falcons that could be observed through a 20-power scope at reasonably close range showed a very light, unmarked upper breast. Occasionally there was a pinkish wash on the remainder of the ventral surface. Dark barring and spotting on the belly and flanks varied from bold to fine, and from dense to widely spaced. Greater diversity of pigmentation and wider extremes in light and dark types were found among the first-year birds. Few spring immatures were streaked on the entire undersurface as they are in the preceding autumn. The majority showed a varying amount of unmarked yellowish-white on the breast, which made them resemble adults. To describe these falcons properly, a good view of the dorsal color Vol. 93 THE CANADIAN FIELD-NATURALIST 298 L 6 iS ¢ I LC ve bv | AOE care C C — oat V ’) 9I 8L6l I C I = ia Vv fk 81 LL6| Vv t a I = 8 8 9| 9L61 = = I = = I I 8I SL6l PS == a ood fo aaa cas tl vLol I | rn I a t 17 cl tL6l I = a 6 a t t Cl cL6l a ae I re Fat I I tl 1L6| Ti I a I ast C t tl OL6I — = — og I I I Ol 6961 poynuopiuy) So]BUd] So] RU So|BU9} So] Ru SdulYsis sdulyais sArq PIA d1N}JBUW | o41N}RUWWY WNpy Wnpy WIN UIT YA WINUITXP Jj Poafop U99q sARY sUOIBoI|dNp pajyoadsns []e ssuHYysis WNuUIW oY) UL “SUOnPOI|dNp apnjout SSUNYSIS WNWIXe QJ “8261-6961 “1940190 ¢] 01 doqtuajdag | potiod oy) Sulinp papsosas sautBaiod Jo sioquinu pur sdep UONRAIISGQ—Z V1HV | ve 99 9€ L9 ee 9CC Stl 6kC L9| Ips 6LC SWAT t Cl C 6 9 GS CC tt St 06 50 pe 8261 S II L S S tt oP 16 IG tol 66 Bt LL6| Ol vl 6 8 te A LI 6C OC 99 COL Ot 9161 L e 61 C It 8 Ol tc Ip 36 6C SL6l Pa L t v C 9| lL 8C II OV c 6 Ne vLol = v v Ol t IC 9 Sl OC IV tc tL6l t t = t S vl ¢ 8 tl 9 6 cL6| 5 t v Cc ¢ vl Sl lI L tt LC 1Z6] I C I v I 6 v v LL S| SC OL6I C t t t I Cl ¢ Ol ¢ OC CC 6961 payjiy sajPud} soyeuu sojeuWld} = Saye SBUNYSIS paljyuapluy, oaingeuwy ynpy ssuysis sinoy SARC IVI, -Udpluy) = daNyRUIW] d1Nj}RVUU] NPY WNpy WNUWHUTYAl WINWIX® JJ Apleq pojajap aie suoyvoydnp poyoodsns |jB ssunysis wnuturt oy} up ‘suoNPodnp apnyout SBUNYBIS WNWIXEY “8/61 6961 ACW [Ee OI [Udy S| pouad ay) SULINP payYysis souLIBosad Jo ssaquunu pur “pjaye sinoy Ajlep adesdrKw “SBP UONRALISGQ —| VIAV 1979 DEKKER: MIGRATING PEREGRINES IN ALBERTA 299 on 8 8 6 G 8 ul FIGURF 1. Maximum numbers of peregrines sighted in spring. Data pooled for 10 years. 1969-1978. 10 PEREGRINES 5 10 15 20 Zs) 30 1 5 10 15 SEPTEMBER OCTOBE FIGURE 2. Maximum numbers of peregrines sighted in autumn. Data pooled for 10 years. 1969-1978. 300 had to be obtained. Their brown backs ranged in tone from blackish-brown to sandy. The pale brown F. p. tundrius immatures were easily mistaken for Prairie Falcons (Falco mexicanus), which were only rarely seen in the study area during spring and fall. In fall, second-year peregrines had attained adult coloration although their wings and backs retained some brownish feathers. Of interest was the occurrence in the study area of “red-tailed” peregrines. About one-third of the adults showed a buffy or brownish cast on the upper tail. Also some immature falcons had “red” tails. The color varied from dark reddish brown to a strikingly pale buffy-orange. To my knowledge, there are no records in the literature of such “red-tailed” peregrines. Age Ratios During 10 springs, 102 immature peregrines were recorded among a total of 202 falcons classified as to age. This age ratio cannot be considered representa- tive of the population as a whole. Shor (1970, p. 56) reported that peregrine populations contain more adults than first-year individuals “even at the time of the fall migration.” This difference should be even more pronounced in spring, as Enderson (1969) calculated a 70% mortality among first-year birds from band returns. The very small sample of fall data obtained included 14 immatures and 6 adults. Fall counts from coastal migration routes indicate immature to adult ratios of 4 to | and 6 to! (Berry 1971; Ward and Berry 1972: Hunt et al. 1975). These ratios suggest dif- ferential migratory behavior among the age classes. Berry (1971) argued that adults, being more experi- enced in foraging, could fly directly to their wintering grounds. My observations suggest that during spring both age classes hunt the same prey and ina similar way. Hunting success ratios of adults and of immature falcons migrating through central Alberta during spring are not significantly different (Dekker, un- published data). Hence, the observed prominence of first-year young in central Alberta in spring cannot be explained as a consequence of different hunting ability as suggested by Berry (1971) for fall migrants. The majority of peregrines known to linger in the study area, however, were in immature plumage. The duplication rate of immatures was 2.3 (102 to 239), that of adults was 1.6 (100 to 167). It appeared that adult peregrines on their way north through central Alberta were less inclined to tarry on attractive feeding grounds than the immature birds. Sex Ratios The smallest male peregrines are about two-thirds the size of the largest females (Godfrey 1966). Hunt et al. (1975), however, have recognized the difficulty of THE CANADIAN FIELD-NATURALIST Vol. 93 accurately estimating sex ratios on the basis of sightings. I agree that the size of a peregrine cannot be judged reliably while the bird is resting unless it can be compared to another bird. In flight, however, a male peregrine usually appears slenderer than a female. Small to average-sized males and large to average- sized females are readily distinguished when seen in flight by an experienced observer. Additionally, foraging habits may provide clues and assist in classification. For example, males are more given than females to pursue and swoop repeatedly at fleeing sandpipers. About 85% of the peregrines included in the minimum count and classified to sex were observed under conditions that allowed me to make such classification with confidence. An element of doubt is present in about 15% of cases when individual variation in size made the sexes look alike. Rather than assigning these puzzling birds to the unidentified category, I classified them on the basis of impressions. As only one observer was involved these impressions may be expected to show a degree of consistency. Sex ratios obtained for adult (33%) and immature peregrines (35%) in fact show a striking similarity. Adult males are reported to be most elusive along the fall migration routes. Only eight out of 639 peregrines trapped on Assateague Island from 1939 to 1971 were identified as adult males (Ward and Berry 1972). No adult males were among 250 peregrines captured on the Texas beaches from 1952 to 1973 (Hunt et al. 1975), and only one among 20 classified in fall in this study. Percentages of immature male peregrines trapped along the major fall migration routes along the Texas and Maryland coasts respec- tively were reported as 23.4% (Hunt et al. 1975) and 30% (Ward and Berry 1972). Hunt et al. (1975) claim that there is no obvious reason why immature sex ratios should be un- balanced. The observed preponderance of females in the Texas survey may reflect differential trap- response and/or foraging habits. The males are believed to be able to exploit inland populations of small birds better than are the larger females, and are less often seen on the beaches. In my opinion, aggression between sexes may force the smaller males away from localities where female peregrines congre- gate. I frequently saw females chase males. Males were observed to carry their prey long distances before alighting to eat it. Two were observed to soar to great altitudes and feed while on the wing. By contrast, females routinely consumed their prey at the site of capture. On two occasions an adult female was seen to rob an immature male of his freshly caught prey. Attempts at stealing each other’s prey were seen also between immature females and between immature and adult females. 1979 Weather and Daily Routine The influence of weather on the movements of migrating raptors in spring and fall has been discussed in detail by Mueller and Berger(1961) and Haugh and Cade (1966). They concluded that the majority of raptorial birds migrate when tail winds are blowing. Ward and Berry (1972) and Hunt et al. (1975) correlated highest numbers of peregrines in fall with days of heavy cloud, light tail winds, and low temperatures. Those conditions usually accompanied the influx of some falcons into my study area in fallas well as in spring. The birds were easily located then as they rested on exposed perches (fence posts) or hunted at low elevations. Mueller and Berger (1961) have remarked that hawks have rarely been observed in the process of departing on migration. During this study many resting falcons were watched until they flew away of their own accord. In spring they consistently left the study area in a north-northwest direction. Such departures were seldom noticed on days of heavy cloud and low temperatures. Of particular interest was the observation that light to moderate head winds at ground level did not prevent some falcons from departing. They did not migrate close to the ground, as stated by Fischer (1967), but maintained high altitudes attained by soaring. Brown and Amadon (1968, p. 65) write that “falcons usually fly rather than sail or soar” while migrating. Both migrating and foraging falcons observed in this study not only soared during optimum conditions of sun and wind, but also on windy days when skies were cloudy, and even when light rain was falling. When nearly straight overhead of an observer, some of these soaring falcons were just visible in 10-power binoculars. On two occasions soaring peregrines were seen to enter the cumulus clouds that top thermal currents. Falcons that were seen to leave the study area ona steady northerly course on spring days when head winds were blowing, habitually winged upwind for a short distance before beginning to soar. During their circling ascent the falcons drifted downwind until they flexed their wings and glided through the wind rapidly. Even head winds gusting to50 km per hour at ground level did not deter them from high-altitude travel. All falcons observed to migrate during these contrary airflows were adults. First-year falcons that were seen to travel northward did so when tail winds were blowing or when head winds were only light. Most observed departures of adults and immatures took place during mid-morning after the air had warmed sufficiently to allow the formation of rising air currents. During the remainder of the day some migrating peregrines were located when they descended to attack prey species such as feeding shorebirds. After DEKKER: MIGRATING PEREGRINES IN ALBERTA 301 consuming a kill, a peregrine would resume flight by soaring to great heights and sailing away on slightly flexed wings. Such interruption and resumption of daytime migratory travel was observed from 11:00 to 18:00 hours. In spring, during the late afternoon and early evening, falcons sometimes flew in from a southerly direction as if they were just arriving. Aftera rest, these falcons began hunting in the hour preceding sundown. Several immature falcons were seen chasing and capturing shorebirds shortly before dusk. One immature female falcon that had been watched from late afternoon to nightfall began attacking ducks that flew inland from the lake one hour after sundown. Through binoculars, the birds were just visible, silhouetted against a cloudless sky. Crepuscular hunting was also observed by Beebe (1960), Clunie (1976), Fischer (1967), and Rudebeck (1950-1951). Late foraging activity may explain the relative scarcity of falcons on the Texas beach at sundown, which prompted Enderson (1965) to speculate on the likeli- hood of nocturnal migrations. During this study, several falcons were observed roosting at nightfall. They were sitting on posts surrounded by water or on shoreline rocks. Of three adult peregrines that were seen roosting at night only one was still on its post when it was checked at sunrise. It remained there until 10:00 hours when it flew upwind, began to soar and sailed northward. I have seldom found evidence of hunting at dawn. An hour after sunrise was the usual start of foraging flights. When a large food item such as a duck was captured, the falcon would eat for about 45 min and rest until mid-morning. A second meal of the prey remains was frequently taken before the falcon flew away, soared to great height, and glided northwards until lost to view. Some spring migrants that soared and sailed northwards during mid-morning were seen to stoop at prey species that flew below them: it appeared that these peregrines did their foraging along the way. NOTE: Formerly the study area was part of the summer range of peregrines that nested along rivers to the south and west about 100 kmaway. By 1969, at the start of this survey, that central Albertan population of F. p. anatum was all but extirpated (Dekker 1967). In 1977 and 1978 one or two peregrines frequented the area during the summer months. In 1978 a first-year male, recognizable by his banded feet, damaged tail, and lack of shyness, was often seen from April to late August. These birds were believed to have originated from the captive breeding program sponsored by the Canadian Wildlife Service and the Provincial Fish and Wildlife Division. All known and_ suspected sightings of these birds have been eliminated from the tabulations in this paper. 302 Acknowledgments During the springs of 1974 to 1978, this study was financially supported by the Alberta Department of Recreation, Parks and Wildlife under the direction of Wiiliam Wishart, Head of Wildlife Research. I thank D. A. Boag for his valuable suggestions for revising the manuscript. I am also grateful for the constructive criticism offered by A. J. Erskine, D. V. Weseloh, and adie Cade: Literature Cited Beebe, F. L. 1960. The marine peregrines of the Northwest Pacific Coast. Condor 62: 145-189. Beebe, F. L. 1974. Field studies of the Falconiformes of British Columbia. British Columbia Provincial Museum. Occasional Paper Series Number 77. 163 pp. Berry, R.B. 1971. Peregrine falcon population survey. Assateague Island. Maryland. fall 1969. Raptor Research News 5: 31-43. Brown, L. and D. Amadon. 1968. Eagles. hawks and falcons of the world. Hamlyn House. 945 pp. Cade, T. J. 1960. Ecology of the peregrine and gyrfalcon populations in Alaska. University of California Publica- tions in Zoology 63: 151-290. Clunie, F. 1976. A Fiji peregrine in an urban environ- ment. Notornis 23: 8-28. Dekker, D. 1967. Disappearance of the peregrine falcon as a breeding bird of a river valley in central Alberta. Blue Jay 25: 175-176. Enderson, J. H. 1965. A breeding and migration survey of the peregrine falcon. Wilson Bulletin 77: 327-339. THE CANADIAN FIELD-NATURALIST Vol. 93 Enderson, J.H. 1969. Peregrine and prairie falcon life tables based on band recovery data. /nm Peregrine populations — their biology and decline. Edited by J.J. Hickey. pp. 505-509. Fischer, W. 1967. Der Wanderfalk. Die Neue Brehm Bucherei. Ziemsen Verlag. 152 pp. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada. Bulletin 203. 428 pp. Haugh, J.R. and T.J. Cade. 1966. The spring hawk migration around the southeastern shore of Lake Ontario. Wilson Bulletin 78: 88-110. Hunt, W. G., R. R. Rogers, and D. J. Slowe. 1975. Migra- tory and foraging behavior of peregrine falcons on the Texas coast. Canadian Field-Naturalist 89: 111-123. Mueller, H. C. and D. D. Berger. 1961. Weather and fall migration of hawks at Cedar Grove. Wisconsin. Wilson Bulletin 73: 171-192. Rudebeck, G. 1950-1951. The choice of prey and modes of hunting of predatory birds with special reference to their selective effect. Oikos 2: 65-88: 3: 201-231. Shor, W. 1970. Peregrine falcon population dynamics deduced from band recovery data. Raptor Research News 4: 4959. Ward, F. P. and R. B. Berry. 1972. Autumn migrations of peregrine falcons on Assateague Island. 1970-1971. Journal of Wildlife Management 36: 484-492. White, C. M. 1968. Diagnosis and relationships of the North American tundra inhabiting peregrine falcons. Auk 85: 179-191. Received 30 October 1978 Accepted 19 March 1979 Notes Extralimital Record of a Narwhal (Monodon monoceros) in Hall’s Bay, Newfoundland BORA MERDSoy,! JON LIEN, and ANNE STOREY Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland AIC 5S7 \Present Address: 7 Appledore Place, St. John’s, Newfoundland A1lB 2W8 Merdsoy, Bora, Jon Lien, and Anne Storey. 1979. Extralimital record of a Narwhal (Monodon monoceros) in Hall’s Bay, Newfoundland. Canadian Field-Naturalist 93(3): Key Words: Narwhal, Monodon monoceros, Newfoundland, On 1 March 1978 a Narwhal (Monodon monoceros) was seen from a small boat in Hall’s Bay, Newfoundland (49°30’N, 56°00’W). Pack ice, several miles wide, had jammed across the head of the fjord and extended into the bay so that the animal, along with four Humpback Whales (Megaptera novae- angliae), was confined to an area 8 X4km. Humpback Whales were observed in the area earlier in the winter, but the Narwhal was not seen until February. Both local and arctic ice conditions were unstable within the bay, but the entrance to the bay remained blocked so that the Humpback Whales and the Narwhal were unable to leave Hall’s Bay until 6 April. At this time the pack ice, loosened several days earlier by an ice-breaker, drifted to open water enabling the whales to escape. No sightings of the Narwhal or Humpback Whales were made after they left Hall’s Bay although repeated air surveillance was attempted (Cyril Pelly, Springdale, Newfoundland, personal communication). Hall’s Bay contains | m of fresh water at the surface which results in rapid formation of local ice over the entire bay. During the period the whales were confined to the bay the area of open water decreased irregularly. For a period of 3 wk the only open water was 1.6 km NW of Mansfield Head, which varied in size from 120 X 45 m. This confinement allowed for close observation and photography of the Narwhal (Figure 1). The Narwhal was a dark gray color, 2-3 m long with a tusk of approximately 15-20 cm, indicating that it was a male. Generally the Narwhal surfaced in the open water during periods the Humpback Whales were below the surface. In addition to such time- sharing behavior, spatial separation was maintained, as the Humpback Whales typically would follow the edges of the open water while the Narwhal restricted its surface activities to the center. Although ice forced the Narwhal into proximity with human observers, it remained timid throughout its confinement typically maximizing its distance from observers. Food species is) i) 303-304. sighting, range, geographical distribution, pack ice. including Atlantic Herring (Clupea harengus), Rainbow Smelt (Osmerus mordax), and Capelin (Mallotus villosus) were plentiful in the area and hydrophone recordings indicated hunting activity by the animal (Ford and Fisher 1978). Nosurface sounds were heard from the Narwhal. The range of the Narwhal in the Canadian Eastern Arctic extends south to about 66°N (Banfield 1974). Our sighting was approximately 1800 km south of this at 49°30’N. Our report is the second sighting of a Narwhal in Newfoundland in the last decade. The first extralimital record of a mature male Narwhal, found washed on a beach in White Bay in 1969, was reported by Mercer (1973). Several sightings of white whales reported in Newfoundland waters (Sergeant and Fisher 1957; L. M. Tuck, St. John’s Newfoundland, personal communication), could be either Belugas (Delphinapterus leucas) or Narwhals. The south- eastward movement of arctic ice in the late winter brings Harp (Phoca groenlandicas) and Hooded (Cystophora cristata) Seals in large numbers to Newfoundland’s coast. Occasionally Bearded Seals (Erignathus barbatus), Ringed Seals ( Phoca hispida), | Walrus (Cdobenus rosmarus), and Polar Bears ( Ursus maritimus) have drifted south with the ice (L. M. Tuck, personal communication). Narwhals are reported to winter in the Baffin Bay area (Mansfield et al. 1975) and it is probable that, on occasion, they follow ice southward. This could account for the present record, but may not account for the June 1969 record (Mercer 1973) unless that animal came with the ice and survived in Newfoundland through the spring. Because it is possible to mistake the finless dorsum of an immature Narwhal for that of a seal, it may be that more Narwhals frequent the sparsely inhabited shores of Newfoundland than are being recognized. We gratefully acknowledge the help of Francis Hull and Cyril Pelly of Springdale, Newfoundland, during the period of our work in Hall’s Bay. 304 THE CANADIAN FIELD-NATURALIST FIGURE |. The Hall’s Bay Narwhal during the last week of its confinement. The tusk length was approximately 45 cm, which aged the animal at over 2 yr. Photographed by Duncan Findlayson. Literature Cited Banfield, A.W. F. 1974. The mammals of Canada. University of Toronto Press, Toronto. 438 pp. Ford, J. K. B. and H. D. Fisher. 1978. Underwater acou- stic signals of the Narwhal (Monodon monoceros). Canadian Journal of Zoology 56: 552-560. Mansfield, A. W., T. G. Smith, and B. Beck. 1975. The Narwhal, Monodon monoceros, in eastern Canadian waters. Journal of the Fisheries Research Board of Canada 32: 1041-1046. Mercer, M.C. 1973. Observations on distribution and intraspecific variation in pigmentation patterns of odontocete Cetacea in the western North Atlantic. Journal of the Fisheries Research Board of Canada 30: 1111-1130. Sergeant, D.E. and H.D. Fisher. 1957. The smaller Cetacea of eastern Canadian waters. Journal of the Fisheries Research Board of Canada 14: 83-115. Received 19 September 1978 Accepted 30 January 1979 First Canadian Record of the Black Buffalo (Osteichthyes: Catostomidae) E. J. CROSSMAN! and S. J. NEPSzyY2 'Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 2Ontario Ministry of Natural Resources, R.R. #2, Wheatley, Ontario NOP 2P0 Crossman, E. J. and S.J. Nepszy. 1979. First Canadian record of the Black Buffalo (Osteichthyes: Catostomidae). Canadian Field-Naturalist 93(3): 304-305. Ictiobus niger was captured in the Canadian waters of Lake Erie on 20 June 1978. Key Words: Black Buffalo, Jctiobus niger, first Canadian record, Lake Erie, geographical distribution. A specimen of J/ctiobus niger, the Black Buffalo, one of the larger deep-bodied suckers, was captured in the Canadian waters of Lake Erie on 20 June 1978. It had been taken in a commercial trap net set by F.W. 1979 NOTES 305 TABLE 1—Comparison of characters in the three species of Ictiobus Character Ictiobus cyprinellus Ictiobus bubalus Ictiobus niger Body Robust, slightly com- pressed, back not highly elevated Usually 24-28 Large, oblique Dorsal rays Mouth Lips Thinner than others, only faintly striate, upper about level with lower margin of orbit Distance mandibular symph. to tip of maxilla Upper jaw About as long as snout Gill rakers post. face first arch Lower pharyngeal arch Nearly 100 Thin, more than twice as high as wide, teeth weak More slender, but thicker than /. b., less elevated and less sharpened than J. b. Deeper and narrower than Ln., highly elevated, mid dorsal ridge sharper than others Usually 27-30 Smaller, less oblique than Less oblique than /. c., I. c., more inferior than larger and less inferior than Ln. ae: Thicker, more or less Thicker than /.c., more coarsely straite, upper or less coarsely striate, far below lower margin upper far below lower of orbit margin of orbit Only slightly greater than As much as twice diam. of diam. of orbit in adults orbit in adults Distinctly shorter than Shorter than snout snout Fewer than 60 Usually 30 Fewer than 60 Heavy. about as wide as high, teeth strong Heavy. about as wide as high, teeth strong Krause and Sons Fisheries. Leamington, Ontario, in Pigeon Bay. west of Point Pelee (41°56’50’N. 82°31'40”W). The specimen was turned over to Nepszy at the Ontario Ministry of Natural Resources Research Station Wheatley, Ontario, and forwarded to Royal Ontario Museum, frozen, as an unidentified sucker. possibly [ cyprinellus. After fixation in formalin, and preservation in ethy] alcohol the buffalo was 630 mm in total length. and weighed 3.56 kg. Preliminary examination. including cranial fon- tinelles. clarified that it was [ctiobus but that it was not J. cyprinellus. the commonest species and the only species previously reported as occurring in Canada. Morphologically it seemed to resemble both 1 bubalus and I. niger and was somewhat suggestive ofa hybrid. Neither of those species. however. is common in Lake Erie and published characteristics used to separate the two species are difficult to interpret. Ictiobus niger is basically a species of the Missis- sippi River system and occurs from the Ohio River and its tributaries south to Mexico. It has. however. been said to be native to southern Lake Michigan (Hubbs and Lagler 1964) and therefore in the Great Lakes system. Moore (1968) listed 1. niger but not 1. bubalus from Lake Erie. and Hubbs and Lagler (1964) specified earlier that there was one report only of /. niger, and gave no Great Lakes record for J. bubalus. Trautman (1957) gave no records for the Ohio waters of Lake Erie for either species. Trautman did indicate. however (under /. bubalus), that specimens had been caught in Sandusky Bay. Lake Ene. which had been identified as hybrids between [. cyprinellus and I. bubalus or I. niger. So few specimens of ZI/ctiobus other than 1 cyprinellus are seen from the Great Lakes that the characteristic features of larger specimens of /. bubalus and I. niger are poorly known. A comparison of the features of the three species is given in Table 1. The supposed hybrids may have represented the presence of I. bubalus or I. niger in Lake Erie in the past. After detailed examination of the characteristics, including the cranial bones, of this specimen, G. R. Smith and R. M. Bailey of the Museum of Zoology. University of Michigan confirmed that the specimen represents Ictiobus niger. Therefore the specimen is not only a recent corroboration of the occurrence of the species in Lake Erie but also represents a species new to the fauna of Canada. It has been added to the collection of the Royal Ontario Museum (ROM 34562). We once more thank Rudy Krause for his vigilance and for turning over specimens suggestive of some- thing new. We thank Erling Holm. and G. R. Smith and R. M. Bailey for their help in the identification of the specimen. Literature Cited Hubbs, C. L. and K. F. Lagler. 1964. Fishes of the Great Lakes region. University of Michigan Press. 213 pp. Moore, G. A. 1968. Fishes. Jn Vertebrates of the United States. McGraw-Hill Inc. pp. 21-165. Trautman, M.B. 1957. The fishes of Ohio. Ohio State University Press. 683 pp. Received 13 December 1978 Accepted 30 January 1979 306 THE CANADIAN FIELD-NATURALIST Vol. 93 Thaspium trifoliatum (Meadow-parsnip) in Canada P. W. BALL Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6 Ball, P. W. 1979. Thaspium trifoliatum (Meadow-parsnip) in Canada. Canadian Field-Naturalist 93(3): 306-307. Thaspium trifoliatum (Meadow-parsnip) is reported from a locality near Amherstburg, Essex County, southern Ontario. Although reported from Canada on a number of previous occasions, all the earlier records appear to be based on confusion with the very similar Zizia aptera. The differences between T. trifoliatum and Z. aptera are discussed and it is noted that, in addition to the fruit characters usually used to distinguish between them, the width of the petiole also appears to be diagnostic. Key Words: Thaspium trifoliatum, Umbelliferae (Apiaceae), Essex County, Ontario. Thaspium trifoliatum (Meadow-parsnip) has been recorded from Canada on a number of occasions. Indeed, Macoun (1883) states that the species is abundant in the prairie provinces. Subsequent investigations have shown that all of these records are erroneous. For example, Scoggan (1957) makes it quite clear that all Manitoba records of T. trifoliatum should be referred to Zizia aptera. Soper (1949) lists T. trifoliatum from southern Ontario, this record apparently being based on a specimen from Caradoc, Middlesex County, collected by J. Dearness in 1891, and now in the herbarium of the Biosystematics Research Institute, Ottawa (DAO). This specimen was subsequently redetermined by W. J. Cody as Z. aptera, and, although the specimen is immature, it does appear to be this species. Boivin (1967) did not include T. trifoliatum in the Canadian flora and, ina personal communication, has stated that he has not traced any reliable records since then. The discovery of a location for 7. trifoliatum in southern Ontario seems therefore to be the first authentic record for this species in Canada. The locality is a woodlot about 3 km E of Amherstberg in Essex County. The dominant trees were Quercus alba (White Oak), Ostrya virginiana (Hophornbeam), Carya ovata (Shagbark Hickory), and Fraxinus pennsylvanica (Green Ash). The shrub layer was fairly dense with Zanthoxylum americanum (Prickly Ash), Rhus aromatica (Fragrant Sumach), Viburnum rafinesquianum, and Crategus spp. (hawthorn) prominent in the vicinity of the Thaspium. The stand also contained Geum vernum, another species that is rare in Ontario and largely restricted to Essex County. The population of 7. trifoliatum consisted of over 50 individuals so does not appear to be in immediate danger of extinction; however, any major disturb- ance of the woodlot could seriously jeopardize the survival of this species in Canada. Thaspium trifoliatum and Z. aptera are remar- kably similar in appearance, so that confusion between these two species is not surprising. When in fruit they can be readily distinguished by two characters. Zizia aptera has fruits with narrow ridges (Figure 1A) and the central fruit of each subumbel is sessile; T. trifoliatum has fruits which are distinctly winged (Figure 1B) and which are all distinctly pedicelled. Unfortunately these differences cannot be easily observed when the plants are in flower. The wings on the fruits of T. trifoliatum are not apparent until a fairly late stage in fruit development. The presence of a central sessile flower in each subumbel can be readily observed once all the flowers are open, but when the inner flowers of each subumbel are in bud it is difficult to determine whether the central flower is sessile or pedicelled. An additional character which is useful, although not completely reliable, is the width of the petiole of the cauline leaves (Figure 1C, D). In Z. aptera the petioles of the lower leaves on the flowering stem are 8-16 mm wide and almost forma sheath around the stem, whereas in 7. trifoliatum the petioles of the lower leaves are 4-8 mm wide and scarcely enclose the stem except at the node. The petioles of the upper leaves of both species are somewhat narrower, 5-10 mm wide in Z. aptera and 3-5 mm wide in T. @rifoliatum, but if leaves are compared from similar regions of the stem then the petioles of Z. aptera are always wider than those of 7. trifoliatum. Thaspium trifoliatum is variable in flower color. Var. trifoliatum, which is the predominant variety in the Appalachian region, and which also occurs rarely elsewhere, has purple flowers. Var. flavum, which has yellow flowers, is more western in distribution occurring as far west as Minnesota, Arkansas, and Kansas, but it also occurs rarely in the east. The Ontario material is referable to var. flavum. Specimen Examined Essex County. 3 km (2 mi) E of Amherstberg. 10 June 1975. P. W. Ball 14475 (TRTE, DAO) (Flower, young fruit). Anderdon Township 3 km (2 mi) NE of Amherstberg on east side of Concession II. 26 May 1977. P.M. Catling (TRT) (Flower). June 1977. P. M. Catling (TRT, TRTE, CAN). (Fruit). ISD NOTES 307 G FiGure |. Fruit and upper leaves of Zizia aptera (A, C) and Thaspium trifoliatum (B, D). Acknowledgments My thanks to J. McNeill for checking the identity of some of the specimens and to P. Catling for collecting some specimens in fruit. This work was supported by National Research Council of Canada Grant A6494. Literature Cited Boivin, B. 1967. Enumération des plants du Canada. III. Naturaliste Canadien 93: 583-646. Macoun, J. 1883. Catalogue of Canadian plants. Part 1. Montreal. 192 pp. Scoggan, H. J. 1957. Flora of Manitoba. National Muse- um of Canada Bulletin 140. 619 pp. Soper, J. H. 1949. The vascular plants of southern Ontario. Toronto. 95 pp. Received 25 May 1978. Accepted 19 January 1979. 308 THE CANADIAN FIELD-NATURALIST Vol. 93 Largest Gray Wolf Skulls Found in Alberta JOHN R. GUNSON! and RONALD M. NOWAK2 \Alberta Fish and Wildlife Division, 6909-116 St., Edmonton, Alberta T6H 4P2 2Office of Endangered Species, U.S. Fish and Wildlife Service, Washington, D.C. 20240 Gunson, J.R. and R.M. Nowak. 1979. Largest Gray Wolf skulls found in Alberta. Canadian Field-Naturalist 93(3): 308-309. Measurements of greatest length and zygomatic width of five large skulls of Canis lupus from Alberta are reported. Three, all from the range of C. /. occidentalis, are larger than the previous largest specimen, also occidentalis. Key Words: Gray Wolf, skull measurements, Canis lupus, size. According to Goldman (1944) the largest sub- species of the Gray Wolf (Canis /upus) are found in Alaska and northwestern Canada. With respect to measurements in millimetres of greatest length (g.1.) and zygomatic width (z.w.) of skull, the four largest individuals that he listed (pp. 490-497), all males, were the following: United States National Museum 9001, C. | occidentalis, Fort Simpson, Mackenzie, g.l. 292.8, z.w. 156.5; Royal Ontario Museum 33-9-20-5, C. |. pambasileus, White River, Yukon, g.1. 293.7, z.w. 151.5: Royal Ontario Museum 31-2-16-2, C. 1. pambasileus, White River, Yukon, g.]. 292.8, z.w. 149.0; University of California Museum of Vertebrate Zoology 31043, C. /. columbianus, Iskut Summit, British Columbia, g.1. 288.9, z.w. 155.0. On the basis of both measurements, the specimen from Fort Simpson, which was collected in 1869, has been until now the largest reported skull of a North American Gray Wolf. Several specimens of male C. /upus recently collected in Alberta have skull dimensions that approximate or surpass those of the above-mentioned material (Table 1). The largest of all, number Z78.104.1 in the collection of the Provincial Museum of Alberta (formerly 66-18 of the Alberta Fish and Wildlife Division), was taken at Roche Lake in the Swan Hills area, about 155 km northwest of Edmon- ton. Two others, Z78.104.2 and Z78.104.3, taken during 1972-1974 in northwestern Alberta were also larger than the Fort Simpson wolf. All three were taken in areas within the original range of C. /. occidentalis, as was Z78.104.4. Specimen Z78.104.5 was collected in southwestern Alberta from an area within Goldman’s original range of C. /. irremotus. Determination of the taxonomic position of that specimen and others recently collected from even more southerly locations in Alberta awaits the collection of additional data. Several extensive regional collections of North American Gray Wolves have been assembled in recent decades, but with the exception of the Alberta Fish and Wildlife Division collection (12 measurements on each of 468 skulls), none have been thoroughly analyzed. One of us (Nowak) made cursory examina- tions of a collection from western Canada in the possession of the University of British Columbia and a collection from Alaska in the possession of the University of Alaska. No skulls were found that surpassed in size the larger of the above-described specimens, but one of the Alaska skulls of C. /. pambasileus may be the largest ever recorded for that state. The specimen (University of Alaska 40682) was TABLE 1—Specifics of five large male Canis /upus from Alberta, Canada. PMA = Provincial Museum of Alberta: AFW = Alberta Fish and Wildlife Division Specmontie Greatest Zygomatic : Date of length width PMA AFW Location collection (mm) (mm) Z78.104.1 66-18 54°46’N,114°55’W 30 Jan. 1966 304.5 154.8 Z78.104.2 V441036 55° 4’N,119°50’W 13 Feb. 1973 297.6 158.0 Z78.104.3 V441097 54°43’N,113°17’W Feb. 1974 288.8 162.5 Z78.104.4 V441057 54°48’N,119°20’W 27 Feb. 1973 292.6 155.8 Z78.104.5 V441016 $1°27’N,114°50’W 28 Feb. 1973 285.3 160.4 1979 collected in 1966 in the Wood River drainage of central Alaska, and its measurements (taken to the nearest millimetre) are g.l. 292, z.w. 153. Mean greatest length and zygomatic width for 83 adult male C. /upus taken in Alberta during 1965-1966 to 1977-1978 were 275.6 (range 251.3 to 304.5) and 150.7 (range 128.6 to 162.5). The following data on series of North American wolf skulls (Nowak 1973) are offered for purposes of comparison: 233 Recent male C. /upus from throughout northern and western North America, mean g.l. 259.6 (range 235-293), mean z.w. 141.1 (range 126-164); 20 male C. |. pambasileus, mean g.1. 271.4 (range 248-288), mean z.w. 145.4 (range 130-154): 62 C. dirus(unsexed) from late Pleistocene deposits at Rancho La Brea, Cali- fornia, mean g.l. 294.8 (range 258-316), mean z.w. 163.3 (range 148-177). In comparisons of skull dimensions of wolves from several areas in western North America, Skeel and Carbyn (1977) recorded largest mean g.|. and z.w. in specimens from Prince Albert National Park, Saskatchewan, within the range of C. |. occidentalis (Goldman 1944) or C. /. Aquatic Feeding by a Woodchuck! D. FRASER NOTES 309 griseoalbus (Hall and Kelson 1959). Although there is no comprehensive analysis of specimens throughout North America, the wolves of the boreal-subalpine forest regions of Alberta and adjacent areas appear to be the largest of the North American Gray Wolves. Literature Cited Goldman, E. A. 1944. Classification of wolves. Part II. The wolves of North America. S.P. Young and E.A. Goldman. American Wildlife Institute, Washington, D.C. Hall, E. R.and K. R. Kelson. 1959. The mammals of North America. Volume II. Ronald Press, New York. Nowak, R.M. 1973. North American Quaternary Canis. Ph.D. dissertation, University of Kansas, Lawrence, Kansas. Skeel, M. A. and L. N. Carbyn. 1977. The morphological relationship of Gray Wolves (Canis /upus) in national parks of central Canada. Canadian Journal of Zoology 55(4): 737-747. Received 2 November 1978 Accepted 19 January 1979 Ontario. Ministry of Natural Resources, Wildlife Research Section, Box 50, Maple, Ontario LOJ 1E0 Fraser, D. 1979. Aquatic feeding by a Woodchuck. Canadian Field-Naturalist 93(3): 309-310. A Woodchuck (Marmota monax) was seen eating aquatic plants which it obtained by climbing on fallen trees ona lakeshore. An attraction to sodium-rich plants probably accounts for the behavior. Key Words: Woodchuck, Marmota monax, aquatic plants, sodium, feeding behavior. Although they feed on a variety of vegetation, Woodchucks (Marmota monax) are not generally known to eat aquatic plants. The following observa- tions document such behavior. On 30 June 1978, while canoeing on a small unnamed lake in Sibley Provincial Park, Ontario (48°27’N, 88°45’W), I watched through binoculars as an adult Woodchuck climbed among the tangled branches of fallen Eastern White Cedar trees (Thuja occidentalis) overhanging the water. The animal climbed to a location | to 2 m from shore where it clung, holding its head 1 to 3 cm above the surface of the water and its hindquarters slightly higher. Prominent light areas visible around the nipples indicated that it was a female. From this location the Woodchuck dipped its left front paw into the water and pulled vegetation into its mouth. It fed for about 1 min, then returned to shore, apparently because my canoe had drifted within 10 m of the location. The Woodchuck returned to the water’s edge about 8 min later, and walked about | m from shore alonga half sunken log. There it stopped and chewed on three plants which it dipped from the water with a front paw. It then climbed to two sites among the fallen trees, eating submersed and floating-leafed vegetation at each location. The animal left to shore 16 min after it had reappeared. The feeding sites were dominated by submersed Hippuris vulgaris and Potamogeton zosteriformis, floating-leafed Nuphar variegatum, and emergent Sagittaria sp., growing in 30 cm of water over a soft organic bottom. At the main feeding location were 10 H. vulgaris plants missing their upper portions, one uprooted specimen and one damaged fragment of ‘Ontario Ministry of Natural Resources, Wildlife Research Section Contribution No. 78-22. 310 P. zosteriformis, four leaf fragments and two leafless petioles of N. variegatum, and three damaged Sagittaria. The second feeding site by the log had damaged fragments of P. zosteriformis, N. varie- gatum, and Sagittaria. It appeared that the animal had completely consumed the portions of H. vulgaris which it had picked, but left partially chewed fragments of the other species. I revisited the site five times in the next 18 days. On the last day, 18 July, freshly damaged H. vulgaris, N. variegatum, and Sagittaria were observed. Fresh damage was not evident on the earlier visits. Like many herbivores, Woodchucks show evidence of a sodium-specific hunger in the spring and early summer (Weeks and Kirkpatrick 1978), and aquatic plants are generally far richer in sodium than are terrestrial ones (Jordan et al. 1973). This may explain why a Woodchuck would make such an effort to obtain a small quantity of vegetation. Non-aquatic herbage was readily available on the forest floor, lakeshore, and roadsides nearby. Nonetheless, this behavior is clearly a rarity, since few Woodchucks would be expected to find con- venient access to aquatic plants in shallow water, and the animals are rarely seen swimming (Grizzell 1955). THE CANADIAN FIELD-NATURALIST Vol. 93 Hamilton (1934) does, however, list one sighting of a Woodchuck eating Vallisneria americana in July inan Adirondack Mountain pond. This animal also reached the vegetation by walking on a fallen log. I am grateful to D. R. Voigt, M. C. Smith, and H. P. Weeks for offering suggestions on an earlier draft of the manuscript. Literature Cited Grizzell, R. A. 1955. A study of the southern Woodchuck, Marmota monax monax. American Midland Naturalist 53(2): 257-293. Hamilton, W. J. 1934. The life history of the Rufescent Woodchuck, Marmota monax rufescens, Howell. Annals of the Carnegie Museum 23: 85-178. Jordan, P. A., D. B. Botkin, A. S. Dominski, H. S. Lowen- dorf, and G. E. Belovsky. 1973. Sodium as a critical nutrient for the moose of Isle Royale. Proceedings of the North American Moose Conference and Workshop 9: 13-42. Weeks, H.P. and C.M. Kirkpatrick. 1978. Salt prefer- ences and sodium drive phenology in Fox Squirrels and Woodchucks. Journal of Mammalogy 59: 531-542. Received 20 December 1978 Accepted 12 February 1979 Dispersion of Freshwater Leeches (Hirudinoidea) to Anticosti Island, Quebec RONALD W. DAVIES Department of Biology, University of Calgary, Calgary, Alberta T2N IN4 Davies, Ronald W. 1979. Dispersion of freshwater leeches (Hirudinoidea) to Anticosti Island, Quebec. Canadian Field- Naturalist 93(3): 310-313. Four species of freshwater leeches, He/obdella stagnalis, H. triserialis, Glossiphonia complanata, and Mooreobdella fervida are reported from Anticosti Island. Evidence is present to show that these species dispersed to Anticosti Island passively by sea currents from Quebec North Shore of the Gulf of St. Lawrence. Key Words: dispersion, leeches, Hirudinoidea, Anticosti Island, water currents. Anticosti Island in the Gulf of St. Lawrence is about 216 km long with a maximum width of 48 km. It is separated by the 29-km or more wide Jacques Cartier Passage (Détroit de Jacques Cartier) from the North Shore of the province of Quebec to the north-east, and by the 64-km or more wide Gaspé Passage (Détroit d’Honguedo) from the Gaspé Peninsula to the south- west (Figure |). Having been entirely covered by ice during the Wisconsin glacial period (Stockwell 1957) and without known glacial refugia or landbridges to the mainland during or since the ice recession (Bleakney 1958; Cameron 1958), the fauna of Anti- costi Island provides an ideal location to interpret postglacial migrations and dispersal. Although Verrill (1863), a prolific author of papers on leeches (Hirudinoidea), made no reference to freshwater leeches on Anticosti Island it cannot be assumed that none were present at that time. Subse- quently Schmitt (1904) recorded the presence of Dina (= Mooreobdella) fervida and Johansen (1924) re- corded Helobdella stagnalis. Ina general review of the geographical distribution of freshwater leeches of Canada, Davies (1973) reconfirmed the presence of M. fervida and H. stagnalis on Anticosti Island and 1979 NOTES 311 a LAWRENCE QUEBEC NORTH SHORE ANTICOSTI ISLAND st — N ee NEWFOUNDLAND 50 km FIGURE |. Map of south-eastern Quebec and Newfoundland showing the major currents (based on Cameron 1958) and the collection sites of the freshwater leeches (@). added records for Helobdella triserialis and Glossi- phonia complanata. Davies (1973) recorded seven species from the Gaspé Peninsula to which, from subsequent collections, can now be added Nephelo- psis obscura and Percymoorensis marmorata (Table 1). Although no new species can be added to the 18 recorded in Quebec by Davies (1973) (Table 1) the presence on the Quebec North Shore of H. stagnalis, H. triserialis, G. complanata, M. fervida, N. obscura, Erpobdella punctata, and Dina parva can be con- firmed. Three of the four leech species on Anticosti Island (H. stagnalis, H. triseralis, and G. complanata) belong to the family Glossiphoniidae while M. fervida belongs to the family Erpobdellidae. The Glossi- phoniidae produce thin-walled cocoons which are immediately covered by the parents’ body and to which, while still enclosed within the egg membrane, the young glossiphoniids develop an embryonic attachment. As young glossiphonids prematurely removed from the parent fail to survive, it can be assumed for these three species at least, that dis- persion to Anticosti Island must have been in the adult stage. The Erpobdellidae deposit their cocoons on a firm substratum and the parent has no further participation. It is thus possible for M. fervida to be dispersed in the cocoon stage while attached to stones or plant material. TABLE 1—The distribution of freshwater Hirudinoidea species in the province of Quebec, Gaspé Peninsula, Anticosti Island, and the Quebec North Shore Anti- North Gaspé costi Shore Que- Species bec Piscicola geometra (Linn.) Piscicola milneri (Verrill) Piscicola punctata (Verrill) Bactracobdella picta (Verrill) Glossiphonia complanata (Linn.) Glossiphonia heteroclita (Linn.) Helobdella stagnalis (Linn.) Helobdella triserialis (Blanchard) Placobdella ornata (Verrill) Placobdella phalera (Graf) Dina parva (Moore) Erpobdella punctata (Linn.) Mooreobdella fervida (Verrill) Nephelopsis obscura (Verrill) Bdellerogatis plumbeus (Moore) Macrobdella decora (Say) Mollibdella grandis (Verrill) Percymoorensis marmorata (Say) AA aK MK KK KR KK KK ~*~ AAX~K KK of od 3) The occurrence of four species of freshwater leeches on Anticosti Island requires explanation, and there are three possible hypotheses: (a) passive dispersal by animals, wind, or currents; (b) introduction by man; (c) active dispersion. Because of the absence of H. stagnalis from the Gaspé Peninsula, the leech fauna of Anticosti Island shows greater similarity to that of the Quebec North Shore. The absence of a species record from an area is sometimes a reflection of the degree of collection intensity, but as there have been more collections from the Gaspé than from either Anticosti Island or the Quebec North Shore, the absence of H. stagnalis from the Gaspé appears to be real. All the leech records from Anticosti Island are from the north-east Quebec North Shore-facing coast (Figure |), supporting the supposition of the Quebec North Shore as the source of dispersion. Indeed, of the 17 sites sampled on Anticosti Island (Davies 1973), three of the four sites on the north-east coast contained leeches but none of 13 on the southern half of the island did. All four leech species on Anticosti Island are eurytopic (Davies 1973) benthic feeders, lack jaws, and are very sensitive to disturbances in the water. Thus although passive dispersal by birds has been recorded for benthic leeches (Daborn 1976) it hardly seems credible that a bird traversing Jacques Cartier Passage would not occasionally land on the southern half of this narrow island. Similarly if leeches or cocoons were transported by strong winds, hurri- canes, or tornados to Anticosti Island they would be expected on both halves of the island. As passive dispersal by wind can probably be discounted, dispersal by currents or tides remains a possibility. In a study of the osmoregulatory ability of freshwater leeches, Reynoldson and Davies (1976) showed that H. stagnalis was able to maintain itself hyperosmotically in media between 47 and 112.7 mosmol/L and was a conformer at high- er media concentrations, and thus could survive periods of immersion in sea water. Unfortunately the osmoregulatory abilities of H. triserialis, G. com- planata, and M. fervida have not been studied but if they prove to beat least as good as that of H. stagnalis, it is clear that all four species could survive the sea passage to Anticosti Island. Nephelopsis obscura, a widely distributed species in Canada (Davies 1973), present on the Quebec North Shore but absent from Anticosti Island can only maintain itself hyperosmotically between 15.6 and 59.5 mosmol/ L (Reynoldson and Davies 1976) witha correspondingly low survival rate in salt water. Ball and Fernando (1970) concluded on the basis of the dissimilarities between the freshwater triclad faunae of Anticosti Island, Gaspé, and the Quebec North Shore, and the similarity on Anticosti Island of THE CANADIAN FIELD-NATURALIST Vol. 93 the triclad distribution and amphibian introductions to the island by man in 1899 (Johansen 1924) that triclads were also introduced to Anticosti Island by man. As the triclads and amphibian introductions are located in the southern half of Anticosti Island, however, the restriction of leeches to the northern half “strongly negates the probability of man having introduced leeches to Anticosti Island. Active directional dispersion has been recorded for Percymoorensis marmorata in Quebec (Richardson 1942) and for Erpobdella punctata in Michigan (Sawyer 1970). But as neither species is recorded from the Quebec North Shore nor Anticosti Island and dispersion to Anticosti Island would require an initial downstream movement into the sea rather than the upstream movement recorded, active directional dispersion to Anticosti Island is improbable. By elimination, the only probable mode of dis- persion of leeches to Anticosti Island is passive dispersal by sea currents. For AH. stagnalis, H. triserialis, and G. complanata the adults could either be attached to floating debris or free in the water, but for M. fervida the dispersive place could be either the cocoons or the adults. The theory of passive dis- persion of leeches in sea currents to Anticosti Island from the Quebec North Shore is substantiated by examination of the predominant currents (Figure 1) which flow along the Quebec North Shore with a south-southeast orientation to the mouth of the St. Lawrence River and then along the Gaspé coast (Cameron 1958). It thus appears that the freshwater leeches of Anti- costi Island have reached the island from the Quebec North Shore rather than from the Gaspé passively through the action of sea currents. This is the first probable record of passive dispersion in sea currents for freshwater leeches. The samples from Anticosti Island were collected by Ian Ball(then of the University of Waterloo) for my identification. Literature Cited Ball, I. R. and C. H. Fernando. 1970. Freshwater triclads (Platyhelminthes, Turbellaria) from Anticosti Island. Naturaliste Canadien 97: 331-336. Bleakney, J.S. 1958. A zoogeographical study of the amphibians and reptiles of eastern Canada. National Museum of Canada Bulletin 155: 1-119. Cameron, A. W. 1958. Mammals of the islands in the Gulf of St. Lawrence. National Museum of Canada Bulletin 154: 1-165. Daborn, G. R. 1976. Colonization of isolated aquatic habi- tats. Canadian Field-Naturalist 90: 56-57. Davies, R. W. 1973. The geographic distribution of fresh- water Hirudinoidea in Canada. Canadian Journal of Zoology 51: 531-545. 1979 Johansen, F. 1924. A biological excursion to Anticosti Island. Canadian Field-Naturalist 38: 161-164. Reynoldson, T. B. and R. W. Davies. 1976. A comparative study of the osmoregulatory ability of three species of leech (Hirudinoidea) and its relationship to their distribu- tion in Alberta. Canadian Journal of Zoology 54: 1908-1911. Richardson, L.R. 1942. Observations on migratory be- haviour of leeches. Canadian Field-Naturalist 56: 67-70. Sawyer, R.T. 1970. Observations on the natural history and behavior of Erpobdella punctata (Leidy) (Annelida: NOTES Sis Hirudinea). American Midland Naturalist 83: 65-80. Schmitt, J. 1904. Monographie de I’Ile d’Anticosti. Li- brairie scientifique Paris, A. Hermann, 370 pp. Stockwell, C. H. 1957. Geology and economic minerals of Canada. Economic Geology Series 1: 1-517. Verrill, A. E. 1863. Notes on the natural history of Anti- costi Island. Proceedings of the Boston Society of Natural History 9: 132-135. Received 5 December 1978 Accepted 14 February 1979 Relative Efficiencies of Museum Special, Victor, and Holdfast Traps for Sampling Small Mammal Populations ARTHUR M. MARTELL Canadian Wildlife Service, Great Lakes Forest Research Centre, Sault Ste. Marie, Ontario P6A 5M7 Present Address: Canadian Wildlife Service, 204 Range Road, Whitehorse, Yukon Territory YIA 4Y4 Martell, Arthur M. 1979. Relative efficiencies of Museum Special, Victor, and Holdfast traps for sampling small mammal populations. Canadian Field-Naturalist 93(3): 313-315. Small mammal populations in northern Ontario boreal forest were sampled with paired traps. Relative to Museum Specials, Victors captured proportionally fewer of most species, but Holdfasts captured more soricids, an equal number of cricetids, and fewer zapodids and sciurids. The most important factors determining the capture rate appeared to be the type of bait treadle in Museum Special-Victor pairs and trap size in Museum Special—Holdfast pairs. Significant differences in treadle sensitivity were not reflected in the catch. Key Words: trap selection, trapping bias, small mammals. When studying the demography of small mammal populations it is necessary to obtain relatively unbiased samples. This is often attempted with one, or more, of the several types of kill traps commercially available. Some types have been shown to be significantly more effective than others in capturing some species of small mammals (Pruitt and Lucier 1958; Neal and Cock 1969; Smith et al. 1971; Wiener and Smith 1972). Also the size of the trap has been shown to influence the mean weight of the catch (Neal and Cock 1969). Because body weight is often used as an estimate of age in small mammal studies, estimates of age structure of the population would also be biased. Therefore, the type of trap used may sig- nificantly influence the estimates of the abundance of the population, the age (weight) structure of the population, and the relative abundances of the various species in the community. The purpose of the present study was to determine which trap-related biases were present in the small mammal-forestry research being conducted by the Canadian Wildlife Service, Ontario Region. Materials and Methods Two extensively used small mammal kill traps are the Museum Special and the Victor mouse trap. The former measures 69 X 141 mm and has a 12 X 26 mm wood-covered bait treadle, while the latter measures . 48X99 mm and has an I! X15 mm metal bait treadle. A third, occasionally used type is the Holdfast trap, which is the same size as the Victor but has the same triggering mechanism and treadle as the Mu- seum Special. All three traps are manufactured by Woodstream Corporation, Niagara Falls, Ontario L2E 673, or Lititz, Pennsylvania 17543. I compared the relative efficiencies of these three traps by sampling with a Museum Special and either a Victor or a Holdfast at each trapping point. Strings were attached to the traps so that they could be placed in likely capture locations within a |-m radius of the trapping points, which were approximately 15 m apart and located either in a single line or in a grid. Any line or grid contained only one combination of traps. Traps were baited with a mixture of peanut butter, rolled oats, and rendered bacon fat, and were 314 checked on 3 consecutive days. Trapping was conducted in 1976 and 1977 on a variety of uncut and clearcut boreal forest sites near Manitouwadge, Ontario (May to October), and ona series of uncut and strip-cut boreal forest sites near Beardmore, Ontario (September). A total of 12 324 trapnights was accumulated using Museum Special— Victor pairs and a total of 20 572 trapnights using Museum Special-Holdfast pairs (1 trap set for | night = | trapnight). The sensitivity of the trap types was measured by determining the maximum weight, to the nearest gram, which could be placed on the distal part of the treadle and still allow the trap to be set. Fifty of each type of trap were tested after they had been used in the field for at least four rounds of normal trapping. The weights used were standard balance weights. Results and Discussion The average (x + SE) weight that could be carried on the distal end of the treadle without tripping the trap was 1.8+0.08g for Museum _ Special, 3.6 + 0.18 g for Holdfast, and 7.0 + 0.32 g for Victor traps, which reflects the relative sensitivities of the traps. The Holdfast held significantly (P<0.001) more weight than did the Museum Special, and the Victor held significantly (P<0.001) more weight than the Holdfast. The range in weights carried was also different between trap types, with the Victor showing the greatest range: Museum Special, 1-3 g; Holdfast, DOs NICOIn Sal Ole: For Museum Special-Victor pairs I compared the proportion of Victor-caught animals weighing less than 10 g, the maximum observed weight carried by a Victor, with the proportion weighing more than 10 g and found no significant differences (chi-square test) in those proportions for soricids (P > 0.1), cricetids and zapodids combined (P > 0.9), or all three groups combined (P > 0.1). For Museum Special-Holdfast pairs I compared the proportion of Holdfast-caught animals weighing less than 6 g, the maximum obser- ved weight carried by a Holdfast, with the proportion weighing more than 6g and found no significant difference in that proportion for soricids (P > 0.1), but found that the difference was significant for all three groups combined (P< 0.001). (All cricetids and zapodids captured weighed more than 6g.) The Holdfasts therefore caught a greater proportion of animals weighing less than 6 g than did the Museum Specials. The significant differences in triggering weight were not reflected in the catch. If triggering weight was not a factor, then the captures with Museum Special—Holdfast pairs would be influenced only by trap size, and captures with Museum Special-Victor pairs would be influenced by trap size and treadle composition. Table 1 shows that THE CANADIAN FIELD-NATURALIST Vol. 93 relative to Museum Specials, Holdfasts caught sig- nificantly more Masked Shrews (Sorex cinereus) and total soricids, and significantly fewer Meadow Jump- ing Mice (Zapus hudsonius) and Least Chipmunks (Eutamias minimus). Relative to Museum Specials, Victors caught significantly fewer Masked Shrews, total soricids, Deer Mice (Peromyscus maniculatus), Southern Red-backed Voles (Clethrionomys gap- peri), Heather Voles (Phenacomys intermedius), Rock Voles (Microtus chrotorrhinus), total cricetids, and total small mammals. Eight of the nine other species showed a lower capture rate in Victors than in Museum Specials. Trap size appears to be an important factor in determining the capture rate of three species in Museum Special—Holdfast pairs. Body size in relation to trap size may explain the high capture rates of Masked Shrews in Holdfasts, and Least Chipmunks in Museum Specials. If so, then treadle composition clearly outweighs trap size as a factor influencing the capture rate of Masked Shrews in Museum Special— Victor pairs. Smith et al. (1971) and Wiener and Smith (1972) observed a similar difference in capture rate between Museum Specials and Victors, and suggested that Museum Specials were more efficient than Victors because of their more sensitive triggering mechanism. My data do not support that suggestion; rather, the type of bait treadle is indicated as the main factor influencing the difference in capture rate between Victors and Museum Specials. The influence may be due, at least in part, to the wood-covered bait treadle of the Museum Special which likely retains odors better than the metal Victor treadle. There were no significant differences between trap type in mean body weight of any species captured in Museum Special—Holdfast pairs (Table 1). In Mu- seum Special-Victor pairs, however, Southern Red- backed Voles and Meadow Jumping Mice caught in Museum Specials were significantly heavier than those caught in Victors. Differences in mean weight would be expected to be due to differences in triggering weight, but that does not appear to be the cause in my study. There was no significant difference (chi-square test) in the proportions of species captured 10 or more times in Museum Specials and Victors (P > 0.1), but there was a significant difference between Museum Specials and Holdfasts (P< 0.001). That difference was not apparent within either soricids (P > 0.9) or cricetids (P > 0.1), suggesting that the difference was between taxonomic groups. When the proportions of total captures of soricids, cricetids, zapodids, and sciurids were compared, there was a significant difference (chi-square test) between Museum Specials and Holdfasts (P< 0.001) but not between Museum 1979 NOTES 315 TABLE !—Numbers and mean weights (g) of small mammals trapped in Museum Special-Victor pairs (M—V) and Museum Special-Holdfast pairs (M-H), and the significance (chi-square test and t-test) of the differences within pairs. Sample sizes for weights approximate those given in the number columns, but in some cases are slightly smaller Museum Special-Victor Museum Special—Holdfast Species Number Weight Number Weight M Vv M Vv M H M H Sorex cinereus fal 4|** 355) 3.4 88 160*** 3.5 3.6 Sorex palustris 2 0 15.8 0 1 14.1 Sorex arcticus 2. fed 7.6 3 | 7.1 10.2 Microsorex hoyi 0 l 3.3 0 3 4.1 Blarina brevicauda 7 2 18.5 18.0 8 16 19.9 18.6 Total soricid captures 81 46** 99 Sees Peromyscus maniculatus 176 67 Ett 15.4 15.4 267 235 16.2 15.8 Clethrionomys gapperi 240 142*** 20.8 19.6* 196 196 Me3) 20.4 Phenacomys intermedius 28 10** 21.0 24.5 36 DF 19.7 23.2 Synaptomys cooperi 2 0 24.4 l 3 24.3 19.9 Microtus pennsylvanicus 14 1} 24.4 23.6 17 21 25.4 24.0 Microtus chrotorrhinus 25 12* 23.5 22.4 6 11 24.2 Des) Total cricetid captures 485 242*** 523 493 Zapus hudsonius 6 2 ae he 5 0* 14.9 Napaeozapus insignis 2 21.4 0 2 28.3 Total zapodid captures 8 2 5 2 Tamias striatus 0 0 2 2 68.1 91.8 Eutamias minimus 9 7 41.0 45.1 40 hoe? 49.3 47.4 Total sciurid captures 9 7 42 20-% Total, all species 583 DOU SAX 669 696 *P< 0.05. **#P< 0.01. *** P< (0.001. Specials and Victors (P > 0.5). Relative to Museum Specials, therefore, Victors captured proportionally fewer of all species, but Holdfasts captured more soricids, an equal number of cricetids, and fewer zapodids and sciurids. In contrast to my results, other studies have shown significant differences among species in their capture rate by Museum Specials and Victors. I tested data presented by Smith et al. (1971) and Wiener and Smith (1972) for differences in the proportions of species captured 10 or more times in Museum Special-Victor pairs, and found that the difference was significant in both studies (P< 0.001). Small- mammal trappers, therefore, should be aware of these differences and choose their trap type(s) accordingly. Acknowledgments I am indebted to the Great Lakes Forest Research Centre, the Ontario Ministry of Natural Resources, the Ontario Paper Company and Domtar Ltd. for their cooperation. I thank A. L. Macaulay for her assistance in all aspects of the study, and D. A. Welsh, T. C. Dauphine, J. E. Bryant, and J. F. T. Carreiro for their comments on the manuscript. = Literature Cited Neal, BR. and A.G. Cock. 1969. An analysis of the selection of small African mammals by two break-back traps. Journal of Zoology 158: 335-340. Pruitt, W. O., Jr. and ¢. V. Lucier. 1958. On the relative efficiency of two kinds of traps. Journal of Mammalogy 39: 157. Smith, G. C., D. W. Kaufman, R. M. Jones, J. B. Gentry, and M. H. Smith. 1971. The relative effectiveness of two ! types of snap traps. Acta Theriologica 16: 284-288. Wiener, J. G.and M. H. Smith. 1972. Relative efficiencies of four small mammal traps. Journal of Mammalogy 53: 868-873. Received 30 November 1978 Accepted 5 March 1979 316 THE CANADIAN FIELD-NATURALIST Vol. 93 Probable Hybrids of Cinnamon X Blue-winged Teal from Southern Alberta’ D. V. WESELOH and LINDA MCKEANE WESELOH Provincial Museum of Alberta, 12845 102 Avenue, Edmonton, Alberta T5N 0M6 Present Address: Canadian Wildlife Service, Canada Centre for Inland Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6 Weseloh, D. V. and Linda McKeane Weseloh. 1979. Probable hybrids of Cinnamon X Blue-winged Teal from southern Alberta. Canadian Field-Naturalist 93(3): 316-317. Plumage characteristics of three recent and nine previously reported probable hybrid Cinnamon X Blue-winged Teal are summarized: males are the rufous color of Cinnamon Teal but retain the white facial crescent and often the white flank patch of the Blue-winged Teal. Key Words: Alberta, hybrids, Cinnamon Teal, Blue-winged Teal, Anas cyanoptera, Anas discors. Until recently most of our knowledge on the occurrence of Cinnamon (Anas cyanoptera) X Blue- winged (A. discors) Teal hybridization has come from aviaries. Kortwright (1942: p. 212) states that“... even in captivity crosses between the two species are unknown.” Delacour and Mayr(1945), however, state that the two species freely interbreed. Spencer (1953: p. 20) states that “although extremely uncommon among wild Cinnamon Teal, hybrids are by no means unknown.” Of the Blue-winged Teal, Delacour (1956; p. 170) remarks that “In captivity they cross too readily with Cinnamon Teal, and when both species are kept on the same pond it is usual to rear a majority of hybrids . . . This is the more surprising that these two closely allied Teal seldom cross in the wild state in the rather narrow areas where they co-exist in North America.” He goes on to say that Cinnamon Teal “|. . persecute the weaker Blue-winged Teal...” In recent years there has been an increasing number of reports of wild Cinnamon X Blue-winged Teal hybrids; many of these have come from southern Alberta and Saskatchewan (Wedgewood and Wedge- wood 1975; Lang 1973; Lahrman 1971). Also there have been reports of male Cinnamon Teal and Blue- winged Teal simultaneously courting a single female of one of the species (Butot 1974; Andersonand Miller 1953). In the spring of 1974, we observed three ducks which we believe were male hybrids of Cinnamon and Blue-winged Teal. One individual was observed on 20 May at 16:55, and another at 17:02 at a slough 0.5 km east of the southeast corner of Eagle Lake, near Namaka, approximately 56 km east- southeast of Calgary, Alberta. The first duck was 'Natural History Contribution No. 40, Provincial Museum of Alberta, Edmonton, Alberta alone and swam from view within 2-3 min of being sighted. The second bird was in the company of two normally plumaged male Cinnamon Teal and a female typical of the nearly identically appearing Blue-winged and Cinnamon Teal females. The third individual was observed by us on 9 June at 19:45, at Second Vermilion Lake, Banff, Alberta. It was in the company of two normally plumaged male Cinnamon Teal and a female of either teal species. A search of literature as well as present sightings yields 11 field descriptions of apparently wild Cin- namon X Blue-winged Teal hybrids. It is not sur- prising that all individuals have been male as hybrid females would be virtually undistinguishable in the field. We have described the major plumage char- acteristics in Table 1. In summary, the plumage of male Cin- namon X Blue-winged Teal hybrids may be char- acterized as follows: a body having the rufous coloration of the Cinnamon Teal and the white facial crescent of the Blue-winged Teal; the white flank patch of the latter is often present and the breast is sometimes spotted. We express our appreciation to Bob Brown, Michael Bradstreet, and two anonymous reviewers who made comments on earlier versions of this manuscript. Literature Cited Anderson, W. and A. W. Miller. 1953. Hybridization of the Cinnamon and Blue-winged Teal in Northern Cali- fornia. Condor 55: 152-153. Bent, A.C. 1923. Life histories of North American wildfowl. Part 1. U.S. National Museum Bulletin 126. Reprinted 1951. Butot, R. 1974. Ménage a trois, or the end of a species. Calgary Field Naturalist 5(11): 281-282. 1979 NOTES SHl7/ TABLE 1—Plumage features of male Cinnamon X Blue-winged Teal hybrids White facial White flank Breast Body coloration crescent patch spotting Location and source 1. Dark cinnamon Distinctive Distinctive Utah; Wilson and Van den Akker 1948 2. Cinnamon Snow-white Colorado; Bent 1923 3. Cinnamon Distinct Believed lacking Utah; Spencer 1953 4. Cinnamon Prominent Alberta; Spencer 1953 5. Cinnamon Clear Indistinct California; Anderson and Miller 1953 6. Reddish Washed out* Prominent* Present Saskatchewan; Lahrman 1971 7. Cinnamon Faint Alberta; Lang 1973 8. Rusty red Small Light Saskatchewan; Wedgewood and Wedgewood 1975 9. Cinnamon Prominent Prominent Alberta; this paper 10. Cinnamon Washed out Absent Alberta; this paper 11. Rufous-cinnamon Washed out Washed out Alberta; this paper *Authors’ interpretation from photo. Delacour, J. 1956. The waterfowl of the world. Country Life Ltd., London. Volume 2. 232 pp. Delacour, J. and E. Mayr. 1945. The Family Anatidae. Wilson Bulletin 57: 3-55. Kortwright, F. H. 1942. Ducks, geese and swans of North America. Stackpole Company, Harrisburg, Pennsy]l- vania and Wildlife Management Institute, Washington. llth printing. 476 pp. Lahrman, F.W. 1971. Hybrid Cinnamon Teal X Blue- winged Teal at Regina. Blue Jay 29: 28. Lang, V. 1973. Observation of probable Cinnamon Teal (Anas cyanoptera) X Blue-winged Teal (Anas discors) hybrid. Calgary Field Naturalist 5(4): 139. Spencer, H.E., Jr. 1953. The Cinnamon Teal (Anas cyanoptera Vieillot): Its life history, ecology and man- agement. M.Sc. thesis, Utah State Agricultural College, Logan, Utah. 184 pp. Wedgwood, S.L. and J. A. Wedgwood. 1975. Probable Cinnamon Teal — Blue-winged Teal cross. Blue Jay 33(3): 181-182. Wilson, V.T. and J.B. Van den Akker. 1948. A hybrid Cinnamon Teal - Blue-winged Teal at the Bear River Migratory Bird Refuge, Utah. Auk 65: 316. Received 20 October 1976 Resubmitted 28 February 1979 Accepted 5 March 1979 Common Garter Snake Predation on Ring-billed Gull Chicks PETER M. FETTEROLF Department of Zoology, University of Toronto, Toronto, Ontario MSS I1A1 Fetterolf, P.M. 1979. Common Garter Snake predation on Ring-billed Gull chicks. Canadian Field-Naturalist 93(3): 317-318. A Common Garter Snake was observed eating two freshly hatched Ring-billed Gull chicks. Key Words: Common Garter Snake, Ring-billed Gull chicks, predation, Thamnophis sirtalis, Larus delawarensis. In this note I describe an act of predation upon Ring-billed Gull (Larus delawarensis) chicks by a Common Garter Snake, Thamnophis sirtalis. The gull colony is on Mugg’s Island, Toronto Harbor, Toronto, Ontario. The observation was made froma blind situated in a Ring-billed Gull colony of approximately 6000 pairs. About 50 pairs of Herring Gulls, L. argentatus, also nest on the island. The incident began at 07:35 EST on 25 May 1978 as the snake moved from beneath the platform which supports the blind. The 80-cm reptile moved approxi- mately 4 m to the edge of a gull nest which contained three chicks. Two chicks were about 48 h old and the third was less than 24 h old. Freshly hatched chicks average 35-45 g. The snake tested one half of an eggshell lying at the nest rim with its tongue, grasped the shell between its jaws, and moved it about 20 cm from the nest. The snake then raised its head approximately Scm above the substrate, froze momentarily, and lunged rapidly at a chick, seizing it by the bill. The snake engulfed the chick up to the wings within 4 min and then writhed several times. 318 The chick was gone from view 14 min after capture, and the snake remained stationary for a few minutes more. As it contorted slowly and rubbed its nose in the sand, the obvious bulge moved posteriorly. Following this respite, the reptile inched its way toward a nest about | mdistant which contained eggs. The contents were tested with the tongue, and the snake continued to two adjacent nests with eggs where it performed the same operation. Finally, a nest with two 48-h-old chicks was reached. The two chicks were different in size, and the Garter Snake seized the smaller one by the throat. The snake released the chick long enough to grasp it by the bill as it had done with the first victim. The swallowing behavior which followed was similar to that in the first predatory act 23 min earlier, and |! minafter it grasped the chick the snake disappeared beneath the blind. At that time, 80% of the chick had disappeared within the snake’s jaws. The gulls’ reaction to the snake was interesting. Hatching had just started in the area and during the entire incident all chicks remained in the nest bowls or on their rims. Adult gulls stood about 0.75 m from the snake and called loudly, using the “Kow-Kow’ call described by Moynihan (1956). Adults displayed no anti-predator behavior (Krunk 1964) and none took flight at the sight of the reptile. Gull chicks are consumed by avian and mammalian predators such as the Great Horned Owl, Bubo virginianus (Burger 1974); Marsh Hawk, Circus cyaneus (Burger 1974); Mink, Mustela vision (Burger 1974); Hedgehog, Erinaceus europaeus (Kruuk 1964); Red Fox, Vulpes vulpes (Kruuk 1964; Burger 1974). Campbell (1969) reported that the Wandering Garter Snake, 7. elegans vagrans, preyed upon Glaucous- winged Gull, L. glaucescens, chicks. Birds are rare in the diet of 7. sirtalis (Hamilton 1951; Fouquette 1954; Fitch 1965; Gregory 1978). Thus, consumption of Ring-billed Gull chicks may be a special case of opportunistic feeding. Campbell’s observations of Wandering Garter Snake predation on Glaucous- winged Gull chicks occurred in an ecological situation similar to that on Mugg’s Island. In both cases, the snakes were the only observed reptilian inhabitants of the islands used by nesting gulls. Perhaps the snakes, which use olfaction to detect food (Fox 1952; Burghardt 1970), were originally attracted to the colonies by the odor of fish scraps dropped by the gulls. The Wandering Garter Snake ate food pellets regurgitated by gulls (Campbell 1969). The fact that THE CANADIAN FIELD-NATURALIST Vol. 93 the snake on Mugg’s Island struck at and grasped the fresh eggshell, which was unlikely to smell of fish, suggests that the snake may have utilized chick odor in its hunting behavior. It is difficult to assess the frequency of gull chick predation by the Common Garter Snake on Mugg’s Island, for it has only been observed once in over 1200 h of observations in the past 4 yr. Therefore the incident reported here is probably an example of a predator taking a locally abundant, easily obtained food source not normally in its diet. Acknowledgments My thanks go to D. W. Dunham who provided financial support from a National Research Council of Canada operating grant, to A. R. Gibson who provided some garter snake references, and to my wife who did the typing of the manuscript. Literature Cited Burger, J. 1974. Breeding adaptations of Franklin’s Gull (Larus pipixcan) to a marsh habitat. Animal Behavior 22: 521-567. Burghardt, G. M. 1970. Intraspecific geographical varia- tions in the chemical food cue preference of newborn garter snakes. Behavior 36: 246-257. Campbell, R. W. 1969. Notes onsome foods of the wander- ing garter snake on Mitlenatch Island, British Columbia. Syesis 2: 183-187. Fitch, H. S. 1965. An ecological study of the garter snake, Thamnophis sirtalis. University of Kansas Publications, Museum of Natural History 15: 393-464. Fouquette, M. J., Jr. 1954. Food competition among four sympatric species of garter snakes. Texas Journal of Science 1964 (2): 172-188. Fox, W. 1952. Notes on the feeding habits of Pacific coast garter snakes. Herpetologica 8: 4-8. Gregory, P. J. 1978. Feeding habits and diet overlap of three species of garter snakes (Thamnophis) on Vancouver Island. Canadian Journal of Zoology 56(9): 1967-1974. Hamilton, W. J., Jr. 1951. The food and feeding behavior of the garter snake in New York state. American Midland Naturalist 46: 385-390. Kruuk, H. 1964. Predators and anti-predator behavior of the Black-headed Gull (Larus ridibundus L.). Behavior Supplement I1. Moynihan, M. 1956. Notes on the behavior of some North American gulls. 1. Aerial hostile behavior. Behavior 10: 126-179. Received 3 November 1978 Accepted 3 February 1979 1979 NOTES 319 The Rock Vole (Microtus chrotorrhinus) as a Transition Zone Species GORDON L. KIRKLAND, Jr. and CHARLES M. KNIPE Vertebrate Museum, Shippensburg State College, Shippensburg, Pennsylvania 17257 Kirkland, Gordon L., Jr. and Charles M. Knipe. 1979. The Rock Vole (Microtus chrotorrhinus) as a Transition zone species. Canadian Field-Naturalist 93(3): 319-321. Rock Voles (Microtus chrotorrhinus) were collected in mixed northern hardwood forests in New York’s Adirondack Mountains at elevations between 460 and 700 m. These voles were much more common in the Transition zone habitats than previously thought. Data are presented on the habitat, relative abundance, and ecological associates of the 43 specimens. Key Words: Rock Vole, Microtus chrotorrhinus, Adirondack Mountains, Canadian zone, Transition zone. Rock Voles (Microtus chrotorrhinus) have tradi- tionally been considered to be boreal small mammals. Recent research at the periphery of their range indicates that Rock Voles are not only geographically more widespread than previously believed, but they have greater ecological amplitude as well (Buech et al. 1977; Kirkland 1977; Roscoe and Majka 1976; Timm 1974: Timm et al. 1977). In the northeastern United States, Rock Voles have been thought to be limited in distribution to higher elevations and more boreal habitats. Martin (1971a) states that the Rock Vole is most closely associated with the Canadian life zone. Furthermore, he notes that records of the Rock Vole in the Transition zone may be explained on the basis of expansion of the Rock Vole’s distribution under suitable conditions from the sanctuary of more favorable Canadian zone habitat. Examination of the distribution of captures of the Rock Vole in New York’s Adirondack Mountains prior to 1970 tends to confirm this traditional view of the ecological distri- bution of the Rock Vole. With the exception of a series of 29 specimens collected by C. F. Batchelder at Keene Heights (elevation 400 m), 61 of 91 specimens examined in museum collections were taken at elevations above 976 m, which is the lower limit of Red Spruce (Picea rubens)-dominated forests in the Adirondacks (Braun 1964). Since 1970, however, 43 Adirondack Rock Voles collected by Shippensburg State College Vertebrate Museum field personnel have been trapped at elevations below 1100 m. In addition, F. J. Jannett, Jr. (Cornel! University) and R. Rosen (University of Vermont) (personal com- munications) have obtained for use in laboratory studies 20 live Rock Voles at elevations between 640 and 760m in the Adirondacks. These recent records reveal that Rock Voles are more widespread and abundant in Transition zone habitats than was previously believed. This paper presents information on the habitat, relative abundance, and ecological associates of 43 M. chrotorrhinus captured from 1973-1978 at elevations of 460 to 700 m in Essex County, New York. Methods The 43 Rock Voles described in the paper were collected in 10 traplines (Table 1) representing seven localities, as follows: New York, Essex County, St. Hubert’s, 2.7 km SE, elevation 457 m (1); Tahawus, 0.40 km SE, elevation 549 m (1); 1.9 km W, elevation 549m (1); 7.9km NNW, elevation 640m (26); 8.7 km N, 3.2 km W, elevation 670 m (9); Wallface Mountain, 0.8km SW, elevation 700m (4); 3.2 km SW, elevation 625 m (1). The specimens were obtained as part of a small mammal survey of Essex County, New York. Sampling procedures involved establishing traplines with variable numbers of sta- tions, usually with three snap-traps per station. Mu- seum Specials baited with rolled oats were used princi- pally, although each line might have up to 10% rat and/ or mouse traps. At each station, the traps were set with- in 2 m of each other, and traplines were operated for either 2 or 3 d. The 10 traplines yielding Rock Voles had a total sampling effort of 2691 trapnights (TN). Results and Discussion The trapline and specimen records for the 43 Rock Voles reveal that 42 were captured in typical Transi- tion zone forests dominated by Yellow Birch (Betula | lutea), Sugar Maple (Acer saccharum), and American Beech ( Fagus grandifolia). The only exception was the single capture along a stream, 2.7 km SE St. Hubert’s, in a Red Spruce — Eastern Hemlock ( Tsuga canaden- sis) association. All specimens were captured at sites where rocks or boulders were conspicuous habitat components and the canopy was semi-open. Six of the seven localities at which 42 of the Rock Voles were captured had streams of varying sizes flowing through them. Martin (1971a) notes that 47 of 83 Rock Voles he collected were caught within 30 ft (9 m) of running water, either surface or subsurface. The microhabitats of the Rock Voles in this study tended to be characterized by the presence of rocks, mosses, ferns, and forbs (e.g., Oxalis, Clintonia, Viola, and Smilacina). The percentages of Rock Voles collected within 1m of these four microhabitat 320 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Small mammals captured in 10 traplines yielding Rock Voles in Transition zone habitats of Essex County, New York Trapline number, year, elevation (m), sampling effort (TN) Ie 2 4; 73, NOS. 549, 457, Species 450 44] Masked Shrew (Sorex cinereus) Long-tailed Shrew (Sorex dispar) 2 Smokey Shrew (Sorex fumeus) 3) Water Shrew (Sorex palustris) Short-tailed Shrew (Blarina brevicauda) 5 4 Chipmunk (Tamias striatus) 2 Red Squirrel (Tamiasciurus hudsonicus) Northern Flying Squirrel (Glaucomys sabrinus) Deer Mouse (Peromyscus maniculatus) 6 8 Red-backed Vole (Clethrionomys gapperi) ] 16 Rock Vole (Microtus chrotorrhinus) I I Meadow Vole (Microtus pennsylvanicus) Woodland Jumping Mouse ( Napaeozapus insignis) 3} I Meadow Jumping Mouse (Zapus hudsonius) Totals 18 35 a 4, 5, 6. i 8, 9, 10, Totals 1975, P1975," 1975, 1977, 1977, 1978, ealo7sammnenee 549, 625, 700, 640, 640, 640, 640, 700, 540 300 300 240 240 120 120 240 2691TN 6 3 I 2 Ai, LEG I 2 I 2 I 9 2 I 6 2 2 24 12 5 7 10 I 1 69 3 6 I I I 2 16 14 il 4 2 2 2 4 69 14 8 in uae 825 84 I 4 14 10 I Omens I I 1 4 5 4 2 2 2 34 I I [Sie merase 143 OES 9 9 19 343 components were as follows: rocks (66%), mosses (88%), ferns (85%), and forbs (91%). Live ground cover at individual capture sites varied from less than 25 to more than 75% but averaged 50-60%. The Rock Voles appeared to spend a considerable portion of their time in subterranean activity. In this survey, 53.5% of the specimens were caught in traps set under rocks or below the surface of rocky sites, and 65% of the total were caught in “unexposed” sites, either under rocks, logs, roots, or overhangs. The 43 Rock Voles were captured in association with 13 other species of small mammals (Table 1). They were most frequently captured in association with Deer Mice (Peromyscus maniculatus gracilis), Short-tailed Shrews ( Blarina brevicauda), and Wood- land Jumping Mice (Napaeozapus insignis). Although collected in only 7 of 10 traplines, Red- backed Voles (Clethrionomys gapperi) were the most abundant ecological associate with a total of 84 specimens. Martin (1971b) noted that he collected a “relatively unchanging proportion of approximately three C. gapperi for every M. chrotorrhinus in favorable rock vole habitat.” In this study, the proportion of C. gapperi to M. chrotorrhinus was 2.0:1.0; however, this varied considerably between traplines. In individual traplines, the ratio of these two species ranged from 16:1 in favor of C. gapperito 9:0 in favor of M. chrotorrhinus. In October of 1965 and 1966, J. N. Layne and students from Cornell University sampled small mammals on Whiteface Mountain, Essex County, New York. They collected 20 M. chrotorrhinus at elevations from 1128 to 1433 min 10 of their traplines, with a total sampling effort of 1069 TN. These previ- ously unpublished data permit some comparisons of Rock Voles inhabiting Canadian and Transition zones in the Adirondacks. The catch per unit sampling effort of 1.87/100 TN for the Whiteface Mountain sampling was slightly higher but did not differ significantly from the 1.60/ 100 TN for the Transition zone sampling in this study (0.75 > P > 0.50). This indicates that in the Adirondacks, Rock Voles may have comparable population levels in suitable Can- adian and Transition zone habitats. The 20 Rock Voles on Whiteface Mountain were captured in traplines yielding seven other species of small mam- mals as follows: Masked Shrew (Sorex cinereus) (3), Long-tailed Shrew (Sorex dispar) (6), Smokey Shrew (Sorex fumeus) (1), Short-tailed Shrew (3), Deer Mouse (12), Red-backed Vole (27), and Meadow Vole (Microtus pennsylvanicus) (5). It is of interest that on Whiteface Mountain, the ratio of Red-backed Voles to Rock Voles was only 1.4:1.0. A Spearman Rank Correlation analysis (Siegel 1956) of the species of small mammals collected on Whiteface Mountain and in the Transition zone habitats (Table 1) reveals a significant correlation (rs = 0.696, P< 0.05) between the relative abundance of species in small mammal communities containing Rock Voles in the two ecological zones. Thus, although the Rock Voles are occupying dissimilar plant communities, they are members of qualitatively similar small mammal communities. The recent capture of Rock Voles at a variety of locations in the Transition zone of the Adirondack Mountains suggests that traditional assessments of 1979 this species’ ecological distribution should be re- evaluated. Rather than being restricted primarily to boreal habitats characteristic of the Canadian zone, the Rock Voles also occupy sites in northern hardwood forests where appropriate microhabitat components are present, specifically rocks, flowing water, mosses, ferns, and forbs. Acknowledgments We thank James N. Layne for permitting us to use data he and his students had collected on Rock Voles at Whiteface Mountain and for critically reviewing a draft manuscript. Our thanks are also extended to Walter M. Chapman, Manager, of the MacIntyre Development, N L Industries, Tahawus, New York for providing housing and access to private lands sampled in this study. We acknowledge Carol J. Kirkland, Henry W. Setzer, and the students from Shippensburg State College who assisted in collecting the specimens described in this paper. Literature Cited Braun, E. L. 1964. Deciduous forests of eastern North America. Hafner Publishing Company, New York. 596pp. Buech, R. R., R. M. Timm, and K. Siderts. 1977. Asecond population of the Rock Vole, Microtus chrotorrhinus, in NOTES 321 Minnesota with comments on habitat. Canadian Field- Naturalist 91: 413-414. Kirkland, G.L., Jr. 1977. The Rock Vole, Microtus chrotorrhinus (Miller) (Mammalia: Rodentia) in West Virginia. Annals of the Carnegie Museum of Natural History 46: 45-53. Martin, R. L. 1971a. The natural history and taxonomy of the Rock Vole, Microtus chrotorrhinus. Ph.D. thesis, University of Connecticut, Storrs. 123 pp. Martin, R.L. 1971b. Interspecific associations of Rock Voles. Beta Kappa Chi Bulletin 30(2): 5-7. Roscoe, B. and C. Majka. 1976. First record of the Rock Vole (Microtus chrotorrhinus) and the Gaspé shrew (Sorex gaspensis) from Nova Scotia and the second record of Thompson’s pygmy shrew (Microsorex thompsoni) from Cape Breton Island. Canadian Field-Naturalist 90: 497-498. Siegel, S. 1956. Nonparametric statistics for the behavioral sciences. McGraw-Hill Book Company, New York. 312 pp. Timm, R.M. 1974. Rediscovery of the Rock Vole (Microtus chrotorrhinus) in Minnesota. Canadian Field- Naturalist 88: 82. Timm, R.M., L.R. Heaney, and D.D. Baird. 1977. Natural history of Rock Voles (Microtus chrotorrhinus) in Minnesota. Canadian Field-Naturalist 91: 177-181. Received 21 December 1978. Accepted 12 March 1979. Eastern Limit of the five-lined Skink, Ewmeces fasciatus, in Ontario RICHARD D. USSHER! and FRANCIS R. COOK2 IRR 1, Delta, Ontario KOE 1G0 2Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8 Ussher, Richard D. and Francis R. Cook. 1979. Eastern limit of the Five-lined Skink, Eumeces fasciatus, in Ontario. Canadian Field-Naturalist 93(3): 321-323. New records are reported for the Five-lined Skink, Eumeces fasciatus, from Frontenac County (Palmerston Lake; 2.4 km W of Snow Road: and 1.6 km SE of Snow Road Station), Lanark County (Robertson Lake), and Leeds and Grenville (Landon Bay) County. The similarities between the eastern range limit in Ontario of the skink and those of the Northern Ribbon Snake and Black Rat Snake are discussed; the adjacent more poorly-drained lowlands are suggested as a distribution barrier. Based on collections reported by Patch (1934), Logier and Toner (1961, p. 59) cited Arden and Mountain Grove in Frontenac County as the most eastern localities at which the Five-lined Skink, Eumeces fasciatus, had been taken in Canada. Records accumulated since this publication allow a better definition of the eastern limit of its distribu- tion. On 30 August 1961, seven skinks were found by Cook and M. Gordon Foster 1.5 mi (2.4 km*) W of Highway 509 at Snow Road, Frontenac County (these are now in the National Museums of Canada, Collection Number 5618). They were inhabiting a rocky outcropping in the vicinity of a deserted hillside farmsite. One individual was discovered under a board at the farmsite: the others were under generally flat rocks in the adjacent woods and hillside. The woods were primarily White Pine (Pinus strobus) but a mixture of deciduous trees was also present. A thick layer of pine needles ‘covered most of the ground in the wooded area. Old lumber piles and scattered loose boards were prevalent at the farm site, and loose rocks were common in the woods and on the hillside. Additional records in this region include a single 322 individual taken by D. Scobie on 31 July 1960 from near Palmerston Lake, about 2 mi (3.2 km*) up the lake from Ompah (NMC 5110): one specimen collected on 10 August 1961 by A. G. Walker | mi (1.6 km*) SE of Snow Road Station, Palmerston Township, Frontenac County (Carleton University Museum of Zoology; D. A. Smith, personal com- munication); and one specimen collected by F.C. Zufeldt in May 1962 from Robertson Lake (NE of Lavant), Lanark County (NMC 6507). The map in Conant (1975) is based on the National Museum of Natural Sciences records, but individual localities have not been previously published. Luciuk and McCabe (1971) noted that the species is reported to occur at “3rd Depot Lake” and at Perth Road Bay near Chaffey’s Locks. The latter locality is about midway between Snow Road Station and Gananoque. On 22 August 1973 at the Landon Bay Campsite on the north shore of the St. Lawrence River, Ussher observed a single individual. The locality was a bare patch of Precambrian rock, some 30 m above the water of Landon Bay, 8.5 km E of Gananoque in Leeds and Lansdowne Township, Leeds and Gren- ville County. This sighting is the only one from the area by Ussher, although he became a resident of Leeds and Grenville County in 1970, and visited this locality and similar ones on many occasions. East of these records, in the Ottawa District, (a 48- km radius around the National Capital) where herpetofaunal surveys have been undertaken by the National Museum of Natural Sciences for several decades, and near Bishop’s Mills in Leeds and Grenville County, where Cook has resided since 1970, there have been neither sightings nor reports of skinks. Of interest is the roughly similar eastern limit of two other Ontario reptiles, the Black Rat Snake, Elaphe obsoleta obsoleta, which is known to occur from just south of Smith’s Falls to Mallorytown (NMNS files including personal communications from R. V. Lind- say, John Woods, Roger E. Roy, Harold Parsons, and Al MacDonald), and the Northern Ribbon Snake, Thamnophis sauritus septentrionalis, which reaches the “Horseback Mountains” near Pakenham (Cook 1968) and slightly beyond (Tony Tobias, personal communication) and Mallorytown Landing (Woods and Cook 1976). In his review of the herpetofauna of the Thousand Islands Region, New York, Werner (1959) reported both the Black Rat Snake and the Ribbon Snake, but not the Five-lined Skink. None of these three species have been reported from east of the rough and relatively well-drained elevated terrain of the Precambrian Shield topography, onto *Collectors originally stated distances in whole or half miles. THE CANADIAN FIELD-NATURALIST Vol. 93 the adjacent low-lying, and relatively more poorly- drained area which was covered by the postglacial Champlain Sea of the Ottawa and St. Lawrence valleys. Nor have any been reported on the eastern side of this presumed barrier, in the Precambrian of the Gatineau and Laurentian regions between Ottawa and Montreal. The populations of these three species reported in Leeds and Grenville, Frontenac, and Lanark counties may be prevented from colonizing the lowlands at the northeastern edge of their range by a lack of well- drained hibernating sites and the lack of sufficient insulation that is provided by deep accumulations of snow in the hollows of the adjacent broken top- ography. The moderating effect of the Rideau Lakes system on local microclimates, possibly promoting warm pockets, and the less intensive agriculture on its rough terrain, leaving more of the area in forest, are additional factors that may contribute to the survival of these species in the area. Their eastern limit roughly corresponds to the eastern boundary of Herpeto- faunal Section | defined by Bleakney (1958), which is based on mean July temperature and the length of the growing season. A fourth species, the Stinkpot, Sternotherus odor- atus, reaches its eastern limit in the general area, but it invades the lowlands along the Rideau and Missis- sippi rivers at least as far as Beckets Landing (NMC 3938) in Leeds and Grenville counties and Innisville (NMC 2157) and Pakenham (NMC 13777) in Lanark County. Perhaps its different overwintering habitat, in these rivers does not place on it the same constraints for hibernation sites that are suggested for the terres- trially hibernating species discussed above. Additional observations of any of these species at the eastern edge of their range, giving locality, habitat, date, number observed, and observer(s) name(s) would be appreciated by the authors. Information on the status of the Black Rat Snake is also being collected for the Ontario Ministry of Natural Re- sources by Harold Parsons and Al Macdonald as part of a detailed study of it in the Rideau Lakes region, and by the Federation of Ontario Naturalists in a general survey of its Ontario range. Because sightings of any of these species are often a matter of chance, and because populations may be scattered and fluctuating in abundance from year to year, the cumulative interest and efforts of all observant naturalists is needed further to define their ranges. A more precise discussion of these limiting factors must wait until such observations are available, and should also include comparison with the northern limit of these species. The latter may roughly correspond (with the notable exception of the Black Rat Snake) with the southern boundary of the 300-m contour and a large height of land covering the Haliburton and 72) Algonquin highlands (see map in Weller and Palermo 1976: W.G. Sprules and W.F. Weller, personal communication). Literature Cited Bleakney, J. Sherman. 1958. A zoogeographic study of the amphibians and reptiles of eastern Canada. National Museum of Canada Bulletin 155: 1-119. Conant, Roger. 1975. A field guide to reptiles and am- phibians of eastern and central North America. Houghton Mifflin Company, Boston. xvii + 429 pp. Cook, Francis R. 1968. Lizards and snakes of the Ottawa District. Trail & Landscape 2(4): 99-106. Logier, E. B.S. and G. C. Toner. 1961. A check list of the amphibians and reptiles of Canada and Alaska. Second Edition. Royal Ontario Museum, Life Sciences Division, Contribution 53: 1-93. Barn Owls in Quebec NORMAND DAVID NOTES 323 Luciuk, Lubomyr and Jim McCabe. 1971. The Five-Lined Skink (Eumeces fasciatus). Canadian Herpetologists Society Quarterly 1(2): 3. Patch, C. L. 1934. Euwmeces in Canada. Copeia 1934(1): 50-51. Woods, John G. and Francis R. Cook. 1976. Range extension of the Ribbon Snake in Eastern Ontario. Canadian Field-Naturalist 90(1): 69-70. Werner, William E. 1959. Amphibians and reptiles of the Thousand Islands area, New York. Copeia 1959(2): 170- 172. Weller, Wayne F. and R. Victor Palermo. 1976. A northern range extension for the Western Chorus Frog, Pseu- dacris triseriata triseriata (Wied), in Ontario. Canadian Field-Naturalist 90(1): 163-166. Received 10 November 1978 Accepted 16 March 1979 Centre de recherches écologiques de Montréal, Case postale 6128, Succursale “A,” Montréal, Québec H3C 3J7 David, Normand. 1979. Barn Owls in Quebec. Canadian Field-Naturalist 93(3): 323-324. An evaluation of Barn Owl records in Quebec shows that there is no conclusive evidence that the species has actually bred within the province. Key Words: Tyto alba, Quebec. Information from F. Créte, then Director of the Museum of the Deaf and Dumb Institute of Mon- treal, provided the only suggestion of breeding of the Barn Owl (Tyto alba) in Quebec, at Berthierville, Berthier County, in 1931 (Cayouette 1947; Godfrey 1966). The unpublished ornithological notes of the late Victor Gaboriault, however, show that breeding of Barn Owls in Berthierville in 1931 was never substan- tiated. An evaluation of that record and of other reported occurrences in the province indicates that the Barn Owl is a casual visitor, but gives no indisputable evidence that it breeds here. Early in 1945, R. Cayouette learned that a “dozen Barn Owls” had been seen in Berthierville around 1931, and he was referred to Créte for further details. Créte indicated that a pair, discovered by A. Paquette, had nested in the church steeple, that from the four young he had received one in the flesh which was mounted by a local taxidermist, and that Paquette had given him a mounted specimen on 29 December 1931. Créte added that he kept the second specimen and gave the first to the museum of College de Sainte- Anne-de-la-Pocatiére. With his letter Crete included copies of the label and of the accession card of the specimen given to him by Paquette. According to the label the specimen (No. 31/16) is an adult male captured on 10 November 1931; on the accession card, however, Créte wrote that this bird was killed in November 1931, from a brood of four whose parents had settled in the church steeple. On 28 March 1946, Gaboriault interviewed Paquette and was told that high winds had been blowing for several days when about 20 Barn Owls appeared in the © Berthierville Islands in 1931. After a while they took refuge in the church steeple where Paquette killed eight birds. He mounted one which is at the Deaf and Dumb Institute, and sent the others to various museums. As the steeple floor was covered with droppings, it was evident several birds had been there for some time. Paquette could not give more definite details. Paquette’s recollection is confirmed by a post- humous article (Paquette 1961) which he read in French at the fiftieth meeting of the AOU at Quebec City in 1932 (see The Auk S50, p. 74). Relating his observations on birds of prey in captivity he said that about 15 Barn Owls had settled in Berthierville, and that he kept one bird for a week in order to study its postures before mounting it. Gaboriault’s list of Barn Owl specimens taken in 324 TABLE 1—Records of occurrence of the Barn Owl in Quebec THE CANADIAN FIELD-NATURALIST Vol. 93 Locality Date Remarks! References Longueuil October 1915 Captured by W. J. Low V. Gaboriault L’Assomption November 1926 Captured Auk 64, p. 631 Longueuil 1 November 1931 Deaf and Dumb Institute Museum V. Gaboriault November 1931 10 November 1931 15 June 1936 Berthierville Berthierville St.Félix-de-Kingsey Beauharnois 12 May 1944 Captured by V. Gaboriault Thetford Mines 3 September 1944 Captured in a barn Saint-Hubert 1961 One “pair” seen in summer Giffard 18, 24 June 1963 Sight records 14 April 1967 30 May 1968 20 December 1970 22 December 1970 Berthierville Sight record Mont-Carmel Cap Tourmente Cap Tourmente Found dying Sight record Rigaud 12 June 1971 Captured in a barn Cap Tourmente 8 May 1974 Found dead Near Montreal Spring 1975 Shot Huntingdon Fall 1975 Killed by a car Saint-Coeur-de-Marie 1975 Captured in a barn Masson July 1977 condition) Captured by J. Desjardins At least 3 captured by A. Paquette Captured in a barn Captured in a barn Captured (female in breeding V. Gaboriault V. Gaboriault PSNHC? Annual Report 1938, p. 161 V. Gaboriault Auk 64, p. 631 PQSPB3 Annual Report 1961, p. 25 Bulletin ornithologique 8(4), p. 3 Bulletin ornithologique 12, p. 19 Bulletin ornithologique 13, p. 37 Bulletin ornithologique 15, p. 95 Bulletin ornithologique 15, p. 95 Bulletin ornithologique 16, p. 58 Bulletin ornithologique 19, p. 24 American Birds 29, p. 830 American Birds 30, p. 696 Bulletin ornithologique 21, p. 54 Henri Ouellet (pers. comm.) ‘One individual only unless otherwise stated. 2The Provancher Society of Natural History of Canada. 3The Province of Quebec Society for the Protection of Birds. Quebec shows only three extant specimens from the birds killed in Berthierville by Paquette in 1931: one of unstated age and sex in a Berthierville school museum, another of unstated age and sex in the museum of College de Sainte-Anne-de-la-Pocatiére, and an adult in the museum of the Deaf and Dumb Institute (No. 31/16), “from a group of 4 or 5 which had taken refuge in the church steeple.” This annotation shows clearly that Gaboriault was con- vinced that the species had not bred. None of the extant specimens was a flightless young, and Paquette never said that the birds had nested. Paquette may have exaggerated the number of birds seen, and Créte perhaps assumed that only breeding could explain such a number of birds. What appears to be the correct evaluation of the record is found in Gaboriault’s notes. On a map of Quebec he had marked records of occurrence with different keyed symbols according to their seasonal status. The symbol indicating the Berthierville record on the map of the Barn Owl corresponds to a fall visitor. The records of occurrence to date (Table 1) show only a pattern of casual vagrancy in the St. Lawrence lowlands, with exceptional records eastward to Kamouraska County (Mont-Carmel) and northward in the Lake St. John area (Saint-Coeur-de-Marie). The most recent record though suggestive, did not provide conclusive evidence that the Barn Owls actually bred within the limits of the province. A bird in breeding condition was captured in a field near Masson, Papineau County, only 1.5 km north across the Ottawa River from the Ontario border; it was seized by game wardens | or 2d later, on21 July 1977. The specimen, now preserved in the National Mu- seum of Natural Sciences of Canada (No. 67191), isa female having a refeathered brood patch; its reproductive tract contained 10 empty follicles and several ova, the largest measuring 16 mm (Henri Ouellet, personnal communication). I thank Wilfrid Gaboriault who kindly lent me the ornithological notes of his late brother. I also thank Raymond Cayouette who allowed me to examine his correspondence with F. Créte. Henri Ouellet reviewed the manuscript. Victor Gaboriault died before he could write a planned account on the distribution of birds in Quebec; for this important contribution, he should be considered as the author of this note. Literature Cited Cayouette, R. 1947. Barn Owl in Quebec. Auk 64: 631. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin 203. Paquette, A. 1961. J’ai gardé en captivité des oiseaux rapaces. Jeune Naturaliste 11: 129-132. Received 20 November 1978 Accepted 24 February 1979 1979 NOTES 325 Communal Roosting of Song Sparrows under Snowbank MARTIN K. MCNICHOLL Beak Consultants Ltd., 3530 11A St. N.E., Calgary, Alberta T2E 6M7 MeNicholl, Martin K. 1979. Communal roosting of Song Sparrows under snowbank. Canadian Field-Naturalist 93(3): 325-326. Fifteen Song Sparrows ( Me/ospiza melodia) were found roosting under an ice-encrusted snowbank on Prince Edward Island. The birds appeared to be keeping warm by a combination of roosting under snow and reducing individual distance. Key Words: Individual distance, Prince Edward Island, Song Sparrow, sub-nivean environment, winter survival. At 06:55 on 7 April 1977 I was about to cross a stretch of snowbank 50 m long by | m wide in a ditch in front of a farmhouse near French River, Prince Edward Island, when 15 Song Sparrows ( Melospiza melodia) emerged in rapid succession from a hole approximately 4 cm in diameter in the bank. Exam- ination of the hole showed that it led via a short tunnel (approximately 10 cm long) to a small cavity, ap- proximately 18-20 cm long by 11-14 cm wide, and 8 cm high, with no other exit. Previous melt had caused the snowbank to become encrusted with ice. The temperature was —2°C at the time, but with strong north winds felt much colder. (I was uncomfortably cold in an “Eskimo-style parka in which I am normally warm in -20°C weather in the prairie provinces.) Nice (1943, p. 106) stated that Song Sparrows roost in “weeds, hedges and small evergreens,” and I have found no references to their roosting in holes or under snowbanks. Sub-niveal roosting has been recorded for Common Redpolls (Carduelis flammea) (Cade 1953: Novikov 1972), Dark-eyed Juncos (Junco hyemalis) (Linsdale 1928), Tree Sparrows (Spizella arborea) (Thompson 1934), Snow Buntings ( Plectro- phenax nivalis) (Bagg 1943), and several Eurasian passerines (Novikov 1972). In a recent summary of several earlier accounts of passerines roosting under the snow in Finland and the Soviet Union, Novikov (1972) concluded that several species which regularly winter in snow-covered areas frequently roost under snow. Unlike most of the above-mentioned species, Song Sparrows do not usually winter in snow-covered areas, although many return to northern nesting areas before all snow is gone, and there are many records of individual birds wintering in northern areas. Kendeigh (1961) demonstrated the energetic ad- vantage to birds of roosting in cavities, and the well known insulative properties of snow (Formozov 1946; Elsner and Pruitt 1959) would enhance such an advantage. This insulative ability of snow, however, is poorest when the snow has been penetrated by water or converted to ice, as in the present case. Thus, even under the snowbank, the sparrows may have been under considerable cold stress, a condition known to induce breakdown of individual distance, resulting in clumping together (Beal 1978). Although such crowd- ing together is reported for several colonial or flocking species in cold weather (e.g., Grubb 1973; Meservey and Kraus 1976), | am not aware of reports of such behavior in a species such as the Song Sparrow which shows intraspecific intolerance all year (see Nice 1943). As territorial behavior tends to be particularly strong shortly after return to the nesting area in spring, this observation of communal roosting ap- pears particularly remarkable. Thus, the sparrows I observed appear to have combined the strategies of roosting under snow and clumping together to keep warm. In such a territorial species, such a breakdown of individual distance likely occurs only rarely. I thank Anthony J. Erskine, William O. Pruitt, Jr., and two anonymous referees for helpful comments on the manuscript. Literature Cited Bagg, A.M. 1943. Snow Buntings burrowing into snow- drifts. Auk 60: 445. Beal, K.G. 1978. Temperature-dependent reduction of individual distance in captive House Sparrows. Auk 95: 195-196. Cade, T. J. 1953. Sub-nival [sic] feeding of the Redpoll in interior Alaska: a possible adaptation to the northern winter. Condor 55: 43-44. Elsner, R. W. and W. O. Pruitt, Jr. 1959. Some structural and thermal characteristics of snow shelters. Arctic 12: 20-27. Formozoy, A. N. 1946. Snow cover as an integral factor of the environment and its importance in the ecology of mammals and birds. Boreal Institute, University of Alberta, Edmonton, Occasional Publication No. 1. (Translated by W. Prychodko and W. O. Pruitt, Jr.) Grubb, T. C., Jr. 1973. Absence of “individual distance” in the Tree Swallow during adverse weather. Auk 90: 432-433. 326 Kendeigh, S. C. 1961. Energy of birds conserved by roost- ing in cavities. Wilson Bulletin 73: 140-147. Lindsdale, J. M. 1928. Birds of a limited area in eastern Kansas. University of Kansas Science Bulletin 73: 517- 626. Meservey, W.R. and G. F. Kraus. 1976. Absence of “in- dividual distance” in three swallow species. Auk 93: 177-178. Nice, M. M. 1943. Studies in the life history of the Song Sparrow. Volume II. Transactions of the Linnaean THE CANADIAN FIELD-NATURALIST Vol. 93 Society of New York 6. Novikov, G. A. 1972. The use of under-snow refuges among small birds of the sparrow family. Aquilo Ser Zoologica 13: 95-97. Thompson, S. L. 1934. Unusual roosting of Tree Sparrow. Canadian Field-Naturalist 48: 142-143. Received 2 January 1979 Accepted 6 March 1979 Status of Eastern White Cedar, Thuja occidentalis, in Western Nova Scotia GORDON S. RINGIUS Department of Botany, University of Alberta, Edmonton 16G 2E9 Ringius, Gordon S. 1979. Status of Eastern White Cedar, Thuja occidentalis in western Nova Scotia. Canadian Field- Naturalist 93(3): 326-328. The occurrence of Eastern White Cedar, Thuja occidentalis, in western Nova Scotia, in doubt since the initial report published in 1877, is confirmed. A brief account of four of the five known stands in Cumberland County is presented and a map showing the distribution of the species in Nova Scotia is included. Key Words: Thuja occidentalis, rare, Cumberland County, distribution. The earliest record of Eastern White Cedar, Thuja occidentalis, from western (= Cumberland County) Nova Scotia was published in 1877 by Lindsay. Confirmation of Lindsay’s report has not been forth- coming, however, and as a result, subsequent authors (Table 1) have either cited Lindsay (Bentley and Smith 1962), included it without explanation (Maher et al. 1978: Saunders 1970), suggested that the species may be present (Roland 1947), or have excluded it (Loucks 1962; Roland and Smith 1969). The purpose of this note is to confirm the presence of Thuja occidentalis in western Nova Scotia and to indicate where the known stands are located. On | April 1977, I observed a small stand of Thuja occidentalis along the Trans-Canada highway about 5.8km E of the River Phillip - Oxford junction (45°43’N, 63°50’W). The stand was located near the base of a west-facing slope adjacent to a Speckled Alder (Alnus rugosa) — Tamarack (Larix laricina) swamp and consisted of less than 50 mature trees scattered over an area of approximately 5 ha. The trees were sparsely branched, many had single and double bifurcate stems, and most showed reddish brown discoloration of the lower and _ peripheral foliage. The presence of numerous previous years’ cones indicated that the stand was producing seed. Because snow still covered much of the ground, however, no attempt was made to locate seedlings or saplings. A voucher specimen was collected and has been placed in the Smith Herbarium at Acadia University (Ringius 796). To determine whether other stands of Thuja occi- dentalis have been located in western Nova Scotia, I examined the collections at ACAD, DAL, DAO, and NSPM (acronyms from Holmgren and Keuken 1974). In addition, I contacted several people who are involved with forestry in Nova Scotia. As a result of these investigations, I discovered four additional stands, and a previous unpublished account of the stand described above. The locations of these stands are shown in Figure |. On file at NSPM are the following two reports taken from the Chignecto Peninsula Inventory: Eatonville near Apple River, Cumberland Co., a cedar swamp less than one acre [0.4 ha] in size. Sandy MacGregor, Forestry Superintendent, Scott Paper Company, New Glasgow. A few miles southeast of Oxford, Cumber- land Co., scattered cedar trees located along the trans-Canada [highway] in this area. Also a few cedar swamps of about 5 or 6 acres [2.0 — 2.4 ha] each between the 1979 NOTES 307 lec aa a5 47° 46° ‘\ —~ 4) D S Gig 6 ZA 4 » ire i a \iror : % .@ K \ ) ) 45 et ‘ ° Fp ean ~ X o y Ss Thuja occidentalis zh ‘ ie f LA @ Specimens Examined =| 44° Vv, f ‘ 44° # © Other Reports al SCALE L ry 80 Kilometers ge 64° 62° 60° FIGURE 1. The distribution of Eastern White Cedar, Thuja occidentalis, in Nova Scotia, based on specimens examined at ACAD, DAL, DAO, and NSPM, and on reports obtained through correspondence. A — Annapolis County, C — Cumberland County, D — Digby County, K — Kings County, Y — Yarmouth County. highway and the railroad. Cedar, which used to be very common in Cumberland Co., is now rare and there are only a few cedar stands remaining. Department of Lands and Forests, Oxford. S. MacGregor (personal communication 1977) stated that “the population of cedar at Apple River is native as far as I know. There are only a few stunted trees growing ona very tiny wet area.” Similarly, J. H. Beattie, Supervisor Forest Resources, Department of Lands and Forests, Oxford (personal communication 1977) stated the “cedar southeast of Oxford is native.” He also said there are small stands at Lorneville, Pugwash Junction, and near Joggins. All three sites are in Cumberland County. The Lorneville stand was at one time about 4 ha in size but has been seriously reduced over the years. As a result of cutting in the summer of 1977 only a few small trees of this stand remain. On his suggestion 1 contacted Harold V. Hatfield, Supervisor Forest Resources, District of Cumberland West, Parrsboro, who was acquainted with the Joggins stand. He provided information on its location and referred to it as a “small amount of wild cedar. [The trees] are young [and] scattered among spruce and hardwood on an old burn.” It is apparent that the stands of Thuja occidentalis in Cumberland County are small, scattered, and rare. In these respects they are similar to those in Anna- polis, Digby, Kings, and Yarmouth counties in south- western Nova Scotia where the species has been known to occur since about 1801 (Table 1). In an ecological study of the southwestern stands, Bentley | and Smith (1962) found that although reproduction was ample as indicated by seedling density, seedling mortality was apparently high because few young trees were present in any of the stands. They concluded that competition from other tree species was the limiting factor controlling the distribution of Thuja occidentalis in Nova Scotia. It should be mentioned that the species grows well enough when planted; as an ornamental it is found throughout the province with the possible exception of Cape Breton Island. I thank S. P. Vander Kloet of Acadia University, Wolfville, Nova Scotia, for reviewing an early draft of this note and offering helpful suggestions. Informa- tion supplied by J. H. Beattie, H. V. Hatfield, and S. MacGregor, and access to the various herbaria are appreciated and gratefully acknowledged. 328 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Previous reports of Eastern White Cedar, Thuja occidentalis, in Nova Scotia Author Date Locality Smith! ca. 1801 Annapolis Valley (not plentiful) Lindsay 1877 Cumberland County Macoun 1886 Swamp within 3 mi[4.8 km] of Annapolis[ Annapolis County], but said to grow on the Bay of Fundy near [Annapolis]; rare in Nova Scotia Fernald 192] Digby County, Yarmouth County (lakesides) Roland 1947 Annapolis Valley, Digby, Yarmouth counties; possible existence on the isthmus between Nova Scotia and New Brunswick [Cumberland County] Bentley and Smith 1962 Annapolis, Digby, Kings counties; reported from Cumberland County [cite Lindsay 1877] Loucks 1962 Wentworth Lake District [= portions of Digby and Yarmouth counties]; local Roland and Smith 1969 Annapolis Valley [includes Kings County], Annapolis, Digby counties Saunders 1970 Annapolis Valley, Digby, Yarmouth counties; slight ‘spillover’ from New Brunswick on the Chignecto Isthmus [Cumberland County] Maher et al. 1978 Annapolis, Cumberland, Digby, Kings, Yarmouth counties 'Cited by Gorham (1955). Literature Cited Bentley, P. A. and E. C. Smith. 1962. A study of northern white cedar and jack pine in Nova Scotia. Proceedings of the Nova Scotian Institute of Science 24: 376-398. Fernald, M. L. 1921. The Gray Herbarium expedition to Nova Scotia. Rhodora 23: 89-111. Gorham, E. 1955. Titus Smith, a pioneer of plant ecology in North America. Ecology 36: 116-123. Holmgren, P. A. and W. Keuken. 1974. Index herbar- iorum. Part I. The herbaria of the world. 6th edition. Oosthoek, Scheltema, and Holkema, Utrecht, Nether- lands. Lindsay, A. W. H. 1877. A catalogue of the flora of Nova Scotia. Proceedings of the Nova Scotian Institute of Science 4: 184-222. Loucks, O. L. 1962. A forest classification for the maritime provinces. Proceedings of the Nova Scotian Institute of Science 25: 85-167. Macoun, J. 1886. Catalogue of Canadian plants. Part III. Apetalae. Dawson Brothers, Montreal. Maher, R. V., D. J. White, G. W. Argus, and P. A. Keddy. 1978. The rare vascular plants of Nova Scotia. Syllogeus Series Number 18. 37 pp. Roland, A. E. 1947. The flora of Nova Scotia. Proceed- ings of the Nova Scotian Institute of Science 21: 97-644. Roland, A. E. and E. C. Smith. 1969. The flora of Nova Scotia. Nova Scotia Museum, Halifax. 743 pp. Saunders, G. L. 1970. Trees of Nova Scotia. Department of Lands and Forests, Nova Scotia. 102 pp. Received 21 November 1978 Accepted 7 April 1979 1979 NOTES 329 Snowy Egret in the Northwest Territories RICHARD W. QUINLAN Canadian Wildlife Service, #1000, 9942 - 108 Street, Edmonton, Alberta T5K 2J5 Quinlan, Richard W. 1979. Snowy Egret in the Northwest Territories. Canadian Field-Naturalist 93(3): 329. Key Words: Snowy Egret, Northwest Territories, first record. On 23 June 1977, a Snowy Egret ( Egretta thula) was seen near Fort Simpson (61°53’N, 121°25’W). This is the first recorded sighting in the Northwest Territories and the most northerly in North America. This species breeds from the central United States through the West Indies and Central America to Chile and Argentina. It is a rare non-breeding wanderer to Alberta, Saskatchewan, and British Columbia (W. E. Godfrey, 1966. The birds of Canada. National Museum of Canada Bulletin 203). There are six records of Snowy Egrets in Alberta, all in May and June (C.S. Houston and M.I. Houston, 1976. American Birds 29: 74-77; W. R. Salt and J. R. Salt, 1976. The birds of Alberta. Hurtig, Edmonton); the most northerly was at Sandy Lake (52°47’N, 111°00’W) in May 1909. The most northerly of four Saskatche- wan sightings was at Saskatoon (52°07’N, 106°38’W) on 15 May 1977(E. M. Serr, 1977. American Birds 31: 1013-1016). Near Juneau, Alaska (58°20’N, 134°20’W), a Snowy Egret was observed 18-24 May 1957 (I. N. Gabrielson and F. C. Lincoln, 1959. The birds of Alaska. Stackpole Co., Harrisburg, Pa.) The bird reported here was observed on the downstream (northwest) end of Fort Simpson Island in the Mackenzie River. It was first sighted on 23 June by Richard Quinlan and Yvonne Desilets, who watched it for 45 min from a distance of approxi- mately 50 m. Quinlan, with James Steele and (once) Bruce Reilly, also saw the egret on each of the next 4d. All observations except the last (1 km down- stream on a small island) were at the original location. A super-8 movie film taken on 25 June was viewed by Henri Ouellet of the National Museum of Natural Sciences, who verified the identification. Bernard Gollop provided a number of additional records from Alberta and Saskatchewan. Received 11 December 1978 Accepted 19 March 1979 Letters Is Biology Unknown? The recent Guest Editorial by Yorke Edwards, “Biology — The Unknown Science?” (Canadian Field-Naturalist 93(1): 6-9; 1979), invites a reply and several counter-proposals. Of all the sciences, Biology is probably the best known today. It has its Suzuki, and the media are full of “gee-whiz” articles on genetic engineering, tissue culture, test-tube babies, and Cinderella crops. What is missing, according to Edwards, is the synthesizing scientist who will put all these marvelous bits and pieces together “so that people may understand their world.” There are a couple of serious problems with this diagnosis. Implicit is the assumption that science is engaged in discovering reality, not making it. But suppose that at least some science (and certainly all the technology that science nurtures) is motivated not so much by a desire to understand as to control and change? Then people may rightly question whether scientists are all admirable, and whether every old kind of science is A-OK. They might, for example, ask what exactly biologists are up to when they study the deep-diving abilities of seals, or the cold resistance of wolves’ feet. “Why,” your innocent scientist will answer, “we mean to help our fellow men to explore safely the ocean floor for mineral treasures, and warmly to run around on the snow while searching for good Canadian oil!” “But surely,’ the layman responds, “you investigate these things with an enrich- ing sense of wonder and awe, and this you will transmit to me so that I too may understand and marvel?” “Will you please go away and don’t bother me,” comes the reply, “I’m preparing a scientific article (on which my next promotion and merit increase depend) for one of the NRC journals and tomorrow I’m off to Ottawa as a consultant to the Department of National Defence.” A related questionable assumption is that the synthesis of knowledge needed to understand the world will come from a change in attitude of those who know the facts (the analytical scientists) rather than from a newly constituted science that sets as its goal the sympathetic understanding of world eco- systems. After all, how realistic is it to expect nuclear physicists to go to bat for Ecological Reserves, or mining geologists to take up the cudgels for a decentralized solar society? Their training is wrong, and so is the training of most biologists. It is not that Biology is unknown but that the proper subject is unknown. Finally, we hear much today about morality in science, meaning that no matter what is buzzing around in their Pandora’s box, scientists should carefully explain to the public the undoubted advan- tages of taking off the lid. Such morality is largely self- serving. There needs to be, as Aldo Leopold said long ago, a land ethic, acommitment to something external to the human race, something equally if not more important. The main problem with science, and the reason for the lack of Earthcare, is the narcissism of the human race. J. STAN ROWE 9 March 1977 Department of Plant Ecology, University of Saskatchewan, Saskatoon, Saskatchewan 2N4 0WO Never one to be uninterested in reactions to what I write, I gladly accept this opportunity to comment on Dr. Rowe’s thoughtful comments, and I do so briefly. ] read them with complete sympathy and agreement, for most of his thoughts are not far from those I hold, or have held. Inevitably the world we see is the part of it that we are in, and my current views must therefore be colored by a long association with government and primary industry. I can only say in addition to what I have said, that I find my part of the world quite frightening because of the values used in deciding destructions of the living parts of Earth as well as of the mineral bases for that life. As Dr. Rowe probably knows, my copy of Leopold is as old and as well thumbed as his. In sucha brotherhood as this, there can be only relatively minor differences in views of land and its life. YORKE EDWARDS 30 March 1979 British Columbia Provincial Museum, Victoria, British Columbia V8V 1X4 330 Tribute to HOYES LLOYD, 1888-1978 Hoyes Lloyd, who died in his 89th year in Ottawa on 21 January 1978, was the central figure in Canadian nature conservation activities during the critical 25 years from 1918 to 1943. During that period, nature conservation grew from a personal interest in the minds of a few people toa responsibility taken seriously by government. The first person given a professional role in nature conservation by the Canadian Government, Lloyd carefully expanded and consolidated activities aimed at the conservation of migratory birds and of wildlife in the national parks and the Territories, laying the groundwork for the establishment of the Canadian Wildlife Service, which was formed (as the Dominion Wildlife Service) in 1947. His accomplishments rested upon his thorough knowledge of wildlife, his keen sense of what was possible and practical and, perhaps most importantly, his ability to pick a path through political and bureaucratic tangles with calmness, diligence, and humor. For anyone of Hoyes Lloyd’s generation, there was scant opportunity for a professional career in wildlife conservation. Almost all of those who broke into the field before 1930 were first obliged to seek means of livelihood less closely matched to their real interests. Lloyd, born in Hamilton in 1888, and educated at Harbord Collegiate Institute in Toronto and at the University of Toronto, found his first full-time employment in applied chemistry. From 1909 to 1911, while working for his Master’s degree in Chemistry, he served as an assistant in the department, and during 1911 and 1912 he was employed by a manufacturer of chemicals. He entered public service in 1912 when he was appointed chemist in charge of milk control for the City of Toronto, and a few years later he played an important role in bringing about compulsory pas- teurization of the city’s milk supply. Not until 1918 was he able to give his full time to conservation. But Lloyd was an active student of natural history long before that. Just after the turn of the century, he began to keep records of the occurrence and behavior of birds and to collect plants and birds. During the next few years he roamed the woods, fields, and marshes around Toronto, often with young friends who shared his interests, building up his knowledge of nature and through his collections establishing per- manent records of the occurrence of birds and plants. In 1903 he happened ona copy of The Auk ina book store and learned that bird study was a serious thing pursued by serious persons. In 1904, 1905, and 1906 he won prizes for plant collections entered in the Canadian National Exhibition. In the summer of 1909, which he later described as one of the most exciting times of his life, he hiked and paddled the wilderness as a forest ranger and deputy game warden on the Temagami Forest Reserve. In the same year he won a gold medal at the Canadian National Exhibi- tion for his collection of bird skins. Full-time employment as a chemist in the field of public health from 1917 to 1918 did not diminish his activities as a field naturalist. During that period he continued to observe and collect. In 1916 he became an Associate Member of the American Ornithologists’ Union and in 1917 he wrote “Ontario Bird Notes” his first paper for The Auk. Lloyd’s opportunity to work in the field he loved came as attitudes to nature and the use of natural resources began to change. Early in the century, a few North Americans began to see the folly of profligate use of resources. In Canada, the Commission on Conservation issued its series of thoughtful reports in the years 1910-1913. Among its recommendations was one that called for an agreement between Canada and the United States for the protection of migratory birds. Discussions and negotiations proceeded, lead- ing to the signature of the Migratory Birds Con- vention in 1917, and the enactment of enabling legislation in Canada, the Migratory Birds Conven- tion Act in 1918. Within the year Hoyes Lloyd wona competition to head the one-man Migratory Birds Unit in the Department of the Interior. Lloyd remained with the Department of the Interior and its successor, the Department of Mines and Resources, for 25 years, in charge of a wildlife unit that was gradually given broader responsibilities but was always too small to meet its responsibilities to his satisfaction. The early years were particularly diffi- cult: the task immense, the resources infinitesimal. At first Lloyd served as policy-maker, field man, chief — clerk, and everything in between, but within three years he had a clerical assistant and three migratory bird officers to help him cover all of Canada. An urgent requirement then as now was to tell people about the Migratory Birds Convention Act, and the North West Game Act, for the administration of which he was also responsible, and to persuade them that the Acts and Regulations must be observed. This meant writing letters and leaflets; meeting sportsmen and naturalists formally and informally; advising teachers, wardens, police and magistrates; and con- ferring with officials of other government depart- ments, provincial governments and the Government of the United States. It also meant travelling from one end of Canada to the other to familiarize himself with conservation problems in the field, of which there were many, and frequent meetings with his superiors 331 332 in Ottawa to seek more support for the work that had to be done. Progress was slow but steady. Migratory bird sanctuaries were established; the Migratory Bird Regulations were improved and their enforcement became more effective; cooperation with the United States in studying the migrations and populations of migratory birds was initiated and developed. In 1922, Hoyes Lloyd organized and served as secretary to the first Federal-Provincial Wildlife Conference, an institution that soon came to meet annually and enabled effective exchange of informa- tion and views among the wildlife authorities of all the provinces and the federal government. The “Federal- Provincials,” which are still held each year, have been highly effective in the development of wildlife con- servation in Canada. At that first conference Lloyd no doubt displayed the capacity for unobtrusive but effective group leadership that he was later to employ so often in so many different settings. He was President of The Ottawa Field-Naturalists’ Club from 1923 to 1925; President of the International Associa- tion of Game, Fish and Conservation Commissioners in 1929-30; Chairman of the Canadian Section of the International Council for Bird Preservation from 1927 to 1954 and Vice-President of the Council itself from 1938 to 1950. He regularly attended and, as the senior Canadian official, was often a key-note speaker at the annual meetings first known as the North American Game Conference (he was chairman in 1935) and now as the North American Wildlife and Natural Resources Conferences. He organized and chaired innumerable less formal meetings in many parts of Canada and in the United States. By 1943 when Lloyd retired after 25 years of public service, wildlife protection in Canada was widely if not universally accepted and wildlife management asa more broadly based and purposeful discipline was beginning to emerge. Much of the credit for that development is his. Retirement gave Hoyes Lloyd the opportunity to extend his professional interest in birds. This had not been neglected in his years in the Civil Service; he was increasingly active in the American Ornithologists’ Union during the 1920s and 1930s and his bird collection slowly continued to grow. In 1942 he became a Vice-President of the Unionand he served as its President from 1945 to 1948. His participation in the work of the International Council for Bird Preservation became more active in his post-retire- ment years during which he attended and contributed actively to each of its quadrennial conferences until 1972. All this work was, of course, voluntary; though he often represented the Canadian Government abroad in places suchas Helsinki, Caracas, Bulawayo, and Tokyo, he paid his own way and he did so without complaint. Old associates in the American Ornitholo- THE CANADIAN FIELD-NATURALIST Vol. 93 gists’ Union and the Wildlife Society continued to welcome him to their meetings which he attended until 1973. Even after travel became less easy for him Lloyd kept up an active correspondence with his many friends and received occasional contributions to his collections. His collections of plants, birds, and mammals along with his field journals covering a period of 71 years and his extensive library, are now in the care of the Royal Ontario Museum. Hoyes Lloyd wrote over 120 papers and notes for publication. Of these, over 100 were contributions to The Canadian Field-Naturalist; he did much to develop its reputation as a worthy source of informa- tion about Canadian wildlife. In 1944 he published in our journal a 32-page annotated list “The Birds of Ottawa,” which consolidated and added to notes on Ottawa birds published in 1923, 1924, 1925, 1932, 1936, and 1937. In 1945, 1949, and 1951 he again added to his regional list. This work provided a major reference for the bird group of The Ottawa Field- Naturalists’ Club and contributed occurrence records to many later works on the distribution of North American birds. Lloyd’s bibliography includes papers dealing with many other records of bird distribution in other parts of Canada as well as records of migrations derived from the recoveries of banded birds and accounts of the introduction of Ring-necked Pheasant and Hungarian Partridge to various parts of Canada. The only Canadian to be the recipient of the two premier honors in the wildlife field in North America was Hoyes Lloyd. He was presented with the Leopold Award by the Wildlife Society in 1956 and the Seth Gordon Award by the International Association of Fish, Game and Conservation Commissioners in 1974. He was made an Honorary Member of both those organizations and he was elected a fellow of the American Ornithologists’ Union and of the New York Zoological Society. He was an Honorary life member of the Quebec Zoological Society, the Outdoor Writers Association of America, and The Ottawa Field-Naturalists’ Club. Lloyd was married in 1913 to Wilmot Lockwood of Memphis, Tennessee. His two daughters and his son, and later his grandchildren and great-grandchildren all lived in Ottawa, and with his wife and himself formed a close-knit group that was rarely long separated. Their home in Rockcliffe Park, selected because he heard a Wood Thrush sing when he first inspected the spacious grounds, provided an ideal site for frequent family parties. A warm and friendly person, unfailingly courteous, Hoyes Lloyd was always generous with his time and knowledge, particularly with younger people. It was often my personal pleasure to benefit from his counsel HOyYES LLOYD, 1888-1978 333 1979 TTT ATEN jp L/h Hi WOT Hoyes Lloyd On the occasion of his receipt of the 1974 Seth Gordon Award, the highest honor given by the International Association of Game, Fish and Conservation Commissioners, for his outstanding work in wildlife conservation. 334 THE CANADIAN FIELD-NATURALIST and encouragement. While he knew that the continu- ing progress of nature conservation would be well served by the help he gave to those who would take up his cause, his personal relationships went far beyond such mere calculation. He truly loved his fellow man. DAVID A. MUNRO IUCN (International Union for Conservation of Nature and Natural Resources), 1110 Morges, Switzerland. Publications of Hoyes Lloyd 1917 1918 1918 1919 1919 1919 192] 1921 1921 192] 1922 1922 1922 1922 1922 1922 1923 1923 1923 1923 1923 1923 1924 1924 1924 Ontario bird notes. Auk 34: 457-460. Spring in Birdland. Youth and Services. C.S.E.T. Department of Ontario, Volume 20(32). The extraction of fat from bird-skins. 164-169. The Ornithological collector and the law. Canadian Field-Naturalist 33: 93-94. Permits to collect birds for scientific purposes in Canada. Auk 36: 621-623. Protection of bird neighbours. Leaflet. March. Bird notes by the way in the Maritime Provinces. Canadian Field-Naturalist 35: 11-13. The Town of Yarmouth, N.S. buys a bird sanctuary. Canadian Field-Naturalist 35: 35-36. An aquatic habit of the pigeons. Canadian Field- Naturalist 35: 98-99. Attracting birds to the prairie home. Canadian Forestry Magazine 17: 76-78. Some of Captain Henry Toke Munn’s observations on birds of Baffin Island and vicinity. Canadian Field- Naturalist 36: 49-50. Larvae of Phormia chrysorrhea Macq. found upon nestling Bluebirds. Canadian Field-Naturalist 36: 116. Review of “Some useful Australian birds.” Canadian Field-Naturalist 36: 118-119. A recent breeding record of the Trumpeter Swan in Alberta. Canadian Field-Naturalist 36: 143-144. Popular names for birds. Auk 39: 144. Some northern records of the Turkey Vulture. Canadian Field-Naturalist 36: 178-179. Protecting nesting sites. Auk 40: 571. Observations on the wintering flocks of Canada Geese in Nova Scotia. Canadian Field-Naturalist 37: 26-28. Brown Creeper the victim of strange accident. Canadian Field-Naturalist 37: 75. The birds of Ottawa. Canadian Field-Naturalist 37: 101-105; 125-127; 151-156. The eating of birds by chipmunks. Canadian Field- Naturalist 37: 118. Loss of Whistling Swans at Niagara Falls. Canadian Field-Naturalist 37: 138. The birds of Ottawa. Revised to March 20, 1923. Canadian Field-Naturalist 38: 10-16. Christmas bird census 1923. Canadian Naturalist 38: 17. An unusual sight description of a bird verified. Canadian Field-Naturalist 38: 38. Auk 35: Dominion Parks Field- 1924 1924 1924 1924 1924 1925 1925 1925 1925 1925 1925 1925 1925 1925 1925 1926 1926 1926 1926 1926 1926 1926 1927 1927 1927 1927 1927 Vol. 93 A flock of Trumpeter Swans. Canadian Field- Naturalist 38: 57-58. Another mystery bird. Canadian Field-Naturalist 38: 76. Field Sparrow and Towhee at Christie Lake, Ontario. Canadian Field-Naturalist 38: 180. An old breeding record for the Herring Gull at Lac des Iles, Labelle Co., Quebec. Canadian Field-Naturalist 38: 193-194. A Fulmar at Arnprior, Ontario. Canadian Field- Naturalist 38: 194. The birds of Ottawa — Addenda to October 2, 1924. Canadian Field-Naturalist 39: 16-17. Mystery bands. Canadian Field-Naturalist 39: 43-44. The recording of acclimatization experiments. Canadian Field-Naturalist 39: 63-64. A second record of Scissor-tailed Flycatcher at York Factory, Manitoba. Canadian Field-Naturalist 39: 86. A juvenile Richardson’s Owl from Grand Manan, New Brunswick. Canadian Field-Naturalist 39: 113. A mystery band solved. Canadian Field-Naturalist 39: 114. The acclimatization of the Fox Squirrel at Pelee Island, Ontario. Canadian Field-Naturalist. 39: 138. The introduction of the Muskrat and the Rocky Mountain Goat on Vancouver Island. Canadian Field-Naturalist 39: 151-152. The introduction of the European Grey Partridge in Manitoba. Canadian Field-Naturalist. 39: 172. Christmas bird censuses in Canada. Canadian Field- Naturalist 39: 173. Christmas bird census Naturalist 40: 12-13. Yellow-bellied Sapsucker infested with tape worms. Canadian Field-Naturalist 40: 67. More mystery birds. Canadian Field-Naturalist 40: 88. Canadian Government motion picture films showing natural history subjects. Canadian Field-Naturalist 40: 100-102. Review of “Animal life in the Yosemite” by Joseph Grinnell and Tracy Irwin Storer. Canadian Field- Naturalist 40: 114-115. The 44th stated meeting of the American Orni- thologists’ Union, Ottawa, October 11-17, 1926. Canadian Field-Naturalist 40: 189-190. Review of “The relation of birds to woodlots in New York State’? — W.L. McAtee. Canadian Field- Naturalist 40: 194. A mystery solved. Bulletin of the Northeastern Bird- Banding Association 3: 17-18. Point Pelee should have a bird banding station. Canadian Field-Naturalist 41: 89-90. Identification of flies found infesting Bluebirds. Canadian Field-Naturalist 41: 111. Transfers of Elk for re-stocking. Canadian Field- Naturalist 41: 126-127. Additional comments regarding Mallards wintering at Buffalo Lake, Alberta, 1923-24. Canadian Field- Naturalist 41: 138. 1925. Canadian Field- IST) 1927 1927 1927 1927 1927 1928 1928 1928 1928 1928 1929 1929 1929 1930 1930 1931] 1932 1932 1933 1933 1933 1933 1934 1934 1935 1936 1936 1937 HoyYES LLOYD, 1888-1978 Another mystery band. Canadian Field-Naturalist 41: 139. Rocky Mountain Sheep transferred from Banff to Spence’s Bridge, B.C. Canadian Field-Naturalist 41: 140. A mystery band. Canadian Field-Naturalist 155-156. Work of the Canadian Section, The International Committee for Bird Protection. Canadian Field- Naturalist 41: 156. European Widgeon crosses from Iceland. Canadian Field-Naturalist 41: 171. Notes on the introduction of the pheasant in some of the provinces of Canada. Canadian Field-Naturalist 42: 143-145. The spread of the European Starling. Canadian Field- Naturalist 42: 178. Bird banding in Japan. Canadian Field-Naturalist 42: 178. Trans-Atlantic banded pigeon recovery. Canadian Field-Naturalist 42: 178. A method for cleaning large bird skins. Canadian Field-Naturalist 42: 207-208. Review of “Birds of Western Canada.” 2nd Edition. By P. A. Taverner. Canadian Field-Naturalist 43: 109. What surgeon to the birds treated this Pine Grosbeak? Canadian Field-Naturalist 43: 167-168. Who banded this Gull? Canadian Field-Naturalist 43: 189. Whistling Swan at Rigaud, Quebec. Canadian Field- Naturalist 44: 167. Henderson’s “The practical value of birds.” Auk 47: 302-304. Progress of conservation in Canada. Bird Banding II: 105-115. Starlings bathing in Naturalist 46: 22. The birds of Ottawa — Addenda. Canadian Field- Naturalist 46: 123-127, 162-166. An occurrence of the Clay-coloured Sparrow, Spizella pallida, in Renfrew County, Ontario. Canadian Field- Naturalist 47: 36. Wild Geese killed by lightning. Canadian Field- Naturalist 47: 72-73. Obituary Norman Criddie Field-Naturalist 47: 145-147. Review of “Game management” by Aldo Leopold. Canadian Field-Naturalist 47: 159-160. The conservation of wild bird life. Beaver, Outfit 265, June 1934. p. 22. An Ontario sand-fall. Canadian Field-Naturalist 48: 93-95. Review of “Sheep and bear trails” by John P. Holman. Canadian Field-Naturalist 49: 152. The late Norman Criddle’s record of the Snowshoe Rabbit (Lepus americanus) at Aweme, Manitoba. Canadian Field-Naturalist 50: 129-130. The birds of Ottawa — Addenda to February 28, 1935. Canadian Field-Naturalist 50: 143-144. A nest-hunting Broad-winged Hawk. Canadian Field- Naturalist 51: 12. 41: winter. Canadian Field- 1875-1933. Canadian 1937 1937 1937 1937 1938 1939 1939 1939 1939 1939 194] 1943 1943 1943 1944 1944 1945 1948 1949 1951 1951] 1951 1953 1953 1953 1954 1955 1956 335 Short-eared Owl (Asio flammeus flammeus) at Baker Lake, N.W.T. Canadian Field-Naturalist 51: 13. A wintering Towhee at Ottawa. Canadian Field- Naturalist 51: 77. Banded European Widgeon recovered in Prince Edward Island. Canadian Field-Naturalist 51: 77. Twenty year old Ferruginous Rough-legged Hawk. Canadian Field-Naturalist 51: 137. Some aspects of Canadian Bird-life. Canadian Geo- graphic Journal 16: 265-283. Review of “Canoe country” by Florence Page Jacques. Canadian Field-Naturalist 53: 13. Banded Kittiwake from Russia recovered in New- foundland. Canadian Field-Naturalist 53: 29-30. Andrew Halkett, naturalist, 1854-1937. Canadian Field-Naturalist 53: 31-32. Lorene Squire studies prairie birds. Canadian Geo- graphic Journal 18: 19-25. Deciphering numbers on much-worn bird bands. Canadian Field-Naturalist 53: 131. A Kittiwake Gull (Rissa tridactyla) banded near Murmansk, U.S.S.R., recovered in Newfoundland. Canadian Field-Naturalist 55: 138. Review of “Ornithologists of the United States Army Medical Corps: Thirty-six biographies” by Edgar Erskine Hume, Colonel. Canadian Field-Naturalist 57: 14. Review of “Wildlife of Idaho” by William M. Rush. Canadian Field-Naturalist 57: 14-15. Dr. T. Gilbert Pearson. Canadian Field-Naturalist 57: 141. Review of “Some aspects of Canadian birds” by A. L. Rand. Canadian Field-Naturalist 58: 18. The birds of Ottawa. Canadian Field-Naturalist 58: 143-175. Birds of Ottawa, 1944, an inadvertent omission. Canadian Field-Naturalist 59: 150. Review of “Wings in the wilderness” by Allan D. Cruickshank. Canadian Field-Naturalist 62: 166. Bird records for the Ottawa District. Canadian Field- Naturalist 63: 31-34. Duck Hawk at Blue Sea Lake, Quebec. Canadian . Field-Naturalist 65: 46. James Bay Sparrow at Ottawa. Canadian Field- Naturalist 65: 46. Recent Ottawa District bird records. Canadian Field- Naturalist 65: 46. Dead Golden Eagle at Perkins Canadian Field-Naturalist 67: 44. Review of “Proceedings of the Xth International Ornithological Congress, Uppsala, June 1950.” Canadian Field-Naturalist 67: 45-46. Wilson’s Petrel, Oceanites oceanicus oceanicus (Kuhl) at Lake Deschenes, Quebec. Canadian Field- Naturalist 67: 140. Clyde Louis Patch, 1887-1952. Canadian Field- Naturalist 68: 125-126. Mid-Atlantic migration of Long-tailed Jaegers and Terns. Canadian Field-Naturalist 69: 24. Ralph Emerson DeLury, 1881-1956, Canadian Field-Naturalist 70: 169-170. Mills, Quebec. obituary. 336 1959 1961 1961 1963 1963 1965 1920 1921 THE CANADIAN FIELD-NATURALIST William Henry Lanceley (1893-1958). Canadian Field-Naturalist 73: 183. A Manitoba occurrence of the Black-headed Gros- beak. Canadian Field-Naturalist 75: 109. Faith Fyles, artist-naturalist, 1875-1961. Canadian Field-Naturalist 75: 220. Review of “The cry of a bird” by Dorothy Yglesias. Canadian Field-Naturalist 77: 124. Catharacta skua Brunnich sighted in North Pacific. Canadian Field-Naturalist 77: 127. A review of Thoreau on birds. Naturalist 79: 261. J. H. Fleming and Hoyes Lloyd. Ontario bird notes. Auk 37: 429-439. Hoyes Lloyd and F.C. Hennessey. Annual 1920 Christmas bird census for Ottawa and Hull. Bird Lore 7p Gy). Canadian Field- 1929 1948 1950 Vol. 93 Hoyes Lloyd and Norman H. H. Lett. Obituary — George Rivers White, 1856-1927. Canadian Field- Naturalist 43: 103-104. Hoyes Lloyd and Robert G. Lanning. Observations on the birds of Renfrew County, Ontario. Canadian Field-Naturalist 62: 47-65. Hoyes Lloyd, Robert G. Lanning, and Sheldon Davis. Additional notes on the birds of Renfrew County, Ontario. Canadian Field-Naturalist 64: 140-142. Compiled by V. M. HUMPHREYS National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8. News and Comment Robert Lister Receives Award The fourth Loran L. Goulden Memorial Award for outstanding work on natural history within the province of Alberta was awarded to Robert Lister at the annual meeting of the Edmonton Natural History Club on 9 January 1979. Bob Lister, Honourary President of the Edmonton Bird Club, served as an indispensible assistant to the late Prof. William Rowan for many years, while the latter conducted his world-renowned experiments on influence of photo- period on migration; however, he is recognized in this award primarily for his long-term role in encouraging other naturalists in Alberta, through his newspaper column, his role in founding the Edmonton Bird Club, his social activities, and other means. His forthcoming book on the birds of Beaverhill Lake, based largely on Rowan’s work, marks a fitting occasion on which to recognize Bob through this award. Further details on his career and a list of his publications will appear ina future issue of the Alberta Naturalist. Martin K. McNicholl Chairman, Loran L. Goulden Memorial Award Selection Committee Notice of Change to the By-laws of The Ottawa Field-Naturalists’ Club A motion to change By-law 15 of The Ottawa Field-Naturalists’ Club was passed unanimously by the Council at the meeting of 12 February 1979. This By-law now reads as follows: 15. Membership Dues The schedule of dues shall be as follows: Individual $10.00 Family $12.00 Sustaining $25.00 Life Membership shall be granted upon payment of a single sum of $200.00. Article or Note? For The Canadian Field- Naturalist the assigning of an accepted manuscript to one category or the other is currently done solely on the basis of the manuscript’s length. We do not differentiate Articles from Notes on the basis of the quality or importance of the observations or research, although these criteria are stated or implied for some other scientific journals. Notes are run on one after the other so as not to be wasteful of available costly space because we are charged on a per page basis for every page that contains any printing whatsoever. Sometimes referees and editors request that authors condense their papers. The aim of the constructive advice is that the new information is presented as clearly and concisely as possible. Thus if an author focuses on the objectives of his or her study and the purpose of his or her report, Diana R. Laubitz, Recording Secretary the manuscript will be more succinct and its message more easily grasped. Beginning with the 1977 issues, I eliminated the main difference between the earlier printed versions of Articles and Notes by starting to use the same basic format for both. With this issue I have further eliminated the previous difference in type sizes. Therefore, shorter papers will be published as Notes and longer ones as Articles but they will both be in the same format. No differentiation will be made by the Editor nor should be construed by authors or readers regarding the relative significance of the Notes as compared to Articles in The Canadian Field-Natur- alist. LORRAINE C. SMITH, Editor 337) 338 THE CANADIAN FIELD-NATURALIST Voly93 Notice of Motion to Amend the Constitution of The Ottawa Field-Naturalists’ Club Notice of a motion to amend the Constitution of The Ottawa Field-Naturalists’ Club was received, in accordance with Article 23 of the Constitution, for presentation at the next Annual Business Meeting. It is proposed that: Article 11, now reading: “AUDITORS. Two Auditors shall be elected by open vote at the Annual Business Meeting. They shall examine the Treasurer’s accounts and certify as to their correctness.” be amended to read as follows: AUDITOR. An Auditor shall be elected by open vote at the Annual Business Meeting. The Auditor shall examine the Treasurer’s accounts and certify as to their correctness. Motion proposed by Courtney Gilliatt, seconded by William Cody. This motion is in accordance with a recommendation made at the last annual meeting. Diana R. Laubitz, Recording Secretary Request for Participants — International Shorebird Surveys 1979 A cooperative International Shorebird Survey scheme was organized by the Canadian Wildlife Service and the Manomet Bird Observatory in 1974 to obtain information on shorebird migration and to identify and document areas of major importance. Much very valuable information has come from contributors throughout eastern Canada and the USA, the Caribbean Islands, and Central and South America, and this is being used in assessing require- ments for the future protection and conservation of the birds and their habitat. Continuation of the project is planned for 1979. Any observer who will participate in regular counts of shorebirds during migration periods, as well as during the winter in shorebird wintering areas, 1s asked to contact one of the undersigned. Occasional counts from observers visit- ing shorebird areas on an irregular basis would also be most welcome. For areas in Canada: Dr. R. I. G. Morrison, Canadian Wildlife Service, 1725 Wooa- ward Drive, Ottawa, Ontario, Canada KIG3Z7. For areas in USA, Caribbean Islands, Central and South America: Brian A. Harrington, Manomet Bird Ob- servatory, Manomet, Massachusetts, U.S.A. 02345. Request for Information — Shorebird Color-marking In 1979 the Canadian Wildlife Service will be continuing a large-scale program of banding and color-marking shorebirds in James Bay. Since 1974 over 38500 shorebirds have been captured, resulting in more than 1700 ‘bird days’ of sightings of dyed birds ranging from eastern Canada to South America. Considerable valuable information on migration routes and strategies is being obtained and observers are again asked to watch for and report any color- dyed or color-banded shorebirds. Reports should include details of species (with age if possible), place, date, color-marks and, if possible, notes on the numbers of other shorebirds present. For color-dyed birds, please record the color and area of the bird that was dyed. For color bands and standard metal leg bands, please record which leg the bands were on, whether they were above or below the “knee,” the colors involved (yellow or light blue), and the relative position of the bands if more than one was on a leg (e.g., right lower leg, blue over metal, etc.). All reports will be acknowledged and should be sent to: Dr. R. I. G. Morrison, Canadian Wildlife Service, 1725 Woodward Drive, Ottawa, Ontario, Canada K1G 3Z7. The Douglas Pimlott Scholarship in Environmental Studies Innis College, University of Toronto, has estab- lished a scholarship fund in memory of Professor Douglas Pimlott, who for many years taught ecology, resource management, and environmental studies at the university. Among his many contributions, Dr. Pimlott established an Environmental Studies pro- gram, designed not for in-depth specialist training but in the spirit of a liberal education concerned with the environmental issues that face us all. The scholarship will be awarded to an under- graduate student who, like Doug Pimlott in his own life, combines high academic achievement with dedi- cated social involvement in environmental concerns. It is hoped that sufficient funds can be raised to endow the award. Contributions should be made payable to the Douglas Pimlott Fund, University of Toronto, and sent to: The Principal’s Office, Innis College, 2 Sussex Avenue, Toronto, Ontario MS5S 1J5. i Book Reviews ZOOLOGY Animals in Peril: a guide to the endangered species of Canada and the United States By D.C. Grainger. 1978. MacMillan, Toronto. 192 pp., illus. $14.95. This is another in a long line of books examining the plight of animals in North America. The title is misleading, however, because the book deals strictly with mammals. Furthermore, I do not see this book as “a guide to endangered animals of Canada and the United States.” It is a slightly less than erudite review of twenty mammals exploited by man at one time or another since his arrival in North America from Europe. In fact, some of the mammals considered are no longer endangered, or have never appeared on an endangered species list. The mammals considered include the Black-tailed Prairie Dog, Kaibab Squirrel, Spotted Bat, Fisher, Pine Marten, Black-footed Ferret, Sea Otter, Wolver- ine, Gray (timber) Wolf, Red Wolf, northern Kit Fox, Mountain Lion (eastern cougar), Grizzly Bear, Wood Bison, Muskox, Pronghorn Antelope, Florida Key Deer, Big Horn Sheep, Walrus, and the northern Manatee. Grainger has geared his book for young readers or novice biologists. His treatment of habits, habitat, and life cycle biology is very rudimentary and often suffers from too much brevity. The saving grace of this book is the author’s treatment of the cause and effect relationships The Great Arc of the Wild Sheep By James L. Clark. 1978. 4th Edition. University of Okla- homa Press, Norman. 247 pp. Paper US $11.50. “Many are inclined to think of the wild sheep as just scattered here and there around the world with little reason — but this is not so. Although they are rather thinly dispersed, the sheep are confined to a relatively narrow arc of mountain ranges which sweep across three continents.” The Great Arc of the Wild Sheep examines the wild sheep of the world. “All wild sheep of the world may be divided into five general groups: moufloniforms of the Mediterranean Islands and Middle Eastern countries, Argalis or Ammons of Central Asia, Asiatic Bighorns of Eastern Siberia, Dall’s of Alaska and Northern British Columbia, and North American Bighorns of western North America.” Each species is treated separately involved in the decline of some of these mammals. He examines the role man has played in their decline and documents efforts made to reverse the trend. For example, by 1940 there were only thirty Florida Key Deer left in existence, the result of overhunting. Interest in this small deer on behalf of the U.S. Fish and Wildlife Service, the Florida Game and Fresh Water Fish Commission and many conservative organizations, the National Audubon Society to name one, was instrumental in preventing its extinc- tion. Land was set aside as refuge and today there are approximately 400 animals in existence. To this end, Grainger leaves his readers with a glint of optimism. Essentially, the book examines the history behind the environmental manipulation of 20 mammals, and the consequences of such action when poorly planned and executed. The text is accented with color portraits and black line drawings of the mammals. A_ bibliography, although broad in scope, is provided for those who wish to continue on with the study of these mammals. PAUL A. GRAY Hough, Stansbury and Associates Ltd., Suite 409, 1265 East Arthur Street, Thunder Bay, Ontario P7E 6E7 with each treatment generally containing information on the species’ physical features, its habitat require- ments, and its distribution. Scattered throughout the text are stories of hunting expeditions. Some of these hunting expeditions were mounted in an effort to collect specimens for museum collections; some merely hunting adventures. The Great Arc of the Wild Sheep was first printed in 1964. This fouth edition does not include a great deal of new information over the first edition. To enjoy this book, one must be truly interested in wild sheep. PETER CROSKERY Ontario Ministry of Natural Resources Ignace, Ontario POT 1TO 389 340 THE CANADIAN FIELD-NATURALIST A Guide to the Birds of Venezuela By Rodolphe Meyer de Schauensee and William H. Phelps, Jr. 1978. Princeton University Press, Princeton, New Jersey. xxii + 424 pp., illus. Cloth US $50.00; paper US $19.95. The publication of this guide is a noteworthy addition to the ornithological literature of South America in that it summarizes in one volume descriptive, distributional, ecological, and taxonomic information about the 1296 species of birds known to have occurred in Venezuela. The book comprises an introduction in which the Venezuelan avifauna is summarized too briefly (114 page): the geographical and ecological features of the country are dealt with briefly also (4 pages). Travel to and in Venezuela is treated in a few words. A plan of the book and short comments about the illustrations conclude the section. A “Chart of a bird” and a “List of families of Venezuelan birds” follow. The scope of the book is limited to Venezuela but nevertheless encompasses a very broad topic. The book, to quote an excerpt from the introduction, “contains a description of all the species of birds known at present to inhabit Venezuela and its adjacent islands ....” Most of those species, except for some North American migrants, are depicted in the 53 plates assembled in the center of the book. A map of Venezuela appears on both inside covers and cor- responds to geographical entities mentioned in the text. Each family is introduced by a short paragraph in which are given general characteristics of the family, a short summary of the habitat preferences, other ecological facts, and general information of interest. The information dealing with each species is short but factual. The authors have provided in addition to the scientific name, Spanish and English common names for all species. The latter are particularly well chosen and descriptive. Descriptions are given in telegraphic style and, to my knowledge, are accurate, at least for the 25-30 species that I have checked against specimens. Characteristic details peculiar to a species are sometimes given in italics and are provided for both sexes in species where sexual dimorphism is present. Total length and occasional measurements of certain body parts are provided (metric and imperial systems). Descriptions appear to be adequate for a large number of species but one wishes that more distinguishing features had been given for similar species. This would certainly be appreciated in the field. Zoogeographical data are contained in a short paragraph, while in addition to indicating the “life zone” of the species, it summarizes the range in Venezuela and in peripheral countries. The ecological Vol. 93 and distributional data are necessarily brief but are adequate in orienting the observer in the field. In addition, this part often contains behavioral infor- mation as well as descriptions of calls or call notes. Line drawings of an exceptional quality by Michel Kleinbaum supplement several descriptions. The other illustrations are grouped in the center of the book and “show virtually all of the resident species of birds found in Venezuela.” Thirteen black-and-white plates (i-xili) by Guy Tudor are of excellent quality and are particularly efficient in depicting raptors in flight. The 40 color plates have been contributed by Tudor, Wayne Trimm, John Gwynne, and Kathleen Phelps. They are of very good quality in general but Tudor’s are exceptional. Field characters are given for each species opposite each plate, or ina few cases, on following pages where species are too numerous on a single plate. The layout is attractive and species have been arranged in a logical manner on the basis of similarities between them; this is undoubtedly the most practical approach to field identification. Of course, additional details on distinguishing characters appear in the textual des- criptions, which can be located rapidly, the page number being indicated next to the species name. All illustrations should make it easy for the field observer to identify rapidly a large number of species, at least the males. Flycatchers will no doubt continue to bea nightmare! A species list based on specimen records deals with the birds of Isla de Aves on page 369. Subspecies depicted in the illustrations are listed in an appendix, followed by a short bibliography, which should have more correctly been called “selected references.” Indices to English and Spanish names, as well as to genera and species are given in the last pages of the book. The cloth copy is not solidly bound and it appears that it may not resist intensive use in the field. Furthermore, the price of the cloth edition is very high in comparison with the paper edition and I wonder why hard covers alone cost $30.00 (USA)! In spite of this shortcoming, this guide is the best available to date for South America, and the authors and artists are to be congratulated on their excellent work. I highly recommend the book to anyone interested in South American birds or planning to watch birds in northern South America. HENRI OQUELLET National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario K1A 0M8 7) 1979 North American Moose By Randolph L. Peterson. University of Toronto Press, Toronto. 280 pp., illus. Paper $13.50. This book, first published in 1955, has been re- released in paperback. At the time the author did his research it represented the only serious long-term study on Moose in North America. It is still well worth reading, both for the uninitiated and for those of us actively working with this species. North American Wild Geese By M. A. Ogilvie. Buteo Books, Vermillion, South Dakota. 340 pp., illus. US $25. Recently there have been several books written dealing with waterfowl in a general manner (Johns- gard 1975; Bellrose 1976; Palmer 1976). Unlike these former works, “Wild Geese” deals only with geese, and ona world scale as opposed to one continent. The format is considerably different from the usual and somewhat dreary collection of individual treat- ments by species. Instead, subjects are examined including classification, identification, ecology, breeding, population dynamics, migration, distribu- tion, and status. Within this framework, the various species of geese are discussed. This tends to make the text flow in a manner that is pleasant to read. Also contributing to the ease with which this book may be read is the author’s considerable talent in dealing with often highly technical material in a fashion readily understood by the amateur ornithologist. It is well- illustrated, by Carol Ogilvie, with color plates; the pen-and-ink sketches throughout are especially good. The major criticism the reviewers had, of a general nature, was the tendency towards too much detail on certain subjects and limited coverage of others. For example, ringing (banding) notably of Pinkfeet, numbers in subpopulations, and molt migrations all received extensive discussion. On the other hand, behavior (other than during breeding), nutrition, and physiology perhaps should have been covered in greater detail. Bergmann’s rule is mentioned in several locations. This rule is at best tenuous when applied to geese, a fact noted by the author on p. 121. Ross’ Geese and Lesser Snow Geese do not illustrate the rule as stated (p. 25). Over 65 percent of Lesser Snow Geese breed south of most Ross’ Geese. This discussion should have been limited or omitted altogether. The book contains many small errors. Map 25 should include heavily used areas at James Bay and North Dakota. Snow Geese migrate directly to the Gulf of Mexico from James Bay, not Hudson Bay. BOOK REVIEWS 34] Moose presents a foundation on which biologists have built their knowledge of Moose. Asa follow-up to this book and for a more recent compendium of the state- of-the-art on Moose, I recommend Alces, Moose Ecology, Les Presses de Université Laval, 1974. IAN D. THOMPSON P.O. Box 895, Cochrane, Ontario POL 1C0 Map 24 shows no migration out of areas 7 and 10 while some movement further south is known to occur. Ross’ Geese, not using the west coast, winter primarily in Texas as well as Louisiana (Palmer 1976). Numbers of Canada Geese in the Mississippi Flyway are over 700000 not 60000 (Hanson 1974). Cap Tourmente has been used for closer to at least 500 than 400 years. North American waterfowl hunters do not have to report their kills. The second reviewer was surprised to see data quoted from his doctoral thesis, yet not referenced (Prevett 1973). Minor errors such as these can be expected in a book of this type, particularly when a British author tackles the North American waterfowl literature. On the whole, this book is informative yet easy to read. The author has done an excellent job of compiling the extensive literature on geese into a straightforward concise work. We recommend it particularly for amateur ornithologists and biologists not familiar with geese. Biologists will especially appreciate the chapters on ecology and breeding. References Bellrose, F.C. 1976. Ducks, geese and swans of North © America. Harrisburg, Stackpole Books. 544 pp. Hanson, R. 1974. Report on the December, 1973 inventory of Canada Geese in the Mississippi Flyway States. U.S. Fish and Wildlife Service Report. 13 pp. Johnsgard, P. A. 1975. Waterfowl of North America. Bloomington, Indiana University Press. 575 pp. Palmer, R.S. (Editor). 1976. Handbook of North American birds. Volume 2. New Haven, Yale University Press. 521 pp. Prevett, J. P. 1973. Family behaviour and age dependent breeding biology of the blue goose, Anser caerulescens. Ph.D. thesis, University of Western Ontario, London, Ontario. IAN D. THOMPSON! and J. P. PREVETT? 'P.O. Box 895, Cochrane, Ontario POL ICO 2Moosonee, Ontario POL 1Y0 342 Crows of the World By Derek Goodwin. 1976. Cornell University (Comstock), Ithaca, New York. 354 pp. US $28.50. Press It is fitting that Goodwin should contribute this authoritative book on another of his favorite groups. He began work on the jay about 30 years ago, and his long tenure with the British Museum of Natural History has allowed him further opportunities to explore the fascinating crow family. His earlier work, Pigeons and Doves of the World (1967), has recently been reissued as a second edition (Cornell University Press, 1977). The style, a welcomed combination of scientific yet non-technical writing, is similar for the two books. Because it summarizes much of what is known and clearly shows what is unknown, it is an excellent reference for ornithologists yet is very readable for amateur naturalists. The general format also is similar to his earlier work. It begins with four general chapters on nomenclature, adaptive radiation, plumage and coloration, and behavior. The 41-page behavior chapter discusses a myriad of behavioral characteris- tics and is especially interesting. Detailed species accounts are given in Chapter 5. An introductory two-and-one-half pages of Chapter 5 provide an interesting account of man’s relationships with corvids, touching upon their role in our folklore, legend, and literature; our feelings toward their predation on eggs and young of small birds; the crop damages they cause; and their use as food. Where species comprise a distinct group, an introductory preface of taxonomic relationships and habits is provided. Each of the 116 species is then covered in detail — description, field identification, distribution and habitat, feeding and general habits, nesting, voice, display and social behavior, other names, and a list of references. Length of the accounts varies from about one-half page for the Banggai Crow, Flores Crow, and Life of the Harp Seal By Fred Bruemmer. 1977. Optimum (Prentice-Hall, Tor- onto). 170 pp., illus. $25. Fred Bruemmer has produced a very readable work on the life of the Harp Seal. His style is pleasant and flows easily as he provides accurate accounts of the environment, culture, social structure, and history of the Atlantic region where the sealing activity takes place. His descriptions of the ice-environment are especially good. His commentary on some aspects of the biology of the Harp Sealare at times insightful and competent. Sometimes, however, Bruemmer’s sci- entific commentary is mixed with ethical considera- THE CANADIAN FIELD-NATURALIST Vol. 93 Sumatran Tree Crow to over eight pages for such well- studied species as the jay and magpie. For several species, there is no information provided other thana description and range. The many “No information” statements should certainly spur young ornithologists to go “afield.” As the author states, one of the book’s purposes is to indicate what is not known. About one- fourth of the pages in the species accounts chapter deal with the 16 species occurring in Canada and the continental United States. Robert Gillmore has done an excellent job illustrating many of the species with line-drawings. Three colored plates showing adults of 22 species and juveniles of 4 species are nicely done and show the variety of shapes, colors, and patterns within the Corvidae. Range maps are provided for each species. These are small, general, and do not give any indication of seasonal occurrence. Migration or movements, in general, is the only subject I feel is slighted in the entire book. I would suspect that because of their size, the maps may not be totally accurate nor reflect up-to-date distributions. No mention is made of the regular occurrence of the Brown Jay insouth Texas. One other omission I noted was the absence of a discussion of hybridization in New World corvids, a rare but documented event. Common and scientific names are indexed following Chapter 5. This fully referenced volume stands as an extremely useful and important contribution. I believe that ali readers will attest to the accuracy of the author’s statement that the crow family is a numerous, diverse, and successful group of passerine birds. NOEL J. CUTRIGHT Wisconsin Electric Power Company, 231 W. Michigan, Milwaukee, Wisconsin 53201 tions which seem out of place. For example in a discussion of the seal’s underwater hearing ability, he points out that the seal is “as one with the sea, ... but when she must leave the sea... then she and her kind are at the mercy of man.” The text is ripe with anthropomorphic references to “sad and worried mothers” and “jealous mates” which might be forgiven as poetic licence. The cause of the seal is taken up with such emotion, however, that the reader is left to question whether Bruemmer is offering the promised clear and unbiased account of the life of the Harp Seal. Photographs are as important as the text 1979 of this coffee-table-size book. Ninety-eight color and black-and-white photographs which range in size from 2 X 3 inches to double-page spreads are con- tained within the book’s 155 pages and many of them are excellent, as is characteristic of the author’s other books. But a seal pup, in various appealing poses, appears in 61 of these pictures, as if one animal had been photographed many times from different angles. Most of the photographs are technically good, but many are only fair. In the few underwater sequences detail is difficult to resolve, and the large plate on page 101 is upside down. Federal fisheries scientists will be disappointed with Bruemmer’s half-page comment on Harp Seal management. Most of it is cynical and disparaging, and refers to the maximum sustainable yield as the Holy Grail of management biologists. Bruemmer spends much more time describing the “interfering” activities of man (sealers, tourists, aircraft) in the seal’s life. Bruemmer’s description of sensory functioning in Harp Seals includes an able discussion of much of the data on the vision of seals and how it relates to their behavior in air and under water. His descriptions of audition and of vibrissae functioning, however, are inaccurate and at times patently false. He states that the seal’s audiogram in air is similar to, if not better than, that of the human. In fact, though seals can hear higher pitches than humans, they are at all fre- quencies less sensitive to quiet sounds. His suggestion that a seal could hear a man walking on the ice a mile away is unlikely when one considers the relative acoustic impedances of ice and water. As far as whisker functioning is concerned, his statement that seals use their vibrissae to detect changes in water pressure, or the “tap of water against blocks of ice pressed far beneath the floes,” and his implication that the whiskers can perceive the turbulence of fish movements over some distance are totally unfounded. Current evidence indicates their vibrissae are not sensitive enough for this. His confident elaboration about the sense of smell being the primary means by which mothers and pups identify each other, and his contention that mothers can recognize the call of their own pups is completely unwarranted since such hypotheses have never been tested. Similarly, to our knowledge, his statement that pups are able to locate their mother’s teat by an increase in skin temperature in the area of the nipple is unsubstantiated. Also, he implies that the mothers initiate weaning; however evidence from Harbor Seals BOOK REVIEWS 343 shows that pups wean themselves. Bruemmer describes in some detail the navigational methods Harp Seals use to traverse the distance between Greenland and their breeding grounds. In fact, very little is understood about the way in which any seal species plots its course over long distances. In a similar vein he states without qualification that Harp Seals can stay submerged for at least thirty minutes and can dive to depths as great as 600 feet, when in fact these numbers were obtained from the Antarctic Weddell Seal. In his discussion of the social behavior of Harp Seals, Bruemmer asserts that these animals are gregarious, and that during the breeding season the female, when hauled out on the ice maintains a “moveable territory” about herself that only her pup can encroach upon. These are probably reasonable hypotheses although they have not been directly investigated. In further descriptions of reproductive behavior, however, his portrayal of the events leading to copulation are at best misleading and at worst completely incorrect. He states that a single male emerges the victor of an often bloody battle, to mate on the ice with a willing female who has been wooed by an hour-long display. In reality, nearly all copulations occur in the water and therefore have rarely been witnessed. The only published under-ice observations of reproductive behavior in these ani- mals suggest that the seals are probably promiscuous, the females engaging in sexual displays with many males. Furthermore, as is the case in most mammalian species, true “bloody” battle is the exception rather than the rule: agonistic behavior in this context has become ritualized. Bruemmer’s description of an hour-long display on the part of the “victorious” male is the first such report. For a decade Canadians have waited for an independent and unbiased commentary on the Harp Seal and its harvest. One might have expected: Bruemmer to have provided it since he has produced a number of good works. Readers hoping to find enlightenment on this contentious issue will find this book a disappointment. TOM H. NORTHCOTT! and DEANE RENOUF? ‘Northland Associates Limited, P.O. Box 1734, St. John’s, Newfoundland AIC 5P5 *Department of Psychology, Memorial University, St. John’s, Newfoundland AIC 5S7 344 THE CANADIAN FIELD-NATURALIST BOTANY Edible Garden Weeds of Canada By A. F.Szczawinski and N. J. Turner. 1978. National Museums of Canada, Ottawa, Ontario. 184 pp. $8.95. This book is the first of a four-book series on the edible wild plants of Canada slated to be fully published by 1980, under the auspices of the Canadian National Museum of Natural Sciences. In this context, Edible Garden Weeds gains a significance that it would not otherwise have. The volume is published in paper wrappers with a wire binding that allows the book to lie flat open at any page, a desirable feature in any cookbook. And in fact, the great inspiration of this work is the combination of the “standard” cookbook format with the equally “standard” edible weeds or wild plants format. In our experience it stands alone amongst similar books on this count, other texts generally immersing recipes in a chatty dialogue with no comparable systematic layout of ingredients and instructions. Pages are odd sized (approximately 21 X 21 cm) and, unfortunately, the use of space is wasteful; in fact Edible Garden Weeds stands alone in this regard also. The bulky size, and useless weight of many '4- to 7- full pages certainly makes the book of less use to “scouts, outdoorsmen, and others concerned with survival.” We hope that this “spacious” format approach is dropped in other books of this series by the National Museum, for if it is carried through it will be no less than a slap in the face for conservation. The main body of discussion of species and recipes is printed on white glossy paper with an attractive type style and a rather nice, very dark green ink. All other pages are of colored paper, brown in front and bright yellow at the back. The many line drawings are clear and crisp, and only rarely have the artists “fudged” the smaller details of the plants. Several color photos add a definite exciting touch to the book. Actual contents of the text provide a good intro- duction to the edible weeds dealt with; however, we feel it is deficient and misleading in several specific and general instances. The fact that many areas where weeds grow prolifically have poisoned soil needs to be emphasized and re-emphasized, especially to a domi- nantly urban audience. This is a high paradox in this book, for while people are warned away from a few clearly edible species on rather slim grounds, no mention is made of the dangers of lead toxicity inherent in many an “unkempt boulevard.” The plethora of dangerous industrial chemicals that today contaminate many rail rights-of-way, roadsides, and Vol. 93 vacant lots is likewise unmentioned. Asa general rule, any weeds growing near urban or industrial develop- ments should be avoided, but in fairness Edible Garden Weeds is not alone in its lack of this warning. Also, we wish that the nutritional virtues of weeds, when growing in uncontaminated sites, had been far more completely explored. Given the small number of species and species groups treated (about 40), wouldn’t it have been possible to produce a more precise discussion of these virtues, and fill some of the waste space in the book with, say, distribution/ abundance maps? Most of the text discussion is a reasonably complete “standard” description of plants, habitats, uses, etc., slightly modified better to fit the Canadian context, and clearly organized under headings. The many recipes (about 130) without doubt contain the most controversial aspects of this book. They range from simple teas, salads, soups, and spreads to lunches and even rather exotic meals. On the one hand, the recipe layout and use of common “supermarket” ingredients will be attractive to many urban and suburban cooks. On the other hand, natural food buffs can object to over forty ingredients as chemical-ridden or simply “non-food” items. At least fifteen of these could readily have more natural or “real” foods substituted, and the complexity of some recipes should be questioned. A certain balance of nutrition and health seems to have been overlooked in the attempt to make “weeds” credible to the general cook. This is not a book for the backwoods or farm, but rather seems designed for suburban or estate resi- dential environments and tastes. The attempt seems to be to make “weeds” acceptable by associating them with heavily flavored and often exotic ingredients within a colorful, open and “artsy” publication style. Much information is presented, but in no way can the text be considered a critical analysis of the values and uses of edible weeds and it stands on relatively equal footing with several other easily available works. Any one who wishes to understand properly the value of edible weeds will still have to purchase a collection of several books, and in this context the acquisition and use of Edible Garden Weeds of Canada becomes a matter of personal taste. R.S.W. BOBBETTE! and LANDA L. KING? 131 Eccles St. N., Barrie, Ontario L4N 1Y1 28 Peel St., Apt. #3, Barrie, Ontario L4M 3K8 7s) Ferns of the Ottawa District By William J. Cody. 1978. Printing and Publishing, Supply and Services Canada, Hull, Quebec KIA 0S9. 112 pp. Soe: The revised edition of this book is quite delightful. There is a modern durable plastic binding with some attractive silhouettes on it and the printer has done a superb job with good modern clean type and the right mix of various sizes of bold-face type, italicized type, etc. By the clever use of dividing lines, the author has solved the problem of associating the figures, the text, and the distribution maps for each species. The illustrations of selected herbarium specimens are by and large the same as in the first edition of 1956, but because they are on glossy paper, have slightly less gray in the background and have a black line on three sides of the plate, they seem infinitely clearer, sharper, and more three dimensional. Another great im- provement is the distribution maps, which are much larger in size and so crystal clear that one feels one could almost rush out to a given locality dot! The keys are simple and workable and the addition of five new species to the District (Pellaea atropurpurea, P. glabella, Woodsia obtusa, W. oregana, and As- plenium platyneuron) plus a plethora of new locality dots, e.g., Ophioglossum from 5 to 21, brings the revision right up to date. In these days of soaring book costs, this book is a tremendous bargain and should be purchased by anyone interested in ferns. What are the shortcomings of this fine identifi- cation manual? Most of my criticisms are a plea for further information from authors. For example, consider the distribution maps. Do the dots each represent a herbarium specimen or do they represent all records known to the author? Should a distribution map represent the status of the species today or should it include historical records (Dryopteris goldiana now under a shopping mall)? What about sight records and specimens in private herbaria? Next the taxonomy, which might be dubbed “authoritative conservative.” Admitting that the names used are familiar ones and this manual is probably not a good vehicle for radical taxonomy, it still seems the author might include a statement of his philosophy, or even a plaintive disclaimer that he is a follower of, or influenced by, Gray’s Manual, Britton and Brown, Wherry or whatever. It is most confusing to the amateur to see Camptosorus in this book and Asplenium in another; also Athyrium vs. Diplazium. Similarly, the Long Beech Fern can be found in Thelypteris or Phegopteris and as P. polypodiodes or P. connectilis. Fortunately the author has supplied a few synonyms for cross reference. I do not like the treatment of Dryopteris, partly of course because of my personal bias and knowledge of BOOK REVIEWS 345 the genus. I feel that there should be six species treated and not four and also wonder why D. X boottii is singled out as the one hybrid recognized for the district. The most common hybrid is alluded to on p. 70 (D. X triploidea Wherry). The common name, Florist’s Fern for var. spinulosa (not D. intermedia) on p. 66 is most unfortunate. I cannot imagine a florist using this fern and would deplore the attempt! Another point concerns the designation of variety. This seems to be a legacy of Gray’s Manual. Surely if one 1s not going to recognize our Ostrich Fern as a distinct species, it is at least a subspecies? Some varieties seem a bit trivial or useless, e.g., Botrychium matricariaefolium var. matricariaefolium, B. multi- fidum var. intermedium, and Polystichum braunii var. purshii, whereas others such as Dryopteris cristata var. clintoniana are major biological species as shown by cytogenetics. I think a few references for the amateur who wishes to delve more deeply would be extremely worthwhile. Amateurs are also apt to ask for the rationale of the sequence of genera as presented. Why is Dennstaedtia between Polystichumand Athyrium? If no grouping is intended, or no authority followed, then why not use an alphabetical approach? Ideally it would be nice if authors were a bit more explicit about field characters or “tricks of the trade.” How will you know a Pellaea when you see one, instantaneously? For instance, emphasis on indusial characters for Cystopteris and Woodsia when these are obscure on mature fronds confuses amateurs. I foresee difficulties for workers distinguishing between Woodsia obtusa and Woodsia oregana with the key. A minor annoyance in manuals is the rather unspoken assumption that one can map a species for all time if one just carefully “covers the ground.” This is far too static a concept for plant populations. Accordingly, when we are told under Dennstaedtia, “Large ferns in colonies” and then later “perhaps it 1s overlooked,” I would prefer the idea that it is spreading and becoming more weedy in the district. Similarly, with Asplenium platyneuron, | doubt it was there until recently and will not be surprised if the locality dots increase in numbers very shortly. I feel the author has done a most commendable job. What I am really saying is, give us more — more references, more explanations, more taxonomic philosophy, and we will gladly pay more! DONALD M. BRITTON Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2WI 346 THE CANADIAN FIELD-NATURALIST Vol. 93 Dyes from Lichens and Plants: a Canadian dyer’s guide By Judith Waldner McGrath. 1977. Van Nostrand Reinhold, Toronto. 144 pp., illus. $14.95. Translated into Inutituut and syllabics. How this book came to be written is as interesting as the information on dyes from native plants. Mrs. McGrath is a dyer, a weaver, a sculptor, and a teacher. In recent years she has lived in several northern settlements following her husband in his work with the Economic Development Council. Spence’s Bay, where the book was written, is a community of 400 people on the Boothia Peninsula, a tiny speck on the bleak arctic landscape when seen from the air. On arrival, it was a shock to find native people in modern clothes and living in ordinary homes, but as Mrs. McGrath made friends with the Inuit she recognized that cultural differences do exist. In the brief arctic summer flowers bloom every- where. Continual daylight encourages growth but high winds, lack of moisture, and occasional snow cause severe stress to the environment. Yet close to the ground, where the air is warmer, grow some 900 species of flowering plants and perhaps 1000 lichens. It was here that Mrs. McGrath began her search for dye stuffs. Each find was shared with her new friends. Soon an experimental workshop was set up, opened throughout the long winter night so that women could work when it was convenient. The following summer a grant made it possible to gather plants farther afield and to experiment with colors that could be obtained from them. To the Inuit women, used to bright commercial dyes, the new muted shades were exciting. Designs made from them won awards, and official parkas for the 1974 winter games were designed and made here. By 1977, when the McGraths left, the craft shop was becoming a co-operative — 32 people were involved in the new cottage industry. Much of the book is devoted to the plants them- selves, to the parts used and the amount of dye material needed for best results. It is impressive that from the beginning there was concern for the ease with which the fragile ecological balance could be destroy- ed. Spot collecting was done over a wide area. Only a part of each plant was taken, leaving some roots for propagation and some flowers to set seeds. Material was preserved by drying, by fast freezing of plants, or by preparing and freezing the dye bath. Inall, 17 basic dye methods or recipes were evolved. Of these nine are for lichens, the rest for flowering plants, seaweeds, and mosses. Few colors remain fast on fiber without the use ofa mordant such as alum, iron nails, or copper. Mordants combine chemically with the dye and the fabric to fix the color. Variation in shade can be obtained by changing the mordant used or the time at which it is added. For the beginner the easiest plants are those with their own mordant such as sumac, which contains tannin, and the lichens which have their own acids. There are chapters on the people and their crafts and on the land itself, brought to life by black-and- white photographs. Beautiful color plates illustrate groups of plants and display small tie samples of wool colors obtained from them. There are lists of useful equipment, of stores that carry supplies for dyers, an index of plants by colors they can produce, another of plants in more temperate parts of Canada that will produce comparable colors, and one giving botanical as well as common names. In preparing the book, government departments in both Ottawa and the Northwest Territories were consulted. Identification of all plants and lichens was confirmed at the National Herbarium. My only complaint concerns the color plates. They are very attractively arranged and colors are good, but there is no explanation of the photographs. In this section, one finds only outlines of illustrations with difficult-to-decipher numbers and letters. These illus- trations would be so much more useful in the part of the book where plants and dye colors are described. Except for the difficulty with color plates the material is well and interestingly prepared. The author knows her subject and has been generous with that knowledge. Mrs. McGrath says that care in each step from washing of the fiber to mordanting and dyeing affects how well and evenly colors take. Don’t worry about the experts: read them and listen to them, but the only sensible course is to experiment yourself. Although the book is presumably intended for weavers and dyers, many people unfamiliar with the craft will find pleasure in it. The book is sturdily bound with an eye-catching jacket. It was written and published in Canada. Highly recommended. MARY I. MOORE Petawawa Forest Experiment Station, Canadian Forestry Service, Chalk River, Ontario KOJ 1J0 1979 ENVIRONMENT Human Activity and the Environment By Statistics Canada. 1978. Statistics Canada, Ottawa. 190 pp., illus. Paper $2.80 in Canada; $3.40 elsewhere. Statistics Canada has attempted to put together a “statistical series detailing human activities which have a potential for imposing stress on the natural environment.” The resultant book has seven chapters which are entitled watersheds; agriculture; forests; fisheries; transportation; manufacturing; energy. It is an interesting and welcome initiative by Statistics Canada. A novel section contains the presentation of certain social economic data by the natural physiographic unit of the watershed. All of Canada is delineated along the boundaries of the major, and some minor, watersheds. Some of the data given by these units include human population; watershed area; human population density; farms and farmlands fertilized, sprayed and irrigated; livestock numbers; and industrial activity. Some interesting examples of data from the text are given below. The large increase in the sales of pest control products, which are also known to some of us as biocides, over the last 30 years is somewhat disturbing. Since 1971 the increase appears to take the shape of an exponential curve! Statistics Canada outlines the striking decline in the weight of fish taken by Canadians off Canada’s shores from 1968 onward. The corresponding increase in landings by other nations, most notably the Soviet Union, provokes thought. An attempt is made to make the text visually interesting. This attempt works reasonably well. One illustration which stands out is the complex map of noise contours around Toronto International Air- port. The noise magnitudes are presented by various shades of blue. The many people that live in the area BOOK REVIEWS 347 and are subject to the noise disturbances often see red on this issue. The data presented is scattered in content and given uneven treatment. Nevertheless one must sympathize with the authors’ considerable task of choosing subjects in such a broad field. There are some obvious biases ina document of this kind. First, the environmental phenomena must be easily measurable and quantifiable. Second, there must be an existing measurement program underway. Most readers of this journal could easily think of many critical environmental issues that badly need methodological measurement. For example, human activities often put considerable stress on wildlife populations but usually we have only partial data on what population changes do occur. Some classes of data that might be given consideration for future editions include the follow- ing: 1) The attitudes of Canadians towards the environ- ment. Who cares? How much? About what? 2) The recreational pursuits of Canadians in terms of dollars and time spent. 3) Canadian land-use changes over time. The existing section in the book should be expanded. 4) Air quality data. 5) Water quality data. 6) Various wildlife populations data. It must be kept in mind that this first attempt has broken considerable new ground. Hopefully, refine- ment and enlargement will take place with future editions. PAUL F. J. EAGLES School of Urban and Regional Planning, University of . Waterloo, Waterloo, Ontario N2L 3Cl Vermilion Lakes Banff National Park: an introductory study By the Bow Valley Naturalists. 1978. Bow Valley Natural- ists, Banff, Alberta. 68 pp., illus. Paper $3.00. The Vermilion Lakes area, immediately west of the town of Banff, in Banff National Park, has long beena focus of interest for visiting and resident naturalists. The lakes themselves are large, shallow ponds which (for the mountains) are highly productive aquatic systems. The view of the north end of Mount Rundle across the lakes is a famous and frequently photo- graphed sight and has become almost synonymous with Banff. The richness of the area, coupled with increased use by visitors and the threat of further impact by highway construction along the northern edge of the lakes has heightened concern for its future. This publication is, in part, an attempt to increase public awareness of the values of the Vermilion Lakes in the face of new pressures on them. The study covers a range of cultural and natural history fields, each treated in a separate section and based on the field studies and research carried out by various members of the Bow Valley Naturalist Club in 348 the mid-1970s. A set of recommendations regarding the future use and development constraints on the lakes are made which result from an assessment of the data assembled during the study. All of this is pulled together in a tight, attractive, paper-covered booklet which ts effectively illustrated by line drawings, photographs, and maps. A some- what cumbersome poem about the lakes starts it all off, and the booklet ends with a fairly extensive list of references. It is well put together, with clear easily readable type, crisp appropriate photographs and some excellent pen-and-ink drawings. In format and production it is a pleasing and effective document to deal with. The study is intended to introduce the reader to the Vermilion Lakes. It does that quite admirably. | suppose my only major criticism is that the authors held themselves to this. Detailed flora and fauna data are not provided despite quite extensive studies in some of these fields. Appendices are maintained by the Bow Valley Naturalist which contain raw lists of the observations reported during the study period, but these are not easily accessible. In the booklet proper, only very general statements of status are included (e.g., “occasional in wet spruce forest” or “fairly common. Nests”). This does introduce the reader to the subject; it would have given him so much more if that extra effort had been made to pull these obser- vational data together into precise, clearly docu- mented discussions. It seems that making such statements as ... “The Cougar appears to be on the verge of extinction in Banff National Park,” and the American Bittern ... “Must nest but no records,” are quite inadequate without more supportive documen- tation and discussion. MISCELLANEOUS Weather Almanac Edited by J. A. Ruffner and F. E. Bair. 1977. 2nd edition. Gales, Detroit. 728 pp. US $25. The Weather Almanac is useful to both the layman who is interested in weather phenomena and the technical people who use weather data in their jobs. The section on “Weather Fundamentals” would be a better section to put first than the “Atlas Format Weather.” I recommend that this section be read first to both interest and inform the reader. I am slightly disappointed that the 1941-1970 tridecade was used for most tables when an upgrade using data from 1941-1976 is easily available from the THE CANADIAN FIELD-NATURALIST Vol. 93 A volunteer effort of this nature is very dependent on the quality and quantity of the individual effort contributed by concerned naturalists. Inevitably, the quality of content varies throughout in such an effort. The herpetological section, for example, is very strong, whereas the botanical material is not so. I understand that this latter area suffered considerably because of Parks Canada’s unwillingness to provide a collecting permit for the study! Although quite free of typographical errors, a number of spelling errors and omissions do occur (e.g., Gallium for Galium, incomplete scientific name for Alder Flycatcher, Spirea for Spiraea). These are infrequent enough, however, neither to affect seri- ously the credibility of the document nor greatly to impair its readability. The Vermilion Lakes study is a fine effort by a number of concerned and dedicated naturalists who have obviously gone to considerable personal expense and effort to increase our awareness and appreciation of an important natural area in Banff National Park. I hope it inspires other groups to embark on similar ventures and I would certainly recommend it to those readers who would like to know more about Banff. I recommend it even more strongly to those of us who often feel that individuals and small groups of naturalists can’t do much to document and publicize the values of an area which we cherish. The Bow Valley Naturalists have shown that it can be done and they are to be congratulated for their fine efforts. D. F. BRUNTON Kananaskis Provincial Park, Alberta Department of Recreation and Parks, Canmore, Alberta TOL 0MO weather service. This is especially important since new extremes were set in 1976 and 1977 and the almanac does not, therefore, explain the harsh winters for example. The “Glossary of Weather Terms” is certainly useful as it explains in general terms most of the meteorologist’s jargon. The section on “Climates of the World” is very useful for travellers but lacks an overview of “Climate” in general. It would be better to put our present world climate into an historical perspective. We are presently in an abnormal period of climatic 1979 history. We are coming to the end of an interglacial warm period and the tridecade data reflects only this period. Throughout most of the world’s existence, the climate has been “glacial” and the fact that it is not now is both interesting and gives man a sense of the enormous impact of climate. This leads naturally into the “Health and Weather” section and the awareness that there are scientific relationships between health and the weather. The descriptions and safety rules for “storms and severe weather” are clear and especially useful for self- protection in the winter. Folklore (such as p. 119) and rules of thumb (distance to storms, be your own forecaster) add immeasurably to the dry textual information. Earthquakes are not strictly speaking a weather phenomenon, but the tables do allow the reader in a specific area to review this phenomenon both historically and currently. The section on “Air Pollution” and its effects (Table 5.2) is slightly misleading in that there is conflicting evidence on effects and levels of pollution at which they occur. Similarly Figure 5.14 does not cover the case of a plume that is hot enough to break through a weak inversion, thus trapping pollutants above it. In this case almost none of the pollutant comes down to the ground to affect receptors. Introduction to Mathematics for Life Scientists By E. Batschelet. 1976. 2nd edition. Springer-Verlag, New York. 643 pp. Cloth US $27.60; paper US $11.95. Most frequently, book reviewers possess some ex- pertise in the subject area of the book they are reviewing. This review departs from that tradition. My mathematics is best described as shakey. For precisely this reason I feel qualified to recommend Introduction to Mathematics for Life Scientists as a study guide and reference book for those who feel uncomfortable with the steadily increasing number of non-linear functions, differential equations, and ma- trix operations to be encountered in the current literature. To facilitate study the chapters are comprised of bite-sized sections with alternating theory and ex- amples or applications. Most examples are from the recent literature and referencing i$ thorough through- out. Each chapter is concluded by a section of problems, graded in difficulty, with answers for odd- numbered ones. Most commendably the author starts at the beginning with a review of basic algebra, a knowledge of which is all too often erroneously assumed by other BOOK REVIEWS 349 The section on “Energy and Weather” is an excellent introduction to the fundamentals of energy conservation for the days of restraint ahead. The “Weather of Selected U.S. Cities” is a good general summary of climate information and is well documented and presented. The “Atlas Format Temperatures,” on the other hand, is confusing and, because of the type size, almost unreadable. It would be more useful to have one map showing the climate station locations and tables of highs, lows, and averages. The “normal monthly precipitation and snowfall” maps give easy reference to the amount of rain and snow visually but the type is too small to read for quantitative information. Although there are a few typographical errors in the text, on the whole the almanac would be a very useful addition to either a layman or weather scientist’s library. JAMES W.S. YOUNG Atmospheric Environment Service, 4905 Dufferin Street, Downsview, Ontario M3H 514 texts of similar intent. Subsequently, linear, power, periodic, and logarithmic and exponential functions are dealt with. Chapters on ‘limits’ and ‘differential and integral calculus’ build up to a treatment of differential equations. Chapters on graphical methods and probability are essentially self-contained. On the other hand the chapter ‘matrices and vectors’ requires considerable familiarity with earlier sections of the book. A detailed list of contents and a thorough index make this a book to which one can refer easily. The clear concise text constantly refers back to the precise place where a difficult concept or step, likely to be troublesome, was treated. My own fruitful experiences with this book have led me to the opinion that it provides an excellent starting point for any biologist who feels restricted by lack of mathematical expertise. ROBIN MAHON Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1 350 NEW TITLES Zoology Adventure of the stone man. 1978. By F. Hamerstrom. Illustrated by W. Kimber. F. Hamerstrom, Plainfield, Wisconsin. US $6.95 plus 50¢ postage. Alaska whales and whaling. 1978. Alaska Geographic Volume 5, Number 4. Alaska Northwest, Anchorage. 144 pp., illus. US $11.95. Animals in danger. 1979. By Joseph Lucas and Susan Hayes. Clarke Irwin, Toronto. Paper $5.95. *Biology of fishes. 1979. By Carl E. Bond. Saunders, Toronto. 514 pp., illus. $22.75. Biology of insect eggs: Volume I, general biology; Volume II, the eggs of insect families; Volume III, indexes and biblio- graphy. 1979. Pergamon, Elmsford, New York. cl500 pp., US $330. Biology of Reptilia: Volume 7, ecology and behavior. 1977. Edited by A. C. Gans and D. W. Tinkle. Academic Press, New York. xvi + 720 pp., illus. US $60.60. {Birds of the Gainsborough-Lyleton Region (Saskatchewan and Manitoba). 1979. By Richard W. Knapton. Special Publication Number 10. Saskatchewan Natural History Society, Regina. 72 pp. Paper $3. Bird sounds and their meaning. 1977. By R. Jellis. British Broadcasting Corporation. 256 pp., illus. US $11. The Bobcat of North America. 1978. By S. P. Young. University of Nebraska Press, Lincoln. xv + 193 pp. Cloth US $11.50; paper US $3.50. tCamel quest. 1978. By Anne Innis Dagg. York Press, Toronto. 150 pp., illus. Contrasts in behavior: adaptations in aquatic and terrestrial environments. 1979. Edited by E.S. Reese and F. J. Lighter. Wiley-Interscience, New York. 406 pp. US $27.50. Field guide to the seashells of the world. 1978. By G. Linder. Translated from 1975 German edition by G. Vevers. Van Nostrand Reinhold, New York. 272 pp., illus. Cloth US $12.95; paper US $8.95. tFossils of Ontario: Part 1, the trilobites. 1979. By Rolf Ludvigsen. Royal Ontario Museum, Toronto. 96 pp., illus. $7.50. +General patterns of invertebrate development. 1975. By Gary J. Brusca. Mad River Press, Eureka, California. 134 pp., illus. TA guide to the jellyfish of Canadian Atlantic waters/ Guide des méduses des eaux canadiennes. 1977. By C. T. Shih. Natural History Series de l’Atlantique. Number 5. National THE CANADIAN FIELD-NATURALIST Vol. 93 Museums of Canada, Ottawa. 90 pp. English + 90 pp. French, illus. Free. Handbook of common New Guinea beetles. 1977. By J. L. Gressitt and R. W. Hornabrook. Wau Ecology Institute Handbook No. 2. Bishop Museum Press, Honolulu. vin + 88 pp., illus. Paper US $4.50. The history and breeding biology of Canada Geese of Marshy Point, Manitoba. 1978. By James A. Cooper. Wildlife Monographs No. 61. Wildlife Society, Washington. 87 pp. US $3.15. Insect clocks. 1979. Edited by D. S. Saunders. Pergamon, New York. 288 pp., illus. US $11. {The insects and arachnids of Canada: Part 5, the crab spiders of Canada and Alaska (Araneae: Philodromidae and Thomisidae). 1978. By Charles D. Dondale and James H. Redner. Canada Department of Agriculture Publication 1663. Supply and Services Canada, Hull. 255 pp., illus. $7.50 in Canada; $9 elsewhere. } Love affair with a Cougar. 1978. By Lyn Hancock. Double- day, Toronto. 252 pp., illus. $9.95. Mayflies of Michigan trout streams. No date. By J. W. and F. A. Leonard. Bulletin 43. Reprinted. Cranbrook Institute of Science, Bloomington Hills, Michigan. illus. $6.95. Mexican wilderness and wildlife. 1978. By B. Tinker. University of Texas Press, Austin. x11 + 131 pp. US $9.95. Muskrats and marsh management. 1978. By P. L. Erring- ton. University of Nebraska Press, Lincoln. 183 pp. Cloth US $10.95; paper US $3.25. Origins: what new discoveries reveal about the emergence of our species. 1979. By Richard Leakey and Roger Lewin. Dutton (Canadian distributor Clarke Irwin, Toronto). Cloth $29.50; paper $11.95. Patterns of primary production in the biosphere. 1978. Edited by Helmut F. H. Lieth. Academic Press, New York. 368 pp. US $27.50. Planning for wildlife in cities and suburbs. 1978. By D. L. Leedy, R. M. Maestro, and T. M. Franklin. U.S. Fish and Wildlife Service, Washington. 64 pp. Paper US $2.50. Populations of small mammals under natural conditions. 1978. Edited by D. P. Snyder. Papers from a symposium, Linesville, Pennsylvania, May 1976. University of Pitts- burgh Pymatuning Laboratory of Ecology, Linesville, Pennsylvania. xiv + 238 pp., illus. US $8.50. Social insects Volume 1. 1978. Edited by Henry R. Her- mann. Academic Press, New York. 456 pp. 1979 {Standard common and current scientific names for North American amphibians and reptiles. 1978. By J. T. Collins, J. E. Huheey, J. L. Knight, and H. M. Smith. Miscellaneous Publications, Herpetological Circular No. 7. Society for the Study of Amphibians and Reptiles, Lawrence, Kansas. 36 pp. Paper US $2.50. Order from Douglas H. Taylor, Department of Zoology, Miami University, Oxford, Ohio 45056. Statistical inference from capture data on closed animal populations. 1978. By D. L. Otis, K. P. Burnham, G. C. White, and D. R. Anderson. Wildlife Monographs No. 62. Wildlife Society, Washington. 135 pp. US $3.50. Strategies in cold. Natural torpidity and thermogenesis. 1978. Edited by Lawrence C. H. Wangand Jack W. Hudson. Academic Press, New York. 734 pp. US $33. Texas wildlife. 1978. By D. Baxter, T. L. Clark, and J. Jefferson. Texas A & M University Press, College Station, Texas. 196 pp. US $22.50. Wildlife management in wildernesses. 1978. By C. A. Schoenfeld and‘ J.C. Hendee. Boxwood, Pacific Grove, California. 172 pp. Paper US $3.95. Botany The biology and chemistry of the Compositae. 1977. Edited by V.H. Heywood, J.B. Harborne, and B. L. Turner. Papers from a symposium, Reading, England, July - 1975. Academic Press, New York. 1190 pp., illus (2 volumes). US $107.50. A checklist of names for 3000 vascular plants of economic importance. 1977. By E.E. Terrele. Superintendent of Documents, Washington. 11 + 202 pp. Paper US $3.50 + 25% foreign handling. Checklist of vascular plants of the Ottawa-Hull Region, Canada/ Liste des plantes vascularies de la région d’Ottawa- Hull, Canada. 1978. By John M. Gillett and David J. White. National Museums of Canada, Ottawa. 155 pp. Color in plants and flowers. 1978. By J. and S. Proctor. Everest, New York. 116 pp., illus. US $9.95. Embryology of gymnosperms. 1978. By Hardev Singh. Borntraeger, Berlin. 302 pp., illus. DM 138. Essays in plant taxonomy. 1978. Edited by H. E. Street. Academic Press, New York. xxiv + 304 pp., illus. US $24.50. Ferns of north-western Himalayas. 1979. By K. K. Dhir. Cramer, Lohmar, Germany. cl60 pp., illus. cDM 40. Fruits of angiosperms. 1977. By Ingrid Roth. Borntraeger, Berlin. 675 pp., illus. DM 288. +How to identify mushrooms to genus. I: macroscopic features. 1973. By David L. Largent. Mad River Press, Eureka, California. 86 pp., illus. BOOK REVIEWS 3)3)II + How to identify mushrooms to genus. II: field identification. 1977. By David L. Largent and Harry D. Thiers. Mad River Press, Eureka, California. 32 pp. +How to identify mushrooms to genus. III: microscopic features. 1977. By David L. Largent, David Johnson and Roy Watling. Mad River Press, Eureka, California. 148 pp., illus. +How to identify mushrooms to genus. IV: keys to families and genera. 1977. By Daniel E. Stuntz. Mad River Press, Eureka, California. 94 pp. *Identifying grasses: data methods and illustrations. 1977. By H. T. Clifford and L. Watson. University of Queensland Press (Distributed by Prentice-Hall, Herts, England). 146 pp. £ 11.20. Introduction to world vegetation. 1978. By A. S. Collin- son. Allen and Unwin, Boston. 202 pp., illus. Cloth US $16.25; paper US $8.75. +Lehrbuch der Pflanzenphysiologie: 3, vollig Neubearbeitete und erweiterte Auflage. 1978. By H. Mohrand P. Schopfer. Springer-Verlag, Berlin. 639 pp., illus. DM 78. A monograph of the genus Jubula Dumortier. 1978. By W.R. Guerke. Cramer, Lohmar, Germany. 144 pp., illus. DM 40. The mosses of arctic Alaska. 1978. By W.C. Steere. Cramer, Lohmar, Germany. 508 pp., illus. DM 150. Nuclear division in the fungi. 1978. Edited by I. Brent Health. Academic Press, New York. 256 pp. US $16. The pine tree book. 1979. By Russel Peterson. Illustrated by Patricia Wynne. Based on the Arthur Ross Pinetum, New York. Brandywine (Canadian distributor Clarke Irwin, Toronto). Cloth $19.50; paper $10.50. Plant cold hardiness and freezing stress. 1978. Edited by P. H. Liand A. Sakai. Academic Press, New York. 428 pp. - US $21.50. Plant life in anaerobic environments. 1978. Edited by D. D. Hook and R. M. M. Crawford. Ann Arbor Science, Ann Arbor, Michigan. x + 564 pp., illus. US $28. The pteridophytes of Kansas, Nebraska, South Dakota and North Dakota, U.S.A. 1979. By A. J. Ott-Petrik. Cramer, Lehr, Germany. 350 pp., illus. cDM 80. Recognition of tree species on aerial photographs. 1978. By L. Sayn-Wittgenstein. Forest Management Institute In- formation Report FMR-X-118. Environment Canada, Ottawa. 97 pp., illus. Free. The species problem in Cannabis: science and semantics. 1979. Ernest Small. 2 Volumes. Corpus, Toronto. 224 and 160 pp. Cloth $16.95 and $14.95 or $28 set; paper $10.95 and $9.95 or $18 set. 352 A taxonomic monograph of Equisetum subgenus Equise- tum. 1978. By R. L. Hauke. Cramer, Lohmar, Germany. 72 pp., illus. DM 40. *Vascular plant families. 1977. By James Payne Smith, Jr. Illustrations by Katheryn E. Simpson. Mad River Press, Eureka, California. 320 pp., illus. US $7.85. Water deficits and plant growth. 1978. Edited by T.T. Kozlowski. Academic Press, New York. 320 pp. US $31.50. Environment tAnalysis of ecological systems. 1979. Edited by David J. Horn, Rodger D. Mitchell, and Gordon R. Stairs. Ohio State University Press, Columbus. 1x +312 pp., illus. US $27.50. Applications of ecological (biophysical) land classification in Canada. 1979. Edited by C. D. A. Rubec. Proceedings of the Second Meeting Canada Committee on Ecological (Biophysical) Land Classification, 4-7 April 1978, Victoria. Ecological Land Classification Series Number 7. Lands Directorate, Environment Canada. Supply and Services Canada, Hull.. xxix + 396 pp., illus. $7. in Canada; $8.40 elsewhere. Biogeography and adaptation: patterns of marine life. 1978. By G. J. Vermeij. Harvard Press, Cambridge, Massachu- setts. xvi + 332 pp., illus. US $25. + Catalogue of type invertebrate, plant, and trace fossils in the Royal Ontario Museum. 1978. By J. Waddington, P. H. von Bitter, and D. Collins. Life Sciences Miscellaneous Publications. Royal Ontario Museum, Toronto. 151 pp. Paper $7.50. +Contrasts in behavior. 1979. Edited by Ernst S. Reese and Frederick J. Lighter. Wiley, Somerset, New Jersey. x1 + 406 pp., illus. US $27.50. The dark range: a naturalist’s night notebook. 1978. By David Rains Wallace. Sierra Club, San Francisco. xii + 132 pp., illus. Cloth US $15; paper US $8.95. Discovery processes in modern biology. 1979. By W.R. Klemm. Krieger, New York. US $14.95. The economics of environmental quality. 1978. By E. S. Mills. Norton, New York. 304 pp., illus. US $11.95. }Energy, economic and ecological relationships for Gotland, Sweden: a regional systems study. 1978. By A. M. Jansson and J. Zucchetto. Ecological Bulletins 28. NFR, Stockholm. 152 pp., illus. Paper SwCr 70 (c$16). Fungi, man and his environment. 1978. By H. J. Brodie. University of Toronto Press, Toronto. xii + 132 pp., illus. $10. Georgian Bay Islands National Park integrated resource survey. 1978. By R. Hirvonen and R. A. Woods. Forest THE CANADIAN FIELD-NATURALIST Vol. 93 Management Institute Information Report FMR-X-117. Environment.Canada, Ottawa. vill + 82 pp. + appendices, illus. Free. Joy of nature: how to observe and appreciate the great outdoors. 1979. By Reader’s Digest, Montreal. 352 pp., illus. $16.96. {Muskeg and the northern environment in Canada. 1977. Edited by N. W. Radforth and C. O. Brawner. By the Muskeg Committee, NRC Associate Committee on Geo- technical Research. University of Toronto Press, Toronto. 399 pp., illus. $35. The natural history of the land of the bible. 1978. By A. Alon. Doubleday, New York. 276 pp. US $12.95. {Naturalist’s seashore guide: common marine life of the northern California coast and adjacent shores. 1978. By G. J. Brusca and R. C. Brusca. Mad River Press, Eureka, California. 215 pp., illus. US $8.95. *Nature detective. 1978. By Hugh Falkus. Gollancz (Cana- dian distributor Clarke Irwin, Toronto). 256 pp., illus. $22.50. Nature’s oddballs. 1978. By L. Zappler, Doubleday, New York. vill + 148 pp. US $5.95. Oil, ice and climate change: the Beaufort Sea and the search for oil. 1978. By Allen Milne. Edited by R. J. Childerhose. Institute of Ocean Sciences, Sidney, British Columbia. 103 pp., illus. Paper $2.50. Operating manual for spaceship earth. 1978. By R. B. Fuller. Dutton, New York. 143 pp. US $2.95. Pond littoral ecosystems. 1978. Edited by A. Dykyjova and J. Dvet. Springer-Verlag, New York. 480 pp. US $49. Population regulation. 1978. Edited by R.H. Tamarin. Benchmark Papers in Ecology, Volume 7. Dowden, Hutchin- son and Ross, Stroudsburg, Pennsylvania. xvi + 390 pp., illus. US $27.50. St. Lawrence Islands National Park and surrounding areas integrated resource survey. 1978. By R. Hirvonen and R. Woods. Forest Management Institute Information Report FMR-X-114. Environment Canada, Ottawa. viii + 57 pp. + appendices, illus. Free. Miscellaneous The aurora borealis. 1979. By S. I. Akasofu. Alaska Geo- graphic Volume 6, Number 2. Alaska Northwest, Anchor- age. 96 pp., illus. US $9.50. Beyond the green revolution: the ecology and politics of global agricultural development. 1979. By Kenneth A. Dahlberg. Plenum, New York. c215 pp. US $17.95 + 20% in Canada. 1979 Camping and back packing: a guide to information sources. 1979. Edited by Cecil F. Clotfelter and Mary Clotfelter. Gale, Detroit. c350 pp. US $22. Diary of John Clifford Higgins, 1906-1914: a diary of observations on birds. 1978. Edited by William W. Judd. Phelps, London, Ontario. 68 pp. $5. Early naturalists and natural history societies of London, Ontario. 1979. By Wiliam W. Judd. Phelps, London. 216 pp., illus. $7.50. Energy from the waves. 1979. By D. Ross. Pergamon, Elmsford, New York. 80 pp., illus. Cloth US $15; paper US $7.50. Hierarchical analysis of water resources systems: modeling and optimization of large scale systems. 1977. By Y. Y. Haimes. McGraw-Hill, Hightstown, New Jersey. 478 pp. US $39.50. Introduction to marine pollution control. 1979. By Jerome Williams. Wiley-Interscience, Somerset, New Jersey. c256 pp. c US $30. *Introduction to mathematics for life scientists. 1976. By E. Patschelet. 2nd edition. Springer-Verlag, New York. 643 pp. Paper US $11.95; cloth US $27.60. +Migratory game bird hunters and hunting in Canada. 1978. Edited by H. Boyd and G. H. Finney. Canadian Wildlife Service Report Series Number 43. Supply and Services Canada, Ottawa. 125 pp., illus. $7.50 in Canada; $9 elsewhere. The naturalists: pioneers of natural history. 1979. By Alan K. Jenkins. Nelson, Toronto. 185 pp., illus. $17.95. BOOK REVIEWS 353 +The outer shores: part 1, Ed Richetts and John Steinbeck explore the Pacific coast. 1978. Edited by Joel W. Hedg- peth. Mad River Press, Eureka, California. xii + 128 pp., illus. +Pest management. 1978. Edited by G. A. Nortonand C. S. Holling. Proceedings of a Conference, October 1976, Austria. Pergamon, Oxford. 358 pp., illus. US $40. Phosphorus in the environment: its chemistry and bio- chemistry. 1978. By numerous authors. Ciba Foundation Symposium 57 (New Series). Elsevier, Amsterdam. c300 pp., illus. Dfl. 72.60. Resources and planning. 1979. Edited by B. Goodalland A. Kirby. Pergamon, Elmsford, New York. 373 pp., illus. Cloth US $33; paper US $15. Sun: mankind’s future source of energy. 1979. Edited by F. deWinter. Pergamon, Elmsford, New York. 3 volumes, 2250 pp. US $250. The tides of the planet Earth. 1978. By Paul Melchior. Pergamon, New York. 609 pp. US $75. Yukon-Kuskokwim Delta. 1979. By various authors. Alaska Geographic Volume 6, Number |. Alaska Northwest, Anchorage. 96 pp., illus. US $11.95. Wood as an energy resource. 1978. By D. A. Tillman. Academic Press, New York. xiv + 252 pp., illus. US $13.50. *Assigned for review +Available for review Call for nominations for the Council of The Ottawa Field-Naturalists’ Club A Nominating Committee has been chosen by the Council to nominate persons for election to offices and membership of the Council for the year 1980, as required by the Constitution. Club members may also nominate candidates as officers and other members of Council. Such nomi- nations require the signatures of the nominator and seconder, and a statement of willingness to serve in the specified position by the Nominee. Nominations should be sent to the Nominating Committee, The Ottawa Field-Naturalists’ Club, Post Office Box 3264, Postal Station C, Ottawa, Ontario K1Y 4J5, to arrive no later than 15 November 1979. The Committee will also consider any suggestions for nominees which members wish to submit to it by | November 1979. It would be helpful if some relevant background on the proposed nominees were provided along with the suggested names. Chairman, Nominating Committee Instructions to Contributors Content The Canadian Field-Naturalist is a medium for the publication of scientific papers by amateur and professional naturalists or field-biologists reporting observations and results of investigations in any field of natural history provided that they are original, significant, and relevant to Canada. All readers and other potential contributors are invited to submit for consideration their manuscripts meeting these criteria. As the journal has a flexible publication policy, items not covered in the traditional sections (Articles, Notes, Letters, News and Comment, and Book Reviews) can be given a special place provided they are judged suitable. Readers are encouraged to support regional, provincial, and local natural history publications as well by submitting to them their reports of more restricted signifi- cance. Manuscripts Please submit, in either English or French, three complete manuscripts written in the journal style. The research reported should be original. It is recommended that authors ask qualified persons to appraise the paper before it is submitted. Also authors are expected to have complied with all pertinent legislation regarding the study, disturbance, or collection of animals, plants, or minerals. Type the manuscript on standard-size paper, if possible use paper with numbered lines, double-space throughout, leave generous margins to allow for copy marking, and number each page. For Articles and Notes provide a bibliographic strip, an abstract, and a list of key words. Articles also require a running head. Generally words should not be abbreviated but use SI symbols for units of measure. Underline only words meant to appear in italics. The names of authors of scientific names should be omitted except in taxonomic manuscripts or other papers involving nomen- clatural problems. Authors are encouraged to use “proper” common names (with initial letters capitalized) as long as each species is identified by its scientific name once. Although we prefer the names of journals in the Literature Cited to be written out in full, these may be abbreviated following the Bibliographic Guide For Editors & Authors, The American Chemical Society, Washington, D.C. (1974). Unpublished reports should not be cited here. Next list the captions for figures (numbered in arabic numerals and typed together on a separate page) and present the tables (each Reviewing Policy of The Manuscripts submitted to The Canadian Field-Naturalist are normally sent for evaluation to an Associate Editor(who reviews it himself or asks another qualified person to do so), and at least one other reviewer, who is a specialist in the field, chosen by the Editor. Authors are encouraged to suggest names of suitable referees. Reviewers are asked to give a general appraisal of the manuscript followed by specific titled, numbered consecutively in arabic numerals, and placed ona separate page). Mark in the margin of the text the places for the figures and tables. Extensive tabular or other supplementary material not essential to the text, typed neatly and headed by the title of the paper and the authors name and address, should be submitted in duplicate on letter-size paper for the Editor to place in the Depository of Unpublished Data, CISTI, National Research Council of Canada, Ottawa, Canada KIA 0S2. A notation in the published text should state that the material is available, at a nominal charge, from the Depository. The Council of Biology Editors Style Manual, 4th edition (1978) available from the American Institute of Biological Sciences, is recommended as a guide to contributors. Webster’s New International Dictionary and le Grand Larousse Encyclopédique are the authorities for spelling. Tllustrations—Photographs should have a glossy finish and show sharp contrasts. Photographic reproduction of line drawings, no larger than a standard page, are preferable to large originals. Prepare line drawings with India ink on good quality paper and letter (don’t type) descriptive matter. Write author’s name, title of paper, and figure number on the lower left corner or on the back of each illustration. Special Charges Authors must share in the cost of publication by paying $45 for each page in excess of six journal pages, plus $5 for each illustration (any size up toa full page), and up to $45 per page for tables (depending on size). Reproduction of color photos is extremely expensive: price quotations may be obtained from the Business Manager. When galley proofs are sent to authors, the journal will solicit on a voluntary basis a commitment, especially if grant or institutional funds are available, to pay $45 per page for all published pages. Authors may also be charged for their changes in proofs. Limited journal funds are available to help offset publica- tion charges to authors with minimal financial resources. Requests for financial assistance should be made to the Editor when the manuscript is submitted. Reprints An order form for the purchase of reprints will accompany the galley proofs sent to the authors. Canadian Field-Naturalist comments and constructive recommendations. Almost all manuscripts accepted for publication have undergone revision—sometimes extensive revision and reappraisal. The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. TABLE OF CONTENTS (concluded) Eastern limit of the Five-lined Skink, Eumeces fasciatus, in Ontario RICHARD D. USSHER and FRANCIS R. COOK Barn Owls in Quebec NORMAND DAVID Communal roosting of Song Sparrows under snowbank MARTIN K. MCNICHOLL Status of Eastern White Cedar, Thuja occidentalis, in western Nova Scotia GORDON S. RINGIUS Snowy Egret in the Northwest Territories RICHARD W. QUINLAN Letters Tribute to Hoyes Lloyd, 1888-1978 DAvID A. MUNRO News and Comment Book Reviews Zoology: Animals in peril: a guide to the endangered species of Canada and the United States — The great arc of the wild sheep — A guide to the birds of Venezuela — North American Moose — Wild geese — Crows of the world — Life of the Harp Seal. Botany: Edible garden weeds of Canada — Ferns of the Ottawa district — Dyes from lichens and plants: a Canadian dyer’s guide. Environment: Human activity and the Environment — Vermillion Lakes Banff National Park: an introductory study. Miscellaneous: Weather almanac — Introduction to mathematics for life scientists New Titles Mailing date of previous issue 23 April 1979 1979 Council — The Ottawa Field-Naturalists’ Club President: Roger Taylor Elisabeth Beaubien Fran Goodspeed hah Charlie Beddoe Peter Hall Vice-President: Courtney Gilliatt Ron Bedford Hue MacKenzie Treasurer: Barry Henson Frank Bell Frank Pope Bill Cody lan Sneddon Recording Secretary: Diana Laubitz Jane Diceman Ken Strang : : Ellaine Dickson Ken Taylor C ov ; orresponding Secretary: Valerie Hume Honey icieon eee ee Past President: Roger Foxall Marc Forget 321 323 325 326 329 330 331 333) 7/ 359 344 347 348 350 Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. For information on Club activities telephone (613) 722-3050. THE CANADIAN FIELD-NATURALIST Volume 93, Number 3 1979 Editorial To a Bigot LORRAINE C. SMITH Articles Seasonal growth, food and feeding habits of young-of-the-year Black Crappie in the Ottawa River JOHN MARK HANSON and S. U. QADRI Nesting and food-storage behavior of Peromyscus maniculatus gracilis and P. leucopus noveboracensis C. CRAIG TADLOCK and HAROLD G. KLEIN Life history characteristics of Little Brown Bats (Myotis lucifugus) in Alberta DAVID B. SCHOWALTER, JOHN R. GUNSON, and LAWRENCE D. HARDER Winter denning of the Striped Skunk in Alberta JOHN R. GUNSON and RONALD R. BJORGE Range extensions of vascular plants in northern Yukon Territory and northwestern District of Mackenzie JOHN A. NAGY, ARTHUR M. PEARSON, BERNARD C. GOSKI, and WILLIAM J. CODY Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, Ontario HUGH R. MACCRIMMON and ROBERT W. PUGSLEY Song pattern of the Cypress Hills population of White-crowned Sparrows M. Ross LEIN Nesting biology and development of young in Ontario Black Terns ERICA H. DUNN Summer, autumn, and winter diets of Elk in Saskatchewan - HUGH M. HUNT Taxonomy, distribution, and ecology of the cliff-brake ferns ( Pellaea: Polypodiaceae) in Alberta DANIEL F. BRUNTON Characteristics of Peregrine Falcons migrating through central Alberta, 1969-1978 DIcK DEKKER Notes Extralimital record of a Narwhal (Monodon monoceros) in Hall’s Bay, Newfoundland BorRA MERDSoyY, JON LIEN, and ANNE STOREY First Canadian record of the Black Buffalo (Osteichthyes: Catostomidae) E. J. CROSSMAN and S. J. NEPSZY Thaspium trifoliatum (Meadow-parsnip) in Canada P. W. BALL Largest Gray Wolf skulls found in Alberta JOHN R. GUNSON and RONALD M. NOWAK Aquatic feeding by a Woodchuck D. FRASER Dispersion of freshwater leeches (Hirudinoidea) to Anticosti Island, Quebec RONALD W. DAVIES Relative efficiencies of Museum Special, Victor, and Holdfast traps for sampling small mammal populations ARTHUR M. MARTELL Probable hybrids of Cinnamon X Blue-winged Teal from southern Alberta D. V. WESELOH and LINDA MCKEANE WESELOH Common Garter Snake predation on Ring-billed Gull chicks PETER M. FETTEROLF The Rock Vole (Microtus chrotorrhinus) as a Transition Zone species GORDON L. KIRKLAND, Jr. and CHARLES M. KNIPE Zl 232 316 ST) 319 concluded on inside back cover ISSN 0008-3550 aay The CANADIAN ges HARV ote A i UNIVERSI Ty FIELD-NATURALIST Published by THE OT TAWA FIELD-NAT - URALISTS’ CLUB , Ottawa, Canada The CANADIAN FIELD-NATURALIST ublished by THE INA Sis 2 5 awa, a D-NATI STS -D-NA RALI CLUB Pi THE OTTAWA FIEI Ottawa, Canada Contents " Hibernation of bats in southeastern Ontario and adjacent Quebec. By Harold B. Hitchcock 47 “Christmas bird eas 8 rn 59 . Propo als in potanical nomenclature. By Bernard Bova 66 Apparent observations of the whooping crane gn Central Saskatchewan. By Lawrence Ho Walkinshaw as 73 A new Antennaria from Sorthern Ungava- By A. E. Fors d 80 "Salix petioloris 4. Smith in Edmonton Pisirict of Alberts. By George ovum Be $2 The accurrence oi the leech Batrachabdella picta (Verrill) in the dorsal sub-cutane- ‘ous lymph spaces of Rana catesbiana. By Laurence R. Richardson - 85 ‘Squirrel cache of funei By George A. mady se Present state of the chestnut, Castanea dentata (Marsh.) Borkh., 1» Ontario. C By W. Sherwood De Oe 3s A puzzling Labrador reference. By W. L. MeAtee a9 Notes and Observations: — : | . “Breeding of the Sora in New Bumswick. By George F. Boyer de 90 . Ground-nesting robins. By W- H. Whitehead” oe pe OL Rana p. pipiens. By T. 9. Norris-Elye. < 91 ‘Breeding killdeers in Northern New Brunswick. ‘ By Austin Ww. Cameron. 92 Book Reviews Oe ee 92 Published by the 7 OTTAWA FIELD-NATURALISTS CLUB Entered at ‘the Post Office at Olawe, Ont., as second class matter, ‘ Volume 91, Number 1 January-March 1977 Centennial Year Volume 93, Number 4 October-December 1979 The Ottawa Field-Naturalists’ Club FOUNDED IN 1879 Patrons Their Excellencies the Governor General and Mrs. Edward Schreyer The objectives of this Club shall be to promote the appreciation, preservation and conservation of Canada’s natural heritage; to encourage investigation and publish the results of research in all fields of natural history and to diffuse information on these fields as widely as possible; to support and cooperate with organizations engaged in preserving, maintaining or restoring environments of high quality for living things. The Members of Council are listed on the inside back cover. The Canadian Field-Naturalist The Canadian Field-Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas expressed in this journal, however, are private and do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency. Editor: Lorraine C. Smith Assistant to the Editor: Donald A. Smith Book Review Editor: J. Wilson Eedy Associate Editors Gy Dy Bird A. J. Erskine George H. La Roi E. L. Bousfield Charles Jonkel David P. Scott Francis R. Cook Charles J. Krebs Stephen M. Smith W. O. Pruitt, Jr. Copy Editor: Marilyn D. Dadswell Chairman, Publications Committee: J. K. Strang Production Manager: Pauline A. Smith Business Manager: W. J. Cody Subscriptions and Membership Subscription rates for individuals are $10 per calendar year. Libraries and other institutions may subscribe at the rate of $20 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $10 includes a subscription to The Canadian Field- Naturalist. Subscriptions, applications for membership, notices of changes of address, and undeliverable copies should be mailed to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada KIY 4J5. Second Class Mail Registration No. 0527 — Return Postage Guaranteed. Back Numbers Most back numbers of this journal and its predecessors, Transactions of The Ottawa Field- Naturalists’ Club, 1879- 1886, and The Ottawa Naturalist, 1887-1919, may be purchased from the Business Manager. Business Manager: Mr. W. J. Cody, Box 3264, Postal Station C, Ottawa, Ontario, Canada KIY 4J5 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 Address manuscripts on birds to the Associate Editor for Ornithology: Dr. A. J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick EQOA 3C0 All other material intended for publication should be addressed to the Editor: Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0 Urgent telephone calls may be made to the Editor's office (613-996-5840), the office of the Assistant to the Editor (613-231- 4304), or their home on evenings and weekends (613-836-1460), or to the Business Manager's office (613-995-9461). Cover: Since The Ottawa Field-Naturalists’ Club was founded in 1879, it has published a scientific journal. Covers of earlier issues of The Canadian Field- Naturalist are illustrated: the contents were listed on covers from 1945-1969: the larger format with a photograph was introduced in 1970. The Canadian Field-Naturalist Volume 93, Number 4 October-December 1979 Some New and Interesting Vascular Plant Records from Northern Ontario J. L. RILEY Department of Botany, Royal Ontario Museum, Toronto, Ontario MS5S 2C6 Riley, J. L. 1979. Some new and interesting vascular plant records from northern Ontario. Canadian Field-Naturalist 93(4): 355-362. Locations and habitats are described for recent additions to the native arctic-subarctic flora of Ontario, and for selected subarctic species which had been previously known from only one or two sites. Additions to the provincial flora are Agrostis borealis, Aster alpinus, Calamagrostis deschampsiodes, Carex bigelowii, Hierochloe alpina, Kobresia myosuroides, Luzula confusa, Minuartia groenlandica, Phyllodoce coerulea, and Vahlodea atropurpurea. Comments are offered on the occur- rence of more widespread species such as Hierochloe pauciflora, Kobresia simpliciuscula, Pinguicula villosa, Agropyron violaceum, Ledum decumbens, and arctic-subarctic species of the genus Pedicularis. Cypripedium calceolus var. plani- petalum is also recorded. Key Words: northern Ontario, vascular plants, arctic-subarctic, phytogeography, mapping, rare or restricted. During fieldwork directed towards a compilation of the vascular flora of the Hudson Bay Lowland, over 350 different Ontario sites have been visited. Many species new to this flora have been documented. The most interesting group are those restricted to the Hudson Bay area. The purpose of this paper is to report and discuss these new records and the distributions of restricted species now recorded from significantly more sites, in order to provide a useful data base from which to speculate on the character and extent of Ontario’s “maritime tundra.” Species Accounts Agrostis borealis Carex bigelowii Hierochloe alpina Luzula confusa Minuartia groenlandica Kenora District: 54°28’N, 84°54’W:; summit of ab- ruptly rising Precambrian instrusive cuesta west of Hawley Lake, Birch Hill. Riley, 29 July 1978. 54°18’N, 84°30’W: summit of ab- ruptly rising Precambrian intrusive cuesta east of Aquatuk Lake, elevation 900+ ft. [275 m] asl Riley, 29 July 1978. The Sutton Ridges are a discontinuous series of Precambrian outcrops of various elevations and exposures extending across the northeastern Hudson Bay Lowland from the Nowashe River watershed northwestward to within 20 km of the Hudson Bay coast just east of Winisk. These prominent outcrops (Figure 1) are Precambrian diabase-gabbro sills overlying sedimentary Precambrian rocks consisting of a lower dolomitic unit overlain by a unit consisting of iron formations, greywacke, and other materials (Bostock 1968). Two of the most prominent of these, Birch Hill and Aquatuk Lake, had open rock barren summits supporting arctic species characteristic of such Precambrian habitats in northern Quebec. Other outcrops, such as at the Sutton-Hawley Lakes Narrows and near the Hudson Bay coast (55°N, 84°43’W) apparently do not support such species, although the flora of the former includes arctic- subarctic species such as Potentilla nivea (known elsewhere in Ontario only from near the lower. Shagamu River: Sims 2258; SSM, TRT) and from Cape Henrietta Maria (CAN) (Figure 1). As well as providing suitably exposed upland outcrop habitat, these summits were the first islands to emerge from the postglacial Tyrrell Sea in the northeastern Lowland, about 6000 yr BP (Webber et al. 1970). As a result, they have been available to the long-distance dispersal of plants for a much longer period of time than any of the surrounding Lowland except for areas far to the south and west, where none of these species now persist. Agrostis borealis (Riley 9511, 9464) is a circum- polar low arctic species characteristic of “dry rocky and turfy places on acid rock” (Porsild 1957) and of “late-snow-free areas” (Hultén 1968). It was mapped by Porsild (1964) from the northeastern Hudson Bay Lowland but without apparent voucher reference 355) 356 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE |. The Sutton Ridges, a series of Precambrian outcrops penetrating the Paleozoic wetlands characteristic of the Hudson Bay Lowland. This Aquatuk Lake ‘cuesta’ has an elevation ca. 275 m asl. (CAN, DAO, TRT) nor with any mention made of it in his northern Ontario fieldnotes (CAN). It is not included by Scoggan (1978) in the flora of Ontario. The specimens collected (CAN, TRT) are typically awned, with the narrow leaves of var. borealis. It is also known from a third Ontario location (55°N, 84°43’W: abandoned Mid-Canada Line radar station 510, occupying a small inlier of Precambrian diabase- gabbro rock; Sims 2462A, 10 August 1978 (SSMF, Xerox TRT)) (Figure 2). Carex bigelowii ( Riley 9536, 9460) (Figure 2) is an amphi-Atlantic member of an arctic circumpolar Species complex (refer map, Hultén 1964), charac- teristic of dry tundra and rocky habitats. Although also mapped by Porsild (1964) from the northeastern Hudson Bay Lowland, Scoggan (1978) reports it absent from the Ontario flora. Again, no reference is made to it in Porsild’s fieldnotes from northern Ontario (CAN), nor is it represented in major Ontario herbaria (CAN, DAO, TRT). The Birch Hill (CAN, TRT) and Aquatuk Lake (TRT) specimens are typical; collections from outcrop near the Hudson Bay coast (55°04’N, 84°50’W; Riley 9666, 31 July oe ae Luzula confusa Carex bigelowii ee Agrostis borealis _ ,» Minuartia groenlandica Pay Ty & eA 5 r . 0 2 4 6 80 100 M1 SS o @ 8 I~ wo Km ‘ ( te a *. yl at FIGURE 2. Ontario distribution of Luzula confusa, Carex bigelowii, Agrostis borealis, and Minuartia groen- landica. 1979 1978 (TRT)) and from willow thickets near North Point, James Bay (51°28’N, 80°27’W; Tessier, 15 July 1976 (TRT), det. Ball) confirm Porsild’s (1957) observation that the species’ habit is “more lax in wetter sites” (Figure 2). Hierochloe alpina (Riley 9458) was collected only on the summit of the outcrop at Aquatuk Lake(CAN, TRT) (Figure 3). Specimens collected belong to the subspecies orthantha, having untwisted awns attached above the middle of the second glume (Weimarck 1971). This taxon occurs from Greenland to the east coast of Hudson Bay and James Bay south to several mountain areas of New England and upper New York State. It reoccurs sporadically in Beringia, but this collection represents the western limit of its major range; the closest westward location, at Cape Churchill, is of ssp. a/pina. This collection represents the first Ontario record of this circumpolar arctic- alpine species. John Macoun’s reference (1888) to its occurrence at Moose Factory, James Bay, is un- substantiated and highly improbable (Dutilly et al. 1954) (Figure 3). Luzula confusa (Riley 9512, 9467) is a circum- polar arctic species, occurring southward in the Canadian Rockies and ona few mountain tops in New England, in dry turfy tundra heath, and on rocky slopes and ledges (Porsild 1957; Hultén 1968). These first Ontario records (TRT) complete the distribu- tional gap between occurrences at Churchill, Mani- toba and the northeastern shore of James Bay, Quebec (Figure 2). Minuartia groenlandica (Arenaria g.) ( Riley 9526, 9471), the attractive Greenland sandwort, is an 82° 80 100 MI 120 160 Km e@ Hierochloe alpina a ~ Agropyron violaceum % Calamagrostis deschampsioides %t Aster alpinus FIGURE 3. Ontario distribution of MHierochloe alpina, Agropyron violaceum, Calamagrostis deschamp- sioides, and Aster alpinus. RILEY: NEW NORTHERN ONTARIO PLANTS 357 eastern North American subarctic endemic, occurring from Greenland, southwestward through Newfound- land, Nova Scotia and in New England and northern New York State mountains, northwestward to the east coast of Hudson Bay north of James Bay, and now known as far westward as the Sutton Ridges in Ontario (for Quebec distribution, refer to Rousseau 1974). As with the other arctic-subarctic species disjunct on outcrop summits of the Sutton Ridges, the nearest major ranges of species are across the mouth of James Bay, in the Precambrian uplands of northern Quebec (Figure 2). Agropyron violaceum (Hornem.) Lange (A. /atiglume (Scribn. & Sm.) Rydb.) Kenora District: Vicinity of the mouth of the Severn River. Moir 1416, 14 Aug. 1952 (CAN) (sub A. trachycaulum var. novae-angliae; rev. Porsild 1959). Vicinity of the mouth of Goose Creek. Moir, 18-20 Aug. 1952 (CAN) (sub A. t¢. var. n.-a.: rev. Porsild). 15 mi [25 km] from the mouth of the Black Duck river. Moir 2087, 25-27 July 1953 (CAN) (sub A. t. var. n.-a.; rev. Porsild 1959). 56°00’N, 87°38’W; Fort Severn, near H.B.C. post. Hustich 1256, 28 July 1956 (CAN) (det. Porsild). 55°14’N, 85°05’W; Winisk, disturbed gravel- clay areas near airport. Riley 5872, 24 August 1976 (TRT). 55°14’N, 85°07’W, Winisk airport area, near fuel depot. Riley 6/40, 26 August 1976 (TRT). 55°52’N, 86°46’W; coast of Hudson Bay, im- mediately east of mouth of Shagamu River. Riley 7483, 11 August 1977 (TRT). 55°57’'N, 89°57’W; littoral habitat along Black Duck River. Riley 8693b, 19 July 1978 (TRT). 55°N, 84°43’W: abandoned Mid-Canada Line radar station 510. Sims 2472, 10 August 1978 (SSMF, TRT). 55°04’50”N, 85°39’30”W; upland thicket along river levee. Sims 2/30, 23 July 1978 (SSMF, IRA). This species, first reported for Ontario by Hustich (1957. Comparison of the flora and vegetation in the Fort Severn and in the Big Trout area. 2nd report concerning fieldwork in 1956. Arctic Institute of North America, University Library Tower, 2920- 24th Ave., N.W., Calgary, Alberta), occurs north to Ellesmere Island and is transcontinental from Alaska to Greenland, south to northern Manitoba and Ungava; these specimens represent southern limits for this arctic species (refer map, Porsild 1964). None of the specimens seen have the pubescent glumes of the 358 more northern variety hyperarcticum. At Winisk it occurs in disturbed gravel areas, having colonized them since the air base was built in the 1950s. The other locations are from riverbanks of major spring- flooding rivers, in habitats such as low shrub willow thickets (Riley 7483; Hustich 1957) and graminoid meadows. Such calcareous alluvial sites are among the most floristically rich in the coastal area of southwestern Hudson Bay; the author has recorded up to 60 species within 30 m of closed and open low willow thickets along the lower Shagamu River (Figure 3). Aster alpinus Kenora District: 55°52’N, 86°46’W: coast of Hudson Bay, mouth of Shagamu River. Riley 6972, 29 July 1977 (CAN, TRT). This improbable find was made among the gravel bars at the mouth of the Shagamu River, where it grew with such species as Chrysanthemum bipinnatum ssp. huronense, Epilobium latifolium, and Elymus mollis. It represents the first record of the species between the Great Slave Lake area and the Pyrenees in Europe. The North American phase, ssp. vierhapperi, is widely distributed in the Rocky Mountains through to Siberia (refer map, Hultén 1968) (Figure 3). Calamagrostis deschampsioides Kenora District; 55°44’10”N, 86°19’55”W; SW coast of Hudson Bay, about 15 km SW of the Shagamu River mouth. Sims 2245, 28 July 1978 (SSMF, TRT). This Ontario record for this subarctic circumpolar coastline species helps to confirm its distribution around Hudson Bay, filling a gap between Churchill and York Factory, Manitoba (Scoggan 1957), and northern James Bay and Hudson Bay, Quebec (Rousseau 1974; Hultén 1968). It was found in a brackish marsh dominated by Carex aquatilis and Scirpus hudsonianus, about 0.5 km above the high- tide level, in peat 38 cm deep (Figure 3). Cypripedium calceolus var. planipetalum Kenora District: 52°57’30’N, 83°10’W; limestone outcropping across the Attawapiskat River about 40 mi [65 km] upstream of Attawapiskat townsite. Riley 6328, 3 July 1977 (TRT). 52°52’N, 83°45’W: northward facing lime- stone outcrop ... along south shore Attawa- piskat River. Riley 6573b, 5 July 1977 (TRT). This variety was first described by Fernald (1926) from “turfy limestone barrens,” “peaty . . . lime- stone barrens,” etc., in northwestern Newfoundland. It was distinguished by its broad and short sepal, broad flattish petals, and a more cordate stamino- dium. Luer (1975) recognizes the taxon, although his distribution map inexplicably extends its range from THE CANADIAN FIELD-NATURALIST Vol. 93 Newfoundland up the St. Lawrence River to eastern Ontario. It was found growing in open limestone White Spruce (Picea glauca) forest along the lower Attawa- piskat River, associated with Juniperus horizontalis, J. communis, Alnus crispa, and Arctostaphylos uva- ursi. In the second site, it was found close to var. pubescens (Willd.) Corr. It is suggested that the status of this taxon requires closer consideration, especially in sites in which it is sympatric with other intraspecific taxa. Hierochloe pauciflora Kenora District: 54°40’N; 34 mi [55 km] au sud du cap Henrietta Maria. Dutilly, Lepage 3141/1, 17-18 August 1953. (Dutilly et al. 1954). 55°07’N; Cap Henrietta Maria. Dutilly, Lepage 31124, 10 August 1953. (Dutilly et al. 1954). 55°52’N, 86°48’W; mouth of Shagamu River, two miles inland, one mile west of river. Riley 294, 3 August 1977 (TRT). 56°47’N, 88°53’W: West Pen Island. Riley 8438, 16 July 1978 (TRT). 55°N, 82°18’W; Cape Henrietta Maria. Riley 9292, 27 July 1978 (TRT). 55° 14’N, 84°49’10”W; SW coast of Hudson Bay, approx. 10 km E of Winisk, 1.25 km from coast. Sims 2417, 9 August 1978 (SSMF, Xerox TRIP). This high arctic species (Porsild 1957), circumpolar except for Greenland and Scandinavia, is a wet tundra associate. Porsild’s observation (1957), that the species is, “although not littoral, a distinctly coastal and lowland species,” is apparently true along Ontario’s Hudson Bay coast. It is found in mossy inter-ridge tundra situations, such as at Cape Hen- rietta Maria, and in inter-ridge mossy willow thickets. It appears to achieve highest dominance values in open graminoid fen-meadows close to the coast with high pH values (6.2 and 7.2 recorded) and shallow peats (30 to 45cm over permafrost or silts). One collection (Riley 8438) was made in a graminoid association dominated by Ca/amagrostis stricta, with Hierochloe pauciflora having cover value approach- ing 10%. Other species present at this site included Carex rariflora, Pedicularis sudetica, Salix candida, S. brachycarpa, and Saxifraga hirculus (Figure 4). Kobresia myosuroides (Vill.) Fiori & Paol. (K. bellardii (All.) Degl.) Kenora District: 55°52’N, 86°46’W; coast of Hudson Bay, immediately east of mouth of Shagamu River. Riley 74806, 11 August 1977 (TRT). 55° 22’35”N, 85° 22’50” W; SW coast of Hud- son Bay approx. 7 km NW of Winisk, 2 km from 84° 82 Hierochloe pauciflora coast. Sims 2180b (mixed coll’n with Carex bicolor All.), 26 July 1978 (TRT). Easily overlooked, this tiny sedge is undoubtedly more widespread along Ontario’s Hudson Bay coast. A circumpolar arctic-subalpine species occurring to northernmost Ellesmere Island, it is “a pronounced calciphile, preferring dry, sandy heath and windswept ridges” (Porsild 1957). The Shagamu River collection was from a silted, riparian low willow thicket association, while the Winisk collection was from an immature low shrub thicket with shallow peat (7 cm) , and a dense graminoid ground cover including Carex microglochin, C. rariflora, C. capillaris, Scirpus cespitosa, and Kobresia simpliciuscula (Figure 5). 2 0 20 © 6 80 100 Mi ee oe o 0 44 80 120 160 Km 4 Kobresia myosuroides e K. simpliciuscula FIGURE 5. Ontario distribution of Kobresia myosuroides and K. simpliciuscula. RILEY: NEW NORTHERN ONTARIO PLANTS 358 Kobresia simpliciuscula Kenora District: 55°07’N; Cap Henrietta Maria. Dutilly, Lepage 31135, 10 August 1953 (Dutilly et al. 1954). 54°34’N, 84°38’W; Hawley Lake, Sjors, 1961 (Lepage 1966). 54°48’N, 82°23’W; Runway 415, south of Cape Henrietta Maria, Riley 5850, 23 August 1976 (TRT). 56°22’N, 88°08’W. Riley 8465, 16 July 1978 (TRT). 55°29’10”N, 85°41’1S5”W: SW coast of Hudson Bay, approx. 16km NW of Winisk, 7.25 km from coast. Sims 2043, 20 July 1978 (SSMF, TRT). 55° 22’35”N, 85°22’50”W; SW coast of Hud- son Bay, approx. 7 km NW of Winisk, 2 km from coast. Sims 2182, 2185, 26 July 1978 (SSMF, (TRT). 55°02’N, 82°41’W. Riley 9169, 27 July 1978 (TRT). 54°47’'N, 82°39'W. Riley 9358, 28 July 1978 (TRT). 54° 50’N, 82°46’W. Riley 9372, 28 July 1978 (TRT). 55° 12’N, 84° 17’'W. Riley 9401, 29 July 1978 (TRT). 54° 19’N, 84°30’W. Riley 9476, 29 July 1978 (TRT). 54° 38’N, 84°37’W; slope of Sutton River about | km from the river. Riley 9567, 29 July 1978 (TRT). 54° 38’N, 85° 20’W. Riley 9631, 30 June 1978 @iRsh): 53° 38/Ni 83229. W. Riley, 9737, 3 August 1978 (TRT). The distribution in Ontario (Figure 5) of this circumpolar arctic-alpine sedge is much more wide- spread than indicated by the few early records. The dozen subsequent records are all from calcareous habitats; six were made from open graminoid or open low shrub fens with pH’s between 5.8 and 7.2, peat depths from 7 to 50cm. These fen sites all had a vascular plant ground cover dominated by Scirpus cespitosa, with Carex rariflora, C. livida, C. limosa, Betula glandulifera, Juniperus horizontalis, and Myrica gale as common associates. The most com- mon bryophytes recorded were Scorpidium scor- poides, Tomenthypnum nitens, and Drepanocladus spp. The non-wetland sites in which Kobresia sim- pliciuscula occurs include a variety of ericaceous tundra heath, frost-shattered limestone, frost-sorted till, and algal marl habitats not uncommon through the interior of the Hudson Bay Lowland in an area extending from the lower Ekwan River to the lower Winisk, northeastward to Cape Henrietta Maria. 360 THE CANADIAN FIELD-NATURALIST Current studies in such interior areas confirm that terrestrial arctic habitats are not confined simply to a narrow coastal strip but extend to non-arboreal terrestrial (usually till upland or bedrock outcrop) habitats over a much wider area. Phyllodoce coerulea Kenora District: 55°07’N, 83°52’35”W; SW coast of Hudson Bay approx. 6 km SW of Sutton River mouth, 16 km from coast. Sims 2397a, 4 August 1978 (SSMF, TRT). The closest records of this circumpolar subarctic- alpine mountain heather are from Cape Jones, Quebec (Rousseau 1974) to the east and from the Precambrian Shield of northern Manitoba to the west (Scoggan 1957; Ritchie 1962). It was collected from an upland open White Spruce-lichen woodland, with trees reaching 15 m (230 yr old), shallow lichen peat (less than 10 cm), and a water table ca. 50 cm deep. Lichen woodland on such beachridge complexes appears to burn regularly; those supporting forests older than 200 yr should be considered very rare (Figure 6). Cody (1953) mapped and discussed its North American distribution and concluded that P. coerulea has widely re-occupied its glaciated northeastern range. Its occurrence 16 km from the Hudson Bay coast suggests a maximum age for the migration of the species into Ontario. The site is estimated to have undergone postglacial isostatic uplift over the last 1000 yr of about 1.2 m per century, and to have been above sea level for only about 1500 yr (Webber et al. 1970). 84° 5; : ae Z Phyllodoce coerula A | “( 0 Ledum decumbens Vol. 93 Pinguicula villosa Kenora District: 54°34’N, 84°38’W; Hawley Lake, Sjors, 1961 (Lepage 1966). 56° 16’N, 88°05’W. Riley 8478, 16 July 1978 (TRT). 56° 48’N, 88°52’W. Riley 8485, 16 July 1978 (ERD): 55°48’N, 87°31’W. Riley 9042, 23 July 1978 (TRT). 55°20'05’N, 85°39’15”W; approx. 12 km W of Winisk, 17 km from coast. Sims 2089, 13 July 1978 (SSMF, TRT) (sub. P. vulgaris). 55° 11’N, 85°40’W. Riley 9/42, 26 July 1978 (TRT). 54° 28’N, 84°54’W. Birch Hill. Riley 95/3, 29 July 1978 (TRT). 54° 38’N, 85°20’W. Riley 9633, 30 July 1978 (TRT). This circumpolar subarctic butterwort is not an infrequent acidophile of the fen and bog wetlands developing well back of the Hudson Bay coast (Figure 7). Within open graminoid fen patterns of the lowlands, it is found on bog (sphagnum) hummocks, palsas or on the bog ‘laggs’ or ‘bibs’ of peat plateau patterns. On the summit of Birch Hill, it was found growing beside a confined wet sphagnum pool. f 7h a 2 0 2 40 60 80 100 ml | SSeS eS ee 0 4 80 12 160 Km | we ie fi 3 | Vahlodea atropurpurea Kenora District: 55°07’N, 83°52’35”W; SW coast of Hudson Bay approximately 6 km Sw of Sutton River mouth, 16 km from coast. Sims 2397, 4 August 1978 (SSMF, Xerox TRT) (det. Rez- nicek). This subarctic circumpolar grass is known as a rare tundra species in Manitoba (Scoggan 1957, 1978) and 0 2 4 6 80 00 ml ~” 4 80 120 wo Km Ay F FIGURE 6. Ontario distribution of Phyllodoce coerula and Ledum decumbens. IT) a widespread species of Quebec’s alpine meadows and open conifer forests on sandy soils (Rousseau 1974). This first Ontario record is from an open White Spruce-lichen woodland site also supporting Phyl- lodoce coerulea (see further site comments with that species) (Figure 6). This specimen is of the typical, eastern variety. Recent field work in the Hudson Bay Lowland of Ontario has thus resulted in the addition of 10 vascular plant species to the provincial flora. These are Agrostis borealis, Carex bigelowii, Hierochloe alpina, Luzula confusa, Minuartia groenlandica, Vahlodea atropurpurea, Aster alpinus, Calamagrostis deschampsioides, Kobresia myosuroides, and Phyl- lodoce coerulea. The first five of these species are reported from the highest and most exposed of the outcrop summits of the Precambrian Sutton Ridges. Other than their nearly complete inaccessibility, these two stations do not currently enjoy any protective status. The other species are all from widely dispersed locations and, with the exception of Cypripedium calceolus var. planipetalum (lower Attawapiskat River), all are now known from sites within the Polar Bear Provincial Wilderness Park. Discussion All of the species reported here as new to the provincial flora should be included in the listing of rare vascular plants (Argus and White 1977). Of the other taxa discussed or mapped, Pinguicula villosa, Pedicularis labradorica, and Cypripedium calceolus var. planipetalum should also be listed as rare. Because of the rarity of suitable habitat in Ontario, it appears that some species are undoubtedly rare: Agrostis borealis, Carex bigelowii, Hierochloe alpina, Luzula confusa, Minuartia groenlandica, Phyllodoce coerulea, Aster alpinus, Cypripedium calceolus var. planipetalum. Others are widespread but regionally restricted: Pedicularis spp., Ledum decumbens, Agropyron violaceum, Kobresia simpliciuscula, Pin- guicula villosa. Some are species of greatly under- sampled habitats: Hierochloe pauciflora, Calama- grostis deschampsioides, Kobresia myosuroides. In light of the currently accepted rough equivalence between a ‘regionally restricted’ status and a ‘provin- cially rare’ status, a caution may be in order. The “narrow band of tundra along the Hudson Bay coast” (Argus and White 1977) differs from the Carolinian zone (Ontario’s southern zone of coincident restricted species) in that its wildlife remains largely intact, continuous and unendangered, and in that its geo- graphic extent remains largely undefined, but almost certainly far larger than the Carolinian zone of southwestern Ontario. Mapping of most of Ontario’s arctic-subarctic species confirms a ‘maritime tundra’ RILEY: NEW NORTHERN ONTARIO PLANTS 361 concept of Ontario’s Hudson Bay shore (Figure 4, Hierochloe pauciflora; Figure 8, Pedicularis spp.). The distributions of species such as Kobresia sim- pliciuscula (Figure 5), Pinguicula villosa (Figure 7), and Ledum decumbens (Figure 6), however, indicate a much expanded zone. 2 4 60 80 100 mI 1eo Am Sy Pedicularis flammea P. lapponica P. labradorica P. sudetica 2 ELS ar FIGURE 8. Ontario distribution of Pedicularis flammea, P. lapponica, P. labradorica, and P. sudetica. Ritchie (1962) considered the subarctic zone in Manitoba to be characterized by a vegetation of primarily open conifer forest or a mixture of forest and tundra, and by a climate with a growing season not over 650 degree-days Centigrade and with a mean July temperature not over 14°C. This included all of the Hudson Bay Lowland in Manitoba, as well as the Precambrian Shield westward. Meteorological data from extreme northern Ontario are not presently of sufficient detail to delimit such a climatic zone. The ‘openness’ of Hudson Bay Lowland associations is not a function of dominant lichen woodland associations or of tundra thinning forest sites, but the result of the overwhelming dominance of wetland or peatland vegetation on clay-silt marine substrates. Zonal mapping by winter satellite photography (Hare and Ritchie 1972) shows the Hudson Bay Lowland to have albedo levels higher than, or comparable to wood- lands, but this is not a result of any ‘openness’ of terrestrial systems. For similar reasons, the standard latitudinal zones, as summarized by Scoggan (1978), are not valuable in proposing boundaries for an arctic-subarctic floristic province in northern On- tario. As the collected floristic data base is completed, it is hoped that floristic criteria themselves will define such a zone. °362 THE CANADIAN FIELD-NATURALIST Acknowledgments This work was made possible through the field support and encouragement of several individuals and agencies; S. Pala and A.N. Boissonneau of the Ontario Center for Remote Sensing; R. K. Ross and R. I. G. Morrison of the Canadian Wildlife Service, and D. W. Cowell of the Federal Lands Directorate, Burlington. I especially thank R.A. Sims of the Canadian Forestry Service, Sault Ste. Marie, for sharing an enthusiasm for botanizing this wilderness and for sharing his discoveries. Mapping is based on specimens at CAN, DAO, SSMF, and TRT (1978), and on records as indicated in the text. J. M. Gillett (CAN) and W. J. Cody (DAO) kindly checked herbarium specimens and the field notes of A. E. Porsild (CAN) for references to the species new to Ontario. P. W. Ball of Erindale Campus, University of Toronto, assisted with the identification of some sedge specimens, and J. McNeill of the Biosysematics Research Institute, Agriculture Canada, verified determinations of Agro- pyron violaceum. Review of an earlier draft by R. A. Sims and J. H. Soper was very much appreciated. Literature Cited Argus, G. W. and D. J. White. 1977. The rare vascular plants of Ontario. National Museum of Natural Sciences, National Museums of Canada, Syllogeus Series Number 14. Bostock, H. H. 1968. Precambrian sedimentary rocks of the Hudson Bay Lowlands. /m Earth science symposium on Hudson Bay. Edited by P.J. Hood. Geological Survey of Canada Paper 68-53: 206-214. Cody, W. J. 1953. Phyllodoce coerulea in North America. Canadian Field-Naturalist 67: 131-134. Dutilly, A., E. Lepage,and M. Duman. 1954. Contribution a la flore du versant occidental de la baie James, Ontario. Contribution of the Arctic Institute, Catholic University of America, Number 5F. 144 pp. Fernald, M.L. 1926. Two summers botanizing in New- Vol. 93 foundland. Rhodora 28: 49-63, 74-111, 115-129, 145-155, 161-204, 209-225, 234-241. Hare, F.K. and J.C. Ritchie. 1972. The boreal bio- climates. Geographical Review 62: 333-365. Hultén, E. 1968. Flora of Alaska and neighboring terri- tories. A manual of the vascular plants. Stanford University Press, Stanford, California. 1008 pp. Lepage, E. 1966. Apergu floristique du secteur nord-est de V’Ontario. Naturaliste Canadien 93(3): 207-246. Luer, C. A. 1975. The native orchids of the United States and Canada excluding Florida. New York Botanical Garden, New York. 360 pp. Macoun, J. 1888. Catalogue of Canadian plants. Part IV. Endogens. Geological and Natural History Society of Canada. Dawson Bros., Montreal. Porsild, A. E. 1957 (1964). Illustrated flora of the Cana- dian Arctic Archipelago Bulletin 146, Biological Series — 50, National Museum of Natural Sciences, National Museums of Canada. Reprinted 1973. (Including update reprint: Distribution records as amended to 1964, .. . includes the supplement, etc.) Ritchie, J.C. 1962. A geobotanical survey of northern Manitoba. Arctic Institute of North America, Tech- nical Paper Number 9. 46 pp. + maps. Rousseau, C. 1974. Géographie floristique du Québec- Labrador. Les presses de l'Université Laval, Québec. 798 pp. Scoggan, H.J. 1957. Flora of Manitoba. Bulletin 140, Biological Series 47, National Museum of Natural Sciences, National Museums of Canada. 619 pp. Scoggan, H. J. 1978. The flora of Canada. Volumes 1, 2, 3. Publication in Botany Number 7(3), National Museum of Natural Sciences, National Museums of Canada. 1115 pp. Webber, P.J., J. W. Richardson, and J.T. Andrews. 1970. Post-glacial uplift and substrate age at Cape Henrietta Maria, southeastern Hudson Bay, Canada. Canadian Journal of Earth Sciences 7: 317-325. Weimarck, G. 1971. Variation and taxonomy of Hier- ochloe (Gramineae) in the northern hemisphere. Botani- ska Notiser 124: 129-175. Received 22 January 1979 Accepted 23 April 1979 fags Su ad Yearly Variations in the Population Dynamics of Richardson’s Ground Squirrels GAIL R. MICHENER Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E1 Present address: Department of Biological Sciences, University of Lethbridge, Lethbridge, Alberta T1K 3M4 Michener, G. R. Yearly variations in the population dynamics of Richardson’s Ground Squirrels. Canadian Field-Naturalist 93(4): 363-370. Population fluctuations were followed from 1975 to 1978 in a group of Spermophilus richardsoniiin southern Alberta. Adult males ranged widely in early spring but they restricted the extent of their activities once all females were mated, and consequently fewer males were located on the study area after the breeding season than in early spring. Adult females made only minor changes in the location of their ranges from year to year and the major loss of adult females occurred overwinter. Each year fewer juveniles entered the active population than expected from the number of resident females and their reproductive potential. There was net juvenile immigration in the year with the lowest initial density and net juvenile loss in the years with highest density. Both predation and dispersal contributed to juvenile losses, particularly of males in 1976. The adult sex ratio favored females regardless of the sex ratio of the juvenile population in the previous fall. Telemetric studies are required to distinguish mortality in late summer from entry underground to hibernate and to distinguish death underground during winter from early emergence in spring followed by dispersal or death. Key Words: Ground squirrels, Spermophilus richardsonii, dispersal, overwinter mortality, sex ratio, population dynamics, annual variations. Large scale mark-release-recapture studies, by compare the results obtained by this method with Dorrance (1974), Michener and Michener(1977),and those of previous studies. Schmutz et al. (1979), of the population dynamics of Richardson’s Ground Squirrels, Spermophilus Methods richardsonti, have revealed that there is high loss of The Richardson’s Ground Squirrel population was squirrels between years, that male losses are greater located on fescue grassland in Fire Guard Coulee than female losses, that juvenile losses exceed adult (50°34’N, 114°18’W, elevation 1235 m)5.5 km NW of losses, and that the adult sex ratio is biased infavor of Longview, Alberta. An area of 0.72 ha was gridded females. Discrepancies exist between these workersin with flags at 20-m intervals, and a record was kept of accounting for both the magnitude of losses and the the identity and numbers of squirrels, and of their differential nature of losses. Dorrance (1974) con- location within this area. Squirrels were individually cluded that overwinter mortality and loss of pups marked with a numbered metal tag in each ear and prior to weaning were the main determinants of with black dye marks on the fur. As part of a population size, whereas differential mortality favor- behavioral study conducted in 1975 all squirrels ing adult females in early spring and favoring juvenile caught that year were further identified with a colored females in fall and winter contributed to the biased _ plastic disc attached to one ear tag. adult sex ratio. Michener and Michener (1977) Following initial trapping, which began on 13 April concluded that overwinter mortality was a major 1975, the squirrels on the area were observed for 147 h factor accounting for both the extent of interyear on 55d between 5 May and 11 October; in general, losses and the differential nature of losses in their squirrels were retrapped only when redying of the population. Schmutz et al. (1979) concluded that identifying marks was necessary. In 1976, squirrels overwinter mortality, while reducing population size, | were observed for 32 h on 13 d between 28 Marchand did not account for the disparate sex ratio but that 4 July. Thereafter, the area was trapped at approxi- predation, differentially directed toward dispersing mately 2-wk intervals until 11 October to determine adult males in spring and dispersing juvenile males in | which squirrels were still resident. In 1977 and 1978, summer, did. all data were collected by trapping at approximately The aim of this study was to examine the population _10-d intervals between 20 March and 2 October, and dynamics of a small group of individually identifiable 19 March and 24 April, respectively. Under the squirrels by observation and trapping at more observation regime the area was surveyed 5 times an frequent intervals than used in previous studies inan hour for approximately 3 continuous hours, and all attempt to distinguish residents from transients, to active squirrels were identified by their colored disc follow the fate of each resident squirrel, and to and dye marks. Under the trapping regime an attempt 364 THE CANADIAN FIELD-NATURALIST was made to capture every squirrel present; in 1976 this was done by placing an additional dye mark on each captured squirrel until all squirrels were so marked, and in 1977 and 1978 captured squirrels were held in the live-traps until no other squirrels remained active. As there were relatively few adult squirrels, all of which could also be identified without capture, errors in determining the number of residents were likely only within the juvenile cohort. Juveniles were sometimes not caught during one trapping episode but captured during the next one; such animals were assumed to have been in continuous residence and were included in the population count for the day on which they were not caught. On average, such ‘missed’ squirrels accounted for less than 8% of the juvenile population present during any trapping episode. Squirrels adjacent to the study area were trapped ona sporadic basis each year. Except where otherwise indicated, the term ‘resi- dent’ refers to a squirrel present on the area for a minimum of 2 wk, whose major burrow system was within the boundaries of the area, and which spent the majority of its-active time on any day within the area. Vol. 93 Results Adults Adult males trapped in spring could be classed into three categories: transients which were present for at most a day or two before moving off the area, temporary residents which were present on the area for at least a week but did not remain resident for the summer, and permanent residents which remained on the area beyond the breeding season. Table | indicates the yearly residency status of each male classed, in at least one year, aS a permanent resident or as a temporary resident. In the 4 yr of this study there was only one instance of an adult male remaining as a permanent resident in 2 consecutive years and one instance of a male born on the area remaining resident as a yearling. Three other yearlings that had been born on the area were recovered; one was classed as a transient in 1976 and two as temporary residents in 1977. Although at least half of the males classed as permanent residents had moved on to the area in early spring, there were no instances of adult male immigration during the remainder of the active season. TABLE | —Yearly residency status of adult males classed as permanent or temporary residents in at least one year (see text for criteria of classification). For those males also caught as juveniles their status as a juvenile is given Residency status Male 1975 1976 1977 1978 RO permanent RB permanent RBSS permanent 6201 temporary ; 6210 temporary LW transient temporary LB temporary LR temporary RR permanent temporary ILC born on area permanent transient LG born adjacent* permanent permanent RBSH transient permanent LPSS born on area temporary STS born on area temporary 6404 temporary 72 born off area* transient temporaryt 6418 transient temporary 6477 temporary 6462 born adjacent # permanentt 6487 permanent Total adults Permanent 3 3 2 2 Temporary 2 3 4 3 *Immigrated on to study area as a juvenile. t Alive in 1979. #Immigrated on to study area as an adult in 1978. IOVS All losses of permanently resident males during the summer were attributed to entry into hibernation either because the males were recovered the following year, or because they became progressively fatter and less active, confining their movements to smaller areas in the days immediately preceding their disappear- ance. After their emergence from hibernation, there were always more resident females than males (Tables | and 2). Females did not exhibit a pattern of spring movement and temporary residence, and all adult females were resident on or near the study area, either as juveniles or adults, in the previous year (Table 2). Of the 11 original adult females resident in 1975, 10 remained for the summer, hibernated onthe area, and reappeared in 1976. On 3 June a Badger (Taxidea taxus) invaded the main burrow system of the 11th female and she was not subsequently seen. A neighboring, non-lactating female whose range was adjacent to the study area extended her area to include part of that formerly used by the missing female. This female was not recovered in 1976. Fourteen of the I5 females resident in 1976 remained through April and May and produced litters. The missing female, when last captured on 8 April, had a scarred face, weighed 210 g compared with an average of 276 + 29 g for eight other females caught that day, and probably died in mid-April. Of the remaining 14 residents, six were recovered in 1977 and must have hibernated on the area. Four other females disappeared at a similar time to those that were recovered and apparently hibernated, but did not survive over winter. Four females disappeared in June, too early to have gone into hibernation on the area. Of the 12 females resident in 1977, 11 remained on TABLE 2—Composition of the population of adult females resident on the study area each year Number resident Origin 1975 1976 1977 1978 Resident on area 1975 11 10 3 0 Born on area 1975 5 Ds | Born on area 1976 6 3 Born on area 1977 0 Born adjacent 1975+ | Resident adjacent 1976+ I l Resident adjacent 1977+ I Total permanent residents 11 IS) 12 i *A third 2-yr-old was recovered but she established residency adjacent to the study area. +Females in these categories were captured at least once within 40 m of the study area prior to the year they became resident on the study area. MICHENER: GROUND SQUIRREL POPULATION DYNAMICS 365 the area until the time of entry into hibernation, and five of these were recovered in 1978. The 12th animal, born on the area in 1975, disappeared in late May when her pups were about 3 wk old. Death rather than dispersal is the most likely explanation for her disappearance during the lactation period. All seven females resident in 1978 were still present when this study ended on 24 April. The number of adults hibernating on the area was similar in each year: 3 males and 11 females in 1975, 3 males and 10 females in 1976, and 2 males and 11 females in 1977. Recovery rates in the following spring varied: in 1976, 0 males and 10 females were recovered, in 1977, 3 males and 6 females, and in 1978, 0 males and 5 females. Juveniles The numbers of adult female squirrels that re- mained on the area throughout the breeding, gesta- tion, and lactation periods were 10, 14, and 11 for 1975, 1976, and 1977, respectively. Of these 9, 14, and 8 respectively gave birth, and 6, 12, and 7 reared at least one pup to weaning age. I attributed the complete and partial losses of four litters in 1975 to Long-tailed Weasel (Mustela frenata) predation be- cause I saw a weasel hunting in the natal burrow systems of the mothers of these litters. Two of the mothers subsequently extended their ranges and spent less time in the area of the natal burrow, behavior patterns normally seen after the young are weaned (Michener 1979a), and they reared no young. The other two weaned only one young each. Such losses contributed to the smaller size of emerged litters in NOS: Average litter size + SD (range; N) OWS) 2s) ae\ 5) Cl=s2 G) 1976 4.4 + 1.5 (1-6; 11) 1977 4.0 + 0.6 (3-5; 7) The average litter size at birth for females captured when pregnant from the surrounding grassland in 1976 was 4.9 + 1.0 (3-6; 15) (Michener 1977), suggesting a loss of 0.5 young per litter from field litters in 1976 during the 30d between birth and weaning. No attempt was made to estimate in-burrow losses of the 1977 litters. Because 1976 had both the largest number of females rearing litters and the largest emerged litter size, the number of juveniles reared on the area that year was greater than in 1975 and 1977 (Figure !). There were 48 juveniles (28 males and 20 females) in the 11 of the 12 litters for which adequate data were available, compared with 15(7 males and 8 females) in 1975 and 28 (13 males, 13 females, 2 unknown) in 1977. In 1975 only one of the 15 juveniles born on the area failed to remain resident until fall; on 14 July I killed a 366 THE CANADIAN FIELD-NATURALIST Vol. 93 11 1:3 1:3 16° 1S we2i 5.5 0) OH) 2-0 1977 i) 6 litters rae es Chemie ats 10 E | (0) LJ | ao BG T | i — 14 1:3 1-0 0-8 0-8 1 7-0 3 1976 30 11 litters wm Sie2d ine He > born on —) => 20 B area Ba Mis a 7a ws :] S a = 10 a fo H ae = e : | a foo) 0 Il [] [sat 0-9 1-3 4 14 1-4 14 16 9:0 1975 coc = 6 litters co 10 : age ieoml ad Eels jae =m = ] 0 4 R= = 15 31 15 30 15 31 15 31 15 30 15 MAY JUNE SURIAY, AUGUST SEY OCT FIGURE |. Numbers of juvenile males (left bar) and juvenile females (right bar) resident on the area in 1975, 1976, and 1977. The sex ratio (males per female) is given above each pair of bars. The arrows indicate the time in each year at which juveniles first began leaving the population to hibernate. female that had severe myiasis of the neck and shoulder due to Sarcophaga. citellivora maggot infestation. The remaining juveniles disappeared in mid- and late September (Figure 1) but only five females and two males were subsequently recovered as yearlings in 1976. Of the 28 male and 20 female juveniles born on the area in 1976 none of the 19 males and 9 females that disappeared before 28 August was recovered in 1977. I assume that these losses were due to death or dispersal and that the pre-hibernation population consisted of those juveniles still resident on 28 August. There was only one accountable loss: I killed a juvenile male with Sarcophaga citellivora myiasis of the leg on 12 August. The initial sex ratio at litter emergence in 1976 was male-biased (Figure 1), but male loss was greater than female loss, particularly during July and early August when males were lost ata rate of 0.31-d | compared with 0.13-d' for females. By mid-August the sex ratio was female-biased and remained so until juveniles began entering hibernation. Of the 9 males and I]! females resident after 28 August two and six, respectively, were recovered in 1977. Losses from the 28 juveniles that emerged on the area in 1977 commenced with the disappearance of four young, comprising an entire litter, between their emergence on 25 May and my next visit on 29 May. Because the mother remained resident at the same location, discounting the possibility of removal of her litter to a new burrow, and because juveniles 30 to 33d old are unlikely to disperse, these young probably had died. A further three males and seven females NOS disappeared by the end of July. One of these males when last captured on 12 July had a swollen tongue, showed labored breathing, weighed 160 g compared with 223 + 28 g for nine other juvenile males captured that day, and presumably died. Because none of the juveniles born in 1977 was recovered as a yearling in 1978, I could not attribute disappearance in late summer to immergence as opposed to death or dispersal for any individual. Assuming that im- mergence did not commence before mid-August, at most one female and seven males hibernated on the area. Thus there was a total loss of at least 6 males, 12 females, and 2 newly emerged young of undetermined SeX. Each year juveniles appeared that had not been born to resident females. If they used a burrow system on the area for at least 2 wk they were classed as resident immigrants (Figure 1). There were more immigrant juveniles in 1975, the year with the lowest squirrel density, than in 1976 or 1977. Of the five males and one female classed as immigrants in 1975, three of the males were known to have been born to females that were resident immediately adjacent to the area. Their appearance on the area may therefore reflect selective use of the mother’s total range rather than true dispersal away from the natal area. Inadequate information is available on the other immigrants to determine whether any had moved over a large distance. Transient juveniles were also seen each year; some were caught several times over the summer and were probably nearby residents making excursions. Causes of Loss Disappearance of a ground squirrel could be due to three factors: death, dispersal, or entry into hiber- nation. Usually cause of loss could be accurately given only for those squirrels which reappeared the next spring; they left the active population and hibernated. In this study, of the 32 adult females, 8 adult males, 22 juvenile females, and 30 juvenile males resident (including immigrants) at the time of entry into hibernation, 21 (66%), 3 (38%), 12 (55%), and 6 (20%), respectively, were recovered the following year, indicating that greater male than female loss occurred between the time the first squirrel in each cohort disappeared underground and the time the last squirrel in each cohort reappeared in spring. I believe that the majority of the missing animals died after they left the active population to hibernate. Because some males were already active in 1976 and 1977 when the area was first visited in the spring, more males may have survived the hibernation period but moved off the area. A suspected, though not verified, source of mortality during the hibernation phase was predation by terrestrial predators, particular Badgers, on torpid MICHENER: GROUND SQUIRREL POPULATION DYNAMICS 367 squirrels over the several-month period between immergence and freezing of the soil. Extensive Badger digging on the study area occurred between 13 September and 28 October 1975, and between 28 August and 10 October 1976. Such predation could be age or sex selective if there are any differences in the quality or depth of hibernacula used by adults and juveniles or by males and females. A possible source of mortality of adult males during early spring was intraspecific fighting. Once females emerged from hibernation the frequency and extent of wounds on the lower back, legs, and shoulders of males increased; I do not know whether the damage was inflicted by other males or by females. Such wounding associated with the weight loss males normally undergo during the breeding season could result in death, directly or through susceptibility to disease and predation. Of the 7 adult females, 22 juvenile females, 25 juvenile males, and 2 juveniles of unknown sex lost during the active season, two were known deaths (killed because of Sarcophaga citellivora infestation) and three were suspected deaths due to Badger predation or poor health. None of the remaining 5 adults and 48 juveniles that disappeared during the summer was subsequently recovered and cause of disappearance was not known for any of them. Some losses to predators undoubtedly occurred. I saw Long-tailed Weasels, Badgers, and Coyotes (Canis latrans) on the study area on 19%, 11%, and 15% of the days I spent there and I found 0, 13, and 17 scats in 1975, 1976, and 1977 respectively. Twelve of the 13 scats found in 1976 and 13 of the 17 found in 1977 contained the remains of Richardson’s Ground Squirrels. Although these remains may not have come from squirrels on the study area, they do indicate that Richardson’s Ground Squirrels are a usual source of prey for these predators. Red-tailed Hawks (Buteo Jamaicensis), Swainson’s Hawks (B. swainsoni), and Golden Eagles (Aquila chrysaetos) were often sighted and Bald Eagles ( Haliaeetus leucocephalus) were seen every spring during their northward migration. One hawk pellet found on the area in 1976 contained the remains of an adult squirrel. In 1976 one carcass and in 1977 two carcasses were found on the area; there were insufficient remains to verify the cause of death or whether these were resident squirrels. I assume that some of the losses over the active season were due to dispersal but, with the exception of four adult males that were not summer residents but were captured on the area in two consecutive springs, there were no verified cases of dispersal from the area. Dispersal onto the area occurred but often involved only a small-scale movement of a nearby resident. Of 4 adult females, 3 juvenile females, and 10 juvenile males that moved onto the area 4, 0, and 3 368 respectively were known to have been previously resident within 50 m of the area. Four of the seven males classed as permanent residents in 1976, 1977, and 1978 had not been resident the previous year and thus had immigrated in spring. Discussion The adult sex ratio among Richardson’s Ground Squirrels is biased in favor of females (Nellis 1969; Sheppard 1972; Michener and Michener 1977) but virtually all females are bred (Michener 1974), indicating that males range over an area sufficient to encounter three or four females during the breeding season. Although Yeaton (1972) stated that male Richardson’s Ground Squirrels established territories in spring that encompass the burrows of three to five females, implying exclusive access to those females, he neither defined his use of the term territory nor his method of determining territorial size and bound- aries. From my observations of squirrels in early spring of 1976 I found that the ranges (see Michener 1979a for details of calculating range size) of each of the six males classed as temporary or permanent residents overlapped not only with the ranges of at least five females but also with the ranges of at least two males, while the ranges of the females overlapped the ranges of between two and five males. Apparently no male had exclusive access to any female. Because females are bred soon after emergence in spring, exclusive access may occur if males make day-to-day variations in their ranges relative to the location of receptive females. This study indicated that in spring males remained on the study area for periods ranging froma few hours to several days to the entire season. Because of the presence of transients and temporarily resident males, more males were trapped in early spring than subsequently established residency within the area. If males had not been classed according to their status on the area, trapping data alone would have revealed male loss in spring suggesting differential mortality favoring females, as reported by Dorrance (1974) and Schmutz et al. (1979). The size of the ranges occupied by males declined after the breeding season (Michener 1979a) because of a decrease in the length and frequency of movements made by males. Although some loss of males in spring was probably due to mortality, the ‘loss’ of at least seven males that were transients or temporary residents was related to this decrease in movements resulting in a smaller range that no longer overlapped the study area. By the time females began giving birth no further changes in range usage by males occurred and all males still present remained as summer residents. Adult males that had been born on the area also changed their residency status with respect to the study area within any one THE CANADIAN FIELD-NATURALIST Vol. 93 breeding season and between consecutive breeding seasons. Because female Richardson’s Ground Squir- rels remain in the same area over several years one effect of male movement in spring is to reduce the amount of inbreeding. This study confirmed previous observations (Yea- ton 1972; Michener 1979a) that adult females, . including yearlings, make only small-scale changes in the location of their ranges between years and within years. Female losses from the area during the active season were low in 1975 and 1977, and were attributed to mortality. In 1976, however, four adult females disappeared in June. The period immediately after lactation would be most suitable for dispersal of adult females; dispersal during the breeding, gestation, or lactation periods would be disadvantageous to suc- cessful rearing of a litter, and dispersal later in the - summer would interfere with the weight gain that follows weaning of the litter and might not allow sufficient time to locate a suitable hibernation burrow. This study and those of Dorrance (1974) and Michener and Michener (1977), however, have found no evidence for dispersal by adult females. The higher recovery rate of females hibernating over 1975-1976 (91%) compared with 1976-1977 (60%) and 1977- 1978 (45%) may be related to the higher proportion of females raising no young or only one young in 1975. Weather conditions unsuitable for feeding in the late summer of 1976 and the more severe winter over 1977-1978 (Michener 1979b), combined with the demands of rearing larger litters in 1976 and 1977, probably contributed to the reduced survival in the 1976-1977 and 1977-1978 winters. Whatever the causes, mortality during the overwinter period does appear to be the main factor affecting the size of the population of adult female Richardson’s Ground Squirrels. Each year fewer juveniles emerged than expected on the basis of the reproductive potential of the popula- tion, and there was greater between-year variation in the size of the juvenile population at emergence than expected from the differences in the number of adult females resident each spring. Normally 100% of yearlings and older females are successfully bred, and between 90% and 100% lactate (Nellis 1969; Sheppard 1972; Michener 1974; this study in 1975, 1976, and 1978): the unexplained failure of three yearlings to produce litters in 1977 reduced the potential juvenile population by about 12. Comparison of litter size at birth (Michener 1977) or in utero (Schmutz 1977) with size of field litters at emergence indicates that there is usually a small loss from most litters. Predation on litters by weasels, however, caused major losses and largely accounted for the appearance of only 15 juveniles in 1975 when about 36 were expected from the number of resident lactating females. 1979 There was a net incease of juveniles between emergence from the natal burrow and the time of entry into hibernation in 1975 but net loss in the other two years. In 1975 no juveniles left the area, suggesting that there are densities below which dispersal does not occur, but the immigration rate was highest, indicat- ing that dispersal can affect the size of the juvenile population. Although indirect evidence suggests that terrestrial predators took a larger number of squirrels in the years with the highest squirrel densities, | could not estimate the magnitude of losses or determine whether such losses were sex-related. Luttich et al. (1970) reported that the sex ratio of juvenile Richard- son’s Ground Squirrels killed by Red-tailed Hawks approached equality, whereas Schmutz et al. (1979) found that Swainson’s Hawks and Ferruginous Hawks (Buteo regalis) captured disproportionately more juvenile males. I believe that a combination of dispersal and predation accounted for the majority of losses of juvenile males in 1976 and 1977. Male loss exceeding female loss, as in 1976, has previously been reported for juvenile Richardson’s Ground Squirrels (Dorrance 1974; Schmutz et al. 1979) and other hibernating ground squirrels (Rongstad 1965; McCarley 1966: Slade and Balph 1974: Morton et al. 1974: Dunford 1977). Greater loss of females is unusual and I do not know why 90% of juvenile females disappeared in the summer of 1977. With the exception of females in 1977 the major loss of juveniles from the area and the major immigration of juveniles onto the area occurred during July, as in the studies by Dorrance (1974) and Schmutz (1977), suggesting that there is a minimum age juveniles attain before dispersing and that dispersal is completed at least a month before hibernation commences. Although the sex ratio (males per female) among juveniles in the pre-hibernation population varied from 1.4 in 1975, to 0.8 in 1976, to 9.0 in 1977, the sex ratio among yearlings recovered was female-biased in 1976 (0.6) and 1977 (0.4); with no recoveries in 1978, suggesting greater loss of males than females over- winter. Mortality occurring at the end of the active season, however, when juvenile males were predomin- ant in the active population, or at the beginning of the next active season, when adult males predominantly constituted the active population, would also affect the sex ratio. Schmutz et al. (1979) considered that mortality, mainly due to predation by migrating avian predators, at these times was significant in readjusting the sex ratio, whereas Michener and Michener (1977) believed overwinter mortality to be more important in the readjustment. Laboratory studies of hibernating sciurids show that there are basic differences between males and females, with males having lower survival rates, longer bouts of continuous torpor but a shorter hibernation season, and a greater likelihood of MICHENER: GROUND SQUIRREL POPULATION DYNAMICS 369 permanently arousing when stressed or disturbed during hibernation (Twente and Twente 1967; Morri- son and Galster 1975; Pengelley et al. 1978), so higher mortality of males during the hibernation phase is a possibility. As with previously reported studies (Dorrance 1974: Michener and Michener 1977; Schmutz et al. 1979) of Richardson’s Ground Squirrels I found that there was high inter-year loss of squirrels, with male losses exceeding female losses and juvenile losses exceeding adult losses, and that adult female squirrels rarely dispersed, with the major loss from this cohort occurring over winter. Here I have indicated that observation or trapping at frequent intervals during early spring is necessary to reveal the residency status of adult males. The contraction in size of range used by a male once all females are bred contributes to the apparent loss of adult males from a local area. Some losses result, not from dispersal or predation, but from the male confining his activity to a smaller range outside the study area and thus being overlooked. | found, as did Dorrance (1974), that variations in the overwinter survival of females and in the extent of loss of pups prior to weaning determined the density of the population in mid-summer. Michener and Michener (1977) predicted that when the adult female popula- tion is small or the reproductive success of females is low, there will be no juvenile dispersal. In this study weasel predation, reducing juvenile addition to the population, resulted in the lack of dispersal of juveniles from the area in 1975. The size of the population of adults in spring and of juveniles in summer varied each year as a result of these interac- tions between overwinter survival, reproductive suc- cess, predation, and dispersal. Discrepancies among Dorrance (1974), Michener and Michener (1977), Schmutz et al. (1979), and this study in accounting for the greater losses of both adult and juvenile males compared with females can, in part, be attributed to variations in the populations under investigation. Populations located in aspen parkland (Dorrance 1974) and on short-grass prairie in Saskatchewan (Michener and Michener 1977), in Alberta (Schmutz et al. 1979), and at the western limit of the species’ range (this study) have been studied for 2,3, or 4 continuous years between 1969 and 1978, and the densities of adults have ranged from about | per ha (Michener and Michener 1977) to 10 or more per ha (Dorrance 1974). If, as seems likely, a combination of dispersal, predation, and overwinter mortality poten- tially contribute to total losses and to differential losses among sexes, then in any given location in any year one factor may predominate without eliminating the others as important factors under other condi- tions. Disputes between the relative importance of predation and overwinter mortality currently revolve 370 around persuasive arguments rather than empirical evidence, and will only be resolved by studies in which every individual is traced, by telemetry, to distinguish entry underground from late-season mortality and to distinguish death underground from early emergence in spring followed by death or dispersal. Acknowledgments The first two years of this study were conducted while I held an Isaac Walton Killam Memorial Post- doctoral Fellowship at the University of Alberta. Support and equipment were provided by National Research Council of Canada grants to A. L. Steiner and J. O. Murie of the University of Alberta. J. O. Murie and D. R. Michener reviewed the manuscript. I thank L.S. Davis for providing information on survival of males in 1979, and Mr. and Mrs. J. Bews of the Y-Cross Ranch Ltd. for permitting access to their land. Literature Cited Dorrance, M. J. 1974. The annual cycle and population dynamics of Richardson’s Ground Squirrels. Ph.D. thesis, University of Wisconsin, Madison. 150 pp. Dunford, C. 1977. Behavioral limitation of Round-tailed Ground Squirrel density. Ecology 58: 1254-1268. Luttich, S., D. H. Rusch, E. C. Meslow, and L. B. Keith. 1970. Ecology of Red-tailed Hawk predation in southern Alberta. Ecology 51: 190-203. McCarley, H. 1966. Annual cycle, population dynamics and adaptive behavior of Citellus tridecemlineatus. Jour- nal of Mammalogy 47: 294-316. Michener, D. R. 1974. Annual cycle of activity and weight changes in Richardson’s Ground Squirrel, Spermophilus richardsonii. Canadian Field-Naturalist 88: 409-413. Michener, G. R. 1977. Effect of climatic conditions on the annual activity and hibernation cycle of Richardson’s Ground Squirrels and Columbian Ground Squirrels. Can- adian Journal of Zoology 55: 693-703. Michener, G. R. 1979a. Spatial relationships and social organization of adult Richardson’s Ground Squirrels. Canadian Journal of Zoology 57: 125-139. Michener, G. R. 1979b. The circannual cycle of Richard- son’s Ground Squirrels in southern Alberta. Journal of THE CANADIAN FIELD-NATURALIST Vol. 93 Mammalogy 60(4). /n press. Michener, G.R. and D.R. Michener. 1977. Population structure and dispersal in Richardson’s Ground Squirrels. Ecology 58: 359-368. Morrison, P. and W. Galster. 1975. Patterns of hibernation in the Arctic Ground Squirrel. Canadian Journal of Zoology 53: 1345-1355. Morton, M.L., C.S. Maxwell, and C.E. Wade. 1974. Body size, body composition, and behavior of juvenile Belding Ground Squirrels. Great Basin Naturalist 34: 121-134. Nellis, C. H. 1969. Productivity of Richardson’s Ground Squirrels near Rochester, Alberta. Canadian Field- Naturalist 83: 246-250. Pengelley, E. T., R. C. Aloia, B. Barnes, and D. Whitson. 1978. Comparative temporal aspects in hibernation be- tween male and female Golden-mantled Ground Squirrels, Citellus lateralis. Journal of Thermal Biology 3: 88. Rongstad, O. J. 1965. A life history study of Thirteen-lined Ground Squirrels in southern Wisconsin. Journal of Mammalogy 46: 76-87. Schmutz, S. M. 1977. Role of dispersal and mortality in the differential survival of male and female Richardson’s Ground Squirrels. M.Sc. thesis, University of Alberta, Edmonton. 97 pp. Schmutz, S.M., D. A. Boag, and J. K. Schmutz. 1979. Causes of the unequal sex ratio in populations of adult Richardson’s Ground Squirrels. Canadian Journal of Zoology. /n press. Sheppard, D.H. 1972. Reproduction of Richardson’s Ground Squirrel (Spermophilus richardsonii) 1n southern Saskatchewan. Canadian Journal of Zoology 50: 1577- 1581. Slade, N. A.and D. F. Balph. 1974. Population ecology of Uinta Ground Squirrels. Ecology 55: 989-1003. Twente, J. W. and J. A. Twente. 1967. Seasonal variation in the hibernating behavior of Citellus lateralis. In Mam- malian hibernation III. Edited by K.C. Fisher, A. R. Dawe, C. P. Lyman, E. Schonbaum, and F. E. South. Elsevier, New York. pp. 47-83. Yeaton, R.I. 1972. Social behavior and social organiza- tion in Richardson’s Ground Squirrel (Spermophilus richardsonii) in southern Saskatchewan. Journal of Mam- malogy 53: 139-147. Received 24 February 1979 Accepted 30 May 1979 Man?’s Influence on Potential Nesting Sites and Populations of Swallows in Canada A. J. ERSKINE Canadian Wildlife Service, Sackville, New Brunswick. E0OA 3CO0O Erskine, Anthony J. 1979. Man’s influence on potential nesting sites and populations of swallows in Canada. Canadian Field-Naturalist 93(4): 371-377. The nesting of the seven species of swallows (Hirundinidae) occurring in Canada was examined using the nest records assembled in Canadian nest records schemes. All species have been influenced by the actions of modern man, and more have been favourably affected through increased availability of nest sites than have declined, despite adverse competition from exotics introduced by man. Changes in breeding range related to environmental changes caused by man have occurred and are still occurring. On a examiné la nidification de sept espéces d’hirondelles (Hirundinidae) au Canada, utilisant des fiches rassemblées dans les fichiers de nidification des oiseaux canadiens. Les activités de homme depuis la colonisation européene ont influencé toutes les sept espéces, mais la plupart des espéces ont profité a cause de la disponibilité augmenteée des sites de nidification. Les alterations de l'environnement causée par "homme ont aussi influence la distribution de quelques espéces d’hirondelles, et tout ¢a continue toujours. Key Words: Hirundinidae, swallows, nesting sites, Canada, nest records, populations. Seven species of swallows (Hirundinidae) are widespread in Canada and the United States, includ- ing two which also breed in Europe, Asia, and North Africa, and another which breeds also in South America. All are at least moderately gregarious, and several nest colonially. Nesting is restricted more to particular situations, sites, or substrates than to special habitats. This paper explores some of the effects of actions by man, particularly since white settlement, on availability of nesting sites for North American swallows, and speculates on their indirect effects on numbers of those birds. Materials and Methods I examined all records through 1974 of swallow nests in the five major Canadian nest records schemes (NRS) (see Acknowledgments). The few records (less than 20 per species) in the Newfoundland NRS were omitted. Each nest was categorized as in a natural site or one affected by man’s actions; some were not certainly assignable to either category. Obvious duplication within a year was eliminated, but not that between years. Published summaries (e.g., Bent 1942: Graber et al. 1972) and authoritative works on each species (q.v.) were consulted. Those sources described the entire spectrum of nest sites reported on the nest record cards without suggesting that any important types had been missed, and consequently no exhaustive literature review was attempted. The relative abundance of each species in different parts of its range in Canada was inferred from results of the co-operative Breeding Bird Survey (BBS) (Erskine 1978, and unpublished data). Those data are believed suitable for broad comparisons of relative density within a species, but not for comparisons between species. Results Data for each species are summarized below. The species are arranged according to type and construction of nests, rather than in taxonomic order. The sequence followed starts with species that use existing cavities, first those that were originally (as far as we can infer) obligate tree-hole nesters, followed by more tolerant species that accept cliff crevices and bank burrows as well as tree holes. Last come species that actively prepare their own nests by excavating burrows or by building fully or partly enclosed structures from clay pellets. PURPLE MARTIN (Progne subis) Our largest swallow is the northernmost represen- tative of a South American group. Its history of association with man extends back before white settlement, when martins already nested in gourds hung up for them by Amerindians (reviews by Allen and Nice 1952: Jackson and Tate 1974). Its use of natural nest sites is now seldom reported in the east: presumably those were in tree cavities, with no more than four or five nests in any tree, as was still the case recently in the west (Richmond 1953; Finlay 1975). Among roughly 3000 nests reported by Canadian nest records schemes, only seven were stated to be in trees or stubs, and five of those were from British Columbia’s small coastal population, from which only one nest box record was also available. Now most martins use artificial nest boxes, includ- ing multiple-unit “hotels” as well as single boxes. Sil S72 Many Canadian nest records reported only the number of “hotels,” which may have up to 30 or more compartments; in such cases the number of occupied nests could only be guessed at. Mean colony size, calculated from reported data, varied from five pairs in Ontario to nine in New Brunswick, ranging from isolated pairs up to clusters of hotels housing over 50 pairs. In Canada, martins are found mostly in towns and cities, and 24 old records from Ontario referred to nests within the walls or roofs of buildings. TREE SWALLOW (Jridoprocne bicolor) The arrival each year of this relatively boreal swallow represents “spring” to many Canadians. It nests singly in tree holes and nest boxes, often near human habitations. Its natural sites are relatively difficult to find and to inspect; only 14% of 4370 reported sites were in tree holes, largely of wood- pecker origin (Table |). Natural sites made up an appreciable fraction of the available records only in British Columbia (Table 1), where several thesis studies had concerned themselves with hole-nesting birds (Erskine 1960; McLaren 1963: Kelleher 1963). The vastly higher proportion of natural sites in British Columbia may also reflect the relatively higher density there of flickers (Co/aptes) (cf., Erskine 1978, Tables 3-8), whose nest cavities are often used by Tree Swallows. There were only two records, both by experienced observers, of Tree Swallows nesting in holes in earth or rock cliffs; cavities in the walls of buildings were seldom used by this species. Tree Swallows evidently find cliff sites unacceptable, and should be viewed as having been obligate tree-hole nesters originally, unlike the following species. Most nest sites classified as affected by man’s actions were nest boxes, usually erected for bluebirds (Sialia spp.), diving ducks (Bucephala spp.), or for Tree Swallows specifically. Cavities excavated by woodpeckers in poles or posts erected by man were also listed as affected by man. THE CANADIAN FIELD-NATURALIST Vol. 93 VIOLET-GREEN SWALLOW (Tachycineta thalassina) A species confined to western North America, this swallow either breeds as single pairs in tree cavities, openings within the walls or roofs of buildings, or in artificial nest boxes, or else in groups in rock crevices (Edson 1943). All of the 550 Canadian nest records were from British Columbia, and 42% were in natural sites, chiefly as colonies in cliffs. Nest boxes and the walls and eaves of buildings accounted for most of the other 58%. Cliff-nesting might be expected to have been originally more prevalent in the sparsely vegetated interior areas of British Columbia, with tree cavities predominating in the coast forests. Unfortunately, I did not segregate the different types of nest sites used by geographic area within the province. We do not know whether birds reared in tree holes will accept holes in buildings (artificial cliffs) or only nest boxes (artificial tree holes), and conversely whether birds reared in cliff sites will or will not accept other sites. ROUGH-WINGED SWALLOW (Stelgidopteryx ruficollis) This brown-backed swallow breeds from South America to southern Canada, and most nest records examined were from southern Ontario and British Columbia. It nests in small groups and as isolated pairs in various bank and cliff situations, using burrows (some or all made by other species) (Lunk 1962) in earth banks, or existing openings in rock walls. Excluding unassignable records, 68% of the 725 nests reported were in natural sites and only 32% in burrows in gravel pits or road cuttings, or (especially in Ontario) holes in cement, brick, or stone walls (cf.., Lewis 1944). Whereas burrows by rivers predominated in British Columbia, man-made sites were most commonly used in Ontario, the most settled (by humans) and altered part of Canada (Table 2). Colony size was higher in British Columbia than elsewhere (Table 2), and the TABLE 1—Regional variations in proportions of natural and man-made sites used by Tree Swallows in Canada % of recorded nests in Natural sites Man-affected sites Woodpecker holes Woodpecker holes Nest Other Region in trees and stubs in poles and posts boxes sites Total nests Maritime provinces 4 5) 88 2 466 Quebec and Ontario 9 4 86 1 2113 Prairie provinces 4 | 93 2 1059 British Columbia 46 4 44 6 752 All regions 14 3 81 2 4370 1979 ERSKINE: CHANGES IN SWALLOW NESTING SITES Sif) TABLE 2—Regional variations in proportions of natural and man-made sites used by Rough-winged Swallows in Canada % of assignable nests Uncategorized Natural Man-made Total number nests Region sites sites (mean number/ colony) (% of gross) Maritime provinces 100 0 I (1) 0 Quebec and Ontario 50 50 213 (2) 19 Prairie provinces 78 22 18 (2) 0) British Columbia US DS 493 (15) 12 All regions 68 32 125 (4) 14 species is also relatively more common in that province, judged by BBS data. BANK SWALLOW ( Riparia riparia) In Eurasia as well as North America, this small swallow nests colonially in burrows which it excavates in near-vertical banks of earth, clay, and fine gravel (Petersen 1955). Nearly one-third of the nest records could not be categorized as natural or affected by man. Of those assigned (27718 in all) about 40% were in natural sites, mainly along sea coasts, rivers, and lake shores where erosion by water or wind led to exposure of earth slopes. The remaining 60% were in cut banks created by man — sand and gravel pits, road and railway cuttings, and various excavations, except for about 1% in piles of gravel stored for road repairs (cf., Nero 1968), and a few in old sawdust heaps (cf., Greenlaw 1972). The data for various regions of Canada are summarized in Table 3. Natural sites were more often reported where human densities are relatively lower, and particularly in the Maritimes where coastal cliffs are also frequent. Man-made sites made up a larger proportion of the records in Quebec and Ontario, where human population, and presumably also habitat disturbance, is greatest. Man-made situations are predominant in British Columbia, where most records reported nests in roadside cut banks. Munro’s monograph (1945) on birds of the Cariboo Parklands, then an undeveloped area with rather primitive roads, omitted Bank Swallows, which now are regular there (Erskine and Stein 1964). Road-building may thus have permitted more general distribution of a formerly localized species. Mean colony size was much lower in the prairies than elsewhere. The relative paucity of nest records as well as the relative density data from BBS suggest that Bank Swallows are scarcer there too. CLIFF SWALLOw ( Petrochelidon pyrrhonota) Cliff Swallows build up their nests from clay pellets plastered against vertical or overhanging walls of cliffs or structures. The nests are fully enclosed, gourd- shaped structures, often assembled in large colonies (Mayhew 1958). Such nests depend on virtually complete overhead protection from rain and runoff water, whether on cliffs or buildings. Of 21096 Canadian nest records, only 23% were in natural sites on cliffs, the rest being on buildings, bridges, culverts, and dams. Cliffs are presumed to be more prevalent in the western mountains than in most of the flatter regions farther east, and records of natural cliff nest sites were almost all from the west (Table 4), with colonies of 1000 or more nests along the Bow River near Calgary, Alberta. The latter biased the regional comparisons (Table 4) of natural vs. man-affected sites and of mean colony size, as no comparably large colonies were: reported in British Columbia. The BBS density indices, however, were as high for those parts of British Columbia where the species was recorded as for Alberta. TABLE 3—Regional variations in proportions of natural and man-made sites used by Bank Swallows in Canada % of assignable nests Uncategorized Natural ~ Man-made Total number nests Region sites sites (mean number/colony) (% of gross) Maritime provinces iS 25 8207 (56) l Quebec and Ontario 35 65 9934 (38) 39 Prairie provinces 57/ 43 509 (5) 0 British Columbia 13 87 8568 (59) 35 All regions 40 60 27218 (42) 29 374 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 4—Regional variations in proportions of natural and man-made sites used by Cliff Swallows in Canada % of recorded nests in Total number Natural Man-made of nests Region sites sites (mean number/ colony) Maritime provinces trace* 100- 1305 (19) Quebec and Ontario trace* 100- 1702 (8) Prairie provinces 57 43 6735 (102) British Columbiat 9 91 11354 (31) All regions 23 V7) 21096 (30) *Trace = much less than 0.5%. tIncluding a few from Yukon Territory and Northwest Territories. BARN SWALLOW (Hirundo rustica) This composition species is “the swallow” of English proverbs, as familiar in Eurasia as in North America. Barn Swallows build their bracket-shaped mud-pellet nests solitarily or in loose colonies inside barns and other open buildings and under the porches and eaves of occupied homes (Samuel 1971), although the original sites were presumably all on overhanging cliffs or in caves. Of the eight locations in Canada where natural sites were reported in use, all in the eastern provinces, four were in national parks and two in a provincial park. Possibly such parks are among the few areas frequented regularly by man where buildings are scarce or lacking. Among nearly 5000 nests recorded in Canada, only 48 (1%) were on cliffs or in caves, the rest being in (54%) or on the outside of buildings (33%), or on other structures (12%). Perhaps more than any of our other swallows, its range may be still changing rapidly in response to man’s activities, as is discussed later. Discussion (1) Biases. The nest records reflect the sites that observers were able to find and inspect. Those are certainly biased towards man-altered habitats and man-associated nest sites (cf., Erskine 1971), so that nests located by roadsides, around houses and settlements, and in agricultural areas are certainly reported in larger proportions than the natural sites in undeveloped, and unvisited, areas. When the vast majority (over 99%) of sites reported are man- influenced, however, as for Purple Martins and Barn Swallows, this is likely to reflect the real situation. (ii) Effects of man on natural nest sites of swallows. Man’s impact on natural nest sites probably affected tree-hole nesters most. At the time of European settlement, eastern Canada south of the tundra was almost entirely forested. Subsequently, southern Ontario and the St. Lawrence lowlands of Quebec, as well as parts of the Maritime Provinces, were largely cleared for settlement and agriculture (cf., Edwards 1969). Similar clearing of formerly forested land took place all along the northern edge of the prairies, and in the lower Fraser Valley of British Columbia. Every- where, the large trees were the first to go. More recently, dead trees and snags are being eliminated, on the grounds of unsightliness in settled areas, and as foci of infection by forest insects or as fire hazards in cutover and regenerating forest lands. Thus, the large and dead trees in which woodpeckers most often make cavities — usable by swallows — have been reduced much more than the present extent of forest lands might suggest. Man has probably had little overall influence on availability of other types of natural nest sites used by swallows. (iii) Trends in use of new nest sites resulting from man’s activities. Despite man’s effect in reducing availability of natural sites for some swallows, man- made situations undoubtedly have provided many additional nesting opportunities. Purple Martins are perhaps the most “domestic” of swallows now, as most of the sites they use are deliberately provided for them by man, in close proximity to human dwellings. This association began before white settlement, and adaptation to artificial sites may have been gradual. Although martins often assemble in far larger colonies in man-made boxes than can have existed in natural tree sites, the size of colonies still averages only 5.8 nests (calculated from data in Jackson and Tate 1974). Tree Swallows were originally no less obligate as tree-hole nesters than martins, but their dependence on man-made sites is much less. Possibly their association with man is of more recent origin, since their more northern distribution would have brought them in contact, before white settlement, mainly with Amerindians of nomadic habits, whereas the early “adoption” of martins was by sedentary agricultural tribes (Allen and Nice 1952). The importance of nest boxes to Tree Swallows can easily be overestimated. I suggest, largely on the basis of “bluebird trail” reports, that nest boxes systemati- 1979 cally erected for bluebirds and other birds may accommodate perhaps 10000 pairs of Tree Swallows annually across Canada, and backyard nest boxes may add another 10000 pairs, say 20000 pairs in all. This looks impressive, but I have also estimated, from density data given by Graber and Graber (1963) and Stewart and Kantrud (1972) and the BBS density indices (Erskine 1978, Tables 3-8), that the total Canadian population of Tree Swallows may be roughly one million pairs. Neither of these estimates is precise, but they should serve to provide the necessary perspective. The increase effected by use of nest boxes is apparently in the order of 2%, which is almost certainly insufficient to counterbalance the losses of natural tree nest sites throughout the settled parts of Canada in the last 200 yr. The apparent ability of Violet-green Swallows to use both tree and cliff sites deserves study. That species is tolerant of a wide spectrum of nesting cavities, which presumably has eased its acceptance of man-made sites. Its domesticity in using backyard nest boxes and holes in the eaves of houses is in marked contrast to its nesting in remote cliffs. Rough- winged Swallows, which also use crevices in rock cliffs, are possibly the least domestic of all Canadian swallows: they have made no real adaptation in using existing cavities in man-made as well as natural banks and cliffs. It seems likely that man has not had much influence on nesting by those two species. Bank Swallows excavate their burrows in man- made banks exactly as they do in natural situations. The much greater availability of cut banks and excavations, even in flat areas, undoubtedly allows more general distribution of that species than in former times. Nesting sites for Cliff Swallows have become far more generally available as a result of man’s actions. That species is much more widespread and abundant in the west than ineastern North America (Erskine 1978, Tables 3-8), as was also the case in the early 1800s (Bent 1942). In contrast, Barn Swallows have an eastern predominance today (Erskine 1978), but natural sites cannot originally have been numerous /in eastern North America, as is also suggested by the present scarcity of the species in areas without human settlement. The readiness of Barn Swallows to build nests on newly-constructed buildings suggests that they may have colonized permanent dwellings as soon as these were built by European settlers in America. Barn Swallows have recently spread southwards in the southeastern United States, as shown by comparison of recent BBS data with the range given in the AOU Checklist (1957) only 15 yr earlier (D. Bystrak, United States Fish and Wildlife Service, unpublished data; Reid 1975). Prior to 1964, the Barn ERSKINE: CHANGES IN SWALLOW NESTING SITES 3/5 Swallow also had not been recorded in northern Alberta, an area with no cliffs and few settlements (Erskine 1968), but with opening of that area for agriculture and oil prospecting it is now seen regularly. Samuel (1971) thought there was little competition between Barn and Cliff Swallows in West Virginia, although they frequently occurred together. (iv) Effects on swallows of competition by introduced exotics. The introduction to America by European settlers of the House Sparrow (Passer domesticus) and Starling (Sturnus vulgaris) has had adverse effects on most native hole-nesters with which the newcomers compete for nest sites. In the southern states, where Purple Martins are relatively more common and Starlings and sparrows less common than farther north, Jackson and Tate (1974) considered such competition a minor influence on martin numbers. But farther north, near the limits of their range, some martin colonies have been abandoned as a result of competition with sparrows and Starlings when no efforts were made by man to protect them (e.g., Brinkman, in Graber et al. 1972). In the west (Oregon and Alberta), Purple Martins did not accept nest boxes until after the introduced competitors had arrived. House Sparrows were extremely local on the west coast until 1900-1910 (Robbins 1973), and Starlings did not breed in British Columbia until about 1948 (Myres 1958). One or other of the Tree or Violet-green Swallow was the most common bird species in three northern frontier towns in which both House Sparrows and Starlings were present (Erskine 1977, Table 12), but census data show that in southern Canadian cities House Sparrows and Starlings are much more common than any swallow species (Speirs et al. 1970; Weber 1972). Detailed studies of competition are lacking, but competition with the introduced pests has presumably influenced both Violet-green and Tree Swallows, especially in urban areas. Bent (1942) quoted a few records of House Sparrows and Starlings using Bank Swallow burrows, and this is evidently rare in Canada too. Competition from House Sparrows adversely affects Cliff Swal- lows nesting on farm buildings (Samuel 1969), but probably has less influence on those nesting in other situations. It seems likely that the rapid initial spread in range and increase in numbers of Cliff Swallows in the east, following their acceptance of buildings as nesting situations in the early 1800s (Bent 1942), was reversed less than a century later by the influence of House Sparrows. Barn Swallows are very seldom bothered by House Sparrows, which prefer a fully enclosed nest site. (v) Effects on swallows of possible changes in insect populations. In the Maritime Provinces and probably 376 also in eastern Ontario, I have the impression that Tree Swallows have become markedly scarcer over the past 25 yr, and especially between 1950 and 1960, both absolutely and relative to Bank and Barn Swallows (cf., my comments in Boyer 1966). The low numbers I found in the Maritimes in 1960 could have reflected losses in 1958 and 1959, but that leaves unexplained their failure to recover subsequently, and their similar scarcity in eastern Ontario after | moved there in 1968. Competition with House Sparrows and Starlings is unlikely to be implicated, as both exotics were well established in those areas before 1950. I believe the decrease more likely to reflect a reduction in flying (and biting) insects in and around urban areas as a result of the widespread use of DDT and other insecticides for forest spraying and mosquito control. Such insects were still conspicuously present in the frontier towns (with abundant swallows) that I censused in 1970-1975, although I have no quant- itative data on insects. Increased use of cars, with resultant increase in air pollution, is also well correlated in time and space with the apparent recent decline of Tree Swallows in urban areas. The continued relative success of Purple Martins in urban areas may reflect their tendency to forage higher above the ground than other swallows. Local insecticide spraying and air pollution in cities and towns may have less effect in reducing the high-flying insects on which martins feed. (vi) Conclusions. Historical records generally pro- vided little information other than presence inan area as opposed to absence from it, which is of use in assessing changes in distribution, but little help in detecting trends in numbers. The changes in numbers which I am suggesting in Table 5 are based on my experience in using nest records and in interpreting THE CANADIAN FIELD-NATURALIST Vol. 93 population data, and on my personal impressions from 25 yr of field work across Canada. My suggestions are certainly speculative, and some people will question my summation of the various influences acting on swallows. The order in which the species were discussed reflected their success in coping with man’s effects on their environment as well as the types of nests they used. The obligate tree-hole nesters, discussed first, depended on trees large enough to accommodate cavities. Those birds seem to have come off least well, losing potential nest sites with the cutting of trees for settlement and agriculture as well as for timber, and suffering from competition by introduced hole- nesting species. The more tolerant cavity-nesting swallows have been able to cope better with man- caused changes, but have neither decreased nor increased. Finally, the species that actively construct their own nests have been able to take advantage of either man’s penchant for excavation or his buildings, and those species all have prospered. None of our swallows, however, can be considered extreme ex- amples of response to man’s actions; none has been reduced to threatened status or extirpated, either through direct persecution or damage to its environ- ment; and none has burgeoned to the point of becoming a pest. Few other families of passerine birds have become as closely associated with man without achieving either domestic or pest status. I believe it desirable to show that not all species of birds are necessarily affected adversely by changes brought on by man. Even highly migratory and esthetically pleasing species, including several of the swallows, have become almost totally dependent on man for nest sites. We should be aware of the extent of such dependence when we make environmental decisions. TABLE 5—Man’s effects on swallow numbers in Canada, judged from nest-site selection and availability. Key to impacts: — major decrease; — moderate decrease; 0 little or no change; + moderate increase; ++ major increase Impact of man’s actions on swallows Availability of natural Species sites Purple Martin = Tree Swallow — Violet-green Swallow 0; =] Rough-winged Swallow (0) Bank Swallow (0) Cliff Swallow 0) Barn Swallow oO Suggested changes from original Status Availability of new sites Competitors introduced* Qe | 2 © +++ +44 t++oo) | *House Sparrow, Starling. +Cliffs and trees listed separately, also buildings and nest boxes. ——— F 1979 Acknowledgments I thank the co-ordinators of the Canadian nest records schemes and their co-operators for the use of data. The nest records are stored at Canadian Wildlife Service, Sackville, New Brunswick (Maritimes NRS); National Museum of Natural Science, Ottawa, On- tario (Quebec NRS); Royal Ontario Museum, Toronto, Ontario (Ontario NRS); Manitoba Museum of Man and Nature, Winnipeg, Manitoba (Prairie NRS); and British Columbia Provincial Museum, Victoria, British Columbia (British Columbia NRS). C.S. Robbins read and commented on an earlier draft, and A. L. A. Middleton and especially M. T. Myres provided valuable suggestions which aided in preparation of the final version of this manuscript. Literature Cited Allen, R.W. and M.M. Nice. 1952. A study of the breeding biology of the purple martin (Progne subis). American Midland Naturalist 47: 606-665. American Ornithologists’ Union. 1957. Check-list of North American birds. 5th Edition. Ithaca, New York. Bent, A.C. 1942. Life histories of North American fly- catchers, larks, swallows, and their allies. United States National Museum, Bulletin Number 179. pp. 374-510. Boyer, G. F. 1966. Birds of the Nova Scotia~New Bruns- wick border region. Canadian Wildlife Service Occasional Paper Number 8. 41 pp. Edson, J. M. 1943. A study of the Violet-green Swallow. Auk 60: 396-403. Edwards, R. Y. 1969. The hardwood forests that vanished. Ontario Naturalist 2/69: 9-12. Erskine, A.J. 1960. A discussion of the distributional ecology of the Bufflehead (Bucephala albeola; Anatidae: Aves) based upon breeding biology studies in British Columbia. M.A. thesis, University of British Columbia, Vancouver. Erskine, A. J. 1968. Birds observed in north-central Al- berta, summer 1964. Blue Jay 26: 24-31. Erskine, A. J. 1971. Some néw perspectives on the breeding ecology of Common Grackles. Wilson Bulletin 83: 352-370. Erskine, A. J. 1977. Birds in boreal Canada: communities, densities and adaptations. Canadian Wildlife Service Report Series Number 41. 71 pp. Erskine, A. J. 1978. The first ten years of the cooperative Breeding Bird Survey in Canada. Canadian Wildlife Service Report Series Number 42. 59 pp. Erskine, A. J. and R. C. Stein. 1964. A re-evaluation of the avifauna of the Cariboo Parklands. British Columbia, Report of Provincial Museum 1963. AAI8—-AA35. Finlay, J.C. 1975. Nesting of Purple Martins in natural cavities and in man-made structures in Alberta. Canadian Field-Naturalist 89: 454-455. Graber, R.R. and J.W. Graber. 1963. A comparative study of bird populations in Illinois, 1906-1909 and 1956-1958. Illinois Natural History Survey, Bulletin 28: 383-528. ERSKINE: CHANGES IN SWALLOW NESTING SITES S17 Graber, R. R., J. W. Graber, and E. L. Kirk. 1972. Illinois birds: Hirundinidae. Illinois Natural History Survey, Biological Notes Number 80. 36 pp. Greenlaw, J.S. 1972. The use of sawdust piles by nesting Bank Swallows. Wilson Bulletin 84: 494-496. Jackson, J. A.and J. Tate, Jr. 1974. Ananalysis of nest box use by Purple Martins, House Sparrows, and Starlings in eastern North America. Wilson Bulletin 86: 435-449. Kelleher, K. E. 1963. A study of the hole-nesting avifauna of southwestern British Columbia. M.Sc. thesis, Univer- sity of British Columbia, Vancouver. Lewis, H. F. 1944. Recent breeding of the Rough-winged Swallow near Ottawa. Canadian Field-Naturalist 58: 15-16. Lunk, W.A. 1962. The Rough-winged Swallow, Ste/gi- dopteryx ruficollis (Vieillot), a study based on its breeding biology in Michigan. Publications of the Nuttall Ornitho- logical Club, Number 4. 155 pp. Mayhew, W. W. 1958. The biology of the Cliff Swallow in California. Condor 60: 7-37. McLaren, W.D. 1963. A preliminary study of nest-site competition in a group of hole-nesting birds. M.Sc. thesis, University of British Columbia, Vancouver. Munro, J. A. 1945. The birds of the Cariboo Parklands, British Columbia. Canadian Journal of Research 23D: 17-103. Myres, M.T. 1958. The European Starling in British Columbia: 1947-1957. Occasional Papers of the British Columbia Provincial Museum, Number 11. 60 pp. Nero, R. W. 1968. Bank Swallows in gravel stock-piles in Manitoba. Blue Jay 26: 23. Petersen, A.J. 1955. The breeding cycle in the Bank Swallow. Wilson Bulletin 67: 235-286. Reid, R.R., Jr. 1975. Range expansion and habits of the Barn Swallow in Alabama. Alabama Birdlife 23(3, 4): 3-11. Richmond, S. M. 1953. The attraction of Purple Martins to an urban location in western Oregon. Condor 55: 225-249. Robbins, C.S. 1973. Introduction, spread, and present abundance of the House Sparrow in North America. A symposium on the House Sparrow (Passer domesticus) and European Tree Sparrow (P. montanus) in North. America. Edited by S.C. Kendeigh. Ornithological Monographs (A.O.U.), Number 14. pp. 1-9. Samuel, D. E. 1969. House Sparrow occupancy of Cliff Swallow nests. Wilson Bulletin 81: 103-104. Samuel, D. E. 1971. The breeding biology of Barnand Cliff Swallows in West Virginia. Wilson Bulletin 83: 284-301. Speirs, J. M., G. Markle, and R. G. Tozer. 1970. Popula- tions of birds in urban habitats, Ontario County, 1969. Ontario Field Biologist 24: 1-12. Stewart, R. E. and H. A. Kantrud. 1972. Population esti- mates of breeding birds in North Dakota. Auk 89: 766-788. Weber, W.C. 1972. Birds in cities: A study of popula- tions, foraging ecology and nest-sites of urban birds. M.Sc. thesis, University of British Columbia, Van- couver. Received 22 December 1978 Accepted 2 May 1979 Distribution and Habitats of Four Annual Smartweeds in Ontario RICHARD J. STANIFORTH! and PAUL B. CAVERS2 'Department of Biology, University of Winnipeg, Winnipeg, Manitoba R3B 2E9 2Department of Plant Sciences, University of Western Ontario, London, Ontario N6A 5B7 Staniforth, Richard J. and Paul B. Cavers. 1979. Distribution and habitats of four annual smartweeds in Ontario. Canadian Field-Naturalist 93(4): 378-385. Key Words: smartweeds, Polygonum lapathifolium, Polygonum pensylvanicum, Polygonum persicaria, Polygonum scabrum, distribution, habitats, Ontario. Polygonum lapathifolium, P. pensylvanicum, P. persicaria, and P. scabrum are closely related and morphologically similar smartweeds. They often occur together on riverbanks in southern Ontario; outside of riverbanks, their habitats and distributions differ. Polygonum lapathifolium is found on damp disturbed substrates throughout the province (variety lapathifolium predominates in the south, and variety salicifolium in the north); P. pensylvanicum is found (as varieties pensylvanicum and laevigatum) on well drained soils of lakeshores, riverbanks, and occasionally in farmland in southern Ontario. Polygonum persicaria and P. scabrum have been introduced from Eurasia; in Ontario their distributions are more or less restricted to wasteland, and cultivated regions where they are locally common as weeds in cereal crops. A key and line drawings are provided for identification purposes. Annual smartweeds or persicarias (Genus Poly- gonum, Section Persicaria) are a group of similar species which inhabit croplands, as well as wet and disturbed sites in many parts of the world. In Ontario, several species are widespread and occur with high population densities in many locations. The similarity between species has resulted in their frequent mis- identification, or in their being lumped together as “smartweeds.” Furthermore, two or more species may be found together in close proximity, and this raises the question of how very similar species can occur together without one succeeding at the expense of the others. One possible answer to this question is that each is predominant in at least one kind of habitat. The purpose of this study was to clarify and compare the distribution and habitats of four species and their varieties in Ontario. The species studied were Pale Smartweed ( Polygonum lapathifolium L.), Pennsylvania Smartweed (P. pensylvanicum L.), Lady’s-thumb (P. persicaria L.), and Green Smart- “weed (P. scabrum Moench). We follow the nomen- clature and classification of Scoggan (1978) by recog- nizing four species and a number of varieties. A similar classification has been used by Fernald (1950): however, others (e.g., Gleason and Cronquist 1963) have preferred to treat P. scabrum Moench as a variety of P. lapathifolium (P. lapathifolium variety incanum (Roth) K. Koch). By referring to herbarium records, we have documented the distribution of these smartweeds in Ontario. We have also documented, after an intensive survey, their habitats in Middlesex County, Ontario. Methods Specimens from Ontario were examined from the following herbaria during 1972 and 1973: OAC, UWO, TRT, CAN, DAO, SLU, HAM, WLU, WAT, QK, LKHD, MT, MTJB, MT MG, MICH, MSC, and the University of Windsor. Morphological data recorded from these herbarium specimens was later (1978) used to check specimen identification against keys in the recently published Flora of Canada (Scoggan 1978). Each specimen was identified to variety where possible and its geographical location was then plotted. A study area (approximately 3500 ha) was selected near the village of Delaware, Middlesex County, Ontario (42°55’N, 81°25’W). A significant criterion in the selection of this area was that it possessed a variety of habitats and soil types (Anonymous 1931). The habitats included cropland (cereals, corn, tobacco, hay, beans, vegetables), pasture, deciduous wood- land, roadsides, and river bottomlands. Soil types included “sands” (Berrien and Fox-fine sandy loams, Oshtemo and Plainfield sands), “loams” (Guelph, London, and Parkhill loams and Burford gravelly loam), and “clays” (Haldimand, Huron, and Perth clay loams). Sands, loams and clays, as defined above, occupied approximately equal proportions of the study area. The Thames River flows southwards through the center of the area. Many tributaries enter this portion of the river via gulleys on its east and west banks. A survey of the study area was designed to examine the distribution of the smartweeds in different 378 I) habitats and on different soil types. The study area was subdivided into three edaphic types: sand, loam, and clay; 300 sampling units (each circular with diameter of 3 m) were placed randomly in each soil type in such a way that !00 were in cropland, 100 in woodland, and 100 in disturbed sites other than cropland (e.g., roadside verges). The presence or absence of each smartweed species was recorded in each of the 900 sampling units. If the sampling unit fell in the wrong habitat, it was moved to the nearest example of the appropriate habitat. A field check of the soil type was made at the time that a plot was placed in the field. In each sampling unit the soil type was the same as that shown on the Department of Agriculture soil map (Anonymous 1931). The frequency of each smartweed species was determined from 300 sampling units placed along both sides of a 6.5-km section of the Thames River, near London, within the study area. Each bank was divided into 50 sections each 130 10m. Three circular sampling units, each | m in diameter, were placed at random within each section. This habitat was treated separately from those of the previous survey because riverbank soils could not be classified as sand, loam, or clay and because the sampling procedure differed. Results and Discussion Distribution in Ontario Table | gives the numbers of herbarium specimens STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS 37/9) examined; most of these were identified to variety level. Three varieties (P. lapathifolium variety pros- tratum Wimm., P. pensylvanicum variety durum Stanford, P. persicaria variety ruderale (Salisb.) Meisn.) are not recorded by Scoggan (1978) for Ontario and this is probably the first report of these taxa for the province. A key (Table 2) and line drawings (Figure |) are provided for the identification of Ontario taxa. The key has been modified from keys in Scoggan (1978) to include only those taxa (to variety level) found in Ontario, and expanded to include the previously unreported P. pensylvanicum variety durum and a key to allow the separation of P. persicaria variety persicaria from P. persicaria variety ruderale (Salisb.) Meisn. Polygonum lapathifolium was the most widespread species (Figure 2). The most northerly specimen (CAN 244332) was from Big Trout Lake (53°49’N, 89°53’W). Polygonum lapathifolium has been. col- lected from all bedrock types, soil types (except tundra soils), vegetation zones (except tundra), land use types, and climatic regions of the province. Variety /apathifolium appears to be the commonest variety in southern Ontario. Specimens have been collected from riverbanks, damp agricultural land, roadsides, ditches, and wasteland. Variety salici- folium Sibth. is locally distributed in the south, but is the commonest variety in the northern part of the province. Specimens of this variety have been col- lected from sandy beaches of lakes and rivers, and TABLE |—The varieties of four annual smartweed species found in Ontario, based on herbarium specimens examined by the authors No. of specimens Species Variety examined Polygonum lapathifolium L. lapathifolium 218 : salicifolium Sibth. 106 prostratum Wimm. eel 325 Polygonum pensylvanicum L. pensylvanicum 81 laevigatum Fern. 46 durum Stanford! 2 eglandulosum Myers? Av 129 Polygonum persicaria L. persicaria 349 ruderale (Salisb). Meisn. as 350 59 Polygonum scabrum Moench 'Specimens appeared to be hybrids between variety durum and variety pensy/vanicum. 2Not encountered in the specimens examined, but reported by Scoggan (1978) for islands in Lake Erie. 380 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 2—Key to Polygonum lapathifolium, P. pensylvanicum, P. persicaria, and P. scabrum, and their Ontario varieties. ! This key has been adapted from Scoggan (1978) 1 Ocreae (leaf-sheaths) normally fringed with bristles at summit. Spikes at least 7 mm thick; mature calyx prominently reticulate at base; achenes prevailingly lenticular; leaves often purplish blotched above; plants mostly of weedy habitats; (GimMEROG TCE A) ce cree ch etene ss eroe chao ree ates ones ny tue lees ave) oi ceoANfayo shale keane eS PSEIG) OG Casio Lot eR RARE P. persicaria L. 2 Stem ascending or merely decumbent-based; the leaves narrowly to broadly lanceolate; the spike to over 4 cm long var. persicaria 2 Stem prostrate or depressed; the relatively short leaves rhombiclanceolate; the spike usually less than 1.5 cm long var. ruderale (Salisb.) Meisn. 1 Ocreae nearly or quite lacking apical bristles; styles usually 2 (sometimes 3); achenes usually lenticular (sometimes trigonous) 3 Peduncles and axis of inflorescence with obvious stalked glands or strigose-hispid; spikes pink to purplish (rarely WANE), Onis? sapAlls Ouseumaly MEA! ocaccosossvenesGesedunooonoOCDOEODLODODDOO EONS P. pensylvanicum L. 4 Peduncles strigose-hispid, without or with very few glands var. durum Stanford? 4 Peduncles and axis of inflorescence covered with gland-tipped hairs. Si leavesndistinctlyastneoseronmbothisuiblacesm rn ie rice iio ears var. pensylvanicum? 5 Leaves glabrous or at most sparsely strigose on the midrib beneath . var. laevigatum Fern.? 3 Peduncles and axis of inflorescence glabrous or with sessile inconspicuous glands; outer 3 sepals in fruit strongly 3-nerved, each nerve terminating in an anchor-shaped fork (except in P. pensylvanicum var. eglandulosum). 6 Spikes green (rarely purplish), to 5 cm long, erect, the lateral ones mostly sessile or short stalked; floral-axis copiously glandular; achenes about 3 mm long and about equalling the mature calyx; this not constricted at the tips(introduced) ei, se iseseisadaitale + se oe eeeoneia tele crs Geeeeysks tees a olsvannere P. scabrum Moench2 6 Spikes pink to purplish. 7 Plant glabrous throughout; calyx pale, about equalling the somewhat shining achene, this is 3-5 mm longaispikerenectstonl—oremethickeveeemeemiaeeae P. pensylvanicum var. eglandulosum Myers 7 Plant often bearing sessile glands on the peduncles and lower leaf surfaces; calyx pink to purplish, constricted above into a thick beak overtopping the achene, this is not much over 2 mm long; spike often somewhat pendulous, not over | cm thick P. lapathifolium L. 8 Leaves lanceolate, broadest near the base, attenuate to tip. 9 Leaves green on both sides, to 2-5 cm long; spikes to 8 cm long, arching or droop- TTB ses es coe a Sud obese om ae atreri oh en mathe I SU Ree poh oer re var. lapathifolium? 9 Leaves white-pubescent beneath, at most about | cm long; spikes less than 4 cm long: REC aace ee ean Rea ete ne oe ree var. salicifolium Sibth.? 8 Leaves broadest well above the base, not long attenuate. Plant prostrate or depressed; leaves subrhombic, mostly not over 7 cm long; spikes to aboutidicmilom pi cicrenve crstyacle checeher eter mien enna extern var. prostratum Wimm. 'Other annual smartweeds occur in Ontario (e.g., P. caespitosum, P. careyi, P. hydropiper, P. orientale, P. punctatum). These species are either rare, or more or less restricted to swampy ground (see Scoggan (1978) for treatment of these taxa). *Specimens of this variety or species which possessed some characteristics of a second variety or species have been seen by the authors (see text). from exposed muds. Many intermediates between these two varieties were noted; for the purposes of mapping, these were assigned to the variety which they most closely resembled. A specimen of the variety prostratum Wimm. (prostrate plant with broad leaves) had been collected from the vicinity of Waterloo (WLU 2689). Polygonum pensylvanicum has the most restricted distribution of the four species studied (Figure 2). The most northerly specimen (WAT 718a) was from Sudbury (45°30’N, 81°00’W). The Ontario distribu- tion is patchy with three main centers of density: (a) south of a line from Goderich to Toronto, (6) east of a line from Kingston to Renfrew, and (c) the Georgian Bay shoreline. Isolated colonies occur at Nestorville (46° 18’N, 83°36’W; MICH 5165), North Bay (46°19’N, 79°28’W; TRT 16735), and Sudbury (46°30’N, 81°00’W; WAT 718a). Variety pensyl- 1979 POLYGONUM LAPATHIFOLIUM OCREA PEDUNCLE SPIKE STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS 381 3 MM ACHENE PERIANTH FIGURE |. Ocreae, peduncles, spikes, perianths, and achenes (surface and cross sectional views) of annual smartweeds showing major distinguishing features used in the key (Table 2). Spikes of (a) P. lapathifolium variety lapathifolium, and (b) P. lapathifolium variety salicifolium. Peduncles of (c) P. pensylvanicum, varieties laevigatum and pensy/vanicum, (d) P. pensylvanicum variety eglandulosum, and (e) P. pensylvanicum variety hybrid durum X pensylvanicum. Cross sections of (f) trigonous and (g) lenticular achenes of P. persicaria. vanicum and variety /aevigatum Fern. are both common and have similar distributions; however, their main centers of density differ. Variety pensyl- vanicum predominates on the Niagara peninsula, but variety /aevigatum is the most frequent variety in the vicinity of Ottawa and in Elgin and Middlesex counties. Many intermediates between these varieties were noted and, for mapping purposes, were assigned to the variety which they most closely resembled. Two specimens were identified by the authors as inter- mediates between variety durum Stanford and variety pensylvanicum. These had been collected from Point Edward, Lambton County (DAO 1862) and Birch Island, Lake Huron (CAN 44437). No mention is made of the occurrence of variety durum in Ontario by Scoggan (1978); however, Gleason (1958) reports a 382 THE CANADIAN FIELD-NATURALIST POLYGONUM LAPATHIFOLIUM L. var. tapathifolium POLYGONUM LAPATHIFOLIUM L. var. salicifolium Sibth| Vol. 93 POLYGONUM SCABRUM Moench (Pale Smartweed ) POLYGONUM PENSYLVANICUM L. var. pensylvanicum ( Pale Smartweed, willow-leaved variety ) POLYGONUM PENSYLVANICUM L. var. laevigatum Fern (Green Smartweed ) POLYGONUM PERSICARIA L. ( Pennsylvania Smartweed ) Bi FIGURE 2. Distribution of four annual smartweeds and their common varieties in Ontario, based on examination of herbarium specimens. similar variety intermediate from Indiana. Specimens of P. pensylvanicum have been collected from riverbanks, lakeshores, and croplands. The distribution of P. persicaria was most con- centrated in the southern half of the province although specimens have been collected from the clay plains of Kenora, Dryden, Thunder Bay, Kapuskas- ing, and New Liskeard. The most northerly specimen was from Sioux Lookout (50°06’N, 91°55’W). Most habitats given on specimen sheets were described as cultivated land or disturbed sites. The species was recorded from all soil types including Precambrian deposits. A single specimen of P. persicaria variety ruderale (Salisb.) Meisn. was identified by the ( Pennsylvania Smartweed, smooth variety) (Lady's —thumb) authors. This specimen (WAT 39) had been col- lected from Conestogo Lake, Waterloo County. Polygonum scabrum had not been separated from P. lapathifolium in certain of the herbaria visited. Data pertaining to peduncle, spike, perianth, and achene characteristics were used by the authors to discriminate between these two species using the key provided by Scoggan (1978). Many specimens pos- sessed characteristics of both P. scabrum and of P. lapathifolium (either variety salicifolium or variety lapathifolium). Such specimens were assigned to the taxon which they most closely resembled. Polygonum scabrum is of widespread but local occurrence in Ontario; it is particularly abundant in the vicinity of I) Thunder Bay. The most northerly specimen (TRT 107340) had been collected from Onakawana on the Moose River. Information on herbarium sheet labels showed that specimens had been collected from clay or loam soils of farmland and roadsides, and a few from riverbanks. Soil and Habitat Survey Percentage frequencies for P. lapathifolium and P. persicaria in each of the nine habitat-soil categories are shown in Table 3. Polygonum pensylvanicum and P. scabrum were not encountered in this survey, and with the exception of one plant of P. persicaria, no smartweeds were found in woodland. TABLE 3—Frequency and total number of two smartweeds in three habitats and on three soil types in Middlesex County, Ontario. Number of sampling units in each soil type in each habitat = 100 Species, Brcauencyaa) Total habitat Sand Loam Clay number P. lapathifolium Cropland ] 0 0 | Disturbed sites 4 2 0 6 Woodland 0 0 0 0 Total number 5 2 0 7 P. persicaria Cropland 9 24 28 614 Disturbed sites 25 24 24 734 Woodland 1 0 0 ie Total number 35° 4g>-c Se 135 *4R ow or column totals associated with the same letter are not significantly different from each other (P > 0.05). Polygonum lapathifolium was scarce (encountered in only 7 of 900 sampling units); most occurrences were in disturbed sites. Frequency values were too low for statistical comparison. All specimens were of variety lapathifolium. Polygonum persicaria was common in croplands and in disturbed sites on sandy, clay, and loam soils. The frequency values were sufficiently large to allow statistical testing of four null hypotheses: i) That the frequency of P. persicaria on different soils was independent of its frequency in different habitats was rejected (P< 0.05) using the G-test (Sokal and Rohlf 1969). Polygonum persicaria was most frequent in croplands where these were situated on clay soils. ii) That the frequency of P. persicaria did not differ between soil types, or between cropland and disturbed sites was tested by means of pairwise comparisons using the simultaneous testing pro- cedure (Sokal and Rohlf 1969). The frequency onclay STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS 383 soils was significantly higher (P< 0.05) than that on sandy soils. Frequency values for other pairs of soil types, and between habitats, cropland, and disturbed sites were not significantly different. iii) That the proportion of occurrences of P. persicaria in cropland increases as one progresses from sandy through loam to clay soils (i.e., with decreasing particle size) was accepted (P > 0.05) after testing of linear proportions (Snedecor and Cochran 1967). iv) That P. persicaria occurs with equal frequency in different crop types was tested using an R X C test of independence based on the G-statistic, and by pairwise comparisons using the simultaneous testing procedure (Sokal and Rohlf 1969). These tests took into account the total numbers of sampling units in each crop type, in such a way that the scarcity or abundance of sampling units in a particular crop species did not affect the results of the test or provide misleading information. The frequency of P. persi- caria in small-grained cereals was shown to differ (P< 0.05) from that in other crops (Table 4). Small- grain cereals are usually grown onclay or loam soils in the study area. From these studies it is not possible to separate the effects of clay and loam soils from the effects of small-grained cereals on the frequency of P. persicaria. A further complicating factor is that the principal crops on drier soils (corn and tobacco) are usually heavily sprayed with herbicides (such as Atrazine), to which smartweeds are susceptible. TABLE 4—Frequency of Polygonum persicaria in various crop types Small-grained Other Crop type Corn cereals! crops? Number of sites examined 146 85 69 Number of occurrences of ic 54> 22 P. persicaria *’Frequencies associated with the same letter are not significantly different from each other (P > 0.05). ‘Wheat, barley, oats, rye. *Tomato, tobacco, beans, melon. Riverbank Survey Polygonum lapathifolium variety lapathifolium, P. pensylvanicum variety laevigatum, and P. persicaria were encountered in the riverbank survey (Table 5). The null hypothesis that their frequency values did not differ was tested by means of a G-test and by simultaneous testing procedure (Sokal and Rohlf 1969). The null hypothesis was rejected (P< 0.05) 384 TABLE 5—Frequencies of smartweed species along banks of the Thames River, Middlesex County, Ontario Species Frequency (%)! Polygonum lapathifolium 49 Polygonum pensylvanicum 22 Polygonum persicaria 10 Poilygonum scabrum 0 ‘Each value is significantly different from each other value (P< 0.05). indicating that the values were different from each other. It is of interest that P. persicaria, the common- est species in cropland and disturbed sites (habitat and soil survey), was the rarest of the three riverbank species. Polygonum pensylvanicum, which was absent from the previous survey, was common onriverbanks. Polygonum scabrum was absent from riverbank sites, as it was from the habitats examined in the previous survey. Summary The distributions and habitats of the four smart- weeds based on our herbarium studies and field surveys, are summarized in Table 6. Polygonum lapathifolium has long been associated with man’s agricultural activities (Bertsch 1954) and with man’s assistance has increased its range to most temperate parts of the world. It is probably not sufficiently abundant in the croplands of southern Ontario to be rated as a serious weed in that region. It is more usually found in wet and naturally disturbed sites, such as riverbanks, lakeshores, and exposed mud. It may be abundant in these habitats even at locations well away from farmland, e.g., in northern Ontario. When found as a weed, it is usually associated with damp, disturbed substrates, 1.e., conditions very similar to those of the riverbank. Of THE CANADIAN FIELD-NATURALIST Vol. 93 the two common Ontario varieties, variety salicifol- ium is considered to be native but variety /apathi- folium is believed to be partly introduced (Scoggan 1978). It is not known, however, whether the introduced element of this variety is the weedy component. The range of the native species Polygonum pensyl- vanicum is more or less restricted to eastern North America, extending northwards to the Sudbury district of Ontario (for map, see Staniforth 1975). It is a serious agricultural weed throughout its range in the United States, but in Canada it is mostly restricted to riverbanks and lakeshores. When found as a weed in Ontario (locally in Kent, Essex, and Elgin counties), it is associated with crops grown on sandy soils (especially corn and tobacco). One may speculate that its northern distribution is correlated with locations offering a mild climate and light warm soils, such as those of riverbanks, beaches, and sandy farmland. Varieties /aevigatum and pensylvanicum are both common in the province. Polygonum persicaria, like P. lapathifolium, has long been associated with man’s agricultural activi- ties. It has become successfully established in North America from Eurasia. In Ontario, it is more restricted to agricultural and man-disturbed habitats than are the two species discussed above. It does occur in isolated farming regions such as the clay belt and the small farmed clay pockets of northern Ontario. Polygonum persicaria 1s associated particularly with damp, clay soils in Ontario, especially those which support crops of small-grained cereals, e.g., oats, barley, rye, and wheat. Polygonum scabrum was originally introduced from Eurasia. It has become widely distributed in Canada and is now found in all provinces and in the Northwest Territories (Scoggan 1978). In Ontario, it is widespread and locally common on damp clay and loam soils of disturbed substrates, including cropland. TABLE 6—Summary of the distributions and habitats of four annual smartweeds in Ontario Distribution, Species relative abundance Throughout Ontario; common Polygonum lapathifolium Southern Ontario; locally common Polygonum pensylvanicum Polygonum persicaria province; common Polygonum scabrum All cultivated parts of the Throughout Ontario; local Principal habitats Principal soil type Riverbanks, lakeshores, sometimes farmland and waste places Damp clays and loams Riverbanks, lakeshores, locally in crops Well-drained loams, sands, and gravels Wasteland, cultivated land (cereal crops), occasionally riverbanks Damp clays and loams Wasteland, cultivated land, occasionally riverbanks Damp clays and loams ee 1979 It was not encountered during habitat surveys in Middlesex County, Ontario. Acknowledgments We thank the Canadian Northern Sportsmen’s Show for financial support of this project. Comments offered by the referees were appreciated. The assis- tance and co-operation of curators of herbaria in Ontario, Quebec, and Michigan are also gratefully acknowledged. In addition, we thank H. Scoggan for permission to use his keys to species and varieties of Polygonum. Literature Cited Anonymous. 1931. Soil survey map of the County of Middlesex, Province of Ontario. Soil Survey Report Number 6, Experimental Farm Services, Department of Agriculture, Ottawa. Bertsch, K. 1954. Vom neolithischen Feldbau auf der Schwabischen Alb. Berichte der Deutschen Botanischen Gesellschaft 67: 18-21. Fernald, M. L. 1950. Gray’s manual of botany. 8th Edition. STANIFORTH AND CAVERS: ONTARIO SMARTWEEDS 385 American Book Company, New York. 1632 pp. Gleason, H. A. 1958. The new Britton and Brown illustrat- ed flora of the northeastern United States and adjacent Canada. Volume 2. Revised edition. New York Botanical Garden. 655 pp. Gleason, H. A. and A. Cronquist. 1963. Manual of vascu- lar plants of the northeastern United States and adjacent Canada. Van Nostrand Company, Princeton. 810 pp. Scoggan, H. J. 1978. Flora of Canada. Part 3. National Museums of Canada, Publications in Botany Number 7 (3). 568 pp. Snedecor, G.W. and W.G. Cochran. 1967. Statistical methods. 6th Edition. Iowa University Press, Ames. 593 pp. Sokal, R. R. and F. J. Rolf. 1969. Biometry. The principles and practice of statistics in biological research. W. H. Freeman and Company, San Francisco. 776 pp. Staniforth, R. J. 1975. The comparative ecology of three riverbank annual plants. Ph.D. thesis, University of Western Ontario, London, Ontario. 368 pp. Received 6 September 1978 Accepted 29 May 1979 Apparent Differences in Aquatic Macrophyte Floras of Eight Lakes in Muskoka District, Ontario from 1953 to 1977 G. E. MILLER and H. M. DALE Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2W1 Miller. G. E.and H. M. Dale. 1979. Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka District. Ontario from 1953 to 1977. Canadian Field-Naturalist 93(4): 386-390. Forty-eight species of aquatic macrophytes were recorded in two surveys made 23 yr apart of the same areas in eight Muskoka, Ontario lakes. Only 69% of 265 sightings were duplicates in the surveys. Improved sampling methods resulted in additional records of deep-water species whereas certain floating-leafed species were no longer extant in some lakes. Some currently obvious species are thought to have changed sufficiently in abundance to produce the new records. In the shallow- water records. the changes in several subterranean and inconspicuous species were attributed to presence or absence of obvious flowers. Key Words: aquatic plants, macrophyte, flora, biological surveys. freshwater lakes, long-term changes. Muskoka District, Ontario. The process of eutrophication brings about changes in the aquatic flora in a short time (Lind and Cottam 1969: Harman and Doane 1970). Undisturbed aquatic stands, rephotographed in Scotland by Spence (1964) to compare with earlier photographs by West (1905) showed no significant successional change in aquatic communities after more than 50 yr. Dramatic changes more frequently result from man’s activities, because they may alter the water chemistry, clarity, and temperature: these factors have been linked with species change (Dale and Miller 1978: Stuckey 1971: Volker and Smith 1965). The scarcity of accurate floristic surveys of lakes hinders the weighing of the reality of an apparent change in a flora. Too often published records are based on traditional techniques using remote sam- pling from the water surface, with a rake or a grappling hook (Shields!: Soper: Stuckey 1971), or an Ekman dredge (Rich et al. 1971). Direct examina- tion of aquatic vegetation (ina manner comparable to those in which observations are made in terrestrial ecology) using SCUBA was introduced by Schmid (1965) and was the method used for this study. The eight lakes studied are of the unbuffered, slightly acid type, characteristic of the Precambrian Shield of central Ontario. Three Mile Lake has an alkalinity of 9 mg CaCO,/L and an electrical con- 'Shields, J.K. 1953. A survey of the aquatic and marsh vege- tation of some lakes in the Muskoka and Parry Sound Districts of Ontario. A report prepared for the Toronto Fish and Game Protective Association, Toronto. "Soper, J.H. 1948. A survey of the aquatic vegetation of Whitewater Lake, with special reference to its suitability to waterfowl. A report prepared for the Department of Lands and Forests of Ontario, Toronto. 13 pp. mimeo. ductivity of 56 wzmho/cm. The other seven lakes have alkalinities below 5 mg CaCO;/L and conductivities in the range of 29-41 wmho/cm. All pH readings were in the range 6.0-6.6. Three Mile Lake has a notice- ably larger clay component in its substrate; reduced water clarity resulted in a 2.1-m Secchi disc reading compared with the 3.9—9.4 m in the other lakes. Lakes such as these characteristically have both a low productivity of aquatic macrophytes and a low standing crop when compared with lakes richer in dissolved solids. It 1s not known whether these values have changed between the two sampling dates. Differences in the recorded flora are assessed as attributable to sampling technique, to man’s influence on the environment, or to successional trends in the vegetation. Methods Three two-day surveys (by motor boat) were made of each lake by a team of three in the summer of 1976. Several stands (10 to 20) were carefully surveyed using SCUBA or skin-diving equipment. Sampling sites were chosen so that areas fully described in the former survey on the maps of Shields! were revisited. In addition, sites of unique features as well as ones characteristic of the lake were included. The vegeta- tion in an area 1000 m2? was surveyed and all submersed and floating aquatic plant species were rated for abundance on the scale: 0, absent to 4, abundant. Each species was given a mean abundance rating for each lake using the same scale. This index indicated the frequency of an expected encounter. The herbarium at the University of Guelph (OAC) houses a sample of each species. Five of the eight lakes were revisited in 1977 to check typical locations for species apparently ‘lost’ since 1953. 386 1979 MILLER AND DALE: MACROPHYTE FLORAS OF MUSKOKA LAKES 387 80°W 79°'W (1) \ NORTH BAY LAKE NIPISSING BEATTY LAKE ro) 46°N t) 4 Trout Creek 46°N EAGLE L HORN LAKE D Burk’s Falls RAINY LAKE P BUCK LAKE PARRY SouND 4%. 44KE SKELETON ey) LAKE THREE MILE LAKE GEORGIAN BAY 45°N 45° N Gravenhurst 80°W 79°W FIGURE |. Location of the eight lakes sampled. Alternate names for some of these lakes are Rainy (Bartlett). Horn (Sollman). and Eagle (Machar). 388 Results Forty-seven species were identified; this is 12 more than in 1953! (Table 1). There was a loss of a single species of Pondweed (Potamogeton obtusifolius). Five species present in the total flora were missing from two or more lakes; of these, four produce floating leaves and the fifth, a bladderwort, is chiefly subter- ranean but becomes noticeable when it produces yellow aerial flowers. Discussion Although the methods of sampling and recording the data differed, the two studies are comparable. The thorough survey, using a rowboat and rake to determine species present and using symbols to record on maps, produced a complete sampling for the parts of the lake visible from the surface. The recent survey used the same maps to check stretches of shallow water barren of plants in 1953 as well as areas in which a complex of species was found. These lakes have long stretches with few plants and the richest communities consisted of less than a dozen submersed and floating species. The numerous sites were surveyed by wading and swimming by a team of three but there is no reason to suggest that this was an incomplete survey of the shallow-water communities. The major difference in the surveys was in a more complete sampling of the deeper water using SCUBA equipment. Of the plant species that were missing from two or more lakes, Batchelder (G/yceria borealis), a plant of small scattered patches, is difficult to identify after the release of seed, as is the bladderwort (Utricularia cornuta) if it is not in flower. One site of the latter was lost when a highway was built over a small part of Eagle Lake. Three others, the floating-leafed species, must be lost from the lake floras. These very visible species could not be overlooked and their loss was due to severe reduction of their populations, perhaps because of the increased use of outboard boat motors. The effect of the propellors would be similar to that of a mowing machine constantly cutting away the leaves. The number of buildings on one stretch of shore in Skeleton Lake increased from 102 1n 1956-1960 to 177 in 1970 (Map 31 E/3 west. Ist edition, 1960 and new edition, 1974. National Topographical System, Canada). The decades between surveys were also the years of great water skiing activity in the Muskoka lakes. Plants recorded only in the second survey were considered indicators of succession if found in two or more lakes. There has been no study to show whether Quillwort (/soetes macrospora), Big-leaf Pondweed (Potamogeton amplifolius), and Arrowhead (Sagit- taria graminea) could have increased dramatically in abundance in the interval. It is probable that these THE CANADIAN FIELD-NATURALIST Vol. 93 species were present but only in deep water and were not reached by the rake in the first sampling. For the same reason, six species — Quillwort, Arrowhead, Water moss ( Fontinalis sp.), Bushy Pondweed ( Najas flexilis), Big-leaf Pondweed, and Purple Bladderwort (Utricularia purpurea) — could also have been un- recorded. The last species was unrecorded in Law- rence Lake from a surface-based sampling (Rich et al. 1971) although it was common in part of the lake in 1976 (H. M. Dale and D. H. N. Spence, unpublished data). Two of the new species, the bladderworts U. resupinata and intermedia, are obscure and could have been overlooked in 1953. The remaining species, however, are widespread shallow-water species, read- ily seen and identified. These species, therefore, have become more conspicuous or are new elements in the flora. Along the southern shore of Three Mile Lake there is a horseshoe-shaped bay described by Shields! on his maps as containing a dozen species including six with floating leaves. This location was found to be unchanged in 1976 and matched the earlier descrip- tion exactly. In contrast, a site along the western shore of Eagle Lake was marked clearly in 1953 as occupied by only Seven-angled Pipewort (Eriocaulon sept- angular) and Water Lobelia (Lobelia dortmanna), yet in 1976, a very conspicuous population of Narrow Floating-leaf Bur Reed (Sparganium angustifolium) was growing from a dense mat of the former two species. Also present were Needle Rush (Eleocharis acicularis), Rush (Juncus pelocarpus), and Waterwort (Elatine minima), members of the promiment wide- spread new species. Similarly in Horn Lake, Floating- heart and Floating-leaf Bur Reed had invaded a Pipewort-Lobelia mat. In this type of sandy oligotrophic lake, Seven- angled Pipewort and Water Lobelia act as colonizing species and form patches that stabilize the coarse substrate, trap fine particles, and increase the organic matter of the substrate. These patches grow and coalesce, forming a modified environment suitable for colonization by Bur Reed. Waterworts, Rush, Needle Rush, and Floating-heart. Such a successional pro- cess in the vegetation produces a more complex vegetation by increasing the abundance of the rarer species. Some differences between the surveys 23 yr apart may be explained by improved sampling methods: sampling with the use of a rake is less accurate than using SCUBA equipment. The floating-leafed Yellow Water Lily, Water Shield, and Floating-heart have been lost as a result of recreational use of the water: other species increased in abundance as a result of succession. 1979 MILLER AND DALE: MACROPHYTE FLORAS OF MUSKOKA LAKES 389 TABLE |—Prescence (+) or abundance (R—rare, O—occasional, C—common, A—abundant) of aquatic macrophytes at the two sampling dates (1953, 1976) in each of eight lakes. Common names are from Fassett (1957) Three Skeleton Axe Buck Rainy Horn Eagle Beatty Mile Lake Lake Lake Lake Lake Lake Lake Aquatic macrophytes os Ho 33 WO s3 oO 33 Oo 933-10 ~ 33 10 33 1 38 16 Water Moss (Fontinalis sp.) cnn Rec erect tome FY O (S O Quillwort (/soetes macrospora) O C C A Cc A + Narrow Floating-leaf BurReed (Sparganium angustifolium) R C + Floating-leaf BurReed (Sparganium fluctuans) + R + O OF Big-leaf Pondweed (Potamogeton amplifolius A Narrow-leafed Pondweed ( Potamogeton berchtoldii) Ribbon-leaf Pondweed (Potamogeton epihydrus) ap JR ae Variable Pondweed ( Potamogeton gramineus) + Floating Brownleaf (Potamogeton natans) tone Ati: Small Floating-leaf Pondweed ( Potamogeton oakesianus) Pondweed (Potamogeton obtusifolius) + Sago Pondweed (Potamogeton pectinatus) R Clasping-leaf Pondweed ( Potamogeton richardsonii) + Robbins’ Pondweed ( Potamogeton robbinsii) Pondweed (Potamogeton spirillus) Bushy Pondweed (Najas flexilis) Arrowhead rosettes (Sagittaria graminea) + Canada Waterweed (Elodea canadensis) Wild Celery (Vallisneria americana) Batchelder (Gl/yceria borealis) Wild Rice (Zizania aquatica) Needle Rush (Eleocharis acicularis) A Cc R Triangle Spike Rush (Eleocharis robbinsii) Water Bulrush (Scirpus subterminalis) Water Arum (Calla palustris) Seven-angled Pipewort (Eriocaulon septangulare) romeo O) bmi ttt ane Cpa Pickerel Weed (Pontederia cordata) Bayonet Rush (Juncus militaris) Rush (Juncus pelocarpus) Smartweed (Polygonum amphibium) + Coontail (Ceratophyllum demersum) Water-shield (Brasenia schreberi) + Yellow Water Lily (Nuphar variegatum) White Water Lily (Nymphaea odorata) Creeping Spearwort (Ranunculus reptans) Water Starwort (Callitriche heterophylla) 1 Waterwort (Elatine minima) Mares-tail (Hippuris vulgaris) + Farwell’s Milfoil (Myriophyllum farwelli) Milfoil (Myriophyllum tenellum) + Floating-heart (Nymphoides cordatum) + Bladderwort (Utricularia cornuta) Bladderwort (Utricularia gibba) Bladderwort (Utricularia intermedia) Purple Bladderwort (Utricularia purpurea) Bladderwort ( Utricularia resupinata) Common Bladderwort (Utricularia vulgaris) an Reet Water Lobelia (Lobelia dortmanna) + erie Cathe ae 3 Number of changes/ total records 6/22 7/20 10/1 QO 7 non OS) + rm), Oc) cd + O + Oo + ‘o) Chat: QAANFONF OH nw >aAFRXHO > nw + > + + Oe. © + ++ + ARN S + fe OS a Op Of + a + ee) + + Aw + of + A000 + >On + y@ eRe) On i @Q gw cw 2 + AO + + + + + > SOS AaQae CeECoQaaqd@a o) + o) + + © NOFFRFONN 12/25 18/32 9/17 l oS) tO tN oo ~ 390 Acknowledgments We are grateful to R. Dreimanis and B. Elder for their cheerful assistance during the initial sampling. This study was partially supported by National Research Council of Canada grant 3402. Literature Cited Dale, H. M. and G. E. Miller. 1978. Changes in the aquatic macrophyte flora of Whitewater Lake near Sudbury, Ontario from 1947 to 1977. Canadian Field-Naturalist 92: 264-270. Fassett, N.C. 1957. A manual of aquatic plants. 2nd edition, with revision appendix by E. C. Ogden. Uni- versity of Wisconsin Press, Madison, Wisconsin. Harman, W.N. and T.R. Doane. 1970. Changes in the aquatic flora of Otsego Lake between 1935-1939. New York Fish and Game Journal 17: 121-213. Lind, C. T. and G. Cottam. 1969. The submerged aquatics of University Bay: A study in eutrophication. American Midland Naturalist 81: 359-369. Rich, P.H., R.G. Wetzel, and N. Van Thuy. 1971. Distribution, production and role of aquatic macrophytes THE CANADIAN FIELD-NATURALIST Vol. 93 in a southern Michigan marl lake. Freshwater Biology 1: 3-21. Schmid, W. D. 1965. Distribution of aquatic vegetation as measured by line intercept with SCUBA. Ecology 46: 816-823. Spence, D.H.N. 1964. The macrophytic vegetation of freshwater lochs, swamps and associated fens. Jn The vegetation of Scotland. Edited by J. H. Burnett. Oliver and Boyd, Edinburgh and London. pp. 306-425. Stuckey, R. L. 1971. Changes of vascular aquatic flowering plants during 10 years in Put-in-bay Harbour, Lake Erie. Ohio. Ohio Journal of Science 71: 321-342. Volker, R. and S. G. Smith. 1965. Changes in the aquatic vascular flora of Lake East Okoboji in historic times. Proceedings of the Iowa Academy of Science 72: 65- 12. West, G. 1905. A comparative study of the dominant phanerogamic and higher cryptogamic flora of aquatic habit in Scottish lakes. Proceedings of the Royal Society of Edinburgh 25: 967-1023. Received 17 January 1979 Accepted 28 March 1979 Utricularia geminiscapa at Mer Bleue and Range Extensions in Eastern Canada* ERICH HABER Botany Division, National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8 Haber, Erich. 1979. Utricularia geminiscapa at Mer Bleue and range extensions in eastern Canada. Canadian Field- Naturalist 93(4): 391-398. The first Ontario occurrence of Utricularia geminiscapa at the Mer Bleue peat bog just east of Ottawa is reported. The main features that help to distinguish this species from U. vulgaris are tabulated and the problem of differentiating vegetative specimens of these two species is discussed. Additional range extensions for U. geminiscapa in eastern Canada of up to 600 km north of previously known localities are also reported and plotted on a distribution map of the species for North America. Key Words: Utricularia geminiscapa, morphology, range extensions, Ontario, Canada. The discovery of an Ontario locality for the Research Council fellow with the National Museum bladderwort Utricularia geminiscapa occurred onone _ of Natural Sciences, and I were returning to our access of several excursions made in 1973 to Mer Bleue, a __ point at the tip of Dolman Ridge when hé noticed peat bog situated approximately 16km east of some flowering Utricularia in a small bog pool. Parliament Hill, Ottawa (Figure 1). David R. Given Because it was obviously different from the U. from New Zealand, who was then a National intermedia we had seen earlier, we made a repre- FIGURE 1. Mer Bleue peat bog, Ontario. Dolman Ridge is the northern spit of land extending into the bog from the left, Borthwick Ridge the southern. *Contribution Number 9 to “Scientific and Cultural Studies of the Mer Bleue.” 39 392 sentative collection. The identity of this species was readily determined at a later date because it had the characteristic chasmogamous and_ cleistogamous flowers (Figure 2a). Indeed the seeds from the cleistogamous flower capsules (Figure 2b) proved to be extremely characteristic and easily referable to U. geminiscapa by comparison with seed illustrations in Muenscher (1944). This initial encounter with U. geminiscapa, a species new to the province, prompted me to examine the morphology and habitat character- istics of this bladderwort in more detail and to search for additional range extensions in eastern Canada. Morphological Distinctions Although U. geminiscapa is usually quite distinct from U. vulgaris, particularly when flowers or capsules are present, purely vegetative specimens of these species may at times be difficult to identify unequivocably. The vegetative plumes of foliage of small specimens of U. vulgaris can approach in width those of U. geminiscapa, which are usually much narrower than those of U. vulgaris. The identification of vegetative material is further complicated by the lack of a clear-cut distinction in the form and nature of the leaf branching in the two species. In U. gemi- niscapa, the branching is dichotomous (Figure 3a) and is usually distinct from the pseudopinnate branching found in normal robust specimens of U. tT eee “rf eee 2 aN — | 4 va . é vg ’ \ ae e \ Sie 2 Sem THE CANADIAN FIELD-NATURALIST Vol. 93 vulgaris (Figure 3d). This distinction, however, is not always evident because smaller specimens of U. vulgaris may exhibit leaf branching similar to that of U. geminiscapa. Such vegetative characteristics are often almost obliterated by careless mounting pro- cedures in the preparation of herbarium specimens. Fresh specimens should be float-mounted prior to drying to display clearly the nature of the leaf branching. Several features that can be used to distinguish these two species are given in Table 1. The supposed absence of marginal leaf spines in U. geminiscapa has been used as a criterion for distin- guishing this species from U. vulgaris which is characteristically spinulose-margined (Muenscher 1944: Fassett 1957: Roland and Smith 1969). The fresh specimens from the Mer Bleue locality leave no doubt that the leaf segments can be spinulose-margined (Figure 3a). The abundant herbarium specimens from the Farnham bog (Missisquoi County, Quebec) also confirm this observation. The leaves of the winter buds are also slightly spinulose and are terete in section, as in normal leaves. They forma loose, one-sided aggregate and maintain their green color when in bud (Figure 3b). In U. vulgaris the bud leaves are usually conspicuously flattened, highly dichotomized and very spinulose with the whole bud appearing grayish- white owing to the abundance of the solitary or fasciculate spines. The bud forms a dense, round to a - & \ — \ oe \ Pe t et eA # Y ae = 3 © x : ty . 2 * te es tte : ia, ; Y a _ x x SO . - * * oe 12 7 3 aes a yes ‘ Lae & = ys ‘A ‘\ : ov h : = 2 wt : Pe 4 * Oy < 9 hve a ees Y a FIGURE 2. Utricularia geminiscapa: (a) whole plant with normal flowers and cleistogamous capsules, the latter indicated by arrows; (b) seeds from cleistogamous capsules. 1979 - HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA 393 FIGURE 3. Leaf branching pattern and winter bud characteristics in Utricularia. Utricularia geminiscapa: (a) spiny leaf segments in a fresh specimen; (b) portion of leafy plume of a fresh specimen with terminal winter bud (the one-sided aspect of the bud is not evident in this photo). Utricularia vulgaris. (c) portion of leafy plume of a dried specimen with terminal, highly spinulose winter bud; (d) leaf branching pattern in a dried specimen. 394 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Characters useful in distinguishing the two species of Utricularia Character filiform, without scales corolla 0.5-0.8 cm long without basal lobes Aerial scapes Flowers Flower bracts Cleistogamous flowers U. geminiscapa present, apetalous, scattered along stem U. vulgaris coarse, with several scales corolla 1.5-2.5 cm long with basal lobes absent axils and producing abundant seeds discoid 1.0-3.5 cm in diameter dichotomous Seeds Foliage plumes Leaf branching Leaf segments spinulose Winter buds ingly spinulose long acuminate with or without terminal spine(s); margins glabrous to sparingly 0.2-0.5 cm in diameter, green, one-sided; leaf segments terete, glabrous to spar- angular with many facets 3-12 cm in diameter dichotomous to usually + pinnate spine-tipped with single or fasciculate spines; margins + abundantly spinulose 1-3 cm long; dense, round to oval mass, occasionally lobed; leaf segments; flat- tened and highly spinulose oval, sometimes lobed mass of bud leaves (Figure 3c). The best way of ensuring that vegetative specimens of these two species can be identified with some certainty is to collect from a given locality samples including as wide a size variation of the vegetative plumes as possible. Although U. intermedia also occurs in bog pools at Mer Bleue, it can be readily distinguished from U. geminiscapa and as well from U. vulgaris by its flattened, dichotomous branching and by the presence of a mid-vein in the leaf segments. Characteristics of the Mer Bleue Locality A search of herbaria in Ontario and Quebec indicated that the Mer Bleue site is probably the first verified locality for U. geminiscapa in Ontario (see also Morris 1922). In order to gain an insight into its frequency of occurrence at the bog, I returned during October and early November of 1978. The water level in the fall of 1978 was somewhat lower than when the first collection was made (17 July 1973) with the result that the original shallow pool was no longer present. In its place were several small potholes with little standing water and no discernible evidence of the bladderwort. The yearly and seasonal water table fluctuations, therefore, appear to have some influence on the localized development of U. geminiscapa in shallow bog pools. Presumably low water levels would not eradicate the species from such shallow, superficial bog pools because of the presence of the resistant winter buds which could lie dormant in the moist Sphagnum mat until open water was again available. Potential sites for the bladderwort could be readily located by inspecting aerial photographs of Mer Bleue. The pools of varying size that are evident on the aerial photographs are distributed primarily in the vicinity of the wooded islands in the central region of the bog (see Figure 4). Most if not all of these are bomb craters formed when the bog was used as a practice bombing range between 1942 and 1945 (S. Ashley, 1979. The Mer Bleue — The evolution of an urban bog. An oral history of the Mer Bleue and surrounding communities. An in-house publication, National Capital Commission, Ottawa, Ontario; Freeman 1969). The latest aerial photographs taken in 1978 (Figure 4 in part) indicate that virtually all of the pools evident on the earliest photographs taken in 1945 are still present. x Five of the six bomb crater pools surveyed briefly in the fall of 1978 contained the bladderwort (sites 6, 9, 12, 13, 14; Figure 4). One of these pools (site 14) located in a predominantly open area with scattered Tamarack (Larix laricina) is illustrated in Figure 5. These deep, bomb crater pools contain few aquatics and are usually fringed by a relatively firm Sphagnum mat stabilized by ericoid shrubs, in particular by Leatherleaf (Chamaedaphne calyculata). The pres- ence of U. geminiscapa in nearly every pool sampled would indicate that the unorthodox use of the bog asa bombing range may have benefited the species by increasing the number of habitats suitable for coloni- zation. These deep pools would also serve as favored habitats for this bladderwort because low water-table levels would not eliminate such pools. Water samples were collected from each pool surveyed and the pH compared with bog water obtained from within the bog mat at various sites. The location of these samples is indicated in Figure 4. Summarized in Table 2 are the pH measurements which were obtained in the laboratory at room temperature with a Beckman 180 pocket pH meter. The pH values obtained ranged from 3.9 to 6.1. Sites 1-5 and 7 with pH values ranging from 3.9-4.] are perhaps most representative for the bog mat, judging from reports by Terasmae and Mott (1964, pH 3.8) 1979 HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA 395 FIGURE 4. Original collection locality for Utricularia geminiscapa (asterisk). It was subsequently found at sites 6, 9, 12, 13, 14. Each numbered site represents a sample locality for pH determinations. FIGURE 5. Northward view of one of the deep, bomb crater pools (site 14) in which Utricularia geminiscapa was found. Photographed in the fall of 1978. 396 TABLE 2—Acidity (pH values) of pools and bog mat at various sites in the Mer Bleue peat bog Site pH I open mat 4.1 2 open mat 4.0 3 forested bog 39) 4 forested bog 38) 5 sedge zone 4.0 6 bog pool 4.5 7 sedge meadow 4.1 8 bog pool 6.1 9 bog pool 5.4 10 open mat 4.7 11 sedge-leatherleaf mat 4.6 12 bog pool 49 13 bog pool 4.9 14 bog pool 47 15 sedge-leatherleaf mat 4.4 16 marginal channel (lagg) 4.6 and Joyal (1971, pH 3.8 and 3.9). The bog pools ranging from 4.5 to 6.1 had consistently higher values than sites in the bog mat. The remaining sites (10, 11, 15, 16) also had higher values (4.44.7) than the more usual value of about 3.9 for the bog mat. Measure- ments previously taken in June 1974 in the region east of site 16 and extending to the first small island yielded pH values between 5.1 and 5.4. The significance of the higher pH levels of the bomb crater pools and other sites in the central region of the bog is not clear until one also takes into account the localized enrichment of the bog flora that occurs in this area. Some of the species that occur here are also common elements of the peripheral eutrophic zone. This central, primarily treeless, mesotrophic region is represented in Figure 4 as the dark elongate band extending eastward from the southeastern tip of Dolman Ridge to the first small island and again as the major band surrounding and trailing eastward from the large central islands. Cat-tails (Typha latifolia), Bogbean (Menyanthes trifoliata), Water Arum (Calla palustris), Chain Fern (Woodwardia virginica), and Scheuchzeria palustris occur 1n this mesotrophic region. The White Fringed Orchid (Platanthera blephariglottis) is also common along the southern, forested edge of this area east of the large islands (see also Lafontaine 1971). Speckled Alder (Alnus rugosa), a common bog fringe species, occupies a sizable shrubby area between the western ends of the two large islands, and species of sedges form extensive meadows in this central region and add to the fen-like appearance. The generally higher pH values and apparent mesotrophic nature of this central region may be THE CANADIAN FIELD-NATURALIST Vol. 93 directly related to the ground-water levels and shallow nature of the underlying central basin (maximum depth about 2 m) (see map of peat depths in Nystrom and Anrep 1909). During periods of drought as in 1877 and 1878 (The Ottawa Naturalist 1888, 2: 74) and low ground-water levels as in 1970 and 1974 (M. Bik, 1975. In Proceedings of the Mer Bleue Seminar. An in-house publication, National Capital Commission and Central Research Forest, Ottawa, Ontario), the water level would decrease over the central shallow basin and aeration and substrate temperatures would increase resulting in a temporary eutrophication. Subsequent periods of high ground-water levels would cause a reversai of these conditions. This mesotrophic zone lying between two deeper basins (maximum depths about 6 m) may in effect represent a tension zone providing suitable habitats for certain species of plants. The relatively large band of Typha within this zone east of the large islands may represent, together with the peripheral 7ypha zone, the remnants of an earlier, more extensive marshland habitat. In Camfield’s pollen diagram (1969), Typha pollen was most abundant in one of the oldest zones in her Mer Bleue core. Although the earliest organic sediments in Mer Bleue are approximately 7500 yr old (Camfield 1969), the arrival of U. geminiscapa probably dates to a much more recent time commensurate with the formation of suitably acidic bog pools. Throughout its range this species occurs most commonly in quiet, acidic waters and in particular in bog pools. The difference in habitat preferences between U. gemi- niscapa and U. vulgaris is readily apparent at Mer Bleue where the latter occurs in shallow ditches adjacent to the bog but not directly influenced by the acid Sphagnum mat. Throughout its widespread range, U. vulgaris occurs in ponds, lakesides, and sluggish streams. The occurrence of U. geminiscapa at Mer Bleue is simply one more interesting species addition to an increasing list of novelties either first described from the bog or represented regionally only at this locality. Additional information regarding various aspects of this locality can be found in the papers by Baldwin and Mosquin (1969), Corbet and Walley (1969), Dunston (1970), Smith (1970a, b), Hobson(1970), Ouellet et al. (1976), and Small (1976). Range Extensions in Eastern Canada A search of herbaria in Ontario and Quebec (CAN, DAO,. TRT, QFA, QSA, MT, MIMG, and SEs: acronyms according to Holmgren and Keuken (1974)) resulted in the validation of specimens from seven Canadian localities in addition to the Mer Bleue record. The eight validated localities of U. gemi- niscapa plotted on the map in Figure 6 are as follows: FIGURE 6. North American distribution of Utricularia geminiscapa. The generalized range of this species, derived from maps published by Fernald (1933), Thomson (1940), Muenscher (1944), and Roland and Smith (1969), is represented by the hatched areas. The solid dots represent new localities based on verified specimens. Ontario: Mer Bleue peat bog, 45°24’N, 75°30’W, The Regional Municipality of Ottawa-Carleton, bog pool, E. Haber & D. R. Given 1963, 17 July 1973 (CAN). Quebec: plaine de la Riviere Ouelle, cté Kamouraska, E. Campagna, 13 July 1928 (QSA); tourbiére de Farnham, cté Missisquoi, M. Raymond, 20 Aug. 1933 (CAN) [the earliest of many collections from this site]; L. Hibou, 5 mi[3 km] southwest of Taschereau, W. Abitibi Co., small boggy lake, W. K. W. Baldwin & A. J. Breitung 4287, 29 Aug. 1952 (CAN); entre St- Faustin et Lac Supérieur, Lac artificiel Dupré, cté Terrebone, Frére Rolland-Germain 422, 23 July 1957 (CAN); Ilets-Jérémie, cté Saguenay, dans les petites mares, tourbiére a cypéracéés, J. Cayouette 73-719, 14 Aug. 1973 (QFA); Hopkins Hole, between Ramsay L. and Hawley L., Gatineau Pk., Gatineau Co., A. & J. Reddoch, 19 Aug. 1978 (CAN). The most northerly range extensions for U. gemi- HABER: UTRICULARIA GEMINISCAPA IN EASTERN CANADA 397 niscapa occur at Lake Hibou (48°38’N, 78°48’W) and at Ilets-Jérémie (48°53’N, 68°48’W) jn Quebec. Both localities are approximately 600 km north of pre- viously documented localities in upper New York State and Maine, respectively. Although the Lake Hibou specimens were originally determined as U. minor by Baldwin, they are sufficiently characteristic in spite of their vegetative condition that they were validated as U. geminiscapa. A number of perplexing collections made by Frére Rolland-Germain at Lake Monroe in Montcalm County, Quebec and identified as U. geminiscapa (CAN, MT, SFS) are considered to be merely small vegetative specimens of U. vulgaris. This decision was reached only after seeing a whole series of vegetative specimens of U. vulgaris from the same locality at the Marie-Victorin herbarium (MT). These specimens ranged from the more typical large-plumed specimens to those that were much reduced in size approaching U. geminiscapa in appearance. Utricularia geminiscapa may indeed be much more widely spread in eastern North America than is at present documented. The vegetative characteristics of U. vulgaris such as overall size of leaf plumes, manner of leaf branching, form and pubescence of winter bud leaves, and leaf pubescence are extremely variable and require further investigation over the whole range of this cosmopolitan species. The cultivation of this species under controlled environmental conditions may prove useful in accounting for some of the observed natural variation. Literature Cited Baldwin, W. K. W. and T. Mosquin. 1969. Scientific and cultural studies of the Mer Bleue. Canadian Field- Naturalist 83(1): 4-6. Camfield, M. 1969. Pollen record at the Mer Bleue. Canadian Field-Naturalist 83(1): 7-13. Corbet, P.S. and G.S. Walley. 1969. The dragonflies (Odonata) of the Mer Bleue. Canadian Field-Naturalist. 83(1): 14-16. Dunston, J. 1970. The Mer Bleue bog. Trail & Landscape 4(1): 26-31. Fassett, N. C. 1957. A manual of aquatic plants. University of Wisconsin Press, Madison. 405 pp. Fernald, M. L. 1933. Recent discoveries in the Newfound- land flora. Rhodora 35: 1-16, 47-63, 80-107, 120-140, 161-185, 203-223, 230-247, 265-283, 298-315, 327-346, 364-386, 395-403. Freeman, T. N. 1969. Some historical notes on collecting Lepidoptera in the Mer Bleue. Canadian Field-Naturalist 83(1): 16-18. Hobson, G. D. 1970. Bedrock features of the Mer Bleue area by seismic methods. Canadian Field-Naturalist 84(1): 35-38. Holmgren, P. K.and W. Keuken. 1974. Index herbariorum Part 1. The herbaria of the world. 6th edition. Regnum Vegetabile 92: 1-397. 398 Joyal, R. 1971. La tourbiére a sphaignes Mer Bleue prés d’Ottawa. II. Quelques facteurs écologiques. Canadian Journal of Botany 50: 1209-1218. Lafontaine, J.D. 1971. What ever happened to Habenaria blephariglottis? Trail & Landscape 5(5): 134-136. Morris, F. 1922. Bladderworts of Ontario. Canadian Field- Naturalist 36(7): 123-126. Muenscher, W.C. 1944. Aquatic plants of the United States. Comstock Publishing Co. Inc., Cornell University, Ithaca, New York. 374 pp. Nystrom, E. and S.A. Anrep. 1909. Investigation of the peat bogs and peat industry of Canada, during the season 1908-1909. Canada Department of Mines, Mines Branch, Bulletin 1, Report 30. Ouellet, H., S. J. O'Donnell, and R. A. Foxall. 1976. Gray Jay nesting in the Mer Bleue Bog, Ottawa, Ontario. Canadian Field-Naturalist 90(1): 5-10. Roland, A. E. and E. C. Smith. 1969. The flora of Nova THE CANADIAN FIELD-NATURALIST Vol. 93 Scotia. Part 2. The dicotyledons. Proceedings of the Nova Scotian Institute of Science 26(4): 277-746. Small, E. 1976. Insect pollinators of the Mer Bleue peat bog of Ottawa. Canadian Field-Naturalist 90(1): 22-28. Smith, D. A. 1970a. Notes on bog ecology. Trail & Land- scape 4(5): 135-143. Smith, D. A. 1970b. Observations on nesting Hawk Owls at the Mer Bleue, near Ottawa, Canada. Canadian Field- Naturalist 84(4): 377-383. Terasmae, J. and R. J. Mott. 1964. Pollen deposition in lakes and bogs near Ottawa, Canada. Canadian Journal of Botany 42: 1355-1363. Thomson, J. W., Jr. 1940. Preliminary reports on the flora of Wisconsin. 27. Lentibulareaceae. Wisconsin Academy of Sciences, Arts and Letters 32: 85-89. Received 13 February 1979 Accepted 24 April 1979 atl al Movements of Sympatric Species of Snakes at Amherstburg, Ontario B. FREEDMAN! and P. M. CATLING2 'Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario L5L 1C6 Present address: Department of Biology and Institute for Resource and Environmental Studies, Dalhousie University, Halifax, Nova Scotia B3H 4J1 2Department of Botany, University of Toronto, Toronto, Ontario M5S 1A1 Freedman, B. and P. M. Catling. 1979. Movements of sympatric species of snakes at Amherstburg, Ontario. Canadian Field-Naturalist 93(4): 399-404. In order to investigate the movements of Butler’s Garter Snakes (Thamnophis butleri), Eastern Garter Snakes ( Thamnophis sirtalis sirtalis), Brown Snakes ( Storeria dekayi), and Eastern Fox Snakes (Elaphe vulpina gloydi), we systematically searched for, captured, and individually marked by scale-clipping these species. Recaptured snakes were found to have moved relatively short distances, with over 50% of marked snakes being recaptured less than 50 m from the initial point of capture after various recapture intervals. There was no evidence of large-scale emigration of snakes from areas inhabited in the early spring. For all species, only a small percentage of the population demonstrated relatively long-distance (about 400 m) unidirectional movements. There were no consistent differences in movements between sexes. More female 7. butleri were recaptured than males. No directional trends in movements were observed, but a tendency to avoid road-crossing was noted for T. butleri. In cases of multiple recapture, minimum activity ranges are presented. Key Words: Thamnophis butleri, Thamnophis s. sirtalis, Storeria dekayi, Elaphe vulpina gloydi, movements, activity range, southwestern Ontario. Gregory and Stewart (1975) described dispersal of the Red-sided Garter Snake (Thamnophis sirtalis parietalis) following spring emergence from karst hibernacula in the Interlake District of Manitoba. There are no other published data on the movements of snakes in Canada, although these are an important part of general ecological understanding and are critical for the conservation of populations of rare animals. The purpose of our study was to describe move- ments of four sympatric species of snakes and to analyze these with respect to the extent of suitable habitat required, and the sizes of areas that must be protected to ensure the survival of populations. The study area, 2.4km NE of Amherstburg, Essex County, Ontario, has been described previously (Freedman and Catling 1978). Two of the species studied, Butler’s Garter Snake (Thamnophis butleri) and the Eastern Fox Snake (Elaphe vulpina gloydi) are considered rare or endangered in Canada (Cook 1970, 1977; Gregory 1977). The other two species, the Eastern Garter Snake (Thamnophis sirtalis sirtalis) and the Brown Snake (Storeria dekayi), are wide- spread and common. Methods We searched systematically for snakes along pre- determined census routes during a series of visits to the study area in the spring and summer of 1976 (14 and 30 May, 15 June, 10 and 24 July). All snakes captured were marked by clipping scales (Freedman and Catling 1978), and the point of capture was 399 recorded on _ large-scale aerial photographs (1 cm = 10 m). Sex and total length were also record- ed for each snake to allow comparisons among sexes and size classes (Freedman and Catling 1978). All snakes were released where captured within 15 min. Young snakes less than | yr old and young-of-the- year (defined by body length) were difficult to mark by scale-clipping, and were not included. Snakes less than | yr old were relatively uncommon (Freedman and Catling 1978). For all captured snakes the distance between the recapture and the most recent point of capture was determined to give a minimum distance travelled. In cases of more than one recapture of an individual snake, these minimum distances were also summed to. give a total minimum distance travelled. In the cases of multiple recapture, minimum activity ranges were calculated from the areas enclosed by the joined points of capture and recapture. We also considered the possibility that roads could restrict movements of snakes; we noted whether snakes crossed an open 10- m-wide gravel road. Results and Discussion Out of a total of 250 individuals of T. butleri, 72 T. s. sirtalis, and 102 S. dekavi, a minimum of 10 recaptures was made per species, and some indi- viduals were recaptured more than once. All of these species exhibited a strong tendency for relatively short-distance movements, with more than 50% of all marked individuals being recaptured less than 50 m from the initial point of capture after various intervals 400 of time (Figures 1-3). In general, total distance moved by all three species increased the longer the time interval between initial capture and subsequent recapture(s). This tendency is illustrated for T. butleri (Figure 1b-d), for which there were 25 recaptures. For each of the three species, there were no consistent differences in movement between the sexes (Table 1), Although female S. dekayi appear to move less than males, the sample of males is only two. More female 7. butleri were recaptured than males (Table () uJ a =) fS a q (S) uJ a ep) uj x dq a 2p) iw (e) (e) 2 THE CANADIAN FIELD-NATURALIST Vol. 93 1),, even though the proportion of males in the population was about 0.56 (Freedman and Catling 1978). Possibly female 7. butleri are less mobile and have a greater tendency to bask in the open (Gregory 1975) and are consequently more often captured than males. The S. dekayi covering the greatest distances were a male 31 cm long covering 374 m in 30 d, and a gravid female 39 cm long covering 226 m in 30 d, both single recaptures. The 7. butleri covering the greatest Thamnophis butleri All recaptures N=24 LI Recaptures after |3-I6days N=7 pay Recaptures after 30-40 days Recaptures after 54-70 days N=1l0 P in 430-440 510-520 MINIMUM MOVEMENT FIGURE 1. Minimum movements of marked Thamnophis butleri: (a) all recaptures during 1976; (b,c, d) recaptures at specific time intervals following release. Data include 20 individuals, of which two were recaptured twice and one was recaptured three times. 1979 FREEDMAN AND CATLING: DISTANCE MOVEMENTS, SYMPATRIC SNAKES 401 (a) a5 Thamnophis sirtalis sirtalis > a, eee = ae dq Oo uJ a wn > N=12 uJ x = 2 (ep) w oc | fe) 2 Oo O O oO O oO O O oO Oo O O O = aN) no) t wo op) O = (aN) ~m Tt wo oO Sir ee ee eee ie Ia oe thane ap sh Sli ona (e) O Oo (e) FIGURE 2. Minimum movements of marked Thamnophis sirtalis sirtalis from all 1976 recaptures. Data include 11 indi- viduals, one recaptured twice. (a) uJ : or Storeria dekayi = par tia Panacea ceo = au 6 oO uJ rs N=10 op) uJ x 3200 ZWE 12 0 0 0 ZPF 157 132 8 2 ZMF 16 1] 0 0 ZWP 52 16 3 0 ZMP lS 7 l 0 SWE 38 4 0 0 SPF 300 85 9 2 SWP 6 2 0 0 'Species/location code given in Table 1. TABLE 7—Frequency of girdling of conifers by Porcupines on the Skelton area Frequency of leader death Species, location! 0 ; | 2 3 SWF 29 8 0 0 SPF 172 105 30 4 SWP 6 3 0 0 'Species/location code given in Table |. (Bannan 1955; Grillos and Smith 1959), and provide more foliage and a greater volume of inner bark than crowded trees. Because Porcupines generally feed on inner bark that is accessible from positions that can be maintained without undue exertion (Taylor 1935; Spencer 1964), open-crowned trees, which provide more large branches, allow Porcupines greater access to food. The phloem and foliage of conifers contain the greatest concentrations of fats and carbohydrates HARDER: TREES UTILIZED BY PORCUPINES 409 of the aerial tissues (see Kramer and Kozlowski(1960) for a review), and the tendency of Porcupines to feed on these structures within the tree crown where growth is most rapid may result in the consumption of the most nutritious food available. The quality of a food item is a complex property that involves not only its energy and nutrient content, but also the digestion and assimilation efficiency of its consumers (Longhurst et al. 1968), and the influence of secondary plant compounds (Freeland and Janzen 1974). In addition, an animal’s response to a particular stimulus, such as food with a specific nutrient content, can be greatly modified by its physiological state (Cabanac 1971). An awareness by Porcupines of chemical qualities is not required to explain the observed preferences for particular trees, because vigorous trees may be identified by obvious physical features. Chemical attributes, however, may be important within trees because Porcupines appear to select food items through olfaction (Murie 1926; Taylor 1935). The average diameters of used trees from both study areas (Table 3) fall within the range of size preferences reported for Porcupines from various regions of North America (Taylor 1935; Curtis and Wilson 1953; Krefting et al. 1962; van Deusen and Myers 1962). A predilection for a particular size class irrespective of geographic location or tree species lends support to the suggestion of Curtis and Wilson (1953) that Porcupines may be most adept at climbing trees with a DBH between 15 and 25 cm. Presumably trees can become too large to be readily climbed (Taylor 1935; Curtis and Kozicky 1944; Curtis and Wilson 1953), resulting in a decline in their use by Porcupines even though they may contain large quantities of high-quality food. Local differences in the size of used trees of the magnitude observed between the Zoratti and Skelton areas would not be expected, however, if Porcupines exercised size preferences solely related to the ease of climbing trees, unless the preferred size was unavailable. Storm and Halvorson (1967) have presented convincing evidence that the removal of bark by Porcupines greatly affects the vigor of a tree for up to 10 yr. If a forest, such as the leeward pure community on the Zoratti area, was heavily used for several years, the number of rapidly growing trees within the preferred size class would be reduced. Porcupines feeding in this forest during subsequent winters could, therefore, face a choice between slower-growing trees of the appropriate size or smaller vigorous trees. Consumers cannot afford to be selective when resources are limited (Emlen 1966, 1968). Relaxation of selectivity may have occurred on the more extensively used Zoratti area and resulted in the apparent preference for smaller trees. The trees used 410 on the Skelton area consequently may provide a better indication of the physical qualities of trees preferred by Porcupines. Acknowledgments This study was supported in part by the National Research Council of Canada. J. O. Murie and F. C. Zwickel kindly reviewed various drafts of this paper. Barney and Anne Zoratti, and Bill and Mary Skelton generously consented to my activities on their lands. V. A. Loewen and V. Aziz assisted with the collection of data on the Skelton area. Literature Cited Bannan, M. W. 1955. The vascular cambium and radial growth in Thuja occidentalis L. Canadian Journal of Botany 33: 113-138. Brander, R. B. 1973. Life-history notes on the porcupine in a hardwood-hemlock forest in upper Michigan. Michigan Academician 5: 425-433. Brown, C. L. 1971. Secondary growth. Jn Trees — Struc- ture and function. Edited by M. H. Zimmermann and C. L. Brown. Springer-Verlag, New York. pp. 67-123. Cabanac, M. 1971. Physiological role of pleasure. Science 173: 1103-1107. Curtis, J. D. and E. L. Kozicky. 1944. Observations on the eastern Porcupine. Journal of Mammalogy 25: 137-146. Curtis, J. D.and A. K. Wilson. 1953. Porcupine feeding on Ponderosa Pine in central Idaho. Journal of Forestry 51: 339-341. Daubenmire, R. F. 1974. Taxonomic and ecologic rela- tionships between Picea glauca and Picea engelmannii. Canadian Journal of Botany 52: 1545-1560. Emlen, J. M. 1966. The role of time and energy in food preference. American Naturalist 100: 611-617. Emlen, J. M. 1968. Optimal choice in animals. American Naturalist 102: 385-389. Freeland, W.J. and D.H. Janzen. 1974. Strategies in herbivory by mammals: the role of plant secondary com- pounds. American Naturalist 108: 269-289. Gabrielson, I. N. 1928. Notes on the habits and behavior of the Porcupine in Oregon. Journal of Mammalogy 9: 33-38. Gabrielson, I. N. and E. E. Horn. 1930. Porcupine control in the western states. United States Department of Agri- culture, Leaflet Number 60. 8 pp. THE CANADIAN FIELD-NATURALIST Vol. 93 Grillos, S. J.and F. H.Smith. 1959. The secondary phloem of Douglas-Fir. Forest Science 5: 377-388. Husch, B., C.I. Miller, and T. W. Beers. 1972. Forest ~ mensuration. 2nd Edition. Ronald, New York. 410 pp. Kramer, P. J. and T. T. Kozlowski. 1960. The physiology of trees. McGraw-Hill, New York. 642 pp. Krefting, L. W., J. H. Stoeckeler, B. J. Bradle, and W. D. Fitzwater. 1962. Porcupine-timber relationships in the Lake States. Journal of Forestry 60: 325-330. Longhurst, W.N., H. K. Oh, M.B. Jones, and R. E. Kepner. 1968. A basis for the palatability of deer forage plants. Transactions of the North American Wildlife and Natural Resources Conference 33: 181-189. Mitchell, K. J. 1969. Simulation of the growth of even aged stands of White Spruce. Yale University School of Forestry Bulletin 75. 48 pp. Moss, E. H. 1944. The prairie and associated vegetation of southwestern Alberta. Canadian Journal of Research 22, (es UleSi. Murie, O. J. 1926. The Porcupine in northern Alaska. Journal of Mammalogy 7: 109-113. Reukema, D. L. 1959. Missing annual rings in branches of young-growth Douglas-Fir. Ecology 40: 480-482. Rudolf, P.O. 1949. Porcupines’ preferences in pine planta- tions. Journal of Forestry 47: 207-209. Schulman, E. 1956. Dendroclimatic changes in semiarid America. University of Arizona Press, Tucson. 142 pp. Shapiro, J. 1949. Ecological and life history notes on the Porcupine in the Adirondacks. Journal of Mammalogy 30: 247-257. : Sokal, R.R. and F. J. Rohlf. 1969. Biometry. W. H. Free- man, San Francisco. 776 pp. Spencer, D. A. 1964. Porcupine population fluctuations in past centuries revealed by dendrochronology. Journal of Applied Ecology 1: 127-149. Storm, G. L. and C. H. Halvorson. 1967. Effect of injury by Porcupines on radial growth of Ponderosa Pine. Journal of Forestry 65: 740-743. Taylor, W. P. 1935. Ecology and life history of the Porcu- pine (Erethizon epixanthum) as related to the forests of Arizona and southwestern United States. University of Arizona Bulletin 6: 1-177. van Deusen, J.L. and C.A. Myers. 1962. Porcupine damage in immature stands of Ponderosa Pine in the Black Hills. Journal of Forestry 60: 811-813. Received 3 February 1979 Accepted 29 May 1979 Interactions between Snowy and Short-eared Owls in Winter M. Ross LEIN and PETER C. BOXALL Department of Biology, University of Calgary, Calgary, Alberta T2N IN4 Lein, M. Ross and Peter C. Boxall. 1979. Interactions between Snowy and Short-eared Owls in winter. Canadian Field- Naturalist 93(4): 411-414. Observations on habitat use, food habits, and behavioral interactions of Snowy Owls (Nyctea scandiaca) and Short-eared Owls ( Asio flammeus) in winter near Calgary, Alberta, suggest that no serious competition occurs between them. They differ in habitat use, obtaining different types of prey as a result. Kleptoparasitism by Snowy Owls upon Short-eared Owls may be the most frequent interspecific interaction. Reports of Short-eared Owls in the diet of Snowy Owls probably reflect Opportunistic feeding, rather than a mechanism to reduce competition for food. Key Words: Snowy Owl, Nyctea scandiaca, Short-eared Owl, Asio flammeus, habitat use, food habits, kleptoparasitism, Alberta, winter. Levin et al. (1977) described a case of probable predation by the Snowy Owl ( Nyctea scandiaca) upon the smaller Short-eared Owl (Asio flammeus), and considered the complex relationship between two possible competitors that may also interact as predator and prey. They suggest that, besides ob- taining food, the larger raptor may benefit by eliminating a potential competitor. We report find- ings that clarify the interactions between these species in winter. Our observations were made near Calgary, Alberta (SI°N, 114° W), where Snowy Owls regularly winter (Bird 1972). Short-eared Owls breed near Calgary, and some overwinter in certain years (Salt and Salt 1976). Hence the two species sometimes come into contact, providing opportunities to witness inter- actions. Methods Our study area 1s a 185-km? block of agricultural land southeast of Calgary. Approximately 70% of the area is cultivated for cereal grains, and about 20% is hayfields and pasture.. Numerous small sloughs contain water in spring but support a dense cover of grasses, sedges, and weeds during the rest of the year. During visits to the area we recorded the habitat in which each owl was located when first sighted. If an owl was on a boundary between two habitats a value of 0.5 owl was assigned to each. We analyzed pellets obtained at known roosting sites and around hunting perches. Most pellets were readily assigned to one or the other species by differences in size and by association with known perches. The identity of a few pellets was question- able; these were omitted from the analysis. Frag- mented remnants of an estimated four to six pellets from a Short-eared Owl roost were also analyzed. Observations of interspecific interactions were made whenever possible from 1973 through 1978. Between November 1977 and March 1978 over 130 h of behavioral observations of Snowy Owls were made. Results The habitat distributions of sightings of the two species (Table |) are not totally comparable because of behavioral differences. Many of the Snowy Owl TABLE |—Habitat distributions of Snowy and Short-eared Owl sightings on the study area in the winter of 1977-1978, plus the habitat distribution of prey-capture attempts by Snowy Owls Number of cases (% total) Habitat Snowy Owl Short-eared Owl Snowy Owl sightings sightings hunting attempts Stubblefield 87.5 (43.8) 30.5 (51.7) 14.5 (51.7) Summerfallow 60.0 (30.0) I (5) 0.5 (1.8) Hayfield 28.5 (14.3) 7.0 (11.9) 7.0 (25.0) Pasture 14.0 (7.0) IS (25) 1.0 (3.6) Ungrazed grassland 6.0 (3.0) ND (loz) 1.5 (5.4) Slough 4.0 (2.0) 17.5 (29.7) 3.5 (12.5) Total 200.0 (100.1) 59.0 (100.0) 28 (100.0) 4} 412 sightings probably were of non-hunting birds, be- cause they roost in conspicuous locations such as on poles or hilltops. Short-eared Owls use communal roosts in winter, in locations offering protection from both weather and easy detection (Clark 1975; personal observation). Therefore, most Short-eared Owls seen were flying and probably were hunting. The distri- bution of prey-capture attempts by Snowy Owls (Table 1) is also not fully comparable with the Short- eared Owl sightings because we cannot assume that all the Short-ears were hunting. Both species were observed most often in stubble- fields, which comprised 37% of the study area. While most hunts by Snowy Owls were also in this habitat (Table 1), many of the Short-eared Owls recorded in stubble actually appeared to be hunting over road ditches or sloughs and were assigned to stubble only because they were crossing that habitat when first sighted. Data on prey-capture attempts by Short- eared Owls are unavailable, but would probably indicate a lesser importance of this habitat than it has for Snowy Owls. The high number of Snowy Owls sighted in summerfallow relates to their roosting habits, des- cribed above. They rarely hunted in this habitat (Table 1). Sloughs, comprising only 3.6% of the study area, are favored hunting locations for Short-eared Owls, which frequently are seen coursing back and forth over them. In contrast, few hunting attempts by Snowy Owls were observed around sloughs (Table 1). Table 2 lists the prey items recovered from pellets. The proportions of Deer Mice (Peromyscus manicu- latus) and Meadow Voles ( Microtus pennsylvanicus) in the diets of the two species differ significantly (chi- square test, P<0.001). Microtus comprised 79% of prey individuals in Short-eared Owl pellets but were only half as frequent as Peromyscus in Snowy Owl pellets. Remains (feet and skull fragments) of Short-eared Owls were recovered from two Snowy Owl pellets found on Snowy Owl territories adjacent to Short- eared Owl roosts. The range of prey size is greater for the Snowy Owl, as expected from its larger body size (Schoener 1969). Although both Snowy and Short-eared Owls were seen repeatedly in the same areas, only a few actual encounters were witnessed. These are summarized as follows: 1) 18 February 1975. At 15:57 (MST) a Snowy Owl (SO) flew from a power pole toward a Short-eared Owl (SEO) hunting 300-400 m away. As it approached, the SEO dove at it, coming within 2-3 m. The SO landed and assumed a crouched defensive posture with the body axis at 15-20° from the horizontal, plumage ruffled, and wings partly spread. The SEO circled, making repeated shallow stoops, for 2-3 min before flying away. THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 2—Prey of Snowy Owls and Short-eared Owls on the study area in the winter of 1977-1978, as determined from examination of pellets Number of individuals (% total) Prey species Snowy Owl! Short-eared Owl? Mammals Peromyscus maniculatus 79 (51.0) 8 (16.7) Microtus pennsylvanicus 39 (25.3) 38 (79.2) Spermophilus richardson? WZ (HT) 0 Mustela frenata* 3) (9) 0 Birds Perdix perdix> 14 (9.0) 0 Asio flammeus A (V3), — Passerine spp. 6 (3.9) Di (AD) Total 155 (100.1) 48 (100.1) ‘Based on analysis of 72 pellets. 2Based on analysis of 22 pellets plus numerous fragments. 3Richardson’s Ground Squirrel. 4Long-tailed Weasel. 5Gray Partridge. 2) 1 December 1977. A SO was perched on a powerline tower ina pasture. At 16:07 a SEO, flying high above, dove at it, coming within 2 m. It circled about 10 m above the SO, then stooped again. There was no observable response. The SEO then flew off and began hunting ina field 800 m away. 3) 6 December 1977. At 13:45, a hunting SEO landed ina stubblefield 200 m from a perched SO. The SO flew and supplanted the SEO, which then circled 8-10 m above it, uttering “keeow” calls for 20-30 s before flying away. The SO was not disturbed by this, but appeared to be watching another SO flying in the distance. 4) 14 December 1977. A SO was perched ona knoll ina stubblefield. At 13:32 a SEO flew towards it and stooped several times within | m of the perched SO, which ducked while watching its attacker. The SEO circled several times, then flew to a fencepost 900 m away. It resumed hunting at 13:41. 5) 21 December 1977. At 15:00 a SO flew to a power pole near a SEO that was hunting in a stubblefield. As the SO landed, the SEO flew toward it and dove twice, coming within 3 m. Three other SEOs immediately joined the first and circled 25-30 m above the SO. After | min the SEOs scattered and resumed hunting. No reactions by the SO were seen. 6) 21 December 1977. A SO was sitting on the ground 100 m from a slough where a SEO was hunting. Between 17:25 and 17:30 the SEO made three unsuccessful pounces at prey, without landing for longer than 10 s. The SO watched the SEO but did not appear aroused. At 17:30 the SEO landed in the grass, whereupon the SO flew and supplanted it. The SEO took flight, circled briefly 10-15 m above the SO, and flew away. The SO then flew back and forth across 179 the slough several times and landed on a fencepost about 150 m away. 7) 19 December 1978. At 15:00 a SO flew from a power pole toward a SEO flying, with a mouse in its feet, about 500 m away. As the SO approached, both owls disappeared behind a low rise. Almost immediately the SEO rose straight up to about 15m with empty feet. It circled the area, stooping toward the ground several times, before flying away. The SO, which presumably was perched on the ground with the mouse, was not re-sighted. Discussion The interspecific differences in habitat use and diet are closely interrelated. Snap-trapping surveys at the beginning of the winter of 1977-1978 indicated that Microtus were most common in sloughs and roadside ditches (Boxall, unpublished data), where Short-eared Owls frequently were observed hunting. The pref- erence of Microtus for dense cover is well-known (e.g., Birney et al. 1976). Peromyscus were more abundant than Microtus in stubblefields and hay- fields, where most Snowy Owl hunting attempts were observed (Table 1). The failure of Snowy Owls to prey more heavily on Microtus may be related to hunting behavior. Snowy Owls are “sit and wait” predators, launching hunting attempts from perches after prey has been detected (Hohn 1973; personal observation). In contrast, Short-eared Owls hunt primarily by flying back and forth low over vegetation (Clark 1975), pouncing on any prey located. A small mammal moving within the dense cover in sloughs or ditches would be screened from a Snowy Owl perched some distance away, but would be visible to a flying Short-eared Owl looking directly down into the vegetation. The data on habitat use and food habits suggest that considerable ecological segregation exists between the two species and that interspecific competition for food is unlikely under the conditions of our study. Our behavioral observations support this conclusion. Snowy Owls initiated four interactions. Two (see examples 3 and 6) were in apparent response to a Short-eared Owl landing after prey-capture attempts that might have been successful. In observation 6 the Snowy Owl did not react to the Short-ear hunting nearby until, after 5 min, the latter landed for longer than a few seconds. We suggest that these actions by Snowy Owls were attempted piracy, triggered by an apparently successful hunt by a Short-eared Owl. Observation 7 appears to be a definite example of kleptoparasitism. Duffy et al. (1976) describe klepto- parasitism by a Snowy Owl ona Marsh Hawk (Circus cyaneus) and Tullock (1968) reported Snowy Owls robbing Hooded Crows (Corvus corone). Snowy Owls supplanted Short-eared Owls in several cases but did not pursue them. These probably LEIN AND BOXALL: OWL INTERACTIONS 413 were not attempts at predation because Snowy Owls will pursue flying prey (Meinertzhagen 1959: Nero 1964). Similarly, interspecific territoriality is an unlikely explanation for such encounters because Snowy Owls ignored Short-eared Owls unless the latter appeared to have captured prey. Snowy Owls are territorial in winter (Keith 1964), but the postures and modified flight patterns seen in intraspecific territorial encounters (personal observation) were absent in these interspecific interactions. The encounters initiated by Short-eared Owls (see examples 2, 4, 5), and the reaction of the Short-eared Owls in observations | and 7, are examples of the mobbing response of many birds to predators (Hinde 1970). Once mobbing by one Short-eared Owl attracted three conspecifics to join in briefly. The “keeow” vocalization noted in observation 3 is an alarm call of the Short-eared Owl (Clark 1975). The displays used by Short-eared Owls in intraspecific territorial encounters (Lockie 1955; Clark 1975) were not seen. Breeding Short-eared Owls direct such displays towards various raptors (Clark 1975), and we have seen a wintering Short-ear display to a Rough- legged Hawk (Buteo lagopus). Thus the behavior of Short-eared Owls towards Snowy Owls is an alarm response rather than an aggressive or territorial one. The posture of the Snowy Owl while being mobbed in observation | resembles that of owls attacked by jaegers (Stercorarius spp.) (Tullock 1968) or by a Goshawk (Accipiter gentilis) (personal observation). The lack of territoriality between Snowy Owls and Short-eared Owls in winter, along with differences in habitat use and diet, suggests little competition between them. Short-eared Owls winter in areas with high populations of small mammals (Clark 1975), suggesting that winter contact with Snowy Owls will occur primarily in situations where high food density alleviates competition (see also Lack 1946). Pitelka et al. (1955) recorded no interactions between these two species at Barrow, Alaska, when they bred together in’ response to a high lemming population. We suggest that the major relationship between the two species is that of kleptoparasite and victim. The mobbing reactions of the Short-eared Owls demon- strate that they recognize Snowy Owls as potential pirates or predators. In most records of Short-eared Owls in the diet of Snowy Owls (e.g., Uttendorfer 1952: Mikkola 1976) it is unknown whether predation or scavenging had occurred. The Snowy Owl has remarkably catholic food habits (Gross 1944) and actual predation may be of irregular or exceptional occurrence. The rarity of documented predation and the weakness of the Short-eared Owl’s mobbing reaction suggest that Snowy Owl predation upon Short-eared Owls is not significant in the biology of either species. 414 THE CANADIAN FIELD-NATURALIST Acknowledgments We thank R.M. Fisher, D. P. Nugent, C. C. Mamo, and M. Rothfels for assistance in the field, and M. T. Myres and J. B. Gollop for their comments on the manuscript. R. M. Fisher made his notes on observation 6 available to us. This work was supported by the National Research Council of Canada and the University of Calgary. Literature Cited Bird, C. D. 1972. The Snowy Owl in the Calgary area, 1964-1972. Calgary Field Naturalist 3: 106-107. Birney, E. C., W. E. Grant, and D. D. Baird. 1976. Impor- tance of vegetative cover to cycles of Microtus popula- tions. Ecology 57: 1043-1051. Clark, R. J. 1975. A field study of the Short-eared Owl, Asio flammeus (Pontoppidan), in North America. Wild- life Monographs, Number 47. Duffy, D. C., B. Beehler, and W. Hass. 1976. Snowy Owl steals prey from Marsh Hawk. Auk 93: 839-840. Gross, A. O. 1944. Food of the Snowy Owl. Auk 61: 1-18. Hinde, R.A. 1970. Animal behaviour. 2nd_ Edition. McGraw-Hill Book Company, New York. Hohn, E. O. 1973. Winter hunting of Snowy Owls in farm- land. Canadian Field-Naturalist 87: 468-469. Keith, L. B. 1964. Territoriality among wintering Snowy Owls. Canadian Field-Naturalist 78: 17-24. Vol. 93 Lack, D. 1946. Competition for food in birds of prey. Journal of Animal Ecology 15: 123-129. Levin, S. A., J. E. Levin, and R. T. Paine. 1977. Snowy Owl predation on Short-eared Owls. Condor 79: 395. Lockie, J. D. 1955. The breeding habits and food of Short- eared Owls after a vole plague. Bird Study 2: 53-69. Meinertzhagen, R. 1959. Pirates and predators. Oliver and Boyd, Edinburgh. Mikkola, H. 1976. Owls killing and killed by other owls and raptors in Europe. British Birds 69: 144-154. Nero, R. W. 1964. Snowy Owl captures duck. Blue Jay 22: 54_55. Pitelka, F. A., P.Q. Tomich, and G. W. Treichel. 1955. Ecological relations of jaegers and owls as lemming predators near Barrow, Alaska. Ecological Monographs 25: 85-117. Salt, W.R. and J. R. Salt. 1976. The birds of Alberta. Hurtig Publishers, Edmonton, Alberta. Schoener, T. W. 1969. Models of optimal size for solitary predators. American Naturalist 103: 277-313. Tullock, R. J. 1968. Snowy Owls breeding in Shetland in 1967. British Birds 61: 119-132. Uttendorfer, O. 1952. Neue Ergebnisse uber die Ernahrung der Greifvogel und Eulen. Eugen Ulmer, Stuttgart. Received 26 February 1979 Accepted 9 June 1979 — The Biological Flora of Canada 1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry* IVAN V. HALL,’ LEwWIs E. AALDERS,! NANCY L. NICKERSON,! and SAM P. VANDER KLOET? 'Research Station, Agriculture Canada, Kentville, Nova Scotia B4N 1J5 *Department of Biology, Acadia University, Wolfville, Nova Scotia BOP 1X0 Hall, Ivan V., Lewis E. Aalders, Nancy L. Nickerson, and Sam P. Vander Kloet. 1979. The biological flora of Canada. 1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry. Canadian Field-Naturalist 93(4): 415-430. This contribution on Vaccinium angustifolium Ait., Sweet Lowbush Blueberry (Ericaceae), is the first of a series presenting biological information on plants that are components of the flora of Canada. Vaccinium angustifolium is a deciduous low shrub endemic to North America, whose range in Canada extends from the east coast of Newfoundland to Lake Winnipeg in Manitoba. It occurs from Quebec 57°N to Virginia 38°N. To produce fruit the flowers usually require cross-pollination by wild bees or related insects. The fruit of V. angustifolium is edible and matures during late summer. During the past 30 yr inthe Atlantic Provinces of Canada many fields abandoned from other forms of agriculture have been brought into stands of this species by burn-pruning and selective weed control. Key Words: Vaccinium angustifolium, Sweet Lowbush Blueberry, biology, ecology, physiology, distribution, economic importance. 1. Name Vaccinium angustifolium Ait.; section Cyanococcus; Ericaceae; V. lamarckii Camp (Camp 1943, 1945); V. pennsylvanicum Lam. (Robinson and Fernald 1908); V. pensilvanicum Lam. (Fernald 1950); Sweet Lowbush Blueberry; airelle a feuilles étroites (Marie-Victorin 1964). 2. Description of the Mature Plant (a) Raunkiaer life-form. Chamaephyte. Winter-deciduous, broad-leaved low shrub with ascending branches, edible blue fruits, deep tap root; reproduces by seeds and rhizomes. (6) Shoot morphology. Stems woody, average height 20 cm with maximum of 50 cm, generally glabrous, with raised lenticels, the bark variously pigmented from yellow to deep red in autumn; buds of two types, the larger flower buds borne terminally, the more lanceolate vegetative buds born proximally; leaves alternate ina spiral, simple, pinnately netted, serrate glandular, elliptic or ovate-oblong, apex acute, base obtuse, the ventral surface waxy green, the dorsal surface pale green and sometimes with light bloom; rhizomes woody bearing numerous shoots, new growth white or pinkish (Figure la, b). (c) Root morphology. The radicle of the seedling develops into an extensive tap root system (Hall 1957): the root system is finely divided at the extremities and several authors state that there are no root hairs (e.g., Addoms and Mounce 1931). (d) Inflorescence. The members of Vaccinium section Cyanococcus are characterized by flowers borne in racemes; members of section Vaccinium by contrast have flowers borne singly in the leaf axils. Flowers of V. angustifolium are typically pentacyclic with five sepals, five petals fused into a bell-shaped corolla, 10 stamens in two whorls of five and fused to the corolla, and a single pistil with inferior ovary. Fruit is a true berry bearing 10 pseudolocules each with a few to many small seeds. Bell (1957) found that the average number of perfect seeds was 13 and of imperfect seeds was 50 per berry. Some mutants were reported by Hall, Aalders, and Lockhart (1964). Both Vaccinium and Gaylussacia have inferior ovaries but the seeds of the former are much smaller. (e) Subspecies. None. *See La Roi (1977) for notice, guidelines and schedule for contributors. 415 416 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE |. Sweet Lowbush Blueberry Plant. A: Above, close-up of flowers: below, flowering plant. B: Above, close-up of fruit: below, fruiting plant. (f) Varieties and forms. Fernald (1950) recognized a puberulent var. hypolasium and a smooth-leaved var. laevifolium House, but we do not, considering these to be part of the variation found in the species. We recognize the following forms: V. angustifolium Ait. forma angustifolium with green leaves, fruit with a heavy bloom, consequently a blue color, and straight stems, is the most frequent taxon (Vander Kloet 1978); V. angustifolium forma nigrum (Wood) Boivin is characterized by blue-green leaves, fruit with little or no bloom giving it a black color, and zigzag stems (Aalders and Hall 1963a); V. angustifolium forma leucocarpum (Deane) Rehder is a white-fruited form. (g) Ecotypes. Forma nigrum tends to increase more rapidly than forma angustifolium in stands that are burned regularly (Hall et al. 1975). (h) Chromosome numbers. Longley (1927) first reported 24 bivalent chromosomes for V. angustifolium. Newcomer (1941) sub V. pennsylvanicum also reported that five selections had n = 24 chromosomes. Darrow et al. (1942) are the only workers to report a diploid condition (2m = 24) but no voucher specimen for this count can be located (Vander Kloet 1978). Counts from clones of Canadian and Maine V. angustifolium all have 2n = 48 (Hall and Aalders 1961; Bent and Vander Kloet 1976; Hersey and Vander Kloet 1976; Whitton 1964). 3. Distribution and Abundance (a) Geographic range. Vaccinium angustifolium is a North American endemic, extending from Cape St. Francis on the eastern tip of Newfoundland to the Pine Falls area of Lake Winnipeg in south central Manitoba (Figure 2). It extends from 57°N in northern Quebec southward to isolated uplands in the Appalachian Mountains of Virginia, 38°N. A complete distribution map appears in Vander Kloet (1978). 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 417 °/ AS 800 Kilometres Vaccinium angustifolium Ait. FiGuRE 2. Canadian distribution of Vaccinium angustifolium from specimens in the Department of Agriculture Herbarium, Ottawa, Ontario (DAO): National Herbarium, National Museums of Canada, Ottawa, Ontario (CAN): and Acadia University Herbarium, Wolfville, Nova Scotia (ACAD). (b) Altitudinal range. Vaccinium angustifolium extends from ca. sea level to 1300 m asl in Eastern Canada to 1300-1500 m asl in Virginia. On the higher points of the Adirondack Mountains V. angustifolium is replaced by V. boreale (Sweet Hurts) (Vander Kloet 1977). 4. Physical Habitat (a) Climatic relations. Vaccinium angustifolium occurs ina large area of eastern North America and tolerates a wide range of climatic conditions. The variability of this species with regard to productivity is well established (Aalders and Hall 1963b: Hall, Aalders, and Wood 1966) but the adaptiveness of the species to different climatic conditions requires further study. A comparison of growth under different climatic conditions was conducted on uniform plant and soil media in 1965 and 1966 at Kentville, Nova Scotia; Normandin, Quebec; and St. John’s West, Newfoundland. Linear shoot growth of 20 plants from seven clones at the three respective stations averaged 183, 141, and 111 cm per plant in 1965; and 195, 153, and 70 cm in 1966. The number of flower buds produced per plant averaged 126, 94, and 66 in 1965; and 88, 54, and 23 in 1966. Thus climatic conditions for growth varied widely among the three stations. Vegetative growth was not notably different between 1965 and 1966 at Kentville and Normandin, but much poorer in 1966 at St. John’s West; reproductive growth was much better in 1965 than 1966 at all three stations. In Eastern Canada, winter temperatures for V. angustifolium and other chamaephytes are generally ameliorated by a snow cover. Inadequate snow cover may be followed by shoot dieback to ground level in the Lac de St. Jean area (Poirier and Dubé 1969). Unusually warm weather and salt spray from a February hurricane, followed by cold temperatures, were probably responsible for extensive mortality of FV. angustifolium flower buds (Figure 3).in the Fox River area of Nova Scotia in 1976. Cold winds off the Northumberland Strait are a factor in delaying the beginning of plant growth in Inverness County, Nova Scotia (Hall et al. 1963). 418 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE 3. Winter injury on shoots of Vaccinium angustifolium. In New Brunswick, Hall (1955) found that 0.5% of full sunlight under a Picea rubens — Abies balsamea canopy (Red Spruce —- Balsam Fir) gave a minimum growth or just survival, 10% sunlight under a Berula populifolia canopy (Oldfield Birch) gave moderate vegetative growth but no flowering, and 50% sunlight (Openings in canopy) gave flowering and fruiting. Seed germination occurs only after periods of prolonged rainfall in late summer or early fall. Vander Kloet (1976a) found that seedling density was higher in a field sampled in Pictou County, Nova Scotia than in an old field at Leeds County, Ontario and he correlated this with higher precipitation and soil moisture. Relative humidity may play an important role in fruit production. In dry years areas along the Atlantic coast have produced exceptional crops. Plants in the coastal fog belt are frequently covered with water droplets while plants farther inland experience stress from lack of moisture (e.g., withered berries, browned leaf margins). (b) Physiographic relations. One of the most important factors limiting flowering and fruit development is the occurrence of late spring frosts on low-lying areas (Jackson et al. 1972b). Laboratory tests have shown that 6 h of —2.2°C were detrimental and a further reduction to —3.3°C for 6 h markedly reduced fruit set (Hall, Aalders, and Newbery 1971). Vaccinium angustifolium grows, yields, and reproduces well by rhizomes on both mineral and organic soils provided adequate moisture and aeration are available. It tolerates a wide range of soil conditions with best growth at low pH levels (Hall, Aalders, and Townsend 1964). In soils with much humus, most rhizomes grow in the top 5 cm. Jackson et al. (1972a) reported that soils such as Westbrook loam and Southampton sandy loam witha high percentage of stone or gravel provide the conditions most favorable to the emergence, growth, and development of seedlings. 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 419 (c) Nutrient and water relations. At the time of flower bud initiation the following suggested levels of nutrients (as % dry weight) should exist in the leaves (Townsend and Hall 1970): N, 1.50-2.00: P, 0.08-0.12: K, 0.40-0.55: Ca, 0.40-0.65; and Mg, 0.15-0.20. Lockhart (1959) first described the symptoms of mineral deficiency in V. angustifolium. Rayment (1965) clearly demonstrated the beneficial effects of applications of nitrogen to stands of V. angustifolium in Newfoundland. Townsend et al. (1968) reported that sugars were lower, and starch was notably higher, in the rhizomes than in the leaves. Barker et al. (1963) found that the reducing sugars in fruits of five different clones varied between 7.4 and 7.9% of fresh weight. ; Once established, the rhizone system of this species plays an important role in preventing slope erosion. If soil particles are washed into the network of rhizomes and shoots, new roots and shoots develop in the additional soil, favoring the plant and retaining the soil. The taproot system (Hall 1957) may penetrate to >1 msoil depth, allowing use of subsoil moisture reserves. Laycock (1967) states that the concentration of roots and rhizomes near the surface in the pine barrens of New Jersey allowed V. angustifolium and other species to absorb much of the water from light rains that fell during the growing season. The nutrient and pH requirements and competitive ability of this polymorphic species invite further investigation. 5. Plant Communities. Within the largely undisturbed part of its range, V. angustifolium is an important member of exposed headland vegetation, raised bogs, high moors, outcroppings on the Canadian Shield, mountain summits in the Gaspé, as well as in the herb—dwarf shrub stratum of open to moderately shaded pine to spruce woodlots and forests on coarse textured, mainly infertile soils in the Acadian, Great Lakes — St. Lawrence and Boreal Forest Regions of Rowe (1972), from Lake Winnipeg to the Atlantic coast. But the species is most abundant in disturbance communities which result from clear-cutting, forest fires, and abandonment of agricultural land. The latter is.especially noteworthy in the Maritime Provinces where, witha decline in soil fertility, land was abandoned during the early part of this century, providing many old field habitats for this species to colonize. Conners (1972) reports that by the late 1940s several of these fields were being cultivated. With the introduction of frozen foods on the American market a rapid development of the stands and industry took place. Table | shows the contrasts among the floristic composition of three community types: (1) blueberry barrens on mainland Nova Scotia, (2) granite outcroppings near Kaladar, Ontario and, (3) the boreal conifer-hardwood forests of the Great Lakes region. Aside from V. angustifolium, the three community types have floristically littie in common. Out of 71 species, excluding lichens and those with less than five occurrences, only V. myrtilloides (Sour-top Blueberry), Gaultheria procumbens (Teaberry), Fragaria virginiana (Wild Strawberry), Rumex acetosella (Sheep Sorel), and Maianthemum canadense (Wild Lily-of-the-Valley) are shared. Maycock and Curtis (1960) have shown that V. angustifolium is an important component in both dry and wet sites of the boreal coniferous forest, but is unimportant on mesic sites. In Wisconsin, Curtis (1959) showed that it is the most prevalent groundlayer species in the northern dry forest and also an important constituent of northern dry-mesic forest, northern wet forest, bracken-grassland, and open bog. Lamoureux and Grandtner (1977) showed that this species was important in dune formation on Iles-de-la-Madeleine. Lavoie (1968) has described the Jack Pine ( Pinus banksiana) forests witha Kalmia angustifolia-Vaccinium understory for the Lac de St. Jean area of Quebec. Community descriptions and tabulations of V. angustifolium in the Maritime. Provinces are given by Hall (1955, 1959, 1975), Hall and Aalders (1968), and Hall et al. (1973, 1974, 1976). 6. Growth and Development (a) Morphology. In the lowbush blueberry seedling (Figure 4) the oblong-elliptical cotyledons are about 2 mm long. The first seedling leaves are much smaller and usually more elliptical than mature leaves, making identification of young seedlings difficult. Stages in seedling development are given by Eaton and Hall (1961). Plants rarely flower or produce rhizomes until 4 yr after germination (Hall 1953). After plants reach a total width of 30 cm and rhizome growth occurs in several directions, expansion of the clone is more rapid. Rhizomes may grow up to 10 cm/yr on mineral soils and up to 50 cm on organic soils. Clones with intact rhizomes 10 m long have been excavated in Kings County, Nova Scotia. The minimum age of a parent clone may be estimated by counting growth rings from rhizome cross-sections (Figure 5). The importance of rhizome growth in the expansion of the clone has been outlined by Barker and Collins (1963b). (b) Physiology. Forsyth and Hall(1965) have shown that photosynthesis is more rapid in the early morning and that temperature, CO, and leaf age affect photosynthetic rates. Rates of apparent photosynthesis in shoots at different light intensities have been reported by Bonn et al. (1969). Red leaf infected leaves (see 9(c)) and genetic mutant leaves had abnormally lower photosynthetic rates (Hall, Forsyth, Lockhart, and Aalders 1966). Strong 420 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1— Vaccinium angustifolium Aiton, associated species on six granitic outcroppings from Kaladar, Lennox-Addington County to Mount Fitzsimmons, Leeds County, eastern Ontario: two barrens* on mainland Nova Scotia: and I! dry Boreal Conifer — Hardwood Forests of the Great Lakes Region** Frequency of occurrence (%) Eastern Mainland Great Lakes Species Ontario Nova Scotia region Bare ground, rock, or litter 789 (21.6) 1716 (17.2) NA Epipetric lichens 522 (14.3) — NA Fruticose lichens PON GSES) —- NA Polytrichum commune 28 ( 0.8) 199 ( 2.0) NA Polytrichum jJuniperinum ISU ( 41) — NA Polytrichum piliferum 33 ( 0.9) = NA Lycopodium annotinum*** — = (12) Lycopodium clavatum _ 6 (21) Lycopodium obscurum a 44 ( 0.4) (22) Lycopodium complanatum — 6 ( 0.06) ( 1) Pteridium aquilinum _ 59 ( 0.6) (78) Juniperus communis @il ( 25) — — Danthonia spicata 159 ( 4.4) 536 ( 5.4) = Deschampsia flexuosa 144 ( 3.9) = == Poa compressa 30 ( 0.8) —- — Agrostis scabra 6 ( 0.2) — — Panicum subvillosum = 14 ( 0.1) Oryzopsis asperifolia — — ( 8) Carex pensylvanica 228 ( 6.2) — Carex nigromarginata 23 ( 0.6) — = Carex umbellata 8 ( 0.2) = = Luzula multiflora = 6 ( 0.06) a Clintonia borealis — — (17) Maianthemum canadense 15 ( 0.4) 8 ( 0.08) (66) Salix bebbiana — 335) ((_ )315))) = Betula populifolia _— 30 ( 0.3) — Corylus cornuta — = (23) Rumex acetosella AV) (( 1.3) 4( 0.04) (ey) Comandra umbellata 66 ( 1.8) i Anemone quinquefolia = a Dalibarda repens — — Spiraea alba 18 ( 0.5) — Spiraea latifolia _ 60 ( 0.6) — Spiraea tomentosa — 29 ( 0.3) Amelanchier sanguinea ihe @.33)) — — Amelanchier spicata 19 ( 0.5) — — Prunus virginiana 9 ( 0.3) — —- Prunus serotina 7 (( O22) — — Pyrus melanocarpa 163 ( 4.5) -- a Rubus arundelanus 12 ( 0.3) — = Rubus hispidus = 41 ( 0.4) — Rubus idaeus -- — ( 6) Waldsteinia fragarioides — — (18) Fragaria virginiana 5) (( W310) 39 ( 0.4) (#5) Potentilla tridentata — 46 ( 0.5) = Potentilla simplex = 84 ( 0.8) — Polygala paucifolia _ — (22) Rhus copallina 9 ( 0.2) — — Rhus typhina 11 ( 0.3) = — Aralia nudicaulis — — (33) Cornus canadensis — 389 ( 3.9) (61) Ledum groenlandicum _- 14 ( 0.1) — Rhododendron canadense — 106 ( 1.1) — 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 421 TABLE |—(Concluded) Frequency of occurrence (%) Eastern Mainland Great Lakes Species Ontario Nova Scotia region Kalmia angustifolia — STM (B21) — Arctostaphylos uva-ursi Ol @ ey) — — Gaultheria procumbens 23 ( 0.6) 348 ( 3.5) (21) Vaccinium myrtilloides 32 ( 0.9) 1 ( 0.01) (20) Vaccinium angustifolium 528 (14.5) 2139 (21.4) (51) Vaccinium vitis-idaea — Po (O27) — Gaylussacia baccata sy) (IS) = = Trientalis borealis — — (39) Melampyrum lineare — — (19) Linnaea borealis = — (12) Viburnum rafinesquianum 21 ( 0.6) — = Lonicera canadensis — 6 ( 0.06) = Diervilla lonicera 8:((022) = == Helianthus divaricatus 26 ( 0.7) = = Aster cordifolius 8 ( 0.2) — — Aster macrophyllus — = (33) Solidago puberula - 648 ( 6.5) Solidago rugosa = 6 ( 0.06) = Solidago graminifolia — 15 ( 0.15) ( 3) Antennaria neglecta = 120812) (a) Prenanthes trifoliolata = 5 ( 0.05) — Hieracium aurantiacum = 8 ( 0.08) (12) Hieracium pratense — 5 ( 0.05) — 34 spp. with fewer than 5 occurrences 46 ( 1.3) 10 ( 0.1) NA Totals 3650 (100) 10,000 (100) 11 Stands *Data from Hall and Aalders (1968) slightly modified and reduced. **Data from Maycock and Curtis (1960) slightly modified and reduced. ***Scientific nomenclature of vascular plants follows Gray’s Manual of Botany (Fernald 1950). vegetative growth results when plants of different provenances are given 16 h light and 8 h dark periods per 24 h at 18°C. Flower buds are formed when plants are given 12 h light and 12 h dark per 24 h at 18°C. Hall et al. (1970) have shown that leaf anthocyanin concentration increases as the temperature decreases. Vaccinium angustifolium leaves develop brilliant red or yellow coloration in autumn. (c) Phenology. Vaccinium angustifolium overwinters in a leafless state, with twigs yellow to reddish brown. At Kentville, vegetative and flower buds swell in early May if air temperatures have exceeded 10°C for 3-4 d. Flowering occurs from late May to mid-June. In cooler coastal areas, flowering may be delayed 2-3 wk (Bell 1953). Leaf development precedes, is concomitant with, or follows flowering depending on the particular clone. Leaves harden by mid-July, turn red in late August, and absciss by late October. In seasons following a burn- prune, vegetative growth commences a week after flower bud expansion and terminates in early July. Termination of shoot growth as evidenced by a black tip in the apical meristem (Bell 1950) occurs earlier on older unpruned bushes (Barker and Collins 1963a). Berries ripen in early August at Kentville, 2-3 wk earlier than in Cumberland County, Nova Scotia (Aalders et al. 1972). Flower primordia begin to develop shortly after cessation of vegetative growth (Bell and Burchill 1955a), but continue until late October if air temperatures remain >0°C with extended periods greater than 10°C. Thus temperature is an important regulator of plant phenology. But other factors including day length are important (Hall et al. 1970). At Lac de St. Jean, Quebec, flower bud formation is poor in some years, probably owing to early frosts. Halland Ludwig (1961) have shown that different clones react differently to day length and temperature. By winter, primordia ofall the floral organs are microscopically recognizable (Bell and Burchill 1955b). Kender (1968) found that growth potential of rhizome buds was greater in early spring and late summer than in July. 422 THE CANADIAN FIELD-NATURALIST Vol. 93 FIGURE 4. Sweet Lowbush Blueberry seedlings. Approximate age 3 yr upper left and 4 yr lower right. The latter has begun to form a rhizome (indicated by arrow). 7. Reproduction (a) Floral biology. The flowers are insect pollinated, but not very attractive to bees, probably because the nectar volatiles are simple acetaldehyde and ethanol (Hall, Forsyth, Lightfoot, and Boch 1971). The stamens and pistil are functional as soon as the flower fully opens. The pistil remains highly receptive under field conditions for4 d but fertility drops to about 20% by the 7th d (Wood 1962). Wood (1961a) also found that nectar volume and weight of nectar sugar increased with the flower age. After pollination the corolla turns pink and deteriorates very rapidly. This is associated with a marked increase in ethylene production (Hall and Forsyth 1967) anda high rate of respiration (Forsyth and Hall 1969) which persists through the small green fruit stage. Glasshouse studies indicate that flowers selfed using pollen from the same flower or any flower of the same clone do not generally set fruit (Aalders and Hall 1961). The results of intercrossing and selfing six clones of V. angustifolium are given in Table 2. The vascular anatomy of the ovary has been described by Bell and Giffin (1957). (b) Seed production and dispersal. It is rare to find a seedless fruit of V. angustifolium even in highly productive clones. Table 2 shows the number of seeds per berry in crosses completed in 1977 and 1978 among six clones. 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 423 Seeds are spread in the droppings of birds and mammals. The American Robin (Turdus migratorius) isa major seed disperser in southwestern New Brunswick as it often feeds and migrates just before fruit harvest by man (Eaton 1957). We have found seeds of V. angustifolium in the droppings of Black Bear ( Ursus americanus), Red Fox (Vulpes vulpes), and Raccoon (Procyon lotor). TABLE 2—Cross- and self-fertility of six selected clones of Vaccinium angustifolium in glasshouse trials, Kentville, Nova Scotia, 1977-1978 Percent of pollinated No. seeds Percent flowers forming No. seeds per 100 seed Female Male fruits! per berry? pollinations germinations3 Augusta ™X Brunswick 100 31 3100 88 x Chignecto 70 24 1680 97 x 510 65 25 1625 98 xX ME3 81 55 4485 93 x ME4161 61 49 3026 88 x Self 56 10 568 89 Brunswick X Augusta 0 — 0 = x Chignecto 0 = 0 -- x 510 2 16 30) 94 x ME3 2 20 49 90 x ME4161 0 os 0 — x Self 0 — 0 = Chignecto X Augusta 74 30 2249 88 x Brunswick 65 17 1131 91 x 510 5) 24 1298 90 xX ME3 56 28 1548 98 x ME4161 45 25 1125 95 xX Self 0 — 0 = 510 x Augusta 12 46 580 84 x Brunswick 95 34 3230 80 x Chignecto 80 33 2632 86 x ME3 1] 4] 461 82 x ME4161 0 oo 0 = x Self 0 — 0 = ME3 x Augusta 18 37 644 89 x Brunswick ~ 65 39 2535 94 x Chignecto 14 38 521 89 x 510 6 22 140 95 x ME4161 l 14 18 93 xX Self 0 = ) — ME4161 X Augusta 42 21 892 97 x Brunswick 29 23 673 93 x Chignecto 26 16 410 90 x 510 39 23 907 91 99% of germinating seeds died even under partially controlled conditions. 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 425 The shoot crown diameters of plants established by seed can be converted to age classes according to a formula devised by Eaton and Hall (1961). Using this size-age relationship, Vander Kloet (1976b) found a marked difference in the age structure of V. angustifolium in old fields in eastern Ontario and eastern Nova Scotia. In the former, all seed-established plants exceeded 40 yr of age: in the latter the 11-20 yr age class was the most frequent. On stable habitats such as mountain summits, exposed headlands, or outcroppings in the Canadian Shield, colonies of V. angustifolium may attain considerable age. Clones attain large size (>10 m diameter) and age (150 yr). Vander Kloet (unpublished data) studied the species composition and population patterns of granitic outcrop communities in the Thousand Island region of Ontario and New York over a 10-yr period. Little or no change in species composition or abundance was observed; however, the positions of colonies of V. angustifolium, Juniperus communis (Common Juniper), and Pyrus melanocarpa (Black Chokeberry) shifted laterally. Vigorous peripheral branches of Juniperus shaded out colonies of Vaccinium and Pyrus, while the latter two invaded the senescing centers of Juniperus colonies. Similarly Pyrus and Vaccinium invaded the senescent portions of each other’s colonies. Turnover rates in V. angustifolium populations may be more rapid in the maritime provinces where the species is often a common component of old field succession, reaching peak abundance 15-25 yr after abandonment. By the judicious use of fire, this seral stage can be maintained as blueberry barrens for many years. Ordinarily the low growing V. angustifolium is soon shaded out by a variety of hardwood or conifer tree species. Once the forest canopy fully develops, it becomes uncommon and/or sterile. Vaccinium myrtilloides has a much higher survival potential than V. angustifolium in the mature Acadian boreal forest (Hall 1959). 9. Interaction with Other Species (a) Competition. Competing plant species may be treated in four ecological site groups: (1) undisturbed natural; (2) post-logging; (3) old field; (4) blueberry crop. On undisturbed natural sites within its range of suitable habitats, V. angustifolium has many competitors, as may be seen in Table 1. In relatively stable and oligotrophic Pinus banksiana woodlands and Picea mariana (Black Spruce) forests of the Boreal Forest and Great Lakes — St. Lawrence Regions (Rowe 1972) east of Lake Winnipeg, its major competitors for nutrients, water, and light in the herb-dwarf shrub stratum include I. myrtilloides, Cornus canadensis (Bunch Berry), Gaultheria hispidula (Creeping Snowberry), Coptis groenlandica (Goldthread), Pteridium aquilinum (Bracken Fern), Clintonia borealis (Corn-lily), Comptonia peregrina (Sweet-fern), Maianthemum canadense, and Lycopodium spp. Major competitors in the low shrub stratum include Kalmia angustifolia (Lambkill), Ledum groenlandicum (Labrador-tea), and Diervilla lonicera (Bush Honeysuckle). On post-logging sites in eastern Canada, surviving and/or invading V. angustifolium populations must compete with surviving species that respond favorably to clearing, e.g., Pteridium aquilinum (Cody and Crompton 1975), Dennstaedtia punctilobula (Hay-scented Fern) (Cody et al. 1977), and Cornus canadensis (Hall and Sibley 1976), as well as with aggressive invader species, e.g., Epilobium angustifolium (Fireweed). On old field sites in the Acadian Forest Region (Rowe 1972) V. angustifolium is an important seral species in the transition stage between field and forest. On coarse-textured soils in old fields of Prince Edward Island, Myrica pensylvanica (Bayberry) succeeds V. angustifolium before the Picea glauca excludes both (Hall 1975). On light sandy soils of the Lac de St. Jean of Quebec, Comptonia peregrina is a major invader of blueberry stands (Lavoie 1968). Members of the blueberry crop group offer competition because they thrive under the cultural practices pertaining to lowbush blueberry culture. Probably the strongest competitor for space is Kalmia angustifolia (Hall et al. 1973). Its stem growth following burn-pruning exceeds that of V. angustifolium (Hall and Aalders 1968). Fertilizing stands of lowbush blueberry increases the growth of Pyrus melanocarpa relative to that of FV. angustifolium (Hall et al. 1978). On poorly drained areas Rhododendron canadense (Rhodora) and Spiraea latifolia (Meadow-Sweet) (Hall et al. 1974) replace V. angustifolium. (b) Symbiosis. The principal native pollinators in Maine and eastern Canada are species of Halictidae and Andrenidae although a few species of Bombidae, Anthophoridae, Colletidae, and Xylocopidae are of some importance (Boulanger et al. 1967). In eastern Ontario Vander Kloet (1976c) found that the solitary bees Andrena vicina and A. carlini, and the bumblebees Bombus bimaculatus, B. terricola, and B. ternarius were the important oe Wood (1961b) found honeybees effective during a short period of bloom in New Brunswick. Mycorrhizal associations in V. angustifolium were described by M. MacArthur (1955. Mycorrhiza in the 426 THE CANADIAN FIELD-NATURALIST Vol. 93 blueberry. /n Horticulture Division, Central Experimental Farm, Ottawa, Progress Report 1949-53. pp. 71-72) but the fungi involved were not identified. (c) Predation and parasitism. Foliage of V. angustifolium is eaten by Black Bear ( Ursus americanus), Eastern Cottontail (Sy/vilagus floridanus), and White-tailed Deer (Odocoileus virginianus). Fruits are eaten by a number of mammals and many birds (Martin et al. 1951). The important insects are Blueberry Maggot, Rhagoletis mandax Cn.; Black Army Cutworm, Actebia fennica (Yausch.); Chainspotted Geometer, Cingilia catenaria (Drury); Blueberry Flea Beetle, Altica sylvia Mall.: Blueberry Casebeetle, Chamisus cribripennis (LeConte) (see Wood 1978): Blueberry thrips, Frankliniella vaccinni Morgan and Catinathrips kainos O'Neill: Blueberry Tipworm, Contarinia vaccinii Felt.: sawflies, Neopareophora litura Klug, Pristiphora idiota Nort., Pristiophora sp.; Red-striped Fireworm, Aroga trialbamaculella Chamb.; and Stem Galler, Hemadas nubilipennis Ashm. The stage of the life cycle of these insects which affects V. angustifolium, the manner of infection, the symptoms for recognition, and other details are given in Hall et al. (1975). The important fungal diseases (Conners 1967) are monilinia blight caused by Monilinia vaccinii-corymbosi (Reade) Honey: botrytis blight (Borrvtis cinerea Pers.): red leaf (Exobasidium vaccinii Wor.); witches-broom (Pucciniastrum goeppertianum (Kuhn) Kleb.); dieback (Diaporthe vaccinii Shear); powdery mildew (Microsphaera penicillata (Wallr. ex Fr.) Lev. var. vaccinii (Schw.) W. B. Cke.): leaf rust (Pucciniastrum vaccinii (Wint.) Jorstad); leaf spot (Septoria sp.); Gloecsporium leaf spot (G/oeosporium minus Shear); and canker (Godronia cassandrae Pk. f. vaccinii Groves). A V. angustifolium plant which showed symptoms of shoestring virus disease was reported by Lockhart and Hall (1962). (d) Toxicity and allelopathy. None reported to date. 10. Evolution and Migration. Camp (1942), on the basis of meager and unsubstantiated evidence (Vander Kloet 1978), assumed that V. angustifolium was a diploid species (1.e., gametes had 12 chromosomes) but as was pointed out in section I(/) this cannot be confirmed. Therefore his allopatric speciation model for the species is also doubtful since Camp (1942) argued, rightly, that processes leading up to speciation in diploid populations differ markedly from those which give rise to tetraploid species. Vander Kloet (1977) has postulated a recent hybrid origin for V. angustifolium. He proposes that V. boreale migrated south or moved down mountain slopes where it came into contact with V. pallidum Ait. (Upland Low Blueberry) of oak-pine woods. Both of these species are diploids and by spontaneous chromosome doubling in a hybrid, an allotetraploid species is plausible. Probably V. angustifolium migrated into the northern part of its habitat from the southern United States following the retreat of the last glacial ice. Birds doubtless accelerated this migration by dispersing seeds across water bodies and other habitat barriers. An initial attempt has been made to separate clones of V. angustifolium on the basis of chlorophyll and anthocyanin content of bark from shoots (Wood and Barker 1963). 11. Response Behavior (a) Fire. In natural communities or managed forests V. angustifolium survives wild fire or controlled burning below ground. Recolonization occurs by rhizome sprouting. Commercial stands are burn-pruned every second year resulting generally in unbranched stems which have more flower buds per stem and more flowers per bud than on older wood. There were no significant differences between fall- and spring-burned plants with respect to amount of shoot growthand number of flower buds per shoot (Hall, unpublished). Burning after the plants were in full leaf was detrimental to new shoot growth and flower bud formation (Eatonand White 1960). Black (1963) found that total fruit production over a 9-yr period was greater from burning every second year than from burning every third year. Smith and Hilton (1971) found that improved lowbush blueberry performance in Ontario after burning resulted mainly from the stimulative effects of nutrients in ash deposited on the surface soil. (b) Grazing and harvesting. In New Brunswick, sheep were observed to graze grasses and sedges selectively rather than feed on V. angustifolium and V. myrtilloides (Hall, I. V. 1954. Ecological studies. Ja Dominion blueberry substation, Tower Hill, New Brunswick progress report 1949-1953. Canada Department of Agriculture. pp. 18-23). Removal of shoot tips by browsing White-tailed Deer (Odocoileus virginianus) results in lateral branching. 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 427 (c) Flooding. During the dormant period V. angustifolium can withstand considerable flooding such as that which occurs in many bogs, but it is not characteristic of wet habitats such as marshes or lake margins. (d) Drought. During prolonged drought in early summer shoot growth is reduced. Dry weather later in the season results in shriveling of fruit and reduced flower bud formation. Irrigation prior to harvest substantially increases fruit size. (e) Herbicides. 2,4-D causes a twisting of the terminal growth followed by browning and leaf fall. No data are available on other herbicides. 12. Relationship to Man The fresh fruits of V. angustifolium have been a part of man’s diet in North America since prehistoric times. Indians dried and pulverized fruits for blending with meat (Hedrick 1919). European settlers collected and preserved the fruit for jam, jelly, and preserves as well as eating them raw. These are still the main uses, but new products such as muffin mixes, ice cream, yogurt, and wine (Hope 1965) are using considerable quantities of fruit. For commercial use the fruits are quick-frozen by passing them through a tunnel of air at -29° to -34°C. For many years the only fruit markets for the species were in Canada and the United States. Recently the fruit has found some acceptance in western Europe where it competes with the European V. myrtillus L. The area of lowbush blueberries under management is expanding each year in the Atlantic Provinces, which have programs for assisting growers to develop existing stands of V. angustifolium. Agriculture Canada is also providing funds through the New Crop Development Fund to determine the cost and feasibility of establishing plantings of this species within its range. Acknowledgments The authors acknowledge the technical assistance of A. T. Lightfoot, A. C. Brydon, M. A. Spicer, and B. S. Brooks. Literature Cited Aaiders, L. 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American Book Company, New York, New York. 1632 pp. Forsyth, F. R. and I. V. Hall. 1965. Effect of leaf maturity, temperature, carbon dioxide concentration, and light intensity on rate of photosynthesis in clonal lines of the lowbush blueberry, Vaccinium angustifolium Ait. under laboratory condi- tions. Canadian Journal of Botany 43: 893-900. Forsyth, F. R.and I. V. Hall. 1969. Ethylene production with accompanying respiration rates from the time of blossoming to fruit maturity in three Vaccinium species. Naturaliste Canadien 96: 257-259. Hall, I. V. 1953. Successional studies primarily concerned with three species of Vaccinium. Ph.D. thesis, Cornell University, Ithaca, New York. 119 pp. Hall, I. V. 1955. Floristic changes following the cutting and burning of a woodlot for blueberry production. Canadian Journal of Agricultural Science 35: 143-152. Hall, I. V. 1957. The tap root in lowbush blueberry. Canadian Journal of Botany 35: 933-934. Hall, I. V. 1959. Plant populations in blueberry stands developed from abandoned hayfields and woodlots. Ecology 40: 742-743. Hall, I. V. 1975. The biology of Canadian weeds. 7. Myrica pensylvanica Loisel. Canadian Journal of Plant Science 55: 163-169. Hall, I. V. and F. R. Forsyth. 1967. Production of ethylene by flowers following pollination and treatments with water and auxin. Canadian Journal of Botany 45: 1163-1166. Hall, I. V., F. R. Forsyth, C. L. Lockhart, and L. E. Aalders. 1966. Effect of time of day, a parasitic fungus, and a genetic mutation on rate of photosynthesis in the lowbush blueberry. Canadian Journal of Botany 44: 529-533. Hall, I. V., F. R. Forsyth, H. J. Lightfoot, and R. Boch. 1971. Volatiles of lowbush blueberry nectar. HortScience 6: 493-494. Hall, I. V., F. R. Forsyth, and R. J. Newbery. 1970. Effect of temperature on flower bud and leaf anthocyanin formation in the lowbush blueberry. HortScience 5: 272-273. Hall, I. V.,G. W. Wood, and L. P. Jackson. 1978. The biology of Canadian weeds. 30. Pyrus melanocarpa (Michx.) Willd. Canadian Journal of Plant Science 57: 499-504. Hall, I. V. and J. D. Sibley. 1976. The biology of Canadian weeds. 20. Cornus canadensis L. Canadian Journal of Plant Science 56: 885-892. Hall, I. V.and L. E. Aalders. 1961. Cytotaxonomy of lowbush blueberries in Eastern Canada. American Journal of Botany 48: 199-201. Hall, I. V. and L. E. Aalders. 1968. The botanical composition of two barrens in Nova Scotia. Naturaliste Canadien 95: 393-396. Hall, I. V., L. E. Aalders, A. C. Brydon, and M. A. Spicer. 1977. Rooting lowbush blueberry cuttings. Canadex 235: 21. Hall, I. V., L. E. Aalders, and C. F. Everett. 1976. The biology of Canadian weeds. 16. Comptonia peregrina (L.) Coult. Canadian Journal of Plant Science 56: 147-156. Hall, I. V., L. E. Aalders, and C. L. Lockhart. 1964. Note on some putative lowbush blueberry mutants. Canadian Journal of Botany 42: 124-125. Hall, I. V., L. E. Aalders, and G. W. Wood. 1966. Female sterility in the Common Lowbush Blueberry, Vaccinium angustifolium Ait. Canadian Journal of Genetics and Cytology 8: 296-299. 1979 HALL ET AL.: VACCINIUM ANGUSTIFOLIUM 429 Hall, I. V., L. E. Aalders, L. P. Jackson, G. W. Wood, and C. L. Lockhart. 1975. Lowbush blueberry production. Canada Department of Agriculture Publication 1477. 42 pp. Hall, I. V., L. E. Aalders, and L. R. Townsend. 1964. The effects of soil pH on miineral composition and growth of the lowbush blueberry. Canadian Journal of Plant Science 44: 433-438. Hall, I. V., L. E. Aalders, and R. J. Newbery. 1971. Frost injury to flowers and developing fruits of the lowbush blueberry as measured by impairment of fruit set. Naturaliste Canadien 98: 1053-1057. Hall, I. V., L. E. Aalders, and W. G. Barker. 1963. A preliminary investigation of factors limiting lowbush blueberry production on Cape Breton Island. Canadian Journal of Plant Science 44: 491-492. Hall, I. V., L. P. Jackson, and C. F. Everett. 1973. The biology of Canadian weeds. |. Kalmia angustifolia L. Canadian Journal of Plant Science 53: 865-873. Hall, I. V.and R. A. Ludwig. 1961. The effects of photoperiod, temperature and light intensity on the growth of the lowbush blueberry (Vaccinium angustifolium Ait.). Canadian Journal of Botany 39: 1733-1739. Hall, I. V.,R. A. Murray, and L. P. Jackson. 1974. The biology of Canadian weeds. 2. Spiraea latifolia. Canadian Journal of Plant Science 54: 141-147. Hedrick, U. P. 1919. Sturtevant’s notes on edible plants. J. B. Lyon Company, State Printers, Albany, New York. p. 587. Hersey, R. and S. P. Vander Kloet. 1976. IOPB chromosome number reports LI1. Taxon 25: 241-346. Hope, G. W. 1965. A review of the suitability of the lowbush blueberry for processing. Food Technology 19: 115-119. Jackson, L. P., I. V. Hall, and L. E. Aalders. 1972a. Lowbush blueberry seedling growth as affected by soil type. Canadian Journal of Soil Science 52: 113-115. Jackson, L. P., L. E. Aalders, and I. V. Hall. 1972b. Berry size and seed number in commercial lowbush blueberry fields of Nova Scotia. Naturaliste Canadien 99: 615-619. Kender, W. J. 1968. Rest period in rhizome buds of lowbush blueberry, Vaccinium angustifolium Ait. Proceedings of the American Society for Horticultural Science 93: 254-259. Lamoureux, G. and M.M. Grandtner. 1977. Contribution a l’étude écologique des dunes mobiles. I. Les éléments phytosociologiques. Canadian Journal of Botany 55: 158-171. La Roi, G. H. 1977. The Biological Flora of Canada — a new series. Canadian Field-Naturalist 91(3): 269-272. Lavoie, V. 1968. La phytosociologie et l'aménagement des bleueti¢res. Naturaliste Canadien 95: 397-412. Laycock, W. A. 1967. Distribution of roots and rhizomes in different soil types in the pine barrens of New Jersey. United States Department of the Interior, Geological Survey Professional Paper 563-C. Lockhart, C. L. 1959. Symptoms of mineral deficiency in the lowbush blueberry. Plant Disease Reporter 43: 102-105. Lockhart, C. L. and I. V. Hall. 1962. Note on an indication of shoestring virus in the lowbush blueberry, Vaccinium angustifolium Ait. Canadian Journal of Botany 40: 1561-1562. Longley, A. E. 1927. Chromosomes in Vaccinium. Science 66: 566-568. Marie-Victorin, Frere. 1964. Flore laurentienne. Deuxiéme édition. Les Presses de L’Université de Montréal, Montréal 3, Québec. p. 442. Martin, A. C., H. S. Zim, and A. L. Nelson. 1951. American wildlife and plants. Dover Publications, Inc., New York. pp. 356-357. Maycock, P. F. and J. T. Curtis. 1960. The phytosociology of boreal conifer-hardwood forests of the Great Lakes region. Ecological Monographs 30: 1-35. Newcomer, E. H. 1941. Chromosome numbers of some species and varieties of Vaccinium and related genera. Proceedings of the American Society for Horticultural Science 38: 468-470. Nickerson, N. L. 1978. Callus formation in lowbush blueberry fruit explants cultured in vitro. Horticultural Research 18: 85-91. Nickerson, N. L. and I. V. Hall. 1976. Callus formation in stem internode sections of lowbush blueberry (Vaccinium angustifolium Ait.) cultured on a medium containing plant growth regulators. Horticultural Research 16: 29-35. Poirier, G. and P. Dubé. 1969. Importance de la couverture de neige. /n La recherche sur le bluet. Université Laval. ‘Faculté d’Agriculture. pp. 139-140. Rayment, A. F. 1965. The response of native stands of lowbush blueberry in Newfoundland to nitrogen, phosphorus and potassium fertilizers. Canadian Journal of Plant Science 45: 145-152. Robinson, B. L. and M. L. Fernald. 1908. Gray’s new manual of botany. American Book Company, New York. 926 pp. Rowe, J.S. 1972. Forest regions of Canada. Department of the Environment, Canadian Forestry Service, Publication Number 1300. 172 pp. Smith, D. W. and R. J. Hilton. 1971. The comparative effects of pruning by burning or clipping on lowbush blueberries in North-Eastern Ontario. Journal of Applied Ecology 8: 781-789. Townsend, L. R. and I. V. Hall. 1970. Trends in nutrient levels of lowbush blueberry leaves during four consecutive years of sampling. Naturaliste Canadien 97: 461-466. Townsend, L. R., I. V. Hall, and L. E. Aalders. 1968. Chemical composition of rhizomes and associated leaves of the lowbush blueberry. Proceedings of the American Society for Horticultural Science 93: 248-253. Vander Kloet, S. P. 1976a. A comparison of the dispersal and seedling establishment of Vaccinium angustifolium (the lowbush blueberry) in Leeds County, Ontario and Pictou County, Nova Scotia. Canadian Field-Naturalist 90: 176-180. Vander Kloet, S. P. 1976b. A novel approach to sampling Vaccinium populations. Canadian Journal of Botany 54: 669-671. 430 THE CANADIAN FIELD-NATURALIST Vol. 93 Vander Kloet, S. P. 1976c. Nomenclature, taxonomy, and biosystematics of Vaccinium section Cyanococcus (the blue- berries) in North America. |. Natural barriers to gene exchange between Vaccinium angustifolium Ait. and Vaccinium corymbosum L. Rhodora 78: 503-515. Vander Kloet, S . P. 1977. The taxonomic status of Vaccinium boreale. Canadian Journal of Botany 55: 281-288. Vander Kloet, S. P. 1978. Systematics, distribution, and nomenclature of the polymorphic Vaccinium angustifolium Aiton. Rhodora 80: 358-376. Whitton, L. 1964. The cytotaxonomic status of Vaccinium angustifolium Aiton in commercial blueberry fields of Maine. Ph.D thesis, Cornell University, Ithaca, New York. 150 pp. Wood, F. A. and W. G. Barker. 1963. Stem pigmentation in lowbush blueberry. Plant Physiology 38: 191-193. Wood, G. W. 196la. The association between age of inflorescence and nectar production in the lowbush blueberry Vaccinium angustifolium. Canadian Journal of Botany. 39: 1037-1040. Wood, G. W. 1961b. The influence of honeybee pollination on fruit set of the lowbush blueberry. Canadian Journal of Plant Science 41: 332-335. Wood, G. W. 1962. The period of receptivity in flowers of the lowbush blueberry. Canadian Journal of Botany 40: 685-686. Wood, G. W. 1978. Studies on Neochlamisus cribripennis (Le Conte) with particular reference to its reproductive biology. PhD. thesis, University of New Brunswick, Fredericton, New Brunswick. 95 pp. Received 5 June 1978 Accepted 17 April 1979 Notes Mosses New to Ontario and Quebec ROBERT R. IRELAND! and GILDA BELLOLIO-TRUCCO? ‘National Museum of Natural Sciences, National Museums of Canada, Ottawa, Ontario KIA 0M8 239 Commanche Drive, Ottawa, Ontario K2E 6E8 Ireland, Robert R. and Gilda Bellolio-Trucco. 1979. Mosses new to Ontario and Quebec. Canadian Field-Naturalist 93(4): 431-433. Five mosses are reported new to the bryoflora of Ontario (Cinclidium latifolium, Drepanocladus revolvens var. intermedius, Hypnum hamulosum, Myurella tenerrima, Orthothecium strictum) and five new to Quebec (Atrichum tenellum, Fissidens obtusifolius, Hypnum hamulosum, Orthotrichum strangulatum fo. lescurii, Pseudoleskeella catenulata). Key Words: mosses, Ontario and Quebec flora, geographical distribution, new records. Recent collecting by the authors and others have accounted for several new additions to the moss flora of Ontario and Quebec. The Ontario records are new to the checklist of Ireland and Cain (1975), while the Quebec mosses are not listed by Lepage (1947), Kucyniak (1948, 1949, 1950a, b, c, 1952a, b, c, 1961), Love et al. (1958), LeBlanc (1949, 1951, 1954, 1963), or Masson (1967). In the case of Fissidens obtusifolius Wils., Lepage (1947) reported it for Quebec but LeBlanc (1963) later deleted it from the flora. The purpose of this paper is to report four species and one variety new to Ontario and four species and one form new to Quebec. Voucher specimens are deposited in the National Herbarium of Canada (CANM) in Ottawa. New to Ontario Cinclidium latifolium Lindb. Kenora District: 56°45’N, 88°45’W, coastal raised beach-sedge meadow system, Kershaw, July 1972 (CANM 242656). An interesting moss collected by Kershaw (1974) in his studies of the raised beaches in northwestern Ontario, this species is a circumpolar arctic-alpine calciphile that was previously known in Canada from northern Manitoba, Yukon Territory, and Northwest Territories. Mogensen (1973) has mapped its world- wide distribution. Drepanocladus revolvens var. intermedius (Lindb. ex C. Hartm.) Grout Bruce County: Cemetery Road, Caesar’s Bog, C. Williams 1107. Cochrane District: island in Moose R. near mouth of Abitibi R., 51°03’N, 80°57’W, Baldwin B-22. Rainy River District: 5 mi [8 km] NE of Gameland, Garton 9263. Thunder Bay District: Sibley Peninsula, bog at NW corner of Middlebrun Bay, 2.75 ml [4.4 km] NE of Silver Islet, Garton 4362. 431 Ireland and Cain (1975) considered this variety a synonym of the typical variety; however, there has been so much debate (Steere 1978) regarding its taxonomic status that we have decided to recognize it as a distinct taxon until a detailed study can be made. The variety intermedius is a calciphile that is also known from British Columbia, Newfoundland, Yukon Territory, and Northwest Territories. Hypnum hamulosum B.S.G. Kenora District: 56°45’N, 88°45’W, coastal raised beach-sedge meadow system, Kershaw, July 1972 (CANM 242653). Verified by H. A. Crum. Another calcicolous arctic-alpine species, like Cin- clidium latifolium, that was collected by Kershaw (1974) in his raised beaches studies, this species has been reported for all Canadian provinces and terri- tories except Prince Edward Island and New Bruns- wick. Its presence in Newfoundland (Macoun and Kindberg 1892) and Nova Scotia (Erskine 1968) seems doubtful, however. Ando (1972) has mapped the worldwide distribution of this Hypnum, as well as others that occur in North America. . Myurella tenerrima (Brid.) Lindb. Thunder Bay District: Ourmet Canyon, 10 km W of Dorion, 48° 43’N, 88°40’W, Garton 11634, 15365, 15371; canyon at NW corner of Cavern Lake, 16 km NNW of Dorion, 48°50’N, 88°41’W, Garton 14997, 15098. This calcicolous arctic-alpine species is now known from all provinces and territories in Canada except Saskatchewan, Newfoundland, New Brunswick, Nova Scotia, and Prince Edward Island. Its occurence in Ouimet and Canyon Lake canyons is extremely interesting as other arctic mosses, such as Au/acomni- um acuminatum (Lindb. & H. Arnell) Kindb. (Wil- liams 1968) and A. turgidum (Wahlenb.) Schwaegr. also occur there, reaching their southern limit in these two Ontario localities. These mosses are arctic relicts 432 that have been able to survive at the bottom of these canyons far south of their present ranges because of the cold microenvironment created among huge boulders. Orthothecium strictum Lor. Kenora District: south shore Attawapiskat River, 52°52’N, 83°45’W, Riley 6512. This is a circumboreal arctic-alpine calciphile that is also known in Alberta, Quebec, Newfoundland, Yukon Territory, and Northwest Territories. New to Quebec Atrichum tenellum (Roehl.) B.S.G. Gatineau County: Gatineau Park, trail N of Cham- plain Lookout, NW of Hull, 45°30’N, 75°54’W, Treland, Iwatsuki & Kuc 9684. Determined by G. L. Smith. This collection was originally identified as A. crispum (James) Sull. & Lesq. but G.L. Smith discovered the misidentification and informed the senior author (personal communication) that it represented a species new to North America. Ireland (1971) subsequently reported the species for southern British Columbia and Quebec but without specific collection data. The species is now known from scattered localities in British Columbia, Manitoba, Ontario, Labrador, Newfoundland, Prince Edward Island, and New Brunswick. Fissidens obtusifolius Wils. Gatineau County: Stag Creek, near Low, ca. 45°48’N, 76°07’W, Ireland & Bellolio-Trucco 18099. This distinct calciphile, which was reported by Lepage (1947) for Quebec, was later deleted from the flora by LeBlanc (1963). It is probably rare in the province but it may eventually be found in other localities in southern Quebec. It seems to be restricted to eastern Canada where it is also known from southern Ontario. Hypnum hamulosum B.S.G. James Bay region, 54°53’N, 79°08’W, Lethiecq OFB-E 11498. This species is also reported as new to Ontario in this paper. Orthotrichum strangulatum fo. lescurii (Aust.) Vitt Gatineau County: Gatineau Park, Luskville Falls, NE of Luskville, 45°32’N, 76°00’W, Ireland & Bellolio-Trucco 17922. Determined by D. H. Vitt. This form differs from the typical form strangu- latum, which is also known from Quebec, in several minor variations of the gametophyte and sporophyte (Vitt 1973). The form /escurii has been reported for British Columbia and Ontario (Grout 1935) but its occurrence in British Columbia is doubtful according to Vitt (personal communication). THE CANADIAN FIELD-NATURALIST Vol. 93 Pseudoleskeella catenulata (Brid. ex Schrad.) Kindb. Gatineau County: Paugan Falls, 1 mi[1.6 km] E of Low, ca. 45°49’N, 75°5S’W, Ireland & Bellolio- Trucco 18050. This calcareous species is also known from Ontario, Labrador, and Northwest Territories. Lewinsky (1974), who mapped the worldwide distribution, showed a record in the Gaspé Peninsula but we have seen neither a literature report nor a herbarium specimen for that locality. Acknowledgments We thank H. A. Crum, University of Michigan, G. L. Smith, New York Botanical Garden, and D. H. Vitt, University of Alberta, for identifying or verifying some of the collections. Literature Cited Ando, H. 1972. Distribution and speciation in the genus Hypnum in the circum-Pacific region. Journal of the Hattori Botanical Laboratory 35: 68-98. Erskine, J. S. 1968. An introductory moss flora of Nova Scotia. Nova Scotia Museum, Occasional Paper 6, Science Series 4. 110 pp. Grout, A. J. 1935. Moss flora of North America, north of Mexico. Newfane Volume 2(2). pp. 67-138. Ireland, R. R. 1971. /n Moss flora of the Pacific Northwest. By E. Lawton. Hattori Botanical Laboratory, Nichinan. 362 pp. Ireland, R.R. and R.F. Cain. 1975. Checklist of the mosses of Ontario. National Museums of Canada, Publi- cations in Botany 5. 67 pp. Kershaw, K. A. 1974. Studies on lichen-dominated sys- tems. X. The sedge meadows of the coastal raised beaches. Canadian Journal of Botany 52(8): 1947-1972. Kucyniak, J. 1948. Sur trois additions a la flore bryolo- gique du Québec. Annales de !1ACFAS 14: 67. Kucyniak, J. 1949. On the occurrence of Tortula latifolia Bruch in Québec. Bryologist 52(1): 32-33. Kucyniak, J. 1950a.. Mise au point sur deux muscinées québécoises: Hygrohypnum smithii et Oncophorus virens var. serratus. Annales de TACFAS 16: 154-156. Kucyniak, J. 1950b. Muscinées nouvelles pour le Québec: Calypogeia fissa, Odontoschisma elongatum, Didymodon rufus et Hypnum bambergeri. Naturaliste Canadien 77(1 1-12): 305-312. Kucyniak, J. 1950c. Le genre Splachnum dans le Québec. Revue Bryologique et Lichénologique 20(1—2): 38-42. Kucyniak, J. 1952a. The occurrence of Grimmia teretiner- vis Limpr. in North America, with notes on its distribu- tion. Bryologist 55(1): 35-47. Kucyniak, J. 1952b. Notes sur les Pohlia du Québec. I. Deux espéces peu fréquentes: P. filiformis et P. proligera. Naturaliste Canadien 79(6—7): 233-238. Kucyniak, J. 1954. Notes sur les Pohlia du Québec. Il. P. bulbifera et P. drummondii. Naturaliste Canadien 81(10- 11): 197-202. Kucyniak, J. 1955a. Mousses nouvelles ou intéressantes de la bryoflore du Québec. Annales de ?ACFAS 21: 104-108. 1979 Kucyniak, J. 1955b. Le genre Seligeria dans le Québec. Annales de PACFAS 21: 110-115. Kucyniak, J. 1955c. An overlooked moss in the Quebec flora: Drepanocladus brevifolius. Svensk Botanisk Tids- skrift 49(1-2): 325-328. Kucyniak, J. 1955d. Précisions sur le genre Thelia dans le Québec. Naturaliste Canadien 82(2-3): 45-48. Kucyniak, J. 1957a. Notes sur les Pohlia du Québec. III. Addition du P. sphagnicola et extension d’aire du P. bulbifera. Naturaliste Canadien 84(12): 249-253. Kucyniak, J. 1957b. Un Fissidens nouveau pour le Québec. Annales de ! ACFAS 23: 94. Kucyniak, J. 1958a. Notes sur les Pohlia du Québec. IV. Ré-intégration du P. cucullata dans notre flore. Natur- aliste Canadien 85(4): 94-100. Kucyniak, J. 1958b. Une mousse inattendue pour le Qué- bec: Prerigoneurum ovatum. Naturaliste Canadien 85(10): 217-224. Kucyniak, J. 1958c. On Drepanocladus badius in continen- tal North America. Bryologist 61(2): 124-132. Kucyniak, J. 1961. Un autre Desmatodon pour le Québec. Naturaliste Canadien 88(6—7): 161-165. LeBlanc, F. 1949. Additions a la bryoflore du Québec. Naturaliste Canadien 76(8-10):; 223-228. LeBlanc, F. 1951. Ulota drummondii (Hook, et Grev.) Brid. en Amérique du Nord. Revue Bryologique et Liché- nologique 20(3—4): 260-262. LeBlanc, F. 1954. Additions a la bryoflore du Québec (2e série). Naturaliste Canadien 81(3—4): 90-99. LeBlanc, F. 1963. Notes sur les mousses du Québec. I. Naturaliste Canadien 90: 41-50. NOTES 433 Lepage, E. 1947. Les lichens, les mousses et les hépatiques du Québec et leur réle dans la formation du sol arable dans la région du bas de Québec de Lévis 4 Gaspé. Naturaliste Canadien Extract (fascicle 4. Les mousses): 125-283. Lewinsky, J. 1974 (1975). The genera Leskeella and Pseu- doleskeella in Greenland. Bryologist 77(4): 601-611. Love, D., J. Kucyniak, and G. Johnston. 1958. A plant collection from interior Quebec. Naturaliste Canadien 85: 25-69. Macoun, J. and N. C. Kindberg. 1892. Catalogue of Cana- dian plants. Part 6, Musci. Geological Survey of Canada. Montreal. 295 pp. Masson, P. 1967. Notes sur quelques muscinées du Québec. Annales de PACFAS 33: 39. Mogensen, G.S. 1973. A revision of the moss genus Cinclidium Sw. (Mniaceae Mitt.) Lindbergia 2: 49-80. Steere, W. C. 1947. 6. Musci. Jn Botany of the Canadian Eastern Arctic. Part II. Thallophyta and Bryophyta. Edited by N. Polunin. National Museum of Canada Bulletin 97: 370-490. Steere, W. C. 1978. The mosses of Arctic Alaska. Bryo- phytorum Bibliotheca 14. 508 pp. Vitt, D. H. 1973. A revision of the genus Orthotrichum in North America, north of Mexico. Bryophytorum Bibli- otheca |. 208 pp. Williams, C. 1968. New and additional moss records for Ontario. Bryologist 71(3): 282-284. Received 5 March 1979 Accepted 31 March 1979 Swans Wintering on Vancouver Island, 1977-1978 RICHARD W. MCKELVEY Canadian Wildlife Service, P.O. Box 340, Delta, British Columbia V4K 3Y3 McKelvey, Richard W. 1979. Swans wintering on Vancouver Island, 1977-1978. Canadian Field-Naturalist 93(4): 433-436. An aerial survey of swans wintering on Vancouver Island, British Columbia in February 1978 revealed a minimum of 1065 birds. Most were believed to be Trumpeter Swans (Olor buccinator) although some Mute Swans (Cygnus olor) were present on southern Vancouver Island. Cygnets accounted for 21.5% of the total. Changes in wintering distribution were noted: the largest population shift was into the agricultural areas of Comox and Nanaimo. Key Words: Trumpeter Swan, Olor buccinator, Mute Swan, Cygnus olor, wintering, Vancouver Island, estuaries. Swans winter along the coast of western North America, from Alaska to southern Washington (Bellrose 1976). Counts of winter populations have generally been sporadic owing to the extent and remoteness of the winter habitat, except on Van- couver Island. Estimates of the number of swans wintering there have been made periodically over the last eight years. In 1977-1978 I conducted an aerial survey of Vancouver Island areas frequented by swans. Objectives of this survey were further to assess winter swan populations, determine the proportion of cygnets, and document changes in major concentra- tion areas. Methods The survey, conducted 16-18 February 1978, covered most areas previously surveyed (D. Trethe- wey, personal communication) including most estu- aries and lakes found to be ice-free. Northern Vancouver Island, from Cape Scott to Port Hardy, 434 was not surveyed because of poor weather: its omission is not considered important, because previ- ous surveys recorded few swans there and habitat is limited. A float-equipped Cessna 185 was flown at an altitude of approximately 215 m. Observations were made from both sides of the aircraft. In areas of high concentration the flocks were counted as the aircraft circled, and the counts were verified later from photographs. Results and Discussion Population Status Table | compares results of the 1977-1978 survey with those of previous surveys. Vancouver Island has been divided into eight regions based on landforms (after Holland 1964). The regions and sighting locations are shown in Figure 1. In the present study 1065 swans were counted, a result which is not substantially different from those reported previously. Smith and Blood (1972) esti- mated a peak population of 1076 in the period 1969-1971. This was based on the results of a single aerial count of 1013 birds in February 1971, and several estimates made by others (I. Smith and D. Trethewey. unpublished data). I. Smith and D. Trethewey recorded only 892 swans in 1972-1973. The NUMBER OF SWANS e 1-10 @ \i\-25 @ 26-50 @ 51-100 KILOMETERS 20 0 20 40 60 80 100 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Swans counted during aerial surveys on Van- couver Island Number of swans (cygnets) Region 1970-71! 1972-73 1977-78 Parksville-Victoria 62 48(4) 2292(45) Barkley Sound 90 35(1) 109 (30) Alberni Basin 1128 195(25) 97 (12) Ucluelet-Quatsino 308 207(27) 166 (27) Cape Scott 107 90(10) 44 (6) Kelsey Bay 140 129(20) 76 (38) Comox Valley 112 108(7) 246 (52) Adults and immature 760 789 836 cygnets 2463 103 229 'From Smith and Trethewey (unpublished data). 2Including two Mute Swans. 3Based on an average of 24.3% cygnets, after Smith and Blood (1972). variability between counts 1s reduced, however, when cygnets are eliminated from the comparison(Table 1). There seems to have been little or no increase in numbers of adult and subadult swans since 1969- 1971. The estimate by Smith and Blood (1972) of a 129% increase over the 1969-1971 period may have resulted from incomplete surveys, or from the VANCOUVER ISLAND FIGURE |. Locations of swan sightings in the 1977-1978 winter survey. 1979 inclusion of cygnets in their calculations. The year-to- year variation in cygnet production can be large. so estimates of population increases may be biased if cygnet numbers are included. Much of the apparent increase on this survey may be the result of previous surveys not having con- sidered resident Mute Swans (Cygnus olor) estab- lished locally from escaped park birds. I believe most swans I observed were Trumpeter Swans (Olor buccinator) as did Smith and Blood (1972). Both Mute Swans and Whistling Swans (O. columbianus), however, are regularly recorded on Vancouver Island in winter (British Columbia Provincial Museum sight records (BCPM)). Records of Mute Swans on Vancouver Island extend back to at least 1945 (BCPM). The feral population had increased to at least 100 birds by 1972 (Tatum 1973). Christmas Bird Counts indicate that - the population is still increasing but is confined to the Victoria-Ladysmith area (American Birds, Volume 25-32). Although only two Mute Swans were iden- tified in the 1977-1978 survey, on Somenos Lake near Duncan, it seems likely that many more of the swans recorded in that area were of this species. Few swans were reported in previous aerial surveys between Ladysmith and Victoria (Table |). Because it is unlikely that Mute Swans were identified and excluded from the reported results of those surveys, they may have been concentrated in areas not thoroughly surveyed, such as Greater Victoria (D. Trethewey, personal communication). If Mute Swans in the Ladysmith-Victoria area were missed in previous surveys, their inclusion in my survey may be responsible for the apparent increase. The 65 adults I counted in that area is close to the number of Mute Swans reported (67) in the 1976-1977 Christmas Bird Count (Weseloh 1977, p. 457). Lowering the adult and subadult total for 1977-1978 by 65 gives a figure very close to those for 1970-1971 and 1972-1973 (Table 1). Weekly counts of swans at Comox Harbour and Port Alberni in 1977-1978 showed 1-3% were Whis- tling Swans. Other records (BCPM) indicate that Whis- tling Swans are usually seen only on southern Vancouver Island. That may be because most ob- servers are from the southern part of the island, or it may reflect the more southerly winter distribution of Whistling Swans (Bellrose 1976). If Whistling Swans were uniformly distributed a conservative estimate of their numbers would be 2% or 20 birds. Cygnets accounted for 229 or 21.5% of the swans counted in this study. The proportions of cygnets reported by Smith and Blood (1972) ranged between 22% and 26%, which they suggested was indicative of an expanding population not yet limited in breeding habitat. Although productivity between years 1s NOTES 435 variable (Table 1), factors other than availability of breeding habitat may be limiting the population. King (1976) reported a yearly breeding population growth of only 3.5%. Mortality may be high during the winter (J. King, personal communication) or during spring migration in order for the population to be expanding so slowly. Major Concentrations Large concentrations in 1977-1978 were at Nanai- mo Harbour (58), Mitchell Lake (55), Nanaimo Lakes (24), Somenos Lake (28), Cowichan River estuary (35), Port Alberni (72), Sarita River estuary (53). Kelsey Bay (76), Northy Lake (91), and Comox Harbour (92). Areas of major use seem to have changed between surveys (Table 2). The largest decrease was at Holberg, from 60 in 1970-1971 to only 8 in 1977-1978, but no regular ground counts exist for that area: local movements may have affected the counts in one or more surveys. TABLE 2—Numbers of swans wintering in areas reported to have large concentrations in previous surveys Number of swans Area 1969-71' 1972-732 1977-783 Kelsey Bay 140 129 76 Pt. Alberni 83 195 97 Kyuquot Sound 81 39 35 Tlupana Inlet 78 57 8 Comox Harbour 74 79 185(274)4 Holberg 50 37 8 Herbert Inlet 51 4 1] Nanaimo 51 48 15] Chewat River 48 45 28 ‘Smith and Blood (1972). 2Smith and Trethewey (unpublished data). 3This study. 4Maximum observed in weekly ground counts at Comox. 1977-1978. The largest increases have been in the Comox and Nanaimo areas. Both locations have estuaries, but the major surrounding land use is agricultural. More swans were seen on adjacent farmland than on either estuary in this study. In both areas there has been a trend in the last 10 yr towards more intensive dairy farming. Consequently pastures are now planted with fast-growing grasses of high nutrient content. I believe the attractiveness of these pastures and their prox- imity to open water are the main reasons swans are concentrating in these areas. Although adjacent estuaries continue to be important when snow or frost keeps the swans off the pastures, conflicts with farmers seem inevitable, should swan use of pasture land continue to increase. 436 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 3—Numbers of swans wintering in areas subject to frequent ground counts or regular Christmas Bird Counts Location Date Comox! Nanaimo? Kelsey Bay? Port Algerni3 1972-73 5] 23 No data No data 1973-74 56 14 98 No data 1974-75 144 28 83 No data 1975-76 No data 36 73 178 1976-77 No data 50 59 No data 1977-78 264 75 75 80 'Data from D. Trethewey (personal communication) and this study. 2Data from American Birds. Vol. 27-32. 3Data from Copland (1976. p. 212) for 1974-75. and this study in 1977-78. Other data support my belief that swans may be relocating into certain areas (Table 3). Ground counts conducted by the Canadian Wildlife Service at Comox showed that peak counts have increased. Data from Christmas Bird Counts, while not as rigorous as the ground counts at Comox, show similar trends. Nanaimo also showed a general increase, while Kelsey Bay showed a decrease in swan use. Local British Columbia Fish and Wildlife Branch personnel also believe that these trends are real: the locations of major concentrations of swans are changing. More research is needed to determine the habitat and food requirements of wintering swans. Alter- native means of reducing swan use of pasture grasses, including lure crops and scaring, willalso need study if such use increases. Acknowledgments The assistance of D. Smith, B. Whitehead (pilot), D. Trethewey, R. W. Campbell, J. P. Kelsall, and the reviewers is greatly appreciated. Literature Cited Bellrose, F. C. 1976. The ducks, geese and swans of North America. Stackpole Books, Harrisburg, Pennsylvania. Copland, H.W.R. (Compiler). 1976. Christmas bird count for 1975-76; Western Canada. American Birds 30: 208-216. Holland, S.S. 1964. Landforms of British Columbia. A physiographic outline. British Columbia Department of Mines and Petroleum Resources, Bulletin 48. King, J. G. 1976. The current status and future of the Alaska Trumpeter Swan population. Proceedings of the Fifth Trumpeter Swan Society Meeting (in press). Smith, I. D. and D. A. Blood. 1972. Native swans winter- ing on Vancouver Island over the period 1969-71. Can- adian Field-Naturalist 86: 213-216. Tatum, J.B. 1973. Annual bird report for southern Van- couver Island 1972. Victoria Natural History Society. Victoria, British Columbia. Weseloh, D. V.(Compiler). 1977. Christmas bird count for 1976-77: Western Canada. American Birds 31: 457-466. Received 18 September 1978 Accepted 27 March 1979 Interspecific Vocal Mimicry by Pine Grosbeaks PETER TAYLOR P.O. Box 597, Pinawa, Manitoba ROE 1L0 Taylor, Peter. 1979. Interspecific vocal mimicry by Pine Grosbeaks. Canadian Field-Naturalist 93(4): 436-437. Mimicry of the calls of the Hairy Woodpecker, Gray Jay, American Robin, and Redpoll (sp.) by adult male Pine Grosbeaks, during “whisper singing” on the species’ winter range, is described. Key Words: Pine Grosbeak; Pinicola enucleator; vocal mimicry. This note describes vocal mimicry during “whisper singing” by adult male Pine Grosbeaks (Pinicola enucleator) on the species’ winter range in south- eastern Manitoba. Interspecific vocal mimicry has been reported for several bird species (e.g., Adkisson and Conner 1978; Armstrong 1973), but the only suggestion of such behavior by Pine Grosbeaks is a reference to four birds uttering “an amplified version of the common flight call of the goldfinch” (Bent 1968, p. 333). Soft, warbled “whisper singing” is docu- 1979 mented for many North American fringillids in- cluding Pine Grosbeaks (Bent 1968). At 09:00 on 26 February 1978 I noted several Pine Grosbeaks in Pinawa; three of these were uttering soft warbling songs. One adult male sang particularly softly, and interspersed the warbled phrases with realistic imitations of several other birds’ calls, uttered clearly but softly. | approached within 5 m of the bird as it perched alone in a small aspen, so there is no doubt that it gave all the calls noted. These included the ‘pic’ call-note of a Hairy Woodpecker ( Picoides villosus), the harsh *kuk-kuk-chikkikik’ call of a Gray Jay (Perisoreus canadensis), the clucking call and rattling alarm-cry of an American Robin (Turdus migratorius), and the call-note, with rising inflection, of a Redpoll (Carduelis sp.). Some slurred whistles were Starling-like (Sturnus vulgaris), but not un- equivocally recognizable. A muted version of the Pine Grosbeak’s own triple-noted flight call was also included. These various calls were interjected, with no apparent ordered sequence, at intervals of about 3 s into the warbled song, and the medley was sustained for periods of 20 s or more. All of the imitations were not included in each bout of singing. The bird sang for about half of the observation period of about 15 min, and was still singing when I left. At 11:30 on 5 March 1978 I heard what was presumably the same bird uttering similar vocaliza- tions at the same location. The song was less sustained on this occasion, and interference from extraneous noise made observation difficult. NOTES 437 At 12:00 on 4 February 1979 I observed an adult male Pine Grosbeak uttering a “whisper song” while foraging in a spruce tree in Pinawa. This song continued, with one 15-s break, for about 6 min. It consisted of a variety of whistled and warbled phrases, and included imitations of American Robin and Hairy Woodpecker call-notes. Although it is possible that the same individual Pine Grosbeak was involved in both the 1978 and 1979 records, the observation of this behavior in two winters does suggest that vocal mimicry may not be uncommon in Pine Grosbeaks. All of the birds imitated, except the American Robin, winter regu- larly in the Pinawa area. If the Pine Grosbeaks had wintered in southeast Manitoba, then they would have learned the robin calls the previous fall at the latest. I thank Richard W. Knapton for helpful comments on this note. Literature Cited Adkisson, C.S. and R.N. Conner. 1978. Interspecific vocal imitation in White-eyed Vireos. Auk 95: 602-606. Armstrong, E. A. 1973. A study of bird song. Dover, New York, New York. 347 pp. Bent, A.C. (and collaborators). 1968. Life histories of North American cardinals, grosbeaks, buntings, towhees, finches, sparrows and allies. U.S. National Museum Bulletin 237. Received 14 November 1978 Accepted 3 February 1979 7 May 1979 (amended) Influence of Weather on Aggression in Tree Swallows! REID N. HARRIS Department of Zoology, University of Maryland, College Park, Maryland 20742 USA Harris, Reid N. 1979. Influence of weather on aggression in Tree Swallows. Canadian Field-Naturalist 93(4): 437-438. Cool foggy weather significantly reduced territorial aggression in a Tree Swallow (/ridoprocne bicolor) population in New Brunswick. Key Words: aggression, weather, Tree Swallow, /ridoprocne bicolor. Weather can influence the behavior of animals in nature. For example, Grubb (1978) and Travis (1977) have found that weather affected the daily and seasonal foraging habits of birds. This note reports on the effects of weather on territorial aggression in the 'This paper is a contribution from the Bowdoin Scientific Station, Kent Island, New Brunswick, Canada. Tree Swallow, /ridoprocne bicolor. Study Site and Methods From 17 May until | August 1978, I conducted a study of aggression, territoriality, and reproductive success in Tree Swallows on Kent Island, New Brunswick, Canada. Tree Swallows have nested in uniformly spaced man-made boxes in two open fields on Kent Island for 45 yr (Paynter 1954). In 1978, two 438 new spatial patterns were created by moving nest- boxes from their original 30-m uniform spacing, leaving 12 uniformly spaced boxes as a control comparable with previous years. The first spatial pattern consisted of alternating, at 30-m intervals, a pair of boxes, | m apart, with a single box. Twenty- eight boxes were arranged in this manner. The second pattern consisted of two replicates of a spiral arrangement of five boxes placed so that boxes were 1 m,2 m,4 m,and 8 maway from the central box. The two spirals were 30 m apart: also 30 m separated the nearest boxes of adjacent spatial patterns. All breeding females and several breeding males were color-marked for individual recognition (see Hoogland and Sherman 1976). Adult Tree Swallow behavior was recorded for 4 wk, including the approximately 20-d nestling period of Tree Swallows on Kent Island (Paynter 1954: this study). Twenty-five occupied nest-boxes from all spatial patterns were selected at random (by drawing numbers from a hat) from a total of 36 active boxes for observation. Two or three active nest-boxes were watched simultaneously for 45-min periods. Each box was observed for about 7h during the 4-wk observational study: a total of 182 h of observation was accumulated. Observations were conducted in all kinds of weather except driving rain from 09:00 to 14:30 Atlantic Daylight Time. Warm days (>12°C) with high visibility (>100 m) were classified as “weather 1” days. Cool days (S<12°C) with low visibility (>100 m:; i.e., fog) were Classified as “weather 2” days. All observational data fit into one of these categories: foggy days were always cool. During observations, the amount of territorial aggression and the locations of territorial boundaries were carefully monitored. Fights were defined as two or more birds actually contacting one another in an aggressive territorial encounter. Chases were similar to fights, but lacked actual contact. All aggression occurred in the context of territorial defense of nest- boxes, 1.e., marked and unmarked resident birds of both sexes excluded unmarked swallows from their territories throughout the nestling period. Results and Discussion Cool foggy weather significantly reduced territorial aggression in Tree Swallows (Table 1). One striking example of the influence of weather on aggression was observed on 11 July. From the beginning of observa- tions at 09:00 until 10:00, cold foggy weather prevailed, and aggression was non-existent. At 10:00, a rapid clearing began. Concomitantly, swallows at the two active boxes being watched began defending THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE I—Aggression rates, mean + SE of aggressive inter- actions per pair of Tree Swallows during a 45-min period, by weather are based on 182 box h. All 25 watched pairs were observed on weather | days, but only 19 pairs on weather 2 days Aggression Weather | Weather 2 ies Fights AA) 22 O22 0.18+0.16 <0.001 Chases 0.61 + 0.28 0.11 + 0.03 >0.05 Total 2.6 + 0.43 0.29+0.07 <0.001 ‘t-test; variances unequal. their nest-boxes against unmarked intruders. Fifteen separate fights that involved actual contact between birds were observed between 10:10 and 10:30. Aggres- sive activity was directly associated with weather conditions. The exact proximal causes of the reduction in aggression during cool foggy weather are unknown. Lowered visibility associated with fog, however, could interfere with the swallows’ cues of individual recogni- tion so that potential aggressors may not be recog- nized except at very close distances. Alternatively, during periods of low visibility, foraging for the same quantity of insects must have taken longer than foraging in clear weather. Under foggy conditions, and given a parental commitment to feed nestlings, most of the swallows’ time would have been taken foraging. Whether these explanations can be distin- guished or demonstrated is the subject for future work. Acknowledgments This work was supported in part by the A. O. Gross Fund. I thank D. E. Gill, G. L. Holroyd, P. Wood- ward, and an anonymous referee for helpful com- ments on the manuscript, and R. Robertson for suggesting the spiral pattern for spacing nest-boxes. Literature Cited Grubb, T. C. 1978. Weather-dependent foraging rates of wintering woodland birds. Auk 95: 370-376. Hoogland, J.L. and P. W. Sherman. 1976. Advantages and disadvantages of Bank Swallow coloniality. Ecologi- cal Monographs 46: 33-58. Paynter, R.A. 1954. Interrelations between clutch size, brood size, prefledging survival and weight in Kent Island Tree Swallows. Bird-Banding 25: 35-58, 102-111, 136-149. Travis, J. 1977. Seasonal foragingina Downy Woodpecker population. Condor 79: 371-375. Received 19 February 1979 Accepted 11 April 1979 1979 NOTES 439 Abandoned Windmill Used as a Nesting Site by Great Blue Herons JEAN-LUC DESGRANGES Canadian Wildlife Service, 2700 Laurier Blvd., P.O. Box 10100, Ste-Foy, Québec G1V 4H5 DesGranges, Jean-Luc. 1979. Abandoned windmill used as a nesting site by Great Blue Herons. Canadian Field-Naturalist 93(4): 439-440. An active Great Blue Heron (Ardea herodias) nest was discovered on the platform of an abandoned windmill at Batture aux Loups-Marins, Quebec. Among the hundred or so heronries known in Quebec, this was the only one where a man-made structure was used. Un nid actif de Grand Héron (Ardea herodias) installé sur la plateforme d’une éolienne désaffectée a été découvert a la Batture aux Loups-Marins, Québec. Parmi la centaine de héronniéres connues au Québec jusqu’a ce jour, cest la premiére fois que Yon observe utilisation d’une structure artificielle par cette espéce. Key Words: Ardea herodias, nesting platform. On 25 April 1978, while conducting anaerialsurvey —_ any of the nests located in trees, the usual nesting site of aquatic bird colonies in the St. Lawrence estuary for this species. for the Canadian Wildlife Service, Pierre Dupuis and I discovered an active Great Blue Heron (Ardea herodias) nest on the platform of an abandoned windmill at Batture aux Loups-Marins (47°14’N, 70°25’W) (Figure 1). We located four other nests being built in a grove of tall willows (Sa/ix sp.) in the vicinity of the windmill. There were no Great Blue Herons at this site prior to 1978 (Reed 1973; personal observation 1977). On | June, the heronry contained eight active nests, four of which contained only eggs (K + SD = 3.0 + 1.6) whereas the others, including the nest on the windmill, held both eggs and young. On 3 July, six nests remained, all with young herons (K + SD = 2.0 + 1.2). The nest-located on the windmill held four young herons whereas the others held fewer than three each. Although Great Blue Herons have been recorded nesting on duck-hunting blinds (Stotts 1959) and navigational buoys (Henny 1978), it is nonetheless unusual for them to nest on artificial platforms. Among the hundred or so heronries known in Quebec, this was the only one in which a man-made structure was used (DesGranges, unpublished data). Neverthe- less, other species of herons have been known to nest on artificial structures (MclIlhenny 1934; Finkenstaedt and Heckenroth 1974; Wiese 1976). This suggests that the erection of elevated platforms where Great Blue Heron natural nest sites have been destroyed could prove to be an effective remedial technique, providing other features of the habitat remain suitable. At Batture aux Loups-Marins, the artificial platform was met occupied from the time the heronry was first establish- Figure 1. Great Blue Heron nest built on platform of ed and the nest found on it produced more young than abandoned windmill. I thank Transport Canada for making a helicopter available for our use in the St. Lawrence estuary survey. 440 Literature Cited Finkenstaedt, C. and H. Heckenroth. 1974. Eine kunstliche kolonie grundung beim Graureiher (Ardea cinerea). Vogelwelt 95: 227-230. Henny, C. D. 1978. Great Blue Herons respond to nesting habitat loss. Wildlife Society Bulletin 6: 35-37. Mclihenny, E. A. 1934. Bird city. Christopher Publishing House, Boston. 203 pp. Reed, A. 1973. Aquatic bird colonies in the St. Lawrence THE CANADIAN FIELD-NATURALIST Vol. 93 estuary. Service de la faune du Québec, Bulletin 18. 54 pp. Stotts, V. D. 1959. Offshore duck blinds: Their use by wild- life and how to improve them for wildlife use. Maryland Conservationist 36: 23-26. Wiese, J.J. 1976. Courtship and pair formation in the Great Egret. Auk 93: 709-724. Received 8 January 1979 Accepted 9 June 1979 Unusually Late Pregnancy of a Muskrat in Southeastern New Brunswick G. R. PARKER Canadian Wildlife Service, P.O. Box 1590, Sackville, New Brunswick E0OA 3C0 Parker, G. R. 1979. Unusually late pregnancy of a Muskrat in southeastern New Brunswick. Canadian Field-Naturalist 93(4): 440-441. A Muskrat (Ondatra zibethicus zibethicus) containing nine embryos was trapped on 6 November 1978 on the Tantramar marshes in southeastern New Brunswick. The projected birth date for the embryos was the last week of November. This is the latest reported pregnancy for a muskrat in eastern Canada. Key Words: Muskrat, late pregnancy, New Brunswick, Ondatra zibethicus zibethicus. As part of a study of Muskrats (Ondatra zibethicus zibethicus) on the Tintamarre National Wildlife Area, Westmorland County, New Brunswick, carcasses of Muskrats caught during the 1978 fall trapping season (4 November — 31 December) were examined for sex, age, productivity, and other morphological measure- ments. Productivity was measured by placental scar counts from macroscopically examined female uterl. The uterus of one female caught on 6 November contained nine embryos; these averaged 5-7 mm in length and were believed to be approximately 10-14 d old. The normal gestation period for Muskrats in northeastern North America is 25-30 d (Errington 1963). This Muskrat would have given birth during the last week of November. This Muskrat had been live-trapped and ear-tagged 2 August; it weighed 1000 g. When caught in November it weighed 1490 g. Its exact age is uncertain. When tagged in August, it was subjectively classified as an adult, based upon size and appearance. When examined in November the molar fluting was at bone line, which left the age of the animal in question. The uterus showed no other scars of earlier preg- nancies. It seems most reasonable that this Muskrat was born late in the autumn of 1977, and that possibly it too came from a fall litter. This would account for its adult appearance in August, the questionable age from molar fluting, the absence of previous breeding, and its substantial weight gain in late summer and fall. The Tantramar marshes of the New Bruns- wick — Nova Scotia border region con- tain some of the best habitat for Muskrats in the Maritimes. Under favorable conditions, it seems reasonable that autumn or fall breeding may oc- casionally occur, and that some kits survive the winter to enter the next year’s breeding population. Such late-born females, however, probably do not breed until late the following summer. Muskrats may give birth during all months of the year in California (Dixon 1922), Louisiana (Svihla and Svihla 1931), and Texas (Lay 1945). Further north, in Maryland, breeding normally ceases by late October (Forbes 1942). Late litters have been reported in September from Wisconsin (Beer 1950), east Tennessee (Schacher and Pelton 1975), Connecticut (Smith and Jordan 1976), and Manitoba (McLeod and Bondar 1952), and in October from Idaho (Errington 1963). In New Brunswick (Dilworth 1967) and Prince Edward Island (Dibblee 1970) the latest observed litters were born in September. Errington suggested that it is quite possible litters might rarely be sired during winter in lowa. This specimen from the Tantramar marshes of New Brunswick, however, is the first documented evidence of a November preg- nancy of a Muskrat in eastern Canada. 1979 Literature Cited Beer, J. R. 1950. The reproductive cycle of the Muskrat in Wisconsin. Journal of Wildlife Management 14(2): 151-156. Dibblee, R. L. 1970. The reproduction and productivity of Muskrats on Prince Edward Island. Proceedings of the Canadian Society of Wildlife and Fisheries Biologists, Atlantic Chapter, Fredericton, November 4-6. pp. 114— 122. Dilworth, T. G. 1967. The life history and ecology of the Muskrat under severe water level fluctuations. Proceed- ings of the Canadian Society of Wildlife and Fisheries Biologists, Atlantic Chapter, Charlottetown, November 28-29. 17 pp. Dixon, J. 1922. Rodents and reclamation in the Imperial Valley. Journal of Mammalogy 3(3): 136-146. Errington, P. L. 1963. Muskrat populations. Iowa State University Press, Ames. 665 pp. Forbes, T.R. 1942. The period of gonadal activity in the Maryland Muskrat. Science 95: 382-383. NOTES 44] Lay, D. W. 1945. Muskrat investigations in Texas. Journal of Wildlife Management 6(1): 56-76. McLeod, J. A. and G. F. Bondar. 1952. Studies on the biology of the Muskrat in Manitoba. Part I. Oestrous cycle and breeding season. Canadian Journal of Zoology 30(4): 243-253. Schacher, W. H. and M.R. Pelton. 1975. Productivity in Muskrats in East Tennessee. Proceedings of the 29th Southeast Game and Fish Commission, St. Louis, Mis- souri, October 12-15. 26 pp. Smith, H.R. and P.A. Jordan. 1976. An unexploited population of Muskrats with unusual biomass, produc- tivity, and body size. State Geological and Natural History Survey of Connecticut, Report of Investigations Number 7. 16 pp. Svihla, A. and R. D. Svihla. 1931. The Louisiana Muskrat. Journal of Mammalogy 12(1): 12-28. Received 11 January 1979 Accepted 2 May 1979 Lynx Movements and Habitat Use in Montana!? GARY M. KOEHLER, MAURICE G. HORNOCKER, and HOWARD S. HASH Idaho Cooperative Wildlife Research Unit, University of Idaho, Moscow, Idaho 83843 Koehler, Gary M., Maurice G. Hornocker, and Howard S. Hash. 1979. Lynx movements and habitat use in Montana. Canadian Field-Naturalist 93(4): 441-442. Movements of two Lynx (Lynx canadensis) were monitored by radio telemetry: an adult male for 74 mo from March to October and an adult female during January. Home range area for the male, determined from 21 radio locations, was 36 km2. Most locations were in densely stocked stands of Lodgepole Pine (Pinus contorta) resulting from the 1910 fires. Snowshoe Hares (Lepus americanus), their principle prey, were also most abundant in these stands. Key Words: habitat, Lynx (Linx canadensis), Snowshoe Hare (Lepus americanus), activity patterns. Little is known about seasonal habitat use or home range size of Lynx (Lynx canadensis). Most previous studies used snow-tracking techniques (Saunders 19632) Nelhis et al. 1972; Brand) et al. 11976): Berrie (1973) used radio telemetry to determine home range size in Alaska. In conjunction with a Wolverine (Gu/o gulo) study in northwest Montana, we monitored movements of two Lynx seasonally during 1977 by radio telemetry. Methods Twenty live-traps were set from January through ‘Contribution of the Idaho Cooperative Wildlife Research Unit; the United States Fish and Wildlife Service, the Idaho Department of Fish and Game, the University of Idaho, and the Wildlife Management Institute cooperating. *University of Idaho College of Forestry, Wildlife and Range Sciences Publication No. 168. April along 40 km of Forest Service trail. Traps, measuring 40 X 40 X 65 cm, were constructed of tubu- lar steel frames, chain link fence sides, and sliding steel _ doors. Lynx were immobilized with ketamine hydro- chloride (“Ketalar,” Parke Davis) in dosages approxi- mating 21 mg/kg body weight. Lynx were eartagged and tattooed in the lip and on the body under the foreleg, measured, evaluated for general physical condition, and fitted with collars containing radio transmitters. Radio signals were monitored from fixed-wing aircraft and the ground. The relative abundance of Snowshoe Hares ( Lepus americanus) in various forest types was obtained by counting the number of tracks crossed per kilometre of trail after fresh snowfall. To help differentiate tracks only those traveling to the west of the trail and spaced 3-m apart were recorded. A vegetative descrip- tion of each section of trapline included dominant overstory species, age class, and relative density. 442 THE CANADIAN FIELD-NATURALIST Vol. 93 TABLE 1—Relative abundance of Snowshoe Hare in four vegetative cover types along trapline during 14 d of survey , Tracks Distance Stand age, Stand surveyed Total ty Vegetative cover yr density (km) no. No. km -d Lodgepole Pine, pure stands < 80 Dense 9.0 Sil 7/ 2.54 Grassland islands in dense Lodgepole Pine < 80 Dense 37) 96 Daisy Subalpine Fir-Englemann Mature Medium 2.4 15 0.45 Spruce > 100 Islands of Lodgepole Pine, Douglas Fir, Ponderosa Pine (Pinus ponderosa) in Mature Sparse 5.6 11 0.14 > 100 grasslands Results and Discussion One adult male (weight 10 kg), one adult female (7 kg), and one juvenile female (4 kg) were captured. The juvenile accompanied the adult female and was not radio-collared because it was too small. The adult female was captured on 14 January and located 8 times prior to being found dead on 31 January. Death was believed due to predation by a Mountain Lion (Felis concolor). The male was captured on 10 March and located 21 times by 27 October. Most locations for the male and female were in young densely stocked stands of Lodgepole Pine (Pinus contorta). Twenty-six of the 29 locations (90%) occurred in timbered areas burned in 1910 and the remainder occurred in mature Douglas Fir ( Pseudo- tsuga menziesii) — Western Larch (Larix occidentalis) stringers along stream bottom within the 1910 burn. Of the locations burned in 1910, 23 (88%) occurred on xeric sites where Lodgepole Pine was dominant and 3 (t2%) on mesic sites where Subalpine Fir (Abies lasiocarpa) and Englemann Spruce ( Picea engleman- nii) were dominant. No locations occurred in open grassland or semi-open areas, 90% were in densely stocked stands, and 10% in medium-stocked stands. The estimated home range size for the male was 36 km?. This figure is similar to that found in other studies. Winter tracking in Newfoundland showed a home range size between 15.5 and 20.7 km? (Saunders 1963). In Alberta they were from 11.1 to 49.5 km? (Brand et al. 1976). In Alaska, Berrie (1973) found Lynx to range from 12.8 to 25.5 km2. Snowshoe Hares were most abundant in densely stocked stands of Lodgepole Pine (Table 1). Brand et al. (1976) and Adams (1959) found that hares were most abundant in dense stands. Winter ground tracking during this study and in Alberta (Brand et al. 1976) indicate that Lynx concentrate hunting activity within areas of high hare activity because hares are their main food (Brand et al. 1976) and 90% of Lynx radio locations were in these stands. Saunders (1963) found that Lynx activity and the location of the home range boundary coincided almost exactly with a tract of 10- to 20-yr-old growth timber. Our data suggest that Lynx concentrate activity in areas of high Snowshoe Hare activity, particularly in young dense stands of Lodgepole Pine. Radio tele- metry indicates that there is no change in range areas or habitat use throughout the seasons. Acknowledgments The United States Forest Service provided field facilities. The cooperation of D. Minister, D. Owen, and K. Granrud of the United States Forest Service is sincerely appreciated. We express special thanks to T. W. Koehler for assisting with the field work. Financial support was provided by the National Science Foundation, United States Forest Service, National Geographic Society, New York Zoological Society, National Wildlife Federation, Audubon Society, National Rifle Association, Wildlife Man- agement Institute, Montana Department of Fish and Game, and the Boone and Crockett Club. Literature Cited Adams, L. 1959. An analysis of a population of Snowshoe Hares in northwestern Montana. Ecological Monographs 29(2): 141-170. Berrie, P. M. 1973. Ecology and status of the Lynx in interior Alaska. Jn The world’s cats. Edited by R.L. Eaton. Volume |. Proceedings of the International Sym- posium on the World’s Cats, March 1971. Published by World Wildlife Safari, Winston, Oregon. pp. 4-41. Brand, C. J., L. B. Keith, and C. A. Fischer. 1976. Lynx response to changing Snowshoe Hare densities in central Alberta. Journal of Wildlife Management 40(3): 416- 428. Nellis, C. H., S. P. Wetmore, and L. B. Keith. 1972. Lynx-— prey interactions in central Alberta. Journal of Wildlife Management 36(2): 320-329. Saunders, J. J., Jr. 1963. Movements and activities of the Lynx in Newfoundland. Journal of Wildlife Management 27(3): 390-400. Received 28 February 1979 Accepted 30 May 1979 1979 NOTES Decline of Summering Bald Eagles in Central New Brunswick R. F. STOCEK The Maritime Forest Ranger School, Fredericton, New Brunswick E3B 4X6 StocekeuRe F: 443-445. 1979. Decline of summering Bald Eagles in central New Brunswick. Canadian Field-Naturalist 93(4): The numbers of Bald Eagles ( Haliaeetus leucocephalus) that summer in central New Brunswick have decreased considerably. This reflects chiefly a decline in the proportion of immature birds. Key Words: Bald Eagle, Haliaeetus leucocephalus, New Brunswick, abundance, age composition. The Bald Eagle ( Haliaeetus leucocephalus) in New Brunswick is listed as endangered under the provincial Endangered Species Act of 1976. It was considered an uncommon resident and transient by Squires (1976), who reported that numbers of the bird had decreased drastically; however, there do not appear to be any published accounts giving details of that decline. This paper describes the decrease in numbers and change in age composition of summering Bald Eagles in the lower St. John River basin and other parts of central New Brunswick (between 45°30’N and 47°00’N). The 104-km?2 French Lake district, west of Grand Lake, Queens County, and centered at 45°50’N, 66°17’W, is of particular interest here (Figure 1). It includes the shores of French, Indian, and Maquapit Lakes, the thoroughfare joining them, and Loders and Porto- bello Creeks in Sunbury County. Although distributed over the entire province in the past, the resident Bald Eagle, H. /. alascanus, was never considered numerous (Moore 1928). More recently, banding returns suggested that most sum- mering eagles belonged to the southern subspecies, H. |. leucocephalus (Squires 1952). Those birds, mostly immatures coming from Florida (Broley 1947), wandered into the Maritime region in April and May, and returned southward in August and September (Wright 1953). The wintering birds of the resident form apparently move inland from coastal areas in February and March (R. F. Stocek and P.A. Pearce, unpublished data), and return to the coast late in the fall (R. F. Stocek, unpublished data). Methods Bald Eagle records on file in the Northeastern Wildlife Station at the University of New Brunswick and at the New Brunswick Museum were compiled, with observations from the various naturalist clubs and provincial and federal resource agencies. Rela- tively few quantitative data were available prior to 1959 except for the French Lake district records of the Northeastern Wildlife Station, extending from 1949 to 1962. The French Lake sight records are treated FiGuRE |. Map of New Brunswick showing the major areas mentioned in the text. The French Lake district, shown in dark shading, lies next to Grand Lake in Queens County. The lower St. John River extends downstream from Fredericton (the dark circle) and is shown enclosed by the solid outlines of the four basin counties. The central part of the province is defined by the marginal lines of latitude, 45°30’N and 47°00'N. separately, rather than being combined with the scattered data from the rest of the province, because so much more effort was spent on this relatively small area. Numerous low-level aerial flights (both fixed- wing and helicopter) were made over the lower St. John River basin during the spring and summer of 1974 and 1975. As used here, winter refers to December, January, and February; spring to March, April, and May; summer to June, July, and August, and fall to September, October, and November. 444 Results and Discussion Cumulative sight records between 1959 and 1975 show that central New Brunswick accounted for 55% of the total spring observations and 60% of the summer observations of Bald Eagles in the province. Typically most of the eagles in this region were seen in the summer (38% of the area total) and spring (30%), and fewer in the winter and fall (22% and 10%, respectively). Thirty years ago the basin of the lower St. John River was thought to be one of the most important summer habitats of H. 1. lewcocephalus in the north- east maritime region of the continent. The peak summer population of Bald Eagles in the French Lake district was estimated at 54 in 1949 and 45 in 1950 (Wright 1953). The Foshay Lake - Grimross Neck area 20 km to the southeast appeared to support a similar number in 1949. During those years a United States Fish and Wildlife Service aerial survey team saw more eagles in this area than in all the rest of the maritime provinces. Practically all the eagles counted in New Brunswick were seen here, with as many as 18 in the air at one time near French Lake. A total population of at least 100 eagles summering in the lower St. John River basin was suggested (B.S. Wright, unpublished data). Large numbers of sum- mering eagles had also been reported there in May and June, 1937 to 1944, especially along Portobello Creek and from Fredericton to Jemseg (H. S. Peters, un- published data). During the 1930s and 1940s, it was not uncommon to see 20-30 eagles in a day on the waterfowl breeding grounds in central New Bruns- wick (Squires 1976). The mean numbers of summering eagles seen in the French Lake district declined during the 1950s and early 1960s. According to data from the North- eastern Wildlife Station, 2.3 birds were seen per successful day* during June-July, 1953-1954. By 1961-1962 this figure was down to 1.3; eagles then were being seen only infrequently by station per- sonnel. B.S. Wright (personal communication), who had studied waterfowl in the St. John swamps and marshes since 1945, believed that there were no more than 10 eagles on the entire area in the summer of 1964. On | August 1974, I surveyed by helicopter 125 km of shoreline in the French Lake district and another 179 km of adjacent Grand Lake and Jemseg River; no Bald Eagles were seen. A_ provincial naturalist conducting a group canoe trip through the district and the nearby Oromocto River drainage reported that only one eagle was seen during the 192-km trip in mid-July 1974. The only eagles I saw *A day on which at least one eagle was sighted; unfor- tunately statistics on total effort (days) are not available. THE CANADIAN FIELD-NATURALIST Vol. 93 during the many flights I made through the French Lake area in 1974 and 1975 were the one pair nesting there. The decrease of Bald Eagles in certain parts of North America has been accompanied by a decline in the proportion of immature birds in those populations (Sprunt 1969). The percentage of immatures in the French Lake district summering population for 1953 to 1962 is shown in Table |. The influx of wandering birds probably accounted for the high May values, while birds leaving the area in August reduced the number seen. B.S. Wright (unpublished data), re- ported a high 1:4 adult to immature ratio for the district during the spring-summer period in 1949. TABLE 1—The monthly distribution of immature Bald Eagles in the French Lake district of New Brunswick from 1953 to 1962, expressed as a percent of the total known-age birds seen Month Q% Immature Total no. seen May 44 4] June 23} 73 July 2) 70 August 10 19 By 1953-1962 the ratio had changed to 4:1. Values of 30% immatures recorded in 1953-1954 and 13% in 1961-1962 suggested a decline during that period. Although data are fragmentary for the intervening years, June and July sightings provide some insight. From 1953 to 1956, 25% of the 79 eagles seen in the district were immatures. Station personnel reported 19% immature birds (of 64 seen) from 1959 to 1962. New Brunswick sight records show that, in the province as a whole, but excluding data from the French Lake area, the proportion of immature eagles seen In spring and. summer decreased from an average of 32% during 1967-1973 to 22% in 1974-1975 (Table 2). These comparative seasonal values very likely reflect a real difference between the two time periods, because the increased effort in 1974-1975 TABLE 2— The seasonal distribution of immature Bald Eagles in New Brunswick from 1967 to 1973 and 1974-1975, expressed as a percent of the total known-age birds seen (in parentheses). French Lake district statistics are not included here % Immature, % Immature, Season 1967-1973 1974-1975 Winter 20 (56) 20 (35) Spring 25 (76) 14 (50) Summer 37 (104) 27 (93) Fall 22 (49) 20 (25) 1979 would have been expected to reveal a greater rather than a smaller proportion of the less conspicuous immature birds. The decline in the summering population of Bald Eagles in central New Brunswick and probably throughout the province was to be expected. The decrease in Bald Eagle reproduction in other regions of North America during the last two decades is also well documented (e.g., Sprunt 1969; Sprunt et al. 1973). Both Broley (1950, 1958) and Howell (1958) reported increasing nest failures in Florida, which was an important source of birds summering in the Maritimes. Acknowledgments I am grateful to the late Bruce Wright, to Peter Pearce, and John Baird for their helpful suggestions and discussions. The interest of many individuals and agencies, and particularly the New Brunswick Department of Natural Resources in contributing valuable data, is greatly appreciated. The Canadian Wildlife Service provided financial support for this work. Literature Cited Broley, C. L. 1947. Migration and nesting of Florida Bald Eagles. Wilson Bulletin 59: 3-20. NOTES 445 Broley, C.L. 1950. The plight of the Florida Bald Eagle. Audubon Magazine 52: 42-49. Broley, C. L. 1958 The plight of the American Bald Eagle. Audubon Magazine 60: 162-163, 171. Howell, J.C. 1958. Further history on some Bald Eagle nest sites in east-central Florida. Auk 75: 96-98. Moore, W.H. 1928. A list of the birds of New Brunswick, Canada. 67th Annual Report, Department of Lands and Mines 1927: 91-112. Sprunt, A., IV. 1969. Population trends of the Bald Eagle in North America. /n Peregrine Falcon populations: their biology and decline. Edited by Joseph J. Hickey. University of Wisconsin Press, Madison. pp. 347-351. Sprunt, A., 1V, W. B. Robertson, Jr., S. Postupalsky, R. J. Hensel, C. E. Knoder, and F. J. Ligas. 1973. Compara- tive productivity of six Bald Eagle populations. Trans- actions of the North American Wildlife and Natural Resources Conference 38: 96-105. Squires, W. A. 1952. The birds of New Brunswick. New Brunswick Museum Monographic Series Number 4. 164 pp. Squires, W. A. 1976. The birds of New Brunswick. New Brunswick Museum Monographic Series Number 7. 210 pp. Wright, B.S. 1953. The relation of Bald Eagles to breeding ducks in New Brunswick. Journal of Wildlife Manage- ment li 55—02. Received 30 October 1978 Accepted 10 May 1979 Lesser Black-backed Gull, Larus fuscus, in Labrador Waters KEVIN D. POWERS Manomet Bird Observatory, Manomet, Massachusetts 02345 USA Powers, Kevin D. 1979. Lesser Black-backed Gull, Larus fuscus, in Labrador waters. Canadian Field-Naturalist 93(4): 445-446. An adult Lesser Black-backed Gull, Larus fuscus, sighted 140 km E of Nain, Labrador, adds to evidence that the species occurs in small numbers through eastern North American waters. Key Words: Lesser Black-backed Gull, Larus fuscus, Labrador. On 21 July 1978 I observed an adult Lesser Black- backed Gull (Larus fuscus) from the CCGS Narwhal at 57°11’N, 59°20’W approximately 140 km ENE of Nain in the Labrador Sea. Sightings of Lesser Black- backed Gulls include Greenland: Godthadb (ca. 64°30’N) and Godhavn (69°20’W) (Salomonsen 1967); at sea 100 to 675 km SSE of Cape Farewell, Greenland (ca. 54°30’N to 60°00’N) (Brown 1968): Northwest Territories (cf., American Birds 1978, 32: 1186; Alsop and Jones 1973); Churchill, Manitoba (Ross and Cooke 1969): Grand Banks (Brown et al. 1975); Nova Scotia (cf., Nova Scotia Bird Society Newsletter 1977, 19: 100; E. L. Mills, unpublished data); and northeastern United States (W.R. Petersen, unpublished manuscript; Bull 1964; K. D. Powers, unpublished data). The gull approached the ship from astern at 16:40 EST and “ship followed” for 55 min before departing. It did not appear to be associated with any other species in the area. The winds were WNW at 25 knots, and seas were 1-2 m. The skies were overcast with no precipitation. The bird was observed with 8 X 40 binoculars in good light within 50 m of the ship. All distinguishing field marks of the species were seen, 446 including the bright yellow legs and feet. The mantle was slate-gray becoming black in the outermost primaries, suggesting L. f. graellsii. A color photo- graph was obtained showing the mantle color pattern of the bird. The sighting is the first record of Lesser Black- backed Gull for Labrador waters, and it adds to evidence of increasing regularity off eastern North America of the subspecies L. f. graellsii, which breeds in Iceland, Faeroe Islands, the British Isles, and Brittany (Witherby et al. 1941). The new record is only about 2000 km distant from Iceland, well within the range of sightings along American coasts from Davis Strait to Cape Hatteras. I express my appreciation to R.G. B. Brown, W.R. Petersen, and T. Lloyd-Evans who helped to improve this note. R. G. B. Brown of the Canadian Wildlife Service arranged my passage aboard the CCGS Narwhal, and traveling expenses. I am also grateful to the United States Department of Energy (DOE Contract No. EE-78-S-02-4706) for support. Occurrences of the Red Phalarope Adjacent States! J. PAUL GOOSSEN2 and DANIEL G. BUSBY3 THE CANADIAN FIELD-NATURALIST Vol. 93 Literature Cited Alsop, J. F. and E.T. Jones. 1973. The Lesser Black- backed Gull in the Canadian Arctic. Canadian Field- Naturalist 87: 61-62. Brown, R.G.B. 1968. Sea birds in Newfoundland and Greenland waters, April-May 1966. Canadian Field- Naturalist 82: 88-102. Brown, R. G. B., D. N. Nettleship, P. Germain, C. E. Tull, and T. Davis. 1975. Atlas of eastern Canadian seabirds. Canadian Wildlife Service, Information Canada, Ottawa. 220 pp. Bull, J. 1964. Birds of the New York area. Harper & Row, New York. Ross, R. K.and F. Cooke. 1969. Lesser Black-backed Gull at Churchill, Manitoba. A new bird for Canada. Canadian Field-Naturalist 83: 399. Salomonsen, F. 1967. Fuglene pa grénland. Copenhagen. [Bo QUT. Witherby, H. F., F. C. R. Jourdain, N. F. Ticehurst, and B. W. Tucker. 1941. The handbook of British birds. Volume V. H. F. & G. Witherby Ltd., London. 356 pp. Received 14 February 1979 Accepted 25 April 1979 in the Prairie Provinces and Regent College, 2130 Wesbrook Mall, Vancouver, B.C. V6T 1W6 3Canadian Wildlife Service, P.O. Box 400, Fredericton, New Brunswick E3B 4Z9 Goossen, J. Paul and Daniel G. Busby. 1979. Occurrences of the Red Phalarope in the prairie provinces and adjacent states. Canadian Field-Naturalist 93(4): 446-449. Twenty-six records of Red Phalaropes ( Phalaropus fulicarius) were obtained for the prairie provinces and adjacent states. It is suggested that the predominance of fall records in western Alberta and Montana is due to the guiding influence of the Rocky Mountains for southbound and southwestbound birds, and that the scattered spring records result from the lack of such an influence for northbound birds. It remains possible that the initial displacement from the normal migration route is caused by adverse weather conditions. Key Words: Red Phalarope, Phalaropus fulicarius, extralimital sightings. Our interest in the Red Phalarope (Phalaropus fulicarius) was aroused when we observed a female in alternate plumage in the Assiniboine River Diversion about 3km south of Lake Manitoba and 5 km west of Delta, Manitoba (50°11’N, 98°19’W). This was the third sighting in southern Manitoba; the species is considered a rare spring migrant in the Churchill region (Jehl and Smith 1970). The Red Phalarope breeds along the coasts and 'This is paper number 74 of the University of Manitoba Field Station (Delta Marsh). islands of northern Asia, Europe, and North America. It winters primarily in the southern hemisphere off the coasts of South America and western Africa (Godfrey 1966). In North America, it migrates along both the Atlantic and Pacific coasts (American Ornithologists’ Union 1957; Godfrey 1966). This note summarizes and discusses Red Phalarope records from the Canadian provinces of Manitoba, Saskatchewan, and Alberta, and from the adjacent states of Montana, North Dakota, and Minnesota. Twenty-six records of the Red Phalarope in the area under consideration are summarized in Table 1. 1979 NOTES 447 TABLE I—Red Phalarope sightings for the prairie provinces and adjacent north-central states Location Date No. Source Manitoba East Shoal Lake 12 October 1963 | K. Gardner Delta Marsh 19 June 1969 | R. E. Jones Delta Marsh 30 June 1975 | This paper Saskatchewan Long Lake! July 1879 I Macoun and Macoun (1909) Old Wives Lake? May 1895 l Macoun and Macoun (1909) Sandfly Lake 11 June 1914 I Mitchell (1924) Proctors Lake 21 May 1946 I| 22 Mowat (1946) Alberta Didsbury 3 September 1903 1 Salt and Wilk (1966) Beaverhills Lake September 1925 I Salt and Wilk (1966) North Saskatchewan River near Graveyards Cabin 23 May 1953 4? Banfield (1954) Pigeon Lake 13 July 1960 IPS Salt and Wilk (1966) Carseland Dam? 23 October 1966 | Smith and Klauke (1967) Cochrane Lake} 30 October 1966 | Smith and Klauke (1967) Big Lake 11 November 1975 I Ebel (1976) Stirling _ — Godfrey (1966) Montana Bowdoin National Wildlife Refuge July 1953 | P. D. Skaar Harrison Lake 26 August 1959 | Rogers (1960) Harrison Lake 3 November 1963 ] Rogers (1964) Harrison Lake 11 October 1970 l Rogers (1971) Freezeout Lake 17 August 1976 12 Serr (1977) North Dakota Cando4 Slade National Wildlife Refuge Minnesota Knife River Lake Mille Lacs Lake Mille Lacs Lake Moorhead 23 May 1890 25 July 1963 17 November 1963 29 October 1976 19 November 1977 27 May 1977 'Now called Last Mountain Lake. 2Now called Johnstone Lake. Stewart (1971) Stewart (1971) Hofslund (1964) Savaloja (1977) Harding (1978) Wachtler and Wachtler (1977): Anderson (1977) 3Validity of sighting questioned by Sadler and Myres (1976). 4Considered as hypothetical by Stewart (1971). seven are of females; the others were not sexed. We grouped the 25 observations with dates into three periods: spring (April-June), midsummer (July), and fall (August-November), and plotted them (Figure 1). Two patterns are noticeable. First, seven of the eight spring records and three of the four midsummer records are scattered east of 110° W. Second, 9 of the 13 fall records are from the extreme western part of the area, on or near the eastern slope of the Rocky Mountains. No correlation with weather could be established for the Canadian sightings of Red Phalaropes and P. D. Skaar (personal communication) noticed no unusual, weather conditions associated with the Montana sightings. The spring record for Minnesota, however, was preceded by periods of thunderstorms and unsettled weather with 35 to 40-km south winds. The known eastern spring migration route follows the Atlantic Ocean to the arctic breeding grounds. Observations of Red Phalarope in southern Ontario (W. E. Godfrey, personal communication), however, suggest that some of these birds may pass through the Great Lakes and Hudson Bay, using a similar short- cut to that suggested for Arctic Terns (Sterna paradisaea) (Godfrey 1973) and Sabine’s Gull (Xema sabini) (Lambert 1973). The scattered spring migrants reported on the Great Plains may be phalaropes attempting to migrate overland. Oldsquaw (Clangula hyemalis), also a northern breeder, regularly pass through southern Manitoba in both spring and fall (Sexton and Collins 1977), and the possibility of an overland migration route for another arctic breeder, 448 FIGURE |. Red Phalarope sightings for the prairie provinces and adjacent north-central states. Open circles, spring records: half-filled circles, midsummer records: closed circles, fall records. the Long-tailed Jaeger (Stercorarius longicaudus), has also been suggested (Jehl and Smith 1970). The numerous fall records in Alberta and Montana suggest that some migrants proceed southward or southwestward until they encounter the Rocky Mountains. With reference to the Montana observa- tions, P. D. Skaar suggested that a prolonged Pacific air flow might have deterred the birds from crossing the mountain range (Rogers 1964), which thus served as a leading line. No such guiding influence would be available for northbound birds, which are much more scattered. The fall sightings from Manitoba and Minnesota may be migrants from the Hudson Bay — Great Lakes route. Two of the fall birds were listed as females in Table 1; however, we suggest that generally these are young of the year having no previous migratory experience. Of 18 casual records presented by Bent (1927) for interior Canada and United States, 16 were fall records and one was not dated. This further suggests that dispersal of the young may account for many of the interior records. It seems possible that adverse weather conditions are the initial displacement factor for these extralimital sightings, but that after dis- placement the birds attempt the usual southward or northward orientation, which leads some of them overland. Despite the periodic, infrequent sightings of Red Phalaropes over the prairie provinces and adjacent states, such records constitute an extremely small fraction of the total migration of the species. It is unlikely that enough of these birds survive for THE CANADIAN FIELD-NATURALIST Vol. 93 selection in favor of a regular inland migration to occur. Acknowledgments We thank those individuals who replied to our request for information regarding the Red Phalarope. These include H. W. R. Copland, D. R. Crooke, J. B. Falls, J. Ferrand, Jr., M. B. Fitzgerald, N. L. Ford, K. Gardner, W. E. Godfrey, J. C. Green, E. B. Hazard, J. Hines, P. B. Hofslund, E. O. Héhn,C. S. Houston, R. D. James, R.E. Jones, F. McKanney. MEK: McNicholl, W. J. Maher, D. Maurer, M. T. Myres, R. A:-Paynter, P. D» Skaar, R: E-StewantsRewe Storer, D. V. Weseloh, P. L. Wright, and R. L. Zusi. We thank W. R. Salt and S. G. Sealy for reading an earlier draft of the manuscript and also the reviewers, J.B. Gollop and R.G.B. Brown, for their comments. Literature Cited American Ornithologists’ Union. 1957. Check-list of North American birds. Fifth edition. Port City Press Inc., Baltimore, Maryland. 691 pp. Anderson, E. 1977. Red Phalarope at Moorhead-Loon 49: 1973. Banfield, A. W.F. 1954. Further notes on the birds of Banff National Park, Alberta. Canadian Field-Naturalist 68: 182. Bent, A. C. 1927. Life histories of North American shore- birds. Part 1. United States National Museum Bulletin 142. Ebel, G.R. A. 1976. Sightings of a Red Phalarope near St. Albert, Alberta. Alberta Naturalist 6: 2. Godfrey, W.E. 1966. The birds of Canada. National Museum of Canada Bulletin 203: 1-428. Godfrey, W. E. 1973. A possible short-cut spring migra- tion route of the Arctic Tern to James Bay. Canada. Canadian Field-Naturalist 87: 51-52. Harding, H. 1978. Red Phalarope—Mille Lacs Lake. Loon 50: 45-46. Hofslund, P. B. 1964. The Red Phalarope in Minnesota. Loon 36: 25. Jehl, J. R. and B. A. Smith. 1970. Birds of the Churchill region, Manitoba. Manitoba Museum of Manand Nature Special Publication |. 87 pp. Lambert, K. 1973. The migration of Sabine’s Gulls. Xema sabini, in the Northwest Atlantic. Canadian Field- Naturalist 87: 57-60. Macoun, J. and J. Macoun. 1909. Catalogue of Canadian birds. Canada Department of Mines, Geological Surveys Branch, Ottawa. 761 pp. Mitchell, H. H. 1924. Birds of Saskatchewan. Canadian Field-Naturalist 38: 101-118. Mowat, F. 1946. Bird watching from a “jeep.” Blue Jay 4: 39. Rogers, T. 1960. Northern Rocky Mountain — Intermoun- tain Region. Audubon Field Notes 14: 56-58. Rogers, T. H. 1964. Northern Rocky Mountain Region. Audubon Field Notes 18: 57-60. Rogers, T. H. 1971. Northern Rocky Mountain — Inter- mountain Region. American Birds 25: 80-84. 1979 Sadler, T.S. and M. T. Myres. 1976. Alberta birds 1961-— 1970 with particular reference to migration. Provincial Museum of Alberta Natural History Occasional Paper No. 1. Salt, W.R. and A. L. Wilk. 1966. The birds of Alberta. Second edition. Queens Printer, Edmonton, Alberta. 511 pp. Savaloja, T. 1977. Minnesota’s second Red Phalarope. Loon 49: 44-45. Serr, E. M. 1977. Northern Great Plains. American Birds 31: 190-194. NOTES 449 Sexton, D. A. and K.M. Collins. 1977. Records of the Oldsquaw in southern Manitoba. Blue Jay 35: 96-99. Smith, W. and R. Klauke. 1967. A sight record of the Red Phalarope in Alberta. Blue Jay 25: 25-26. Stewart, R. E. 1971. Check list of birds of North Dakota. Prairie Naturalist 3: 3-12. Wachtler, D. and G. Wachtler. 1977. Red Phalarope at Moorhead. Loon 49: 172. Received 6 April 1978 Accepted 24 May 1979 Nesting of the Calliope Hummingbird in Kananaskis Provincial Park, Alberta DANIEL F. BRUNTON,! SIDNEY ANDREWS.,2 and DAVID G. PATON3 ‘Southwick Drive, R.R. 3, Manotick, Ontario KOA 2N0 2General Delivery, Holland Landing, Ontario LOG 1HO 31481 Jalna Avenue, Mississauga, Ontario LSJ 1S6 Brunton, D. F.. S. Andrews, and D.G. Paton. 1979. Nesting of the Calliope Hummingbird in Kananaskis Provincial Park, Alberta. Canadian Field-Naturalist 93(4): 449-451. A nest of Calliope Hummingbirds (Ste/lu/a calliope) was studied in Kananaskis Provincial Park, Alberta, in 1977 and 1978. Incubation period (16 d) and nestling period (18-21 d) were determined. The nest was constructed in precisely the same location in successive years and nest construction was completed during incubation. These unusual nesting characteristics may be adaptations to severe environmental constraints at high elevations near the northern limit of the species’ range. Key Words: Calliope Hummingbird, Stellula calliope, incubation period, nestling period, nest construction. nest site selection, Alberta. A nest of the Calliope Hummingbird (Sre//ula calliope) was discovered in Kananaskis Provincial Park, Alberta, in mid-June 1977. This bird is found only locally in Alberta (Salt and Salt 1976). A second nesting was discovered at the same site in 1978. The following discussion deals with the results of our observations for both years. Observations The nest was situated on a horizontal limb of a Lodgepole Pine (Pinus contorta) at the base of a steep shale creek bank. It was | km south of the “Church Camp” on the east bank of Pocaterra Creek (50°41’N, 115°5’W) at an elevation of 1660 m. The nest was a tiny cup constructed of lichen fragments and fine plant materials. The 1978 nest was constructed on the weathered remains of the 1977 nest: beneath the latter were the remains of one or more previous nests (Figure |). In 1978 the nest was first observed on 10 June as the first egg was being laid. The nest at that time was a shallow cup with poorly defined walls and an unlined bottom. At 11:45 the bird was observed frequently poking and prodding the edges of the nest and pulling material in towards the center. For about 2 min it performed an action we described as “running on the spot” — ap- parently deepening the cup in preparation for egg- laying. The bird subsequently became very still on the nest for several minutes and then flew off. A single. pinkish-white translucent egg was subsequently found in the nest. We believe the egg was laid as we were watching the bird. On 13 June the nest contained one dull-white, opaque egg and one (freshly laid) pinkish- white, translucent egg. Although the first egg was laid on the bare floor of the nest, both were now nestled into a bed of feathers in a deep high-walled cup. The increased size and depth of the nest was quite striking. By 29 June the nest was larger yet, with substantially thickened wall. The two-egg clutch of the 1978 nest was completed on 13 June when we observed it at 16:20. On 29 June at 17:30 two tiny young were observed, one still wet and lying on fragments of the egg from which it had just hatched. The other young was dry. Incubation had taken 16d. The nestling period ran from 3 July to 21 July(18 d) in 1977 and 29 June to 20 July (21 d) in 1978. On 20 July 1978 we observed the departure of the young from the nest. One of the young was out of the nest but still in the nest tree when observations began. In approximately 20 min, the remaining nestling FiGure 1. Adult (female) Calliope Hummingbird on nest in Kananaskis Park. Alberta. Note remains of previous nest under active nest: the 1978 nest was situated on the remains of this one. Photographed on 23 June 1977 by D. G. Paton. moved to the lip of the nest, made its first flight, and ultimately moved 4m away. The female adult returned to the nest tree four times to feed or preen the young birds during the observation period. The adult (only the female was seen in both years) usually approached the nest from the creek side of the tree, perching near the top of the nest tree and then working down through it in three or four short flights. The bird used the same perches each time. Calliope Hummingbirds also exhibited a set pattern of man- oeuvers to approach a feeder at Bow Valley Pro- vincial Park, Alberta, and when disturbed the birds would flee to precisely the same perch in a nearby Lodgepole Pine (B. Romanyshyn, personal com- munication). During the early stages of incubation (up to approximately 10 to 12 d), the adult would leave the nest when observers approached within 10 m. During late incubation and when young were in the nest, however, the adult would allow the observers to approach within | m of the nest before flying off. THE CANADIAN FIELD-NATURALIST Viole 93 When it was flushed during this latter period, the bird would aggressively “buzz” the observers (usually positioned 3 to 5 m from the nest). The bird would often approach to within 30 to 50 cm of the observers during this aggressive behavior. Discussion There is no published literature on the nesting of Calliope Hummingbird in Canada with which we can compare our observations. Calder (1971) cites 21 to 23 d for the nestling period in Wyoming and this compares well to a figure of 21 d for a nest at Hood River, Oregon (on file with the North American Nest Record Scheme). Figure 2 indicates the timing of egg and young observations from cards on file with the British Columbia Nest Record Scheme. None of the British Columbia cards recorded a complete incubation or nestling period observation. By summing the total of days in which young or eggs were in these nests and placing these data in quarter-month periods, the nesting chronology of the Calliope Hummingbird in British Columbia is illustrated. The total nesting period is roughly 37 d. The difference between starts of young and egg peaks is about 15 d. The dates for Kananaskis Park in 1977 and 1978 agree well with the peak of observations of eggs and young in nests in British Columbia. The sample of British Columbia records was too small to subdivide by latitude and/or elevation. When sufficient numbers of observers are filed, this would certainly be worthwhile. The total length of the nesting period for Calliope Hummingbird is approximately 34 to 38 d in north- western North America (Calder 1971: this study). There is some suggestion that the period is shorter at the northern limit of the species range (34-37 d in Kananaskis vs. 36-38 d in Wyoming). We strongly suspect that the same female was responsible for the 1977 and 1978 nestings in Kananaskis Park. As the bird was not banded, we cannot be certain of this. In any case, the use of precisely the same site in successive years 1s apparently atypical in hummingbirds. W. Ray Salt (personal communication) observed this practice with Rufous Hummingbirds (Selasphorus rufus) in the 1960s in British Columbia, and has a nest collected by J. Grant of Vernon, British Columbia, which was used in 1938 and again in 1939. Pearson (1953) notes repeated use of nests by the Estella Hummingbird ( Oreotrochilus estella) in high mountain caves in the Peruvian Andes. He indicates, however, that these nests are “recon- ditioned” each year: the Calliope Hummingbird nests we observed were completely rebuilt upon the previous year’s remains. We found no published reports of nests being completed during the period of egg laying, although Skutch (1951) does report 1979 451 Nestin NOTES 25 20 3 ¥ 8 een 6 : — O: \ ° =! 5 i | 6 2 : \ £ 10 HE + 2 2; ; : \ 5 \ ° \ \ e OP Pcccecce, June ( quarter July months ) FIGURE 2. Frequency of occurrence of eggs and young in nests of the Calliope Hummingbird in British Columbia. renewal of some nesting material during incubation by some tropical hummingbirds. The utilization of the same nest site for a succession of years, completion of nest construction during incubation, and the possibly shorter nesting period of the species in this area may be adaptations to near- marginal breeding conditions in Alberta at the northern limit of its range. Acknowledgments Field assistance and additional data were provided by C. Backer, A. Masters, and B. Romanyshyn. H. W. R. Copland and J. B. Gollop provided nesting data from the Prairie Nest Record Scheme, as did R. W. Campbell from the British Columbia Nest Record Scheme and J. Crump from the North American Nest Record Card Program. R. D. Strick- land contributed an extensive collection of humming- bird literature. W. R. Salt provided unpublished data. and observations from earlier studies. Several earlier drafts of the manuscript were typed by B. Wackerle. Our thanks to all these people for their valuable assistance. Literature Cited Calder, W. A. 1971. Temperature relationships and nesting of the Calliope Hummingbird. Condor 73: 314-321. Pearson, O. P. 1953. Use of caves by hummingbirds and other species at high altitudes in Peru. Condor 55: 17-20. Salt, W.R. and J. R. Salt. 1976. The birds of Alberta. Hurtig Press, Edmonton. Skutch, A. F. 1951. Life history of Longuemare’s Hermit Hummingbird. Ibis 93: 180-195. Received 24 February 1978 Accepted 3 April 1979 News and Comment FON Conservation Award for The Canadian Field-Naturalist A Federation of Ontario Naturalists (FON) Con- servation Award has been presented to The Ottawa Field-Naturalists’ Club in recognition of its out- standing contribution to the cause of conservation. The citation for the FON Award read as follows: “for the continued excellence of the journal The Canadian Field- Naturalist which has a well deserved national and international reputation and which provides a vital forum for reports on natural history and the environment by ama- teur and professional observers alike.” The FON Conservation Award was accepted by Dr. Lorraine C. Smith, Editor of The Canadian Field- Naturatist, at the Federation of Ontario Naturalists Conference at Toronto, Ontario on 5 May 1979. Dr. Peter A. Peach, President of the FON expressed the appreciation of all FON members for the continued excellence of the journal and personally congratulated the Editor. Mike Singleton, General Manager of the FON, added his congratulations and commendation to those of the Awards Committee, FON Directors, and participants at the annual conference and pointed out that the award in recognition of The Canadian Field- Naturalist was well earned. Moreover, he wrote: “It seems to me that we do not often enough recognize the great contribution the Ottawa club has made through this journal in facilitating natural history reports by amateur and professional alike, and most importantly in stimulating research by amateurs across the country.” He asked that the commendation and congratulations be drawn to the particular attention of all members of The Ottawa Field- Naturalists’ Club. ° Notice of The Ottawa Field-Naturalists’ Club Annual Business Meeting The 101st Annual Business Meeting of The Ottawa Field-Naturalists’ Club will be held in the auditorium of the National Museum of Natural Sciences, Metcalfe and MacLeod, on Tuesday, 15 January 1980, at 8:00 p.m. 452 Diana R. Laubitz, Recording Secretary Book Reviews ZOOLOGY Guide to the Study of Animal Populations By James T. Tanner. 1978. University of Tennessee Press, Knoxville. 186 pp. U.S. $8.95. Many of the mathematical and statistical tech- niques applied to the study of animal populations elude all but a specialist in the field. Therefore, I was somewhat elated to discover that this book is comprehensible to an average biologist having some experience in biometrics. The book contains ten chapters describing a variety of techniques for studying animal populations. Sub- jects such as Density and Dispersion, Mark-Re- capture Methods, Sex and Age Composition, Mortality and Survival, Reproduction, Migration and Recruitment are presented. Considerable discussion is devoted to mathematical relationships between population characteristics and methods of predicting population changes. As the title suggests, all examples given deal with animal populations and many deal with big game species. Several problems are worked out in each chapter and suggestions are given for the arrangement of the raw data. These examples do much to dispel the mystique of seemingly incomprehensible formulae. Symbols are used as consistently as_ possible throughout the text and a list is provided at the beginning of the book. The book is replete with references to more detailed discussions of various subjects and methodologies available in the literature. Moreover, the text is conveniently cross-referenced from chapter to chapter, enabling the reader to refresh his memory quickly from previous presentations on a topic. One of the most interesting chapters discusses mark-recapture methods for measuring population characteristics. This is a very practical presentation and provides valuable insight into research design compatible with statistical analysis of the data. The publication is intended for use by graduate students and field biologists. I am certain that it could be a very useful addition to any biological bookshelf. The straightforward presentation and numerous examples bolster the reader’s confidence that he can apply statistical methodologies and _ techniques competently and that studies of animal populations need not bea mathematical maze. The author’s candid critiques of techniques are valuable. Also, the list of literature citations is a useful tool for non-specialists wishing. to delve further into some aspect of population studies. DAN MURPHY Wildlife Service, Government of the Northwest Territories, Yellowknife XOE 1HO Quaternary Vertebrate Faunas of Canada and Alaska and Their Suggested Chronological Sequence By C. R. Harington. 1978. Syllogeus Number 15. National Museums of Canada, Ottawa. 105 pp., 15 illus. Available from Library of National Museum of Natural Science, Ottawa KIA OMS8. Most natural historians working with recent faunas are not especially familiar with the literature dealing with Pleistocene faunas and natural history. This is understandable, considering the present volume of literature dealing strictly with modern faunas. Any- one working with modern groups, however, can appreciate the historical perspective provided by the paleontological literature, especially that of the Pleistocene. Often this historical perspective provides insight into modern animal and plant distributions and geographic patterns as well as an understanding of changes in their geography in response to changes in ancient drainage patterns or patterns of glaciation. A good synoptic review serves two functions. First, it provides a broad generalized background on a subject. Secondly it introduces the nonspecialist to the literature dealing with that subject. With regard to the Pleistocene faunas of Canada, Harington’s paper successfully fulfills both objectives. The information is presented in a concise, straight- forward manner that makes data retrieval simple. The faunas are listed for each province and Alaska. Each section opens with a narrative concerning various fossil discoveries and their significance with regard to understanding Pleistocene events in the province. Both marine and terrestrial vertebrates are described. There is a good balance in the text between descriptions of the history of collecting, outstanding specimens that have been found, and the type of information that these finds provide. Following the 453 454 opening narrative of each section the Pleistocene faunas of specific provincial localities are listed. It is at this point that the paper becomes a reference work. Each faunal description begins with the fauna’s location. The locality information varies in detail from the very general such as “Hand Hills, Alberta near Delia,” to the very specific: “Acasta Lake, Northwest Territories (65°24’N, 115°31’W).” The locality description is followed by a faunal list which includes all known vertebrates from the site. This part alone will be of use to anyone interested in determin- ing the Pleistocene distribution of a particular species. The mammals are well covered; the birds, reptiles, amphibians and fish are not as complete, but this isa reflection of the state of the art and the availability of specimens. It points out sharply the areas where more research is needed. Each faunal list is followed by a suggested age for the locality. Radiocarbon dates are included when available. The last part of each faunal description, entitled “Remarks” includes a diverse amount of information, such as descriptions of the bone deposits, environment of deposition, interpre- tation of habitat and the first occurrences of particular species. Finally, there is a listing of references for that particular fauna. The paper has sixteen illustrations. One is a map showing the location of all the faunas discussed, one 1s a chart with the suggested chronological sequence of Birds of Man’s World By Derek Goodwin. 1978. British Museum (Natural History) and Cornell University Press, London and Ithaca. villi + 193 pp., illus. U.S. $10.95. Although the title may suggest to some another “gloom and doom” volume on Man’s destruction of wildlife, the subject matter concerns the ability and inability of birds to adjust to human environmental changes. Through a series of examples, Goodwin illustrates how the natural habits of various species suit them for adapting to, and thriving in Man’s world, while those of other species doom them to extinction. This book appears to be written primarily for the layman, but as pointed out by D. W. Snow in the foreword, it also contains much of value to the professional ornithologist. The majority of Goodwin’s examples are European, with considerable emphasis on personal experience; however, other continents are not neglected, with many examples coming from the literature. Unfortun- ately; the sources are rarely given, and then usually vaguely. Thus, the serious student who wishes to pursue a topic in greater depth has little help in THE CANADIAN FIELD-NATURALIST Vol593 the faunas, three are artistic reconstructions of specific Pleistocene vertebrates, and eleven are photo- graphs of specimens. All of the photographs occupy a full page. The map could have been a little larger as the amount of reduction tends to make reading the numbers of the localities difficult, but with a little squinting they can be deduced. The final discussion and summary places the entire history of the Pleistocene of Canada in perspective. It discusses chronologically first appearances, migra- tions, changing environments and their significances with regard to changes in the fauna. The reference section has plenty of citations, which will permit anyone interested to pursue a particular subject further. The appendix contains a listing of nine faunas from northeastern Siberia for comparison with those from Canada and Alaska. This section is of interest since the faunal lists include species that did not manage to cross the Bering Straits into North America. This well-ordered, concise review will makea handy starting point for anyone who wishes to obtain an overall view of the Pleistocene in Canada and Alaska. H. GREGORY MCDONALD Department of Vertebrate Palaeontology, Royal Ontario Museum, Toronto, Ontario M5S 2C6 locating original material. In his introduction, Goodwin points out that his book is not intended as a thorough review covering all aspects of Man’s interactions with birds. The six numbered chapters cover various aspects of this topic as outlined below. The text is chatty and non- quantitative but written well and apparently free of printing errors. Several photographs and line illustra- tions by Robin Prytherch enhance the book, although neither are of exceptional quality. Chapter |, “Man and the environment,” is concern- ed with the effects on birds of habitat alteration. These include obvious direct effects, such as destruction of habitat for some species, and expansion of habitat for others. Indirect effects are also discussed. For example, although many species benefit from the provision of artificial nest sites, the increase in aggressive hole-nesters such as Starlings could result in a decrease in less aggressive hole-nesting species. Although not intended as a complete review, this chapter seems comprehensive, with a few minor omissions. A long discussion about birds coming to 1979 drink at artificial watering-holes in desert areas might well have mentioned the large number of recent occurrences of marine birds far inland at large reservoirs in the southern USA. I found Goodwin’s discussion of Purple Martins and Tree Swallows misleading. After mentioning the benefit to the martins of numerous houses built for them, Goodwin notes that Tree Swallows have also been incidental beneficiaries by using nest boxes which have not attracted martins. This is true, but surely the Tree Swallow has benefitted far more from the extensive bluebird house programs in the prairie provinces and other parts of North America. The appearance of hybird bluebirds and swallows during range expan- sions caused by the provision of artificial sites is also omitted. Chapter 2, “Birds in towns,” consists of a compari- son of major cities in various parts of the world in relation to their avifauna. Interactions of human behavior and bird behavior are shown to determine both the extent birds will live in cities and which species will do so under various circumstances. Chapter 3, “Birds fed by Man,” includes discussion of birds as pests, birds as scavengers on garbage, and deliberate feeding of birds by people. As in other chapters, Goodwin emphasizes the natural behavior of species as it adapts them to respond to food provided by Man intentionally or otherwise. Missing from the discussion of birds in crops are cases of birds saving crops, such as California Gulls in Utah and Franklin’s Gulls in Manitoba. This chapter overlaps to some extend both chapters 2 and 5. Introduced birds on all continents form the subject BOOK REVIEWS 455 matter of chapter 4. After pointing out that most species that do well on being introduced into a new place are species that did well in their places of origin, where they were frequently somewhat of a pest, Goodwin reviews the success and failure of various introductions, and comments on factors which may determine rate of success. This chapter is slightly outdated for North America: the House Finch now inhabits much more of the East than indicated: the Skylark has spread to the San Juan islands of Washington (at least temporarily), and the predicted action to control the Monk Parakeet in the USA has taken place. I was also surprised to find only brief mention of the Black Swan of New Zealand with no reference to its adverse effects on native waterfowl there. In chapter 5, Goodwin compares several species as to their habit of eating an unnatural food — bread. He ponders how each might come to recognize bread as food, howeach behaves in obtaining and eating bread, and how much competition for bread among species takes place. In his final chapter, Goodwin offers a cautiously optimistic outlook on the future. A list of scientific names of birds discussed includes page numbers, and thus also serves as a species index. I recommend this book as a highly readable essay on an important topic. MARTIN K. MCNICHOLL Beak Consultants Ltd., 3530 11 A St. N.E., Calgary, Alberta T2E 6M7 Statistical Inference from Band Recovery Data: a handbook By C. Brownie, D. R. Anderson, K. P. Burnham, and D. S. Robson. 1978. Resource Publication Number 131, U.S. Fish and Wildlife Service, Washington, D.C. No price given. Many workers with fish, mammals and birds have need of mark-recapture estimation techniques for assessment of population sizes and mortality rates. It is not that better estimates are provided by such techniques, but rather that direct sampling of concealed and highly mobile populations is impossible. Given the need for them, mark-recapture methods have evolved to levels of sophistication that would greatly impress early workers in the area. This handbook represents the latest state of the art, reflecting several decades of improvements in two areas: precision and accuracy. Students encountering the simplest mark-recapture methods for the first time tend to react with shock at the low precision (i.e., wide confidence intervals) of their estimates, and often are convinced that they must have made a mistake in their calculations. The methods also tend to be vulnerable to biases in estimation caused by failure of assumptions, and such biases result in low accuracy. Methods such as those presented in this book improve precision by using data from repeated marking and capturing overa long series of time intervals (usually years), and they deal with biases by presenting a hierarchical series of models which differ from each other in their assumptions about the parameters of the populations being estimated. The authors emphasize that results are still dependent on assumptions for any particular model, but this approach allows sequential testing of the validity of various assumptions, which leads to 456 selection of the model satisfying the conditions with which the researcher is dealing. Although the authors hope they have produced “a simple, easy-to-read primer.” have emphasized examples from real data, and have put some of the mathematics in appendices, this handbook is not for the beginner. Differential calculus and two or three courses in applied statistics are assumed, and anyone without prior knowledge of the basic principles of mark-recapture methods is advised to start with the treatment of this subject in a general quantitative ecology text. For the professional working on wild populations that can not be assessed directly, however, this handbook will be a necessary addition to his library. The discussion and examples are in terms of bird-banding studies, but the models are generally applicable. Equal time intervals and recovery of dead animals (marked or unmarked) are assumed, with some discussion of unequal intervals and of capture and release of live animals. All methods described are related to FORTRAN computer programs designed for IBM systems, with The Gannet By Bryan Nelson. 1978. Buteo Books, Vermillion, South Dakota. 336 pp., illus. U.S. $25. It is obvious that Nelson has spent many hours watching Gannets. His readable style combines dry and factual with informal and anecdotal accounts. An incredible amount of detail is found throughout the book with emphasis placed on fact, integration, and interpretation. Nelson assures the reader that this book is not an extract of his recent monograph (The Sulidae: Gannets and Boobies. 1978. Oxford Uni- versity Press. 1012 pp. U.S. $98) but has been written afresh and with a different approach. The book is organized into seven chapters. Addi- tional introductory and final sections include a preface, acknowledgments, a thumbnail sketch intro- duction, two pages on names, references, bibliog- raphies, and an index. The 32 tables are grouped together and follow chapter 7. Major points are set out in numbered statements in a summary for each chapter. Numerous illustrations by John Busby are scattered throughout the book and are particularly valuable in the behavior chapter; a two and one-half page schematic representation of the Gannet’s life history is especially nice. Black-and-white photo- graphs covering 32 pages are located in the middle of the book. Although figures are numbered, the text often does not refer to them. The map numbers corresponding to the locations of the six Canadian THE CANADIAN FIELD-NATURALIST Vol. 93 versions converted for use on CDC systems also available. Detailed instructions for data coding are given, and small jobs can be sent (as punched data card decks) to the Patuxent Wildlife Research Center in Maryland for analysis and commentary at no charge. The loan of tapes can be requested by those wishing to obtain the programs, and they will be sent without charge to be copied and returned. The authors strongly emphasize the need for advance planning of any study based on a thorough understanding of the methods and their assumptions. Chapter 8 summarizes analysis methods proposed by previous workers, and discusses interpretation of mark-recapture analysis results. Chapter 9 considers the planning of a study, including timing of sampling and necessary sample size. There is a comprehensive bibliography. R. H. GREEN Department of Zoology, University of Western Ontario, London, Ontario N6A 5B7 gannetries in Figure 10 also do not correspond to the numbers in the text. In general a better correlation between text citations and figures, tables, and illustra- tions would have been an improvement. Over 100 pages are devoted to breeding behavior and ecology where the author presents a myriad of interesting facts and ideas. An interpretation of the function and survival value of various behavioral traits 1s usually provided. The numbers and distribu- tion chapter contains excellent sections on how, when, and what to count. Another interesting section discusses fishing techniques and includes quotes from Scottish fishermen. Chapter 6 on the Gannet family and the order seems to come too late in the book. Chapter 7, on the other hand, is a perfect final chapter with subchapter headings on such fascinating topics as early accounts of the Gannet’s natural history, its use as food, man’s practice of culling in breeding colonies, oiled birds, its behavior in captivity, and its role in art and literature. I believe that those who read the book will find immense enjoyment. Buteo Books should be com- mended for publishing and making available several important avian works originally published in Europe. NOEL J. CUTRIGHT Wisconsin Electric Power Co., 231 W. Michigan, Mil- waukee, Wisconsin 53201 1979 BOTANY Vascular Plant Families By James Payne Smith, Jr. 1977. Mad River Press, Inc., Route 1, Box 151-B, Eureka, California 95501. 320 pp., illus. $9.55. Students and teachers of plant taxonomy alike will welcome this elementary book which contains every- thing from an outline of the taxonomic hierarchy and scientific names, through family descriptions, lists of regional floras for the United States, notes on how to collect plants, to a useful glossary. Following the system of Cronquist, Takhtajan, and Zimmerman (1966) for divisions, classes, subclasses and orders, and Cronquist (1968) for the sequence of families, the author presents “an introduction to the families of vascular plants native to North America, and selected families of ornamental or economic importance.” This introduction consists of short descriptions of the families, an indication of the number of genera and species, a few selected genera, BOOK REVIEWS 457 some recognition characters, and a floral formula. Over one hundred plates of fine line drawings by Kathy Simpson accompany this text. Of special interest to the student is the chapter entitled “Vege- tative morphology of the flowering plants.” This is an illustrated glossary which is divided into sections that give the terminology used to describe various parts of a plant and their arrangement, e.g., roots, stems, leaves, etc. The book is written for students by a teacher who has tried to present his subject as simply as possible. The elimination of all but a few of the non-North American families has certainly made this easier. The author is to be congratulated on a job well done! WILLIAM J. CODY Biosystematics Research Institute, Canada Agriculture, Ottawa, Ontario KIA 0C6 An Atlas of Airborne Pollen Grains and Common Spores of Canada By I. John Bassett, Clifford W. Crompton, and John A. Parmelee. 1978. Agriculture Canada, Ottawa. 350 pp., illus. $12 in Canada; $14.40 elsewhere. This attractively and robustly bound research monograph represents the culmination of over two decades of careful, well reported investigations by the authors into the problems of airborne allergenic pollen and spores. It isaddressed to workers who wish to identify airborne pollen and spores because of their interest in either allergy problems, or in general and applied palynology. Its effectiveness therefore should be measured by the clarity and quality of the illustrations; the rigor and decisiveness of the keys: and the degree to which it will become indispensable to the laboratory scientist in the appropriate dis- ciplines. But its format implies a wider readership target. The first 39 pages introduce the subject of allergy pollen. They vary from a few fascinating pages (15-16), to some dreary tables (16-39), which should be appendices. Pollen grains are exciting objects. A lively up-to-date chapter on their development, role in reproduction, life-cycle strategies, paleoenvironmen- tal reconstructions, etc., as well as a more animated discussion of the intriguing material on p. 13 on diurnal patterns and the spectacular data on the decline of ragweed fallout since the 1950s might have captured a wider readership. And what about allergic reactions to pollen? Couldn’t we have had a few pages summarizing the state of the medical arts on that? We are plunged into the dichotomous identification keys to pollen, the heart of the book for the user. Presumably, the user will be the technician in the allergy laboratory, because other pollen work deals with ranges of material incompletely covered in this book. Pollen keys at best are unsatisfactory and they are usually fully comprehensible and usable only by the authors themselves. The keys in this book are useful but uneven. For example, the distinctions between important species (Fagus, Quercus) are not clearly handled. Comptonia peregrina is not consistently tetraporate, as Figure 107 C and D demonstrate, and in fact is indistinguishable from Myrica. On the other hand A/nus could be separated readily to species, if it were useful to do so from the allergy viewpoint. The photomicrographs are of very uneven quality. Some taxa have no light microscope photographs (Carex), others have interference and SEM but no non-interference light microscope illustrations though they are essential (e.g.. Fraxinus, Luzula, Triglochin, Zea, and others). The magnification varies widely from plate to plate. And no attempt has been made to achieve uniformity of positioning of the grains in the SEM, photographs essential if they are to be used diagnostically. We are told that interference 458 photomicroscopy was employed because of its great use in such groups as Urtica. Possibly, but it is far from convincing that its use in most of the other groups illustrated adds anything to the book. Part 2 deals with fungus spores, but it differs markedly from Part | in having no keys, in offering only a selection of representative types and illustra- tions, and in providing nothing more than a glossary to initiate the novice into the distinctive descriptive ENVIRONMENT THE CANADIAN FIELD-NATURALIST Vol. 93 terminology of mycology. It creates the impression of being an afterthought. This book will find its way into most North American pollen-spore labs, but whether it becomes dirty and dog-eared with use is another question. J.C. RITCHIE Division of Life Sciences, Scarborough College, University of Toronto, West Hill, Ontario MIC 1A4 A Vanished World: the dinosaurs of western Canada By Dale A. Russell. 1977. Natural History Series Number 4. National Museum of Natural Sciences, National Museums of Canada, Ottawa. 142 pp. $12.95. In his introduction to A Vanished World, Dale Russell states that “The purpose of this volume is to visualize, so far as possible, the vanished world of Canadian dinosaurs . . . it is hoped that this work will help others to understand what western Canada may have been like during a very interesting and relatively well documented period of its physical history.” Upon reading the book, one cannot help but conclude that in the short space of 142 pages, the author has managed to manifest successfully a most original approach to the study of the dinosaur era. The book is filled with superb photographs (by Susanne M. Swibold) of the Canadian badlands, and of modern vegetation and habitats that compare closely with the projected environments of the Late Cretaceous in Canada. The photographs are com- plemented by equally striking artistic reconstructions (by Eleanor M. Kish) of many of the Canadian dinosaurs described in the text. All photographs and illustrations are in color. The first chapter deals with the history of dinosaur collecting in western Canada, and as most published histories of palaeontology are concerned mainly or only with American localities and collectors, much of the material will be new to the general reader. The second chapter provides a general survey of the geology of the Canadian badlands, witha brief insight into the methods of interpreting the prehistoric geography of an area from its sediments. The third chapter describes Dinosaur Provincial Park in Alberta, an area of about 15000 acres of badlands containing exposed sediments that are from 76 to 73 million years old. At that time, Dinosaur Provincial Park was part of an ancient alluvial plain situated between the Rocky Mountains and the great inland sea which then covered much of the interior of the North American continent. Considering in turn the sediments themselves, then their plant, inverte- brate, and vertebrate remains, the author logically and precisely reconstructs the environment of that period. A short section at the end of the chapter invites the reader to imagine himself back in those times and then to consider the possibility of his own survival under such conditions. The fourth chapter treats the badlands around Drumheller, Alberta in a similar manner. The sediments of the Drumheller region contain fossil remains ranging from 72 to 70 million years in age. At that time, Drumheller was located on the southern edge of a large delta that extended more than 200 miles from the Rocky Mountains to the inland sea. The delta was “an enormous low-lying wetland of shallow lakes, swamps and marshes, traversed by streams that fanned out from the major river courses to the edges.” Various aspects of the ecology of the delta are discussed, from the patterns of deposition of sediments to the vegetation and the dinosaurs that inhabited the area. Over a period of a few hundred thousand years, the ecosystem changed and the delta developed the characteristics of a coastal plain, as stream beds shifted and the swamps drained. The vegetation became markedly different from that of the wetland period, with forests of broad-leaved trees, many of which are familiar to us today. The kinds of dinosaurs inhabiting this drier, hardwood-forested area were somewhat different from those that lived in the swamps. The fifth chapter takes us through the sediments of 70 to 65 million years ago into the geological period that signalled the end of the dinosaurs. The vegetation and the dinosaurs that lived on the large subtropical floodplain of this period are discussed. Because the floodplain extended into the United States, con- temporary dinosaurs from Montana, Wyoming, and the Dakotas are also reviewed. 1979 The last chapter considers the extinction of the dinosaurs. The final testimony of the late Cretaceous/ Early Paleocene rocks is discussed, and various general aspects of the extinction problem are ex- amined. In a concluding section, entitled “From the Depths of Space,” the author takes the opportunity to elaborate upon the extinction theory he himself favors most: that of a cosmic event (probably a supernova) that bombarded the Earth with “waves of high-energy radiation comparable to those occurring near powerful thermonuclear explosions.” The bibliography, while comprehensive, is not overwhelming to the non-scientist, and many of the Biogeography and Ecology of Southern Africa Edited by M. J. A. Werger. 1978. W. Junk, The Hague. xvi + 1439 pp., illus. (in 2 volumes). DFL 365. Southern Africa as referred to in these two volumes covers an area of some 6100000 km?. The geo- political areas included are: Angola, part of Shaba, Zambia, Malawi, Mozambique, South West Africa, Botswana, Rhodesia, South Africa, Swaziland, and Lesotho. Southern Africa varies in altitude from sea level to 3482 m and has a wide range of biotic and abiotic features. There is an extensive published and unpublished literature on the biogeography and ecology of southern Africa. Much of this material appears in local journals, in various government reports, and as theses. As such it is not readily available to interested biogeographers and ecologists outside southern Afri- ca. These volumes successfully review this large body of knowledge. As Werger points out, the ecological aspects covered are mainly of a descriptive nature, concentrating on diversity and variation in species and on a structural-functional evaluation; physio- logical aspects of the ecology, and energy and nutrient flow systems remain virtually untouched. Research in these latter fields has only recently started in southern Africa. The two volumes are divided into 41 chapters. Approximately 75 percent of the contributing authors were living and working in South Africa when they wrote their chapters. Chapters | to 9 cover the physical and geological environment, past and pres- ent. Chapters 8 to 14 each deal with a major botanical biome while Chapters 16 to 31 each cover a major zoological taxonomic group. The next nine chapters deal with the biogeography and ecology of special habitats such as lakes, rivers, marine habitats, high termitaria, and heavy metal and other toxic soils. The final chapter discusses the conservation of southern BOOK REVIEWS 459 references are government publications which are reasonably accessible to the general public. A Vanished Werld is written in a clear and logical style, and the ancient environments reconstructed from the rocks and their fossil remains become strong visual images within the mind of the reader. This book will be a valuable addition to the library of any person who has an interest in natural science. JOANNE E. LINDSAY Department of Vertebrate Paleontology, Royal Ontario Museum, Toronto, Ontario MSS 2C6 African ecosystems. An extensive use of photographs and illustrations is made throughout the text. Typo- graphical errors were few and the largest single printing mistake (Figure 9, Chapter 39) is corrected with an erratum. A major pitfall of such reviews can be the length of time required to get them published. Although it is not possible to eliminate completely this lag phase, the publishers can do much to make it as short as possible, thereby increasing the value of the review. Dr. W. Junk, publishers, should be commended for the speed in which these two volumes were brought to press. References in the text date as recently as 1977. Dr. Werger’s contribution to the volumes as editor has been considerable. Throughout the text he has constructed a network of cross-references to earlier or later chapters in which additional information or a difference of interpretation can be found. Unfortunately, in many chapters readers are refer- red to a specific geological formation or plant community located near a town not likely to be known beyond southern Africa or to individuals who have never lived there. A much greater use of maps such as Figure | in Chapters 30 and 38 would have been very useful. Readers who have only a lay interest in the biogeography and ecology of southern Africa will find general information on many subjects concerning a most interesting part of the world. For example, in Chapter 16 we are informed that the world’s largest earthworms, measuring 7 m in length by 7.5 cm in diameter, occur near Debe Nek in the Cape Province in South Africa and that some savanna areas have rodent populations that exceed those of the Brown Lemming in the tundra of Alaska at cycle peaks (Chapter 31). 460 These two volumes should attract many readers in academic or research institutions with an interest in Africa. Although, as Dr. Werger notes, the books are not all-encompassing (both in subject or in literature citations), they are an excellent source of information for any amateur or professional ecologist, naturalist, or biogeographer beginning a project on southern Africa. Purchase of these volumes by interested The Ecology of North America By V.E. Shelford. 1978. University of Illinois Press, Ur- bana. Paperback edition, originally published 1963. 610 pp., illus. No price given. This book was first published in 1963, and is now resurrected in paperback form. Undoubtedly an important contribution to ecology in its time, the purpose of the book is“. .. to describe North America from an ecological viewpoint as it appeared in the period A.D. 1500 to 1600 before European settle- ment.” Taking over half a century to complete, Shelford undertook the monumental task of reconstructing the ecological story of primeval North America using results from his own studies, those of his students and his colleagues. This book is designed to assist the ecologist in understanding the habits, biotic com- munities, and the distribution and abundance of plants and animals in primeval North America. To this end, Shelford focuses on the biogeographical aspects of primeval ecology. The text is organized into 19 chapters. The first chapter reviews the meaning of major underlying ecological principles. Among them are habitat, community, dominance, community development, population densities and distribution, life span, life cycles, solar radiation, temperature and moisture, natural communities of North America, and com- munity terminology. Many of these principles con- tinue to form the underlying components of modern Chlorinated Phenoxy Acids and their Dioxins Edited by C. Ramel. 1978. Ecological Bulletins Number 27, Swedish Natural Sciences Research Council (NFR), Stockholm, Sweden. 302 pp. 75 SwCr. This bulletin is a collection of papers from a conference arranged by the Royal Swedish Academy of Sciences in Stockholm, 7-9 February 1977. The intent of the symposium was to provide an up-to-date summary of objective scientific data on chlorinated phenoxy acids and their dioxins, and to make a risk- benefit evaluation of this data in a series of work THE CANADIAN FIELD-NATURALIST Vol. 93 individuals, however, is unlikely since the approxi- mate Canadian price of $200 will be prohibitive in most cases. RICHARD D. ROBARTS Schultz International Limited, 1155 West Georgia Street, Vancouver, British Columbia V6E 3H4 ecology; however, a decade and a half of research and development has improved and expanded our knowl- edge of them. In this regard it is important that the reader be aware of these changes. The following 18 chapters cover all of the major communities in North America, from the high Arctic to southern Florida and Cuba. Shelford not only attempts to document the abiotic and biotic components of each community, he attempts to interrelate these components and develop a comprehensive review of the mechanics of the natural systems in primeval times. Despite his valiant attempts to do this there are inherent defects. The interrelationships of animals and plants were not well understood during Shelford’s time and consequently are poorly documented. In addition, the book fails to present quantitative data on animal populations and their food habits, and plant population dynamics. The book is accented with maps, graphs, tables and photographs. The references are extensive, and are useful to those who wish to examine some of the earlier works in ecology. I recommend this book to biologists who are well read in general ecology and who are capable of recognizing its inadequacies. PAUL A. GRAY Hough, Stansbury, Michalski Ltd., 1265 Arthur Street, Suite 409, Thunder Bay, Ontario P7E 6E7 groups in the areas of chemistry, plant physiology, toxicology, genetics, and ecology with economics. A group of 35 internationally recognized experts par- ticipated in the symposium. Unlike so many scientific symposia, this one was results-oriented. Summary papers with recommendations were prepared by each of the working groups. All of the background papers and summaries are given, along with a short overall conclusions and recommendations section. The chemistry section and a considerable propor- 1979 tion of the toxicology section are devoted principally to tetrachlorodibenzo-p-dioxin, the notorious TCDD. TCDD 1s an incidental contaminant formed during the manufacture of trichlorophenol, which in turnisa precursor of the herbicides 2,4,5-T and 2,4,5-TP (fenoprop). The bulletin gives an interesting account of the toxicological problems (real or imagined) from phenoxy acid (largely 2,4-D) use in Sweden. One of the more controversial aspects was the suspected poisoning of a reindeer herd in 1970. The veracity of other alleged cases of poisoning of humans, wildlife, fish, and bees are discussed in some detail. In the section on ecology and economics there is a thorough coverage of the effects of herbicide use in silviculture on vegetation, but not much information is given on how these effects relate to animal or insect ecology (i.e., habitat changes), in spite of the title. The importance of this subject is recognized, however, in the summary and recommendations. A good review of the effect of phenoxy herbicides on soil organisms 1s given. According to the National Research Council of Canada document on the same subject (Phenoxy MISCELLANEOUS BOOK REVIEWS 461 Herbicides, their Effects on Environmental Quality, NRCC Number 16075), several million kilograms of 2,4-D are used in Canada, largely for weed control in the prairies. In the order of 50,000 kilograms (1973- 1974) of 2,4,5-T are used for brush control on rights-of-way, mainly in the eastern provinces. This bulletin therefore addresses a situation in Canada of similar magnitude and import to that in Sweden. Although there are gaps in coverage (in large part due to a lack of information, I suspect), this volume is one of the best overall sources of information available on the subject. The many sections on TCDD have relevance apart from phenoxy herbicides, because trichlorophenol manufacture and dumping have created more TCDD problems than 2,4,5-T use. The summaries and recommendations are succinct and well-balanced from an overall risk-benefit viewpoint. R.J. NORSTROM Wildlife Toxicology Division, Canadian Wildlife Service, Ottawa, Ontario KIA 0E7 Last of the Naturalists: the career of C. Hart Merriam By Keir B. Sterling. 1977. Arno Press, New York. 472 pp., illus. U.S. $23.00. This book should be perused by those interested in the formation and early activities of the American Ornithologists Union, National Geographic Society, American Society of Mammalogists, and U.S. Fish and Wildlife Service, all of which Merriam (1855-1942) was directly involved in founding. Sterling has exhaustively researched the life of Merriam using file letters to and from Merriam, congressional documents where Merriam partici- pated, journal and newspaper articles, interviews with people who knew him, and other detailed records to produce a fine biographical work that doesn’t stop with information on Merriam but includes per- sonalities of people that were around him. This revised book, first issued in 1974, is part of the Natural Sciences in America collection. The text is clut- tered with quite a few typographical errors: entire words and even sentences seem to be missing in some spots. The chapter on speciation is poorly written. As well, errant definitions occur, suchas Bergmann’s rule (p. 205). The Elliott Coues —- Merriam vendettas are probably: not fully covered. Perhaps a better title would be The First American Wildlife Biologist or Mammalogist, as the book’s present title implies that the Leopolds and Muries were not naturalists. Merriam’s career included being an amateur naturalist, a medical school graduate, practicing physician and professional biologist with the govern- ment. He worked under the vigilance and influence of Joel A. Allen, Spencer F. Baird, Jean B. Lamarck, Charles V. Riley, and Theodore Roosevelt. He influenced the professional lives of Vernon Bailey, William Brewster, Frank M. Chapman, Albert K. Fisher, Joseph Grinnell, Francis Harper, Henry W. Henshaw, Waldo L. McAtee, Edward W. Nelson, and Wilfred Osgood, many of whom worked for the U.S. Biological Survey under his leadership. Merriam was highly interested in the biogeography and systematics of mammals and birds (he collected thousands of specimens); published his first paper at age 17, became an authority on birds at 22; was the United States’ first full-time professional ornitholo- gist; did not “follow the herd”; scorned religion; liked to eat meat of cat, eagle, and skunk; had yearly field experience throughout his life; believed that species were basically fixed but all were not yet discovered: named and described many new genera, species, and subspecies, being a splitter when it came to zoological 462 THE CANADIAN FIELD-NATURALIST nomenclature; put forth the life-zone theory, feeling that temperature was the ultimate factor affecting the distribution of plants and animals; worked intensively with gophers and bears; thought life was too short for memorizing terminology, believing that practical application was pertinent to all work; felt that microscopic work in the laboratory was emphasized too much over field work in college; did not get along with others whose drives were as great as his; was involved in the still controversial killing of seals in the north; could not receive criticism well; tried to get deer farming initiated in areas where cows could not graze; liked art and automobiles; despised the words biota and ecology; was against conservation movements that were not scientifically based; had a special ability to obtain capable men to work for him at the Le Naturaliste Canadien, Index By G.-W. Corrivault and P. Morisset. 1979. Les Presses de PUniversité Laval, Québec. 352 pp. $10. Readers of The Canadian Field-Naturalist are familiar with the problems of data retrieval from a long series of volumes. The search for specific data through so many annual indices Is not only tedious, it may also be a poor use of one’s time. Because annual indices are not equally thorough and comprehensive, some being rather skimpy, one is never sure that significant data has not escaped such a search. Cumulative indices are the usual solution to these problems, but not every journal has been able to prepare and publish these. Le Naturaliste Canadien is now in its 105th volume and its data retrieval problems are quite similar to those of The Canadian Field-Naturalist, except for the existence of a series of cumulative indices of which the present one is the fourth. A cumulative index covering the first 20 volumes was published at the end of volume 20. A second cumulation was printed at the end of volume 54 and covers volumes 21-54. A cumulated author index for volumes 1-81 will be found at the beginning of volume 82. The new cumulation carries the coverage forward to volume 100. It is a separate publication, not a particular issue of the serial. It is published in two parts. The first part is analphabetized author index of Vol. 93 Biological Survey; served office in many scientific societies; Was most particular about the proper preparation of skins and skulls; became interested in Indians in the early 1900s, mostly ignoring or even terming as useless most other mammals for the remainder of his life; later in life turned against the Biological Survey’s predator poisoning campaign which he started; was never considered an intellectual; and never accepted the developments that progressed in genetics. He was the father of modern Mammalogy. RICHARD M. ZAMMUTO Department of Zoology, University of Western Ontario, London, Ontario N6A 5B7 2504 papers found in volumes | to 100. By design the author index is not thorough: many minor items of fleeting value have been omitted, such as notices from the editor to his readers, some letters to the editor, society notices, and the like. The second part deals only with volumes 55 to 100. It is a subject and taxon index with double references. The bracketed numbers refer to the bibliography and are followed by the usual volume and page references. There are more than 60000 individual entries. Botanists will be especially interested by this index. There is very little botany in the first 50 volumes of Le Naturaliste Canadien. But in the 1920s and 1930s botanical papers gradually gained in numbers and import. More than 8000 plant names occur in the index, including nearly all the vascular plants of Canada, Greenland, and Alaska; also a majority of Canadian mosses and lichens, and a fair showing of algae and fungi. Taxonomic innovations are underlined in the index: there are more than a thousand of them anda good half of them are plant names. B. BOIVIN Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6 Se 1979 NEW TITLES Zoology The alligator: king of the wilderness. 1977. By William and Ellen Hartley. Nelson, New York. 175 pp. US $7.95. + Animal behavior: an evolutionary approach. 1979. By John Alcock. Second edition. Sinaver, Concord, Maine. xii + 532 pp., illus. US $16. Animals of the oceans: the ecology of marine life. 1977. By M. Angel and T. Harris. Two Continents, New York. 156 pp., illus. US $10.95. Arachnology. 1978. Edited by P. Merrett. Proceedings of the Symposia of the Zoological Society of London, Number 42, Exeter, England, July 1977. Academic, New York. xxxil + 530 pp., illus. US $46.50. Eloquent animals: a study in animal communication. 1978. By Flora Davis. Coward, McCann, and Geoghegan, New York. 223 pp. US $8.95. Field guide to seashells of the world. 1978. By Gert Lindner. Van Nostrand Reinhold, New York. 271 pp., illus. Cloth US $12.95; paper US $8.95. Fifty common birds of Oklahoma and the south great plains. 1977. By G.M. Sutton. University of Oklahoma Press, Norman. xii + 113 pp. US $7.95. Insect magic. 1978. By Kjell B. Sandved and Michael G. Emsley. Viking, New York. 128 pp., illus. US $16.95. t+Management and biology of Pacific flyway geese. 1979. Edited by Robert L. Jarvis and James C. Bartonek. Oregon State University Book Stores, Corvallis, Oregon. 346 pp., illus. Paper US $5.50. Primates of South Asia: ecology, sociobiology and be- havior. 1977. By M.L. Roonwal and S.M. Mohnot. Harvard University Press, Cambridge, Massachusetts. xx + 421 pp., illus. US $25. Responses of Peary Caribou and Muskoxen to helicopter harassment. 1979. By Frank L. Miller and Anne Gunn. Occasional Paper Number 40, Canadian Wildlife Service, Edmonton. 90 pp. Free. Sea squirts of the Atlantic continental shelf from Maine to Texas. 1979. By Harold H. Plough. Johns Hopkins University Press, Baltimore. ix + 118 pp., illus. US $20. t+Studies on Atlantic halocyprid ostracods. 1979. By Martin V. Angel. Pergamon Press, Elmsford, New York. 128 pp.. illus. US $27.50. The vertebrates: their forms and functions. 1978. By Charles G. Crispens, Jr. Thomas, Springfield, Illinois. x + 209 pp., illus. US $15.50. BOOK REVIEWS 463 Water birds of California. 1977. By Howard L. Cogswell. University of California Press, Berkeley. vi + 399 pp.., illus. US $5.75. *Wolf ecology and prey relationships on Isle Royale. 1978. By Rolf Olin Peterson. National Park Service Scientific Monograph Series Number 11, United States Department of the Interior, Washington. 211 pp. Paper. The world of worms. 1978. By Dorothy Hinshaw Patent. Holiday House, New York. 11 + 124 pp., illus. US $6.95. Botany Botany: basic concepts in plant biology. 1978. By Terry L. Hufford. Harper and Row, New York. xviii + 535 pp., illus. US $16.95. Essays in plant taxonomy. 1978. Edited by H.E. Street. Academic Press, New York. 304 pp. US $24.50. *A guide to the literature on the herbaceous vascular flora of Ontario. 1978. By James L. Hodgins. Revised edition. Botany Press, 90 Wolfrey Avenue, Toronto. 73 pp. Paper $4. *How to prepare common wild foods. 1978. By Darcy Williamson. D. Williamson, Box 1032, McCall, Idaho 83638. 110 pp., illus. Paper US $6.95. Introduction to the algae: structure and reproduction. 1978. By Harold C. Bold and Michael J. Wynne. Prentice-Hall, Englewood Cliffs, New Jersey. xiv + 706 pp., illus. US $24. The physiology of woody plants. 1979. By Paul J. Kramer and Theodore T. Kozlowski. Academic Press, New York. Plant strategies and vegetation processes. 1979. By J.P. Grime. Wiley-Interscience, New York. 264 pp. Cloth US $26.50; paper US $10.95. Plantcraft: a guide to the everyday use of wild plants. 1978. By Richard Mabey. Illustrated by Marjorie Blamey. Universe, New York. 176 pp., illus. US $10. The powdery mildews. 1978. Edited by D.M. Spencer. Academic Press, New York. 586 pp. US $54.75. +The rare vascular plants of Saskatchewan. 1979. By Robert V. Maher, George W. Argus, Vernon L. Harms, and John H. Hudson. Syllogeus Number 20. National Museum of Natural Science, Ottawa. 55 pp. + maps English + 55 pp. French. Free. River plants: the macrophytic vegetation of watercourses. 1978. By S.M. Haslam. Illustrated by P.A. Wolseley. Cambridge University Press, New York. xii + 396 pp., illus. Cloth US $62.50; paper US $14.95. 464 Vegetation and production ecology of an Alaska arctic tundra. 1978. Edited by L.L. Tieszen. Springer-Verlag, New York. xvill + 217 pp., illus. US $34.80. Environment Adaptive environmental assessment and management. 1978. Edited by C.S. Holling. IHASA International Series on Applied Systems Analysis, 3. Wiley-Interscience, New York. xx + 378 pp., illus. US $16.50. * Aleta Karstad’s Canadian nature notebook. 1979. By Aleta Karstad. McGraw-Hill Ryerson, Toronto. 144 pp., illus. SIZ295: Biology and ethics: reflections inspired by a UNESCO symposium. 1978. By Bruno Ribes. UNESCO (Canadian distributor Renouf Publishing, Montreal). 202 pp. $14. Coral reefs: research methods. 1978. Edited by D.R. Stoddart and R.E. Johannes. Monographs on Oceano- graphic Methodology, 5. UNESCO (Canadian distributor Renouf Publishing, Montreal). xv + 581 pp. $33.60. The ecology of fossils: an illustrated guide. 1978. Edited by W.S. McKerrow. MIT Press, Cambridge, Massachusetts. 384 pp., illus. US $22.50. Environmental assessment: approaching maturity. 1978. By Selina Bendix and Herbert R. Graham. Ann Arbor Science, Ann Arbor, Michigan. xi + 288 pp. US $20. The environmental impact statement process: a guide to citizen action. 1978. By Neil Orloff. Information Resources Press, Washington. 200 pp. US $10.95. Exploring nature with your child. 1977. By Dorothy Shuttlesworth. Abrams, New York. 240 pp., illus. US $18.50. Freshwater wetlands: ecological processes and manage- ment potential. 1978. Edited by Ralph E. Good, Dennis F. Whigham, and Robert L. Simpson. Academic Press, New York. 392 pp. US $17.50. Handbook of environmental data and ecological par- ameters. 1979. Edited by S.E. Jorgensen. Pergamon Press, Elmsford, New York. 1100 pp. US $150. Handbook of environmental data on organic chemicals. 1977. By Karel Verschueren. Van Nostrand Reinhold, New York. vit+ 659 pp. US $37.50. An introduction to population ecology. 1978. By G. Evelyn Hutchinson. Yale University Press, New Haven. xi + 260 pp. US $17.50. Investigating ecology. 1978. By E.H. Blaustein and R. Blaustein. Arco, New York. 140 pp. Cloth US $11.95: paper US $7.95. THE CANADIAN FIELD-NATURALIST Vol. 93 Key works to the fauna and flora of the British Isles and northwest Europe. 1978. Edited by G.J. Kenrich, D.L. Hawksworth, and R. Sims. Academic Press, New York. 192 pp. US $16.15. Marine biota of the NE Pacific: a bibliography emphasizing systematics and distribution. 1978. By W.C. Austin and M.M. Deutsch. Khoyatan Marine Laboratory, Cowichan Station, British Columbia. Variously paged. Paper $16. Oceanography: concepts and history. 1978. Edited by Margaret B. Deacon. Benchmark Papers in Geology, Volume 35. Dowden, Hutchinson, and Ross (Distributed by Academic Press, New York). 416 pp. US $25. The pollution of flowers by insects. 1978. Edited by A.J. Richards. Academic Press, New York. 262 pp. US $26. Sampling design and statistical methods for environmental biologists. 1979. By Roger H. Green. Wiley-Interscience, New York. 272 pp. US $19.95. Tropical forest ecosystems: a state of knowledge report. By UNESCO, UNEP, and FAO. Natural Resources Research, XIV. UNESCO (Canadian distributor Renouf Publishing, Montreal). 683 pp. Paper $53.50; cloth $66.50. Miscellaneous Suicide or survival? The challenge of the year 2000. 1978. Various authors. UNESCO (Canadian distributor Renouf Publishing, Montreal). 192 pp. $14. Astronomy: from the earth to the universe. 1979. By J.M. Pasachoff. Saunders, New York. 476 pp., illus. $15.95 (Canadian $19.15). + Decision making for national parks in Canada north of 60°. 1978. By Terry Fenge. Working Paper Number 3, Presi- dent’s Committee on Northern Studies. University of Waterloo, Waterloo. 11 + 58 pp. Paper, no price given. +The development of tourism and its potential futures in Canada north of 60° with implications for national parks and related reserves. 1978. By Bryan Smale. Working Paper Number 1, President’s Committee on Northern Studies. University of Waterloo, Waterloo. 11 + 61 pp. Paper, no price given. +Elements of park and recreation administration. 1979. By Charles E. Doell and Louis F. Twardzik. Fourth edition. Burgess Publishing, Minneapolis. vi + 367 pp. US $13.95. +An international comparison of policies and institutional arrangements for national parks and related reserves in hinterland areas. 1978. By Julia Gardner. Working Paper Number 2, President’s Committee on Northern Studies. University of Waterloo, Waterloo. 11 + 61 pp. Paper, no price given. *Assigned for review + Available for review Index to Volume 93 Compiled by W. HARVEY BECK Aalders, L.E., 415 Acer nigrum, 34 rubrum, 34 saccharum, 32 Achillea borealis, 184 Actitis macularia, 188 Adamcik, R.S., A. W. Todd, and L. B. Keith. Demographic and dietary responses of Red-tailed Hawks during a Snowshoe Hare fluctuation, 16 Agabus sp., 171 Aggresion in Tree Swallows, Influence of weather on, 437 Agonum decentis, 80 Agrohordeum macounii, 166 Agropyron violaceum, 355 yukonense, 184 Agrostis borealis, 355 exarata, 163 Alaska, 77 Alaska, Fall foods of Common Snipe on the Copper River Delta, 171 Alaska, Response of wintering Moose to mechanical habitat rehabilitation in, 19] Alberta, 16, 67, 176, 272, 363, 411 Alberta, Black-necked Stilts nesting near Edmonton, 68 Alberta, Characteristics of Peregrine Falcons migrating through central, 1969-1978, 296 Alberta, Habitat utilization and population densities of the amphibians of northeastern, 144 Alberta, Largest Gray Wolf skulls found in, 308 Alberta, Life history characteristics of Little Brown Bats (Myotis lucifugus) in, 243 Alberta, Nesting of Calliope Hummingbird in Kananaskis Provincial Park, 449 Alberta, Probable hybrids of Cinnamon X Blue-winged Teal from southern, 316 Alberta, Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in, 48 Alberta, Taxonomy, distribution, and ecology of the cliff- brake ferns ( Pe//aea: Polypodiaceae) in, 288 Alberta, Winter denning of the Striped Skunk in, 252 Alberta, Winter feeding by Porcupines in montane forests of southwestern, 405 Alces alces gigas, 191 Allium schoenoprasum var. sibiricum, 167 Amblystegium juratzkanum, 140 Ambystoma laterale, 194 maculatum, 194 Amelanchier alnifolia, 167, 184 arobrea, 32 Amphibians of northeastern Alberta, Habitat utilization and population densities of the, 144 Anaphalis margaritacea var. subalpina, 163 Anas cyanoptera, 316 discors, 316 platyrhynchos, 55 Andrews, S., 449 Andropogon gerardi, 35 Anenome drummondtii, 184 multifida, 184 narcissiflora s. lat., 263 parviflora, 184 Anomodon minor, 140 Antennaria Friesiana ssp. compacta, 265 neoalaskana, 265 Aquila chrysaetos, 367 Arabis canadensis, 36 Aralia nudicaulis, 163 Arctic Canada, Ivory Gull colonies in southeastern Elles- mere Island, 173 Arctostaphylos rubra, 184 uva-ursi, 184 Ardea herodias, 439 Arenaria interpres, 188 melanocephala, 188 Arnica alpina, 184 cordifolia, 169 Artemisia alaskana, 184 frigida, 184 hyperborea, 184 rupestris, 184 Article or note?, 337 Asclepias exaltata, 36 Asio flammeus, 411 Asplenium platyneuron, 36 Aster alpinus, 355 alpinus ssp. vierhapperi, 169, 265 ciliolatus, 163 ericoides, 35 macrophyllus, 33 modestus, 169 pilosus, 36 Sagittifolius, 33 sibiricus, 184 yukonense, 184 Astragalus americanus, 168 umbellatus, 184 Athyrium filix-femina ssp. cyclosorum, 165 Atrichum tenellum, 431 Avena sativa, 285 Avocet, American, 68 Award, FON Conservation, for The Canadian Field- Naturalist, 452 Award, Robert Lister receives, 337 Bacidia chlorococca, 140 Badger, 365 Bailey, R.O. Wild Mallard stocking in a large marsh habitat, 55 Ball, P.W. Thaspium trifoliatum (Meadow-parsnip) in Canada, 306 Barclay-Estrup, P. and R. A. Sims. Epiphytes on White Elm, Ul/mus americana, near Thunder Bay, Ontario, 139 465 466 Barkworth, M. E. reviews by, 104, 212 Bassett, I. J., reviews by, 102, 103 Bat, Big Brown, 245 Keen’s, 247 Bat, Big Brown, (Eptesicus fuscus) in Alberta, Reproductive biology of, 48 Bats, Little Brown, (Myotis lucifugus) in Alberta, Life history of, 243 Beam, B., review by, 95 Bear, Black, 423 Beechey, T. J., review by, 100 Beetles, carabid, Examination of overwintering, for asso- ciated mites, 79 Bellolio-Trucco, G., 431 Berberis nervosa, 128 Bidens cernua, 163 Biological flora of Canada. 1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry, The, 415 Biology — the unknown science? (editorial), 6 Bird banding, Inland, 202 Bird, D. M., review by, 205 Bjorge, R. R., 252 Black Bear confrontation, Blue Grouse brood hen, 200 Blackbird, Brewer’s breeding in the Northwest Territories, 76 Blarina brevicauda, 315, 320 Blue Grouse brood hen — Black Bear confrontation, 200 Blueberry, Sweet Lowbush, The biological flora of Canada. 1. Vaccinium angustifolium Ait., 415 Bluebird society, 202 Bobbette, R. S. W., review by, 344 Boissonneau, A. N., review by, 213 Boivin, B., review by, 462 Bonasa umbellus, 18 Bosmina sp., 267 Botrychium virginianum var. europaeum, 163 Boxall, P. C., 411 Boxall, P. C. Interaction between a Long-tailed Weasel and a Snowy Owl, 67 Boyd, H., review by, 105 Brachythecium relfexum, 140 salebrosum, 140 Brasenia schreberi, 389 Breeding areas and overnight roosting locations in the northern range of the Monarch Butterfly (Danaus plexippus plexippus) with a summary of associated migratory routes, 41 Breeding in the Northwest Territories, Brewer’s Blackbird, 76 Brewster, G. J., 132 British Columbia, 70, 71, 200, 433 British Columbia, Giant Cow Parsnip (Heracleum mante- gazzianum) on Vancouver Island, 82 British Columbia, Nesting of Horned Puffins in, 84 British Columbia, Virgin Douglas Fir forest on Saturna Island, 126 Britton, D. M., review by, 345 Bromus ciliatus, 166 Brunton, D. F., review by, 347 Brunton, D. F., Taxonomy, distribution, and ecology of the cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta, 288 THE CANADIAN FIELD-NATURALIST Vol. 93 Brunton, D. F., S. Andrews, and D. G. Paton. Nesting of the Calliope Hummingbird in Kananaskis Provincial Park, Alberta, 449 Buffalo, Black, (Osteichthyes: Catostomidae), First Can- adian record of the, 304 Bufo hemiophrys, 144 Busby, D. G., 446 Buteo jamaicensis, 16, 367 swainsoni, 367 Butterfly, Monarch, (Danaus plexippus plexippus), Breed- ing areas and overnight roosting locations in the northern range of the, with a summary of associated migratory routes, 41 By-laws of The Ottawa Field-Naturalists’ Club, Notice of change to the, 337 Calamagrostis canadensis ssp. Lansdorffii, 262 deschampsiodes, 355 purpurascens, 184, 262 Calathus ingratus, 80 Calidris alpina, 188 bairdi, 188 mauri, 188 minutilla, 188 pusilla, 188 Calla palustris, 262, 389 Callitriche heterophylla, 389 Calosoma frigidum, 80 Cameron, R. D., K. R. Whitten, W. T. Smith, and D. D. Roby. Caribou distribution and group composition associated with construction of the Trans-Alaska Pipe- line, 155 Campbell, C., 28 Campbell, R. W., H. R. Carter, and S. G. Sealy. Nesting of Horned Puffins in British Columbia, 84 Canada, Man’s influence on potential nesting sites and populations of swallows in, 371 Canada, Thaspium trifoliatum (Meadow-parsnip) in, 306 Canada, The biological flora of, |. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry, 415 Canada, The Oriskany sandstone outcrop and associated natural features, a unique occurrence in, 28 Canada, Utricularia geminiscapa at Mer Bleue and range extensions in eastern, 391 Canadian Field-Naturalist, FON Conservation Award for The, 452 Candelaria concolor, 140 Canis latrans, 367 lupus, 308 l. irremotus, 308 l. occidentalis, 308 |. pambasileus, 308 Capella gallinago delicata, 171 Cardamine umbellata, 263 Carex aquatilis, 184 atherodes, 167 bigelowii, 355 canescens var. subloliacea, 262 concinna, 167 filifolia, 183 glacialis, 262 lapponica, 262 laxiflora var. gracillima, 36 1979 membranacea, 262 pedunculata, 34 pensylvanica, 33 Williamsii, 262 Caribou, Barren-ground, 155 Woodland, 71 Caribou distribution and group composition associated with construction of the Trans-Alaska Pipeline, 155 Caribou, Distribution and movements of Selkirk, 1972- 1974, 71 Carpinus caroliniana, 34 Carter, H. R., 84 Carya cordiformis, 34 ovata, 32 Cassiope tetragona, 184 Castilleja caudata, 168 miniata, 163 Catfish, Flathead, First Canadian record of a, 179 Catling, P. M., 399 Catoptrophorus semipalmatus, 188 Catostomus commersoni, 267 Cavers, P. B., 378 Cedar, Eastern White, Thuja occidentalis, in western Nova Scotia, Status of, 326 Celtis occidentalis, 34 Cerastium maximum, 261 Ceratophyllum demersum, 389 Cervus elaphus, 282 Cetraria ciliaris, 140 Chamaedaphne calyculata, 264 Chaoborus albipes, 267 Charadrius alexandrinus, 188 semipalmatus, 188 vociferus, 188 wilsonia, 188 Chipmunk, 320 Least, 314 Chlidonias niger, 276 Cicuta maculata var. angustifolia, 168 Cinclidium latifolium, 431 Cinna latifolia, 163 Circaea alpina, 163 Cirsium arvense, 284 Cladonia spp., 286 Clethrionomys gapperi, 314, 320 Cody, W. J., 163, 259 Cody, W. J., reviews by, 211, 212, 457 Collomia linearis, 168 Conopholis americana, 36 Constitution of the Ottawa Field-Naturalists’ Club, Notice of motion to amend the, 338 Cook, F. R., 321 Cook, F. R. and J. Preston. Two-lined Salamander, Eurycea bislineata, in Labrador, 178 Cormorant, Great, Wind-caused death of, 175 Cornus racemosa, 33 stolonifera, 284 Coyote, 367 Crappie, Black, in the Ottawa River, Seasonal growth, food, and feeding habits of young-of-the-year, 232 Crataegus chrysocarpa, 34. sp., 34 INDEX TO VOLUME 93 467 Crepis nana, 184 Croskery, P., reviews by, 93, 97, 107, 339 Crossman, E. J. and J. H. Leach. First Canadian record ofa Flathead Catfish, 179 Crossman, E. J. and S. J. Nepszy. First Canadian record of the Black Buffalo (Osteichthyes: Catostomidae), 304 Cutright, N. J., reviews by, 99, 207, 342, 456 Cyclops sp., 267 Cygnus olor, 433 Cypress Hills population of White-crowned Sparrows, Song pattern of the, 272 Cypripedium calceolus var. planipetalum, 355 Cyr, A., review by, 106 Cystopteris fragilis ssp. fragilis, 261 Dagg, A. I., reviews by, 109, 217 Dale, H. M., 386 Danaus plexippus plexippus, Breeding areas and overnight roosting locations in the northern range of the Monarch butterfly with a summary of associated migratory routes, 41 Dance, K. W., 180 Danthonia intermedia, 166 spicata, 34 Daphnia longispina, 267 David, N. Barn Owls in Quebec, 323 Davies, R.W. Dispersion of freshwater leeches (Hir- udinoidea) to Anticosti Island, Quebec, 310 Dawe, N. K. and E. R. White. Giant Cow Parsnip ( Hera- cleum mantegazzianum) on Vancouver Island, British Columbia, 82 DDT, Eggshell thickness in American shorebirds before and since, 187 Deer, Black-tailed, 130 Dekker, D. Characteristics of Peregrine Falcons migrating through central Alberta, 1969-1978, 296 Deckker, D., R. Lister, T. W. Thormin, D. V. Weseloh, and L. M. Weseloh. Black-necked Stilts nesting near Ed- monton, Alberta, 68 Dendragapus obscurus, 200 Denning of the Striped Skunk in Alberta, Winter, 252 Densmore, R., 77 DesGranges, J.-L. Abandoned windmill used as a nesting site by Great Blue Herons, 439 DesGranges, J.-L., review by, 94 Desmodium canadense, 34 dillenti, 35 paniculatum, 35 rotundifolia, 35 Desmognathus fuscus fuscus, 193 Development of young in Ontario Black Terns, Nesting biology and, 276 Diaptomus oregonensis, 267 Dicranum fuscescens, 129 montanum, 140 Diets of Elk in Saskatchewan, Summer, autumn, and winter, 282 Dispersion of freshwater leeches (Hirudinoidea) to Anti- costi Island, Quebec, 310 Disporum lanuginosum, 36 Distribution and group composition associated with con- struction of the Trans-Alaska Pipeline, Caribou, 155 468 Distribution and habitats of four annual smartweeds in Ontario, 378 Distribution and movement of Selkirk Caribou, 1974, 71 Distribution, Taxonomy, and ecology of the cliff-brake ferns ( Pellaea: Polypodiaceae) in Alberta, 288 District of Mackenzie, Range extensions of vascular plants in northern Yukon Territory and northwestern, 259 Draba alpina, 263 borealis, 263 crassifolia, 263 incerta, 263 Drepanocladus revolvens var. intermedius, 431 Drosera rotundifolia, 263 Dryas alaskensis, 263 drummondii, 167, 184 integrifolia, 184 octopetala ssp. alaskensis, 263 Dryopteris fragrans, 262 spinulosa, 163 Dunn, E. H. Nesting biology and development of young in Ontario Black Terns, 276 Eagle, Bald, 367 Golden, 367 Eagles, Bald, in central summering, 443 Eagles, P. F. J., reviews by, 102, 347 Earthworm cocoons as a drift component in a southern Ontario stream, 180 Ecology of the cliff-brake ferms (Pellaea: Polypodiaceae) in Alberta, Taxonomy, distribution, and, 288 Editor’s report for 1978, 201 Edwards Y. Biology — the unknown science? (editorial), 6 Edwards, Y. Is biology unknown? (letter), 330 Eggshell thickness in American shorebirds before and since DDT, 187 Egret, Snowy, in the Northwest Territories, 329 Egretta thula, 329 Elaphe obsoleta obsoleta, 28, 322 vulpina gloydi, 399 Elatine minima, 388 Eleocharis acicularis, 388 robbinsii, 389 Elk in Saskatchewan, Summer, autumn, and winter diets of, 282 Ellesmere Island, Arctic Canada, Ivory Gull colonies in southeastern, 173 Ellesmere Island, Ring counts in Salix arctica from, 81 Elm, White, Ulmus americana, near Thunder Bay, Ontario, Epiphytes on, 139 Elodea canadensis, 389 Elymus glaucus, 129 macouniil, 166 Epilobium angustifolium, 184, 284 latifolium, 184 Epiphytes on White Elm, U/mus americana, near Thunder Bay, Ontario, 139 Eptesicus fuscus, 245 f. pallidus, 48 Eptesicus fuscus, Reproductive biology of the Big Brown Bat in Alberta, 48 1972- New Brunswick, Decline of THE CANADIAN FIELD-NATURALIST Vol. 93 Equisetum hyemale var. affine, 166 spp., 284 Erethizon dorsatum, 405 Erigeron caespitosus, 184 compositus, 184 Philadelphicus, 169 Eriocaulon septangulare, 388 Eriophorum angustifolium ssp. triste, 262 russeolum var. albidum, 262 triste, 262 Eritrichium splendens, 264 Erskine, A. J. Man’s influence on potential nesting sites and populations of swallows in Canada, 371 Etheostoma sp., 268 Eumeces fasciatus, Five-linked Skink, in Ontario, Eastern limit of the, 321 Euphagus cyanocephalus, 76 Eurhynchium oreganum, 126 Eurycea bislineata, Ywo-lined Salamander in Labrador, 178 Eurycea bislineata bislineata, 194 Eutamias minimus, 314 Evernia mesomorpha, 140 Fagus grandifolia, 34 Fahselt, D., P. Maycock, G. Winder, and C. Campbell. The Oriskany sandstone outcrop and associated natural features, a unique occurrence in Canada, 28 Falco peregrinus, 296 p. anatum, 297 p. tundrius, 297 sparverius, 198 Falcons, Peregrine, migrating through central Alberta, 1969-1978, Characteristics of, 296 Feeding, Aquatic, by a Woodchuck, 309 Feeding habits of young-of-the-year Black Crappie in the Ottawa River, Seasonal growth, food, and, 232 Feeding, Winter, by Porcupines in montane forests of southwestern Alberta, 405 Festuca occidentalis, 129 ovina ssp. alaskana, 262 subulata, 129 Fetterolf, P. M. Common Garter Snake predation on Ring- billed Gull chicks, 317 Fetterolf, P. M. Intraspecific food theft by the American Kestrel, 198 Field research investigators, Proposals invited from, 201 Fir, Douglas, forest on Saturna Island, British Columbia, Virgin, 126 Fissidens obtusifolius, 431 FON Conservation Award for The Canadian Field-Nat- uralist, 452 Fontinalis sp., 388 Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, Ontario, 266 Foods of Common Snipe on the Copper River Delta, Alaska, Fall, 171 Food-storage behavior of Peromyscus maniculatus gracilis and p. leucopus noveboracensis, Nesting and, 239 Forest, oak-hickory, 28 Fox, Red, 423 Foxall, R. A. One hundred years in perspective — the chang- ing roles and objectives of The Ottawa Field-Natural- ists’ Club, | ITY . Fragaria vesca, 129 Fraser, D. Aquatic feeding by a Woodchuck, 309 Fratercula corniculata, 84 Fraxinus americana, 32 Freddy, D.J. Distribution and movements of Selkirk Caribou, 1972-1974, 71 Freedman, B. and P.M. Catling. Movements of sym- patric species of snakes at Amherstburg, Ontario, 399 Freitag, R., 79 Frisch, T. and W. C. Morgan. Ivory Gull colonies in south- eastern Eilesmere Island, Arctic Canada, 173 Frog, Boreal Chorus, 144 Wood, 144 Frullania bolanderi, 140 eboracensis, 140 Galium aparine, 33, 129 boreale, 184 Brandegei, 261 circaezans, 33 pilosum, 35 triflorum, 169 Gallinago paraguaiae, 189 Gammarus, 267 Garter Snake, Common, predation on Ring-billed Gull chicks, 317 Gaultheria shallon, 126 Geranium maculatum, 33 richardsonii, 168 Geum aleppicum vat. strictum, 168 Glaucomys sabrinus, 320 Glossiphonia complanata, 310 Glyceria borealis, 388 Striata var. stricta, 166 Goodyera repens, 167 Goossen, J. P. and D. G. Busby. Occurrences of the Red Phalarope in the prairie provinces and adjacent states, 446 Gordon, D. M. New localities for the Northern Spring Salamander and the Four-toed Salamander in south- western Quebec, 193 Goski, B. C., 259 Gray, P. A., reviews by, 104, 108, 215, 339, 460 Green, R. H., reviews by, 456 Grosbeak, Evening, Xanthochroism in the, 66 Grosbeaks, Pine, Interspecific vocal mimicry by, 436 Grouse, Ruffed, 17 Sharp-tailed, 17 Gruchy, C., review by, 206 Gull, Ivory, Colonies in southeastern Ellesmere Island, Arctic Canada, 173 Gull, Lesser Black-backed, Larus fuscus, in Labrador waters, 445 Gull, Ring-billed, chicks, Common Garter Snake predation on, 317 Gulls, Herring, and Common Terns in northeastern Sas- katchewan, Colonial nesting, 132 Gunson, J. R., 48, 243 Gunson, J. R. and R. M. Nowak. Largest Gray Wolf skulls found in Alberta, 308 Gunson, J.R. and R.R. Bjorge. Winter denning of the Striped Skunk in Alberta, 252 Gyrinophilus porphyriticus porphyriticus, 193 INDEX TO VOLUME 93 469 Haber, E. Utricularia geminiscapa at Mer Bleue and range extensions in eastern Canada, 391 Habitat rehabilitation in Alaska, Response of wintering Moose to mechanical, 191 Habitat selection by wintering Snowy Owls (Nyctea scan- diaca), 176 Habitat use in Montana, Lynx movements and, 441 Habitat utilization and population densities of the amphibi- ans of northeastern Alberta, 144 Habitats of four annual smartweeds in Ontario, Distri- bution and, 378 Haematopus ostralegus bachmani, 188 oO. palliatus, 187 : Haliaeetus leucocephalus, 367, 443 Hall, I. V., L. E. Aalders, N. L. Nickerson, and S. P. Vander Kloet. The biological flora of Canada. 1. Vaccinium angustifolia Ait., Sweet Lowbush Blueberry, 415 Hanson, J. M. and S. U. Qadri. Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie in the Ottawa River, 232 Harder es D243 Harder, L. D. Winter feeding by Porcupines in montane forests of southwestern Alberta, 405 Hare, Snowshoe, 19, 441 Harms, V. L., review by, 213 Harris, R. N. Influence of weather on aggresion in Tree Swallows, 437 Hash, H.S., 441 Hawk, Red-tailed, 367 Swainson’s, 367 Hawks, Red-tailed, Demographic and dietary responses of, during a Snowshoe Hare fluctuation, 16 Hedysarum alpinum, 184 mackenzii, 184 Helleiner, C. W. Xanthochroism in the Evening Grosbeak, 66 Helobdella stagnalis, 310 triserialis, 310 Hemidactylium scutatum, 193 Heracleum mantegazzianum, Giant Cow Parsnip on Van- couver Island, British Columbia, 82 Herons, Great Blue, Abandoned windmill used as nesting site, 439 Hesperiphona vespertina, 66 Hexagenia sp., 267 Hieracium scabriusculum, 169 Hierochloe alpina, 355 pauciflora, 355 Himanotopus mexicanus, 68, 188 Hippurus vulgaris, 309, 389 Hirudinoidea, Dispersion of freshwater leeches, to Anticosti Island, Quebec, 310 Hirundo rustica, 374 Hoefs, M. Flowering plant phenology at Sheep Mountain, southwest Yukon Territory, 183 Hogan, G. G., 175 Hordeum jubatum, 184 vulgare, 285 Hornocker, M. G., 441 Houston, C. S., review by, 207 470 Hummingbird, Calliope, in Kananaskis Provincial Park, Alberta, Nesting of, 449 Hummingbird, Rufous, 450 Humphreys, G., 63 Hunt, H. M. Summer, autumn, and winter diets of Elk in Saskatchewan, 282 Hybrids of Cinnamon X Blue-winged Teal from southern Alberta, Probable, 316 Hylocomium splendens, 126 Hypericum perforatum, 34 Hypnum circinale, 129 hamulosum, 431 Tchthyomyzon fossor, First record of the Northern Brook Lamprey, in Nelson River drainage, Manitoba, 199 Ictiobus niger, 304 Interaction between a Long-tailed Weasel and a Snowy Owl, 67 Interactions between Snowy and Short-eared Owls in winter, 41] Ireland, R. R. and G. B. Bellolio-Trucco. Mosses new to Ontario and Quebec, 431 Tridoprocne bicolor, 372, 437 Is biology unknown? (letters), 330 Tsoetes macrospora, 388 Tsothecium stoloniferum, 129 IUCN, Books available from, 87 IUCN views on whale management, 88 Jaagumagi, R., review by, 209 Jacana spinosa, 188 Jenkins, R. E., 63 Juglans nigra, 34 Juncus arcticus spp. alaskanus, 262 arcticus spp. ater, sensu, 167 balticus var. alaskanus, 262 balticus var. littoralis, 167 militaris, 389 pelocarpus, 388 Juniperus virginiana, 34 Jyrkkanen, J. and D.G. Wright. First record of the Northern Brook Lamprey, /chthyomyzon fossor, in the Nelson River drainage, Manitoba, 199 eno, 1, 1.5 NG Kennedy, A. J., review by, 218 Kestrel, American, Intraspecific food theft by the, 198 Katies be 187 King, L. L., review by, 344 Kirkland, C. J., 195 Kirkland, G.L., Jr. and C.M. Knipe. The Rock Vole (Microtus chrotorrhinus) as a Transition zone species, 319 Kirkland, G. L., Jr., D. F. Schmidt, and C. J. Kirkland. First record of the Long-tailed Shrew (Sorex dispar) in New Brunswick, 195 Klein, H. G., 239 Knipe, C. M., 319 Kobresia myosuroides, 355 simpliciuscula, 355 Koehler, G.M., M. N. Hornocker, and H. S. Hash. Lynx movements and habitat use in Montana, 441 Kott, E.. R. E. Jenkins, and G. Humphreys. Recent collec- tions of the Black Redhorse, Moxostoma duquesnei, from Ontario, 63 THE CANADIAN FIELD-NATURALIST Vol. 93 Labrador, Two-lined Salamander, Eurycea bislineata, in, 178 Labrador waters, Lesser Black-backed Gull, Larus fuscus, in, 445 Lamprey, Northern Brook, Jchthyomyzon fossor, in the Nelson River drainage, Manitoba, First record of the, 199 Laporte, P., review by, 92 Lappula myosotis, 184 Larus argentatus, 132 delawarensis, 317 fuscus graellsii, 446 Larus fuscus, Lesser Black-backed Gull, waters, 445 Lathyrus ochroleucus, 284 venosus, 284 each erates |79 Lecanora conizaea, 140 impudens, 140 Ledum decumbens, 355 groenlandicum, 264 palustre, 184 palustre ssp. groenlandicum, 264 Leeches, freshwater (Hirudinoidea), Dispersion of, to Anti- costi Island, Quebec, 310 Lein, M.R. Song pattern of the Cypress Hills popula- tion of White-crowned Sparrows, 272 Lein, M. R. and G. A. Webber. Habitat selection by winter- ing Snowy Owls (Nyctea scandiaca), 176 Lein, M.R. and P. C. Boxall. Interactions between Snowy and Short-eared Owls in winter, 411 Lemna trisulca, 262 Lepidozia reptans, 129 Lepraria membranacea, 140 neglecta, 140 Leptodora kindtii, 267 Lepus americanus, 16, 441 Lespedeza capitata, 35 intermedia, 35 Lewin, V., 144 Lien, J., 303 Life history characteristics of Little Brown Bats (Myotis lucifugus) in Alberta, 243 Limosa fedoa, 188 Lindsay, J. E., review by, 458 Limnaea borealis, 184 borealis ssp. americana, 264 Linum perenne, 184 virginiana, 35 Lister, R., 68 Lister, Robert, receives award, 337 Listera cordata, 129 Lloyd, Hoyes, 1888-1978, Tribute to, 331 Lobaria pulmonaria, 140 Lobelia dortmanna, 388 Lophocolea bidentata, 129 Lunda cirrhata, 84 Lupinus arcticus, 184 Luzula arctica, 262 arcuata s. lat., 262 confusa, 355 multiflora s. lat., 262 in Labrador IQ) nivalis, 262 parviflora, 262 Wahlenbergii ssp. Wahlenbergii, 261 Lycopodium annotinum ssp. pungens, 261 complanatum, 261 Lynx canadensis, 441 Lynx movements and habitat use in Montana, 441 MacCrimmon, H. R. and R. W. Pugsley. Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, Ontario, 266 Mackenzie, District of, Range extensions of vascular plants in northern Yukon Territory and northwestern, 259 Mahon, R., review by, 349 Maianthemum canadense, 33 Mallard, Wild, stocking in a large marsh habitat, 55 Manitoba, 55, 75, 446 Manitoba, First record of the Northern Brook Lamprey, Ichthyomyzon fossor, in the Nelson River drainage, 199 Marmota monax, 309 Martell, A. M. Relative efficiencies of Museum Special, Victor, and Holdfast traps for sampling small mammal populations, 313 Martin, K. Common Garter Snake predation on Robin nestlings, 70 Martin, Purple, 371 Matteuccia struthiopteris var. pensylvanica, 163 Maycock, p., 28 McAllister, D. E., review by, 208 McDonald, H. G., review by, 454 McKeating, G. B., review by, 210 McKelvey, R. W. Swans wintering on Vancouver Island, 1977-1978, 433 MecNicholl, M. K. Communal roosting of Song Sparrows under snowbank, 325 MeNicholl, M. K., reviews by, 98, 455 McNicholl, M. K., Robert Lister receives award, 337 MeNicholl, M. K. and G. G. Hogan. Wind-caused death of Great Cormorant, 175 Meadow-parsnip, Thaspium trifoliatum, in Canada, 306 Medicago sativa, 285 Meeting, Annual Business, Notice of The Ottawa Field- Naturalists’ Club, 452 Melospiza melodia, 325 Mephitis mephitis, 252 Merdsoy, B., J. Lien, and A. Storey. Extralimital record of a Narwhal (Monodon monoceros) in Hall’s Bay, New- foundland, 303 Mertensia paniculata, 184 Michener, G. R. Yearly variations in the population dynam- ics of Richardson’s Ground Squirrels, 363 Mickelson, P. G., 171 Micropalama himantopus, 188 Microsorex hoyi, 315 Microtus chrotorrhinus, 314 pennsylvanicus, 18, 315, 320, 412 Microtus chrotorrhinus, The Rock Vole, as a Transition zone species, 319 Miller, G.E. and H.M. Dale. Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka District, Ontario from’ 1953 to 1977, 386 Mimicry by Pine Grosbeaks, Interspecific vocal, 436 INDEX TO VOLUME 93 471 Mimulus guttatus, 168 Mink, 276 Minuartia dawsonensis, 167 groenlandica, 355 Mites, Examination of overwintering carabid beetles for associated, 79 Mnium cuspidatum, 140 Monarda fistulosa, 35 Moneses uniflora, 264 Monodon monoceros, Narwhal, in Hall’s Bay, Newfound- land,Extralimital record of a, 303 Montana, Lynx movements and habitat use in, 44] Moore, M. I., review by, 346 Mooreobdella fervida, 310 Moose, Response of wintering, to mechanical habitat rehabilitation in Alaska, 191 Morgan, W. C., 173 Morrison, M. L. and L. F. Kiff. Eggshell thickness in Ameri- can shorebirds before and since DDT, 187 Mosses new to Ontario and Quebec, 431 Mouse, Deer, 239, 314, 320, 412 Meadow Jumping, 314, 320 White-footed, 239 Woodland Jumping, 239, 320 Movements and habitat use in Montana, Lynx, 441 Movements of Selkirk Caribou, 1972-1974, Distribution and, 71 Movements of sympatric species of snakes at Amherst- burg, Ontario, 399 Moxostoma duquesnei, Recent collections of the Black Redhorse from Ontario, 63 Moxostoma erythrurum, 63 macrolepidotum, 64 Muhlenbergia mexicana, 163 Munro, D.A., Tribute to Hoyes Lloyd, 1888-1978, 331 Murphy, D., review by, 453 Muskrat, 276 Muskrat in southeastern New Brunswick, Unusually late pregnancy of a, 440 Mustela frenata, 67, 365, 412 nivalis, 68 vison, 276 Myosotis alpestris, 184 Myotis keenti, 247 lucifugus lucifugus, 243 Myotis lucifugus, Little Brown Bats, Life history char- acteristics, in Alberta, 243 Myriophyllum farwelli, 389 tenellum, 389 Myurella tenerrima, 431 Nagy, J. A., A. M. Pearson, B. C. Goski, and W. J. Cody. Range extensions of vascular plants in northern Yukon Territory and northwestern District of Mackenzie, 259 Najas flexilis, 388 Napaeozapus insignis, 315, 320 Narwhal (Monodon monoceros) in Hali’s Bay, Newfound- land, Extralimital record of a, 303 Nepszy, S. J., 304 Nesting and food-storage behavior of Peromyscus mani- culatus gracilis and P. leucopus noveboracensis, 239 472 Nesting, biology and development of young in Ontario Black Terns, 276 Nesting of Horned Puffins in British Columbia, 84 Nesting of the Calliope Hummingbird in Kananaskis Pro- vincial Park, Alberta, 449 Nesting site, Abandoned windmill used as a, by Great Blue Herons, 439 Nesting sites, potential, and populations of swallows in Canada, Man’s influence on, 371 Nettleship, D. N., review by, 95 New Brunswick, 437 New Brunswick, Decline of summering Bald Eagles in central, 443 New Brunswick, First record of the Long-tailed Shrew in, 195 New Brunswick, Unusually late pregnancy of a Muskrat in southeastern, 440 New York, 239, 319 Newfoundland, Extralimital record of a Narwhal (Monodon monoceros) in Hall’s Bay, 303 Newt, Red-spotted, 195 Nickerson, N. L., 415 Norstrom, R. J., review by, 460 Northcott, T. H., review by, 342 Northwest Territories, 81 Northwest Territories, Brewer’s Blackbird breeding in the, 76 Northwest Territories, Snowy Egret in the, 329 Notophthalmus viridescens, 195 Notropis atherinoides, 266 Nova Scotia, 67 Nova Scotia, Status of Eastern White Cedar, occidentalis, in western, 326 Nowak, R. M., 308 Numenius americanus, 188 phaeopus, 188 Nuphar variegatum, 309, 389 Nyctea scandiaca, 67, 411 Nyctea scandiaca, Snowy Owls, Habitat selection by winter- ing, 176 Nycticryphes semicollaris, 189 Nymphaea ordorata, 389 Nymphoides cordatum, 389 Odocoileus hemionus columbianus, 130 Olor buccinator, 433 Olynyk, J. E.and R. Freitag. Examination of overwintering adult carabid beetles for associated mites, 79 Ondatra zibethicus, 276 z. zibethicus, 440 Ontario, 28, 79, 198, 306, 309 Ontario and Quebec, Mosses new to, 431 Ontario Black Terns, Nesting biology and development of young in, 276 Ontario, Distribution and habitats of four annual smart- weeds in, 378 Ontario, Eastern limit of the Five-lined Skink, Eumeces fasciatus, in, 321 Ontario, Epiphytes on White Elm, U/mus americana, near Thunder Bay, 139 Ontario, Food and feeding of the Rainbow Smelt (Osmerus mordax) in Lake Simcoe, 266 Thuja THE CANADIAN FIELD-NATURALIST Vol. 93 Ontario from 1953 to 1977, Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka District, 386 Ontario, Movements of sympatric species of snakes at Amherstburg, 399 Ontario, Recent collections of the Black Redhorse, Moxo- stoma duquesnei, from, 63 Ontario, Some new and interesting vascular plant records from northern, 355 Ontario stream, Earthworm cocoons as a drift component in a southern, 180 Oriskany sandstone outcrop and associated natural features, a unique occurrence in Canada, The, 28 Ornithologists, Societies and amateur, 88 Orthothecium strictum, 431 Orthotrichum obtusifolium, 140 speciosum, 140 strangulatum fo. lescurii, 432 Oryzopsis asperifolia, 163 Osmerus mordax in Lake Simcoe, Ontario, Food and feed- ing of the Rainbow Smelt, 266 Osmorhiza depauperata, 163 obtusa, 168 Ostrya virginiana, 34 Ottawa Field-Naturalists’ Club Annual Business Meeting of The, 452 Ottawa Field-Naturalists’ Club Auditor’s report, 226 Balance sheet, 227 Minutes of the ninety-ninth annual business meeting, DDB} Report of council, 224 Statement of income and expenditure, C.F.N., 229 Statement of income and expenditure, O.F.N.C., 228 Ottawa Field-Naturalists’ Club, Call for nominations for the council of The, 353 Ottawa Field-Naturalists’ Club. Notice of change to the by- laws of The, 337 Ottawa Field-Naturalists’ Club, Notice of motion to amend the constitution of The, 338 Ottawa Field-Naturalists’ Club, One hundred years in perspective — the changing roles and objectives of, The, | Ottawa River, Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie in the, 232 Ouellet, H., reviews by, 210, 340 Owl, Snowy, Interaction between a Long-tailed Weasel and a, 67 Owls, Barn, in Quebec, 323 Owls, Snowy and Short-eared, in winter, Interactions between, 411 Owls, Snowy, (Nyctea scandiaca), Habitat selection by wintering, 176 Oxytropis borealis, 264 deflexa var. foliolosa, 168 deflexa var. sericea, 264 glutinosa, 264 huddelsonii, 184 nigrescens ssp. pygmaea, 264 pygmaea, 264 viscida, 184 NTS Oystercatcher, “American”, 189 “black”, 189 Pagophila eburnea, 173 Paleoclimate, Natural fires as an index of, 116 Panicum lanuginosum var. praecocius, 36 linearifolium, 35 Papaver Hultenii, 263 Keelei, 263 Macounii, 263 Parietaria pensylvanica, 163 Parker, G. R. Unusually late pregnancy of a Muskrat in southeastern New Brunswick, 440 Parmelia saxatilis, 140 sulcata, 140 Parsnip, Giant Cow, ( Heraclium mantegazzianum) on Van- couver Island, British Columbia, 82 Partridge, Gray, 412 Paton, D. G., 449 Pearson, A. M., 259 Pedicularis flammea, 361 labradorica, 361 lapponica, 264, 361 spp., 186 sudetica, 361 Pedioecetes phasianellus, 18, 74 Pellaea atropurpurea, 288 occidentalis, 288 suksdorfiana, 288 Pellaea (Polypodiaceae) in Alberta, Taxonomy, distribu- tion, and ecology of the cliff-brake ferns, 288 Pentaneuris monilis, 267 Perdix perdix, 412 Peromyscus maniculatus, 314, 320, 412 Peromyscus maniculatus gracilis and P. leucopus nove- boracensis, Nesting and food-storage behavior of, 239 Petrochelidon pyrrhonota, 373 Phalacrocorax carbo, 175 Phalaris arundinacea, 166 Phalarope, Red, in the prairie provinces and adjacent states, Occurrences of the, 446 Phalaropus fulicarius, 188, 446 lobatus, 188 tricolor, 188 Phenacomys intermedius, 314 Phyllodoce coerulea, 355 — Physalis heterophylla, 35 Physcia adscendens, 140 aipolia, 140 orbicularis, 140 Physconia grisea, 140 Picea glauca, 405 X engelmanii, 405 Pimlott, The Douglas, scholarship in environmental studies, 338 Pinguicula villosa, 264, 355 Pinicola enucleator, 436 Pinus banksiana, 282 flexilis, 405 strobus, 34 Plagiothecium denticulatum, 140 Plantago canescens, 184 | Platydicta subtile, 140 INDEX TO VOLUME 93 473 Plethodon cinereus cinereus, 194 Pluvialis dominica, 188 squatarola, 188 Poa ammophila, 163 glauca, 184 Polygala polygama, 35 verticillata, 35 Polygonatum biflorum, 36 Polygonum amphibium, 389 amphibium var. stipulaceum, 167 lapathifolium var. lapathifolium, 378 lapathifolium var. prostratum, 379 lapathifolium var. salicifolium, 378 pensylvanicum var. durum, 379 pensylvanicum vat. laevigatum, 378 pensylvanicum var. pensylvanicum, 378 persicaria var persicaria, 379 persicaria var. ruderale, 379 scabrum, 378 Polypodiaceae, in Alberta, Taxonomy, distribution, and ecology of the cliff-brake ferns (Pel/aea), 288 Polystichum munitum, 128 Pomoxis nigromaculatus, 232 Pontederia cordata, 389 Population densities of the amphibians of northeastern Alberta, Habitat utilization and, 144 Populations of swallows in Canada, Man’s influence on potential nesting sites and, 371 Populus and Salix seedfall, A trap to measure, 77 Populus balsamifera, 77 grandidentata, 34 tremuloides, 34, 77, 282 Porcupines in montane forests of southwestern Alberta, Winter feeding by, 405 Porella platyphylla, 140 Potamogeton alpinus ssp. tenuifolius, 262 amplifolius, 388 berchtoldii, 389 epthydrus, 389 gramineus, 389 natans, 389 oakesianus, 389 obtusifolius, 388 pectinatus, 389 perfoliatus ssp. Richardsonii, 262 Richardsonii, 262, 389 robbinsii, 389 spirillus, 389 zosteriformis, 309 Potentilla elegans, 264 fruticosa, 184 hookeriana, 184 Hookeriana ssp. Hookeriana, 264 hyparctica, 264 nivea ssp. Hookeriana, 264 rubricaulis, 261 simplex, 34 Powers, K. D. Lesser Black-backed Gull, Larus fuscus, in Labrador waters, 445 Predation on Ring-billed Gull chicks, Common Garter Snake, 317 Predation on Robin nestlings, Common Garter Snake, 70 474 President’s message, | Preston, J., 178 Preston, W. B., reviews by, 94, 98 Prevett, J. P., review by, 341 Prince Edward Island, 175, 325 Prionocera sp., 171 Procyon lotor, 423 Progne subis, 371 Proposals invited from field FessEe investigators, 201 Prunus americana, 36 cerasus, 34 pensylvanica, 284 serotina, 32 virginiana, 33, 284 Pseudacris triseriata maculata, 144 Pseudoleskeela catenulata, 431 Pseudotsuga menziesii, 126 menziesii var. glauca, 405 Pteridium aquilinum, 33, 128 Pteritis nodulosa, 166 Pterostichus adstrictus, 80 pensylvanicus, 80 Puffin, Horned, Nesting of, in British Columbia, 84 Puffin, Tufted, 84 Pugsley, R. W., 266 Pulsatilla patens, 184 Pylaisiella polyantha, 140 selwynii, 140 Plyodictis olivaris, 179 Pyrola secunda ssp. obtusata, 264 Pyrus coronaria, 33 Qadni, S. U., 232 Quebec, 439 Quebec, Barn Owls in, 323 Quebec, Dispersion of freshwater leeches (Hirudinoidea) to Anticosti Island, 310 Quebec, Mosses new to Ontario and, 431 Quebec, New localities for the Northern Spring Salamander and the Four-toed Salamander in southwestern, 193 Quercus alba, 32 bicolor, 35 macrocarpa, 34 muehlenbergii, 34 rubra, 32 velutina, 32 Quinlan, R. W. Snowy Egret in the Northwest Territories, 329 Raccoon, 423 Rana sylvatica, 144 Range extensions in eastern Canada, niscapa at Mer Bleue and, 391 Range extensions of vascular plants in northern Yukon Ter- ritory and northwestern District of Mackenzie, 259 Rangifer tarandus caribou, 71 t. granti, 155 Ranunculus abortivus, 167 Eschscholtzii, 263 hispidus, 36 reptans, 389 Recurvirostra americana, 68, 188 Redhorse, Black, Moxostoma duquesnei, from Ontario, Recent collections of the, 63 Utricularia gemi- THE-CANADIAN FIELD-NATURALIST Vol. 93 Redhorse, Golden, 63 Shorthead, 64 Renouf, D., review by, 342 Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta, 48 Request for information — shore bird color-marking, 338 Request for participants — International Shorebird Surveys IGA), 33s Response of wintering Moose to mechanical habitat re- habilitation in Alaska, 191 Responses, Demographic and dietary, of Red-tailed Hawks during a Snowshoe Hare fluctuation, 16 Rhizomnium glabrescens, 129 Rhus typhina, 34 Rhytidiadelphus triquetrus, 129 Ribes triste, 263 Riley, J. L. Some new and interesting vascular plant records from northern Ontario, 355 Ring counts in Salix arctica from northern Ellesmere Island, 81 Ringius, G.S. Status of Eastern White Cedar, occidentalis, in western Nova Scotia, 326 Riotte, J. C. E., review by, 203 Riparia riparia, 373 Ritchie, J. C., 458 Robarts, R. D., review by, 459 Roberts, W. and V. Lewin. Habitat utilization and popu- lation densities of the amphibians of northeastern Alberta, 144 Robin, American, 423 Robin, Common Garter Snake aredanon on nestlings, 70 Robitaille, G., reviews by, 217, 218 Roby, D: D., 155 Roosting, Communal, of Song Sparrows under snowbank, B25 Rosa acicularis, 184 gymnocarpa, 128 spp., 284 Rowe, J. S., Is biology unknown? (letter), 330 Rubus acaulis, 264 arcticus ssp. acaulis, 264 strigosus, 284 Sagittaria graminea, 388 sp., 309 Salamander, Blue-spotted, 194 Red-backed, 194 Spotted, 194 Northern Dusky, 193 Northern Two-lined, 194 Salamander, Two-lined, Eurycea bislineata, in Labrador, 178 Salamander, Four-toed, in southwestern Quebec, New localities for the Northern Spring Salamander and the, 193 Salamander, Northern Spring, and the Four-toed Sala- mander in southwestern Quebec, New localities for the, 193 Salix alaxensis, 78 alaxensis var. longistylis, 167 arbusculoides, 79 Barrattiana, 167 bebbiana, 79 Thuja 979: interior, 78 lasiandra, 79 longistylis, 167 monticola, 78 novae-anglicae, 78 padophylla, 167 planifolia, 79 spp., 79, 282 Salix arctica from northern Ellesmere Island, Ring counts in, 81 Salix seedfall, A trap to measure Populus and, 77 Sanguisorba sitchensis, 168 stipulata, sensu, 168 Sarcophaga citellivora, 366 Saskatchewan, Colonial-nesting Herring Gulls and Com- mon Terns in northeastern, 132 Saskatchewan, Summer, autumn, and winter diets of Elk in, 282 Savile, D. B. O. Ring counts in Salix arctica from northern Ellesmere Island, 81 Saxifraga caespitosa, 263 hieracifolia, 263 oppositifolia, 184 tricuspidata, 184 Scapania nemorosa, 129 Schizachne purpurascens, 166 Schmidt, D. F., 195 Scholarship, The Douglas Pimlott, in environmental studies, 338 Schowalter, D. B. and J. R. Gunson. Reproductive biology of the Big Brown Bat (Eptesicus fuscus) in Alberta, 48 Schowalter, D. B., J. R. Gunson, and L. D. Harder. Life history characteristics of Little Brown Bats (Myotis lucifugus) in Alberta, 243 Schwert, D. P. and K. W. Dance. Earthworm cocoons as a drift component in a southern Ontario stream, 180 Scirpus subterminalis, 389 Scotter, G. W. and W. J. Cody. Interesting vascular plants from southeastern Yukon Territory, 163 Scutellaria galericulata var. pubescens, 168 Sealy, S.G., 84 Sedum rosea, 184 Selaginella selaginoides, 166 Selasphorus rufus, 450 Selkirk Mountains, 71 Senecio atropurpureus ssp. tomentosus, 265 Kjellmannii, 265 Senner, S. E. and P. G. Mickelson. Fall foods of Common Snipe on the Copper River Delta, Alaska, 171 Sexton, D. A. and M. M. Gillespie. Effects of fire on the location of a Sharp-tailed Grouse arena, 74 Sharp-tailed Grouse arena, Effects of fire on the location of a, 74 Shiner, Emerald, 266 Shorebird color-marking, Request for information, 338 Shorebird Surveys, International 1979, Request for par- ticipants, 338 Shorebirds before and since DDT, Eggshell thickness in American, 187 Shrew, Long-tailed, 320 INDEX TO VOLUME 93 475 Masked, 314, 320 Short-tailed, 320 Smokey, 320 Water, 320 Shrew, Long-tailed, (Sorex dispar) in New Brunswick, First record of the, 195 Sigman, M. Response of wintering Moose to mechanical habitat rehabilitation in Alaska, 191 Silene acaulis, 184 acaulis ssp. subacaulescens, 263 menziesii, 167 Sims, R. A., 139 Skink, Five-lined, Ewmesces fasciatus, in Ontario, Eastern limit of the, 321 Skunk, Striped, in Alberta, Winter denning of, 252 Smartweeds in Ontario, Distribution and habitats of four annual, 378 Smelt, Rainbow, (Osmerus mordax) in Lake Simcoe, Ontario, Food and feeding of the, 266 Smilacina racemosa, 184 trifolia, 167 Smith, L. C. Article or note? 337 Smith, L. C. Editor’s report for 1978, 201 Smith, L. C. Our responsibilities as field-naturalists and biologists (editorial), 113 ; Smith, L. C. Special appreciation expressed to R. Emerson Whiting, 87 Smith, L. C. The Canadian Field-Naturalist — the status quo or a new direction? (editorial), 10 Smith, L. C. To a bigot (editorial), 231 Smith, W. T., 155 Snake, Black Rat, 28, 322 Brown, 399 Butler’s Garter, 399 Common Garter, 70, 317 Eastern Fox, 399 Eastern Garter, 399 Northern Ribbon, 322 Snake, Common Garter, predation on Robin nestlings, 70 Snakes at Amhertsburg, Ontario, Movements of sympatric species of, 399 Snipe, Common, Fall foods of, on the Copper River Delta, Alaska, 171 Snowshoe Hare fluctuation, Demographic and dietary responses of Red-tailed Hawks during, 16 Solidago caesia, 33 canadensis, 35 canadensis var. salebrosa, 169 juncea, 35 multiradiata, 184 nemoralis, 35 Sonchus asper, 284 Song pattern of the Cypress Hills population of White- crowned Sparrows, 272 Sorex arcticus, 315 cinereus, 314, 320 dispar, 320 fumeus, 320 palustris, 315, 320 Sorex dispar, First record of the Long-tailed Shrew, in New Brunswick, 195 476 Sparganium angustifolium, 388 fluctuans, 389 minimum, 262 Sparrows, Song, under snowbank, Communal roosting of, 395 Sparrows, White-crowned, Song pattern of the Cypress Hills population of, 272 Spermophilus franklinii, 20 richardsonii, 17, 363, 412 Sphaeroderus nitidicollis, 80 Sphenopholis intermedia, 163 Spiranthes Romanzoffiana, 263 Squirrel, Franklin’s Ground, 20 Northern Flying, 320 Red, 320 Richardson’s Ground, 17, 19, 412 Squirrels, Richardson’s Ground, Yearly variations in the population dynamics of, 363 Staniforth, R. J. and P. B. Cavers. Distribution and habitats of four annual smartweeds in Ontario, 378 Stelfox, H. A. and G. J. Brewster. Colonial-nesting Herring Gulls and Common Terns in northeastern Saskatche- wan, 132 Stelgidopteryx ruficollis, 372 Stellula calliope, 449 Stepney, P. H.R. Brewer’s Blackbird breeding in the Northwest Territories, 76 Sterna hirundo, 132 Sternotherus odoratus, 322 Stilts, Black-necked, nesting near Edmonton, Alberta, 68 Stinkpot, 322 Stocek, R. F. Decline of summering Bald Eagles in central New Brunswick, 443 Storeria dekayi, 399 Storey, A., 303 Sullivan, M.G. Blue Grouse brood hen — Black Bear confrontation, 200 Sullivan, T. P. Virgin Douglas Fir forest on Saturna Island, British Columbia, 126 Swallow, Bank, 373 Barn, 374 Cliff, 373 Rough-winged, 372 dinees 372 Violet-green, 372 Swallows in Canada, Man’s influence on potential nesting sites and populations of, 371 Swallows, Tree, Influence of weather on aggression in, 437 Swan, Mute, 433 Trumpeter, 433 Swans wintering on Vancouver Island, 1977-1978, 433 Swertia caroliniensis, 36 Systematic and evolutionary biology, Second International congress of, 87 Tachycineta thalassina, 372 Tadlock, C. G. and H. G. Klein. Nesting and food-storage behavior of Peromyscus maniculatus gracilis and P. leucopus noveboracensis, 239 Tamias striatus, 315, 320 Tamiasciurus hudsonicus, 320 Taraxacum officinale, 184 Phymatocarpum, 265 THE CANADIAN FIELD-NATURALIST Vol. 93 Taxidea taxus, 365 Taxonomy, distribution, and ecology of the cliff-brake ferns (Pellaea: Polypodiaceae) in Alberta, 288 Taylor, P. Interspecific vocal mimicry by Pine Grosbeaks, 436 Teal, Blue-winged, 316 Cinnamon, 316 Teal, Cinnamon X Blue-winged, from southern Alberta, Probable hybrids of, 316 Tendipes tentans, 267 Terasmae, J. and N. C. Weeks. Natural fires as an index of paleoclimate, 116 Terns, Black, Nesting biology and development of young in Ontario, 276 Terns, Common, in northeastern Saskatchewan, Colonial- nesting Herring Gulls and, 132 Thalictrum sparsiflorum var. richardsonii, 167 Thamnophis butleri, 399 Sauritus septentrionalis, 322 sirtalis, 70, 317 sirtalis sirtalis, 399 Thaspium barbinode, 36 Thaspium trifoliatum (Meadow-parsnip) in Canada, 306 Thompson, I. D., review by, 341 Thomson, S. C., review by, 101 Thormin, T. W., 68 Thuja occidentalis, in western Nova Scotia, Status of Eastern White Cedar, 326 Thuja plicata, 126 Tilia americana, 34 To a bigot (editorial), 231 To know ourselves, 89 Toad, Canadian, 144 Todd, A. W., 16 Townsendia hookeri, 184 Trans-Alaska Pipeline, Caribou distribution and group composition associated with construction of the, 155 Trientalis latifolia, 129 Triticum aestivum, 285 Tsuga heterophylla, 126 Turdus migratorius, 70, 423 Tyto alba, 323 Ulmus americana, 34 Ulmus americana, Epiphytes on White Elm near Thunder Bay, Ontario, 139 Urquhart, F. A. and N. R. Urquhart. Breeding areas and overnight roosting locations in the northern range of the Monarch Butterfly (Danaus plexippus plexippus) with a summary of associated migration routes, 41 Urquhart, N. R., 41 Ursus americanus, 200, 423 Urtica gracilis, 167 Ussher, R. D. and F. R. Cook. Eastern limit of the Five- lined skink, Eumesces fasciatus, 1n Ontario, 321 Utricularia cornuta, 388 gibba, 389 intermedia, 388 purpurea, 388 resupinata, 388 vulgaris, 264, 389 Utricularia geminiscapa at Mer Bleue and range extensions in eastern Canada, 391 IVE Vaccinium angustifolium forma angustifolium, 416 angustifolium forma leucocarpum, 416 angustifolium forma nigrum, 416 pallidum, 36 vacillans, 36 vitis-idaea, 184 Vahlodea atropurpurea, 355 Vallisneria americana, 389 Vancouver Island, 1977-1978, Swans wintering on, 433 Vander Kloet, S. P., 415 Vanellus chilensis, 189 Vascular plant records from northern Ontario, Some new and interesting, 355 Vascular plants from southeastern Yukon Territory, Inter- esting, 163 Vascular plants in northern Yukon Territory and north- western District of Mackenzie, Range extensions of, 259 Viburnum acerifolium, 33 edule, 184 trilobum, 284 Vicia americana, 163, 284 Viola langsdorffii, 168 pedata var. lineariloba, 36 rugulosa, 163 Vole, Heather, 314 Meadow, 18, 320, 412 Red-backed, 314, 320 Rock, 314, 320 Vole, Rock (Microtus chrotorrhinus) as a Transition zone species, The, 319 Vulpes vulpes, 423 Wapiti, 282 Weasel, Long-tailed, 365, 412 Weasel, Long-tailed, and Snowy Owl, Interaction between a, 67 INDEX TO VOLUME 93 477 Weather, Influence of, on aggression in Tree Swallows, 437 Webber, G. A., 176 Weber, W. C., reviews by, 91, 203 Weeks, N. C., 116 Weseloh, D. V., 68 Weseloh, D. V. and L. M. Weseloh. Probable hybrids of Cinnamon X Blue-winged Teal from southern Alberta, 316 Weseloh, D. V., review by, 96 Weseloh, L. M., 68, 316 Whale management, IUCN views on, 88 White, E. R., 82 Whiting, R. Emerson, Special appreciation expressed to, 87 Whitten, K R., 155 Wildlife film festival, second annual international, 90 Winder, G., 28 Wolf, Gray, skulls found in Alberta, Largest, 308 Woodchuck, Aquatic feeding by a, 309 Wright, D. G., 199 Xanthochroism in the Evening Grosbeak, 66 Xanthoria polycarpa, 140 Young, J. W.S. review by, 348 Yukon Territory and northwestern District of Mackenzie, Range extensions of vascular plants in northern, 259 Yukon Territory, Flowering plant phenology at Sheep Mountain, southwest, 183 Yukon Territory, Interesting vascular plants from south- eastern, 163 Zammuto, R. M., review by, 461 Zapus hudsonius, 314, 320 Zasada, J.C. and R. Densmore. A trap to measure Populus and Salix seedfall, 77 Zizania aquatica, 389 Zonotrichia leucophrys oriantha, 272 Zygadenus elegans, 184 Index to Book Reviews Botany Arditti, J. (ed.). Orchid biology: reviews and perspectives, 1., 104 Argus, G. W. and D. J. White. The rare vascular plants of Ontario/Les plantes vasculaires rares de Ontario, 100 Bassett, I. J. An atlas of airborne pollen grains and common spores of Canada, 457 Brodie, H. J. Fungi: delight of curiosity, 101 Brunton, D. F. Flora of Alberta: a checklist, 213 Cody, W. J. Ferns of the Ottawa district, 345 Dorn, R. D. Manual of vascular plants of Wyoming, 211 Good, R.E., D. F. Wigham, R. L. Simpson, and C. G. Jackson, Jr. (eds.). Freshwater wetlands: ecological processes and management potential, 213 Harrington. H. D. How to identify grasses and grasslike plants (sedges and rushes), 212 Markgraf, V.and H. L. D’Antoni. Pollen flora of Argentina: modern spore and pollen types of Pteridophyta, Gymn- nospermae, and Angiospermae, 102 McGrath, J. W. Dyes from lichens and plants: a Canadian dyer’s guide, 346 Muenscher, W. C. and M. A. Rice. Garden spice and wild pot-herbs: an American herbal, 212 Nilsson, S. Atlas of airborne pollen grains and spores in northern Europe, 103 Shetler, S. G. and L. E. Skog (eds.). A provisional checklist of species for Flora North America (revised), 212 Smith, J. P., Jr. Vascular plant families, 457 Szezawinski, A. F. and N. J. Turner. Edible garden weeds of Canada, 344 Zander, R.H. Floristics and environmental planning in western New York and adjacent Ontario: distribution of legally protected plants and plant sanctuaries, 102 Environment Bow Valley Naturalists. Vermillion Lakes, Banff National Park: an introductory study, 347 Button, J. The Shetland way of oil: reactions of a small community to big business, 105 Colinvaux, P. Why big fierce animals are rare: an ecologist’s perspective, 108 478 Dansereau, P. EZAIM: écologie de la zone de l’aéroport international de Montréal — le cadre d’une recherche écologique interdisciplinaire, 106 Girard, C. M. and M.C. Applications de la télédétection a l'étude de la biosphere, 217 Jurant, M., J. L. Bélair, V. Gerardin, and J. P. Ducrue. L’inventaire du capital nature: méthode de classification et de cartographie écologique du territoire (3 éme approximation), 216 Kinkead, E. Wildness is all around us: urban naturalist, 217 Long, D. R. (ed.). Outdoors Canada: a unique and practical guide to our wilderness and wildlife, 107 McMillan, V. and B. Nature quizzes for Canadians, 104 Pruitt, W. O. Boreal ecology, 215 Ramel, C. (ed.). Chlorinated phenoxy acids and their dioxins, 460 Russell, D. A. A vanished world: the dinosaurs of western Canada, 458 Shelford, V. E. The ecology of North America, 460 Statistics Canada. Human activity and the environment, 347 Wall, G. and C. Wright. The environmental impact of out- door recreation, 215 Werger, M.J..A (ed.). Biogeography and ecology of southern Africa, 459 notes of an Zoology Beebe, F. L. Hawks, falcons and falconry, 205 Blokpoel, H. Bird hazards to aircraft: problem and pre- vention of bird/ aircraft collision, 96 Brownie, C., D. R. Anderson, K. P. Burnham, and D.S. Robson. Statistical inference from band recovery data: a handbook, 455 Bruemmer, F. Life of the Harp Seal, 342 Cayouette, R. Nichoirs d’oiseaux, 210 Clark, J. L. The great arc of the wild sheep, 339 Comstock, A. B. Ways of the six-footed, 94 deSchauensee, R. M. and W. H. Phelps, Jr. A guide to the birds of Venezuela, 340 Eriksson, P. S. The bird finder’s 3-year note book, 99 Erskine, A. J. Birds in boreal Canada, 207 Ferguson, D. C. The moths of America north of Mexico, including Greenland. Fascicle 22.2, Noctuidae (in part): Lymantriidae, 203 Goodwin, D. Birds of Man’s world, 454 Goodwin, D. Crows of the world, 342 - Grainger, D. C. Animals in peril: a guide to the endangered species of Canada and the United States, 339 THE CANADIAN FIELD-NATURALIST Vol. 93 Grassé, P. P. Précis de zoologie: vertébrés. 3 — repro- duction, biologie, évolution et systématique. Oiseaux et mammiféres, 94 Harington, C. R. Quaternary vertebrate faunas of Canada and Alaska and their suggested chronological sequence, 453 Kelley, A. H. Birds of southeastern Michigan and south- western Ontario, 95 Mackenzie, J. P.S. Birds in peril: a guide to the en- dangered birds of Canada and the United States, 210 Mark, D. M. Where to find birds in British Columbia, 203 Merritt, R. W. and K. W. Cummins (eds.). An introduction to the aquatic insects of North America, 209 Nelson, B. The Gannet, 456 Nelson, J. S. Fishes of the world, 206 Ogilvie, M. A. Wild geese, 341 Pelton, M.R., J. W. Lentfer, and G.E. Folk (eds.). Bears: their biology and management, 97 Perry, R. Watching sea birds, 95 Peterson, R.L. North American Moose, 341 Pettingill, O.S., Jr. A guide to bird finding east of the Mississippi, 91 Raveling, D. G. and H. G. Lumsden. Nesting ecology of Canada Geese in the Hudson Bay Lowlands of Ontario: evolution and population regulation, 207 Reynolds, J. W. The earthworms (Lumbricidae and Spar- ganophilidae) of Ontario, 98 Schaller, G. B. Mountain monarchs: wild sheep and goats of the Himalaya, 93 Schorger, A. W. The Passenger Pigeon: its natural history and extinction, 98 Squires, W. A. The birds of New Brunswick, 92 Tanner, J. T. Guide to the study of animal populations, 453 Vaillancourt, J. Lexique anglais-frangais: termes techniques a lusage des biologistes, 208 Miscellaneous Batschelet, E. Introduction to mathematics for life scientists, 349 Blaker, A. A. Field photography: beginning and advanced techniques, 218 Corrivault, G.-W. and P. Morisset. Le Naturaliste Canadien, Index, 462 Inglis, A. Northern vagabond: the life and career of J. B. Tyrrell, 109 Ruffner, J. A. and F. E. Bair (eds.). Weather almanac, 348 Sterling, K. B. Last of the naturalists: the career of C. 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The Editor makes the final decision on whether a manuscript is acceptable for publication, and in so doing aims to maintain the scientific quality and overall high standards of the journal. ee lee pict ae lem: Ai ast : ha ha Vea a my a! e 1 Sear, Oey ready TABLE OF CONTENTS (concluded) Book Reviews Zoology: Guide to the study of animal populations — Quaternary vertebrate faunas of Canada 453 and Alaska and their suggested chronological sequence — Birds of Man’s world — Statistical inference from band recovery data: a handbook — The Gannet Botany: Vascular plant families — An atlas of airborne pollen grains and common spores of Canada 457 Environment: A vanished world: the dinosaurs of western Canada — Biogeography and ecology of 458 southern Africa — The ecology of North America — Chlorinated phenoxy acids and their dioxins Miscellaneous: Last of the naturalists: the career of C. Hart Merriam — Le Naturaliste 461 Canadien Index New Titles 463 Index to Volume 93 Compiled by W. HARVEY BECK 465 Mailing date of previous issue 26 June 1979 Erratum Canadian Field- Naturalist 93(3): 232-238; 1979. Seasonal growth, food, and feeding habits of young-of-the-year Black Crappie in the Ottawa River by John Mark Hanson and S. U. Qadri. The log,, values for total length given on the X-axis of Figure 4 are incorrect; they should be 1.1, 1.2, 1.3, 1.4, 1.5, RGR plevenlese01-9 > and)2-0! 1979 Council — The Ottawa Field-Naturalists’ Club President: Roger Taylor Elisabeth Beaubien Fran Goodspeed Charlie Beddoe Peter Hall Vice-President: Courtney Gilliatt Ron Bedford Hue MacKenzie ¢ Frank Bell Frank Pope Treasurer: Barry Henson Bill Cody eee eaee Recording Secretary: Diana Laubitz Jane Diceman Ken Strang ; . Ellaine Dickson Ken Taylor Corresponding Secretary: Valerie Hume oneuneKson ee Past President: Roger Foxall Marc Forget Those wishing to communicate with the Club should address correspondence to: The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C, Ottawa, Canada K1Y 4J5. For information on Club activities telephone (613) 722-3050. THE CANADIAN FIELD-NATURALIST Volume 93, Number 4 1979 i Articles Some new and interesting vascular plant records from northern Ontario J. LL REE Yearly variations in the population dynamics of Richardson’s Ground Squirrels GAIL R. MICHENER Man’s influence on potential nesting sites and populations of swallows in Canada A. J. ERSKINE Distribution and habitats of four annual smartweeds in Ontario RICHARD J. STANIFORTH and PAUL B. CAVERS Apparent differences in aquatic macrophyte floras of eight lakes in Muskoka District, Ontario from 1953 to 1977 G. E. MILLER and H. M. DALE Utricularia geminiscapa at Mer Bleue and range extensions in eastern Canada ERICH HABER Movements of sympatric species of snakes at Amherstburg, Ontario B. FREEDMAN and P. M. CATLING Winter feeding by Porcupines in montane forests of southwestern Alberta LAWRENCE D. HARDER Interactions between Snowy and Short-eared Owls in winter M. Ross LEIN and PETER C. BOXALL The Biological Flora of Canada 1. Vaccinium angustifolium Ait., Sweet Lowbush Blueberry IVAN V. HALL, LEwis E. AALDERS, NANCY L. NICKERSON, and SAM P. VANDER KLOET Notes Mosses new to Ontario and Quebec ROBERT R. IRELAND and GILDA BELLOLIO-ITRUCCO Swans wintering on Vancouver Island, 1977-1978 RICHARD W. MCKELVEY Interspecific vocal mimicry by Pine Grosbeaks PETER TAYLOR Influence of weather on aggression in Tree Swallows REID N. HARRIS Abandoned windmill used as a nesting site by Great Blue Herons JEAN-LUC DESGRANGES Unusually late pregnancy of a Muskrat in southeastern New Brunswick G. R. PARKER Lynx movements and habitat use in Montana GARY M. KOEHLER, MAURICE G. PeRNaceR. and HOWARD S. HASH Decline of summering Bald Eagles in central New Brunswick Re PB; SiOGEK Lesser Black-backed Gull, Larus fuscus, in Labrador waters KEVIN D. POWERS Occurrence of the Red Phalarope in the Canadian prairie provinces and adjacent states J. PAUL GOOSSEN and DANIEL G. BUSBY Nesting of the Calliope Hummingbird in Kananaskis Provincial Park, Alberta DANIEL F. BRUNTON, SIDNEY ANDREWS, and DAVID G. PATON News and Comment 391 399 405 411 415 43] 433 436 437 439 440 44] 443 445 446 449 452 concluded on inside back cover ISSN 0008-3550 Acme Bookbinding Co., Inc. 100 Cambridge St. Charlestown, MA 02129 CCTV 3 2044 win bx Shty 26 SSSR 25 LSA Se Snes RSnGN nese tase as ts gra: ae ory Rises a) rear i We ae ao Miner Ae a Ws ray * SUS OT ant SEUSS UID ‘, Ne a Xi. 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