The Canadian field-naturalist

a

ee ee
hb eh

Nee wiki
ary

Whee as
Sea)
a ay
al

Agneta
Aestyona
AOS

Pay
Bee hae
Ve in tie ine

vs poe Fi
* lg Ww ge ie
sh wrest
oy Lien acer a as
Ua ee aL
ar an wy thyme
} De rovank!
WI NG
ou HN daa
araerts
‘ rears
al: z Wh edb qoiptte ke
‘ t fy ot
ANB ate At oy Ce AOR are 7 Wes ee Weare rem TBR
on aN | WV Bald bo PV yee alee
woree Tan sin 07 foe IBAYRE fa We ERI GD
eS pyr Sa veNe tw i LPG ARSENE LB feel
wy Wr 09 ‘ ve oa ae aL
oa oe eB Vote ey mek Ws Punt ss: ofp PDN OE None baler
an t revit 10 iPyeant “ SAP wae Ie Ae
cre PRA b HY 3h She Se Wom EW WB Ip ie ayes)
ee me hyd Pr Fevay Soya WRN Epa vbr AP ne eat Hila pe Veco sy
red eer ar ny f ae yte Vp ABV Glas rag) TAC
, ‘ Merah fet Woot WHALER HS TH CNN Be Hehe
‘ne VI wey wwe i eye Sys a fees ype denise yg Wer Ne IWS
ort 4s ts ve gh Ty f 7 } f house FAW ty BPS were te Wipe ps ep femiegbs BeWe Be
aa py t ang ge bey wk, OUT IE spat ¥ [etalon SWS Ve EAR
ii it Ar Ret We ap Puy VR REED? Vo Uthat vate
bot oe are) 7 oa yi Ue Cae ES Wi ate bonne yori
ae) ‘ , 7 CRN eae ee TT
Hell Fee er Cae) . AGMA MO MART RAD ATA TE Ae IN EO GBE ID: Mb IT USS
Ue) 1 z
Pe Rea ,
Fe Uh be blaye ae we t WRB WINANS
FORT Geary ay Ape EW 959 Henrie Gongs AN pervatt DAME e TYE iy
. ty S . eon cn Sy
w ee ety wep wae torte wee
3 ‘ Rn a SSS ETAT TY
seat an j ety ey p fe ry Matron sie hdl
* ff AB} one : Whats prea
, warp i it Mogae Prararsy
aoe ou f 1 wae PUREE ARR SS a
7 Bt) Gey ' Wve a ee ‘whe rer
é } ba . nie weg wey wot’
. theta ra r 1 1 ety a ay epi
Oy ae Ne lee’ re TEU WLI MON Me tp He ye WIpAd WOE enw gagay
Aon) ee Re + , LING APIS pete Hager ier eye TWEE re y
. ten DAUR a SUC RON IEE Wee BCR
‘ veo sae wie Wie Fe ay a BLE AM MW fs
' pip te Po PBN Ce A
Et phe aon ’ Nee Me rn Ye AB Wye es if aya Ca a WY yo a Mage a
15 ' “4
. % ‘ 5 de by vf "
run oni ate 5 He pe it
ay J vs yp! .
' af ‘ on . Pra
+ ys ‘ . ‘ Wye .
ve nod i CDA Teena iyrfioty
, va 1 ny Ve ety eaygel
" ‘ “mi tyne

HARVARD UNIVERSITY
ev
Library of the

Museum of

Comparative Zoology

Ba
hae Oy

ty

)
Bide
a

PNY ashy

1,

me

TUS Ain
ri

im wae oh

Ma i, ii ti

5 tear
tome int

FEB 13 195!

The CANADIA
PIPL D-NATURALISE

: Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

Volume 95, Number 1 January-March 1981

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

C. D. 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: Patricia A. Lalla 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.

Production Manager: Pauline A. Smith, R.R. 3, Wakefield, Quebec JOX 3G0
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, 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 EOA 3C0

All other material intended for publication should be addressed to the Editor:
Dr. Lorraine C. Smith, R. R. 3, Stittsville, Ontario, Canada KOA 3G0

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: Bighorn Sheep ram photographed by Luigi E. Morgantini on the Ya Ha Tinda Ranch, Alberta, during the winter of
1975-1976. See article on page 69.

THE CANADIAN
FIELD-NATURALIST

Volume 95

1981

THE OTTAWA FIELD-NATURALISTS’ CLUB

OTTAWA CANADA

a
eg, Fs i

The Canadian Field-Naturalist

Volume 95, Number | January-March 198]

The Ottawa Field Naturalists’ Club Centennial Symposium
100 Years of Natural History in Canada

Introduction

IRWIN M. BRODO

Botany Division, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8

The late seventies have emerged as a time of retrospectives. There is widespread interest in “heritage” parks
and buildings, historical exhibits, and retrospective collages of movies and television shows. There is no time
more appropriate for a review of past achievements than at a centenary, and The Ottawa Field-Naturalists’
Club, established 100 years ago, has not let the opportunity slip by.

In my opinion, a periodic stocktaking is a worthwhile exercise. It tells you not only where you have been and
where you are, but, if done seriously and thoughtfully, perhaps where you are going or ought to be going as well.
In this spirit, then, the Council of The Ottawa Field-Naturalists’ Club decided to conduct a kind of colloquium,
inviting experts in various fields to tell us what has happened in their own fields of specialization over the last
100 years or so. It was decided that the talks should have a Canada-wide rather than local focus to emphasize the
role this club has played in the study of natural history throughout Canada.

The meeting took place on 19 May 1979, at the Victoria Memorial Museum (home of the National Museums
of Natural Sciences and of Man). The papers, as you will see, related a fascinating history of natural history in
Canada. It was clear from the outset that the history took shape as a result of individual personalities reacting to
events of the time. In fact, what emerged was a mini-history of Canada, with its stories of the explorations for the
transcontinental railways, isolated Hudson’s Bay Company trading posts, and growth of the major Canadian
cities. They traced the change from the strong influence of Europe and Europeans on early endeavors to the
gradual growth of a body of competent Canadian professionals and amateurs. It was pointed out by several
speakers that serious amateurs made many significant contributions throughout this period.

Throughout the talks, it became clear that much of the accumulating information on Canada’s natural history
was revealed in The Canadian Field- Naturalist. Members of The Ottawa Field-Naturalists’ Club have every
right to be proud of the part the journal has played in documenting the history and development of the
field-oriented natural sciences in Canada.

The papers are presented here in the same order in which they were given at the May symposium. Only two of
the papers are not included: the entertaining, informal presentation of /00 Years of natural history in Ottawa
given by William Dore, and Canadian botany since Hooker (1840) by Bernard Boivin which is not yet available
for publication.

Editor’s Note: These symposium papers retain the approaches and styles used by their authors in their oral presentations.
Although not formally reviewed nor as rigorously edited as conventional scientific articles, they have nonetheless been
checked for accuracy and clarity by specialists in the several disciplines.

200 Years of Ornithology in Canada!

FRED COOKE
Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6

The topic, 200 years of ornithology in Canada, is a large one and in the time allotted, I can do no more than
provide a few vignettes which reflect my personal biases on the topic. At the risk of offending both eastern and
western Canadians, all my examples come from central Canada. At the outset I am going to cheat and stretch
beyond the guidelines set by the symposium chairman. The reason for this is simple; the groundwork for
Canadian ornithology was established in the 18th, not the 19th century, and therefore I will consider a time
frame of slightly more than 200 years, rather than 100.

First, there is the discovery phase in which I consider the origins of Canadian ornithology. This is the period
when virtually nothing comprehensive had been written about North American ornithology, nothing in fact
until Wilson and Ord published their American ornithology in 1808. The most important figure in Canadian
ornithology in this period was Andrew Graham, some of whose observations were quoted by Forster in 1772.
Whatever writings in ornithology were available at that time came from Europe.

Second, in the 19th century there occurred what might be called the inventory stage. Tomes on North
American ornithology were becoming available (forexample, Audubon’s Birds of America published in 1826),
and much of the descriptive inventory phase of Canadian ornithology was carried out by visitors to Canada, who
collected and recorded, and filled in details of distribution and taxonomy. Henry Hadfield, who visited
Kingston in the 1850s, was one such visitor with whom I am familiar through the researches of Helen Quilliam in
her delightful book, History of the birds of Kingston, Ontario.

The third trend in the late 19th and early 20th century could be called the conservation phase. Soon after
Canadians became aware of the wildlife and birdlife present in the country, they became aware that in many
species bird numbers were declining at a dangerous rate. Although population size estimates were nothing more
than guesses, it was clear that species such as the Passenger Pigeon, the Eskimo Curlew, and the Golden Plover
were fast approaching extinction. Contrary to what is normal at the present time, many of the conservation
initiatives in early days were taken not at the prompting of naturalist and conservationist organizations but by
alert and effective civil servants, many of them from Ottawa, men like James Macoun, Percy Taverner, Gordon
Hewitt, and James Harkin. One such initiative resulted in the establishment of the Migratory Birds Convention
Act in 1917. The book, Working for wildlife, by Janet Foster, is the source of many of the facts.

It was this Act of Parliament and the recognition that to understand and protect birds in this country it was
necessary to know more details of the population dynamics and overall biology of birds, which led to the fourth
stage of Canadian ornithology which could be called the population studies phase. This is characterized by a
need to know how many individuals of a species exist and whether the species is increasing or decreasing. Much
of the activity of ornithologists, both professional and amateur, at the present time is related to this —Christmas
counts, breeding bird surveys, aerial censuses of colonial birds, bird-banding programs, and the like.

My first vignette, then, is of Andrew Graham, who, but fora mysterious act of plagiarism, would be much
more widely recognized as the father of Canadian ornithology than he is. It is good to remind ourselves that
much of our country was discovered and settled from the north, and an understanding of the avian inhabitants
of the country’s biomes, the taiga and the tundra, was in fact known earlier than that of more densely settled
parts of the country. Andrew Graham was anemployee of the Hudson’s Bay Company and was sent over from
Scotland as a young man in 1741 to work in the three major trading posts on the west coast of Hudson Bay. He
was a trader at Fort Severn, second-in-command at Fort York, 1762-1774, and Chief Factor at Fort Prince of
Wales (Churchill) from 1774 to 1775. He then returned to Scotland where he died in 1815. During his years on
the bay, he traveled extensively between the forts, often carrying the packet, and he kept notes of his
observations, not only on the birds, but on other wildlife, people of the region, and fort life. These observations
are now preserved in 10 separate books in the Hudson’s Bay Company archives in Winnipeg, all carefully
handwritten. Each is slightly different as later copies were added to and subtracted from. Graham’s style is
quaint at times and some of his birds are difficult to recognize from his descriptions. One must remember,
however, that he had no scientific training and no companions except Thomas Hutchins with whom to discuss

'This paper was presented on 19 May 1979 at The Ottawa Field-Naturalists’ Club Centennial Symposium entitled /00 years
of natural history in Canada.

1981 COOKE: ORNITHOLOGY IN CANADA 3

his ideas. He had to rely on European texts such as Pennant’s British zoology, used only his eyes and his gun, and

worked in an extremely inhospitable environment particularly in the winter months. Here is an example of his

work, a description presumably of a Boreal Owl:
Shipomosish, the small owl. This is the smallest owl in the Hudson’s Bay, nearly corresponding to the Small Owl of
Pennant. It weighs four ounces and a quarter. The length is eight inches and a half; the breadth twenty. Irides bright
yellow; the feet and legs are feathered and talons black. There is no material difference between the cock and hen birds,
only the latter being smallest, which is a distinguishing mark of the owl genus. It lives amongst the pines in all seasons
feeding on mice. It builds a nest half way up the pine tree in the month of May; lays two white eggs, and the young fly in
the month of July. They are not plenty, and are the most solitary bird in Hudson’s Bay, seldom moving in the daytime,
but a brisk mouser in the night. They never change colour. It is sport for the Indian children to steal towards them in the

daytime and seize them. Ae
The description does not rule out Saw-whet Owl but by location is almost certainly the Boreal Owl. There are

some errors of fact suchas the observation that female owls are smaller than males, but, on the whole, the report
suggests the interest of an active naturalist who has a personal knowledge of the bird in the field. Graham sent
many specimens to Europe and many of these became the type specimens upon which the original descriptions
were based. Forster in 1772 described for the first time Great Gray Owl, White-crowned Sparrow, Blackpoll
Warbler, and Boreal Chickadee on the basis of specimens sent to Europe by Graham.

The fifth copy of Graham’s Observations in the Hudson’s Bay Company archives is ascribed not to Graham
but to Hutchins. Hutchins was a surgeon who spent 16 years on the Bay. It is clear from researches carried out by
Glyndwr Williams that Hutchins took Graham’s work to Britain and presented it as his own. Whether he did
this with Graham’s knowledge has not been finally resolved, but even in his later life Graham always referred to
“my friend, Mr. Hutchins.”

The major consequence of Hutchins’ plagiarism was that Pennant’s Arctic zoology, the major work of this
time, published in 1785, refers to Hutchins for all of Graham’s observations, and to this day the small arctic race
of the Canada Goose is referred to as the Hutchins Goose, not the Graham Goose.

Let us now turn the clock forward almost 100 years. At this point a number of important treatises on North
American birds had been produced. Visitors to Canada could uncover the local patterns of distribution and sort
out the problems of classification. Graham was confused about various types of sparrow; these had been
clarified in Hadfield’s day. Hadfield visited a Canada already modified by the agricultural activities of man. Let
us take a field trip with him in mid-1 9th century, as quoted in Helen Quilliam’s book from Hadfield’s 1859 paper
in The Zoologist.

Quitting the town of Kingston by the Upper Portsmouth road, leaving the Cathedral on the right and the splendid new

Court-house on the left, you proceed for nearly a mile on the usual raised boarded foot-way, through level, half

cultivated fields almost without a tree or evena stump, then leaving the high road, and striking off obliquely to the right

for half a mile or more over a barren looking common, dotted here and there with clumps of everlasting fir, the first wood
or enclosure is reached . . . there, being endless snake fences to be clambered over, no easy matter when encumbered by
shot and gun, to say nothing of heavy boots. . . and ornithologists afflicted with obesity... should pause ere they trust
themselves astride on the top most bars of suspicious looking snake fences, as they are in these old enclosures very apt to
give way, when great is the fall thereof. I have in my mind’s eye a vivid recollection of a scene of this sort, but the sufferer,
fortunately, was of the leaner kind, or the fall of some five or six feet, gun in hand might have knocked out his own brains

as well as those of the much prized specimens he held.

Birding in the 19th century could bea hazardous business, but less rugged than the life which Graham led, one
feels. This rural scene is at the present day part of urban Kingston, as mancontinues to modify the environment.
Scientific writing in Hadfield’s day portrayed a more leisurely pace of life. What modern editor of The Canadian
Field- Naturalist would tolerate Hadfield’s rambling style?

It was through scientists and collectors like Hadfield that the basic knowledge of bird distribution in Canada
was obtained, and it was from this base of knowledge that the concern for depleted wildlife resources arose. The
depletions arose from three major sources: habitat destruction, market hunting, and the millinery trade. We are
all familiar with the effects of habitat destruction particularly on woodland and marshland species. Market
hunting is now a thing of the past and some of the species which were at that time threatened with extinction
seem to be building up in numbers again now — species such as Hudsonian Godwits, Golden Plover, and
Trumpeter Swans. Perhaps most remote from usat the present time are the depredations of the millinery trade.
Janet Foster (1978) in her book Working for wildlife quotes a lady at the turn of the century as saying, “There is
a great deal of sentiment wasted on birds. There are so many of them they will never be missed, any more than
mosquitos. I shall put birds on my bonnet.” Herons and brightly colored birds were the most vulnerable.

The problem was finally recognized and seen to require for its solution agreement at least between Canada
and the United States. In addition agreement between the Dominion government and the provinces was

4 THE CANADIAN FIELD-NATURALIST Vol. 95

necessary. Dedicated civil servants sensitive to the issues were needed to guide legislation through all the
potential hazards. Fortunately there were such dedicated men available. Gordon Hewitt as Dominion Ento-
mologist was concerned with the economic value of birds and their importance to agriculture. He came to
Canada from England and brought with him the Europeanas opposed to North American wildlife tradition. He
also had a potentially important political advantage in that his wife was the niece of Prime Minister Robert
Borden. She wrote about her husband on one occasion as follows: “Morning and evening he walked around the
garden, bathing himself in greenness, and in the odour of lilacs, roses and new mown grass. Then it was he spoke
to every flower and bird, no matter how small or how shy, and held conversation with the chipmunks and
squirrels, who held a safe tenure within the garden precincts.”

Perhaps it needed devotion like this to put the Migratory Birds Convention Act on the statute books. The act
has now been in place for 62 years and has had a major effect on Canadian ornithology.

This takes us to our final stage, which I call the population phase. If we have the legislation to protect and
preserve migratory bird species, then we need to be able to monitor population changes, to detect population
declines, and to understand population dynamics. We need the facts to know when and how the laws should be
applied.

Much of the energy of the Canadian Wildlife Service (CWS) has gone into the acquisition of population data
through their own research, through contracts with Canadian universities and others, and by encouraging
amateurs. I will give a few examples of this from my own experience.

The Lesser Snow Goose population which nests in the Hudson Bay region migrates through the prairies to the
Gulf Coast. As such it is vulnerable to hunting and destruction of its feeding areas. In 1972 CWS attempted to
make a total count of the nesting geese in this population. Aerial photographs were taken during June when the
birds nest in large colonies. Each colony was visited and photographed. Each individual nesting female
appeared on the photograph as a white dot, and each dot was counted. This was an enormous undertaking but
from this it was determined that more than one million Snow Geese were nesting in the Hudson Bay - Foxe
Basin regions. This seems to be the only Canadian species other than the extremely rare ones such as the
Whooping Crane, for which a reasonably accurate census is available. Such censuses are essential if population
changes are to be assessed.

I have been involved as a university researcher in another aspect of the population monitoring (see Cooke and
Sulzbach 1978). Since 1968 we have been studying closely a self-contained Snow Goose colony at La Pérouse
Bay in northern Manitoba. By collecting data on an annual basis it is possible to assess annual mortality and
annual production. These are necessary measurements if population declines are to be detected. Unfortunately
this information is known for too few species of Canadian birds.

Most of the monitoring of population changes comes, however, not from the professional ornithologist, but
from the amateur: the weekend birdwatchers and birdfeeder watchers. The documentation of the spread of the
Cardinal into Ontario or of the Evening Grosbeak from the West depended largely on the observations of
dedicated amateurs. Christmas bird counts if analyzed cautiously can be used to document changes of status of
some of our birds. More recently breeding-bird surveys have been introduced which give a certain standardiza-
tion to amateur record keeping. This is an early example of government initiatives in harnessing the activities of
amateur birdwatchers.

The assistance which the Federal Government has recently given to Canada’s two bird observatories, Long
Point Bird Observatory and Prince Edward Point Observatory, points also to the importance of data collected
systematically by amateurs. The increase in certain species of warblers at migration points in years of high
Spruce Budworm outbreak has been reported. Most birdwatchers are happy to have their observations used for
such a purpose, and the efforts of government agencies such as the Canadian Wildlife Service in coordinating
the activities are welcomed.

There is much still to be done. Much of our inspiration still comes from Europe where more progress in this
direction has been made. In The Atlas of British birds (1976) the distribution of every British breeding bird is
recorded, and estimates of the numbers of each species plus indications of range expansions or declines are
noted. Because of the geography of Canada and because we have fewer birdwatchers it will be several years
before we can emulate the Atlas here, but there are beginnings. At the 1979 Ontario Ornithologists meeting, a
‘small committee was set up to look into the feasibility of an atlas project at least for the settled parts of southern
Ontario. I welcome these initiatives.

We have come a long way since the “Observations” of Andrew Graham. I am always overwhelmed, however,
by how little we still know. The amateur birdwatcher can still find out new facts himself or become part of a team
doing essential census work. Only by doing this can we ensure that the Migratory Birds Convention Act can be

1981 COOKE: ORNITHOLOGY IN CANADA 5

effectively administered. Our hobby or research can be turned to good use for the protection of those species
we enjoy observing.

Literature Cited

Audubon, J. J. L. 1827. The birds of America from original drawings. 4 volumes. London.

Cooke, F., and D. S. Sulzbach. 1978. Mortality, emigration and separation of mated Snow Geese. Journal of Wildlife
Management 42: 271-280.

Foster, J. 1978. Working for wildlife, the beginning of preservation in Canada. University of Toronto Press, Toronto.

Graham. A. 1772. Observations on Hudson Bay. Manuscript, Hudson’s Bay Company Archives, Winnipeg, Manitoba.

Hadfield H. W. 1859. Birds of Canada observed near Kingston during the spring of 1858. Zoologist 17: 6701-6709;
6744-6752.

Kerbes, R. H. 1975. Lesser Snow Geese in the eastern Canadian Arctic. Canadian Wildlife Service Report Series #35. 49 pp.

Pennant, T. L. 1766. The British zoology. Published under the inspection of the Cymmrodorian Society, London.

Pennant, T. L. 1792. Arctic zoology. 3 volumes. London.

Quilliam, H. R. 1973. History of the birds of Kingston, Ontario. Published privately. 209 pp.

Sharrock, J. T. R. 1976. The atlas of breeding birds in Britain and Ireland. British Trust for Ornithology. Irish Wildbird
Conservancy. Published by T.A.D. Poyser. 480 pp.

Williams, G. 1969. Andrew Graham’s observations on Hudson Bay, 1767-1791. Hudson’s Bay Record Society. Volume
XXVII.

Wilson, A., and G. Ord. 1808. America ornithology or the natural history of the Birds of the United States. 9 volumes.
Philadelphia.

Mammalogy in Canada: a Historical Overview!

DONALD A. SMITH
Museum of Zoology, Department of Biology, Carleton University, Ottawa, Ontario K1S 5B6

Mammalogy is simply the study of mammals and is usually restricted to wild, nonhuman mammals, as I will
do here. The story of the development of mammalogy in Canada is closely interwoven with the history of events
and human beings in Europe and elsewhere in North America as well as in our own country. It isa complex and
interesting story but far too long and varied to relate in the limited time available to me, so I’ll tell you something
of the background, some highlights, some interesting episodes, and something about interrelations of The
Ottawa Field-Naturalists’ Club and Canadian mammalogy over the past century.

Certainly only well within the last 100 years, perhaps only within the last 60, has mammalogy reached the state
of being considered by its practitioners or by scientists in general as a separate field of science anywhere in the
world, but especially in North America, which is in the forefront of mammalogy. Prior to the last six or seven
decades most work in mammalogy was done on the side by ornithologists. The early work was often done rather
casually and most early reports on Canadian mammals are either scattered through the journals of explorers, or
are in combined bird~mammal publications, often as mere appendices to ornithological reports.

Discovery Phase

To outline the saga of the evolution of mammalogy in Canada, I must go back much earlier and set the scene
by dealing with other parts of the world first. lam going to have to wander quite a bit outside Canada because
external cultural and political events have had a great deal to do with Canadian mammalogy.

Certainly all aboriginal peoples who depended on animals for food and clothing knew something about
mammals. This knowledge was essential to their survival. They had practical, often ecological, information
which was passed down from generation to generation usually by the spoken word, although graphics were
sometimes employed. Written reports came much later and at first were based more on superstition, on rumors,
on speculation, than on observation. If you’d like a time reference, I’m talking now about the time of the ancient
Greeks, around 2000 years ago. I'll take a big jump over the Dark Ages during which very little advance was
made anywhere on incipient mammalogy.

The 15thand 16th centuries brought the beginnings of the major voyages of exploration by Europeans to the
New World, but aside from relatively minor mentions of mammals by adventurers, there was no significant
documentation of New World mammals. In the 17th and 18th centuries exploration intensified and coloniza-
tion of different parts of the world became significant. Back in Europe there were many wealthy people witha
great deal of leisure time that could be devoted in part to cultural activities. Many developed an interest in
natural history, and amassing specimens as display objects for their natural history cabinets was one of the
favored occupations of many gentlemen. (I’m sorry to state that I'll have to report exclusively about the
activities of gentlemen rather than ladies in this context; I don’t know what the ladies’ interests in natural history
were in this period — perhaps botany and ornithology — because they’ve been little publicized.) Many of the
European gentry interested in possessing and displaying specimens did not collect them themselves, but sent
naturalist-collectors off to the New World, Africa, and elsewhere to explore and bring back new and interesting
specimens for their private collections and eventually for those of the great institutions in European capitals. As
a result of expanding exploration, commercial enterprises were set up to exploit the newly discovered natural
resources of the colonies, and settlement commenced or accelerated. The major result of significance here is that
there was a flow of specimens of mammals and other organisms back to Europe. Meanwhile, in several
European countries, events occurred that collectively improved the scientific climate for the reception of these
specimens. The organization of the Royal Society in London in 1662 greatly accelerated the acquisition of
knowledge. A landmark date is 1758 when the 10th edition of Systema naturae was published by Karl von Linné
(Carolus Linnaeus). This famous Swedish biologist initiated the system of scientific binomial nomenclature that
is the basis for the scientific names we use today forall plants and animals. He named and described 86 species of

'This paper was presented on 19 May 1979 at The Ottawa Field-Naturalists’ Club Centennial Symposium entitled /00 years
of natural history in Canada.

1981 SMITH: MAMMALOGY IN CANADA 7

mammals in 1758 and others began to use his scheme to name still more species. Thus there was considerable
activity in naming new species from specimens brought back to museums in London, Paris, Berlin, Munich, and
other European centers. Sometimes, unfortunately, other new species were named only from descriptions or
illustrations by earlier writers.

Concurrently there was awakening interest in paleontology in Europe largely as a result of the spark by
Georges Cuvier in France in the late 1700s and early 1800s. At this same time, surprisingly, there was much
activity in vertebrate paleontology in the fledgling United States of America. This was due largely to an eminent
naturalist who wrote on fossils and other branches of sciences — none other than the third President of the
United States, Thomas Jefferson. He and later presidents sent parties of explorers and naturalists into the
western part of North America to survey the animals, plants, fossils, and minerals. For example, most have
heard of the expeditions of Lewis and Clark, for whom Lewis’ Woodpecker and Clark’s Nutcracker were
named. Very quickly there built up in the United States a great tradition of semimilitary explorers and
naturalists. Sometimes the same person filled both these roles; sometimes the surgeon who went along was also
the naturalist. There were many of these expeditions in the first half of the 19th century and they intensified in
the 1850s whena route was sought fora railroad to the Pacific. They all brought data and specimens back east to
new museums that were founded by individuals, universities, and governments at several levels. Clearly this was
a continuation of what had gone on earlier, when the specimens went to Europe. During the latter half of the
19th century, the museums of the United States built up staffs of specialists who studied all this western material;
in particular they described many new species, including new mammals. There was rapid publication and
efficient dissemination of knowledge. Here was the case of aggressive and productive United States enterprise!
In case you wonder what significance this has for Canada, it is that these expeditions were not confined to the
western states, but they entered Mexico, and they went a great deal into British North America. Thus, again
foreigners were collecting, removing, and describing part of Canada’s natural resources! I'll now come closer to
home briefly and quote from the President’s Address to The Ottawa-Field Naturalists’ Club in 1887. John
Macoun (Ottawa Naturalist 1(2): 20. 1887) was deploring, as so many have since, the indifference of Canadians
in contrast to all this aggressive activity south of the border. He deplored also the lack of a great museum in
Canada and said, “It is painful to confess that all our best specimens, whether mineral, animal, or archaeological
are going to Washington, and the apathy of our people is such that I see no help for it.” ll refer to a later stage of
this removal of part of our heritage shortly.

No account of mammalogy in Canada is complete without saying something about the fur trade, which was
based, initially at least, on one species of mammal, good old Castor canadensis, our friend the Beaver! This
industry began in the mid-1 6th century as French explorers along the Atlantic Coast and up the St. Lawrence
River began bartering with the Indians. Other Europeans, largely from the United Kingdom, also began
exploiting this resource, for the fur of Beavers was very popular back home. In 1670 Charles II granted a charter
to the Hudson’s Bay Company to utilize the Hudson Bay area. From that date until Canada was ceded to Britain
in 1763, there was intense competition between the French fur traders based at Montreal and the British traders.
with Hudson Bay as their center of operations. As the populations of Beavers were depleted, trapping activities
and the establishment of trading posts were gradually extended westward, and this led to further exploration
and eventually to settlement. Various species of mammals other than Beavers and other valuable furbearers
were also obtained by the fur traders and sent back to Europe. The process I described earlier of these specimens
being described and named by European scientists who had never seen the living or intact dead animal took
place in Britain, France, and Germany. For instance, J.R. Forster was asked by the Royal Society in the 1760s to
examine “a choice collection of skins of quadrupeds . . .” collected by servants of the Hudson’s Bay Company at
their posts on the bay.

There was sometimes a very anomalous consequence of scientists obtaining specimens from nonscientific
sources such as the Hudson’s Bay Company and its competitors. Some of you students and scientists know that I
frequently complain about inadequate data on specimen labels. Well, the data accompanying specimens from
the fur-trading companies were certainly minimal, if they existed at all, and usually consisted of only the names
of the posts from which they had been shipped. Whena species new to science is being described and named, the
author records the “type locality” as the locality from which the holotype or “type specimen” came. Needless to
say, the type localities of some Canadian mammals described in the 1 8th century by Europeans from specimens
obtained from the fur trade are rather vague and sometimes even outside the natural range of the animals! One
example is Taxidea taxus, the American Badger, which in Canada occupies grassland regions of the prairie
provinces, some valleys in British Columbia, and a couple of specific parts of western and southwestern Ontario.
It also extends, of course, extensively through the western United States and northern Mexico. Its type locality,

8 THE CANADIAN FIELD-NATURALIST Vol. 95

though, is “Labrador and Hudson Bay” as recorded by the German Schreber, who described it in 1778. He
cannot be faulted because he did not know any better, not having been to North America, and not knowing that
American Badgers are prairie-dwellers that have never been recorded in Labrador nor anywhere near Hudson
Bay. I presume that although his specimen must have truly been shipped from a fur post on the shore of Hudson
Bay, it had actually been killed somewhere in the prairies by Plains Indians who traded it to Swampy Crees or
Chipewyans who transported it up the Nelson or Churchill rivers to a post on the bay. We'll never know for sure.
That is just one example; there are comparable ones.

Before leaving the fur trade, I should mention that our knowledge of the natural history of northern Canada
began with the formation of the Hudson’s Bay Company in 1670. Several of the men sent from Britain as factors
in charge of trading posts were keen naturalists and sent reports of their observations and specimens of
mammals, as well as of birds, etc., back to the United Kingdom. Some well-known explorers of northern
Canada who were associated with the fur trade also recorded observations on mammals. For example,
Alexander Mackenzie in 1789 described the “white buffalo” in the mountains of the northwest. His “white
buffalos” are now known as Mountain Goats. In 1796, Samuel Hearne published rather accurate (for his time)
observations on northern mammals that had seen on his travels between 1769 and 1772. Incidentally, the old
records of the fur trade proved to be important to 20th century mammal ecologists such as Elton, Chitty, and
MacLulich, because they provided data on population fluctuations of foxes, lemmings, and hares over many
years.

To finish off my consideration of this discovery or exploration phase of Canadian mammalogy, let me remind
you that our knowledge of the fauna of arctic Canada was greatly increased during an era of exploration
stimulated in 1818 by the British government’s offer of £20 000 for making the Northwest Passage from the
Atlantic to the Pacific. Three British explorers who were prominent in this search were Sir John Franklin; Sir
John Richardson, who was surgeon-naturalist for Franklin’s expeditions of 1819-1822 and 1825-1826; and Sir
William Edward Parry. They were most active in the decade after 1819, but Richardson was active until 1848.
His last trips were to hunt for the ill-fated third Franklin expedition of 1843. One special item of interest to
mammalogists is that all three of these gentlemen have been commemorated by having species of ground
squirrels named after them! So we have Franklin’s (Spermophilus franklinii), Richardson’s (S. richardsonii),
and Parry’s (now Arctic) (S. parryii) Ground Squirrels. I don’t know if they’d be honored; I hope they would; I
certainly would! Of these three men, Richardson contributed by far the most to natural science by describing
mammals (and other animals) he saw and collected on his travels. The first part of his Fauna boreali-americana,
published in Britain in 1829, was a real landmark in describing the quadrupeds of northern Canada. His
collections, like those of many others, including type specimens, went to the British Museum in London.

At the close of the active period of arctic exploration by the British coincidental with the discontinuation of
the search for Franklin’s expedition about 1857, the involvement of the British with Canadian natural history
including mammalogy became almost nonexistent. United States zoologists replaced the British as outside
investigators of the taxonomy and distribution of Canadian animals through their expeditions, identification of
submitted specimens, and taxonomic studies. Otherwise, contributions were largely being made by workers
whose primary responsibilities were in other activities, and whose interest in mammals was personal rather than
official. Thus during the rest of the 19th century, there were various minor accounts of Canadian mammals by
explorers, geologists, missionaries, and others including a few naturalists, all adding bits and pieces of
information, but there would not be a Canadian medium for pulling them all together for some years.

Before proceeding to acompletely different aspect of the story, Ishould report on the increase in numbers of
mammals known. In 1820 Desmarest described all known species of North American mammals — about one
hundred. The first list of Canadian mammals was published by a member of The Ottawa Field-Naturalists’
Club, J.B. Tyrrell, field-geologist of the Geological and Natural History Survey of Canada, in 1888. His 28-page
The Mammalia of Canada listed 123 species with brief annotations. Frank Banfield’s The mammals of Canada
published in 1974 treated 196 species.

Contributions by Canadians in the 19th Century

I shall digress now to tell you about the nature of contributions to mammalogy by Canadians during the 19th
century, and especially, around 100 years ago, the latter in response to a request by the chairman of this
symposium. Among the earliest writings about Canadian mammals by Canadians, and including perhaps the
first records of Ottawa-area mammals, are general accounts of cervids (“Moose Deer,” “Wapite[sic] or
Canadian Stag,” “Common Deer”), carnivores (bears, foxes, skunk), and squirrels published in 1856. They were
all written by Elkanah Billings, amateur paleontologist, naturalist, barrister, former Editor of The Citizen,

1981 SMITH: MAMMALOGY IN CANADA 9

publisher, and from 1856 to 1876, the first paleontologist to the Geological Survey of Canada. He published his
mammalarticles in the first volume of The Canadian Naturalist and Geologist which he founded in Ottawa (and
of which he was at first practically the sole contributor), but he moved it to Montreal later in 1856. This was, I
believe, the first English language natural history serial in Canada, and its publication“. .. marks anepoch in the
history of the progress of scientific research and discussion in Canada” according to President H.M. Amiinan
address to The Ottawa Field-Naturalists’ Club in 1900 (Ottawa Naturalist 14(11): 204. 1901).

My search for information next led me to the first few volumes of the publications of The Ottawa
Field-Naturalists’ Club. In perusing them, I was quickly captivated by the amount of fascinating information
and the literate, readable style, so that I could hardly put them down. Although there is plenty about plants,
plenty about birds, plenty about insects, plenty about fossils, and even plenty about the common local mammal
Homo sapiens, in contrast there is precious little about native Canadian mammals. In Transactions of The
Ottawa Field- Naturalists’ Club No. | for 1879-1880, aside from humans, there is absolutely nothing about
mammals. In Transactions No. 2 the next year, John Macoun mentioned that buffalo and deer had “. . . eaten
the district [of the prairies] bare,” but that’s the entire content about mammals. In Transactions No. 3 and 4, I
could find nothing about Canadian mammals, but lots about geology, plants, insects, birds, and shells, for there
were then highly organized “branches” of the club studying these groups.

In Transactions No. 5 for 1883-1884, in H. Beaumont Small’s President’s Inaugural Address, which was
delivered 6 December 1883, he referred to certain accomplishments of the club over its four years of existence in
publishing much local information on geology, stratigraphy, minerals, plants, birds, fishes, and shells — listing,
then adding to and perfecting their lists. Indeed this was in accord with the “object” of the club to study “the
natural history of this locality.” But he went on to predict that“... before another four years shall have passed,
the resources of this district will be so thoroughly known that we shall be sighing for fresh fields to explore and
conquer.” He was contemplating expanding the club’s efforts to surrounding regions of the country before even
one word had been written about a local mammal. How shocking! In that same address, however, Small did
acknowledge that mammals existed. The first use of the work “mammal” in the club’s publications was in his
profound statement “Mammal orders are according to the hoof and teeth, . . .” [sic]. In the ensuing discussion
Macoun mentioned “marsupials and monotremes, the lowest forms of mammalia [sic]” but this was strictly out
of the book because the former were not then known to occur in Canada and the latter are not known to range
outside Australasia. Finally, late in this same volume, appeared the very first paper on mammals in the club’s
publications. “The deer of the Ottawa Valley” had been delivered on 13 March 1884 by W.P. Lett. It dealt with
Moose, Caribou, Wapiti, Virginia or Red Deer[now White-tailed Deer], and was rather well done. Towards the
end of Transactions No. 5, in the first Report of the new Zoological Branch of the club, was the first note on
small rodents. I was pleased to learn that they finally recognized the existence of some of the wonderful beasts I
study — “white-footed or deer-mouse,” as well as the “meadow mouse” and “jumping mice” (Zapus hudsonius),
which they seem to have called also “kangaroo mouse.”

Transactions No. 6 for 1884-1885 contains the President’s Inaugural Address again delivered by H. Beau-
mont Small, M.D., on 4 December 1884. He referred to what had been accomplished by the club and what
remained “for new workers to begin upon.” This perceptive naturalist now realized that mammals had received
very little attention, and he promoted “the systematic study of our mammals and the preparation of a perfect
list” with annotations. The only paper on mammals was “The Canadian Otter” by William Pittman Lett. The
Report of the Zoological Branch by Small and Lett contained almost two pages on local mammals including the
first mentions in club publications of bats and two reports of seals in the Ottawa River. They hoped to present “a
complete list of the mammals met with in this vicinity” the next year but this hope was not realized until 1945,
when A.L. Rand’s “Mammals of the Ottawa District” appeared in The Canadian Field- Naturalist (59: 111-132).
Indeed the momentum of the previous number did not carry over to Transactions No. 7 for 1885-1886, for the
only significant article on mammals was “The Black Bear” by none other than William Pittman Lett. This man
was still the only club member to put his own observations and others’ published observations on mammals
together into species accounts, and he did it well too!

In 1887-1888 The Ottawa Naturalist commenced as Volume 3 of the Transactions. At this time President
John Macoun was ina complaining mood. He deplored the fact that contemporary Canadian naturalists didn’t
like to walk for miles in the countryside as many English naturalists did; instead they liked to ride to a point in
the country, eat a hearty lunch, take a short stroll, eat again, then ride back to their homes. He also deplored the
lack of botanic gardens and museums, and I quoted earlier his regrets that all the specimens were going to
Washington and that people were so apathetic about it. He complained further that space was limited in the
Geological Survey Museum in Ottawa. There was one small room for mammals and birds [we can’t escape from

10 THE CANADIAN FIELD-NATURALIST Vol. 95

birds!], and the insects and plants were out in the hall! In other words, he advanced some of the perennial
complaints that many of usin the museum community are still voicing. That same year, 1887, the club’s faithful
lone mammalogist William Pittman Lett reported on “The Cougar or Panther.”

In the next President’s Address in December 1887 R. B. Whyte stated, “We have pretty well accomplished our
task of investigating and recording the results; the small additions that have been made to our lists of late years
show that not much remains to be done... .” But he did realize that “next to nothing is known...” about the
mammals, “making it a fertile field for study and investigation.” Furthermore, without giving any details, he
claimed that “every year our animals are becoming scarcer, and papers on the mammals... of the district would
be interesting and valuable.” Well, it took a long time before more than a few people did anything about the
mammals of the Ottawa area.

On 9 February 1888, however, during the discussion after Professor Macoun’s lecture on vertebrates,
including mammals, came the first exhibition to the club of a mammal specimen (although whether it was
actually prepared as a specimen for retention or merely exhibited as a carcass and later discarded is not stated).
“Mr. [W. H.] Harrington exhibited a fine specimen of Hesperomys [now Peromyscus] leucopus, the white-
footed mouse, which he had trapped the previous night in his shed. Attention was called to the great beauty of
this native field mouse and a few remarks were made on its habits. The President, Mr. R. B. Whyte, urged the
collection and study of our small mammals, of which there are many species but little known to the members”
(Ottawa Naturalist 1: 179. 1888).

Shortly afterwards, on | March 1888, J. Ballantyne broke W. P. Lett’s monopoly on presenting papers on
mammals, for he read a paper on “Our Squirrels” at a soirée, and exhibited mounted specimens (from the
museum of the Geological and Natural History Survey of Canada) of the different species (Ottawa Naturalist
2(1): 7. 1888) His paper, published on pages 33 - 44 of this volume, contains many personal observations of his
own, plus information from the literature, about five local species. The remainder of Volume 2 of The Ottawa
Naturalist contains only few brief mentions of mammals.

Volume 3 of The Ottawa Naturalist for 1889-1890, the last early volume I surveyed, contained some
innovations, as well as the second-last contribution by William Pittman Lett, this time on “The American
Skunk.” The scope of the journal was extended to include papers from elsewhere in Canada instead of being
confined to papers on purely local topics. Editor James Fletcher forthwith published a brief notice of the
publication by C. Hart Merriam (from Washington) inan 1888 American Naturalist of the description of what
was then considered a new species of red-backed vole, Evotomys dawsoni, whose type specimen had been
collected in the Yukon by George M. Dawson, a prominent geologist and club member. This specimen, and
others collected in southern British Columbia by John Macoun and his son and mentioned in this same volume,
are the first bona fide museum specimens of mammals referred to in the journal, beyond the mounted squirrels
mentioned earlier.

Anyone who has looked through these early journals knows that there are many accounts of the field trips of
The Ottawa Field-Naturalists’ Club. There are details on the plants, birds, insects, shells, and fossils found, but
up to 1890, at which point I stopped my survey, there was only a single mention of a mammal being observed on
a club outing. Even that observation, of a Red Squirrel “near Meech’s [sic] Lake,” was not ina formal account of
an outing but was buried in Ballantyne’s paper on squirrels (Ottawa Naturalist 2: 38. 1888). Well, I suppose
this is not surprising; in contrast to birds, insects, and plants, most mammals, especially small ones — shrews,
moles, mice, voles — are nocturnal, shy, secretive, and they’re rarely seen. In those days, the little furry creatures
in which I’m so interested were usually brought in by cats or dogs or they were picked out of water pails or rain
barrels into which they had fallen, and usually they were in rather bad condition. Those that were actively
collected had to be caught by hand, shot, or taken in steel traps. Only a few ingenious people seem to have
devised traps to capture small mammals, at least in North America. One such person from the USA is the
Reverend John Bachman, who, with John James Audubon, wrote the three-volume folio The viviparous
quadrupeds of North America which appeared between 1845 and 1848. This was a landmark, the first major
work on North American mammals. Bachman used a figure-4 trigger and an inverted bowl to trap mice.

Mousetraps and the Inventory Phase

This brief digression on relatively crude traps leads me to a story about how a brilliant but simple
technological advance triggered a revolution in mammalogy. One of the conventional features of historical
accounts such as mine is to describe the major “revolutions,” “break-thoughs,” or “turning-points” in the branch
of science or technology being treated — Newton, Watt, Darwin, transistors, sputniks, lunar landings, and so
on. Well, the new device that resulted in the first breakthrough in mammalogy was not nearly as spectacular as

1981 SMITH: MAMMALOGY IN CANADA 11

these, but it was nonetheless important. The quantum leap forward in mammalogy came about asa result of the
invention, mass production, and marketing of small, cheap, efficient, breakback mousetraps! Although this
familiar device doesn’t seem to rate in the earth-shattering category, this little product of Yankee ingenuity, first
called the Cyclone trap when it was marketed in 1887 or 1888 — I’m not sure of the exact year — brought abouta
great revolution in mammalogy when its possibilities were recognized by a fertile mind.

To continue this story I must talk about a very important individual whom I have already mentioned in
passing — C. Hart Merriam from the USA. This eminent pioneer mammalogist was elected a corresponding
member of The Ottawa Field-Naturalists’ Club in 1884 or 1885, and his donation of his book Mammals of the
Adirondacks was the basis for the first published report of a publication on mammals being received by the
club’s library (Transactions No. 6: 159. 1885). He was a medical doctor whose interests lay more in ornithology
and mammalogy than in medicine. He was employed by the United States Department of Agriculture in 1885 as
the first Director of its Branch of Economic Ornithology in the Division of Entomology. The next year he
expanded his branch to include mammalogy. Soon, with the advent of Cyclone traps, he envisaged a continental
campaign of large-scale mammal-collecting; this was the complete revolutionary idea. He had realized that
because animals range according to physiographic or environmental boundaries rather than political ones,
comparative study of the systematics of North American mammals (and other organisms) required specimens
from the entire continent. Accordingly he sent collectors throughout North America, from Mexico to arctic
Canada. Their mission was to collect large series of small-mammal specimens, each with its own label bearing
complete data — precise locality, date, collector’s name, standard measurements, mass, sex, breeding condition,
habitat, remarks, everything! These series of specimens showed both individual and population variation; they
showed variation due to age, to sex, to season, and perhaps most striking of all, to geographic variation. Up to
this period many museum collectors were content to obtain specimens of one male and one female of each
species, hence this so-called “Merriam concept” of collecting many specimens was really very trend-setting and
led to great advances in North American mammalogy. His collectors sent or brought their specimens and
detailed field notes back to Washington, of course, and Merriam identified them. His many descriptions of new
species and subspecies were based on these series rather than merely on single specimens, as was the usual case
earlier. Merriam was a prodigious worker and he and his colleagues — Bailey, Preble, Nelson, Osgood, et al. —
put out vast numbers of papers on mammals and other vertebrates collected on U.S.-sponsored expeditions or
sent to Washington by other collectors. The findings of these early workers on the taxonomy and evolution of
mammals have generally stood up very well in the face of more recent studies using modern techniques of
electrophoresis, chromatography, karyology, and so on. As might be expected, many of their results were
published in U.S. Government series suchas North American fauna (from 1889) and in various publications of
United States museums.

All this activity below the border has had one very striking consequence for Canada. I have already described
how early mammal-collecting by Europeans resulted in the accumulation of specimens of Canadian origin in
European museums. Then in the 19th and early 20th centuries, many Canadian specimens went to the United
States. The result is that most holotypes from Canadian type localities are in museums in these other countries.
Some types are not known, others have been lost, but many are in museums in London [see A. W. F. Banfield’s
1961 paper in National Museum of Canada Bulletin 172 about some of our Canadian treasures in the British
Museum (Natural History) there], in Washington, New York, Chicago, Ann Arbor (Michigan), Cambridge
(Massachusetts), Munich, etc. Even though many North American species have type localities in Canada, our
national collection of mammals in the National Museum of Natural Sciences in Ottawa contains not one single
holotype of a Canadian mammal species. There are some holotypes of geographic races or subspecies there, but
these are of much less importance and interest to most workers.

Canadian Mammal Collections

I have emphasized the large-scale collecting by personnel from United States institutions. Meanwhile, were
there no Canadians also collecting specimens of mammals? Yes, there were. We too had our share of hardy
explorers, missionaries, geologists, and naturalists traveling in the hinterlands of the west and north, especially
during the latter half of the 19th century. Unfortunately many of the mammal specimens they collected were sent
to the big museums in Washington or New York, and relatively few were deposited in the humble museum of the
Geological and Natural History Survey in Ottawa or elsewhere in Canada.

Eventually though, the Merriam type of museum collecting began to be practiced by Canadians, and
Canadian mammal collections were established. Unfortunately, however, the discovery and inventory phases
had been so nearly completed — virtually all the species and almost all the subspecies had been described by

12 THE CANADIAN FIELD-NATURALIST Vol. 95

foreigners — that there remained largely the filling in of details, as far as taxonomy and distribution of
Canadian mammals were concerned.

Nevertheless, through the efforts of devoted curators, technicians, and collectors, often working with
extremely limited resources, valuable collections of mammals have been built up in national and provincial
museums and in some universities across Canada. These collections serve not only as secure repositories for
important specimens, which are part of our national heritage, but as reference collections or “bureaus of
standards” for aid in identifying new specimens. The museums provide a place for researchers, whether
professionals, students, or amateurs, to work with the valuable resources they hold, and some have scientific
publication series for disseminating the new systematic and distributional knowledge emanating from their
researches. The most important museum series of publications are those of the National Museum of Canada,
the Royal Ontario Museum, and the British Columbia Provincial Museum. The development of museum
collections of mammals, particularly those in universities, has advanced the development of mammalogy as an
academic subject. Thus mammalogy in Canada has been positively influenced by the increase in number and
quality of Canadian museum collections of mammals.

The latest information I have on the size and status of Canadian collections of mammals relative to those
elsewhere in North America was collected in 1973 and published in 1975 by J. R. Choate and H. H. Genoways
(Journal of Mammalogy 56(2): 452-502). There were then in eight provinces 21 Canadian collections of
mammals with 100 or more specimens, plus about nine smaller or presumably inactive, nonreporting ones. A
1975 survey of most Canadian mammal collections by Jack Dubois of the Manitoba Museum of Man and
Nature reported no substantial changes among those reporting.

Canada’s largest mammal collection is at the Royal Ontario Museum in Toronto. Six years ago it had around
73 400 specimens, many of which are from foreign countries. This is only the eighth largest mammal collection
in North America. Incontrast, this continent’s largest, in Washington, D.C., contained over 480 000 specimens
in 1973. These are in the combined collections of the Smithsonian Institution’s National Museum of Natural
History and the United States Bureau of Sport Fisheries and Wildlife, Bird and Mammal Laboratories
(formerly the Branch of Economic Ornithology and Mammalogy, mentioned earlier, and later the Bureau of
Biological Survey). The second largest is in the American Museum of Natural History in New York with 250 000
specimens! Both of these huge collections are also worldwide in scope.

Most of the specimens in the rest of Canada’s mammal collections are of Canadian origin, although certain
collections have a remarkable diversity of foreign mammals as well. Our second largest and North America’s
11th largest is at the National Museum of Natural Sciences in Ottawa, with about 43 500 mammals. Then the
next five are at the University of British Columbia, the British Columbia Provincial Museum, Carleton
University (surprisingly enough), the University of Alberta, and the Manitoba Museum of Man and Nature. In
1973 their mammal collections ranged from 9525 down to about 6000 specimens. The others ranged from about
2000 downward. Most important Canadian mammal collections are currently being added to by active but
selective collecting to fillin gaps in our knowledge of Canadian and foreign mammals. Their value in preserving
for posterity for reference and research in perpetuity examples of rare and endangered as well as currently
abundant species of mammals will be enhanced as time goes on.

It is time to pass from the story of Canadian museum mammal collections to a broader aspect of mammalogy
in North America, and to consider some of the ways in which Canadians have contributed.

The American Society of Mammalogists

After the intense activity in mammal study that took place in the last two decades of the 19th century and the
first two of the 20th, mammalogy really came of age as a separate discipline in North America with the founding
of the American Society of Mammalogists (ASM) in 1919 and the start of publication that same year of its
quarterly Journal of Mammalogy, the world’s first journal devoted solely to papers on mammals. Of the 60 men
and women who attended the founding meeting in Washington, only two were Canadians. Among the
approximately 250 charter members who joined the society in 1919 were R. M. Anderson, J. B. Harkin, Hoyes
Lloyd, P. A. Taverner, and M. Y. Williams, all from Ottawa, and all but Harkin were active members of The
Ottawa Field-Naturalists’ Club. Moreover, Anderson was elected to the first Council of the new society. Charter
members of the ASM from elsewhere in Canada included Allan Brooks and J. A. Munro from British
Columbia; Normanand Stuart Criddle from Manitoba; J. H. Fleming, A. B. Klugh, W. E. Saunders, and J. D.
Soper from Ontario; and a sprinkling of others. As mentioned earlier, most students of mammals at that time,
whether professional or nonprofessional, were also interested in birds, if not all other vertebrates, and often
invertebrates and plants as well, and this was certainly true of these Canadian mammalogists of 60 years ago.

1981 SMITH: MAMMALOGY IN CANADA 13

Canada has provided the meeting sites for three of the 59 Annual Meetings of the ASM held up to this year.
Successful meetings were held in Toronto in 1948, Winnipeg in 1965, and Vancouver in 1971. I can state with
confidence that the latter two were successful because my wife Lorraine and I attended both as active
participants. My memories of the 1948 meeting, however, while vivid about the success (for me) of the last
session, which I surreptitiously attended as an unregistered awestruck teen-ager, do not permit me to evaluate
the whole meeting. Others who attended, however, have since assured me that it was a very worthwhile meeting,
as usual. Canadian mammalogists have published many articles in the society’s journal, have reviewed
manuscripts for it, and have served on society committees, on the Board of Directors, and on the executive. One
outstanding Canadian mammalogist, Randolph L. Peterson of the Royal Ontario Museum, served as President
for two years from 1966 to 1968. Two prestigious Canadian mammalogists were elected as honorary members of
the ASM: Ernest Thompson Seton in 1941 and Rudolph M. Anderson in 1947.

In an attempt to quantify the increase in numbers of Canadians seriously interested in mammal study during
the past 60 years, I have used the 15 membership lists for 1920 to 1978 published in the Journal of Mammalogy
by the American Society of Mammalogists. I tallied the number from each province and territory in each list and
compiled the resulting counts as Table |. This shows that except for a marked decline during the Great
Depression of the 1930s and World War II, there was a steady increase in the number of Canadian mammalo-
gists (or at least in the number of mammalogists with Canadian addresses) up to a peak of 231 in 1972, anda
leveling off by 1978. Table | also shows that Ontario has always had an inordinately large proportion of the
total, partly because of the concentration there of mammalogists employed by the federal government. During
about the last decade, however, the numbers of mammalogists in Alberta and British Columbia have risen
significantly, while the numbers in the provinces east of Ontario and in the territories have remained relatively
low.

TABLE |—Numbers of mammalogists with Canadian addresses in the membership lists of the American Society of
Mammalogists*

Province or

territory 1920 1922 1924 1927 1934 1937 1940 1945 1948 1950 1953 1959 19€5 1972 1978
Yukon Territory l I 1
Northwest Territories l 3 l 4 4 1 3 2
British Columbia 6 5 4 10 5 2 3 4 3 13 15 13 30 25 34
Alberta 3 4 5 3 l ! 2 2 5 5 7 12 29 50 49
Saskatchewan 2 | 2 2 | l I l 2 3 5 7 9 17 10
Manitoba 4 5 4 4 4 4 4 2 3 3 3 4 6 17 22)
Ontario 11 13 16 19 15 16 20 16 31 34 44 47 50 81 75
Quebec 7 3 3 4 2 2 ! 4 2 4 11 17 20 23
New Brunswick l 5 6 7 6 6 8 5
Nova Scotia | 2 l l 2 1 3 2 4 4
Prince Edward Island 1 | 1 l

Newfoundland l | l 5 D, 5 4
Total Di 31 34 42 32 27 32 28 Si7/ 70 92) Ns) 1S4) 23) 9229

* Journal of Mammalogy 1(4): 201-212. 1920; 3(3): 203-217. 1922; 5(4): 277-293. 1924; 8(4): 327-349. 1927; 15(4): 336-354.
1934; 18(3): 390-407. 1937; 21(4): 483-505. 1940; 26(3): 327-345. 1945; 29(3): 321-343. 1948; 31(2): 220-244. 1950; 35(2):
284-315. 1954; 40(2): 278-312. 1959; 46(1): 128-180. 1965; Supplement to 54(2): 1-69. 1973; Supplement to 59(4): 1-74. 1978.

Jurisdiction over Mammals in Canada

The history of Canadian mammalogy has pretty well paralleled that of Canadian ornithology as described
earlier by Fred Cooke. I have already outlined the discovery and inventory periods. The conservation period has
been treated at length in the book Working for wildlife — the beginning of preservation in Canada by Janet
Foster (1978). This describes how dedicated government scientists and bureaucrats took initiatives to conserve
wild mammals (and birds) and their habitats, although not always for what we would consider the right reasons.
The Canadian conservation story will be dealt with later by Vic Solman.

One major difference between the legal status of mammals and that of birds in this context is that there is no
major federal act concerning Canadian mammals. Jurisdiction over mammals (and other groups of wildlife) is

14 THE CANADIAN FIELD-NATURALIST Vol. 95

not clearly stated in the British statute that serves as Canada’s constitution at present, The British North
America (BNA) Act of 1867. Asa result there isa complex situation involving divided responsibilities based on
interpretation — a typical Canadian situation! It has been broadly interpreted that the responsibility for natural
resources (including wildlife) is provincial. Interpreting its authority from Section 92(16) of The BNA Act (“In
each Province the Legislature may exclusively make Laws in relation to. . . generally all Matters of a merely
local or private Nature in the Province.”), each province and territory has passed legislation dealing with
mammals and other wildlife. In fact, most mammals in Canada are under provincial or territorial jurisdiction. In
spite of this, there are arrangements whereby federal biologists investigate problems of national interest such as
diseases, effects of pesticides, and the status and management of endangered species among otherwise provin-
cially or territorially “owned” mammals. All mammals in the territories used to be under federal jurisdiction,
but nowadays the only Canadian mammals that are considered a federal responsibility are those in national
parks and most marine mammals.

The scientifically anomalous situation whereby whales, seals, and their relatives are under the jurisdiction of
the federal government is interesting. The BNA Act, Section 91 (12), states that “. . . the exclusive Legislative
Authority of the Parliament of Canada extends to. . . Sea Coast and Inland Fisheries.” S. V. Ozere (1961) in his
background paper for the Resources for Tomorrow Conference explained that “fisheries” applies to sea
mammals as well as conventional fishes, and that the basic statute with respect to fishery regulations — The
Fisheries Act — specifically applies to marine “animals.” It is from this statute that the Government of Canada
derives its power to make regulations for the management and protection of seals and whales, although there are
other, more recent acts involving international conventions as well.

One administrative result of the interpretation of The BNA Act to include seals and whales as “fishes” is that
all Canadian marine mammalogists are employed by the federal Department of Fisheries and Oceans, unless of
course they are associated with universities. Federal, provincial, and territorial biologists cooperate in studying
Polar Bears. The single species of marine mammal that is clearly only under provincial jurisdiction at present is
the Sea Otter, which has been reintroduced to the coast of British Columbia by provincial wildlife biologists. I
suppose we should consider ourselves lucky that Beavers, Muskrats, River Otters, Minks, Water Shrews, and
even Moose have not also been declared “inland fishes” and therefore federal responsibilities!

Canadian Mammalogy from Fifty Years Ago

One of my mandates for this paper was to document some of the trends in Canadian mammalogy from 1879
to 1979. There have indeed been remarkable changes from the situation I have described for 90-100 years ago
when practically no one in Canada was studying Canadian mammals. To assess the status and nature of
Canadian mammal study at the halfway point in the century we are concentrating upon today, I turned once
again to Canada’s primary natural history journal. Volume 43 of The Canadian Field- Naturalist had as
Associate Editor for Mammalogy the aforementioned Rudolph M. Anderson of the National Museum of
Canada, who served our journal in this role from 1918 to 1955!

There were nine numbers in that 1929 volume. In the first, there was a short article about Moose and deer in
Nova Scotia. In the second, there were two notes on Smoky Shrews and Brewer’s [now Hairy-tailed] Moles by
W. E. Saunders of London, Ontario. In the third there was a reply to the Moose-deer article from E. R. Hall, a
prominent United States mammalogist; there was also a note on European Hares at Toronto by my late friend
Jim Baillie from the Royal Ontario Museum. Number 4 had nothing on mammals, but in Numbers 5 and 6 there
were notes on Moose, Black Squirrels, Water Shrews, and Little Brown Bats. In other words, there wasn’t a
great deal about mammals, but there was more than in the early days. Most of the papers in this 50-year-old
volume of The Canadian Field- Naturalist were still about birds, and there was relatively less about plants,
insects, or geology than there had been in the early days of this club.

Number 7 contains “An annotated list of the mammals of Aweme, Manitoba,” by Stuart Criddle, a member
of a famous pioneer family and later an Honorary Member of this club. In browsing through his article, I was
very interested to note, first, that he acknowledged his debt to Merriam, Jackson, and Bailey of the United
States Biological Survey for identifying his specimens years earlier, and second, that while he wrote his paper he
was aided by a grant from the National Research Council of Canada. Remember that he was an amateur
naturalist, a very competent one, and a farmer; unfortunately, the likelihood that such a deserving person would
ever receive support from the corresponding federal agency, the Natural Sciences and Engineering Research
Council, today is virtually nonexistent. This number of the journal also contained notes on the fluctuation in
numbers of rodents and on a Pygmy Shrew record. Number 8 contained a fascinating partial biography of

1981 SMITH: MAMMALOGY IN CANADA 15)

“William Couper — a pioneer Canadian naturalist” by James L. Baillie, Jr. Couper had collected a bat near
Toronto in 1851 and sent it to Professor Louis Agassiz at Harvard for identification, and he recorded notes on
mammals in various publications for the next 30-odd years. Number 8 continued with Norman Criddle’s
recollections of the abundance and habits of mammals (and birds) at Aweme during his boyhood between 1882
and 1890 and comparisons with their 1928 status. The ninth and final number for 1929 contained W. E.
Saunders’ notes on certain shrews and rodents he had trapped that summer.

When the first mammal article I came to in Volume 44 of The Canadian Field- Naturalist for 1930 turned out
to be “Barney Woodchuck,” the story of a family pet, I decided to quit on the previous high points and I
concluded my survey of mammal articles in the club’s journal of 50 years ago. Clearly they were mostly short
reports of specimens, original observations, and opinions written by both amateur and professional naturalists
from several provinces, but they revealed a much greater abundance and diversity of interest in mammals than
40-50 years earlier.

The trends in research on Canadian mammals between 1931 and 1970 were documented in Volume 86 of The
Canadian Field- Naturalist in 1972 (pages 217-221) by Anne Innis Dagg, and I recommend that you look up this
interesting compilation for the details. Not surprisingly, the increase in numbers of papers on Canadian
mammals since 1931 roughly paralleled the increased number of Canadian mammalogists I mentioned earlier.
The Canadian Field- Naturalist published the most papers in that period, but the Journal of Mammalogy, the
Canadian Journal of Zoology, and the Journal of Wildlife Management were also important media for
publication of Canadian mammal papers, the latter two especially in the 1960s. She found that university people
had done more mammal research in these decades than had other categories of researchers, and that the
University of British Columbia (UBC) was the leader by far, because of the impetus of lan McTaggart Cowan.
He was the author or coauthor of fully 49 mammal papers during the four decades Dagg surveyed!

In the early 1950s when I was considering where to do graduate studies in mammalogy, there were only two
serious possibilities in Canada — UBC or the University of Toronto. Both of these great universities are still
active centers of mammalian research, and in addition there are other Canadian universities offering excellent
programs of graduate study in at least some aspects of mammalogy: from the west towards the east, Simon
Fraser, Alberta, Calgary, Manitoba, Laurentian, Windsor, Western Ontario, Waterloo, Guelph, York,
Carleton, McGill, Montreal, Laval, Acadia, and perhaps others.

Dagg reported that the absolute numbers of Canadian mammal papers published by non-Canadians per
decade have remained fairly constant over the four decades, while the outputs by workers at universities and
federal and provincial employees have all steadily increased. Mammal papers by “unaffiliated Canadians”
(= amateurs?) were most numerous in the 1940s and fewest in the 1960s but have generally been scarce, in

.contrast to earlier times!

Canadian Books on Mammals

So far I have said a great deal about reports about Canadian mammals in the scientific periodical literature,
especially in The Canadian Field- Naturalist, and | have mentioned museum research series and other museum
publications, but I have scarcely mentioned books on Canadian mammals. During the discovery phase, reports
on Canadian mammals were all published in Europe and there were no books on Canadian or North American
mammals. During the inventory phase, some early mammal books published in the United States were of use to
the few Canadians interested in the study of mammals, and mammal books published south of the border have
continued to be pertinent to Canadian mammalogy right up to the present time. In fact, some have been
indispensable. But gradually, Canadian books have been written on Canadian mammals, and Id like to
mention a few of the most important.

In 1909 the famous naturalist Ernest Thompson Seton (whose mammal specimens appear in various
museums as collected also by Ernest E. Thompson or Ernest Seton Thompson) published his Life histories of
northern animals: an account of the mammals of Manitoba. He combined his own observations and anecdotes
from many sources to expand this two-volume work into the famous four-volume Lives of game animals which
was completed in 1929, and dealt in detail with northern carnivores, ungulates, and some rodents.

The first modern scientific inventory was R. M. Anderson’s Catalogue of Canadian Recent mammals
hopefully dated 1946, but which appeared early in 1947. The first important species monograph was R. L.
Peterson’s North American Moose, published in 1955. This was followed by two more outstanding books on
northern ungulates: Muskoxen in Canada: a biological and taxonomic review by John S. Tener in 1965, and
The migratory Barren-ground Caribou of Canada by John P. Kelsall in 1968. The most outstanding provincial
handbook on mammals, The mammals of British Columbia by lan McTaggart Cowan and Charles J. Guiguet,

16 THE CANADIAN FIELD-NATURALIST Vol. 95

was published in 1956 and revised in 1965. The next year R. L. Peterson’s The mammals of eastern Canada
appeared and provided students of mammals in the six provinces from Ontario eastward with useful keys for
identification, detailed range maps showing specimen localities, and an authoritative text. The last major book
I'll mention is The mammals of Canada by A. W. F. Banfield, which in 1974 made available a vast amount of
information on Canadian mammals collected by the author and many others.

Modern Methods and Advances in Mammalogy

Earlier I described the snap-trap revolution of 1887 or 1888 that led to the collection of large series of
small-mammal specimens and hence to the rapid growth of mammal collections. Thus it led to a great increase in
knowledge of the taxonomy and distribution of North American mammals, at first in the USA but later in
Canada too. The next important advance or revolution in mammalogy was the adoption of live-trapping, a
practice that started in the United States shortly after World War I. After being examined in detail, a
live-trapped mammal is marked or tagged for individual recognition and released. It may be observed
subsequently in the free-living state or it may be retrapped, sometimes repeatedly at intervals. A great deal of
sequential information on individual living mammals may be gathered, sometimes over periods of years. Thus
much information about the behavior, ecology, and life histories of mammals can be obtained; generally such
information was unavailable from the dead mammals that most earlier mammalogists were forced to study.
Indeed, the late, eminent United States mammalogist Tracy Storer called live-trapping “the most important
single method in mammalogy.” It has certainly been much used in Canada over the past few decades and is-
currently still in vogue and yielding worthwhile information.

A related technique that yields comparable results for one particular order of mammals but which may be
more familiar to students of birds is banding, in this case of bats. Bat-banding began with the banding of four
bats by an ornithologist in the USA in 1916, carried on in a small way there in the 1920s, and started ona
reasonably large scale in 1932 in Germany and the United States. The first Canadian bat-banding was done by
Harold Hitchcock of the University of Western Ontario in the late 1930s and even after he returned to the USA,
he continued to band bats in Ontario and Quebec until 1961. Since I took over his Canadian bat project in 1962,
my students and I have continued to monitor his banded bats (and our own) on our annual visits to their
hibernating sites. Our most remarkable recoveries of Hitchcock’s banded bats were in 1976 and 1977 when we
found hibernating two Little Brown Bats (Myotis /ucifugus) that he had banded in 1947; their minimum ages
were 29.5 and 30.5 years, the world’s record longevities for bats of all species!

There is another technique whose introduction has completely revolutionized the study of bats. In the late
1940s some North American ornithologists picked up from the Japanese the use of fine nylon mist-nets. Before
long mammalogists began using mist-nets at night to capture bats. Soon knowledge of the distribution,
behavior, and ecology of bats expanded by leaps and bounds especially in the tropics, but relatively little netting
was done in Canada. The major advances in knowledge of Canadian bat biology, largely by my colleague Brock
Fenton and his students at Carleton University, and many other advances in mammalogy generally, came in the
1970s as a result of yet another revolution, the last one I will describe.

All the major advances in mammalogy I have outlined have come about through the application of new
techniques made possible by the invention of devices or pieces of equipment. I might also mention that the
increasing use of diverse terrestrial wheeled and tracked vehicles including snowmobiles, of a variety of
watercraft, and of aircraft, both fixed-winged and helicopters, has increased the scope and practicality of field
work in remote places. Over the past decade or so, the use of increasingly sophisticated equipment of a variety of
types has been expanding, and this too has resulted ina striking increase of our knowledge of mammals. | call
this latest revolution the modern technological revolution. Mammalogists, including Canadian ones, have
lately been using radioactive isotopes, Geiger counters, tape recorders, video cameras, night-visionscopes,
fluorescent dyes, ultrasonic detectors, computers, and more. The use of tranquillizing drugs in darts shot by
guns has allowed the collection of information on live mammals somewhat larger than the ones that can be
readily trapped and handled. This technique also permits the attachment of telemetric devices that continue to
transmit information not only on the location of the mammal but also ona variety of physiological variables
and activities long after its release. The miracle of miniaturization, involving increasingly tiny transistors,
microcircuit-bearing chips, and other wonders of modern electronics, has permitted transducers of less and less
bulk and mass to be attached to smaller and smaller mammals so that we are starting to obtain information
about free-ranging mammals of almost the whole gamut of sizes. Allin all, employment of the whole array of
these devices has opened up vast new areas of field research on the behavior and ecology of shy, secretive, and
especially nocturnal mammals.

1981 SMITH: MAMMALOGY IN CANADA lie?

Modern Canadian Contributions to Mammalogy

Canadians have been prominent in the development and use of several of these techniques, from live-trapping
to bat detectors. Canadian mammalogists have also carried out some of the world’s most ambitious integrated
field research projects on large mammals. Over periods of years, teams of government biologists have collected
data on Caribou, Muskoxen, Polar Bears, Gray Wolves, Harp Seals, and others, so as to better understand and
manage our populations of these precious mammals. From the early days of the discovery period, when British
and other European naturalists were the experts on Canadian mammals, to the inventory period when
Americans took over this role, we have progressed to the present situation where we have a sizable group of
active Canadian mammalogists expert in various subdisciplines. Indeed some, suchas R. L. Petersonand Brock
Fenton, who specialize in the study of bats, are world authorities in their fields. The late Douglas Pimlott was
one of the world’s top experts on wolf biology. In the past two decades Canada has reversed the flow and has
become a supplier of mammalogical talent to the world. Among the experienced Canadian mammalogists our
federal government has loaned to Third World countries especially in East Africa to do research, teach, and
advise are Allan C. Brooks, John Tener, J. Bristol Foster, John Bindernagel, and C. H. D. Clarke.

Well, that concludes my story of some of the highlights of Canadian mammalogy. Time has permitted me to
say little of applied mammalogy, part of wildlife management, or about studies of marine mammals, or about
many other aspects of field-oriented mammal study in Canada. But I hope I have put across some of the
tremendous transitions that have taken place in this branch of biology in Canada, and how certain external and
internal situations, certain individuals, and certain inventions have stimulated the progress of mammalogy in
Canada.

The future? We still have most of our Canadian species of mammals, although the numbers of some are
precariously low, and there is still a great deal to be learned about most of them. We need to continue our
inventories of mammals in remote areas of Canada and to work out some of the details of their life histories,
physiological and behavioral adaptations, population dynamics, and management. We need to increase our
recent upsurge of interest in establishing the status of rare and endangered species of mammals in Canada and
elsewhere and to practice sound conservation measures to maintain these species and their habitats.

I referred much earlier to the somewhat premature prediction of one of the early presidents of The Ottawa
Field-Naturalists’ Club to the effect that after about eight years of existence and study they would have
discovered about all there was to know about the plants, birds, fishes, etc. of the Ottawa district. I think that
now, almost a century later, we realize that in spite of all the knowledge we have acquired, we still have a long
way to goto learn about the mammal faunas of Ottawa, of Canada, and of the world. There is still much that can
and should be done locally, nationally, and internationally, and many contributions to be made by both
professional and amateur Canadian mammalogists.

I have tried to show today that The Ottawa Field-Naturalists’ Club, mainly through its publication of The
Canadian Field-Naturalist and its predecessors for the past century, has already played a major role in
stimulating Canadian mammalogy. I hope that by continuing its almost unexcelled longevity, vitality, and
especially its valuable role in integrating the activities and reports of amateurs and professionals, it will go on
contributing to the development of Canadian mammalogy for many more centuries.

Canadian Entomology of the Last Century!

J. R. VOCKEROTH

Biosystematics Research Institute, Agriculture Canada, Ottawa, Ontario KIA 0C6

Entomology has one interesting parallel with ornithology and one marked contrast. The parallel is that both
groups of researchers seem to be concerned with numbers. The ornithologist is trying to determine the number
of individuals of the species in each habitat. The entomologists are still trying to determine the number of species
of insects in Canada. The contrast is that most of the activity and particularly the federal activity and federal
regulation in ornithology is concerned with protection; in entomology it is concerned with extermination.

As Donald Smith did, I would like to go back a little farther than 100 years. As far as I can determine, the first
insect described from a Canadian specimen was a long-horned beetle, Leptura canadensis, described by a
French entomologist in 1795. For the next 30 years or so, I have been able to find two short papers describing
specifically Canadian insects collected on arctic expeditions. Then, in 1837, what I would consider the first
major paper on specifically Canadian insects and their classification appeared.

As with mammals it was in Fauna boreali americana. The material was collected by Richardson between New
York State and the arctic posts of Canada, and it was treated by a British clergyman, William Kirby. He
described in reasonable detail 447 species and appended a list of 103 species which previously had been recorded
in Greenland and the northern parts of North America. This madea total of 550 species, and this is, in effect, the
first catalogue or checklist of Canadian insects.

By the early 1860s there were a number of moderately active amateurs in eastern Canada who published in
two general scientific journals, one in Toronto and one in Montreal. In 1857 the Professor of Chemistry at
Toronto, H. Y. Hind, and Abbé Léon Provancher, in Quebec City, each published a paper on the insect pests
of wheat. This probably marked the beginning of economic entomology in Canada.

In 1862, 10 entomologists held a meeting in Toronto. This sounds like a very insignificant thing, but this was
really the beginning of developed and consistent entomology in Canada. The meeting was arranged by C. J. S.
Bethune, a young clergyman, and William Saunders, a druggist in London, Ontario. The next year, in 1863, they
founded the Entomological Society of Canada with 25 members. Five years later they began publication of their
journal, The Canadian Entomologist. This journal became the entomological publication with the longest
continuous period of publication in North America. There had been several in the United States before this, but
at one time or another they ceased publication and disappeared. The introduction to Volume |, Number |, in
August 1868, is rather interesting.

For a long time the wielders of the butterfly net and beetle bottle in Canada have been longing for some medium of
inter-communication — some mode of telling one another what they have taken, how and where they have takenit, and
what they are in want of. This desire the Entomological Society proposes now to satisfy to some extent by the publication
of Canadian Entomologist. It is but a few years since the Society itself began asa little germ with a few members, and now
we find it rapidly growing into a goodly tree with its main trunk in Toronto, its thriving branches in London and Quebec,
and its scattered adherents all over the country...

I did not check the contributors to the first volume, but the second volume in 1870 had 180 pages and 20
contributors of whom 10 were Canadians and 10 Americans. In contrast, Volume 110 last year, the last complete
volume, had almost 1400 pages and about 250 contributors. The Society began with 25 members; at present it
has about 1000.

The Abbé Provancher in Quebec City was the first important Canadian insect taxonomist. In 1868, the same
year that the Canadian Entomological Society was begun, he began a journal, Le Naturaliste canadien, and he
was the main contributor until he died in the 1890s. He also wrote Petit faune entomologique du Canada
between 1877 and 1890. It totalled almost 3000 pages and contained descriptions of several thousand Canadian
insects, especially parasitic wasps.

Public support for entomology began in 1870 when the Ontario government gave the Entomological Society
of Canada $400 a year, provided it publish an annual report on noxious insects and continue to publish The
Canadian Entomologist. Shortly afterwards, the Entomological Society of Canada was reorganized as the

'This paper was presented on 19 May 1979 at The Ottawa Field-Naturalists’ Club Centennial Symposium entitled /00 Years
of natural history in Canada.

18

1981 VOCKEROTH: ENTOMOLOGY IN CANADA 19

Entomological Society of Ontario, which, in effect, remained the national society until 1950 when the
Entomological Society of Canada was reestablished.

In 1884 James Fletcher, an accountant in the Parliamentary Library in Ottawa, was appointed honorary
entomologist to the Canadian Government. Two years later the Experimental Farm Service was started with
Saunders, the druggist from London, appointed as the first director. Fletcher then became Dominion Entomol-
ogist and botanist. He had almost no staff, but he did have 400 observers scattered across Canada who reported
on noxious insects enabling him to publish a series of annual reports on insect pests.

Legislation aimed at controlling the introduction and spread of insect pests began in Prince Edward Island in
1883 against the Potato Beetle, and in Ontario and Nova Scotia in 1888 against the San Jose Scale.

Instruction in entomology as a subject in universities began in 1877 at the Ontario Agricultural College in
Guelph and in 1888 at the Nova Scotia Agricultural College in Truro. Gradually, many other colleges and
universities began giving formal instruction in entomology.

During this period, entomology was still largely a field for amateurs. There were a few people who did become
professional entomologists, such as Fletcher, who became Dominion Entomologist, and Bethune, the other
cofounder of the society, who became professor of zoology and entomology at Guelph early in this century.
They, as well as the other professionals, had not been trained as entomologists, and usually not as zoologists.

Then, in 1909, Gordon Hewitt, who had trained in zoology and entomology in England, was appointed
Dominion Entomologist. He was the first trained professional in the field and was a very active and forceful
man. The Insect and Pest Control Act was passed by the federal government in 1910 in an attempt to control by
regulation the spread of insect pests within the country. Hewitt, between 1911 and 1919, established federal
government entomological laboratories in all the provinces except Prince Edward Island. He also began to give
attention to forest entomology which previously had been completely neglected in Canada. The initial work by
the employees of the federal government was largely devoted to control measures, but gradually, especially from
1945 on, they began to be concerned with all aspects of insect biology, nutrition, metabolism, toxicology, cold
hardiness, population dynamics, pathogens, etc.

In 1945 the Department of Agriculture had about 140 full-time entomologists employed and in 1956 over 350.
Ido not know the present number, but there has been a substantial and unfortunate decrease from 1966 to 1976.
University entomology, of course, increased greatly at the same time, and most Canadian universities, I believe,
now have courses in entomology; 12 or more give graduate degrees in this field.

Now I would like to say something about the Canadian National Collection of Insects which I regard as the
center of entomological research in Canada (perhaps because I have been associated with it for almost 30 years).
I am sure many people would disagree.

The collection was established as the Canadian National Collection of Insects in 1917, and because of a
peculiar accident it is in the Department of Agriculture rather than the National Museum.

In 1915 it was observed that the Department of Agriculture had a small collection of insects which had, as a
nucleus, Fletcher’s private collection of 1887. This modest collection grew as a few other collections were
donated, and members of the Entomology Branch across the country contributed specimens. The Biological
Division of the Geological Survey also had a small collection. It consisted, in large part, of the material collected
during the Canadian Arctic Expedition during the First World War. Both of these collections consisted mostly
of butterflies and moths. It was considered reasonable to combine the two collections, and they were to go to the
Biological Division of the Geological Survey (now the National Museum of Natural Sciences), but in 1916 the
Parliament Building burned down and space had to be found to house Parliament until the buildings were
replaced. It was decided that the Victoria Memorial Museum would serve the purpose well, and so Parliament
moved here, probably meeting in this very hall. To make as much roomas possible for the law makers, the insect
collection went to the Department of Agriculture, where it has remained ever since.

The first full-time insect taxonomist in the Department of Agriculture was James McDunnough, a Canadian,
who had taken his graduate training in Germany. He was appointed as Head of Systematic Entomology in 1919
and published an account of the collection at that time. The following are some figures indicating growth in the
collection and staff. In 1931 there were three professional taxonomists on staff, in 1947 six, in 1956 twenty-three
— a fourfold increase in nine years — and in 1979 twenty-six, with prospects of slight increase in the future. In
1910 there was about 200 drawers of insects in the combined collections. When McDunnough was appointed,
there were 700 drawers; in 1926, 1550 drawers; in 1956, 8000; and in 1960, 10 500. Ido not have a definite recent
figure, but my estimate of current holdings would be about 17 000 drawers.

As far as individual specimens are concerned, I have no figure before 1956 whena carefully prepared estimate
was 3 million specimens. The present estimate is somewhere between 9 million and 11 million. Much of this

20 THE CANADIAN FIELD-NATURALIST Vol. 95

material was added by members of the staff of what is now the Entomology Section of the Biosystematics
Research Institute. The earlier surveys were rather sporadic. Staff members would go to parts of Canada in
which they had an interest or where it was felt insects were poorly known, and would spend a few weeks ora few
months collecting and preparing specimens for the collection. In 1947, the first and most concentrated effort to
build a Canadian national collection began, largely under the direction of T. N. Freeman. This was the
Northern Insect Survey. The Department of National Defence had established bases in the North. The DEW
(Distant Early Warning) line sites were either being built or being considered, and for some reason they found
the biting flies a serious problem. Asa result, the Department of Agriculture was asked to undertake studies of
the biting flies. This led to survey parties which, between 1947 and the early 1960s, visited about 70 localities in
northern Canada. Their task was to collect and obtain information on biting flies and other insects. The “other
insects” were a great boon, because members of the party could doa reasonable amount of work on biting flies
and then collect the other insects in which they were particularly interested.

At the same time three northern laboratories were established for the study of life histories, habits, ecology,
and control of biting flies. The main laboratory was at Fort Churchill and for a few years similar studies were
done at Whitehorse and Goose Bay. This Northern Insect Survey provided probably a million specimens of
arctic and subarctic insects, which gave the Canadian National Collection by far the largest collection of arctic
insects in the world. I always thought that probably the Russians, with their apparent emphasis on northern
development, were also collecting very large numbers of arctic insects. This seems, however, not to be the case.
In 1966 one of my colleagues, another dipterist, spent a month in Leningrad where the major Soviet collections
_ are housed, and in 1969 I was there fora week. Unless the Russians have large collections which they didn’t show
us, which I think is extremely unlikely, both of us got the impression that their total collection of arctic flies
would be the equivalent of what one reasonably competent party of two people would collect in one summer in
one locality in arctic Canada. This indicates that we have probably 40-60 times as many arctic flies as the
Russians have in Leningrad. Admittedly the arctic tundra is a very much smaller part of the total area of Russia
than it is of Canada, probably 5% as compared to 30%, but the Russians seem to have done surprisingly little
collecting in the arctic and not very much more in the northern boreal forest.

The Northern Survey has never officially been terminated, but from about 1960 on emphasis started to switch
to surveys of more southern parts of Canada. These have continued since. Occasionally rather large parties have
gone toa particular area for the summer. More recently, individual people have gone out to areas of particular
interest, and more and more tend to deal with the particular group thay interests them. The Northern Survey
parties that went out in the 1950s and 1960s made aneffort to collect insects in general. This usually meant that
the members collected groups of insects in which they were interested, and other groups that they obtained using
the same collecting methods.

In 1976, the Biosystematics Research Institute began publishing a series of handbooks of Canadian insects.
The first one was on the bark beetles. Since then, two handbooks on small families of bugs (Hemiptera), and one
oncrab spiders, have appeared. Five more on the plant bugs of the prairies, spittlebugs, mosquitoes, the genera
of Canadian caddis flies, and the fleas of Canada, Alaska, and Greenland will appear this year or next year.

One major project, I believe the largest single work that Entomology in the Biosystematics Institute has
undertaken, is almost reaching its conclusion. In 1934 C. Howard Curran, who from 1922 to 1928 was the
dipterist in the Entomology Branch, published a manual of genera of North American flies. He had been
working as a professional entomologist for 12 years. The section on one major family was contributed by
someone else, but Curran did most of the rest of the work himself. It was a magnificent piece of work, but as is
almost inevitable, it was becoming out of date. In the early sixties one of our staff members suggested that we
should revise it. We postponed this until 1965 when the United States Department of Agriculture published a
catalogue of the known flies of North America. That year or the next year we began working ona manual of the
genera of North American flies. There are about 40 contributors, and this year the first of two volumes are
finally going to press. Maybe there is something to be learned from this. What Curran did by himself in 12 years
from the time he started work as a dipterist now requires 40 people working for 13 years. It is going to bea much
bigger and better work than Curran’s. It will have well over 1000 pages, will treat 2000-3000 genera, will have
roughly that number of illustrations, and will include extensive sections on morphology, biology, classification,
distribution, and so on.

Whereas I have been concentrating in this talk on the Canadian National Collection and its work, there area
number of other major collections in Canada, most of which tend to specialize on one ora few orders of insects.
The Lyman Collection at Macdonald College began largely as a collection of Lepidoptera, but a number of the
staff members there are interested in Orthoptera-grasshoppers, crickets, and so on. It has now become a major

1981 VOCKEROTH: ENTOMOLOGY IN CANADA 21

collection of Canadian, and, in some groups, world Orthoptera. The Royal Ontario Museum has undoubtedly
the largest collection in Canada of Odonata (damselflies and dragonflies), the speciality of E. M. Walker. It has
also a large collection of Orthoptera, and, in recent years, Glenn Wiggins has developed a very large collection of
North American caddis flies. The University of Alberta has good general collections of insects and a very large
collection of North American Carabidae (ground beetles). The University of British Columbia has a good
collection of British Columbian insects. There are, of course, many others, but these are, I believe, the four
largest outside the collection in Ottawa.

I would like to give you some indication of the increase in number of known species of butterflies in Canada.
A brief digression may be desirable here. This is what we are trying to do — to get to know, and to make known,
all the species of insects in Canada. When I stop to think what “known” means, it occurs to me that it means
pitifully little. It means that one or more specimens of a species are in collections, that a name for and description
of the species have been published, and that, hopefully, the description is sufficiently diagnostic for the species to
be easily recognized by later workers. In most cases this is all we know. We all too often know little or nothing of
the life history, the nature of the immature stages, or the economic or biological role of the species. By “known” I
fear I mean that we can apply a name to the species, but even this is a beginning.

In 1894 Bethune reported 133 species of butterflies from eastern Canada. There were undoubtedly a few more
reported in the literature or known to him from western Canada, but I doubt the total would be much more than
150 butterflies from all of Canada. There are now 262 known. In the last 10 years, I was surprised to find that
about 12 butterflies have been added to the Canadian list. One of these was a new Ontario species of hairstreak,
collected, recognized, and described by Donald Lafontaine, who has been an active member of this club. Four
or five species new to southern Ontario have been collected in the last 10 years. Five new to Canada have been
taken in the Cypress Hills. Donald Lafontaine found two species new to Canada from British Columbia in
material that was already in the collection. A new species of Boloria, one of the small fritillaries, is known from
the Yukon; Ken Philip in Fairbanks, who is making a special study of northern butterflies, will be describing
this. Donald Lafontaine’s estimate is that probably another 10 will be found in the next 10 years, and he thinks
that over a long period after, that perhaps another 10 species will be found provided the climate remains
constant. If the warming trend that has been going on for some time and that may be responsible for the
northward movement of birds and insects continues, Lafontaine says we could quite easily have a number of
species moving north from the United States. In 1866a list of insects of British Columbia included 22 species of
butterflies. The last complete list that I could find was from 1927 in which 133 were recorded.

A few words now about the state of knowledge of other Canadian insects. Five years ago the Entomological
Society of Canada proposed the establishment of a biological survey of the insects of Canada. The initial plan
was to study the need for it and the feasibility, and the methods that should be used. This initial work has
resulted in the publication last month of what I think will probably be the most useful work on the Canadian
insect fauna that has yet appeared. It is called Canada and its insect fauna, and apart from having extensive
sections on Canada as an environment for insects and on the geological history of insects in Canada, it has a
lengthy section, order by order and family by family, on the Canadian insect fauna. For almost every family
there is a brief outline of the general biology and distribution in Canada, its economic importance, or its
particular role in other studies. There are tables of known and probable total species numbers for each family of
insects and for the other groups of terrestrial arthropods — the spiders, mites, and crustacea and the smaller
groups such as centipedes, millipedes, etc. One feature of major importance is that for the first time we have a
reasonably reliable estimate of the number of species of each of these groups that probably occur in Canada. For
the insects, earlier estimates were 60 000-80 000; the brief in which the establishment of this pilot project in
connection with the Biological Survey of the Insects of Canada was first proposed had an estimate of 90 000
species. The present estimate is between 54 000 and 55 000; this figure may be off by a few thousand, but it is
probably fairly reliable. Of these, 30 000 are known in Canada and 24 500 are unknown; I have mentioned
earlier what “known” and “unknown” means. We know little more than the appearance of dead specimens for
perhaps 90-95% of the “known” species. On the other hand, it is estimated that about 55% of the species have
been named and recorded, either recorded in the literature or present in the Canadian National Collection or
other collections as identified specimens. Unfortunately the immature stages of only about 10% of the species
have been described.

Three important orders of insects, as well as a few very small and less important orders, are well known, with
90% or more of the estimated species described. These three orders are Orthoptera (grasshoppers, crickets, etc),
Odonata (dragonflies and damselflies), and Siphonoptera (fleas). However, whereas over 90% of the immature
stages of 194 species of Odonata are known, only 4% of the larvae of the 180 species of fleas have been described.

22 THE CANADIAN FIELD-NATURALIST Vol. 95

For the other major orders, it is estimated that 75% of the beetles have been recorded, 45% of the flies, 70% of the
moths and butterflies, and 37% of the Hymenoptera (sawflies, ants, wasps, and bees). The main reason the
Hymenoptera appear to be so poorly known is that of the two large groups of parasitic wasps, the ichneumonids
with an estimated 7000 species, and the chalcids with an estimated 4000 species, only about 25% have been
identified and recorded. In contrast, you have heard figures earlier today of the numbers of species of mammals
and birds in Canada. A recent estimate by Bernard Boivin of the number of species of vascular plants is 3060, but
in contrast one family of parasitic wasps is estimated to have 7000.

For other terrestrial arthropods the figures vary widely. About 90% of the spiders are known, but of the mites
only 2000 of an estimated 9500 species are known (scarcely over 20%), and there are only two full-time mite
taxonomists in Canada. One has been active for about 15 years, the other for three or four — these two people
are trying to cope with some 9500 species of Canadian mites.

Larvae, the immature stages of insects, are about 10% known, but within the flies and within all other orders,
the number varies greatly with different families. All the larvae of the 74 species of Canadian mosquitoes have
been described and can be identified quite readily. On the other hand, in the Dolichopodidae, the long-legged
flies — bright metallic green flies which I am sure all of you have seen in your gardens on leaves, with wings out
at an angle — with an estimated 800 species, only five of the larvae are described, which is just over one-half of
1%.

Donald Smith has referred to a major breakthrough in the collecting of mammals, as a result of improved
techniques. I think there is an equivalent breakthrough in the collecting of insects. This was not developed in the
United States. A Swedish entomologist, Malaise, in 1934 was collecting insects in the mountains of northern
Burma; he noticed that a lot of insects flew into his open tent and collected on the window at the back. He tooka
tent, removed the two end walls, and placed one of them at the middle of the tent to act as a barrier. At the
highest point of the roof he puta collecting jar witha killing agent. Insects flew into the tent, hit the barrier, flew
up to the highest point, and ended up in the jar. His catches were enormous. In 1937 he published an account of
his trap, but it was, I think, only in the 1950s that other entomologists, particularly in Canada and the United
States, began to use them. They are now used everywhere. They collect insects 24 h a day, and even collect
during rain. Endless new designs are being produced: some are X-shaped, some are 10 m long, some are
miniatures for low-flying insects, some can be hoisted into trees. They have revolutionized the collecting of
flying insects, especially flies and parasitic wasps. The number of species new to the Ottawa area that we have
obtained from Malaise traps in the last 15 years is frightening. We consider the Gatineau Hills north of Ottawa
to be the best collected area in temperate Canada (only one area in the high Arctic is probably more thoroughly
collected). About 5 years ago one of my colleagues set two Malaise traps 50 km north of Ottawa — one on top of
a high hill, one ina sphagnum bog. Week after week he would bring his catches in, we would go through them,
and there would be more and more species we had not seen in Canada. Fortunately he gave up long before the
summer ended!

I would, finally, like to say something about the role of amateurs in Canadian entomology. As I said earlier,
they began entomology in Canada in about 1860. They were the entomologists. They continued in some
numbers until the 1940s or 1950s by which time most of them were rather old; very few of the group of amateurs
who were active in the 20s and 30s are alive today.

Ina talk on 100 years of Canadian entomology given by G. J. Spencer of the University of British Columbia in
1963, he commented on this decline in amateurs and pointed out that at least in southern British Columbia the
rising generation had far too many distractions. He mentioned a number of these — sports cars, swimming,
skiing — but surprisingly omitted television and rock groups. I think, however, that this trend may be reversing.
One group of amateurs I would like to mention in particular are those associated with Bourget College in
Rigaud, Quebec. Father André Larochelle has long been a student of the Carabidae (ground beetles) of Quebec,
and he seems to be developing an interest in insects in many of the teachers and students. One of the teachers,
Raymond Hutchison, occasionally visits the Biosystematics Institute here in Ottawa. He tells me he has had
great success in interesting students of 10-14 yr of age, at summer camps where he teaches natural history, in
various groups of insects. He trains them to recognize insects and to observe them; at the end of a few weeks with
a group of students, many of them will recognize the larger and more abundant species in a group. Their
enthusiasm is strong, I sincerely hope they retain their interest.

I do not really think it is necessary to indicate what amateurs can do. There are many things. They can rear
insects and can obtain life history data and material of unknown immature stages. They can collect insects in the
field, make observations on behavior, or, if they wish, specialize in the taxonomic study of a group of insects.
They can do almost anything a professional entomologist can do unless elaborate equipment is needed. Unlike

1981 VOCKEROTH: ENTOMOLOGY IN CANADA 23

birds and mammals, insects can be collected without a permit (except in national and some provincial parks).
There is, as far as I know, only one protected species of insect in Canada, the West Virginia White, Pieris
virginiensis, 1n Ontario. Insects are easy to collect; they are easy to preserve; one does not have to spend hours
skinning or otherwise preparing them, so the field is wide open for amateurs. Amateurs can make endless

contributions.

Geology, 1879-1979!

ROBERT F. LEGGET
531 Echo Drive, Ottawa, Ontario KIS IN7

One hundred years ago, Canada was at a singularly interesting stage in its development. Sir John A.
Macdonald had defeated Alexander Mackenzie in the general election of 1878. Sir John was determined to push
ahead with his dream of a railway spanning the continent, to bring British Columbia truly into Canada.
Visionary the idea was, because there were then only 23 000 people west of Lake Superior, whereas the
population of eastern Canada had already reached three and a half million. But it was a somewhat static
populace because roads were still in their infancy. Canals and rivers provided the main transportation routes
during summer months. The great railway era was beginning, Toronto having been connected by rail with
Montreal only in 1856.

Three years before the year that is in our minds today, in 1876, the Intercolonial Railway, linking Halifax with
Montreal, had been completed. The Chief Engineer of this great pioneer work, was Sandford Fleming (an early
member of The Ottawa Field-Naturalists’ Club). Although construction of the line did not result in any mineral
discoveries (as did other early railways), Fleming had a real appreciation of the importance of geology. This is
clearly shown in his notable book on The /ntercolonial in which he included geological descriptions of the main
divisions into which he had partitioned the line (Fleming 1876). And in the vicinity of Trois-Pistoles, excavation
of necessary cuttings was plagued by landslides in the Leda Clay (already so named by J. W. Dawson). Allearly
travelers — between Montreal and Halifax — could see the scars left by these slides.

Sandford Fleming was also the Chief Engineer of the Canadian Pacific Railway, directing the vast task of
surveying for the route finally to be chosen and supervising the expenditure of the $14 million spent on initial
construction by government, under Mackenzie. He resigned from this post in 1880, geological factors looming
large in the proceedings of the preceding parliamentary inquiry into the railway’s progress.

The Canadian Pacific Railway (CPR) Company was formed on 17 February 1881, all the work done up to
that time being transferred to it under the master agreement. On | January 1882, William Cornelius Van Horne
took up office as General Manager. What followed, completion of the line from Montreal to Vancouver in less
than four years, is part of the folklore of Canadian history. How many Canadians, however, know that Van
Horne was a keen amateur geologist? As a schoolboy he had been fascinated by Hitchcock’s Elements of
geology. He spent the first 25 cents he ever earned for the first of five exercise books into which, over a period of
five weeks, he copied the full text of this book, every illustration, and even the index! (Vaughan 1920).

He became an avid collector of fossils around Joliet, Illinois, where he lived. His collection soon included
crinoids, trilobites, brachipods, and fishes, some not then classified. In later years, nine specimens were named
after their discoverer and are today to be seen in the Field Museum of Natural History in Chicago. Canadians
think of Van Horne, later to be knighted by Queen Victoria, as one of themselves. It may, therefore, be recorded
that with a group of friends he founded the Agassiz Club of Joliet of which he was the first president. He
corresponded with many eminent geologists, including James Geikie of Edinburgh after the latter’s visit to
Canada. Even on his early inspection trips along the CPR, he would spend time searching for fossils. The
pressures of his work soon terminated this pleasant diversion but his interest in geology remained.

This is well shown by the support that he gave to the study of geology along the entire route of the new railway
during its construction, by officers of the Geological Survey of Canada, notably by A. R. C. Selwyn, G. M.
Dawson, and Robert Bell, all of whom were later to be directors. Their reports will be found in Survey
publications, but Principal J. W. Dawson of McGill published a general account in the Quarterly Journal of the
Geological Society of London, after presenting his paper on the CPR to the society on 23 April 1884 (Dawson
1884). Dawson mentions in this paper not only the work of the Survey geologists but also the remarkable
collection of Silurian fossils collected from the banks of the Red River by J. H. Panton, an amateur geologist,
even in those early days and in sucha pioneer community. This tradition of combined official and unofficial,
professional and amateur interests in Canadian geology has continued throughout the century now being
celebrated. It is only to be regretted that few of the amateurs have left any record of their work, but of their
enjoyment in their hobby we may be assured. One would like to know more about the work of Charles F.
Newcombe (1851-1924) on Vancouver Island, and of others like him.

'This paper was presented on 19 May 1979 at The Ottawa Field-Naturalists’ Club Centennial Symposium entitled /00 Years
of natural history in Canada.

24

1981 LEGGET: GEOLOGY IN CANADA 25

Such was the interest of the public in both railways and geology in those early days that there was published in
1879 An American geological railway guide “giving the geological formation at every railway station, and its
altitude above mean high water, with notes on interesting places on the routes, and a description of each of the ~
formations.” The author was James Macfarlane, who acknowledged the assistance of the State Geologists “and
other Scientific Gentlemen.” A second edition was published in 1885 to which George M. Dawson contributed a
32-page section on the Dominion of Canada. The notes accompanying the interesting tables of railway routes,
with mileages and elevations, provide (in the smallest legible type) a fascinating guide to the geology of Canada
as it was then known (Macfarlane 1885).

It was during the construction of the Canadian Pacific Railway that, in a cutting near Sudbury, Tom
Flanagan, a blacksmith in one of the construction gangs, noticed some rock stained with iron oxide and on
digging found copper sulfide. This was in 1884 and proved to be the beginning of the discovery of the vast nickel
ore deposits, a geological phenomenon now known around the world. It is interesting today to note that so little
was its value originally realized that a patent for the lot was granted the next year to a certain Thomas Murray
for $1 a year! (Gibbon 1935).

In a similar way, when the Temiskaming and Northern Ontario Railway (now the Ontario Northland
Railway) was being constructed as a development line, in the face of much skepticism, a similar discovery was
made in 1903 near the crossing of the Montreal River. Fifty years ago when in Cobalt, I was assured by an
eyewitness that a blacksmith threw a hammer at a jackrabbit, missed the animal, but dislodged a piece of
bedrock, revealing an unusual-looking material which proved to be native silver, the beginning of the Cobalt
boom, and the opening up of so much mining in this part of northern Ontario and the adjacent part of Quebec
(Legget 1973).

With geological discoveries such as these “making the news,” it is not surprising to find that in the closing two
decades of the last century, and around the beginning of the 20th century, geology in Canada was a subject of
wide and general interest. A succession of geological disasters served to intensify this public interest. In October
1881, two years after The Ottawa Field-Naturalists’ Club was founded, one of Canada’s most remarkable
landslides took place on the Thompson River in British Columbia, near the small town of Ashcroft. Terraces
above the incised river valley were being irrigated with water from the mountains above, impounded by
amateur-built dams. One of these failed. Resulting seepage through the steep slope above the river and the new
railway track caused a slope failure which precipitated 60 million yd? (45 X 10° m3) of soil and rock into the
river, blocking its flow completely. The artificial dam was 160 ft (48 m) high. Only when it was overtopped by
the impounded river water was flow resumed downstream, initially with a mighty flood that it is difficult to
imagine (Stanton 1897).

Twenty-two years later, on the morning of 29 April 1903, the great rock fall occurred at Frank, Alberta. This
took 70 lives and buried well over | mi(1.6 km) of the Crow’s Nest Railway (Report 1912). News of this disaster
quickly reached eastern Canada, but there had been devastating local landslides which must have had even
greater impact upon the public conscience. All were in the so-called Leda Clay, now better called the Champlain
Clay, well known for its unusual sensitivity. In 1894 occurred the slide at St. Albons involving the rapid flow
sliding of 600 million yd3 (450 X 10° m3); in 1895 that at St. Luc de Vincenne in which five lives were lost; and in
1898 another at St. Thuribe in which a mere 3.5 million yd3 (2.7 X 10° m3) moved. Almost in the outskirts of
Ottawa, on the Liévre River, occurred in 1908 the tragic slide at Notre Dame de la Salette in which 33 lives were
lost (Mitchell and Markell 1974).

When, therefore, the headquarters of the Geological Survey of Canada (GSC) was moved from Montreal to
Ottawa, few residents of this city had to be told what the science of geology was. The Survey had started its work
in 1842 in Montreal as the one-maneffort of William Logan. Under his inspired leadership it had grown in staff
and activity, and had started its own museum. So well was its work appreciated that when the perfectly logical
move to the capital city of Ottawa was proposed, the strongest objections were raised by Montrealers (Zaslow
IO):

It is wryly humorous today to read of the Director at the time, A. R. C. Selwyn, saying that “removal to
Ottawa could not fail to operate in every respect most prejudicially.” But the Geological Survey Bill was passed
by Parliament in 1878 and the move was confirmed. Objections continued. Principal J. W. Dawson of McGill
wrote personally to Sir John A. Macdonald. The Montreal Natural History Society objected strongly in 1881
and the Montreal Board of Trade went so far as to send a delegation to Ottawa in an attempt to stop the move.
The transfer of staff, equipment, and museum collections was made, however, in April and May of that same
year. Principal Dawson received a grant from the Redpath family to start a new museum at McGill. And when
the Geological Survey Museum was reopened in Ottawa, it attracted 9549 visitors in its first year, as compared

26 THE CANADIAN FIELD-NATURALIST Vol. 95

with 1652 in its last operative year in Montreal, proof indeed of the well-developed interest here in natural
history, and especially geology, 100 years ago.

The Geological Survey moved into the fine gray stone building still standing at the corner of Sussex Drive and
George Street, adjacent to the market area. The building served originally as a hotel but had been extended and
was refurbished for Survey use. Some members of the Survey staff worked there until 1959, an occupation of
almost 80 yr: the Survey moved in that year to its own building on Booth Street. The old George Street building
is, therefore, by far the oldest “scientific building” in Ottawa. Happily well preserved, should it not be today the
headquarters for Ottawa’s many scientific societies? In its day, it was the scene of many dramatic encounters, as
all who know the history of our distinguished Survey will appreciate. Many famous geologists passed through
its doors. Early members of The Ottawa Field-Naturalists’ Club with geological interests must have been regular
visitors, for it will have been observed that the Survey moved to Ottawa two years after the founding of the club.
Throughout the club’s history, therefore, those interested in this particular branch of natural history had
available to them the possibility of association with professional geologists, with a geological museum and
library. It is not surprising, therefore, to find geology occupying a prominent place in the early years of the club’s
activity.

Before glancing at this, however, that earlier reference to the Montreal Natural History Society may be noted.
This society was founded ina small way in 1827 and by 1858 had its own spacious quarters and museum at the
intersection of Cathcart and University streets. So well established was the society that it was one of the hosts
when, in 1857, the American Association for the Advancement of Science held its annual meeting in Montreal.
Sessions were held in the Court House, the Natural History Society tendering a soirée as a part of the social
activities. Even older, however, was the Literary and Historical Society of Quebec, founded by that outstanding
(but often misjudged) Governor, the ninth Earl of Dalhousie, the founder also, with John By, of our own city of
Ottawa. In the first volume of the Quebec Society’s journal, published in 1829, there were two papers on geology
(Baddeley 1829; Bayfield 1829).

Down by the sea, the Nova Scotian Institute of Science was founded in 1862 by the amalgamation of the
Halifax Mechanics’ Institute (founded in 1830) and the Halifax Literary and Scientific Society (founded in
1839). The Nova Scotia Museum had its originin the museum started by the Mechanics’ Institute in 1831, a large
proportion of the early exhibits being rocks, minerals, and fossils. The museum participated in the 1867
exhibition in Paris, under the leadership of its curator, the Reverend Dr. David Honeyman, the Presbyterian
Minister at Arisaig, Halifax, who became an authority on the Silurian fossils of his area(R. Grantham, personal
communication).

The Ottawa Field-Naturalists’ Club had its antecedents also, mainly geological. As is well known, Ottawa had
its origin as the main construction camp of the Corps of Royal Engineers when they came in 1826 to start the
building of the Rideau Canal, selected by Lieutenant Colonel John By and approved, on the spot, by Lord
Dalhousie. Named Bytown unofficially but in honor of the Superintending Engineer, Bytown it remained until
the official change in name to Ottawa in 1855. It is difficult today to credit that even at that early date, less than
30 years after the first tree was cut in the unbroken forest, there was here an established Silurian Society. But
Hunter recorded in his Ottawa scenery, published in 1855, that in the library of the Silurian Society he had seen
“a small, but very beautiful illustrative collection of geological specimens; for this the people of Ottawa are
mainly indebted to Mr. Billings...” (Hunter 1855). The original Natural History Society was formed in 1863 but
in 1869 amalgamated with the Mechanics’ Institute (founded originally in 1847) to form the Ottawa Literary and
Scientific Society from which our club was an offshoot. All the bodies mentioned regularly had public lectures
on geology.

There was, therefore, a well-established geological tradition in Ottawa when the club started its work, and so
it is not surprising to find that geology looms large in the early records of the club. As we have been so pleasantly
reminded in Trail & Landscape, by 1893 the club members had prepared a geological map of the Ottawa area
(Reddoch 1979). Members assisted in the preparation of a report on the Stewart Quarry, near Rockland, which
contributed to the settlement of a case before the Exchequer Court of Canada. The club advised the Ottawa
Granite Company about a local outcrop of quartzite which led to successful use of this deposit. Geological field
trips were frequent, quite a number being to sites within the limits of the city, owing to the state of the city’s
development at that time. A favorite location for the collection of fossils was “The Heap” south of Beaver
Meadow, west of Hull, a pile of broken Trenton limestone excavated from the nearby railway cutting. Thirty
specimens were obtained during a single visit in 1900.

With the Geological Survey now well established in Ottawa, with railway construction across the Dominion
of Canada regularly bringing to light new facets of geology, and with occasional natural disasters providing

1981 LEGGET: GEOLOGY IN CANADA ny

tragic reminders of geological hazards, it is small wonder that those early years saw a real flowering of geology as
a major club activity. At the time of the club’s formation, there was a lively interest in the geology of cities. In
1885, for example, there was published in Philadelphia a well-illustrated volume entitled Town geology: “the
Lesson of the Philadelphia Rocks; Studies of Nature along the Highways and Byways of a Metropolitan Town,”
the author being Angelo Heilprin of the Academy of Natural Science of Philadelphia (Heilprin 1885). In
England, there had been published another fine volume with exactly the same title, Town geology, in 1877 (bya
publisher with the Dickensian name of Daldy, Isbister, and Company). The author was the Reverend Charles
Kingsley, eminent Anglican divine who once visited Ottawa, better known, perhaps, as the author of Westward
ho! and The water babies (Kingsley 1887).

There was no Canadian book with the same title, but in 1900 one of the most colorful presidents of this club,
H. M. Ami, presented to the Royal Society of Canada a fascinating paper entitled “On the Geology of the
Principal Cities of Eastern Canada.” Ami pleaded fora continuing study of the geology beneath city streets for,
as he said: “What the drill has to penetrate in any one of our larger centres of activityin Canada. . . isa question
not only of interest but also of economic value” (Ami 1900). If only Canadian engineers had given heed to that
sound advice, this country could today have been a world leader in the study and recording of urban geology.
But as the young country hastened on its way, geology fell into neglect in engineering, just as it did in natural
history circles.

Ami, a graduate of McGill, joined the staff of the Geological Survey in 1882. A palaeontologist of note, he
must have made life a little difficult at times for his superiors. It is recorded of him that he was absent from his
official duties for 284 working days in the space of four years. His wife was a lady of means and so he was able to
undertake private ventures such as leading an excursion to the south of France to study cave remains of early
man. Later, in Ottawa, he rented a room in a downtown building in which to display his finds. He resigned from
the Survey in 1910 because of ill health. It would have been interesting to hear his pithy comments upon all the
troubles that beset the new building into which the Survey moved its headquarters in that same year, 1910.

This was the Victoria Museum in which this meeting is being held. The very odd appearance of the front
entrance is explained by the fact that the extension for the entrance hall was designed, and built, four full storeys
in height, crowned by a magnificent castellated tower (“Victorian wedding-cake architecture at its worst” as it
has been described). The contractor for the building warned that the heavy masonry walls would exert too great
a bearing pressure on the foundation beds, but his warning was not heeded. Excessive settlement took place
before the building was occupied. The front extension broke away from the rest of the building, and there was a
gap (near the roof) of 14 in. (35 cm) when the contractor was ordered to remove the tower and the upper two
storeys before complete failure occurred.

It was ironic that this classical failure should have taken place in the building erected to house the Geological
Survey and its museum, since the cause of the trouble was geotechnical. By chance, the building was located on
one of the “pockets” of Leda Clay that are features of the urban geology of Ottawa. The clay was loaded almost
to the point of failure. Settlements were to be expected. Modern geotechnical studies have been correlated with
the long-term records of settlement, which today is negligible (Crawford 1953). The interior of the building has
been reconstructed so that the wild tales circulating around Ottawa for 60 years — about the building’s
imminent collapse, about the great fault beneath the building which was supposed to be the cause of the trouble
(this, from an eminent hard-rock geologist!) — should now be set at rest for ever.

The building is, however, now known around the world in geotechnical circles as a classical near-failure. The
excitement surrounding its construction and necessary alteration must have been a major topic of conversation
at the time, and further evidence of the importance of applied geology. But the war which started in 1914 soon
put all such local problems in the background of public thinking. Reconstruction during the 1920s followed the
Armistice of 1918. The slow rebuilding of the economy in the 1930s followed the stock market boom and
collapse. Then came the second World War, its termination coinciding with the end of the second third of the
century being marked by these lectures. Just as the first third was full of geological excitement, so the second
third witnessed a remarkable decline in public and private interest in the science.

Mineral prospecting continued unabated, in the inter-war years, a life-saver for many men in the years of
depression. Some new mines were brought into production, notably the Eldorado uranium mine on Great Bear
Lake and the gold mines on Lake Athabasca and Great Slave Lake. The Geological Survey was put under severe
strain during the 1930s, even its official name disappeared at one time. But its work went on, its loyal staff
continuing their work despite all difficulties. One name must be mentioned from those trying years, that of a
great lady who was an ardent member of this club, the late Mrs. Forsey (mother of Senator Eugene Forsey). She
joined the staff of the Survey in 1913 and served as its librarian from 1918 to 1941. Through all the difficult years,

28 THE CANADIAN FIELD-NATURALIST Vol. 95

the Survey library was a lighthouse of hope to all interested in the science and the best source of geological
information in Canada. How Mrs. Forsey managed to keep the library operating to its usual high standard was
one of the mysteries of those years, although I do know that she thought nothing of buying small items of office
equipment with her own money if the economies of the time made official purchase difficult. I can testify to the
kindness and helpfulness which she showed me, a young visitor from Montreal, when I came seeking informa-
tion on my occasional visits to Ottawa.

Geological instruction continued at universities but under severe restrictions. None of these deterrents,
however, explain the great decline in public interest in this branch of natural history. Confirmation of this
subjective impression is given by even a cursory study of the club’s own journal, The Ottawa Naturalist until
1919, and The Canadian Field-Naturalist thereafter, published throughout this long period as a monthly
journal (until 1943). Early volumes carried at least two or three good geological papers, regular items of
geological interest from around Ottawa and, at one time, even a summary of the summer program of the
Geological Survey of Canada. In the third volume of The Ottawa Naturalist appears the first annual report of
the geological branch of the club, and in the fourth there are even annual reports from two branches, those for
geology and mineralogy, and for palaeontology. The Geological Branch was still reporting in 1912. A third ofa
century later almost all references to geology had disappeared, only one or two occasional papers on fossils
appearing, and this, despite the long and loyal service of Fred J. Alcock as Associate Editor for Geology.

There must be an explanation for this significant change which seems to be worthy of detailed study, rather
than the necessarily brief survey made for this lecture. Such study as has been made convinces me that the blame
must be attributed to high-school education. Once science was introduced into school curricula, chemistry,
physics, and biology in some way unknown to me got priority of treatment. A good case can be made for the
suggestion that geology, the science of the world around us, should be the first science to which young minds are
introduced. Its study leads logically into the study of biology, of chemistry, and even of physics. But it did not
receive this recognition. Even today, it is rare indeed to find any course on earth science, as such, in the
curriculum of any North American high school. It is sometimes said to be included in more general courses such
as those on social studies, or geography, but I am firmly of the opinion that a good introductory course in
geology has a rightful place in the science curricula of all high schools, not only as the best of all introductions to
science but also as a liberal influence in itself. As but one example of this continuing neglect, the Ontario
Department of Education produced in 1971 a beautifully printed brochure of 36 pages describing a new course
on urban studies in which the word geology does not appear once!

Despite this official neglect, there are some encouraging signs. There are many devoted teachers who do
appreciate the value and interest of geology for young minds and who are teaching it in their classes. The
American Geological Institute has had a major program related to high-school teaching and has produced much
valuable material for teaching aids. In Canada, Professor Gordon Winder initiated a fine summer training
course for interested high-school teachers, receiving support from industry for this effort at the University of
Western Ontario. Similar courses are now available in other places, so the tide may be turning and we may yet
see geology again recognized as an important part of natural history.

It is significant to note that the Geological Association of Canada (GAC) was not started until 1947. Prior to
that the only national meeting place for professional geologists was at the small annual meetings of the Royal
Society of Canada (to which, incidentally, this club used regularly to send a delegate). The exclusive member-
ship of the Royal Society, junior to this club by four years, meant that younger geologists had to turn to British
or United States journals as outlets for their writings. Today the GAC has a membership of about 3000 and two
regular publications that supplement the Canadian Journal of Earth Sciences, published by the National
Research Council, which has already made its mark, inits 15 yr, as one of the leading geological journals of the
English-speaking world.

On the more general front there has been the welcome establishment of many local groups of “Rock Hounds.”
I use the colloquial expression to indicate, with appreciation, the many clubs in Canada today of mineral and
rock collectors and gem fanciers, almost all of which are of post-war origin. One of the few exceptions is the
Walker Mineralogical Club associated with the University of Toronto, with its own distinguished publication.

One of the post-war turning points, I feel sure, was the series of popular lectures on geology delivered over the
national radio network of the CBC in the late 1950s by David Baird and his later lectures on television. The radio
lectures were reprinted as an attractive booklet. In both spoken and written form these excellent presentations
must have awakened the interest of many Canadians to the fascination of geology. Baird followed up his radio
and television talks with his series of pocket guides, first (in 1960) with A guide to geology for visitors in
Canada’s national parks, whichis a booklet of use and value anywhere in Canada (Baird 1960), In the same year

1981 LEGGET: GEOLOGY IN CANADA 29

the Geological Survey published The story of the mountains in Banff National Park by Helen Belyea, also in
pocket form (Belyea 1960). In succeeding years a series of similar guides by Baird to the geology and scenery of
all the major national parks (and the National Capital District) were published, so successful that some have
been reprinted by a private publisher in association with the Geological Survey. W. H. Mathews of the
University of British Columbia has written, and the GAC has published 4 popular guide to the geology of the
Garibaldi Lake area, that wonderful playground for residents of Vancouver (Mathews 1975).

Many of the regular publications and maps of the Geological Survey are naturally of use to amateur
geologists. Special mention, however, must be made of the 16 guides to Rocks and minerals for the collector
written by Ann Sabina of the Survey staff, some in association with joint authors (Sabine 1963). The first four
gave general coverage for the whole country, subsequent guides being for specific areas. The Survey is also
responsible for the distribution of the 87 guide books that comprise the series prepared for the Twenty-third
International Geological Congress held in Canada in 1972. The tours arranged for participants in this great
gathering covered the country from the High Arctic to the southern tip of Ontario; so do the guide books.
Copies are still available, still in regular demand; and a list is available from the Survey. We now have a special
highway map for Ontario showing the main features of the geology of the province. And suites of minerals and
rocks are readily available, about 6000 sets being sold every year to prospectors and amateur geologists. There
is, therefore, a wealth of information about the geology of all parts of Canada readily available in convenient
form for purse or pocket for the use of all who travel and who want to do more than just admire the scenery.

For those who dwell in cities and are limited in the traveling they can do, there are now available, from coast
to coast, museums, large and small and of varying degrees of excellence, almost all with geological exhibits.
After the museum in this building, pride of place must be given to the Royal Ontario Museum, established in
1914, which now has one of the world’s leading mineralogical collections, having specimens of about 1800 of the
2400 minerals now known. It has also one of the few original copies of William Smith’s geological map of
England and Wales. Its appreciation of the interests of the average citizen was well shown during the
construction of the first section of the Toronto subway when a special exhibit was arranged showing the
geological strata encountered in excavation for the subway beneath the streets of the city, and the methods
followed in excavation, with actual samples of the rocks and soils encountered.

From the many other museums, large and small, two have been of special attraction. At Drumheller, Alberta,
I was surprised to find a most excellent display of the local geology in the local museum, including a complete
skeleton of one of the dinosaurs (the first I believe) excavated in the valley of the Red Deer River. On the other
side of the country is the island of Grand Manan, guardian of the entrance to the Bay of Fundy. A small island,
15 X 7 mi(24.1 X 11.3 km), witha population of less than 3000, it yet has a fine little museum, established by the
islanders in 1967 as acentennial project. It, too, has a first-class exhibit of the island’s geology owing, largely, to
the enthusiasm of the “naturalist-extraordinary” of the island, Elmer Wilcox, to whom the whole realm of
nature is an open book.

There are books, too, about the geology beneath the streets of some Canadian cities, not very many as yet but
some of which I know are exemplars of what every city in Canada should have. G. H. Eisbacher of the
Geological Survey has written, and again GAC has published, Vancouver geology, a popular guide to the
geology of Vancouver (Eisbacher, 1977). The “Saskatoon Folio” is now justly famous far beyond the borders of
Canada (Christiansen 1970). Toronto has, to date, only a well-produced leaflet about its geology, and Montreal
a splendid, but professional, 244-page official report, not easy reading for the amateur (Clark 1972). Ottawa has
been here in the lead, thanks to the publications of this club.

In 1962, The Canadian Field- Naturalist published as a special feature, and the club later reprinted, a splendid
“Guide to the geology of the Gatineau-Liévre district” by Donald Hogarth, one of the best guides of its kind
known to me (Hogarth 1962). Hogarth has continued his studies of minerals and mining on the “other side of the
river,” one product being his delightful booklet on “Pioneer mines of the Gatineau region, Quebec” (Hogarth
1975).

The best of alll have kept to the last so that I may conclude with my tribute to one of the greatest ladies I have
ever been privileged to know, Alice Wilson. There is no need for me to remind this audience of her many
distinctions, of her long membership of this club, covering half of our century, of her devoted service to the
Geological Survey continuing, quite unofficially, until her 80th year. But | would remind you that in 1956 The
Canadian Field- Naturalist published a special monograph issue (and later reprinted) “A guide to the geology of
the Ottawa district, by A. E. Wilson (Wilson 1956). Its writing was a labor of love, as you can tell when reading
it, but one had to be with her in the field to realize the way in which she could make any aspect of geology
interesting and arresting. She indeed knew, and showed in so many ways that geology is, in Charles Kingsley’s

30 THE CANADIAN FIELD-NATURALIST Vol. 95

phrase, The people’s science. So should it be thought of today; so must it be thought of if we are to be true
guardians of the heritage that is ours in this land of Canada.

Knowing Alice Wilson as I did, I feel sure that she would have agreed that the best way to bring toa close this
brief review would be for me to use not my own words, but rather the words which Charles Kingsley used to end
Town geology almost exactly 100 years ago;

If | shall have awakened any townsmen here and there to think seriously of the complexity, the antiquity, the grandeur,
the true poetry, of the commonest objects around them, even the stones beneath their feet . . . if I shall have helped to
open their eyes that they may see, and their ears that they may hear, the great book which is free to all alike, to peasant

as to peer, to men of business as to men of science, even that great book of nature which is, as Lord Bacon said of old,

the Word of God revealed in facts, then I shall have a fresh reason for loving that science of geology, which has been my

favourite study since I was a boy.

Acknowledgments

I gratefully acknowledge the provision of information by the following: A. Baracos, R. Grantham, H. R.
Gulliver, P. Harker, and D. C. Tibbetts; in addition to that provided by the books referenced and, in some case,
by their authors.

Literature Cited

Ami, A. M. 1900. On the geology of the principal cities in Eastern Canada. Proceedings and Transactions of the Royal
Society of Canada, Second Series, 6(4): 125-173.

Baddeley, F. H. 1829. Geology ofa portion of the Labrador coast. Journal of the Literary and Historical Society of Quebec
1: I-

Baird, D. M. 1960. A guide to geology for visitors in Canada’s national parks. Department of Northern Affairs and National
Resources, Ottawa. 144 pp.

Bayfield, H. W. 1829. Outlines of the geology of Lake Superior. Journal of the Literary and Historical Society of Quebec 1:
71-

Belyea, H. R. 1960. The story of the mountains in Banff National Park. Geological Survey of Canada, Ottawa. 42 pp., 35
plates.

Clark, T. H. 1972. Région de Montréal area. Geological Report Number 152, Ministry of Natural Resources, Geological
Exploration Service, Quebec. 244 pp.

Christiansen, E. A. 1970. Physical environment of Saskatoon, Canada. Saskatchewan Research Council and National
Research Council, Ottawa. 68 pp.

Crawford, C. B. 1953. Settlement studies on the National Museum building, Ottawa. Proceedings of the Third International
Conference on Soil Mechanics and Foundation Engineering, Zurich |: 338-345. (Also available as Research Paper Number
11 of the Division of Building Research, National Research Council, National Research Council 3071. Ottawa.)

Dawson, J. W. 1884. Observations on the geology of the line of the Canadian Pacific Railway. Quarterly Journal of the
Geological Society of London 40: 376-388.

Eisbacher, G. H. 1977. Vancouver geology; a short guide. Geological Association of Canada, Vancouver. 51 pp. (Original
edition 1973).

Fleming, S. 1876. The Intercolonial; a historical sketch. Dawson Brothers, Montreal.

Gibbon, J. M. 1935. Steel of empire. The Bobbs-Merrill Company, Indianapolis and New York. 423 pp. (see p. 195).

Heilprin, A. 1885. Town geology: the lesson of the Philadelphia rocks. Published by the author, Academy of Sciences,
Philadelphia. 137 pp., 7 plates.

Hogarth, D. D. 1962. A guide to the geology of the Gatineau-Liévre District. Canadian Field-Naturalist 76: 1-55.

Hogarth, D. D. 1975. Pioneer mines of the Gatineau region, Quebec. Bytown Beavers Publishers Registered, Ottawa. 44 pp.

Hunter, W. S. 1855. Hunter’s Ottawa scenery; Canada West. Published by the author, Ottawa. 19 pp., + plates.

Kingsley, C. 1877. Town geology. Daldy, Isbister, and Company, London, England. 239 pp.

Legget, R. F. 1973. Railroads of Canada. Douglas, David, and Charles, Vancouver. 255 pp.

Macfarlane, J. 1885. An American geological railway guide etc. Second edition. D. Appletonand Company, New York. 83
pp. (With a special section by G. M. Dawson on “The Dominion of Canada.” pp. 51-83.)

Mathews, W. H. 1975. Garibaldi geology; a popular guide to the geology of the Garibaldi Lake area. Geological Association
of Canada, Vancouver. 48 pp.

Mitchell, R. J., and A. R. Markell. 1974. Flowsliding in sensitive soil. Canadian Geotechnical Journal 11: 11-31.

Reddoch, J. 1979. Favourite OFNC excursion places. Trail & Landscape 13: 71-96.

Report of the Commission appointed to investigate Turtle Mountain, Frank, Alberta. 1912. Memoir Number 27, Geological
Survey of Canada, Ottawa.

Sabina, A. 1963. Rocksand minerals forthe collector: Sudbury to Winnipeg. Geological Survey of Canada, Ottawa, Paper
Number 63-18. 69 pp.

Stanton, R. B. 1897. The great landslide on the Canadian Pacific Railway in British Columbia. Minutes of Proceedings of
the Institution of Civil Engineers, London, England 132: 1-20.

Vaughan, W. 1920. The life and work of Sir William Van Horne. Century Company, New York. 388 pp.

Wilson, A. E. 1956. A guide to the geology of the Ottawa District. Canadian Field-Naturalist 70: 1-68.

Zaslow, M. 1975. Reading the rocks. Macmillan Company of Canada, Toronto. 599 pp.

Federal Wildlife Conservation Work in Canada in the Past 100 Years!

VICTOR E. F. SOLMAN
Canadian Wildlife Service, Ottawa, Ontario KIA 0E7

North America was discovered by Europeans who sought the riches of the East Indies. Having encountered
Canada they found that there were great resources there of fish, trees, and fur. They began the harvest of those
resources and transported the products to the markets of Europe. Beaver became symbolic of Canada because
its pelt was one of the first export products to Europe. Wildlife was important from the beginning of Canada.
Most of the people carrying out the exploitation of the resources came from Europe and to them the stocks
available in North America seemed almost without limit. That tradition of abundance lasted fora long time and
indeed still persists in parts of Canada.

As Canada became settled and heavy use was made of wildlife in small areas, game laws were introduced and
have been in effect in parts of Canada since the early 1700s. At the time the provinces were united under the
British North America Act, provincial control of wildlife resources was well established and there was little
federal concern. In fact the whole matter was taken so much for granted that there is no mention of wildlife in the
British North America Act.

That did not indicate a complete lack of federal interest, however, because there were some federal activities
on behalf of wildlife as early as 1887 when the islands and shoreline at the north end of Last Mountain Lake,
Saskatchewan, were created a bird sanctuary by Order-in-Council on 8 June 1887. That was one of the first
activities of the kind in North America. On 23 June 1887 Canada’s first national park was authorized under the
Rocky Mountain Park Act. There were clauses in that legislation relating to wildlife protection, but the main
intent was the creation of a tourist resort and the preservation of a group of hot springs and associated scenery
rather than the management of wildlife. As the parks organization expanded, progressively more attention was
paid to wildlife, but most national parks were created primarily to preserve other values.

On 12 May 1909 a bill to establish a Commission of Conservation in Canada was passed in Parliament. That
Commission, in which both federal and provincial officials participated, was nonpartisan. It was a unique,
autonomous, objective body set up to explore questions pertaining to natural resource conservation in Canada.
It was set up to be responsible directly to Parliament and was so arranged that it would report to the House of
Commons from time to time through the Minister of Agriculture although it was not responsible to him or to
any other minister or government department. The Commission could deal with any and all resources and as
such had many responsibilities beyond those involved in wildlife and related renewable resources. The
Commission kept in touch witha rather similar organization in the United States and considered international
concerns as well as those affecting Canada alone. Among the early projects undertaken by the Commission were
an analysis of fishery resources in Canada, a compilation of statistics on commercial fishing, and summaries of
dominion and provincial laws and regulations governing fisheries with special reference to the oyster and lobster
industries. A paper on fur-bearing animals and how to prevent their extinction was considered in 1910. That was
mainly aimed at the fur-trapping industry. The fur trade, like the fisheries, was a major industry in Canada at
that time. There were also considerations of fur farming which was just beginning to develop. During the first
three years of the Commission’s work, most of the studies involved the economics of fish and fur-bearing
animals. The emphasis was on utilization rather than conservation. An investigation in 1921 suggested that the
Commissions work overlapped that done by other government agencies. During the period of Commission, its
pioneering activities had come to be well done by government agencies in forestry, water power, agriculture,
interior, marine and fisheries, mines, publicity, and external affairs. Wildlife was not included in the resources
listed in that review, possibly because the review committee was not particularly interested in wildlife. The
Commission ceased operations in 1921 after 11 years of activity during which it had helped to encourage public
interest in wildlife and conservation and the development of public sentiment in favor of that activity.

Between 1850 and 1900 a number of influential persons in North America had become concerned about the
declining supplies of Passenger Pigeons and Bison. When they looked ahead they could see the end of certain
species unless actions were taken to prevent it. One group of birds of concern to many people was waterfowl
including ducks, geese, and swans. They were shot for food and for sport and indeed were a part of the

'This paper was presented on 19 May 1979 at The Ottawa Field-Naturalists’ Club Centennial Symposium entitled /00 Years
of natural history in Canada.

31

32 THE CANADIAN FIELD-NATURALIST Vol. 95

commercial food supply in many parts of the country. Although provinces and states had been legislating for the
protection of wildlife for more than 100 years, they found that they could not effectively deal with a migratory
resource like waterfowl that moves from the breeding grounds to the winter grounds in the fall and returns in the
spring, and is harvested in each jurisdiction through which it passes. In that situation it is easy to overharvest the
stock. Not only the annual increment was being taken each year buta part of the parent stock as well. There was
also shooting in the spring and on the breeding grounds which was very destructive of the resource. Farsighted
people then tried to have the harvest reduced by encouraging the provinces and states to cooperate. That was
easy to suggest. Human nature being what it is, it was very difficult to bring about. When the birds were ina
province or state, that province or state tried to secure the best possible opportunity for its citizens to harvest a
“fair” share of the resource, knowing that other political entities would also be involved in the total amount of
the harvest. Invariably each political entity took its full share of the harvest and perhaps a bit more, and the
decline continued.

There was an attempt in the United States to impose federal control on the system, but that was challenged by
some states as unconstitutional and the whole matter was referred to the United States Supreme Court. It was
clear that federal control in that way was unconstitutional in the United States and that a similar attempt in
Canada would probably be unsuccessful for similar reasons. Farsighted people in Canada and the United States
sought a solution that would be acceptable and workable in both countries. The solution they finally arrived at
involved a treaty between the two countries. It worked simply because in both the British North America Act
and the United States Constitution there is provision for the implementation of the terms of an international
treaty as a federal responsibility.

After several years of preparatory work the Migratory Birds Convention (Treaty) between Canada, repre-
sented by Great Britain and the United States, was signed in 1916. Following that, each country passed
legislation implementing the treaty and began to promulgate regulations for the control of the migratory birds
referred to in the treaty. The signing of the treaty, and of the act based upon it, put the federal government
squarely into the wildlife management business for the first time.

There was created, by Order-in-Council on 28 December 1916, an Advisory Board on Wildlife Protection.
That organization had been recommended by J. B. Harkin, the Commissioner of Dominion Parks, and had as
its members James White, D. C. Hewitt, R. M. Anderson, as well as J. B. Harkin. One of the first jobs that
members of the Board did was to draft the Migratory Birds Convention Act and suggest the appointment of an
officer to administer the act and regulations. They also drafted a Northwest Game Act and concerned
themselves with Caribou in the Northwest Territories, Pronghorn, Elk, wolves, and other wildlife, as well as
special habitats such as Point Pelee and Bonaventure Island and a number of areas of importance for birds in
western Canada.

Because of the passage of the Migratory Birds Convention Act, there was a need for an official to be
appointed to carry out the terms of the act. By 1918 there was an active and growing National Parks
organization in the Department of the Interior. It seemed logical that an agency dealing with wildlife should be
located within the National Parks organization. Accordingly the official charged with responsibility for the
Migratory Birds Convention Act, Hoyes Lloyd, reported for duty to that agency on 11 December 1918. His job
was called Ornithologist and he started at a yearly salary of $2200. His basic responsibility was administration of
the Migratory Birds Convention Act, but he had an added responsibility involving the Northwest Game Act as
well. In 1919 the protection of wildlife in national parks was added to his duties and his title was changed to
Supervisor of Wildlife Protection. He had a secretary and, as you can imagine, he and the secretary were kept
pretty busy trying to cope with migratory birds and their problems throughout Canada, game in the Northwest
Territories, and wildlife in national parks. It was not long before he was able to convince his superior, J. B.
Harkin of the National Parks Branch, that he needed some help to carry out a job that covered sucha large area
and involved so many wild creatures of a wide range of species.

His pleas for assistance in carrying out his duties as Superintendent of Wildlife Protection resulted in the
creation of additional positions in the summer of 1920. Competitions were held for the positions of Chief
Federal Migratory Bird Officer for Ontario and Quebec, and for the western provinces. J. A. Munro of British
Columbia was the successful applicant for the western position and Harrison F. Lewis for the position involving
Ontario and Quebec. Prior to the appointments for migratory bird officers for western Canada and for Ontario
and Quebec, R. W. Tuffs of Wolfville, Nova Scotia, was appointed on 19 November 1919 as officer for the
maritime provinces.

As can be seen from the record thus far, although the Commission on Conservation had functioned since 1911
and an advisory board on Wildlife Protection since late 1916, work by qualified professionals in wildlife

1981 SOLMAN: FEDERAL WILDLIFE CONSERVATION IN CANADA 33

conservation did not begin in the federal government until 1920 and only then because of the responsibilities
under the Migratory Birds Convention. As the years passed, there were further subdivisions of Canada in terms
of staffing in the federal service, and eventually J. A. Munro restricted his activities to British Columbia and J.
Dewey Soper came on strength to deal with migratory bird problems in the prairie provinces.

By Order-in-Council on 12 April 1921, H. Lloyd was appointed Secretary to the Advisory Board on Wildlife
Protection to ensure that his work as Superintendent of Wildlife Protection would be fully coordinated with
that of the Advisory Board. The Advisory Board continued until the late 1940s and always had as members or
secretary, members of the organization which began with the appointment of H. Lloyd.

As the little nucleus of wildlife officials expanded within the National Parks organization, their assistance was
frequently requested on behalf of wildlife in the national parks and in the Northwest and Yukon Territories.
Although the basic responsibility was and remains migratory birds, the other responsibilities increased with
time. By the late 1930s there was added staff with expertise in matters involving mammals and freshwater
biology including fisheries, with particular reference to the fishery resources of the national parks. After 1945
the mammal and freshwater activities were expanded. By 1947 there were three mammalogists, a limnologist,
the four chief federal migratory bird officers, and the Superintendent of Wildlife Protection with supporting
staff. At that point the Director of the Lands, Parks and Forests Branch of the Department of Mines and
Resources deemed that a separate federal wildlife agency should be created. The Dominion Wildlife Service
came into being in November 1947. It became the Canadian Wildlife Service in April 1950.

Starting with the Commission of Conservation which involved the provincial governments as well as federal
government officials, there was a tradition of increasing federal and provincial cooperation in the wildlife field.
With termination of the work of the Commission of Conservation and the need for close cooperation with
regard to migratory birds conservation, a series of conferences was begun in 1922 at which federal and
provincial officials came together. They discussed migratory birds and the differing approaches to the use,
conservation, and control of those resources by the two levels of government. They also dealt with other matters
of interest to both federal and provincial wildlife officials. The Federal-Provincial Wildlife Conferences which
began in 1922 were held initially at irregular intervals but eventually became annual fora for the exchange of
ideas throughout the wildlife field; the one held in July 1978 was the 42nd in the series. Over the years those
conferences have been extremely helpful in sorting out the differing approaches to wildlife conservation when
perceived from the federal and from the provincial points of view. As a case in point, which has been debated
frequently, the federal government, under the Migratoryework of the Migratory Birds Convention Act and
Regulations, is responsible for safegarding the migratory bird resources by whatever means are appropriate.
Provincial governments on the other hand look upon migratory birds as resources which pass through their
jurisdictions annually, which are harvested by their citizens, and from which, through the sale of licenses and
other means, they secure revenue for operating at least a part of their wildlife management activities. Because of
the differences in approach there are annual discussions about what harvests of the harvestable resources should
be allowed and under what conditions. Because federal and provincial officials cooperate in annual studies of
the resources, there is usually no argument about the data on populations and distribution which are the key to
decisions about allowable harvest and where, when, and by what means it should be conducted. There are
complicating factors, including those related to the damage which migratory birds may do to agricultural crops
largely as a result of the kinds of technology used in harvesting those crops. To put it bluntly the grain farmers
would like to see fewer ducks. The duck hunters would like to see more, and the provincial and federal
governments are in the middle of the perennial debate about how many there should be and where they should
occur. Enforcement of provincial and federal regulations regarding the harvest of waterfowl by hunters is
carried on jointly by the Royal Canadian Mounted Police and the provincial conservation officers.

The major responsibility of the forerunner of the Canadian Wildlife Service was migratory birds and the
service still has that major responsibility. Over the years through cooperation with provincial governments and
private industry, the Canadian Wildlife Service has become involved ina variety of other projects ranging from
the controlled reproduction of raptors to prevent extinction of such birds as Peregrine Falcons, through studies
of animal diseases which are sometimes very important both to wildlife and to humans (for example rabies), and
in recent years increasing involvement with toxic substances in the environment. Those substances, in all parts
of Canada, may have detrimental effects on migratory birds and in many cases on fish and other wildlife and on
people.

Cooperation has always been good between the federal Canadian Wildlife Service and its provincial
counterparts and with other agencies in wildlife work including those in the private sector of the economy.
Cooperative studies of joint importance have gone on for more than 30 years. To bring a measure of legislative

34 THE CANADIAN FIELD-NATURALIST Vol. 95

authority to such arrangements Parliament passed the Canada Wildlife Act in July 1973. This made it easier to
justify to the Treasury Board expenditures for cooperative work with provinces which in any case had been
going on for a very long time. It has one rather unique feature in that it defines wildlife as any non-domestic
animal. With that definition, it can deal with any kind of wildlife in the country. The new act has not yet resulted
in research on the well-being of frogs or snails or some of our smaller invertebrate friends, but it looks as if there
is the authority to do just that if there is ever need for it.

Federal activity in the wildlife field naturally brought the Canadian Wildlife Service into contact with the
original Canadians. Many of the treaties that were signed between the Government of Canada and the Indian
tribes talk about the right of access to wildlife. When the Migratory Birds Treaty was signed it set limits on when
migratory birds could be taken. Applied to native populations in remote areas, those would have created
hardships and real difficulty. The application of those regulations had to be tempered with the reality of human
survival and so it was for many years. With the rise in concern over native rights in recent years, the native land
claims settlements in the James Bay area, and those under negotiation in the Mackenzie Valley and other parts
of northern Canada, it is probable that native use of migratory birds will come under a different sort of control.
It may be necessary to modify the Migratory Birds Treaty to spell out more clearly the application of the act
based on that treaty to native peoples throughout Canada. That matter is under study.

Public reaction to wildlife as shown by federal government action has changed from a complete lack of
concern for supposedly infinite stocks 100 years ago to great concern for wildlife and its habitat now. Canadians
currently examine the potential impact on wildlife and its habitat of almost any large construction project
proposed. The recent development of public awareness of the worth of wildlife resources now seems to assure
wildlife a continuing place in the Canadian way of life.

When the world human population reaches a sufficiently high level, decisions will have to be made between
using land for production of human food and for the production of wildlife. Then there will be very many
hungry persons who will not opt for the presevation of wildlife habitat. Nevertheless, until we in this fortunate
country reach that unhappy state of affairs, there will be ample land not required for food production. Wildlife if
properly managed can be produced and can continue to offer to Canadian citizens and their guests all of the
rewards which we normally associate with being able to observe, photograph, and harvest wildlife resources.

We have had an almost complete reversal of public attitude to wildlife during the life of The Ottawa
Field-Naturalists’ Club. It is interesting to realize that many people who were instrumental in creating the
agencies I have discussed were, at the same time, active and productive members of the club. In the early years,
only a small, dedicated minority was concerned with wildlife; now, governments at all levels are well aware that
tourism, based onan interest in wildlife and the out-of-doors, is one of the two or three largest earners of foreign
currency for Canada. Those who came together 100 years ago to form the club and to foster their interests in
wildlife and to encourage the public to become interested in wildlife have been well rewarded for their interest
and foresight.

Man evolved as a hunter. Hunters had to know a lot about wildlife to be successful. Although we no longer
must kill wildlife to survive, we are still hunters. As hunters and naturalists we find it rewarding to know where
to find wildlife, how to observe it, and how to pass along our knowledge and capacity for outdoor pleasure to
succeeding generations.

The Ottawa Field-Naturalists’ Club

Honorary Members

C.H. Douglas Clarke
William J. Cody
William G. Dore

R. Yorke Edwards
Clarence Frankton

W. Earl Godfrey
George H. McGee
Hugh M. Raup
Loris S. Russell
Douglass B.O. Savile

Pauline Snure

J. Dewey Soper
Charles M. Sternberg
Mary E. Stuart
Robie W. Tufts

A Canadian Paradox — Private Land, Public Wildlife:
Can it be Resolved?!

J. P. RYDER2 and D. A. BOAG:

2Department of Biology, Lakehead University, Thunder Bay, Ontario P7B 5E]1
3Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9

In Canada wildlife is the sole resource that remains legally the property of the Crown, even on lands that are
privately owned. Asa consequence, wildlife as a resource has traditionally been viewed as expendable whenever
it competed with any other resource that was the property of the landowner. In a recent discussion of this
problem in the United States, Davis (1979) concluded that “[in] the United States where approximately 2/3 of
the land area is owned by the private sector, wildlife is considered not as an economically competitive resource
but merely as a nonessential by-product” and that “[competing] uses for land have been and continue to be one
of the most important influences on the survival of America’s wildlife resources which include all life forms not
dependent upon man for survival.” .

It is not our function to attempt to resolve definitively the problems of just exactly what “wildlife” is. There
are numerous definitions that have historically included only those animals that are in one way or another
harvested by man both for economic and recreational return. Plants are not usually thought of as wildlife and,
until recently, neither were the terrestrial cold-blooded vertebrates and invertebrates (Clarke 1974). Because we
fee] that an abundance of wildlife has other than future consumer interests and benefits, our working definition
will be any undomesticated plant or animal.

Today, throughout much of the settled regions of the prairie provinces of Canada, those modern agricultural
practices, inimical to the survival of wildlife populations which historically inhabited those regions, are often
encouraged by governmental and agricultural agencies. Even on land held in the name of the Crown, wildlife
resources, for the most part, are not given priority or even parity with other resources that have immediate
commercial value, suchas forest production or open cattle grazing. Such disregard for wildlife has a long history
in Canada. Except for fisheries, the wildlife resource is not mentioned specifically in the British North America
Act of 1867. The resource automatically and without challenge became a provincial and territorial responsibil-
ity until a federal involvement was established in 1917 with the passing of the Migratory Bird Convention Act
and the National Wildlife Policy and Program in 1966. The former guaranteed protection to migratory game
birds as well as a large number of nongame species; the latter authorized the Canadian Wildlife Service to
acquire and manage wildlife habitats known as National Wildlife Areas, and to take responsibility for all types
of wildlife in national parks and in provinces and territories where the interests of wildlife species surpassed the
boundaries of such provinces and territories. All Canadian provinces and territories have their own wildlife
agencies and programs, the main thrust of their operations being to manage the wildlife resource, primarily but
not solely, from a consumptive viewpoint.

Thus, although we have created in this country, federal, provincial, and territorial agencies whose mandate it
is to conserve and manage wildlife, those actually responsible for administering the policies that deal with the
resource find themselves in a paradoxical situation. They must attempt to conserve and manage wildlife, yet
they have only very limited jurisdiction over the habitat upon which the resource is wholly dependent. If we are
to succeed in assuring a future abundance of wildlife, it is imperative that those charged with managing and
conserving the resource have more say in the formulation of land use policy both on privately owned lands and
those held in the name of the Crown.

To realize the goal of future wildlife resource abundance, we believe there must be some incentive for the
landowner to retain wildlife habitat. Perhaps the right of the private landowners to receive compensation or
even profit for providing or enhancing wildlife habitat should be recognized by all sectors of Canadian society.

'This statement was written as background foran ongoing discussion within the Wildlife Biologists’ Section of the Canadian
Society of Zoologists. It was prompted by a growing concern among biologists over the loss of wildlife habitat in the more
populated parts of Canada, expressed most effectively for the continent as a whole in A. J. Leopold’s recent book The
California Quail (1978. University of California Press, Berkeley).

39

36 THE CANADIAN FIELD-NATURALIST Vol. 95

Here we look at some possible ways and means of accomplishing, for the private landowner, some form of
financial return for retaining viable wildlife habitat in areas where economically important biological resources
other than wildlife currently form the livelihood for the landowner.

The lack of tradition and incentives for the establishment of wildlife areas on private land represents a very
serious historical omission in Canada. There exists no pride in owning, or encouragement to establish, wildlife
habitat. Within the prime agricultural areas of Alberta, Saskatchewan, and Manitoba, there are two main
classifications of agricultural crop lands: unimproved and improved. Improved land consists largely of
cropland given over to cereal grains. Unimproved land is, in the main, range pasture, 1.e., grazing lands with
natural vegetation, copses, potholes, and streams. The Dominion Bureau of Statistics (1968) and Statistics
Canada (1978) documented the areas of these lands in the three prairie provinces. Between 1961 and 1976 the
total area of improved land increased by 37 316 km? (13 434 mi2) through conversion of unimproved land on
old holdings as well as clearing land on new holdings.

On the prairie provinces it is the conversion of land from the unimproved to the improved status that, in many
ways, is the root cause of the existing paradox. Under the present regulations the Canadian Wheat Board
markets all wheat, oats, and barley delivered to them by the producers, but takes no responsibility in the area of
land-use policies. Each producer obtains a permit to deliver a given quota of the above crops, depending on the
area of improved land under his control. The quota system provides for all areas not cultivated in quota crops
(forexample, field peas, sunflowers, mustard, sugar beets and potatoes, summer fallow, nonnative pasture, and
new breaking) to be transferred to the actual crop area. Area in perennial forage, up to a maximum of one-third
the actual crop area, is also transferable to the quota area. The result is to increase the quota that can be
marketed. The upshot of this system is that it prevents a farmer from claiming unimproved land in establishing
his delivery quota. By simply tilling his unimproved land, the farmer can include it in his crop area entitlement
and thus potentially realize increased financial return.

The Canadian land assessment regulations automatically place a tax value on all unimproved land and
so-called wasteland. Section 30 of the Canadian Income Tax Act (1978) reads “. . . there may be deducted in
computing a taxpayer’s income fora taxation year from a farming business any amount paid by him in the year
for clearing land, levelling land or laying tile drainage for the purposes of carrying on the farming business.”
Section 18(2)e states in effect that no deduction shall be made in respect of land that cannot reasonably be
considered to have been in that year “held primarily for the purpose of gaining or producing income of the
taxpayer from the land for that year.”

The consequences of the Wheat Board and Income Tax regulations are that unimproved land cannot
contribute to raising a farmer’s crop quota nor can it be claimed for income tax purposes; in sum, it potentially
costs the farmer money to leave as is. Sawatzky (1979) noted that this system of agricultural regulations creates
among most farmers the opinion that they are paying taxes on land, the natural products of whichare of little if
any value to them. The result is that the landowner has no incentive to retain his unimproved land in its native
state and, in fact, the incentive may be to improve his “wasteland” to realize some return on the taxes he pays out
annually on it.

There are several ways in which farmers expend energy and money to improve their wasteland. All are
detrimental to wildlife and include the following: intensive cultivation of lands that are marginal for agricultural
production; summer fallowing which reduces soil fertility and tilth, increases soil vulnerability to wind and
water erosion, and leaves a large area of our arable land ina state that is virtually uninhabitable by wildlife;
draining wetlands which removes wildlife habitat and seriously influences groundwater systems and runoff
patterns (Sawatzky 1979).

In most areas the above practices render no benefit to the surrounding lands per se (Miller 1979a, Sawatzky
1979), and in fact can cause deleterious effects particularly related to downhill erosion problems, changed soil
chemistry rendering soils potentially sterile, and lowered water-holding capacity rendering low-lying land
subject to flooding. Such practices are carried out mostly on lands of less-productive soil types with a poorer
potential crop return. It appears that farmers who consider the costs, in terms of energy, time, and money, are
more than compensated for by the current income tax laws and grain quota systems. Sawatzky (1979) detailed
the needless practice of land “improvement” from a land management standpoint. Miller (1979b) noted that
only half of United States landowners contacted have participated in voluntary conservation programs and of
those only a few have adopted and retained all of the recommended soil conservation measures. The basic
reason fora general lack of cooperation is clearly stated by Elderkin (1979); “It is not that farmers and ranchers
are against wildlife, on the contrary the vast majority enjoy wildlife and are avid hunters. But they are in the
business to make money and land not producing a cash crop is losing them money.”

1981 RYDER AND BOAG: VIEWPOINT 37

What then can be done to alleviate these problems? A modification of the regulations of the Canadian Wheat
Board Act to permit transfer of unimproved land into quota entitlement would seem an obvious first step. The
actual amount transferred should be tied to its utility for wildlife populations and this will depend on its extent
and its quality. The farmer would then be able to claim such an area in his quota and at the same time he would
not be required to expend energy, time, or money to maintain the land.

Second, the Canadian Income Tax Act (Section 30) should be modified to allow deduction for land that is
significantly improved for wildlife and no taxes levied or deductions allowed for land that is left untouched or on
which no significant improvements for wildlife have been made. Deductions are envisioned as all costs for
improving the habitat in any given tax year. Additionally, we propose that Section 18(2)e of the Tax Act be
changed to allow freedom from tax any land not held primarily for the purpose of gaining or producing income
from a crop. We suggest that if direct income is obtained from the unimproved lands (see below), that such
income be taxable and not based on area or owned property. It is conceivable that if no funds are received from
unimproved land that no taxes should be levied, as long as the land has received some form of wildlife capability
rating, perhaps along the lines of the Agricultural and Rural Development Act (ARDA) of 1961. Taxes
currently levied on unimproved land, which would be lost to the municipality under the suggested scheme, could
be recouped through an increased level of taxation on the cultivated portion of property where the bulk of
income is attained in any one tax year. Arnolds and Woods (1979) discussed a unique program of direct
financial and tax relief for landowners in Colorado which involved selling or giving a conservation easement toa
portion of land whereby the owner may continue his farming or ranching operations and at the same time
receive either cash for the rights that he sells or an income tax deduction for the value of the rights given as a
qualified charitable donation. A state or federal agency or conservation group may acquire the recreational
right to use the land for a cash price much lower than it would cost to buy outright. In Colorado annual taxation
on the land subject to a conservation easement is assessed with regard to the restrictions on the use of the
property caused by the easement. Thus the owner who sells a conservation easement could have his land
revalued down and therefore pay lower property taxes. In effect the landowner receives two forms of financial
compensation: one from the sale of some rights to his land and the second from lowered property tax. The
people of Colorado have benefited significantly because at a moderate cost to the state they have the guarantee
that the beauty of and accessibility to land will be preserved for the future. This system might work well in
Canada.

Third, the establishment of a recognized legal fee schedule for both the nonconsumptive and consumptive use
of wildlife should be considered. Most provinces neither authorize nor prohibit landowners from charging
trespass fees for access to wildlife. Only Saskatchewan and Alberta specifically prohibit such payments under
their Game Act and Wildlife Act, respectively (Thompson and Lucas 1973). Such fees should be permitted by
law and payable in one form oranother to the landowner for specified uses of habitat untouched, maintained, or
improved for wildlife, in any given tax year. The fee structure established should reflect the value of the land for
wildlife to encourage ultimately the landowner to develop attractive wildlife areas. However, if the landowners
decide not to touch their lands and these lands have wildlife capability, there shall be no additional tax burden.

This suggested fee system runs counter to the philosophy in Canada that wildlife resources are free
(Gottschalk 1977). However, Gottschalk (1977) noted that “[free] hunting is a bankrupt concept that prevents
any but an intangible return on lands devoted to wildlife habitat.” The landowner, unless he is totally motivated
by esthetic considerations, bears the burden of wildlife habitat ownership with little or no compensation.
Indeed, if wildlife habitat is attractive to large numbers of citizens, who frequently trespass on private land in
pursuit of wildlife, whether birdwatchers, naturalists, hunters, or hikers, the maintenance of that habitat
becomes a significant burden for the landowner.

Charges, especially for access of naturalists to wildlife sanctuaries and for hunting, are not novel. A number of
states have established programs and fee structures for access to wildlife resources. In New Jersey, for example,
the Green Acres Tax Exemption Program makes private land available for public recreation and nature
observation with real estate tax relief to the landowner. In Nebraska the legislature has approved a $7.50
Habitat Stamp that is paid when a hunting licence is purchased by hunters (Gottschalk 1977). Funds from the
stamp are used to improve privately owned land for wildlife. This fund does not assure hunter access; the
decision to permit hunting rests with the landowner. If the landowner elects to permit hunter access, the Game
and Parks Commission is authorized to pay him $2.50 per acre for this access. In California, specific fees have
been established for access to private land, and are payable to the landowner depending on the type of services
received or game hunted (Teague 1971). Although trespass fees, which mean direct reimbursement to the
landowner for the right of access to wildlife habitat, isa growing practice in most provinces, payments received

38 THE CANADIAN FIELD-NATURALIST Vol. 95

by the landowner are not established by legislation and recognized as a legal source of income. Appropriate
changes in the existing Acts could easily legalize these procedures. Fees paid for the nonconsumptive and
consumptive use of wildlife could easily be administered through municipal or district game offices. Specific
land permits would be purchased by each user, giving him access to the private land. Monies received by sale of
the permits could then go back to maintain or develop land for wildlife conservation. Additionally, registration
of land for access would be prerequisite for tax relief and crop quota. Such a system would not force landowners
to permit persons on their land. Those that choose to prohibit access would still register their land as a wildlife
area, for the Tax and Wheat Board benefits. Those individuals who wish to use the wildlife on their land would
be entitled to receive additional financial compensation for continued improvement and repair of the habitat
and possible restocking of the resource. Such a system would not penalize the landowner who has a fine habitat
and who wishes to retain it incomplete privacy. The system would cost little in energy and financial output to the
individual and would positively encourage the maintenance of areas suitable for wildlife instead of destroying
them for limited financial gain under the present legislation.

The above proposals are meant to stimulate ideas on solving the Canadian Wildlife Paradox. They will have
to be discussed in depth to evolve mechanisms that are workable and satisfactory to almost all concerned,
especially the wildlife resource. To this end we solicit the readers’ reactions to these proposals in the hope that
their responses will contribute to solving this important conservation problem in Canada.

Acknowledgments

We thank those personnel from Agriculture Canada, the Canadian Wheat Board, and the provincial
governments, who gave us valuable ideas and suggestions. The following persons have made significant input
into this statement: R. N. Brown, A. J. Macaulay, G. J. Mitchell, H. L. Sawatzky, W. D. Wishart, and F. C.
Zwickel.

Literature Cited

Arnolds, D, and L. Woods. 1979. The conservation easement. Colorado Outdoors (Nov.—Dec.): 18-22.

Clarke, C. H. D. 1974. Conservation of wildlife in Canada. /n Conservation in Canada. Edited by J. S. Maini and A.
Carlisle. Department of the Environment, Canadian Forestry Publication 1340. Ottawa. pp. 191-224.

Davis, P. B. 1979. Pigeons to blackbirds: an illustration of the relationships between land use and wildlife resources.
Jack-Pine Warbler 57: 3-7.

Elderkin, R. L., Jr. 1979. A response to Harvey Miller’s dialogue. Wildlifer 176: 41.

Gottschalk, J. S. 1977. Wildlife habitat — the “price-less” resource base. Transactions of the 42nd North American Wildlife
and Natural Resources Conference. pp. 237-245.

Miller, H W. 1979a. Dialogue. Wildlifer 174: 15.

Miller, H. W. 1979b. Counter response to Robert Elderkin’s dialogue. Wildlifer 176: 41.

Sawatzky, H. L. 1979. Prairie potholes. Blue Jay 37: 3-8.

Teague, R. D. 1971. Wildlife enterprises on private land. Jn A manual of wildlife conservation. Edited by R. D. Teague.
Wildlife Society, Washington. pp. 140-143.

Thompson, A. R., and A. R. Lucas. 1973. Landowner-wildlife relationships: a preliminary legal study. Transactions of the
37th Federal—Provincial Wildlife Conference. pp. 36-50.

Dynamics of Moose Populations near Rochester, Alberta, 1975-1978

WILLIAM R. MYTTON and LLOYD B. KEITH
Department of Wildlife Ecology, University of Wisconsin, Madison, Wisconsin 53706

Mytton, William R., and Lloyd B. Keith. 1981. Dynamics of Moose populations near Rochester, Alberta, 1975-1978.
Canadian Field-Naturalist 95(1): 39-49.

A Moose population was studied near Rochester, Alberta, from October 1975 to April 1978. Mean densities of Moose on 228
km? of nonagricultural land were 0.64/ km? in winter 1975-1976. Fifty-nine Moose were radio-collared, and an additional 66
were color-collared. Sixty-one percent of the captured Moose (excluding calves) were yearlings and 2-yr-olds. The combined
yearling-and-adult sex ratio averaged 25 bulls to 75 cows during winter 1975-1977. Radio-collared Moose exhibited both
“migratory” and “nonmigratory” movement patterns. Distances between seasonal range centers of migratory individuals
were significantly greater for bulls, i.e., 13 vs. 7 km. There appeared to be no difference between seasonal home range sizes of
bulls and cows. Long-distance “dispersal” movements occurred among 1.5- to 2.5-yr-olds. Mean annual survival of radio-
collared yearlings and adults was estimated at 0.84. Calf survival estimated from disappearance rates of calves of radio-
collared cows, from demographic data, and from survival of radio-collared calves averaged 0.67 annually. Fecundity of
migratory cows was probably higher than of nonmigratory ones. Calves averaged 39% of the winter population in six aerial
surveys. Moose were located on aspen islands within muskegs during summer months and in open and treed muskeg during
September-November. Movements to tall high-density upland aspen and conifer stands occurred in December and continued
into March. During the last week of March and first week of April there was a marked shift into lowland areas. February and
March were the only months when Moose were not located further from human disturbances than would be expected ina
random distribution. The percentage of Moose having about one-third or more cleared land within a I-km radius of their
observed location increased from 17% in November to 48% in March.

Key Works: population dynamics, Moose, habitat, movements, radio-tracking, Alberta.

In October 1975, as part of the Alberta Oil Sands determined from radio-tracking.
Environmental Research Program (AOSERP), we
intensified our studies of Moose (Alces alces) near Study Area
Rochester, Alberta. During the previous 10 winters The 287-km2 (112-mi2) Rochester Study Area was
we had routinely counted, classified (by sex and age), located 100 km north of Edmonton (Figure 1). About
and plotted locations of Moose on 179 km? while 21% was cleared for agriculture; 43% was upland
conducting aerial surveys of Coyotes (Canis latrans) forest dominated by Trembling Aspen (Populus tre-
and White-tailed Deer (Odocoileus virginianus). muloides) and Balsam Poplar (P. balsamifera) with
These surveys indicated that the Moose population intermixed stands of Jack Pine (Pinus banksiana) and
was increasing rapidly, and that both reproductive White Spruce (Picea glauca); 32% was bog containing
and recruitment rates were high (Rolley and Keith scattered Tamarack (Larix Jaricina) and dense stands
1980). of Black Spruce (Picea mariana); and 3% consisted of

Because the Moose population at Rochester was _ ponds, lakes, and rivers. The western and southern
unhunted, essentially without predation, and onwhat sections of the study area bordered economically
- appeared to be excellent range, it provided a fine marginal farmland. Climate, soils, topography, and
opportunity to measure rates of increase and other vegetation of the area have been summarized else-
demographic data under what were optimal condi- where (Weatherill and Keith 1969; Rusch et al. 1971;
tions for Moose in northcentral Alberta. In other Keith and Windberg 1978).
words, the Rochester population could be used as a There were three “survey” areas within the Roches-
base against which to compare the dynamics and ter Study Area (Figure 1). The initial survey area of
growth potential of other regional populations. The 179 km?(hereafter called the Helicopter Survey Area)
Rochester population also offered opportunities to was established in 1965 for monitoring Coyote and
examine relationships between human disturbances White-tailed Deer populations, but Moose were also
and seasonal distribution and habitat usage by counted each winter. The second area of 228 km? (the
Moose. Moose Survey Area) was selected because Moose

The present paper is based on analysis of data numbers appeared high there, and it was bounded by
obtained between October 1975 and April 1978. It is | easily recognized road systems. The third area of 143
primarily concerned with population demography, km? (the Intensive Survey Area) was mainly forested
movements, distribution, and habitat utilization as and had the highest Moose densities.

a9

40

ALBERTA

STUDY
AREA
*e

EDMONTON

s
PERRYVALE

Fo G8 Gees:

Veqeaqebaqas ga qga geese ease:
OI IIL LL IL I 1

ROCHESTER
a

STONY CREEK

it

HELICOPTER SURVEY AREA1965-77 (179KM2) ~~

THE CANADIAN FIELD-NATURALIST

ROCHESTER MOOSE STUDY AREA

Se SCs gaar

Vol. 95

PINE CREEK

a

PIII IIIS! Oo erent, tnnneneinin

PHPTTF GFF FGFS FFFF8 88S Fe F880

MOOSE SURVEY AREA1975-77 (228KM2) am

INTENSIVE SURVEY AREA1976-77 (143KM2) ‘9

FIGURE 1. Location of Moose survey areas within the Rochester Study Area.

Methods and Materials
Capture, Marking, and Aging

During October 1975 — October 1977, 144 Moose
were darted from Bell 206 and Hughes 500c helicop-
ters, and immobilized with M99 (etorphine hydro-
chloride) or a mixture of fentanyl] citrate, Rompun,
and Wydase, as described by Haigh et al. (1977).
Ninety-eight (39 bulls, 59 cows) Moose were captured
during 29 October — 13 December 1975; 30 (15 bulls,
15 cows) during 22 October 1976 — 19 January 1977;
and 16 (3 bulls, 13 cows) were recaptured during 17
October — 21 October 1977.

Fifty-nine of the above Moose were fitted with
radio-collars; an additonal 66 were given color-coded
collars (21 cm wide) for individual identification dur-
ing aerial observations. All received numbered metal
ear tags bearing our address.

An outside incisor was pulled on each animal
(excluding calves) for age determination by tooth-
cementum annuli (Sergeant and Pimlott 1959; Gasa-
way et al. 1978).

Relocations

Radio-collared Moose were relocated about every
10d from a Cessna 185 equipped with two four-
element yagi antennas connected to a switch box
within the aircraft.We first determined approximate
position by equalizing signal strength from each
antenna, and then located the Moose either visually
(79%) or by maximizing signal strength (21%). During
summer 1976, locations were also obtained from the
ground every 3 d by triangulating radio signals from
an 18-m tower and two truck-mounted paired yagi
antennas.

All locations were recorded on 1000-m-grid maps.
Linear distances between successive locations were
determined; home range size was estimated using
Mohr’s (1947) minimum-perimeter-polygon method.

Seasons were delineated from mean dates of arrival
and departure of Moose on easily recognized seasonal
ranges: winter (22 December to 2 April) and summer
(12 May to 15 September).

During location flights we also recorded Moose

1981

activity and number of associates, whether contact
was visual, and habitat occupied when contacted. Our
habitat descriptions utilized a forest classification sys-
tem (Alberta Forest Service) that included species
composition, density, and overstory height.

Population Surveys

Weemployed three types of aerial surveys to obtain
demographic data on the Moose population: (1) east-
west transects in a Cessna 185 at 0.8-km intervals,
airspeed of 80-100 km/h, and altitude of 95-100 m
agl (above ground level); (2) east-west transects ina Bell
206 helicopter at 0.4-km intervals, airspeed of
85 km/h, and of altitude 75 m agl; and (3) intensive
random searching in a Bell 206 helicopter. Ratios of
marked Moose observed on survey flights to total
marked present were used in Peterson-index calcula-
tions to estimate numbers. Only radio-collared ani-
mals were considered as the marked cohort because
we knew the number present on any given survey date.

Disturbance

An index to three human disturbance factors was
obtained by measuring the nearest distance from each
Moose observation to occupied dwellings, petroleum
development, or graded roads. In addition, the extent
of agricultural clearing tolerated by Moose was inves-
tigated by determining the amount of cleared land
within | km of each Moose location. These observed
values were then compared with theoretical distances
to disturbance and percentage of clearing, as calcu-
lated from a random sample of 425 points on the
1000-m-grid map of the Rochester Study Area.

Results and Conclusions
Population Density

Rolley and Keith (1980) documented the growth of
the Moose population on 179 km? at Rochester from

MYTTON AND KEITH: MOOSE POPULATION DYNAMICS, ALBERTA 41

0.016/km2 of nonagricultural land in winter 1965 to
0.75/km2 by 1978.

We calculated a mean of 0.64 Moose/ km? of non-
agricultural land through four survey flights between
2 January and 20 February 1976 on the 228-km2
Moose Survey Area(Table I, Figure 1). This included
the northern half of the 179-km? Helicopter Survey
Area.

The only flights in winter 1976-1977 were on I 1-12
December, and covered the 143-km2 Intensive Survey
Area (Figure |). The observed marked-to-total ratio
yielded an estimated density of 1.4 Moose/km? of
nonagricultural land.

The above densities were three to five times higher
than those 300 km north near Fort McMurray during
1976-1978 (Hauge and Keith 1981). This difference is
likely related in part to the absence of large predators
and legal hunting at Rochester. Densities similar to or
higher than those at Rochester have been reported
from Newfoundland (1.1—-4.6/km?2, Bergerud and
Manuel 1969), Minnesota (0.4-0.8/km2, Peek et al.
1976), and Isle Royale (1.5-3.0/km2, Peterson 1977).

Sex and Age Structure

Sex ratios of combined yearlings and adults were
determined from six helicopter flights during Decem-
ber-February 1975-1976 and 1976-1977. The main
criterion for sexing was presence or absence of a vulva
patch (Mitchell 1970); antlers or antler scars were
sometimes a convenient alternative method.

We observed a mean bull:cow ratio of 27:73 (n = 83)
on surveys during December—February 1975-1976
(Table 2). The following winter a ratio of 23:79
(n = 117) was obtained. These ratios differed signifi-
cantly from the 47:53 (m = 389) noted on darting flights
during October-December 1975, and the 41:59 on
relocation flights during July-November when 163

TABLE !—Estimated numbers and density of Moose on the Rochester Study Area as determined from aerial surveys

described in text

Area Total no. Moose 95% Moose per km? of
and Aircraft of Moose population confidence nonagricultural
winter Date type observed estimate’ limits land
Moose Survey 2 January Fixed-wing 143 117-247 0.83(0.88)"
Area (228 km2) 19 January Helicopter 64 93 54-188 0.54(0.85)
1975-1976 7 February Fixed-wing 44 110 64-136 0.64(0.38)
20 February Helicopter 67 97 46-146 0.56(0.89)
Means 111 0.64(0.75)
Intensive Survey
Area (143 km?) 11-12 December Helicopter 119 135 75-218 (1.40)
1976-1977

*Peterson-index calculations: estimates based on ratio of radio-collared Moose seen to radio-collared Moose present.
b sae ‘ . .
Densities in parentheses refer to 143 km2 Intensive Survey Area (see Figure 1).

42 THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 2—Sex and age composition of the Moose population as determined from helicopter surveys at Rochester during

winter (December-February)

Winter Total Adult sex ratio .
and survey no. of bulls:cows % of population
date observations (%) Bulls Cows Calves Unknown
1975-1976
10 December 15 29:71 13 33 53 0
19 January 64 28:72 16 41 39 5
20 February 67 25:75 15 45 40 0
Total and
weighted means 146 27:73 15 42 41 2
1976-1977
11-12 December 119 Dye ys 18 49 34 0
16 December 33 15:85 9 52 39 0
1 February 33 17:83 9 45 45 0
Total and
weighted means 185 23:77 15 49 37 0

incidental observations were made. This difference

may indicate the following:

1) A differentially higher egress rate of bulls to adja-
cent game management units in late fall where they
suffer increased mortality through hunting.

2) A differentially lower rate of ingress of bulls due to
bull-only hunting in adjacent management units.

3) Sex-specific movement patterns which temporar-
ily shift more bulls than cows off survey areas
during winter.

The tendency for bulls to make longer seasonal
movements than cows, as discussed later, tends to
support (1) and (3) above. Further evidence of a
higher egress rate among bulls were the reobservation
rates of marked individuals as discussed by Rolley and
Keith (1980). Yearling and adult bulls were reob-
served significantly less often than other sex-and-age

classes, with yearlings evidently disappearing at the .

highest rate.

In ungulate populations, tertiary sex ratios tend to
favor females as a result of greater male mortality
allegedly due to greater activity in general (including
larger home-range size, and higher metabolic rates)
and that associated with reproduction in particular
(Flook 1970). Markgren (1971) reported that in
Sweden, Moose embryos and calves were 55-58%
males, whereas tertiary ratios usually favored females.

Bull:cow ratios in northeastern Alberta (26:74 in
1976, 33:67 in 1977) (Hauge and Keith 1981), where
both hunting and Gray Wolf (Canis /upus) predation
occur, were similar to those at Rochester (Table 2).

The sample of 64 Moose (excluding calves) that we
darted and immobilized in October-December 1975
was composed of individuals taken as encountered,
and should thus be representative of the population’s
age distribution. Thirty-three percent were yearlings,

28% were 2-yr-olds, and 39% were older animals. This
distribution was not significantly different from the
theoretical stable age distribution of 32:22:47 calcu-
lated from observed age-specific fecundity and survi-
val rates (Rolley and Keith 1980).

Aggregations and Associations

Radio relocations enabled us to examine the sea-
sonal aggregation of Moose and the attendant associ-
ation of different sex and age-classes. We used Ber-
gerud and Manuel’s (1969) definition of an
aggregation as any group of animals occurring within
close proximity of one another.

Mean size of bull-dominated groups (more bulls in
group than cows) varied from 1.0 (n = 65) in early
August to 2.3 (nm = 105) in November. During Decem-
ber-July, bulls were usually solitary. Largest bull-
only aggregations occurred immediately after the rut
(mid-October to late-November) when up to nine
were seen together. There were no age-specific differ-
ences in size of bull-dominated groups, although it
appeared that younger bulls (1.5-3.5 yr) remained
with cows longer than did older bulls (August to Jan-
uary vs. September to November).

Cows with calves tended to be solitary throughout
the year except fora brief period in September during
the rut (Figure 2). At that time 57% of cows with
calves associated with other adults or with yearlings;
all of such associates were bulls. Similar groupings
have been noted elsewhere (Geist 1963; Peek et al.
1974; Rounds 1978; Hauge and Keith 1981). Rounds
(1978) found that during fall single cows associated
with other adults up to 2 mo longer than did cows with
calves. Our observations at Rochester suggest that an
even longer period of association (August- December)
is common (Figure 2).

198]

oD
So

a
{e)

% ASSOCIATION WITH
YEARLINGS AND ADULTS
nm w b
oO (e) Oo

—
oO

MYTTON AND KEITH: MOOSE POPULATION DYNAMICS, ALBERTA 43

COWS WITHOUT CALVES
(N - 277) r

x
Aceresrescagy,

COWS WITH CALVES nN
(N- 248)
4

JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR APR MAY

FIGURE 2. Percentage of radio-collared cow Moose, with and without calves, associated with other yearlings and adults at

Rochester during October 1975 — April 1978.

Seasonal Movements and Home Ranges

We analyzed four types of Moose movements at
Rochester: (1) short-term movements (3-14 d)
between successive relocations; (2) seasonal “migra-
tions” between lowlands to the east and uplands along
the Tawatinaw River to the west; (3) seasonal shifts
between lowlands and immediately adjacent uplands
in the eastern portion of the study area; and (4) disper-
sal from the study area.

Movements between successive relocations of year-
lings and adults were usually < 2 km, with 87% being
< 5 km. Bulls made significantly more moves > 5 km
than did cows. This was owing to the greater distances
(means 13.4 vs. 4.9 km) between seasonal ranges
(Table 3) and longer movements during the rut. Phil-
lips et al. (1973) and Best et al. (1978) noted similar
seasonal differences in length of daily or short-term
movements.

As indicated in (2) and (3) above, our radio-collared
Moose at Rochester exhibited two types of seasonal
movements. The first involved five cows that made
annual moves between their winter ranges along the

Tawatinaw River uplands and summer ranges in the
muskeg and lake region to the east. Three characteris-
tics set this “migratory” cohort apart from the four
“nonmigratory” cows that remained in the muskeg
and lake region throughout the year: (1) individuals
moving to the Tawatinaw River area were traditional
in their use of wintering areas during successive win-
ters, i1.e., all were found on the same winter ranges
during three successive years; (2) routes of travel
between seasonal ranges were approximately the same
each year; and (3) four of the five cows had no overlap
between winter and summer ranges, and the other had
only an 8% overlap (Figure 3). The four “non-
migratory” radio-collared cows had winter ranges and
travel routes that differed each year, and a mean over-
lap between winter and summer ranges of 76%.
Three radio-collared adult (2 2.5 yr) bulls made
seasonal movements similar to those of migratory
cows. Two moved southwest to farmland areas, and
one wintered along the Tawatinaw River about 5 km
northwest of Perryvale (Figure 3). In each case, sea-
sonal ranges were separated by at least 6 km, and two

TABLE 3—Mean distances in kilometres between the centers of summer and winter home ranges of radio-collared adult

(= 2.5 yr) Moose at Rochester during 1976 and 1977

Migratory cows

Nonmigratory cows

1966 3

Bulls
No. of pairs Mean distance
Year of ranges (+ SE)
10.0 + 2.5
1977 2 18.5 + 6.7
13.4 + 3.6

No. of pairs Meandistance No. of pairs Mean distance
of ranges (= SE) of ranges (+ SE)
5 6.6 + 0.9 4 2.6 + 0.5
5 7.0 + 1.0 4 2.5+ 0.4
10 6.8 + 0.7 8 2.6 + 0.3

Totals and weighted means 5

44 THE CANADIAN FIELD-NATURALIST

WINTER

@e coo

(Wiese ei eae een

s 7, a CaS
hes jewance— GR aT
/ H

”
a
Pa

NONMIGRATORY

FIGURE 3. Movements between seasonal range centers for
“migratory” and ‘“nonmigratory” radio-collared
adult (2 2.5 yr) Moose at Rochester during
1975-1977. Centers of ranges that were < 0.5 km
apart were combined as one point.

of the three winter ranges were situated off the study
area.

Four calves (one male, three female), radio-collared
concurrently with their mothers along the Tawatinaw
River, returned there as yearlings in the following
winter. These apparently learned movements were not
exhibited by four other calves (three males, one
female) that were permanently separated from their
mothers during tagging operations on summer and
fall ranges. The latter calves were nonmigratory, win-
tering on the summer ranges of their mothers. Two
female calves separated after they had moved to the
Tawatinaw River area returned there the following
winter.

Vol. 95

One radio-collared bull and three cows moved
>50 km from the study area. All four were 1.5 or
2.5 yr old. Such dispersal occurred in late summer and
early fall and was to the north and northwest. One
additional bull, color-collared in 1975 as a yearling,
moved 250 km west of Rochester during a 2-yr period.

In fall 1976, four yearlings (one bull, three cows)
that as calves had migrated to the Tawatinaw River
area with their mothers moved 10-50 km north and
west of the study area. By late winter (mid-January to
February), however, all four had returned to the areas
on which they had previously wintered as calves. As
2-yr-olds in fall 1977, these same individuals moved
directly to their previous winter ranges.

Mean dates of arrival on, and departure from, dis-
tinct seasonal ranges were used to delineate the follow-
ing winter and summer periods:

1) Later winter to spring (22 December to 2 April):
the period when Moose were on winter range.

2) Spring (3 April to 11 May): the period when Moose
were between winter and summer ranges.

3) Summer (12 May to 15 September): the period
when Moose were on summer range.

4) Fall and early winter (16 September to 21
December): the period of rut and movement
between summer and winter ranges.

As Tester and Siniff (1973) pointed out in assessing
the measurement of home range, size generally
increases with the number of observations and/ or the
length of the observation period. At Rochester there
was a direct relationship (r?= 0.71) between the
number of Moose locations and the resulting estimate
of home range size. We therefore compared seasonal
ranges only where numbers of seasonal locations were
similar. Chi-square tests indicated no statistically sig-
nificant differences in home range size of bulls versus
that of cows, but innate variability was high (1 km?to
54 km2); nor was there a difference in size of seasonal
ranges of yearlings and 2-yr-olds versus those of older
individuals. Mean winter and summer home ranges of
15 km? and 16 km? at Rochester, based on 7-12 loca-
tions per animal (Table 4), were larger than those
reported in other areas, from a similar number of
locations (Houston 1968; Goddard 1970; Phillips et
al. 1973), but were smaller than ranges determined by
radio-tracking in northeastern Alberta (22-28 km? in
winter and 37 km2insummer; Hauge and Keith 1981).

Survival Rates
Adult

Survival rates of radio-collared yearlings and adults
were calculated using the method of Trent and Rong-
stad (1974). We used two approaches to calculation
problems imposed by malfunctioning or lost collars:
(1) only individuals whose fates were known to us
were included in the first (maximum) estimate of sur-

1981

TABLE 4—Mean home range size of radio-collared Moose
during winter (December-March) and summer (May-15
September) on the Rochester Study Area as calculated from
minimum perimeter polygons (Mohr 1947). Smallest and
largest home range sizes shown in parentheses

Winter Summer
No. of Mean No. of Mean No. of
locations (km?) Moose (km2) Moose
3-6 3(1-18)° 21 4( 1-16) i
W=12 15(2-54) 45 16( 1-34)’ 16
13-19 25(11-39) 10
20-23 26(12-44) 9

“Home ranges were calculated for these two cells using ran-
domly selected subsets from the 7 to 12 locations per Moose
on winter ranges and the 13 to 23 locations on summer
ranges.

vival; (2) the second (minimum) estimate assumed
that all radio-collared individuals with whom we had
lost contact died, even though radio malfunction was
a possibility.

Four of the 32 yearlings and adults radio-collared
on the Rochester Study Area were known to have
died; one bull and one cow were legally shot in an
adjacent game-management unit, one cow was
poached, and another was struck by a truck. Four
color-collared animals (three bulls, one cow) were
poached on the east and northeast edge of the study
area during September-November 1976 and 1977,
and another was shot by atreaty Indian 250 km to the
west in January 1978.

The annual survival rates of radio-collared year-
lings and adults in 1976 and 1977 were 0.89 and 0.92
(assumption (1) above). Annual survival as calculated
from assumption (2) was 0.76 and 0.79. The mean
annual survival rate from these four different, but not
independent, estimates was 0.84. This is probably our

MYTTON AND KEITH: MOOSE POPULATION DYNAMICS, ALBERTA 45

best estimate of survival; small sample sizes precluded
any meaningful test for age- or sex-specific
differences.

Calf

Information on calf survival was obtained from
three sources; (1) 20 cows were radio-collared and the
presence or absence of their calves noted at each relo-
cation during mid-May to April; (2) winter calf:cow
ratios, coW survival rates, population age structure,
and age-specific fecundity rates were used to calculate
May to January survival; (3) 27 calves were radio-
collared during November—February and relocated
thereafter.

Calves that disappeared from their mothers’ sides
during mid-May to | December were classified as
mortalities because lone calves were not seen during
fall and early winter relocation flights. On the other
hand, disappearance of calves of radio-collared cows
after 1 December could not be entirely equated with
death: we know, for example, that 3 of 12 calves
radio-collared during 24 November - 11 December
had become separated from their cows by mid-
January, but each survived the winter. Survival
among 23 calves of radio-collared cows during the
approximate 6.5-mo period from birthto | December
was 0.73.

Mid-May to mid-January survival rate was esti-
mated demographically from (a) the mean calf:cow
ratio of 85:100(n = 279) observed on six winter survey
flights; (b) age-specific rates of fecundity of 0.57 and
1.67 calves for radio-collared yearling and adult cows;
(c) a cow survival rate of 0.89 for 8 mo (from annual
survival of 0.84 given earlier); and (d) a yearling:adult
cow ratio of 33:77 (given earlier). The calculated calf
survival rate was 0.58 over 8 mo (Table 5).

The foregoing estimates of calf survival are equival-
ent to 7-mo rates of 0.71 and 0.62, respectively. Their
mean of 0.67 is probably our best estimate of survival

TABLE 5—Calculation of calf survival during mid-May to mid-January utilizing the mean calf:cow ratio in winter, age-
specific fecundity, a yearling:adult cow ratio, and adult survival as given in text

Mean winter calf:cow ratio from six aerial surveys (mean, mid-January) X 85 calves:100 cows

Calculated calf:cow ratio at birth in mid-May:

3.33° (2-yr-olds) X 0.57 (calves:2-yr-old) = 0.19
0.67 (adults) 1.67 (calves:adult) = 1.12

1.31 calves:cow

Estimated calf:cow ratio in mid-May = 131 calves:100 cows

Adult survival estimate from mid-May to mid-January = 0.89

Apparent calf survival rate to mid-January = 85:131 = 0.65

Calf survival to mid-January, corrected for cow survival = 0.65 X 0.89 = 0.58

*Proportion of 2-yr-olds in the May population.

46

during the first 7 mo of life. It is also likely a good
estimate of total first-year survival, since all 15 calves
radio-collared between 24 November and 5 February,
and not permanently separated from cows during col-
laring, survived to April. In contrast, 4 of 12 calves
that became permanently separated during radio-
collaring died during February—April, thus support-
ing Markgren’s (1964) suggestion of increased mor-
tality among lone calves.

Calf Production

Calf:cow ratios and twinning rates among radio-
collared adults (23.0 yr) were obtained within the
first 10 d after birth in spring 1976. A calf:cow ratio of
167:100 (15:9) and a twinning rate of 88% (seven of
eight) were observed at that time. By fall and early
winter 1976 (September- December) the calf:cow ratio
was down to 122:100 (11:9), and the apparent twin-
ning rate was just 38% (three of eight).

In 1975 and 1977, calf:cow ratios during September-
December were 175:100 (14:8) and 144:100 (13:9) and
twinning rates were 75% (six of eight) and 44% (four
of nine).

Seven yearling cows radio-collared between
October 1975 and January 1977 as 2-yr-olds produced
four calves: two single calves and one set of twins. This
is a minimum estimate of yearling fecundity because
dates of first visual observation after possible calving
in mid-May were 20 May (2) and 9 July (2) for the four
calfless cows and 25 November, 8 December, and 9
December for the three with calves.

The fecundity of cows that migrated to the Tawa-
tinaw River area may well have been higher than those
that did not migrate. Mean calf:cow ratios in spring

THE CANADIAN FIELD-NATURALIST

Vol. 95

were 200:100 (20:10) vs. 125:100 (5:4), respectively,
and in fall 180:100 (27:15) vs. 100:100 (11:11); by fall
apparent twinning rates were 80 and 9%, respectively.

Survey flights during winter 1975-1976 and
1976-1977 provided additional information on twin-
ning rates and calf:cow ratios. Calves comprised 41%
of the population during December—February
1975-1976 and 37% in 1976-1977 (Table 2). This was
higher than elsewhere in Alberta: 22% in the Swan
Hills (G. Lynch, Alberta Fish and Wildlife Division,
unpublished data) and 20% near Fort McMurray
(Hauge and Keith 1981). The greater percentage of
calves at Rochester vs. Fort McMurray was also
reflected in a markedly higher calf:cow ratio during
December-February (85:100 vs. 51:100) and apparent
twinning rate (39 vs. 12%).

Rolley and Keith’s (1980) analysis of past survey
data from Rochester showed consistently high repro-
ductive rates over the period 1965-1977: winter calf:
cow ratios and twinning rates were higher at Roches-
ter than those observed in utero or immediately after
birth in most other areas of North America.

Habitat Utilization

Availability of cover types was determined through
a stratified random sampling of Alberta Forest Ser-
vice cover maps (1:16 000) of the study area. Observed
rates of cover usage were then compared to known
availability.

Movements of radio-collared Moose within and
between seasonal ranges were generally reflected by
changes in habitat utilization. Both aspen-dominated
uplands and muskeg-willow lowlands were used
extensively during June-September (Table 6), with

TABLE 6—Percentage of use of different cover types as indicated by location of radio-collared Moose on the Rochester Study

Area from October 1975 to April 1978°

Avail-

Cover type June July Aug. Sept. Oct
Aspen 42 51 56 46 37
Aspen — Jack Pine 2 2 |
Aspen - White Spruce ! I 3 l
Jack Pine I 2
Jack Pine - Aspen
White Spruce l l l
White Spruce —- Aspen I

Total upland usage 45 55 58 49 43
Aspen —- Balck Spruce ! l l
Black Spruce 2 l ! 2 l
Muskeg - willow 52 44 40 49 56
Water

Total lowland usage 55 46 42 51 58

Total no. of

observations 143. 203 147 88 137

“Field work suspended from 15 April to 1 July 1977.

Nov. Dec. Jan. Feb. Mar. April May Mean ability
AT) 5 70 70 ~=66 2734 50) 45
2 l 2 2 3 6 2 2
3 I l l 4 2 5
2 l I 3 I 2 l
l 2 2 I

I 5 5 2 I I

6 l 3 2

So) S19 75 80 84 32.44 56 57
l l l

I I I 3

46 43 26 20 15 66> 43 34
4

46 43 26 20 Io OS) 3S 44 42
KO Ce AN Sy TE 8 TO Cy NAS ie 4)

1981

bulls being located in lowlands significantly more
often (47%) than cows (28%). Aspen areas used during
the summer were usually islands and ridges of slightly
higher relief scattered throughout the muskeg and
lake region in the eastern portion of the study area.

During the rut (mid-September to mid-October)
both sexes used adjacent open muskegs more fre-
quently. From mid-October through November, post-
rut aggregations of bulls were commonly in muskegs
(54% of locations), whereas cows utilized aspen
islands (80% of locations). As bull aggregations grad-
ually disintegrated, lowland use decreased to less than
30% by mid-December.

Use of tall high-density aspen stands increased from
December into March (Table 6); such cover was fre-
quented more by bulls (67%) than by cows (56%)
during February-March. Utilization of uplands
peaked in March when over 84% of the radio-collared
Moose were relocated there. Rolley and Keith’s (1980)
1965-1979 surveys during December-March sug-
gested the same overwinter changes in habitat use.
Our comparison of usage and availability indices sug-
gested that Moose actively selected these high-density
aspen stands (Table 6).

A striking shift into open lowlands occurred during
the last week of March and first week of April. The
increased frequency of locations in lowlands (from 6
to 54%) during this 2-wk period coincided with the
loss of snow (reduced to trace amounts by 25 March in
1976 and 7 April in 1977). The 68% utilization of
lowland sites during the last week of April was greater
than at any other time of year. Berg and Phillips
(1974) and Hauge and Keith (1981) found a similar
situation in Minnesota and northeastern Alberta

MYTTON AND KEITH: MOOSE POPULATION DYNAMICS, ALBERTA 47

where movements to low open willow (Salix spp)
habitats occurred in early to mid-April. A synopsis of
the above-described seasonal differences in habitat
use by Moose is given in Table 7.

Moose Distribution
Disturbance
The distribution of radio-collared Moose in rela-
tion to human disturbance factors varied seasonally
(Figure 4). Movements from summer to winter ranges
brought Moose into closer proximity with sources of
disturbance, as most uplands on the Rochester Study
Area were utilized for roads, buildings, and farmland.
Early to late winter movements into remnant den-
sely forested uplands shifted both migratory and
nonmigratory Moose toward the farmland border,
with mean distances to disturbance decreasing from
1.1 km in November to 0.6 km in March (Figure 4).
February and March were the only months when
Moose were not located significantly farther from
disturbance than would be expected on a purely ran-
dom distribution basis. As distance to disturbance
decreased through winter, the amount of cleared land
within | km of each Moose observation increased.
Seventeen percent of Moose relocations in November,
and 48% in March, were in areas having greater than
one-third cleared land (Figure 4). But average utiliza-
tion of cleared land was, during all months of the year,
less than would be expected if distribution had been
random. Rolley and Keith (1980) reported similar
trends in distance to human disturbance in their anal-
ysis of November—March surveys conducted at
Rochester during 1965-1979.
There was great variation in the amount of clearin
that Moose tolerated. A 4.5-yr-old radio-collared

in Relation to Human

TABLE 7—Synopsis of seasonal distribution of Moose at Rochester in relation to major habitat characteristics. Data from

locations of both radio-collared and unmarked individuals

Total no. % of Moose
of in specified
Season observations habitat
Summer
(12 May - 15 September) 475 69
Fall
(16 September - 361 63
21 December)
Winter
(22 December - 2 April) 433 82
Spring
(3 April - 11 May) 290 53

“Forest density rating used by Alberta Forest Service.

Habitat characteristics

Overstory
Overstory height
Topography _ Vegetation density* (m)
Medium to
Uplands Deciduous high =>
Uplands Deciduous
and open islands Low <9
lowlands in muskeg
Deciduous Medium to
Uplands conifer high >9
Muskeg
Lowlands willow Low <9

48 THE CANADIAN FIELD-NATURALIST

oO
(fe)

80)

PERCENT OCCURRENCE
IN| LOWLANDS

OF MOOSE

IN| KM

TO HUMAN DISTURBANCE

DISTANCE

2 9 td Ie aia C
fe)
= =
rr e 0-30% CLEARED LAND
O7 WITHIN 1.0KM
WwW
Oo 6
z
©
a5
=)
3 4
30 : BSS oe.
5 31-50% CLEARED =
Wi 20 ;
x 75 1-100% CLEARED —~_
a epee Bie a

JUN JUL AUG SEP OCT NOV DEC JAN FEB MAR APR MAY
(143) (203) (147) (108) (137) (101) (69) (141) (152) (140) (172) (47)
FiGuRE 4. Monthly distribution of radio-collared Moose in
relation to indices to human disturbances: (A) utiliza-
tion of lowland areas; (B) distance to roads, buildings,
and petroleum development; (C) changes in percent-
age of cleared land found within | km of each Moose
observation at Rochester during October 1975 -
April 1978. Number of monthly observations shown
in parentheses.

bull, for example, was relocated for 3.5 mo during all
three winters (1975-1978) in areas of between 75 and
95% cleared land, while other individuals rarely ven-
tured into areas of more than 25% clearing. Such
inherent variability prevented generalizations about
tolerance of Moose to human disturbance. It seemed,
however, that most animals did not tolerate areas of
> 30% cleared land for extended periods.

Vol. 95

During spring, movements away from disturbance
occurred when snow depths decreased to trace
amounts (25 March — 7 April). These movements
took Moose from the aspen uplands to the more iso-
lated muskegs and other lowlands (Table 7) where
they calved. The mean distance to disturbance
increased through spring and summer (1.2 km in
April, 1.5 km in August), then decreased in mid-
September with the onset of the rut.

Acknowledgments

We are grateful to R. Frokjer, F. Gingerich, J.
Jorgenson, and P. Mytton for field and office assist-
ance; to J. Cary for computer and statistical advice;
and to T. Fuller, T. Hauge, R. Rolley, and the fixed-
wing and helicopter pilots without whose help this
project would not have been possible. We also thank
J. Kemp, G. Lynch, D. Neave, K. Thirwell, R.
Weatherill, and many other members of the Alberta
Department of Recreation, Parks and Wildlife (Fish
and Wildlife Division) for logistical support. This pro-
ject was funded by the Alberta Oil Sands Environ-
mental Research Program and Fish and Wildlife
Division.

Literature Cited

Berg, W.E., and R.L. Phillips. 1974. Habitat use by
Moose in northwestern Minnesota with reference to other
heavily willowed areas. Naturaliste Canadien 101:
101-116.

Bergerud, A. T., and F. Manuel. 1969. Aerial census of
Moose in central Newfoundland. Journal of Wildlife
Management 33(4): 910-916.

Best, D. A., G. M. Lynch, and O. J. Rongstad. 1978. Sea-
sonal activity patterns of Moose in the Swan Hills,
Alberta. Proceedings of the 14th Annual Moose Confer-
ence and Workshop, Halifax, Nova Scotia. 17 pp. In
press.

Flook, D. R. 1970. Causes and implications of an observed
sex differential in the survival of Wapiti. Canadian Wild-
life Service Report Series Number 1|1. 71 pp.

Gasaway, W.C., D. B. Harkness, and R.A. Rausch.
1978. Accuracy of Moose age determinations from incisor
cementum layers. Journal of Wildlife Management 42(3):
558-563.

Geist, V. 1963. On the behavior of the North American
Moose (Alces alces andersoni) in British Columbia.
Behaviour 20: 337-416.

Goddard, J. 1970. Movements of Moose in a _ heavily
hunted area of Ontario. Journal of Wildlife Management
34(2): 439-445.

Hauge, T. M., and L. B. Keith. 1981. Dynamics of Moose
populations in northeastern Alberta. Journal of Wildlife
Management 44. In press.

Haigh, J.C., R. R. Stewart, R. Frokjer, and T. Hauge.
1977. Capture of Moose with Fentanyl and Xylazine.
Journal of Zoo Animal Medicine 8(3): 22-29.

Houston, D. B. 1968. The Shiras Moose in Jackson Hole,

1981

Wyoming. Technical Bulletin of the Grand Teton Natural
History Association 1: 1-110.

Keith, L. B., and L. A. Windberg. 1978. A demographic
analysis of the Snowshoe Hare cycle. Wildlife Mono-
graphs Number 58. 70 pp. i

Markgren, G. 1964. Den ensamma kalven. Svensk Jakt
Number 11. pp. 566-569.

Markgren, G. 1971. The question of polygamy at an unbal-
anced sex ratio in the Moose. IUCN Publication, New

_ Series Number 24. International Union for Conservation
of Nature and Natural Resources, Morges, Switzerland.
pp. 756-758.

Mitchell, H.B. 1970. Rapid aerial sexing of antlerless
Moose in British Columbia. Journal of Wildlife Manage-
ment 34(3): 645-646.

Mohr, C. QO. 1947. Table of equivalent populations of
North American small mammals. American Midland
Naturalist 37: 223-249.

Peek, J. M., R. E. LeRusche, and D. R. Stevens. 1974.
Dynamics of Moose aggregations in Alaska, Minnesota,
and Montana. Journal of Mammalogy 55(1): 126-137.

Peek, J. M., D. L. Urich, and R. J. Mackie. 1976. Moose
habitat selection and relationships to forest management
in northeastern Minnesota. Wildlife Monographs
Number 48. 65 pp.

Peterson, R. O. 1977. Wolf ecology and prey relationships
on Isle Royale. National Park Service Scientific Mono-
graph Series, Number 11. 210 pp.

Phillips, R. L., W. E. Berg, and D. B. Siniff. 1973. Moose

MYTTON AND KEITH: MOOSE POPULATION DYNAMICS, ALBERTA 49

movement patterns and range use in northwestern Minne-
sota. Journal of Wildlife Management 37(3): 266-277.

Rolley, R. E., and L. B. Keith. 1980. Moose population
dynamics and winter habitat use at Rochester, Alberta.
Canadian Field-Naturalist 94(1): 9-18.

Rounds, R. R. 1978. Grouping characteristics of Moose
(Alces alces) in Riding Mountain National Park, Mani-
toba. Canadian Field-Naturalist 92(3): 223-227.

Rusch, D. H., L. B. Keith, and E. C. Meslow. 1971. Natu-
ral vegetative communities near Rochester, Alberta.
Alberta Department of Lands and Forests, Fish and Wild-
life Division Technical Bulletin Number 4. 22 pp.

Sergeant, D. E., and D. H. Pimlott. 1959. Age determina-
tion in Moose from sectioned incisor teeth. Journal of
Wildlife Management 23(3): 315-321.

Tester, J. R., and D. B. Siniff. 1973. Relevance of home
range concepts to game biology, Transactions of the 11th
International Congress of Game Biologists, Stockholm.
pp. 287-296.

Trent, T. T., and O. J. Rongstad. 1974. Home range and
survival of cottontail rabbits in southwestern Wisconsin.
Journal of Wildlife Management 38(3): 459-472.

Weatherill, R. G., and L. B. Keith. 1969. The effect of live-
stock grazing on an aspen forest community. Alberta
Department of Lands and Forests, Fish and Wildlife Di-
vision Technical Bulletin Number 1. 31 pp.

Received 24 September 1979
Accepted 23 May 1980

Régime alimentaire des Orignaux du sud-ouest Québécois
pour les Mois d’Avril a Octobre

MICHEL CRETE! et PETER A. JORDAN?

'Ministére du Loisir, de la Chasse et de la Péche, Service de l’aménagement et de I’exploitation de la faune, 13, rue Buteau,
Hull, Québec J&Z 1V4

Adresse actuelle: Ministére du Loisir, de la Chasse et de la Péche, Direction de la recherche faunique, 9530, rue de la Faune,
Orsainville, Québec GIG 5E5

2Department of Entomology, Fisheries and Wildlife, University of Minnesota, St. Paul, Minnesota 55108

Créte, Michel et Peter A. Jordan. 1981. Régime alimentaire des Orignaux du sud-ouest québécois pour les mois d’avril a
octobre. Canadian Field-Naturalist 95(1): 50-56.

On a analysé le contenu de 64 rumens d’Orignaux (Alces alces) récoltés entre 1977 et 1979 dans la réserve de La Vérendrye,
Québec. La période considérée a été divisée en quatre: printemps, début de l’été, fin de l’été et automne. Les ramilles
dominaient le régime alimentaire des Orignaux au printemps jusque vers le 15 maiet les feuilles le reste du temps; a ’automne,
les ramilles dominaient a nouveau dans les rumens vers la mi-octobre. Le Sapin Baumier (Abies balsamea), V If du Canada
(Taxus canadensis) et ’Erable a Epis (Acer spicatum) représentaient les espéces les plus communes dans les rumens au
printemps; durant tout !’été, l’Erable 4 Epis prédominait dans la nourriture des Orignaux alors qu’il cédait sa place aux
cerisiers (Prunus spp.) a l’'automne. Les petits nés au printemps pouvaient posséder une nourriture de méme nature que celle
des adultes dés la fin de juin. Une proportion importante des femelles a semble ne pas s’alimenter pendant la période durut en
1978. Ona conclu que I’Erable a Epis constitue la base du régime alimentaire annuel des Orignaux dans une bonne partie du
Québec.

Mots clés: Orignal, Québec, érable, régime alimentaire, Alces.

Créte, Michel et Peter A. Jordan. 1981. Régime alimentaire des Orignaux du sud-ouest québécois pour les mois d’avril
a octobre. Canadian Field-Naturalist 95(1): 50-56.

The study was based on the analysis of 64 samples of Moose (A/ces alces) rumen content collected in La Vérendrye Reserve,
Quebec, between 1977 and 1979. The interval between April and October was divided into four periods: spring, beginning of
the summer, end of the summer, and fall. Twigs dominated rumen contents in spring until approximately 15 May and leaves
thereafter; in fall, twigs dominated again by mid-October. Balsam Fir (Abies balsamea), Ground Hemlock (Taxus canaden-
sis), and Mountain Maple (Acer spicatum) composed the bulk of the browse in spring; Mountain Maple dominated Moose
diet in summer and was replaced by cherry (Prunus spp.) in autumn. Calves could havea diet similar to that of adults starting
at the end of June. Many females seemed to stop feeding during the rut in fall of 1978. Mountain Maple appears to represent
the yearlong basis of Moose nutrition in many areas of Quebec.

Key Words: Moose, Quebec, maple, food habits, Alces.

Le bilan énergétique des cervidés d’Amérique du _—_ qualité et la quantité de nourriture disponible en été.
Nord fait voir uncycle annuel quise caractérise parun Mais premiérement, fallait-il connaitre le régime ali-
gain de poids en été et une perte en hiver(Franzmann __mentaire de cette espéce animale. Le but de cet article
et al. 1978; Moen 1978). La fin du printemps et l’été est de décrire les habitudes alimentaires estivales de
constituent par conséquent une période importante _|’Orignal dans la région d’étude; aucune étude n’a
puisque les animaux refont leurs réserves. La qualité encore touché ce sujet en profondeur dans l’est du
et l’'abondance de la nourriture alors disponible pren- Canada. L’analyse du contenu de rumens d’Orignaux
nent encore plus d’importance pour les femelles qui _s’avérait une technique appropriée pour atteindre cet
doivent aussi affronter les exigences de la fin de la _ obyectif.
gestation et de la lactation (Gasaway et Coady 1974;

Verme 1977; Bahnak et al. 1979). Enfin, le régime Région d’étude

alimentaire précédant la période de reproduction La moitié sud de la réserve de La Vérendrye, sise a
influence l’Age a la puberté des femelles(Abler 1974)et environ 300 km au nord-ouest de Montréal, consti-
leur taux d’ovulation (Verme 1969). tuait la région d’étude. Cette réserve est traversée par

Dans le cadre d’une étude sur la dynamique des _ une route asphaltée importante qui relie le nord-ouest
populations d’Orignaux (Alces alces) dusud-ouestdu du Québec aux régions plus au sud. On y compte
Québec (Créte et al. 1981), il fallaitdoncconsidérerla environ 60 accidents routiers impliquant des Ori-

50

1981

gnaux et des véhicules automobiles par été. De plus,
ony tient une chasse contrélée de Orignal au cours de
laquelle 150-200 bétes sont abattues annuellement
(Bouchard et Moisan 1974). Les échantillons analysés
provinrent de ces deux sources.

La réserve de La Vérendrye posséde un relief ondulé
(altitude: 300-500 m), typique du Plateau Laurentien.
Des sols podzoliques s’y sont développés a partir
d’une assise rocheuse surtout granitique (Service de
Gestion de Environnement, 1977. Documentaire sur
le bassin de la riviére des Outaouais. Péche et Envi-
ronnement Canada). Les précipitations annuelles
atteignent en moyenne l’équivalent de 900 mm de
pluie, dont pres de 30% tombe sous forme de neige
(Ferland et Gagnon 1967). En moyenne, le débourre-
ment des bourgeons a lieu vers le 15 mai alors que la
défoliation débute vers le 15 septembre (Villeneuve
1969).

Des foréts mélangées contenant des feuillus tolérant
l’ombre (surtout le Bouleau Jaune, Betula allegha-
niensis et ’Erable a Sucre, Acer saccharum) occupent
souvent la partie supérieure des collines; le bas des
versants et les terrains plats sont surtout couverts de
foréts mélangées contenant des feuillus intolérants
(principalement le Bouleau a Papier, Betula papyrif-
era et le Peuplier Faux-Tremble, Populus tremu-
loides). Les feux de foréts qui étaient historiquement
plus fréquents dans la deuxiéme catégorie de peuple-
ments expliquent en grande partie la distribution des
deux communautés végéetales dominantes (Créte
1977). Les coniféres les plus importants, le Sapin
Baumier (Abies balsamea) et VEpinette Blanche
(Picea glauca) ont été affectés par la Tordeuse du
Bourgeon de l’Epinette (Choristoneura fumiferana)
de fagon parfois trés sévére. L’Erable a Epis (Acer
spicatum) constitue l’arbuste le plus abondant (Créte
1977) et le plus important pour l’Orignal en hiver
(Joyal 1976).

Méthode

Un échantillon d’environ | L du contenu des
rumens d’Orignaux était prélevé dans les 8 h suivant la
mort de l’animal; il était ensuite conservé dans une
solution de formol 10%. Une des deux premieres inci-

sives I, était aussi arrachée et conservée dans une.

solution de formol 3%.

Au laboratoire, chaque échantillon de contenu de
rumen était séparé en trois, selon la taille des frag-
ments, a l’aide de tamis munis de mailles de 6.3-4.0 et
1.7 mm (Bergerud et Russel 1964; McCaffery et al.
1974; Skinner et Telfer 1974). L’eau servait a entrainer
les particules d’un tamis a l’autre; les fragments qui
n’étaient pas retenus par les mailles les plus fines
n’étaient pas considérés. Lorsque l’échantillon était
dominé par des fragments de plantes non ligneux (le

CRETE ET JORDAN: NOURRITURE DE l’ORIGINAL Sil

cas le plus fréquent), on retirait d’abord, par espéce,
les ramilles des trois tamis. On déterminait ensuite
limportance relative des fragments non ligneux des
deux premiers tamis par échantillonnage; on étendait
une partie du contenu d’un tamis dans un bac peu
profond ot lon répartissait systématiquement 100
points-échantillon a laide d’aiguilles a pointe fine
(Chamrad et Box 1964). A chaque visée, on tentait
d’ identifier a ’espéce le fragment cible. Comme l’iden-
tification des fragments non ligneux était pratique-
ment impossible pour le tamis aux mailles de 1.7 mm,
on devait assumer que sa composition végétale était
similaire a celle des deux premiers tamis. On séchait
enfin les ramilles et le contenu des trois tamis a une
température de 70°C pendant 72h et lon pesait.
Lorsque l’échantillon était dominé par des ramilles, il
était tres laborieux de séparer les ramilles et les partic-
ules non ligneuses du troisiéme tamis; aussi la sépara-
tion n’était faite que pour un sous-échantillon et l’on
extrapolait a tout le tamis. De plus, lorsque les
ramilles dominaient, il était souvent plus rapide de
séparer par espéce les fragments non ligneux des deux
premiers tamis que de procéder par échantillonnage.

Lidentification des fragments de plante n’était pas
facile; pour s’aider, le technicien examinait d’abord les
feuilles et/ou ramilles encore entiéres dans l’échantil-
lon, consultait une collection de plantes séchées et
demandait l’avis de personnes plus familiéres que lui
avec la végétation de la région d’étude. De plus, ce
technicien avait développé une série de critéres d’iden-
tification pour plusieurs espéces communes.

L’age des Orignaux a été déterminé par le décompte
des annuli de cément entourant la racine de la pre-
mieére incisive I, (Sergeant et Pimlott 1959); le plan de
coupe était transversal. Cette technique n’était pas
nécessaire pour les petits de l'année que !l’on reconnait
a leur taille.

Enfin, ’équipe affectée a étude de la dynamique
des populations d’Orignaux de la région d’étude
notait, par espéce, la présence de feuilles d’arbres et
d’arbustes broutées par les Orignaux lors de ses
déplacements en pleine forét. Les observateurs
notaient simplement la présence d’espéces broutées
sans €valuation quantitative et deux observations
devaient étre distantes d’au moins 50 m.

Compte tenu des stades de développement de la
végétation et du nombre d’échantillons disponibles, la
période de temps couverte a été divisée en quatre:
printemps (avant le 15 mai), début de l’été (15 mai au
14 juillet), fin de l’été (15 juillet au 14 septembre) et
automne (aprés le 15 septembre). L’analyse statistique
s’est limitée a la comparaison de distributions de
fréquence par le test du chi-carré; les moyennes
présentées sont suivies de l’erreur standard de la
moyenne (sx).

Sy? THE CANADIAN FIELD-NATURALIST

PRINTEMPS
(I9 AVRIL-8 MAI)

Ramilles

n=6
l6%
FIN DE L'ETE
(18 JUIL.- 4 AOUT )
Ramilles
n=6
Inconnu
17 %
Feuilles
60%

Plantes graminoides
I5 %

Vol. 95

DEBUT DE LETE
(18 MAI-12 JUIL.)

Ramilles

AUTOMNE
(ISFSEPT 1 ShOCie)

Ramilles

I9 %
Plantes aquatiques
2%

59 %

Plantes graminoides
3%

FiGuRE 1. Composition en poids sec ducontenu des rumens d’Orignaux selon des groupes de plantes pour quatre périodes de
l'année. Les dates entre parenthéses indiquent le jour de la premiére et de la derniére collection. Les inconnus

n’incluent que des fragments non ligneux.

Résultats

Soixante-quatre échantillons de contenu de rumen
ont été récoltés entre avril 1977 et aout 1979; le tiers
des prélévements provint d’Orignaux tués sur la route,
le reste d’Orignaux récoltés a la chasse. Les jeunes de
l'année représentaient 8% de ces individus alors que
Yon comptait 42% de males parmi les adultes. L’age
moyen des adultes était de 4.2 ans + 0.5 (n = 44). Ces
statistiques suggeérent que |’échantillonnage était re-
présentatif de la population sur pied (Bouchard et
Moisan 1974; Créte et al. 1981).

Méme si l’identification des fragments végétaux des
rumens est difficile, seulement 22% + 2.5 (n = 50) et
27% + 2.7 des parties de végétaux ne purent étre iden-
tifiées dans le premier et le second tamis respective-
ment. En excluant les fragments non identifiés, le test
du chi-carré a révélé une composition végétale iden-

tique des deux premiers tamis pour 64% des 50 ێchan-
tillons contenant plus d’une espéce végétale (ou
groupe d’espéces).

Evolution du régime alimentaire au cours de la saison

La figure | fait voirla nature du régime alimentaire
des Orignaux pour les quatre périodes. Les végétaux
rencontrés dans les rumens d’Orignaux ont été
regroupés selon qu’ils étaient des parties d’arbres ou
d’arbustes (feuilles ou ramilles), des plantes grami-
noides ou des plantes aquatiques. Cette division
permet entre autres d’identifier le type d’habitat
fréquenté par les Orignaux puisque les arbres et
arbustes croissent en milieu terrestre alors que les
plantes aquatiques se retrouvent naturellement dans
les plans d’eau; sans en avoir l’assurance formelle, on
croit que les plantes graminolies (cypéracées, gram-

1981

inées, joncacées) rencontrées dans les rumens crois-
saient aussi dans ou prés des plans d’eau.

Au printemps, ce sont les ramilles qui dominent
largement le régime alimentaire des Orignaux de la
région d’étude; les aiguilles de coniféres constituent la
majorité des feuilles consommeées durant cette saison.
Au début de l’été, on note un changement radical alors
que les feuilles deviennent dominantes dans les
rumens; on note aussi que c’est le moment ou les
plantes aquatiques et les plantes graminoides abon-
dent le plus. Joyal et Scherrer (1978) ont d’ailleurs
montré que la fréquentation des plans d’eau par
’Orignal était la plus grande au début de l’été dans le
sud-ouest du Québec. La nature du régime alimentaire
ne change pas tellement a la fin de l’été: tout au plus,
les feuilles prennent-elles un peu plus d’importance. A
lautomne, les feuilles demeurent encore la compo-
sante principale de la nourriture de l’Orignal. Les
plantes graminoides deviennent passablement moins
fréquentes alors que l’inverse s’observe pour les
ramilles. Dans ce dernier cas, il est bon de noter qu’a-
vant le 10 octobre, les ramilles ne représentent que
12% du contenu des rumens alors qu’entre le 10 et le 13
octobre (dernier jour de la chasse dans la Réserve),
elles comptent pour 32% du poids des échantillons.
C’est donc vers la mi-octobre que les ramilles rede-
viennent majoritaires dans le régime alimentaire des
Orignaux; a ce moment, la défoliation est générale-
ment trés avancée (Villeneuve 1969).

CRETE ET JORDAN: NOURRITURE DE!’ ORIGINAL 53

La plupart des études touchant le régime alimen-
taire des Orignaux au cours de la période sans neige se
sont limitées 4 comparer limportance relative des
feuilles d’arbres et d’arbustes. Toutefois, Belovski et
al. (1973) ont estimé a environ 13% la proportion, en
poids sec, des plantes aquatiques dans la nourriture
estivale des Orignaux de I’Isle Royale, le reste étant
constitué de feuilles; cette estimation est du méme
ordre de grandeur que nos résultats si !on suppose que
la majorité des plantes graminoides rencontrées dans
les rumens croissaient en milieu aquatique. En
Alaska, l’observation, au cours des mois de juillet et
d’aoit, de trois Orignaux apprivoisés a permis d’esti-
mer que les feuilles représentaient environ 65% de la
nourriture de ces animaux, et que, comparativementa
l’étude présente, les plantes aquatiques et graminoides
perdaient de l’importance au profit des plantes
annuelles a feuilles larges (LeResche et Davis 1973).
Dans le méme état, Cushwa et Coady (1976) ont
observé pour leur part que les ramilles dominaient les
contenus de rumen au printemps, que les feuilles
étaient plus importantes en été et que les Orignaux
broutaient occasionnellement des plantes herbacées
au printemps et en été.

Le tableau | permet d’identifier les espéces végétales
qui sont les plus importantes pour l’Orignal pendant
la période considérée. Ainsi, au printemps, le Sapin
Baumier, l’If du Canada (Taxus canadensis) et ? Era-
ble 4 Epis comptent pour plus du trois quarts des

TABLEAU |—Importance relative (%) des végétaux retrouvés dans les rumens d’Orignaux selon la saison, 1977-1979, réserve
de La Vérendrye, Québec. Les espéces représentant moins de 0.5% des fréquences d’observation ont été omises

Printemps’
(n = 6)
Abies balsamea 29
Taxus canadensis 24
Acer spicatum 24
Thuja occidentalis 2
Prunus spp. l
Salix spp. 13
Populus tremuloides I
Typha spp. _—

Dryopteris spp. l

Diervilla lonicera =
Corylus cornuta 3

Acer saccharum =
Eriocaulon septangulare _
Betula spp.* l

Myrica gale

Viburnum alnifolium

Mousses

_Diaprés le poids sec des ramilles.

D’apres le contenu des tamis a maille de 6.4 et 4.0 mm.
“Trace (< 0.5%).
“Surtout Betula papyrifera.

Début de été” Fin de lété” Automne”
(n = 10) (n = 5) (n = 36)
10 = I
T a ili

68 68 21
4 12 52
3 14 6
9) 6 3
6 = 5)
as _ 5
tas 128s 4
= ae T
ts A »)
2 bah ilk
ae an
I ws ii
2 ae if

l
V. cassinoides l
|

54 THE CANADIAN FIELD-NATURALIST

ramilles identifiées dans les rumens. La consomma-
tion en grande quantité de sapin et spécialement dif
est particuliére a cette saison car, en hiver, ces deux
espéces sont beaucoup moins utilisées dans la région
d’étude (Joyal 1976; Créte données non publiées);
d’ailleurs, l’if est peu disponible en hiver a cause de sa
petite taille. Au début de l’été, l’Erable a Epis devient
lespéce dominante dans le régime alimentaire des
Orignaux, représentant prés de 70% des fragments
non ligneux identifiés. Par ailleurs, la présence d’ai-
guilles de sapin et d’if dans les rumens apreés le 15 mai
s’explique en grande partie par leur long temps de
digestion (Bergerud et Russel 1964). Parmi les plantes
aquatiques identifiées, les typhas (7ypha spp.) sont les
plus fréquemment rencontreées. A la fin de ’été, l’Era-
ble a Epis compose encore la majeure partie de la
nourriture des Orignaux. Le Peuplier Faux-Tremble
et les cerisiers (Prunus spp.) possédent quand méme
une importance appréciable. A l’automne, Il’Erable a
Epis céde le premier rang aux cerisiers (surtout P.
pensylvanica). De plus, des espéces tels le Diéreville
Chévrefeuille (Diervilla lonicera) et les dryoptérides
(Dryopteris spp.), qui sont rarement broutées en d’au-
tres temps, font leur apparition dans la nourriture des
Orignaux. Ces changements de préférence sont prob-
ablement attribuables au fait que les Orignaux recher-
chent les espéces qui conservent leurs feuilles vertes le
plus longtemps a l’automne. D’avril a octobre, les
saules (Salix spp.) occupent toujours une fraction
importante du régime alimentaire des Orignaux; si
leur importance est moindre au Québec qu’au Minne-
sota (Peek et al. 1976) ou qu’en Alaska (Cushwa et
Coady 1976), c’est probablement pour la simple rai-
son quils sont moins abondants dans la région
d’étude.

Les observations d’arbres et d’arbustes broutés par
les Orignaux (tableau 2) confirment la grande impor-
tance que revét |’Erable a Epis pour ces cervidés dans
le sud-ouest du Québec en été. Si !importance relative
de l’Erable a Epis est moindre d’aprés les observations
visuelles que d’apres les contenus stomacaux, c’est
surtout que la premiére estimation ne représente
qu’une fréquence d’occurence. De plus, il est fréquent
que ces deux méthodes conduisent a des estimations
différentes, l’observation visuelle étant moins
complete (Smith et Shandruk 1979). L’importance de
l’Erable a Epis s’étend probablement a tout le sud du
Québec car cette espéce est souvent abondante dans
les aires d’hivernage d’Orignaux des autres régions de
la province (Brassard et al. 1974). Ces résultats vont
toutefois a l’encontre des observations de Joyal et
Scherrer (1978) qui, pour la période estivale dans un
secteur situé a environ 100 km a lest de la région
d’étude, calculérent que le Cerisier de Pennsylvanie
dominait la nourriture des Orignaux. Cette contradic-

Vol. 95

TABLEAU 2—Fréquence d’occurrence (%) d’arbres et d’ar-
bustes broutés par l’Orignal entre juin et aott dans le sud-
ouest du Québec, établie a partir d’observations faites lors de
déplacements en pleine forét, en 1978 et 1979

Fréquence d’occurrence

Espéce (n = 118)
Acer spicatum 40
Viburnum cassinoides 9
Prunus pensylvanica 8
Betula papyrifera 8
Corylus cornuta 7
Viburnum alnifolium 7
Acer saccharum 5
Autres* 16

“Comprend par ordre d’importance: Populus tremuloides,
Betula alleghaniensis, Sorbus americana, Salix spp., Acer
rubrum, Diervilla lonicera, Cornus stolonifera, Lonicera
canadensis, Cornus alternifolia, Nenopanthus mucronatus.

tion apparente semble attribuable a la faible superficie
de leur aire al’étude, ot d’ailleurs, l’Erable a Epis était
rare (environ 10 km?2: R. Joyal, Université du Québec
a Montréal, communication. personnelle). Par ail-
leurs, l’Erable a Epis semble jouer un réle de moins en
moins important pour!’Orignal au fur et a mesure que
lon s’éloigne vers l’ouest, comme le suggerent les
résultats de l’Isle Royale (Belovski et al. 1973;
Miquelle et Jordan 1979) et du Minnesota (Peek et al.
1976).

Variation du régime alimentaire en fonction de l'dge et
du sexe

Les trois échantillons provenant de petits de l'année
qui furent récoltés au courant de l’été montraient une
composition semblable a celle des adultes. Les spéci-
mens du 30 juin et du 30 juillet faisaient voir une nette
dominance de l’Erable a Epis alors que l’échantillon
du 12 juillet était dominé par les plantes graminoides.
Dans ce dernier cas, la mére du petit avait été tuée lors
du méme accident; la comparaison statistique du con-
tenu des rumens de la mére et du petit (fragments non
ligneux des deux premiers tamis) a révélé une dif-
férence trés nette (P< 0.005), l’Erable 4 Epis domi-
nant le contenu du rumen de la mere. Ainsi les jeunes
Orignaux peuvent commencer a brouter une nourri-
ture semblable a celle des adultes avant l’age d’un
mois, mais ils ne semblent pas nécessairement imiter
en tout point les habitudes alimentaires de leur mére.

Durant l’automne 1978, six échantillons parmi les
14 prélevés sur des femelles étaient complétement
liquides, ne permettant identification d’aucun frag-
ment. Par contre, pour la méme saison, aucun échan-
tillon provenant de 16 males ne possédait cette carac-
téristique, ni ceux récoltés a l’automne 1977 (8
femelles, 4 males). Cette observation permet

1981

d’avancer Il’hypothése que certaines femelles cessent
de s’alimenter pendant le rut: le fait qu’aucun corps
jaune d’ovulation n’ait été observé dans sept ovaires
provenant de ces six femelles, toutes aptes a la repro-
duction, suggére qu’elles étaient sur le point d’ovuler,
ce qui donne plus de vraisemblance a l’hypothése
avancée. Par contre, en 1977, on comptait cing corps
jaunes et un follicule primaire fraichement rupturé
dans 10 ovaires disponibles provenant des huit
femelles. Le phénoméne de jeine observé en 1978
s’étendit du 21 septembre au 8 octobre.

Discussion

L’analyse du contenu des rumens d’Orignaux
s’'avere particuliérement laborieuse. Puglisi et al.
(1978), confrontés 4 ce probleme, ont proposé des
moyens de rendre la technique plus efficace en rapport
avec le Cerf de Virginie. Dans le méme but, pour
lOrignal, nous recommandons de m utiliser qu’un seul
tamis munide mailles d’environ 6 mm; il faut, en effet,
considérer que dans 64% des cas, les deux premiers
tamis possédaient une composition semblable et que
de toute maniere, cette technique est plus descriptive
que quantitative pour trois raisons: (1) certains
végétaux ou parties de végétaux sont plus facilement
identifiables que d’autres; (2) apres la premiére identi-
fication d’une espéce ou groupe végétal, on anticipe a
nouveau sa présence dans l’échantillon; (3) la vitesse
de digestion des végétaux n’est pas uniforme (Ber-
gerud et Russel 1964).

Miquelle et Jordan (1979) spéculant sur le rdle de la
diversité dans le régime alimentaire de l’Orignal, ont
émis lhypothése que ce cervidé choisissait comme
nourriture de base en été l’espece végétale comestible
la plus abondante d’une région donnée; d’apreés leurs
résultats, ils avanceérent que cette espéce devrait rep-
résenter 60-70% de la prise alimentaire quotidienne.
Nos résultats supportent assez bien cette hypothése
puisque l’Erable a Epis constitue de fait l’espéce com-
estible la plus disponible dans la région d’étude (Créte
et Jordan, données non publiées).

Les faiblesses de la technique employée n’em-
péchent pas de conclure que l’Erable a Epis constitute
la base du régime alimentaire de Original du sud du
Québec. Des travaux antérieurs (Brassard et al. 1974;
Joyal 1976; Bédard et al. 1978) avaient montré l’im-
portance de cette espéce végétale pour les Orignaux en
hiver; les résultats présentés ici montrent qu’en fait,
lErable a Epis est probablement encore plus impor-
tant en été.

Remerciements

Je veux d’abord remercier le personnel de la réserve
de La Vérendrye, les guides, les chasseurs et quelques
consoeurs et confreres de travail pour avoir recueilli
les échantillons. Je veux d’autre part souligner le tra-

CRETE ET JORDAN: NOURRITURE DE lORIGINAL 5)5)

vail méticuleux de K. Levine qui a fait analyse des
contenus de rumen en laboratoire. Je veux aussi
exprimer ma reconnaissance a R. Ouellet quia estimé
lage des bétes. Merci enfin a J.-R. Moreau pour la
figure et a J. Huot et R. Joyal pour leur commentaire
du texte original.

Références

Abler, W. A. 1974. Effects of energy and protein levels on
attainment of puberty and steroid hormone production in
captive female White-tailed Deer. Thése M.Sc., Virginia
Polytechnic Institute and State University. 84 pp.

Bahnak, B. R., M. C. Holland, L. J. Verme et J. J. Ozoga.
1979. Seasonal and nutritional effects on serum nitrogen
constituents in White-tailed Deer. Journal of Wildlife
Management 43(2): 454-460.

Bédard, J., M. Créte et E. Audy. 1978. Short-term influ-
ence of Moose upon woody plants of anearly seral winter-
ing site in Gaspé Peninsula, Québec. Canadian Journal of
Forest Research. 8(4): 407-415.

Belovsky, G.E., P. A. Jordan et D.B. Botkin.
1973. Summer browsing by Moose in relation to prefer-
ence, and animal density: a new quantitative approach.
Proceedings of the North American Moose Conference
and Workshop 9: 101-122.

Bergerud, A. T., et L. Russel. 1964. Evaluation of rumen
food analysis for Newfoundland Caribou. Journal of
Wildlife Management 28(4): 809-814.

Bouchard, R., et G. Moisan. 1974. Chasse contrélée a
l’Orignal dans les parcs et réserves du Québec (1962-1972).
Naturaliste canadien 101(3-4): 689-704.

Brassard, J.-M., E. Audy, M. Créte et P. Grenier.
1974. Distribution and winter habitat of Moose in
Québec. Naturaliste canadien 101(1-2): 67-80.

Chamrad, A. D., et T. W. Box. 1964. A point frame for
sampling rumen contents. Journal of Wildlife Manage-
ment 28(3): 473-477.

Créte, M. 1977. Importance de la coupe forestiére sur l’ha-
bitat hivernal de !’Orignal dans le sud-ouest du Québec.
Canadian Journal of Forest Research 7(2): 241-257.

Créte, M., R. J. Taylor et P. A. Jordan. 1981. Optimiza-
tion of Moose harvest in southwestern Québec. Journal of
Wildlife Management 45 (2): Sous presse.

Cushwa, C. T., et J. Coady. 1976. Food habits of Moose,
Alces alces, in Alaska: a preliminary study using rumen
contents analysis. Canadian Field-Naturalist 90(1): 11-16.

Ferland, M. G., et R. M. Gagnon. 1967. Climat du Québec
méridional. Ministére Richesses Naturelles du Québec,
MP-13. 93 pp.

Franzmann, A. W., R. E. LeResche, R. A. Rausch et J. L.
Oldemeyer. 1978. Alaska Moose measurements and
weights and measurements-—weight relationships. Cana-
dian Journal of Zoology 56(2): 298-306.

Gasaway, W. A., et J. W. Coady. 1974. Review of energy
requirements and rumen fermentation in Moose and other
ruminants. Naturaliste Canadien 101(1-2): 227-262.

Joyal, R. 1976. Winter foods of Moose in La Vérendrye
Park, Québec: an evaluation of two browse survey
methods. Canadian Journal of Zoology 54(10):
1765-1770.

56 THE CANADIAN FIELD-NATURALIST

Joyal, R., et B. Scherrer. 1978. Summer movements and
feeding by Moose in Western Québec. Canadian Field-
Naturalist 92(3): 252-258.

LeResche, R. E., et J. L. Davis. 1973. Importance of non-
browse foods to Moose on the Kenai peninsula, Alaska.
Journal of Wildlife Management 37(3): 279-287.

McCaffery, K.R., J. Tranetzi et J. Piechura, Jr.
1974. Summer food of deerin Northern Wisconsin. Jour-
nal of Wildlife Management 38(2): 215-219.

Miquelle, D. G., et P. A. Jordan. 1979. The importance of
diversity in the diet of moose. Proceedings of the North
American Moose Conference and Workshop 15: 54-80.

Moen, A. N. 1978. Seasonal changes in heart rates, activi-
ty, metabolism, and forage intake of White-tailed Deer.
Journal of Wildlife Management 42(4): 715-738.

Peek, J. M., D. L. Urich et R. J. Mackie. 1976. Moose ha-
bitat selection and relationships to forest management in
Northeastern Minnesota. Wildlife Monograph 48. 65 pp.

Puglisi, M. J., S. A. Liscinsky et R. J. Hurlow. 1978. An
improved methodology of rumen content analysis for
White-tailed Deer. Journal of Wildlife Management 42(2):
397-403.

Vol. 95

Sergeant, D. E., et D. H. Pimlott. 1959. Age determination
in Moose from sectioned incisor teeth. Journal of Wildlife
Management 23(3): 315-321.

Skinner, W. R., et E. S. Telfer. 1974. Spring, summer and
fall foods of deer in New Brunswick. Journal of Wildlife
Management 38(2): 210-214.

Smith, A.D., et L. J. Shandruk. 1979. Comparison of
fecal, rumen and utilization methods for ascertaining
Pronghorn diets. Journal of Range Management 32(4):
275-279.

Verme, L. J. 1969. Reproductive patterns of White-tailed
Deer related to nutritional plane. Journal of Wildlife
Management 33(4): 881-887.

Verme, L. J. 1977. Assessment of natal mortality in upper
Michigan deer. Journal of Wildlife Management 41(4):
700-708.

Villeneuve, G.-O. 1969. Feuillaison et défoliation des feuil-
lus au Québec. Naturaliste Canadien 96(4): 491-505.

Received 2 May 1980
Accepted 11 August 1980

Caribou (Rangifer tarandus) Encounters with

Pipelines in Northern Alaska

WAYNE C. HANSON

Ecological Sciences Department, Battelle Pacific Northwest Laboratory, P.O. Box 999, Richland, Washington 99352

Hanson, W. C. 1981. Caribou (Rangifer tarandus) encounters with pipelines in northern Alaska. Canadian Field-Naturalist

95(1): 57-62.

Behavior of Caribou (Rangifer tarandus) upon encountering an experimental gas pipeline berm in northern Alaska suggested
that a visual barrier effect greater than 1.2 m above ground level had a pronounced effect of deflecting the movements of
Caribou. Animals readily traversed lower berms but avoided high berms and thermokarst areas.

Key Words: Caribou, northern Alaska, pipeline, visual barrier, behavior.

Development of the oil fields on the North Slope of
Alaska has created one of the most serious confronta-
tions of industrial and conservation interests of our
time. The principal center of controversy has been the
Trans-Alaska pipeline, the 1288-km conduit of |.2-m-
diameter steel pipe that moves the oil from Prudhoe
Bay to Valdez. The environmental implications of
such an installation are considerable in view of the
severe local damage to northern Alaska from past
exploration and initial development activities (Brooks
et al. 1971; Klein 1970; Laycock 1970; Weeden and
Klein 1971). Experiments by Child (1974) with Cari-
bou encountering pipeline simulations at Prudhoe
Bay and the observations by Cameron et al. (1979)
and Roby (1978) of Caribou behavior along the
Trans-Alaska pipeline and haul road have verified
that Caribou are sensitive to these human activities
and installations associated with oil development.

Several of these reports have emphasized the lack of ©

direct experience upon which to predict or mitigate
Caribou reactions to pipelines and other installations
in northern tundra environments. I herein report
observations of northern Alaska Caribou encounter-
ing a raised berm resulting from pipeline burial similar
to the general type of construction expected to be
undertaken to transport natural gas from the Prudhoe
Bay area southward, pending the approval of various
government agencies and the surmounting of several
engineering and economic problems.

Study Area and Methods

The study area was a natural gas pipeline installa-
tion and operation of the Gas Arctic Systems Study
Group research facility at Prudhoe Bay, Alaska (Fig-
ure 1). This installation consisted of a 610-m (2000-ft)
rectangular loop of standard “Alyeska pipe” (1.2-m
diameter) through which refrigerated air (the test gas)
was circulated during the 1971-1973 study period. A

Sy

static pipeline test area and a road construction simu-
lation test area were included in the study. The pipe-
line loop was so constructed that the east 243-m (800-
ft) side was half buried in the substrate and covered
with a 0.8-m (2.5-ft) berm and fill of residual material
and gravel, resulting in a raised barrier of 1.0-1.7 m
above the tundra. The west 243-m side was fully bur-
ied in the tundra under a berm ranging in height from
nearly level with the surrounding terrain to about
0.8 m above ground level (agl). This pipeline was
constructed during April-May 1971 in the continuous
permafrost with high ice content which is typical of
much of the North Slope landscape. Most of the
berms were finished with a 3:1 slope. In the middle of
the 61-m south side of the loop, the pipeline rose and
passed through a refrigeration, heating, and power
control center. The pipeline substrate was implanted
with thermocouples, strain gauges, heat flux trans-
ducers, conductivity probes, soil moisture and soil
density instruments to study permafrost regression,
thermal patterns of the berm and fill materials, and
other geological phenomena. The northeast corner of
the pipeline passed through a 400-m? thaw pond in
which temperature and water chemistry were studied.
Eleven 3-m-tall cross-shaped metal deflection poles
(Figure 2) were vertically mounted on the east and
west legs of the pipeline at 15.2-m intervals to provide
means of measuring pipeline movement optically.
Two 36.5-m sections of nonoperating instrumented
pipeline, one section half buried and the other fully
buried, were located about 75 m east of the pipeline
loop to study static pipeline effects. These sections
were within an 8. 1-ha (1 ha = 10 000 m2) ice road con-
struction simulation test area for studying winter tun-
dra preparation for equipment traffic expected during
pipeline construction.

The location of the Prudhoe Bay Test Site within
the oil field area was fortuitously isolated from other

58 THE CANADIAN FIELD-NATURALIST

Volaos

a

FiGureE |. Aerial photo of the Prudhoe Bay Test Site during August 1971. The installation consists of a 610-m rectangular
loop of 1.2-m-diameter pipe, of which the western side (left) was fully buried and the eastern side was half buried.
Control center is in lower left, static test and road construction simulation test areas are located at right. The Prudhoe
Bay spine road crosses lower part of photo. Light-colored barrels outlining the Gas Arctic Systems Study Group leased
property were removed to facilitate Caribou encounters with the pipeline berms. Reticulated ground pattern results
from buried ice wedges forming polygons characteristic of many arctic coastal plain areas of high ice content

permafrost.

industrial installations and oriented in such a manner
that Caribou moving between the Sagavanirktok
River mouth and western portions of their summer
range encountered the pipeline berms. These encoun-
ters were decumented by visual observations made
from anelevated platform constructed adjacent to the
control center and downwind of the pipeline loop and
from animal tracks in the substrates. Berms were
inspected twice daily and raked free of tracks after
photographic documentation of each encounter. Ten
encounters occurred in 1971 and 24 episodes of Cari-
bou-pipeline interactions during 1972 were combined
into 21 separate encounters.

Results and Discussion

The 1971 encounters began a few days after bands
of 36,51, and 60 Caribou were observed 3 km north of
the facility. The animals initially kept their distance
from the test facility probably because the nearby
spine road carried appreciable vehicular traffic. Six
encounters were visually documented and the other
four occurred while I was absent from the facility
(usually late evening until early the following morn-
ing). Seven encounters were by single animals and
three were by two animals each, reflecting the general
dispersal of Caribou on their summer range (Hem-
ming 1971), especially in the Prudhoe Bay area.

HANSON: CARIBOU-PIPELINE ENCOUNTERS IN ALASKA 59

FicurE 2. Photograph of Caribou in 1971 coursing east berm at point where it is 1.1 m high. Animal at this point carried its
head with eye level ca.0.5 m above grade. Cross-shaped metal deflection poles were 3.05 m tall with 1.0-m crossarms.

The 1972 interactions involved a broader spectrum
of animal groups than the 1971 data and ranged from
single animals interacting with the pipeline berms ona
single occasion to a band of five bulls that interacted
on 18 occasions, either singly, in combinations, orasa
group.

Most of the encounters during both seasons
occurred at the north and west sides of the loop, where
berm heights were lower and there was less noise from
the facility diesel power generators and vehicular traf-
fic, although the latter impacts did not appear to affect
the Caribou greatly. Other field experiments with
sound simulators operated in northeastern Alaska
during Caribou migration periods (McCourt et al.
1974) suggest that the vehicular traffic was probably
more important than the generator noise in deterring
the Caribou. The animals readily crossed the berms of
both residual material and gravel, especially where
berm height was <<1.2 m. Encounters at berm heights

> 1.2 m were characterized by animals (1) approach-
ing obliquely, pausing at the top of the berm, and then
descending in the direction from which they had
approached (Figure 3); (2) first crossing lower berms
on their way to revegetation plantings on the higher
berms and usually retracing their paths to leave the
study area over lower berms, as in the southwest
corner of the pipeline loop (Figure 4); (3) interacting
as a group, usually under the leadership of a large bull;
or (4) under severe insect harassment from nostril flies
(Cephenemyia trompe), warble flies (Oedemagena
tarandi), and mosquitoes (Aedes spp.). Caribou often
laid or stood on the gravel berms to seek relief from
insect harassment (Figure 5).

The animals avoided the northwest sector of the
pipeline loop in 1971 (Figure 3) where several pools of
water were formed by melting snow and general sub-
sidence related to melting ice wedges fractured by the
pipeline construction. This behavior was similar to

60 THE CANADIAN FIELD-NATURALIST

PRUDHOE BAY TEST SITE CARIBOU-PIPELINE ENCOUNTERS
1971

Lease BOY NDARY 7
TT ie
|

ll

PLANE D 1.) aie

FULLY-BURIED

STATIC
/ PLANE B 4 TEST

, HALF-BURIED

CARIBOU
ENCOUNTER
NUMBERS

DEFLECTION POLES
15-m INTERVALS

ii, L\HERMOCOUPLE
PLANE A

WEST LEG
FULLY-BURIED

—— EAST LEG N

HALF-BURIED *

a

a

0 25 50
METRES

FIGURE 3. Schematic diagram illustrating routes of Caribou
involved in 10 pipeline encounter episodes from 23
July to 2 August 1971. Detailed routes are shown to
illustrate avoidance of higher (1.7 m) berms in south-
east sector and pools of water in subsidence craters in
northwest sector.

that of Caribou encountering “topographic confine-
ment” at a nearby study area (Child 1974). These areas
were reclaimed by adding topsoil during April 1972,
so that the Caribou then used the areas regularly
(Figure 4).

Most Caribou appeared wary of deflection poles,
thermocouple planes, and boardwalks and usually
avoided them. However, during 1972 a large-antlered
bull walked atop the east berm fora distance of 130 m,
tilting his antlers to avoid the crossarms of eight
deflection poles in his path. Clearance under the cross-
arms ranged from 0.9 to 1.4 m. He then continued a
course toward an elevated (1.0—1.5 m) effluent line,
which he coursed for 56 m before lightly touching his
antlers to the wooden housing 2-3 times at a point
where clearance was 1.1 m. He then continued to a
point where clearance increased to 1.5 m and passed
under the line. The animal was under moderate to
severe insect harassment during the episode.

Vol. 95

PRUDHOE BAY TEST SITE CARIBOU-PIPELINE ENCOUNTERS

STATIC TEST

FULLY-BURIED

HALF-BURIED

BERM HEIGHT IN METRES
_ |=— ABOVE GRADE

FIGURE 4. Schematic diagram illustrating routes used by
Caribou in the 21 encounters from 15 July to 4 August
1972. Large open arrows indicate number of Caribou
and direction of crossing berms, small dark arrows
represent individual animal paths. Numbers in center
of pipeline berms indicate maximum height (m)
above grade. Note increased use of northwest sector
following reclamation of subsidence craters, apparent
selection of north and west berm sectors for entry,
and east berm for exit.

Two bull Caribou in 1971 (Figure 3) obviously
avoided the combination of thermocouple housing
and boardwalk in the static test area. They reacted by
adopting the alarm stance, urinated, alertly watched
the installation, and then bolted and trotted ina tight
circle. Such behavior was earlier described by Pruitt
(1960). This reaction was similar to that noted several
times during summer when Caribou encountered
roadways that they could not see over. In such cases,
they often appeared uneasy and laid down for some
time; others would intermittently graze while watch-
ing the road and vehicles, crossing warily when lulls in
traffic permitted. As such behavior resulted in suc-
cessful crossings, it was misinterpreted by some per-
sons as unqualified acceptance by Caribou of man’s
activities and installations. Roads, airstrips, and

1981

HANSON: CARIBOU-PIPELINE ENCOUNTERS IN ALASKA 61

FicureE 5. Caribou resting on gravel berm to obtain relief from insect harrassment. Animals appeared undisturbed by steady
noise from diesel generator exhaust located about 60 m away and on opposite side of control center.

equipment pads on the North Slope are usually con-
structed of |.5-2 m of gravel to insulate the perma-
frost and to allow winds to sweep them clear of snow
during the winter. This height also makes them attrac-
tive to Caribou harassed by insects in summer, and
once on the elevated structures the Caribou appeared
reluctant to leave them. I watched a cow and her calf
repeatedly trot back and forth over a |-km intercon-
necting gravel road to avoid vehicle traffic rather than
leave the breeze-swept elevated roadway during the
peak of the mosquito season.

Further evidence of an important “visual barrier”
effect operative in Caribou behavior toward the berms
was supplied during a 1971 encounter. A cow, accom-
panied by her calf, climbed the gravel berm and
paused at a point that provided a view of the lake in
her path, and then left the study area over a lower
berm. Why the lake should deter her is a matter of

conjecture; the proximity of an alternative and easier
route over lower berms with minimal visual obstruc-
tion that had been previously used by the animals in
an earlier encounter perhaps provided the stimulus for
her decision.

Caribou were attracted to fertilized revegetation
plantings of Arctared Fescue (Festuca rubra), Nugget
Bluegrass (Poa pratensis), and several other exotic
grasses at several locations on the experimental pipe-
line berms during the 1972 season. The attractiveness
of these plantings was probably a factor in the accep-
tance of the relatively high berms in the southeast
corner of the loop by a five-bull band that contributed
most of the encounters during 1972.

These observations are by no means indicative of
the entire spectrum of Caribou behavior upon
encountering pipeline berms. Herd behavior is differ-
ent from that of individuals or small bands (Child

62 THE CANADIAN FIELD-NATURALIST

1974), and extrapolations from one to the other are
tenuous. Seasonal differences in behavior may also be
a consideration because of weather conditions, physi-
ological variables, and social organization of Caribou
herds. However, it was apparent that Caribou dis-
criminated in their acceptance of certain of the pipe-
line berms studied and a number of encounters were
recorded. The greater acceptance of the pipeline
berms by bulls in 1972 compared to the behavior of
cows and calves during both seasons and the
decreased sensitivity to the installation with increased
experience Is consistent with observations of Caribou
behavior along the oil pipeline and haul road (Roby
1978). However, insect harassment was concluded to
be the greatest inducement for Caribou on their
summer range to accept crossing facilities of the oil
pipeline simulation (Child 1974).

These observations suggest that suitable gas pipe-
line crossings for Caribou might be attainable if they
are knowledgeably located (LeResche and Linderman
1975; McCourt et al. 1974) and properly constructed.
From experience in Scandinavia, Klein (1971)
stressed the importance of utilizing traditional cross-
ings, and in this report I have suggested that the visual
barrier implications may be moderated by construct-
ing pipeline berms of <1.2 m, or otherwise engineer-
ing them so that visual accommodation can be made
by Caribou (Banfield 1974).

Acknowledgments

Amos Mathews, former Executive Vice-President
of Alaskan Arctic Gas, gave approval to use the data
reported herein. K. N. Child, L. E. Eberhardt, J. E.
Hemming, and D. R. Klein made many helpful com-
ments on the manuscript. This report was prepared
under United States Department of Energy Contract
No. DE-AC06-76RLO 1830.

Literature Cited

Banfield, A. W.F. 1974. The relationship of Caribou
migration behavior to pipeline construction. Jn The
behavior of ungulates and its relation to management.
Volume 2. Edited by V. Geist and F. Walther. Interna-
tional Union for Conservation of Nature and Natural
Resources. IUCN Publication Number 24. Morges, Swit-
zerland. pp. 797-804.

Vol. 95

Brooks, J. W., J. C. Bartonek, D. R. Klein, D. L. Spencer,
and A. S. Thayer. 1971. Environmental influences of oil
and gas development in the Arctic Slope and Beaufort Sea.
Bureau of Sport Fisheries and Wildlife. United States
Department of the Interior Resource Publication 96.
Washington, D.C. 24 pp.

Cameron, R. D., K. R. Whitten, W. T. Smith, and D. D.
Roby. 1979. Caribou distribution and group composi-
tion associated with construction of the Trans-Alaska
pipeline. Canadian Field-Naturalist 93: 155-162.

Child, K. N. 1974. Reaction of Caribou to various types of
simulated pipelines at Prudhoe Bay, Alaska. Jn The behav-
ior 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. IUCN
Publication Number 24. Morges, Switzerland. pp.
805-812.

Hemming, J. E. 1971. The distribution and movement
patterns of Caribou in Alaska. Game Technical Bulletin
Number I, Alaska Department of Fish and Game, Juneau.
60 pp.

Klein, D. R. 1970. The impact of oil development in Alaska
(a photo essay). /n Proceedings of the Conference on
Productivity and Conservation in Northern Circumpolar
Lands, Edmonton, Alberta, 15-17 October 1969. Interna-
tional Union for Conservation of Nature and Natural
Resources, Morges, Switzerland. pp. 209-242.

Klein, D. R. 1971. Reaction of reindeer to obstructions and
disturbances. Science 173: 393-398.

Laycock, G. 1970. Kiss the North Slope goodbye?
Audubon 72: 68-75.

LeResche, R. E.,and S. A. Linderman. 1975. Caribou trail
systems in northeastern Alaska. Arctic 28: 54-61.

McCourt, K.H., J. D. Feist, D. Doll, and J. J. Russell.
1974. Disturbance studies of Caribou and other mammals
in the Yukon and Alaska, 1972. Arctic Gas Biological
Report Series, Volume 5. Canadian Arctic Gas Study
Limited and Alaskan Arctic Gas Study Company, Cal-
gary, Alberta. 246 pp.

Pruitt, W. O., Jr. 1960. Behavior of the Barren-ground
Caribou. University of Alaska, Biological Papers Number
3. 44 pp.

Roby, D. D. 1978. Behavioral patterns of Barren-ground
Caribou of the Central Arctic herd adjacent to the Trans-
Alaska oil pipeline. M.S. thesis, University of Alaska,
Fairbanks. 200 pp.

Weeden, R. B., and D. R. Klein. 1971. Wildlife and oil: a
survey of critical issues in Alaska. Polar Record 15:
479-494.

Received 20 May 1980
Accepted 12 August 1980

Analysis of Weight Lost by Eggs of
Eleven Species of Birds during Incubation

T. H. MANNING

R.R. 4, Merrickville, Ontario KOG INO

Manning, T. H. 1981. Analyses of weight lost by eggs of eleven species of birds during incubation. Canadian Field-
Naturalist 95(1): 63-68.

From 4 to 323 eggs of 11 species of birds were weighed at least twice during incubation to determine daily weight loss. Pipped
eggs were excluded. There was an unexplained difference between years in the daily weight loss of two species, the Red-winged
Blackbird, Age/aius phoeniceus, and the Barn Swallow, Hirundo rustica. For six species the variance of daily weight loss was
partitioned into egg and clutch components. Their means were 62.6% foreggs and 37.4% for clutches. The mean coefficient of
variation for nine species, based on total variance (eggs and clutches), was 28.7. The reduced major axis equation for the daily
weight loss (L) on fresh egg weight (W), using data for 56 species from this and other studies was L = 0.143 w?.747 = 0.0255
(r = 0.968). The total weight lost during incubation as a percentage of fresh egg weight averaged 14.1 + 0.64 for 11 species. The
reduced major axis equation for daily weight loss multiplied by length of incubation period (/) on fresh egg weight was
L-1= 0.153 W °?>* 99% (r = 0.987) or on natural scale L-/ = -0.116 + 0.152 + 0.0073 W(r = 0.990). These equations indicate an
average weight loss of 15% of fresh egg weight during incubation.

Key Words: bird, egg, clutch, weight loss, incubation, partition of variance, coefficient of variation.

Weight loss by eggs during incubation is caused _as whole clutches or in the case of the Sora as partial
almost exclusively by diffusion of water vapor clutches (lots). The two to five eggs that made upa lot
through the shell pores (Kendeigh 1940; Rahnand Ar _ were randomly taken from within a single clutch, so
1974). The weight loss therefore varies with the poros- that there was no lot variance component, apart from
ity, or its equivalent (Ar et al. 1974); water vapor that attributable to eggs, and the lot variance there-
conductance of the shell; as well as with humidity, fore provides an unbiased estimate of the egg vari-
temperature, and barometric pressure (Aretal. 1974). ance. At Merrickville I used 10-g and 5-g Pesola bal-
Data on weight loss during incubation has been ances. These read to 0.2 and 0.1 g. respectively, and
recorded fairly frequently and Drent (1970) collected could be estimated to 0.02 g with reasonable consis-
from the literature records of 46 species, including __ tency. All eggs were therefore weighed individually.
one, the Sora, Porzana carolina, also used in this At North Point eggs were usually weighed at 2-d
study. Most of the records used by Drent are casual _ intervals, and insome cases every day; at Merrickville
observations made in connection with other studies, | they were either weighed twice, usually at 6- to 10-d
and none that I have been able to check give standard _intervals, or three times, usually at 4-d intervals.
errors or related statistics. The purpose of the present Times were recorded to the nearest hour at both
paper is to record the fresh egg weights and daily places. At North Point I marked the eggs with
weight losses during natural incubation of eggs of 11 | numbers in India ink, at Merrickville with small dots
common birds, to examine the variability of the — of red nail polish, a more efficient method. Wind was
losses, to partition the variance into egg and clutch oftena problem inthe open marshes; to counter this at
components, and to examine the relationship between North Point, I suspended the scale in a large tin can,
the fresh weight of eggs and both the daily weightloss and at Merrickville in a transparent acrylic cylinder.
and the total weight lost during incubation. Except for the two Eastern Phoebe clutches, which

were laid in the same nest and almost certainly by the
same female, I avoided late clutches, and | think that
Materials and Methods all or almost all eggs measured in any one year were

In 1975, I weighed and measured eggs at North laid by different females. Also the American Robin
Point, Ontario, near the south end of James Bay and __ eggs used in 1978 were in nests at least | km distant
in 1976, 1978, and 1979 near Merrickville, about from the two nests used in 1976, and therefore were
50 kmS of Ottawa, Ontario(see Table | fordatabase probably laid by different females. The 1978 Barn
and scientific and common names of the birds). The Swallow eggs were, however, from the same barn as
50-g Pesola balance used at North Point read to0.5g _ those used in 1976. Similarly, the 1976, 1978, and 1979

and could be estimated to 0.1 g, but was not suffi- | Red-winged Blackbird eggs were from small marshes
ciently accurate to measure weight loss of the smaller within the same area of about 2 km radius.
eggs individually. Small eggs were therefore weighed As the rate of weight loss increases rapidly after an

63

64 THE CANADIAN FIELD-NATURALIST

egg is pipped (Barth 1952; Rahnetal. 1976), eggs with
a visible pipping fracture were excluded from the
study. At Merrickville the first day of incubation was
also excluded and usually balanced by the exclusion
of 1 or 2d at the end of the incubation. The periods
covered by the weight loss measurements were, there-
fore, near the central part of the incubation period
(Table 1) and should give a good estimate of the
average daily weight loss for the whole period prior to
pipping, even if, as has been found in some species
(Kendeigh 1940; Fisher 1969; Sotherland et al. 1980),
the rate of weight loss increases during incubation. As
the Sora begins incubation early in the laying period,
and up to 18 eggs were found in a single nest, some
nests contained eggs that were just starting to be incu-
bated as well as eggs nearing full term. The average
period of weight-loss measurements was, however,
probably not far from the middle of the incubation
period. I had no means of knowing when a Sora had
started incubation, but as only one clutch (five eggs)
had less than seven eggs when first weighed and as
only two more eggs were added to that clutch, it is
reasonable to suppose that all the Sora eggs weighed
were being incubated.

As some nests were not found before the start of
incubation, and some eggs, that had been weighed
when fresh were later destroyed by predators, the
mean fresh egg weight for most species was estimated
from the reduced major axes (RMA) for fresh egg
weight on volume index (length X breadth?) of those

Vol. 95

eggs that were weighed on the day they were laid
(Table 1), and the volume index of all eggs used for
weight-loss determination. Exceptions were (1) Amer-
ican Bittern, for which the ratio rather than the RMA
was used; (2) Wilson’s Phalarope, the eggs of which
lay flat in water, and were therefore thought to have
been incubated from 0 to 2 d (Hays and Le Croy 1971)
and their weight adjusted for | d of incubation; (3)
Common Grackle, all eggs of which were weighed on
the day laid; (4) Savannah Sparrow, for which the
RMA was calculated from eggs weighed in clutches on
the day the clutch was completed.

Weight losses used for calculating the means and
variances were weighted by the length of the period
covered by the observations. Fresh egg weights were
similarly weighted. The 95% confidence intervals for
egg variances are the least unbiased intervals; those
for clutches were calculated by Scheffé’s method
(Sokal and Rohlf 1969) and are not exact, owing to
unequal numbers of eggs in the clutches. The standard
errors of the percentages of the clutch components,
calculated according to Formula 4 of Swiger et al.
(1964) for the variance of interclass correlation, are
approximate for the same reason. The significance of
differences between both means and regression coeffi-
cients were tested by #, differences between variances
by F. The incubation periods are the periods between
the laying of the last egg and the hatching of that egg.
They were derived from various sources including this
study, but for most species precise data were lacking.

TABLE |—Number of eggs (clutches in parentheses) used for weight-loss estimation, number weighed when fresh, and
difference in days between midpoint of time over which weight loss was measured (MWL) and midpoint of incubation (M1)

No. eggs (weight loss)
North Pt.

Species and years

American Bittern, Botaurus lentiginosus (1975)
Sora, Porzana carolina (1975)
Killdeer, Charadrius vociferus (1975)
Common Snipe, Capella gallinago (1975, 1976)
Wilson’s Phalarope, Steganopus tricolor (1975)
Eastern Phoebe, Sayornis phoebe (1978)
Barn Swallow, Hirunde rustica (1975)
(1976)
(1978)
American Robin, Turdus migratorius (1976, 1978)
Red-winged Blackbird, Agelaius phoeniceus
(1975)
(1976)
(1978)
(1979)
Common Grackle, Quiscalus quiscula (1976)
Savannah Sparrow, Passerculus sandwichensis
(1975)

8 (2)
71 (18)
15 (4)
11 (3)
4 (1)
—(-)
19 (3)
—(—)
=)
—(—)

119 (29)
==\(=>)
aa Gar)
(=)
Pail)

19 (4)

No. eggs (fresh)

Merrickville North Pt. Merrickville MWL-MI
= (=) = (=) I (1) etl
= (=) 24 (8) = (=) 0?

4 (1) 3 (2) 4 (1) 4

4 (1) 12 (3) = (=) =4

an Gan) = (=) = (=) =o

9 (2) = (=) 5 (1) =!

al Gua) = (=) —= (=) =2,5)
44 (10) =) 44 (10) =1.3
31 (7) =(—) 24 (5) —0.5
26 (8) —(—) 36 (13) =15
= (5) = (©) == (=) —+0.4
59 (17) =) 69 (19) -1.3
67 (19) — (—) 67 (21) -0.8
78 (22) — (—) 11 (3) -1.9
10 (2) — (—) 10 (2) 0.0
= (=) 14 (3) == (=) -0.4

“See Materials and Methods.

1981 MANNING: WEIGHT LOST BY EGGS DURING INCUBATION 65

Results and Discussion
Differences between Years

The weight-loss measurements of the Red-winged
Blackbird eggs at Merrickville covered three seasons
and those of the Barn Swallow two seasons (Table 1).
Differences (11.4 mg-egg '-d' for the Red-winged
Blackbird and 6.0 mg egg '-d ' for the Barn Swallow)
between 1976 and 1978 weight-loss means (Table 2)
are highly significant (P < 0.01). This was particularly
surprising as all eggs laid in any one year were believed
to have been laid by different females, whereas some
females no doubt contributed eggs in more than one
year. In 1976 most of the Red-winged Blackbird and
Barn Swallow eggs were dipped ina salt (NaCl) solu-
tion for density measurement, then rinsed and dried
before being returned to the nest (Manning 1979). It
appeared therefore that the lower weight losses
recorded that year might have resulted from the
treatment. To test this hypothesis, 36 eggs of the
Domestic Hen, Gallus domesticus, were dipped in a
similar salt solution for 10 min, rinsed, and dried, then
incubated for 12.2 d with 36 untreated eggs. The
treated eggs lost an average of 7.4 + 10.0 mg-egg ‘-d'

more than the untreated eggs. A further test was made
in 1979 when half the eggs in 22 Red-winged Blackbird
nests were dipped in a salt solution for 4 min on two
different days and their weight loss measured over 4-d
periods to give 53 paired-egg periods. The treated eggs
lost 2.07 + 2.17 mg-egg '-d ' less than the untreated.
This difference (P> 0.2) is clearly insufficient to
account for the variation between years, and as the
same balances and techniques were used, and about
the same portion of the incubation period covered in
both years (Table 1), Iam unable to suggest a cause
other than differences in the environment, acting
either directly to cause variation in the nests’ micro-
climate or, perhaps more likely, indirectly to cause
variation in shell porosity through the condition of the
parent. A long severe winter may apparently increase
weight loss in eggs of the Domestic Hen (Landauer
1967). The 1978 Red-winged Blackbird eggs also had a
significantly greater variance than did the 1976 and
1979 eggs (P<0.05). The difference may be asso-
ciated with the greater mean, as the coefficients of
variation for the three years were similar (Table 2).
The variance of the 1978 Barn Swallow eggs was also

TABLE 2—Estimated mean fresh egg weight, mean daily loss of weight by eggs during incubation (with coefficient of variation
(CV)), length of incubation period (LIP), and weight loss during incubation asa percentage of fresh egg weight (WLI). Species

are listed in order of fresh egg weight

Fresh egg weight,’ Mean weight loss + SE,” LIP, WLI,

Species Egg-days N g mg-egg ‘-d' (ON d %
American Bittern 58 8 eggs 31.9 (36) 202.4 + 10.42 14.6 24 Te
Common Snipe 103 15 eggs 16.6 (16.0) 130.0 + 5.55 16.5 20 15.7
Killdeer 206 19 eggs 14.2 (14.5) O71! ae 3.95 23.1 5) 11.8
Wilson’s Phalarope 55 4 eggs 9.5 (9.4) 50.2 + 0.91 3.6 21 11.1
Sora 493 18 lots 9.0 (8.9) 61.0 + 3.09 ZS 18 12.2
Common Grackle 89 10 eggs 6.71 (—) 61.2+ 6.11 31.6 13 11.8
American Robin 176 26 eggs 6.36 (6.25) 75.0 = 5.23 35.6 13 15.3
Red-winged Blackbird

(North Point 1975) 914 29 clutches 4.31 (—) 53.8 + 2.09 20.8 11.2 =©14.0
Red-winged Blackbird

(Merrickville 1976) 397 59 eggs 4.10(—) 45.9 + 1.85 30.9 MS) W333
Red-winged Blackbird

(Merrickville 1978) 460 67 eggs 3.90 (—) 57.3 = 2.24 32.0 IES Os9)
Red-winged Blackbird

(Merrickville 1979) 518 78 eggs 4.05 (—) 48.6 + 1.46 26.6 IZA > Nebo)
Savannah Sparrow 146 4 clutches 2.18 (—) 33.4 + 4.48 26.8 I] 16.9
Eastern Phoebe 94 9 eggs PENBN(2a12) 22.6 + 4.31 57.4 16 17.0
Barn Swallow

(North Point 1975) 116 3 clutches 2.01 (1.85) 15.1 = 2.97 34.1 15 11.3
Barn Swallow

(Merrickville 1976) 440 44 eggs 1.97 (1.85) 16.8 + 0.69 27.1 15 12.8
Barn Swallow

(Merrickville 1978) 296 31 eggs 1.92 (1.85) 22.8 = 1.12 27.4 15 17.8

The figures in parentheses are taken for comparison from Schonwetter (1971-1979).
The standard error and coefficient of variation for the Red-Winged Blackbird (North Point), Barn Swallow (North Point),
and Savannah Sparrow are based on clutch means, those for Wilson’s Phalarope on eggs within a single clutch, and those for
the other species on the total variance of eggs and clutches.

66 THE CANADIAN FIELD-NATURALIST

greater, though not significantly so (P > 0.05), and
the coefficients of variation are again similar.

Partition of Variances
and Coefficient of Variation (CV)

The Red-winged Blackbird and Barn Swallow vari-
ances, partitioned in Table 3 into egg and clutch com-
ponents are the within-year variances of Table 2.
Judged by the standard errors, the percentages of the
variance components do not differ significantly
among the six species listed in Table 3. Their means
weighted according to the degrees of freedom for eggs
and clutches, respectively, are 62.6% for eggs and
37.4% for clutches. The unweighted means are similar,
62.2 and 37.8%. The clutch components are therefore
low when compared with those (85.0 + 7.8% and
56.5 + 18.1%) for the related measurement, water
vapor permeability, of eggshells of the Black-billed
Magpie, Pica pica, and the Yellow-headed Blackbird,
Xanthocephalus xanthocephalus, respectively
(Sotherland et al. 1979, with my standard errors), but
the difference appears to be significant only in the
former.

Most of the personal and instrumental error, which
may be relatively large owing to the small changes in
weight measured under field conditions, is probably

Vol. 95

segregated in the egg component of weight-loss vari-
ance. Segregation of error will not, however, be com-
plete as all.eggs in the clutch were weighed during the
same visit. The CVs given below indicate that the
variation in shell porosity, both of genetic and envi-
ronmental origin (Landauer 1967), is a large contribu-
tor to the total weight-loss variance and it may be
expected to affect both egg and clutch components.
Differences among nests in the humidity of their
microclimate, caused by variation in parental behav-
ior and weather conditions presumably affects only
the clutch component.

The mean of the CVs (Table 2) that are based on
total variance of eggs and clutches, when weighted by
the number of eggs or lots, is 28.7 (n = 384). lam not
aware of any published CVs for weight loss, but the
weighted mean CV for eggs of two species of gulls,
calculated for comparison from Morgan et al. (1978,
Table 2), is 23.7 (n = 31), and for five species of terns,
calculated from Rahn et al. (1976, Table 4), is 26.0
(n = 227). The corresponding CVs for water vapor
conductance derived, respectively, from Morgan etal.
(1978, Table 1) and Rahnet al. (1976, Table 2) are 21.8
(n = 29) and 18.6 (n = 78). The CVs for shell permea-
bility of the Black-billed Magpie and Yellow-headed
Blackbird derived from Sotherland et al. (1979, Table

TABLE 3—Partition of weight-loss variance into egg and clutch components with percentages of total variance and coeffi-

cients of variation (CV). Species in order of egg weight

Variance (95% % of total
Species and component df confidence limits), mg variance + SE CV
Common Snipe
Eggs 11 261 (143-613) 56 12.4
Clutches 201* (0.0-3690) 44 + 30.1 10.9
Killdeer
Eggs 14 129 (66-302) 54 16.9
Clutches 4 111** (5.5-1218) 46 + 26.1 15.7
Sora
Eggs 11 126 (60-337) 73 18.4
Clutches 46** (5.5-274) 27+ 12.8 11.2
American Robin
Eggs 18 424 (234-889) 60 27.5
Clutches i 287** (5.1-1604) 40 + 21.7 22.6
Red-winged Blackbird
(1976, 1978 and 1979)
Eggs 146 141 (114-181) 61 23.4
Clutches 55 89**** (51-157) 39+ 7.6 18.6
Barn Swallow
(1976 and 1978)
Eggs 58 18.6 (13-28) 69 21.8
Clutches 15 8.5*** (2.326) 31+ 13.3 14.7
<I P<<(05)
**—= P'< 025,
*#* = P< 0.005.

cone wa OLOO IE

1981

1) are 52.8 (n = 28) and 26.1 (m = 28), respectively. Ar
et al. (1974) give the means and standard deviations
for water vapor conductance of eggshells of 29 species
of birds, ranging in size from House Wren, Troglo-
dytes aedon, to Ostrich, Struthio camelus. The weigh-
ted mean of the CVs calculated from these figures is
28.2, but as the standard deviations appear to be
derived from differences among eggs and differences
among six to eight measurements per egg, the real
figure is probably larger. These high CVs indicate that
a large variation in weight loss can be tolerated and is
presumably advantageous in a variable environment.
At the same time it has been shown in at least five
species (Wangensteen et al. 1974; Carey et al. 1977;
Packard et al. 1977; Sotherland et al. 1980) that natu-
ral selection can act to induce changes in water vapor
conductance of eggshells of high altitude populations,
so that the average weight loss/fresh egg weight ratio
is similar to that of their sea level counterparts.
Whether such adapted populations show reduced
variability or skewed distributions does not appear to
have been investigated.

Relationship between Weight Loss,
Fresh Egg Weight, and Length of Incubation

The regression coefficient for the exponential curve
of daily weight loss (L) on fresh egg weight (W) for the
11 species recorded in Table 2 is 0.706 + 0.0934
(r= 0.930). This does not differ significantly
(P> 0.05) from 0.742, the figure given by Drent
(1970) for 46 species. The regression equation for our
combined data, using means when the data for a spe-
cies are duplicated, is L = 0.0153 W973 =°55(n = 56;
r = 0.968). The reduced major axis (RMA) equation is
L=0.0143w°”™’*°™* The standard errors of the
exponents in these two equations are much smaller
than the figure (0.22) given by Drent (1970). The latter
was calculated according to one of the formulas of
Feldstein and Hersh (1935), evidently Formula 6,
which omits (n — 2)” from the denominator. The cor-
rect standard erroris+0.0293 and therefore, contrary
to what Drent supposed, the exponent in his equation
is, as in the two equations above, significantly differ-
ent (P< 0.02) from 0.667, the figure expected if the
rate of weight loss varied according to egg surface
area. More recently Rahn and Ar (1974) have shown
that the daily weight loss varies not only with fresh egg
weight but also with length of incubation to make the
total weight lost during incubation a relatively con-
stant fraction of fresh egg weight. The unweighted
mean of the percentages of the fresh egg weights lost
during incubation for the I! species in Table 2 is
14.1 + 0.64. This figure is similar to 14.3 + 0.52, the
mean for seven species of tern (Rahnet al. 1976), 15.6
for two species of gull (Morgan et al. 1978), and 15.2

MANNING: WEIGHT LOST BY EGGS DURING INCUBATION 67

for the Laysan Albatross, Diomedea immutabilis
(Fisher 1969), but well below the 18% estimated by
Rahn and Ar (1974) from their Equation 5 as an
average for all eggs irrespective of weight and length
of incubation period. Their equation was, however,
based on two regressions, and, as all the variables were
subject to error, RMAs give a more exact relationship
(Kermack and Haldane 1950; Ricker 1973). When
RMA values are substituted for those of regression,
the equation becomes

0.75. 0.25
Ff - 0.014W 11W =015
w !.00

where W is the fresh egg weight and F the fraction of
that weight lost during incubation. The exponents in
the numerator of the equation now add up to | instead
of 0.96 in the equation based on regressions, the Ws
cancel out exactly instead of approximately and F is
reduced from 0.18 to 0.15 or 15%. Rahn et al. (1979)
obtained the same figure by the regression of total
weight loss during incubation on fresh egg weight,
using eggs of 65 species and a wide range of egg size
and incubation length. Though they give neither the
regression equation nor the correlation coefficient, it
is clear from their plot that the latter is very high and
the difference between regression and RMA therefore
negligible. The RMA exponential equation for the 11
species listed in Table 2 is L: J= 0.153W°95*0.052
(r = 0.987), where L is the length of incubation and /
the daily weight loss. As the exponent does not differ
significantly from 1, the weight loss during incubation
is again 15% of fresh egg weight and the RMA a
Straight line on natural scale, the equation being
L-T= - 0.116 + 0.152 + 0.0073 W (r = 0.990). The dif-
ference between the RMA coefficient and the mean
(0.141 = 0.0064) of the ratios, given as percentages in
Table 2, is not significant in this small sample.

It should be noted that the term ‘total weight loss
during incubation’ has been used but no allowance
made for the extra rate of weight loss after pipping. As
the crucial factor is probably the size of the airspace in
the egg just before pipping (Rahn and Ar 1974), it
would be better to consider only the period up to that
point, but its length is not available for most species.

Acknowledgments

I am grateful to the Canadian Wildlife Service and
particularly to S. G. Curtis and R. I. G. Morrison for
the opportunity to work at North Point in 1975. I was
ably assisted at North Point by Mitchel Thomlison,
who found many of the nests and recorded all my
measurements. Pamela M. Morse, though in no way
responsible for the results, gave valuable statistical
advice, and Henri Ouellet and three reviewers made
helpful comments on the first draft of the manuscript.

68 THE CANADIAN FIELD-NATURALIST

Facilities for incubation of the hens’ eggs were pro-
vided at the Animal Diseases Research Institute,
Ottawa, through the courtesy of K. J. Betteridge.

Literature Cited

Ar, A., C. V. Paganelli, R. B. Reeves, D. G. Green, and H.
Rahn. 1974. The avian egg: water vapor conductance,
shell thickness, and functional pore area. Condor 76:
153-158.

Barth, E. K. 1952. Incubation period and loss of weight of
eggs of the Common Gull, Larus canus canus, and the
Lesser Black-backed Gull, L. fuscus intermedius. Riista-
tieteellisia Julkaisuja — Papers on Game Research 8:
111-121.

Carey, C., S. Garber, and E. Thompson. 1977. Modifica-
tion of pore area of avian eggs at altitude. American
Zoologist 17: 930.

Drent, R. H. 1970. Functional aspects of incubation in the
Herring Gull. Behaviour, Supplement 17: 1-132.

Feldstein, M. J..and A. H. Hersh. 1935. The calculation of

relative growth constants. American Naturalist 69:
610-613.

Fisher, H.I. 1969. Eggs and egg-laying in the Laysan Alba-
tross, Diomedea immutabilis. Condor 71: 102-112.

Hays, H.,and M. LeCroy. 1971. Field criteria for determin-
ing incubation stage in eggs of the Common Tern. Wilson
Bulletin 83: 425-429.

Kendeigh, S. C. 1940. Factors affecting length of incuba-
tion. Auk 57: 499-513.

Kermack, K.A., and J. B.S. Haldane. 1950. Organic
correlation and allometry. Biometrika 37: 30-41.

Landauer, W. 1967. The hatchability of chicken eggs as
influenced by environment and heredity. University of
Connecticut, Monograph | (Revised). 315 pp.

Manning, T. H. 1979. Density and volume corrections of
eggs of seven passerine birds. Auk 96: 207-211.

Morgan, K.R., C. V. Paganelli, and H. Rahn. 1978. Egg
weight loss and nest humidity during incubation in two

Vol. 95

Alaskan gulls. Condor 80: 272-275.

Packard, G.C., P. R. Sotherland, and M. J. Packard.
1977. Adaptive reduction in permeability of avian egg-
shells to water vapour at high altitudes. Nature (London)
266: 255-256.

Rahn, H., and A. Ar. 1974. The avian egg: incubation time
and water loss. Condor 76: 147-152.

Rahn, H., A. Ar, and C. V. Paganelli. 1979. How bird eggs
breathe. Scientific American 249(2): 46-55.

Rahn, H., C. V. Paganelli, I. C. T. Nisbet, and G. C. Whit-
tow. 1976. Regulation of incubation waterloss in eggs of
seven species of terns. Physiological Zoology 49: 245-259.

Ricker, W. E. 1973. Linear regressions in fishery research.
Journal of the Fishery Research Board of Canada 30:
409-434.

Schonwetter, M. 1971-1979. Handbuch der Oologie.
Edited by W. Meise. Akademie-Verlag, Berlin. Volumes |
and 2.

Sokal, R.R., and F.J. Rohlf. 1969. Biometry. W. H.
Freeman and Company, San Francisco. 776 pp.

Sotherland, P.R., G.C. Packard, and T.L. Taigen.
1979. Permeability of magpie and blackbird eggshells to
water vapor: variation among and within nests of a single
population. Auk 96: 192-195.

Sotherland, P.R., G.C. Packard, T. L. Taigen, and J.
Boardman. 1980. An altitudinal cline in conductance of
Cliff Swallow (Petrochelidon pyrrhonota) eggs to water
vapor. Auk 97: 177-185.

Swiger, L. A., W. R. Harvey, D. O. Everson, and K. E. Gre-
gory. 1964. The variance of interclass correlation involv-
ing groups with one observation. Biometrics 20: 818-826.

Wangensteen, O. D., H. Rahn, R. R. Burton, and A. H.
Smith. 1974. Respiratory gas exchange of high altitude
adapted chick embryos. Respiration Physiology 21:
61-70.

Received 3 May 1977
Accepted 5 June 1980

Sex Differential in Use of the Physical Environment
by Bighorn Sheep (Ovis canadensis)

LUIGI E. MORGANTINI and ROBERT J. HUDSON
Department of Animal Science, University of Alberta, Edmonton, Alberta T6G 2E3

Morgantini, Luigi E., and Robert J. Hudson. 1981. Sex differential in use of the physical environment by Bighorn Sheep
(Ovis canadensis). Canadian Field-Naturalist 95(1): 69-74.

The use of the physical environment by Bighorn Sheep (Ovis canadensis) rams and ewes on a common winter range in western
Alberta showed sex differential in habitat selection and spatial distribution. In winter ram groups occurred on more steep and
rocky ground and were found closer to forests than ewe groups. In spring habitat selection of rams did not change, whereas
ewes shifted to lower elevations and onto more steepand rocky habitats. During this season segregation between the sexes was
ensured through the use of two distinct sections of the available range. These results, when compared with other studies, do
not support previous hypotheses explaining sex segregation in Bighorn Sheep. A new hypothesis, that spatial separation
maximizes individual fitness by reducing energy-expensive sexual-agonistic behavior in rams at times when reproduction is
not possible, is proposed.

Key Words: Bighorn Sheep, Ovis canadensis, habitat selection, sex segregation.

Habitat and spatial separation in winter between _ relatively mild climate. As a result of the effect of the
Bighorn Sheep (Ovis canadensis) rams and ewes on — surrounding mountains on airflows, cloud cover does
the Palliser range in Banff National Park has been _ not develop to the extent that it does over the sur-
studied extensively by several authors (Geist and rounding areas, and western winds keep the grassland
Petocz 1977; Shank 1979). Although the significance and openslopes largely snow-free (Morgantini 1979).
of such sex segregation is not clear, three hypotheses The vegetational characteristics of the mountain
have been proposed. Geist and Petocz (1977) have slopes have been described by Gates (1975). At high
interpreted habitat separation betweenramsandewes elevation (above 2074 m), Hairy Wild Rye (Elymus
as a behavioral pattern evolved to minimize intraspe- innovatus) is the predominant grass species. Sedges
cific competiton. Bromely (1977) and Geist and make up 40% of the floristic composition. At lower
Bromely (1978) have explained segregation of males elevation, Northern Wheat Grass (Agropyron dasys-
from females in winter as an anti-predator strategy. tachyum), Purple Reed Grass (Calamagrostis pur-
Recently, Shank (1979) hypothesized that ecological purascens), and June Grass (Koeleria cristata) are the
separation between the sexes in Bighorn Sheepsimply dominant species in descending order of frequency of
reflects different body sizes and, therefore, different occurrence. Forests of Trembling Aspen (Populus
habitat requirements (Shank 1979). tremuloides), Lodgepole Pine (Pinus contorta), and

This study was undertaken to determine whether spruce (Picea glauca and Picea engelmannii) also are
habitat and spatial separation betweenramsandewes found (Morgantini 1979).
occurred in another geographical region, the Ya Ha The study area is an important Bighorn Sheep win-
Tinda Ranch area, where the two sexes share acom- ___ ter-spring range. At the time of this study it supported
mon winter range. In the event that sexual segregation a population of some 80 individuals.

did occur, the second objective of the study was to
assess whether the results supported any of the preced- Methods

ing hypotheses. From December 1975 to May 1976 data were col-
lected by direct observation of activity, movements,
Study Area and behavior of Bighorn Sheep, conducted from van-

The study area is located in western Alberta inthe tage points situated on the valley floor. Once a group
foothills of the Rocky Mountains, about 10 kmeastof of animals was sighted, time, number of individuals,
the eastern boundaries of Banff National Park sex, and, if possible, age were recorded. Each location
(51°45’-46’N, 115°33’-35’W). It consists of south- was marked on a low oblique aerial photograph to
facing, largely open slopes situated northofthe YaHa within 5 mso that it could be easily located for subse-
Tinda Ranch valley, one of the most important Elk quent description of its physiographic characteristics.
(Cervus canadensis nelsoni) winter ranges in the pro- Emphasis then was placed on behavioral observations
vince. Elevation varies from 1550 m on the valley foraslongas the animals could be observed. In several
floor to 2400 m on higher slopes. The area enjoys a__ instances movements of the same herd, identified by

69

70 THE CANADIAN FIELD-NATURALIST

[-] RAMS (50) n.s.
(5 80
=
ae
E 60
O
n 40
LL
O 20
&

1500: 1701- >1901

1700 1900

ELEVATION (m)

yA [-] RAMS (50)
Bins Fl EWES (15)
= P < 0.0001
F 60
o)
D 40
LL
O 20
&

OE eH ea” S66

PERCENT OF SLOPE

% OF SIGHTINGS

% OF SIGHTINGS

Vol. 95

[-] RAMS (50)
EWES (15)

80
60 P< 0.005
40
20
O
O 1 2 3 4
DEGREE OF ROCKINESS
([] RAMS (48)
80 EWES (15)
P < 0.0001
60
40
20

9-79 76-150

151-225
DISTANCE FROM FOREST COVER (m)

FiGURE!. Comparison of distribution of Bighorn Sheep ram and ewe groups in winter as measured by elevation, percentage
of slope, degree of rockiness, and distance from forest cover (total number of group sightings in parentheses).

its structure and by the characteristics of individuals,
could be recorded for several consecutive days. Each
month from 15 to 20 d were spent in the field.

To detect seasonal changes in distribution, we sub-
divided observations into two groups: winter observa-
tions, from December through 15 March, and spring
observations, from 16 March through May. The
beginning of the spring season was placed at the time
when the first new plant growth was observed.

Seven abiotic environmental parameters were con-
sidered as potential determinants of animal distribu-
tion on open ranges. They were slope (percent), aspect
(degrees), elevation (metres), rockiness (scores 0-4),
distance to forest cover (metres), snow depth (cen-
timetres), and snow cover (percent). With the excep-
tion of distance to forest cover estimated from aerial
photographs, environmental variables were measured
directly in the field. At the end of the study, aspect was
discarded because it was found to change little within
the study area.

To assess sex differences in spatial distribution, the
study area was arbitrarily subdivided in three sections,

using the North-South lines of the Universal Trans-
verse Mercator (UTM) grid system.

During the study period, 151 group sightings were
made. These comprised a total of 2035 animal obser-
vations. Group sightings were placed in three catego-
ries termed “rams,” when the group was composed
only of adult or subadult males; “ewes,” when the
group was composed of adult females and subadults,
either females or males; “mixed,” whenever mature
males, class II rams (Geist 1971), were present in a
“ewes” group. The response of animals to physiogra-
phic factors was assessed on the basis of percentage of
group sightings. Seasonal changes in occupational
patterns were determined by comparing frequency
distributions using chi-square contingency table
analyses.

Results

In winter, both ram and ewe groups were found
mostly on high elevation ranges. However, ewes were
generally observed farther from forests and on less
steep and rocky ground (Figure 1).

1981

RAMS (36)
6 sol & P < 0.0001
z
=
= 60
oO
wm 40
LL
O 20
= ase as

) peo ee
150027 Olle 11901
1700 1900
ELEVATION (m)
es RAMS (36)
O 80 z n.s.
2
F 60 "
S) Be
M 40 ee
Le oes
O 20 E a
PS a ee %
0 es He Bd ee
0-15 16-28 29-50 >50

PERCENT OF SLOPE

% OF SIGHTINGS

% OF SIGHTINGS

MORGANTINI AND HUDSON: BIGHORN SHEEP HABITAT USE val

(RAMS (36)
80 EWES (33) ALS.

DEGREE OF ROCKINESS

[J RAMS (82)
EWES (25)

n.s.

. : a
Sai 76-1150 \1|51-225
DISTANCE FROM FOREST COVER (m)

FIGURE2. Comparison of distribution of Bighorn Sheep ram and ewe groups in spring as measured by elevation, percentage
of slope, degree of rockiness, and distance from forest cover (total number of group sightings in parentheses).

Snow cover and depth were not found to be signifi-
cant in determining daily or monthly distribution on
Open ranges, possibly because of mild winter condi-
tions. In several instances the same herds could be still
found the day after snow storms grazing on the same
sites on medium and high elevation ranges.

In spring, the distribution of rams did not change
(Figure 2), while ewes shifted to lower elevations
(P< 0.001) and onto more rocky (P< 0.0002) and
steep (P< 0.006) ground. In spring, ewes were also
observed closer to forest cover.

With regard to spatial distribution, ram groups
were generally found on the east section of the range
(Figure 3). This section is separated from the central
section by a timbered draw about 250 m wide, and,
compared to the other sections, is generally more
rocky and steep, with cliffs and talus slopes
interspersed with patches of vegetation. Selection of
this area by rams was particularly evident during the
Spring months and coincided with a shift by ewes to
lower elevations onto the main, central range.

During the study period, we observed 6 mixed
groups in winter and I] in the spring. They were
generally found at lower elevations than either ram or

ewe groups. In their response to rockiness, slope, and
distance from forests, they appeared intermediate to
rams and ewes. No seasonal change could be detected
in the distribution of mixed groups. These compari-
sons may not be valid, however, because of the small
sample size.

Discussion

A difference in distribution between ram and ewe
groups was found. In winter, habitat separation
occurred on the basis of rockiness, slope, and distance
from forests. In spring, elevation was the main factor.
The seasonal difference is attributable to the shift of
ewes in spring to lower elevation and onto more rocky
and steep ground. This shift may possibly indicate a
more intense search for high-quality forage caused by
greater nutrient requirements during pregnancy. For
instance, heat retention by rocks during sunny days in
late winter and spring may have produced warmer
microclimatic conditions favorable to earlier forage
growth and higher nutrient content as compared to
adjacent grassy areas. Similarly, the degree of slope
may affect forage quality by influencing the amount of
solar radiation reaching the ground. Although the

THE CANADIAN FIELD-NATURALIST Vol.

WINTER

oO Rams
@ Ewes-Juveniles
O Mixed An

0.250 0.500 km

Scale

CENTRAL SECTION EASTERN SECTION

SPRING

o Rams

@ Ewes-Juveniles

O Mixed By
0.250 0.500 km

SS SJ
Scale

WESTERN SECTION CENTRAL SECTION EASTERN SECTION

FIGURE 3. Locations of Bighorn Sheep ram, ewe, and mixed groups in winter and spring.

1981

preceding hypotheses are speculative, they are sup-
ported by observations of highly selective foraging
behavior in Bighorn Sheep (Hebert 1973; Shannon et
al. 1975). Alternatively, spring occupation by ewes of
more steep and rocky ground could be interpreted as
an anti-predator strategy at a time when animals were
in poor physical condition (Shank 1979). It is worth
noting that the only case when ewes were observed on
a steep cliff in winter was in response to an unsuccess-
ful hunt by a Cougar (Felis concolor).

The behavior of Bighorn Sheep on the Ya Ha Tinda
range suggests that both habitat and spatial factors
contributed to the separation of the sexes. In winter,
when ram and ewe groups were not spatially separ-
ated, 1.e. geographically separated, they used different
habitats. In spring, when such differences were limited
to the elevational gradient, the two sexes were spa-
tially separated and restricted to two distinct sections
of their range.

This study in itself does not lend much insight to the
significance of sex segregation in Bighorn Sheep.
When the results are compared with other studies,
however, an assessment of previous hypotheses
explaining this phenomenon may be made.

Essentially three hypotheses can be distinguished.
In the first hypothesis, ecological separation between
the sexes is thought to reflect differences in body size
and, therefore, different behavioral patterns and habi-
tat requirements. The hypothesis has been extensively
developed by Shank (1979); however, if differences in
habitat selection between rams and ewes simply
reflected their differences in body sizes, we would
expect to find, for the same sex, similar habitat
requirements or, at least, similar adaptive strategies,
in diverse environments. This is not indicated in our
study. We found that in winter, rams selected rockier
and steeper ground than ewes. A different behavior
was recorded by Geist and Petocz (1977) on the Pal-
liser range in Banff National Park. They reported a
preference by rams for grassy slopes and by ewes for
rocky habitats. This difference between the sexes in
habitat preferences was partially explained on the
basis of snow accumulation patterns on slopes. Rams,
being larger and heavier than ewes, could still graze on
snow-covered slopes, whereas ewes had to move to
relatively snow-free cliffs. Lack of deep snow and mild
climatic conditions during our study may account for
the occurrence of ewes on grassy slopes in winter.
However, Geist and Petocz’s explanation suggests that
rams, which in one environment brave deep snow to
graze on grassy slopes, will remain on snow-free
slopes in other ranges. As shown, this does not occur
on the Ya Ha Tinda range. Further, according to
Shank (1979), the utilization by ewes of cliffs and
rocky habitats reflects a higher “. . . susceptibility to

MORGANTINI AND HUDSON: BIGHORN SHEEP HABITAT USE 73

predation as determined by body size” (p.127). This
implies a sex-specific habitat selection strategy that
should maximize security. Yet, the winter distribution
of ewes in our study area does not indicate any distinct
anti-predator strategy. In conclusion, it appears that,
on both the Palliser and the Ya Ha Tinda range, the
two sexes simply tend to separate and that such a
separation takes a different form depending on the
environmental conditions of their common winter
range. This intersexual avoidance is further indicated
in our study by the decreased number of sightings of
rams on the main, central range in spring, at the time
when ewes had begun to use it (Figure 3).

The second hypothesis suggests that the segregation
of males from females has evolved as anti-predator
strategy. Males, exhausted by the breeding season and
easily distinguishable from females, would be sub-
jected to selective predation during winters, unless
they avoid females or shed their sign of maleness as do
Pronghorns (Antilocapra americana) (Geist and
Bromely 1978). On the Ya Ha Tinda range we
observed rams and ewes restricted toa relatively small
winter range with rams strongly associated with a
particular section. Sucha preference by rams has been
observed for three subsequent seasons (1976-1979)
(L. E. Morgantini, personal observation). If predator
avoidance had shaped the sex-segregation behavior of
Bighorn Sheep, we would not expect rams to restrict
activity to specific limited areas where they could be
easily located by predators, unless those areas pro-
vided high-quality escape terrain. But this was not
indicated in our study. Furthermore, even though the
ram section was somewhat more rocky and steep,
rams were generally found closer to forests than ewes
(Figures | and 2) and always reacted to approaching
humans or predators, regardless of their direction, by
escaping west, throughout the timbered draw, across
the main open slope to the higher elevation ranges.

The third hypothesis, proposed by Geist and Petocz
(1977), suggests that males avoid females in winter to
minimize intraspecific competition. That is, to max-
imize reproductive fitness, “. .. males ought not to
compete with their own offspring during ontogeny,
nor with the prospective mothers of their offspring”
(p. 1803). Basically this hypothesis is an extension of a
hypothesis first proposed by Rand (1952), who noted
that ecological segregation between the sexes, differ-
ent food habits, and sexual dimorphism could be seen
as adaptations to reduce intraspecific competiton.
Our results, however, are not consistent also with this
hypothesis. With regard to competition by exploita-
tion, our data show that in spring ewes selected habi-
tats and areas that had been previously used by rams.
Although Shank (1979) reported differences in food
habits between the sexes, such differences were attrib-

74 THE CANADIAN FIELD-NATURALIST

uted by the author to the use of different areas.
Further, diet quality, as assessed by fecal protein
analysis, differed only ‘marginally’ (Shank 1979). It is
therefore difficult to see how, in our study area, pre-
vious use of the ranges by rams could be advantageous
to breeding females. Yet, because the sexes are separ-
ated, minimization of intraspecific competition by
interference, that is, low harassment of ewes by rams,
may be occurring. However, in addition to the theo-
retical problems reviewed by Shank (1979), the
hypothesis of competition minimization presented by
Geist and Petocz (1977) implies the presence of sex-
linked (Whitney 1976; Wickler 1977) altruistic behav-
ior (Hamilton 1964). To not “compete” with ewes,
rams would risk reducing their own reproductive
potential by using alternative, possibly suboptimal
habitats.

On the basis of our results we think that maximiza-
tion of individual fitness by rams is all that is required
to explain the evolution of sex segregation in Bighorn
Sheep and that there is no need to invoke, directly or
indirectly, altruistic behavior or kin selection. In fact,
it can be expected that the occurrence of rams with
ewes in mixed herds after the end of the breeding
season would cause a higher frequency and intensity
of sexual and agonistic behavior among rams, and
consequently higher energy expenditures at times
when reproduction is not possible. Such higher energy
expenditures could be particularly detrimental during
harsh winters. It is therefore possible that, to minimize
energy expenditures associated with sexually related
social interactions, the coexistence of males with
females in periods other than the breeding season was
selected against, and that it is avoided through habitat
and spatial separation. In this context, different phy-
siological and/or behavioral requirements are
expected to shape habitat preferences that adjust to
local environmental conditions. This hypothesis is
consistent with our data and with the present knowl-
edge of social behavior of Bighorn Sheep (Geist 1971).
It accounts for the fact that rams leave rutting ranges
first; that older rams, for which sexual agonistic
behavior is more energy demanding, stay on rutting
ranges for shorter periods of time than younger ones;
and that generally the occurrence of rams in ewe
groups is inversely correlated with their age (Geist
1971; Geist and Petocz 1977). In our hypothesis, min-
imization of intraspecific competition, predators
avoidance, or sexual dimorphism-related niche differ-
ences could be all seen as by-products of the avoidance
by rams of sexual-agonistic stimuli outside the rutting
season.

Vol. 95

Acknowledgments

We acknowledge the cooperation of the Alberta
Fish and Wildlife Division for financial and logistic
support, the Department of Zoology of the University
of Alberta for equipment, and Parks Canada for per-
mission to conduct the study on the Ya Ha Tinda
Ranch. We are also grateful to Raffaella Morgantini
for many helpful discussions and her assistance in the
field, and to the staff of the Ya Ha Tinda Ranch (Slim
Haugen, Earl Hays, and Gordon Patterson) for their
cooperation throughout the field work.

Literature Cited

Bromely, P. T. 1977. Aspects of the behavioural ecology
and sociobiology of the Pronghorn. Ph.D. thesis, Univer-
sity of Calgary, Calgary. 170 pp.

Gates, C. C. 1975. Aspects of the environment —- lungworm
(Nematoda: Metastrongyloidea) — Bighorn Sheep (Ovisc.
canadensis) system. M.Sc. thesis, University of Alberta,
Edmonton. 55 pp.

Geist, V. 1971. Mountain Sheep: a study of behavior and
evolution. University of Chicago Press, Chicago. 383 pp.

Geist, V.,and P. T. Bromely. 1978. Why deer shed antlers.
Zeitschrift fur Saugetierkunde 43: 223-231.

Geist, V., and R. G. Petocz. 1977. Bighorn sheep in winter:
do rams maximize reproductive fitness by spatial and
habitat segregation from ewes? Canadian Journal of
Zoology 55: 1802-1810.

Hamilton, W.D. 1964. The genetic evolution of social
behavior. Journal of Theoretical Biology 7: I-51.

Hebert, D. M. 1973. Altitudinal migrations as a factor in
the nutrition of Bighorn Sheep. Ph.D. thesis, University of
British Columbia, Vancouver. 375 pp.

Morgantini, L. E. 1979. Habitat selection and resource div-
ision among Bighorn Sheep, Elk and Mule Deer in western
Alberta. M.Sc. thesis, University of Alberta, Edmonton.
187 pp.

Rand, A. L. 1952. Secondary sexual characters and ecolog-
ical competition. Fieldiana Zoologica 34: 65-70.

Shannon, N. H.,R. J. Hudson, V. C. Brink, and W. D. Kitts.
1975. Determinants of spatial distribution of Rocky
Mountain Bighorn Sheep. Journal of Wildlife Manage-
ment 39: 387-401.

Shank, C. C. 1979. Sexual dimorphism in the ecological
niche of wintering Rocky Mountain Bighorn Sheep.
Ph.D. thesis, University of Calgary, Calgary. 193 pp.

Whitney, G. 1976. Genetic substrates for the initial evolu-
tion of human sociality. I. Sex chromosome mechanisms.
American Naturalist 110: 867-975.

Wickler, W. 1977. Sex-linked altruism. Zeitschrift fur
Tierpsychologie 43: 106-107.

Received 25 February 1980
Accepted 12 August 1980

Terrestrial Molluscs of Central Alberta

JAMES VAN ESand D. A. BOAG

Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9

Van Es, James, and D. A. Boag. 1981. Terrestrial molluscs of central Alberta. Canadian Field-Naturalist 95(1): 75-79.

The litter from forest habitats dominated by Populus spp. was sampled within a 100-km radius of Edmonton, Alberta, and
searched for the presence of terrestrial molluscs. Eighteen species belonging to eight families in two orders were identified.
Thirteen of these species had not been reported previously from this area. Information on distribution and abundance of these

molluscs is included.

Key Words: terrestrial molluscs, distribution, size, abundance, central Alberta, Populus, habitat.

The terrestrial molluscan fauna of western Canada
is poorly known. Mozley (1934) reported on this
group of invertebrates in Saskatchewan and Alberta,
listing the six families and 11 species known at that
time. For each species (not all were identified beyond
genus), he gave only the name of the town nearest the
collection site, although in an earlier publication
(Mozley 1930) he described the characteristics of some
of the species (those found in Jasper National Park) in
more detail. To our knowledge, these two papers,
which refer to even earlier investigations (Berry 1922;
Mozley 1933), remain the only publications that
attempt to deal with the terrestrial molluscs of Alberta
as a whole. More recently, Russell (1951) described
this fauna in the Cypress Hills, and Platt (1980) and
Boag (unpublished data) have added to the species list
for the east slopes of the Rocky Mountains in Alberta.

As a first step in an attempt to record the distribu-
tion and abundance of terrestrial molluscs in rem-
nants of the native vegetation of Alberta, we report
here on this fauna in the vicinity of Edmonton,
Alberta (Figure 1).

Methods

Forest habitats within the area shown in Figure |
were sampled (Appendix 1) in April through June and
in October 1979. At each of 31 sites (Figure 1), we took
two 0.25-m? samples of forest litter, usually about
10 m apart and representative of the site in general.
The depth of litter taken varied from 5 to 25cm
depending on the state of decay of the most recent leaf
fall. Each sample was bagged, labeled, and, on return
to the laboratory, dried at room temperature before
being sieved and searched for the shells of snails. All
specimens found were removed and placed in labeled
vials. Subsequently, the snail shells were separated (on
the basis of shell morphology) into the various species
present, tentatively identified, and stored in separate
vials. From each site sampled we randomly selected
up to 25 undamaged and non-eroded specimens of

75

each species for measuring. We used an ocular
micrometer in a dissecting microscope to take the
measurements.

In addition 19 forest sites were sampled by search-
ing the litter and overturning the debris on the forest
floor (Figure 1, Appendix 1). Specimens handpicked
by this method were treated in a similar manner to
those taken from the dried litter samples.

Specimens of each tentatively identified species
were sent to the National Museum of Natural Scien-
ces, Ottawa, where identifications were confirmed or
corrected, and where the samples of these species are
now housed.

Results and Discussion

At 85% (n = 50) of the sites sampled, the dominant
trees were Trembling Aspen (Populus tremuloides)
and/or Balsam Poplar (P. balsamifera), whereas at
the remaining 15% of sites, the forests were mixed
with Populus spp. interspersed among either conifers
(White Spruce, Picea glauca and/or Jack Pine, Pinus
banksiana) and/or Paper Birch (Betula papyrifera).

The two methods of sampling produced different
results (Table 1). Handpicked samples produced sig-
nificantly fewer individuals per sample site (post hoc
procedure using pair-wise contrast, P< 0.05). Furth-
ermore, the specimens recovered were biased signifi-
cantly in favor of the largest and against the smallest
species (chi-square test, P< 0.05). Consequently we
did not use the 19 handpicked samples in any of the
quantitative analyses. Handpicked samples, however,
were useful in determining the presence of slugs, as
they were not recovered from the litter samples, hav-
ing been lost during the drying process.

In the 31 samples of forest litter, we recorded terres-
trial molluscs from two orders (Stylommatophora
and Basommatophora) and seven families (Zonitidae,
Endodontidae, Succineidae, Pupillidae, Valloniidae,
Cionellidae, and Carychiidae). Additionally, we
found one species of slug in an eighth family (Limaci-

76 THE CANADIAN FIELD-NATURALIST

eG

@ BaRRHEAD

Vol. 95

@ weESTLOCK

eR 53

©21 @ WETASKIWIN

114°

FIGURE |. Locations where litter samples (numbers) and handpicked samples (letters) were taken in central Alberta. See

Appendix | for descriptions of exact locations.

dae) among the handpicked samples. Inall, 18 species
have been identified (Table 2).

Three zonitids (Euconulus fulvus, Retinella elec-
trina, Vitrina alaskana), one endodontid (Discus
cronkhitei), one pupillid (Vertigo gouldi), and the

TABLE |—Relative numbers and sizes of terrestrial molluscs
recovered in 19 handpicked and 3! litter samples

Mean number of individuals

Size of molluscs Handpicked Litter
(no. dimensions >2 mm) samples samples
0 1.26 82.83

| 20.42 106.68

2 12.16 26.96

only cionellid (Cionella lubrica) were widely distrib-
uted throughout the area and in the various forest
types. Each species was present in at least 90% of the
samples (Table 2). Discus cronkhitei and Vertigo
gouldi were also relatively abundant at each site, aver-
aging about 50 specimens/0.5 m? sample of forest
litter (Table 2). Vitrina alaskana and Cionella lubrica,
although widespread, were found at much lower den-
sities, On average about 12 specimens/0.5 m2 (Table
2). These differences may represent absolute differen-
ces in abundance or differences in rates of disappear-
ance of the shells of the respective species. In most
samples, some shells were eroded and often on the
point of disintegration, the snails presumably having
been dead for some time. Thus abundance figures
include some shells of snails that may have been dead

1981

VAN ES AND BOAG: TERRESTRIAL MOLLUSCS, ALBERTA

UY

TABLE 2— Terrestrial molluscs recovered from 31 littersamples collected in central Alberta forests dominated by Populus spp.
Species marked with an asterisk were previously recorded in this area by Mozley (1934)

Measurements’
Saniple Height (mm)
Taxa size x (95% conf. inter)
Stylommatophora
Zonitidae
* Euconulus fulvus 256 1.95 (1.90-2.00)
* Retinella electrina 71 1.74 (1.65-1.83)
*Zonitoides
arboreus 38 2.18 (2.07-2.29)
Vitrina alaskana 152 2.38 (2.29-2.46)
Limacidae
Deroceras laeve — — —
Endodontidae
* Discus cronkhitei 352 1.90 (1.84-1.95)
Punctum
minutissimum 170 0.68 (0.67-0.69)
Succineidea
Oxyloma sp. 8.83 (6.23-11.42)
Succinea sp. 11 8.96 (5.75-12.18)
Pupillidae
Gastrocopta
tappaniana 21 1.91 (1.81-2.01)
Vertigo gouldi 337 1.73 (1.71-1.75)
Vertigo modesta 3 2.40 (1.97-2.83)
Columella simplex 42 1.80 (1.73-1.87)
Valloniidae
Vallonia
gracilicosta 155 1.22 (1.21-1.23)
Vallonia sp. 85 0.86 (0.82-0.89)
Zoogenetes harpa 24 3.02 (2.91-3.14)
Cionellidae
*Cionella lubrica 116 5.21 (5.13-5.29)
Basommatophora
Carychiidae
Carychium exile 2 2.0 _

“Only undamaged and non-eroded specimens were measured.

Number of snails

Pre quency, per sample (0.5 m2)

Width (mm) of occurrence

b

x (95% conf. inter) (% of sites) x; (range) n
2.67 (2.62-2.72) 97 27.3 (1-167) 30
3.35 (3.20-3.50) 90 22.9 (1-77) 28
4.27 (4.02-4.51) 42 3.1 > (Ee?) 13
4.35 (4.19-4.50) 90 11.6 (1-47) 28
3.48 (3.39-3.58) 94 48.9 (1-266) 29
1.16 (1.14-1.17) 71 22.6 (1-169) 22
4.95 (3.35-6.55) 3 4.0 — l
5.12 (3.90-6.33) 29 2.6 (1-10) 9
1.20 (1.19-1.21) 10 6.7 (1-18) 3
1.01 (1.00-1.02) 90 60.0 (1-497) 28
1.37 (1.22-1.51) 3 3.0 — l
1.21 (1.20-1.23) 19 7.3 (2-14) 6
2.56 (2.54-2.58) 42 33.1 (2-137) 13
1.73 (1.66-1.80) 13 74.5 (13-195) 4
2.49 (2.44-2.54) 10 20.3 (1-57) 3
2.41 (2.39-2.44) 94 13.7 (1-79) 29
1.0 — 3 2.0 — l

b F 0 :
Mean and range are based only on samples in which at least one specimen was found.
“Not recovered from litter samples but only in handpicked samples (n = 19).

for a variable time period. For species like Vitrina
alaskana, the shell of which 1s very fragile, disintegra-
tion time may be much shorter than for other species
with stouter shells. This may explain the lower abun-
dance of Vitrina alaskana despite its high frequency of
occurrence in the samples. The relative acidity of the
soil may also be an important factor influencing the
rate of disintegration of shells.

Of the remaining species, three were reasonably
common. Zonitoides arboreus was found in 42% of
the litter samples but was never abundant, averaging
3.1 specimens per sample (Table 2). By contrast,
Punctum minutissimum and Vallonia gracilicosta
were both reasonably common and abundant (Table
2). The remaining nine species were neither common

nor abundant, with the exception of Va/lonia sp.
which appeared in only 13% of the samples but at an
average of 74.5 individuals per sample (Table 2). The
distribution of Vallonia sp. 1s interesting since it was
found only in samples taken northeast of Edmonton.
Thus, the frequency of occurrence within the limits of
the sampling area may reflect only where the samples
were taken in relation to the species range rather than
a measure of ubiquity within its normal range. This
problem can be addressed only by more intensive
sampling to determine the actual range of the species.

The species diversity at given sites varied consider-
ably (5-14 per sample). In general the abundance of
individuals was positively correlated with number of
species present (5 species and 14 individuals to 14

78 THE CANADIAN FIELD-NATURALIST

200

100

50}

NUMBER OF Discus cronkhitei PER SAMPLE

150

50 100

200

Vol. 95

250 300

NUMBER OF Vertigo gouldi PER SAMPLE

FIGURE 2. The relationship between the numbers of Vertigo gouldi and Discus cronkhitei in litter samples taken from forest

habitats in central Alberta.

species and 291 individuals per sample). This relation-
ship would seem to reflect relative richness of the
habitat rather than interactions among the species. A
comparison of the numbers of the two most abundant
species (Discus cronkhitei and Vertigo gouldi) at each
site, however, suggests some form of interaction that
leaves one species dominant numerically at one site
and the other at a second site (Figure 2). Where the
abundance of each species was less than 50/0.5 m2,
there was no apparent difference in species abundan-
ces. When the number of individuals of each species
exceeded 50, however, it appeared that either one of
the two was relatively much more abundant, possibly
at the expense of the other. If this is so, it may reflect
some form of competitive advantage of one or other of
the species in a given habitat. In this regard it is
noteworthy that Discus cronkhitei greatly outnum-
bered Vertigo gouldi only in the southwest half of the
study area whereas the converse was true only in the
northeast half.

Mozley (1934) recorded the same four species of
zonitids as we did (Table 2) in Alberta; however, he
referred the specimens of Vitrina to the species V.
limpida. He did not record the endodontid, Punctum
minutissimum, a species we have found to be reasona-
bly abundant. Of the succineids, he recorded only the
genus Succinea and he referred the specimens to S.
retusa. Among the pupillids, he recorded neither the
genus Gastrocopta nor Columella in Alberta but did
record Vertigo; however, he referred these specimens

to the species V. ovata. Of the valloniids, he recorded
neither Vallonia gracilicosta nor Zoogenetes harpa. It
is not surprising for the latter as we have found it
infrequently and then only in the southwest corner of
the area surveyed. Mozley (1934), however, did record
Vallonia costata, a species which we have not found.
We are unable to comment on the differences in the
species identified as we have not seen Mozley’s (1934)
material. Mozley (1934) recorded neither the limacid,
Deroceras laeve, a species apparently widespread in
the province (Platt 1980; Boag, unpublished data) nor
the terrestrial basommatophoran, Carychium exile.

Acknowledgments

We are pleased to acknowledge the stimulus pro-
vided by A. N. Spencer, who initially suggested this
study; the logistic help received in the field from R.
Seward and R. Mandryk; the statistical advice and
analyses completed by A. Wroot; and the confirma-
tion of identification of the molluscs by M. F. I.
Smith.

Literature Cited

Berry, S. S. 1922. Land smails from the Canadian Rockies.
Bulletin of the Victoria Memorial Museum 36: I-19.

Mozley, A. 1930. Reports of the Jasper Park Lakes investi-
gations, 1925-26: the Mollusca of Jasper Park. Transac-
tions of the Royal Society of Edinburgh 56: 647-669.

Mozley, A. 1933. The local and geographic distribution of
some Rocky Mountain Mollusca. Proceedings of the
Malacological Society of London 20: 214-221.

1981

Mozley, A. 1934. Mollusca from the provinces of Saskat-
chewan and Alberta, Canada. Proceedings of the Malaco-
logical Society of London 21: 138-145.

Platt, T. R. 1980. Observations on the terrestrial Gastro-
pods in the vicinity of Jasper, Alberta (Canada). Nautilus
94: 18-21.

Russell, L.S. 1951. Land snails of the Cypress Hills and
their significance. Canadian Field-Naturalist 65: 174-175.

Received 29 March 1980
Accepted 29 May 1980

APPENDIX I—Locations at which samples were collected in

the vicinity of Edmonton, Alberta

Sample site.” Location Forest cover
9 3.3 km N of Fallis Point Poplar
14 1.6 km W of Villeneuve Poplar
18 2.1 km N of Mewassin Poplar
19 12 km N of Darwell Poplar
20 9.1 km N of Gunn Poplar
21 Highway 2S
at Pigeon Lake turnoff Mixed forest
23A South end of Battle Lake Willow
24 7.5 km E of Winfield Mixed forest
24A 7.5 km E of Winfield Mixed forest
25 4.5 km NE of Warburg Poplar
26 2.5 km W of Sunnybrook Mixed forest
Dil 11.6 km N of Warburg Poplar
28 6.6 km N of Genesee Poplar
30 4.1kmS of Miquelon Lakes Poplar
32 Miquelon Lakes
Provincial Park Poplar
33 Junction Highway 14
and Highway 21 Poplar
34 Ministik Campground Mixed forest
35 0.8 km E of Tofield Poplar
36 13 km N of Tofield Poplar
37 Junction Highway 16
and Highway 834 Poplar
38 West Baseline Elk Island
Park, where Highway 16
passes through Poplar
39 3.1 km N of Chipman Poplar
40 1.6 km W and 0.8 km N
of St. Michael Mixed forest
4] 1.6 km W and 2.3 km N

of Deerland

Poplar

9)

ye

m~mIOB4 << Cr

NAL

VAN ES AND BOAG: TERRESTRIAL MOLLUSCS, ALBERTA

8.3 km N of Scotford

5 km N and 1.6 km W
of Akenside

8.6 km N and 1.6 km W
of Redwater

2.5 km S of Opal

1.6 km N and 1.6 km W
of Gibbons

7.5 km E and 3.3 kmS$
of Legal

7.5 km E and 1.2 kmN
of Winfield

South Bank N. Saskatchewan
River, Edmonton, near
Highlevel Bridge

South Bank N. Saskatchewan
River, Edmonton, near
University of Alberta

South Bank N. Saskatchewan
River, Edmonton, West
of Groat Bridge

South Bank N. Saskatchewan
River, Edmonton, near
Whitemud Creek

South Bank N. Saskatchewan
River, Edmonton, near
Highlevel Bridge

South Bank N. Saskatchewan
River, Edmonton

8.3 km NE of Clyde Corner

1.6 km W and 3.3 kmN
of Morinville

0.8 km E of Darwell

9.1 km S of town of
Alberta Beach

79

Poplar
Poplar

Poplar
Poplar

Poplar
Poplar

Poplar

Poplar

Poplar

Poplar

Poplar

Poplar

Poplar
Poplar

Poplar
Poplar

Poplar

Sandy Lake, near campground Poplar

6.6 km S of Alexander
Indian Reserve

3.3 km W and 1.6 kmS
of Sundance

8.3 km W of Highvale

6.6 km W of Highvale

4.1 km NW of town of
Battle Lake

South end of Battle Lake

South end of Coal Lake

1.6 km S of Miquelon Lakes

Poplar

Poplar
Poplar
Spruce bog

Mixed forest
Willow
Poplar
Poplar

“See Figure | for positions of numbers and letters.
’Numbers refer to leaf litter samples (nm = 31). Letters refer to
handpicked samples (n = 19).

Deer Mortality in the Pohénégamook Wintering Area, Quebec

FRANCOIS POTVIN,! JEAN HUOT,? and FRANCOIS DUCHESNEAU!

\Ministére du Loisir, de la Chasse et de la Péche, 9530 rue de la Faune, Orsainville, Québec GIG 5E5
2Département de Biologie, Université Laval, Sainte Foy, Quebec GIK 7P4

Potvin, Francois, Jean Huot, and Francois Duchesneau. 1981. Deer mortality in the Pohénégamook wintering area,
Quebec. Canadian Field-Naturalist 95(1): 80-84.

From 1972 to 1979 a study was conducted in the Pohénégamook wintering area to identify the major factors responsible for
mortality of White-tailed Deer (Odocoileus virginianus) under severe environmental conditions. Aerial surveys, pellet-group
counts, and dead-body surveys were used to monitor the population every year and to estimate the natural mortality. During
the period of the study the population decreased from 480 to 140 deer, a 71% decline. This resulted from very high winter
mortality rates which in some years exceeded 40%. The most severe winters were those of 1975 and 1977 when snow depth was
more than 50 cm during almost the entire season. We estimated that respectively 200 and 120 deer died in these winters. Death
by starvation was the highest of all winter mortalities of known causes. Most of the 121 mortality cases during the study were
fawns or adult females. We suggest that these heavy mortality rates result from extremely severe winter conditions, in the
presence of available food, rather than from “over-population.”

Potvin, Francois, Jean Huot, and Francois Duchesneau. 1981. Deer mortality in the Pohénégamook wintering area,
Quebec, Canadian Field-Naturalist 95(1): 80-84.

De 1972 a 1979, le ravage de Pohénégamook servit de cadre a une étude entreprise dans le but d’identifier les facteurs de
mortalité ducerf de Virginie (Odocoileus virginianus) dans des conditions extrémes d’environnement. Des inventaires aériens,
des dénombrements de tas de féces et des inventaires de carcasses ont permis de suivre la population d’année en année et
d’estimer la mortalité naturelle. Au cours de la période, la population est passée de 480 a 140 cerfs, soit une baisse de 71%. Ce
déclin fut provoqué par des taux de mortalité hivernale élevés, atteignant plus de 40% en certaines années. Les hivers 1975 et
1977 s’'avérérent particuliérement néfastes alors que l’enneigement dépassa le seuil critique de 50 cm durant pratiquement
toute la saison. Ona estimé que respectivement 200 et 120 cerfs avaient alors péri. La mort par inanition vient au premier rang
des cas de mortalité dont on a pu établir la cause. Les faons et les femelles adultes représentent la majorité des 121 cas
rapportés au cours de l’étude. Nous croyons que les fortes mortalités observées dépendent des conditions d’hiver extreémement
rigoureuses, méme si la nourriture est présente en quantité suffisante, plut6t que d’un état de “surpopulation.”

Key Words: White-tailed Deer, Odocoileus virginianus; winter, snow, mortality, starvation.

Heavy winter mortality rates for deer have tradi- _is 280 cm and snow thickness or depth reaches 70 cm

tionally been associated with “over-population” by mid-March. Duration of the period with more than
(Leopold et al. 1943; Robinette et al. 1952; Severingh- 50cm of snow on the ground usually exceeds 80 d.
aus 1956, 1972, 1976; Severinghaus and Free 1963). The topography of the area ts gently rolling and the

The traditional attitude of the deer manager under general exposure is in a southwesterly direction. The
such circumstances has been to allow more liberal forest as a whole is characterized by Sugar Maple
hunting seasons to reduce the population. We mayask (Acer saccharum) and Yellow Birch (Betula allegha-
whether this strategy is still valid onthe northernedge —niensis) on the hill tops with Balsam Fir (Abies balsa-
of the species’ range where severe wintersare frequent mea) and White Spruce (Picea glauca) in the valleys
and may cause mortality in absence of overpopula- (Rowe 1972). Following fire both White Birch (Betula
tion. Very few studies have been conducted to docu- papyrifera) and Trembling Aspen (Populus tremu-
ment deer mortality under northern conditions, and /oides) gain numerically. Selective cuttings have been
those are short-term ones (Runge and Wobeser 1975; _ the rule for many years in the southern portion of the
King 1976). In this paper we have summarized the wintering area while large clear-cuttings were exe-
findings of an 8-yr study on the effects of winter cuted in the northern half during the sixties. Since
severity ona White-tailed Deer (Odocoileus virginia- summer 1975, rather heavy mortality occurred in Bal-
nus) population at the northern fringe of its range. sam Fir stands as a result of the recent Spruce Bud-
worm (Choristoneura fumiferana) epidemic.
The annual harvest in this area dropped from an
Study Area average of 70 deer to about 20 bucks after the buck law
The Pohénégamook wintering area is situated was instituted in 1974. No important natural predator
200 kmeast of Quebec city, inthe Lower St. Lawrence was present when we initiated the study but the
region. The most severe winter conditions of Quebec’s Coyote (Canis /atrans) appeared in the area in the
deer range are reported from this area. Total snowfall mid-1970s.

80

1981

POTVIN AND AL.: DEER MORTALITY IN QUEBEC 81

ISSIs72-
1978

>
zZ

Wee

“Ss
AR}

SAINT-ELZE

Ficure1. Extent of the Pohénégamook wintering area in 1972-1974 and 1978.

Materials and Methods

Aerial surveys were used to map the extent of the
wintering area. Each spring or early summer pellet-
group counts were taken to estimate the deer popula-
tion of the previous winter. Sampling plots 2 X 40 m
were systematically laid along transects equally
spaced and perpendicular to contour intervals. Defe-
cation rate was fixed at 13 pellet-groups per day and
number of yarding days was set at 165 (mid-
November to end of April).

Snow depth was monitored every winter in a conif-
erous stand. From 1974 to 1979 systematic dead-body
surveys were conducted in the spring, along the same
transects used for the pellet-group count, and covered
5-10% of the yard. Dead deer were aged by tooth
replacement and wear (Severinghaus 1949) and their

physical condition was evaluated by the femur mar-
row fat content (Cheatum 1949). Deer whose mortal-
ity cause was not evident and with less than 25% fat in
the bone marrow were classified as dead by starvation.

Results

The wintering area decreased from 30 km? in
1972-1974 to only 11 km?in 1978 (Figure 1). The deer
population underwent a 71% decline from 1972 to
1979 passing from 480 to 140 deer (Table 1).

According to snow depth, winters of 1975 and 1977
were the most severe (Figure 2). Under coniferous
cover, snow depth exceeded 50 cm for 17 wk or more
and reached 70 cm from late January or early Febru-
ary. No supporting crust was present during these two
winters except at the end. On the contrary, in 1973 and

82 THE CANADIAN FIELD-NATURALIST

TABLE |—Estimation of deer population and mortality rate in the Pohénégamook wintering area, 1972-1979

No. of

dead deer

Population from the found in

Winter pellet-group count the survey

1972 480 + 120° No survey

1973 450 + 60 No survey
1974 450 + 60 5
1975 360 + 60 14
1976 220 + 50 0
1977 280 + 50 7
1978 165 + 30 lI
1979 140 + 50 4

Vol. 95
No. of Population
deer at the
estimated beginning Mortality
to be dead of winter’ rate (%)
100 480 21
200 430 47
— 220 —
120 320 38
80 190 42
25 150 17

*Population at the beginning of winter = population from the pellet-group count + '/, number of deer estimated to be dead.
In this equation we assume that deer are dead after 110 d on the average, so that they would not have defecated during the last

third of the winter period.
Ninety percent probability confidence limits.

1974 snow depth of 50-60 cm was typical, and a sup-
porting crust facilitated deer movements. The winter
of 1976 was also mild as the 50-cm threshold was
attained for only 8 wk and snow melt occurred as
early as mid-April. However, the winter of 1978 was
rather severe because difficult snow conditions were
reached at the end of the season when deer are in poor
shape. Finally, despite 11 wk with snow depth of
50 cm or more, the winter of 1979 was an easy one
because of the presence of a supporting crust from
early February.

Deer mortality estimates vary from year to year
between 25 and 200, excluding 1976 when no estimate
could be obtained as no carcass was found (Table 1).
During the winters of 1975, 1977, and 1978, the mor-
tality rate was between 38 and 47% of the population,
twice the rate of the winters of 1974 and 1979. Mortal-
ity causes were identified for 46 carcasses: 36 cases of
starvation, 7 of predation, 2 accidental, and | case of
poaching. Five of the seven cases of predation were
found in 1978, but their identification is doubtful
because they were reported by people outside of our
study group. The femur marrow of || additional deer
had a fat content exceeding 25%, but the cause of
death could not be identified. Fawns accounted for
44% (n = 94) of all the carcasses and 80% (n = 25) of
the starvation deaths. The sex ratio of the adults
(1.5 yrand older) was strongly biased toward females,
being 24 males per 100 females (n = 44).

Discussion

Severinghaus (1947, 1976) indicated that winter
conditions become critical for deer survival when
snow depth of 50 cm persists for 50 d or more in the
Adirondack region of New York. Deep snow covers
low-growing vegetation, increases energy expenditure

of deer, and confines them to dense coniferous stands
where the food supply is often limited. In 1975 and
1977, this 50-cm critical level was exceeded even in
coniferous stands during almost the entire season.
Such conditions were associated with mortality rates
of 40-50%, twice the rate of winters of 1974 and 1979,
two mild years. Deep snow at the end of winter, how-
ever, seems to have the same effect because the mortal-
ity rate was the same in 1978, when only a short critical
period was registered in late March and early April. In
April, deer have already exhausted almost all their fat
reserves so that any additional energy demand might
be too high.

A good habitat can certainly buffer the effects of
severe winters; however, the coniferous cover in the
Pohénégamook area had been reduced by large clear-
cuttings and the recent budworm epidemic. We may
expect lower mortality rates for deer facing the same
winter conditions in a better habitat.

Most of the deer that died of starvation were fawns.
This is normal because fawns have a physical han-
dicap, being smaller, and their metabolic energy needs
are proportionally higher. The large preponderance of
females in the adult segment is amazing at first and
cannot be explained by an excessive removal of males
through hunting because the harvest rate on adult
males is only about 15%. Females begin the winter in
better condition than males because the latter may use
a large proportion of their fat reserves during the rut
(Siegler 1968). However, it is possible that lactation
precludes some females from accumulating fat depos-
its. Fall foods with a high energetic content such as
mast are not available in this area. Furthermore,
pregnant females will spend much energy for their
fetuses especially at the end of gestation (Verme 1977).
This may explain why females are more vulnerable.

1981
1973 1974
20 b i
13 wk Il wk
5
<
a
3
c4
°
H
1977 1978
120 -
17 wk
€
a
3
=
2
’p)

1 : 2 . — bennett
Month Month

POTVIN AND AL.: DEER MORTALITY IN QUEBEC 83

1975 1976

18 wk 8 wk

1979

——- Estimated

* Supporting crust

J F M A
Month

FIGURE 2. Snow depth under coniferous cover and number of weeks with more than 50 cm in the Pohénégamook wintering

area, 1973-1979.

In spite of the general belief that overpopulation is
responsible for high winter mortality rates, we do not
believe that such is the case for the Pohénégamook
wintering area. Browse surveys conducted annually
since 1972 indicate that only one stem out of three and
one twig out of five were browsed by deer in the
coniferous forest type which supports the highest deer
densities (F. Potvin, unpublished data). This utiliza-
tion rate is lower than in many other yards in the
province and the browse species comprise mostly
Mountain Maple (Acer spicatum), Beaked Hazelnut
(Corylus cornuta), and Balsam Fir, typical of our deer
range. These twigs were available in winter because
only twigs between 50 and 210 cm from the ground
were included in the surveys and most of them were

above the 70-cm level. Therefore the quantity of food
does not appear to be a limiting factor.

Although good quality browse may be abundant in
winter, deer will use some of their fat reserves each
day, losing 20-30% of their weight during the whole
season (Mautz 1974). This is a normal adaptation to
compensate for the low digestibility of browse and
also to help supply enough energy to face the harsh
climatic conditions. A longer winter or difficult snow
conditions can exhaust these limited reserves before
springtime. Furthermore, a sudden heavy demand
caused by rapidly deteriorating climatic conditions
(snow storm) or harassment may kill an animal even
though its energy reserves are not completely
exhausted. This may explain the |! mortality cases

84 THE CANADIAN FIELD-NATURALIST

that had more than 25% of fat in the bone marrow.
Mitchell and Staines (1976) made similar observa-
tions on Red Deer (Cervus elaphus).

Habitat is usually seen as the ultimate limiting fac-
tor for deer because this species has no self-regulating
mechanism working through behavior or social stress
(Klein 1967). But some authors have suggested that
winter severity can also act as a regulating mechanism
and preclude habitat over-utilization (Klein 1962,
1965; Ransom 1967; Kucera 1976). Potvinet al. (1978)
hypothesized that harvest strategies based on the con-
cept of carrying capacity of the range could not be
applied at the northern limit of the White-tailed Deer
continental range. We believe that under these cir-
cumstances deer population dynamics and levels at a
given time are related to the general environmental
conditions and to the recent history of the herd
(exploitation, predation, etc.). When winter condi-
tions are severe for the area, a minimum mortality rate
by starvation is to be expected even in the absence of
any sign of overpopulation and in a fair to good
habitat. Edwards (1956) and Severinghaus (1972,
1976) also suggest this possibility. It appears that, at
least in Quebec, occasional severe winters can act
almost independently of the deer density and prevent
Over-utilization of the range by deer.

We then view the regulation of White-tailed Deer
populations as an example of the idea suggested by
Huffaker and Messenger (1964) according to which
the importance of the density-dependent factors
decreases as the environmental conditions become
more variable and the favorable microhabitats are
more scattered. This is probably the case at the limit of
the continental deer distribution especially when the
habitat cannot buffer these climatic variations.
Pohénégamook seems to fit perfectly in this case.

Acknowledgements

We thank C. W. Severinghaus for reviewing the
manuscript, M. A. Bédard and G. Boucher for provid-
ing unpublished data, and the many persons involved
in the surveys.

Literature Cited

Cheatum, E. L. 1949. Bone marrow as an index of malnu-
trition in deer. Conservationist 3(5):; 19-22.

Edwards, R. Y. 1956. Snow depths and ungulate abun-
dance in the mountains of western Canada. Journal of
Wildlife Management 20: 159-168.

Huffaker, C. B., and P.S. Messenger. 1964. The concept
and significance of natural control. /n Biological control
of insect pests and weeds. Edited by P. DeBach. Chapman
and Hall Ltd., London. pp. 74-117.

King, D. R. 1976. Estimates of the White-tailed Deer popu-
lation and mortality in central Ontario, 1970-1972. Cana-
dian Field-Naturalist 90: 29-36.

Vol. 95

Klein, D. R. 1962. Rumen contents analysis as an index to
range quality. Transactions of the North American Wild-
life and Natural Resources Conference 27: 150-162.

Klein, D. R. 1965. Ecology of deer range in Alaska. Ecolog-
ical Monograph 35: 259-284.

Klein, D.R. 1967. Interactions of Rangifer tarandus
(Reindeer and Caribou) with its habitat in Alaska. Inter-
national Congress of Game Biologists 8: 289-293.

Kucera, E. 1976. Effects of winter conditions on the White-
tailed Deer of Delta Marsh, Manitoba. Canadian Journal
of Zoology 54: 1307-1315.

Leopold, A., E. F. Bean, and C. Fassett. 1943. Deer irrup-
tions. Transactions of the Wisconsin Academy of Scien-
ces, Arts and Letters 35: 351-366.

Mautz, W. W. 1974. Deer nutritional and physiological
research — So what? Transactions of the Northeastern
Fisheries and Wildlife Conference 31: 113-122.

Mitchell, B.,and B. W. Staines. 1976. Anexample of natu-
ral winter mortality in Scottish Red Deer. Deer 3:
549-552.

Potvin, F., M. Bélanger, and S. Georges. 1978. Deer
decline in Québec and harvest strategies at the northern
limit. Canadian Wildlife Administration 2(3): 43-50.

Ransom, A. B. 1967. Reproductive biology of White-tailed
Deer in Manitoba. Journal of Wildlife Management 31:
114-123.

Robinette, W. L., O. Julander, J. S. Gashwiller, and J. G.
Smith. 1952. Winter mortality of Mule Deer in Utah in
relation to range condition. Journal of Wildlife Manage-
ment 10: 83-89.

Rowe, J. S. 1972. Forest regions of Canada. Department of
the Environment, Canadian Forestry Service Publication
1300. 172 pp.

Runge. W., and G. Wobeser. 1975. A survey of deer winter
mortality in Saskatchewan. Saskatchewan Department of
Tourism and Natural Resources, Wildlife Report 4. 22 pp.

Severinghaus, C. W. 1947. Relationship of weather to win-
ter mortality and population levels among deer in the
Adirondack region of New York. Transactions of the
North American Wildlife Conference 12: 212-223.

Severinghaus, C. W. 1949. Tooth development and wear as
criteria of age in White-tailed Deer. Journal of Wildlife
Management 13: 195-216.

Severinghaus, C. W. 1956. History, management and ecol-
ogy of White-tailed Deer in Allegany State Park. New
York Fish and Game Journal 3(10): 80-87.

Severinghaus, C. W. 1972. Weather and the deer popula-
tion. Conservationist 27(2): 28-31.

Severinghaus, C. W. 1976. The Central Adirondacks. Con-
servationist 30(5): 26-27.

Severinghaus, C.W., and S. Free. 1963. Management
implications of the trend and distribution of the legal kill
in the Adirondack region. New York Fish and Game
Journal 10(2): 201-214.

Siegler, H. R. (editor). 1968. The White-tailed Deer of New
Hampshire. New Hampshire Fish and Game Department,
Survey Report 10. 256 pp.

Verme, L. J. 1977. Assessment of natal mortality in Upper
Michigan. Journal of Wildlife Management 41: 700-708.

Received 25 February 1980
Accepted 30 June 1980

Moose Browse Utilization in Mount McKinley National Park, Alaska

JERRY O. WOLFF! and JOANNE COWLING?2

'USDA Forest Service, Institute of Northern Forestry, Fairbanks, Alaska, and Department of Biology, University of
Virginia, Charlottesville, Virginia 22901
2Institute of Arctic Biology, University of Alaska, Fairbanks, Alaska 99701

Wolff, Jerry O., and Joanne Cowling. 1981. Moose browse utilization in Mount McKinley National Park, Alaska.
Canadian Field-Naturalist 95(1): 85-88.

During winter of 1978-1979, Moose (A/ces alces) browsing intensities in Mount McKinley National Park averaged 65%.
Percentages of available browse consumed were greatest in small, dense, single-species stands of Felt-leaf Willow (Salix
alaxensis) and less in large, mixed-species stands. Moose populations are below but likely approaching the carrying capacity
of the range.

Key Words: Alaska, Alces alces, browse, carrying capacity, habitat, Moose, Salix.

Mount McKinley National Park supportsa Moose _ one at Teklanika River (46.6 km), one at Teklanika
(Alces alces) population of about 1000 animalsinan _ Bridge (48.0 km), and two at Igloo Creek (51.0 and
area of 8000 km? (J. Dalle-Molle, National Park Ser- 52.0 km). All willows sampled in these stands were
vice, Mount McKinley National Park, Alaska, per- __ Felt-leaf Willow (Salix alaxensis); this was a highly
sonal communication). There is a higher density of | preferred willow and an abundant species in the park.

animals in favorable Moose habitat in the park than Numbers of browsed and unbrowsed twigs were
has been recorded in other regions of interior Alaska — recorded on 100 willow stems along a 500-m transect
with similarly favorable Moose habitat (W. C. Gasa- _ in each stand. Two stems closest to the observer (one

way, Alaska Department of Fish and Game, Fair- right and one left) were sampled at 10-m intervals
banks, Alaska, personal communication). During along each 500-m transect. All twigs were less than
winter, Moose congregate in tall shrub communities 4mm in diameter and between 0.5 and 3.5 m above
to feed on several species of willow (Salix spp.) and —_ ground level. Browsing intensity was defined as the
other woody vegetation. As part of an extensive proportion of twigs, available to Moose within the
Moose browsing study in interior Alaska (Wolff 1978; | sample area, which were browsed.

Wolff and Zasada 1979), we measured browse availa- In three stands along Savage River (11.8 km,
bility and utilization by Moose in Mount McKinley 19.8 km, 20.3 km), numbers of browsed and
National Park. It was hypothesized that Moose popu- _ unbrowsed twigs were recorded in 20 plots, 2 X 5 m.

lation density in Mount McKinley National Park All browse species were included in the sampling. The
would be at or near carrying capacity of the habitat Shafer (1963) twig-count method was used to estimate
when hunting is totally restricted and fire absent. The availability and utilization of browse. Diameters at
last fire recorded in the park occurred in 1924. point of browsing were measured on 30 browsed twigs
To assess this hypothesis, we recorded browsing of each species. Thirty unbrowsed twigs of the same
intensities and the biomass of available browse and _ diameter were clipped, dried, and weighed.

estimated carrying capacities in the major wintering A road and aerial census was conducted from 3
areas in the park. through 8 June 1979 to count cows and calves. Addi-

tional data on natality, survival, and predation were
Study Areas and Methods supplied by the National Park Service and Haber

The major wintering areas for Moose inthe park (1977).

are along Savage, Sanctuary, and Teklanika rivers,

Igloo Creek, and near park headquarters. Most of Results

these areas are within 2 km of the park highway. Browsing intensities of S. alaxensis at the 14 stands
Browsing intensities were recorded in 14 stands at areshownin Table |. The mean browsing intensity for
seven sites (Figure |). One transect was sampled near _ all stands was 65.0% ranging from 35.1% at Teklanika
the park entrance by the railroad tracks(1.8 kmfrom Campground to 86.3% at headquarters, 8.3 km from
the highway turnoff to the park), one at headquarters _ the park entrance. Browsing intensities corresponded
(8.3 km), one at Hogan Creek (34.2 km),twoatSanc- to the distribution and abundance of forage. A mean
tuary River (34.5 km), oneat Sanctuary Campground (+SE) browsing intensity of 75.8+2.91% was
(35.8 km),one at Teklanika Campground (45.6 km), recorded in single-species stands and/or stands which

85

86

THE CANADIAN FIELD-NATURALIST

McKINLEY NATIONAL

Sanctuary

ice Hogan

{creek

Vol. 95

Y pyiu0/4aLl

Teklanika
“Campground

Q
\ Teklanika
Cerise
O

Igloo
O Creek

\Sanctuary
Campground

Z0niues

Headquarters
} ee

Pe
Railroad
Tracks

Savage River (1.8km)

>
bE

Be
7)
x
i
io)
a

To Anchorage

FiGuRE 1. Map showing the location of the 14 study sites and the browse sampling sites in Mount McKinley National Park,

Alaska.

were dispersed in small dense patches of less than 5 ha
(1 ha = 10 000 m2), such as on river islands, in clear-
ings, or along streams. Browsing intensities were sig-
nificantly lower (P< 0.01, t-test) in mixed-species
stands greater than 5 ha in size (52.8 + 4.08%) or
when forage shrubs were widely dispersed. The rela-
tive proportions of the stand types in Mount McKin-
ley National Park are not known. There was no corre-
lation between browsing intensities and number of
twigs per stem.

Biomass of browse available, carrying capacities,
and browse utilization at the three Savage River
stands are shown in Table 2. The mean (+SE) diame-
ters at point of browsing were 3.6 + 0.13 mm for S.
alaxensis, 2.8+0.09 mm for Littletree Willow (S.
arbusculoides), and 3.0 = 0.10 mm for Grayleaf Wil-
low (S. glauca). The mean weights per twig were
0.64 + 0.02 g, 0.55 + 0.02 g, and 0.60 + 0.02 g for the
three species, respectively. Balsam Poplar (Populus
balsamifera) is a browse species in other regions of
Alaska and was included in the sampling. Balsam

Poplar has a mean weight per twig of 1.32 + 0.07 g for
a 4-mm twig (Wolff and Zasada 1979). The amount of
browse available ranged from 39.2 to 110.7 kg/ha.
This is within the range reported by Wolff and Zasada
(1979) for 16 stands in interior Alaska. Salix alaxensis
made up 90% of the browse available in all stands
followed by S. arbusculoides, S. glauca, and P. bal-
samifera. Diamondleaf Willow (S. planifolia) was
also present in the study area but not in our plots.
American Green Alder (A/nus crispa) was present in
most study areas but was infrequently browsed. Pref-
erence was shown for S. a/laxensis, followed by S.
arbusculoides, S. glauca, and P. balsamifera.

At a daily consumption rate of 5 kg dry weight per
Moose (Gasaway and Coady 1974), the biomass of
browse available in the three stands would support
between 5.9 and 16.7 Moose:d '*ha' (MD/ha).
Moose are usually food-stressed before they consume
100% of the forage available in an area; therefore,
actual carrying capacity was adjusted to 75% of total
browse available (Wolff and Zasada 1979). The actual

1981

TABLE 1—Numbers of twigs per stem (+ SE) and percentage
of browsing intensity (+ SE) by Moose (Alces alces) on Salix
alaxensis at 14 stands sampled in Mount McKinley National
Park, Alaska, during May and June 1979. Number of stems
(N) = 100, unless otherwise indicated

Browsing
Stand Twigs/stem intensity
Single species stands or small dense stands
Headquarters 1.8 km 4.6 (0.23) 75.7 (3.22)
Headquarters 8.3 km 5.0 (0.25) 86.3 (2.19)
Hogan Creek 6.1 (0.35) 83.7 (3.73)
Sanctuary River I 13.6 (2.22) 75.1 (3.22)
Sanctuary River II 5.3 (0.33) 76.5 (3.15)
Sanctuary Campground 6.0 (0.32) 70.7 (3.12)
Igloo Creek I, 51 km 5.6 (0.26) 63.1 (3.76)

Mean 75.8 (2.91)**

Mixed species stands and/or dispersed stands

Savage River] (N=228) 4.3 (0.17) 68.5 (2.61)
Savage River II

(N = 224) 11.4 (2.28) 59.1 (2.56)
Savage River III

(N = 158) 18.7 (2.70) 50.6 (3.13)
Teklanika Campground 14.9 (2.42) 35.1 (3.88)
Teklanika River, 46.6km 4.9 (0.25) 44.6 (4.03)
Teklanika Bridge 4.6 (0.26) 53.9 (4.21)
Igloo Creek, 52 km 5.3 (0.26) 57.9 (3.80)

Mean 52.8 (4.08)**

**Percent browsing intensities significantly different
(P< 0.01, t-test).

WOLFE AND COWLING: MOOSE BROWSE UTILIZATION, ALASKA

87

utilization of S. alaxensis was about 79% of the —
adjusted carrying capacity. Forall browse species uti-
lization was between 64 and 83% of the adjusted carry-
ing capacity.

We counted 23 calves and 27 cows from 3 through 8
June 1979 for a calf:cow ratio of 85:100 (National
Park Service aerial census and personal observa-
tions). Haber (1977) recorded spring calf:cow ratios
ranging from 58:100 to 126:100 from 1966 to 1974.
Seven cows we observed had twins fora twinning rate
of 26%: the incidence of twinning was between 38 and
55% from 1966 to 1974 (Haber 1977). Fall calf:cow
ratios ranged from 12:100 to 38:100 from 1969 to 1973
(Haber 1977) and from 8.2:100 to 18.9:100 from 1974
to 1978 (W. Troyer, National Park Service, Anchor-
age, Alaska, unpublished report).

Discussion

Earlier Wolff and Zasada (1979) suggested that
actual carrying capacity of a habitat is about 75% of
total browse available; 25% is either of poor quality,
inaccessible, or otherwise too difficult to obtain. They
recorded browsing intensities of greater than 75% in
small homogenous stands, but larger stands had
browsing intensities less than 60%. Similar trends of
heavy utilization of small or single-species stands and
lighter utilization of large mixed-species stands were
noted here. In large mixed-species stands, Moose have

TABLE 2—Production of woody browse and utilization by Moose (Alces alces) at the Savage River stands, Mount McKinley

National Park, Alaska

Stems/ha Twigs/stem
Stand Species (xt SE) (x SE)
Savage Salix alaxensis 11 600 (1700) 4.3 (0.17)
River I S. arbusculoides 2 700 (850) 4.9 (0.33)
Total 14 300
Savage S. alaxensis 11 700 (2040) 11.4 (2.28)
River II S. arbusculoides 2 800 (960) 4.0 (0.29)
S. glauca 300 (250) 5.8 (1.31)
Total 14 800
Savage S. alaxensis 8 500 (1500) 18.7 (2.70)
River III 9S. arbusculoides 2 800 (960) 4.0 (0.29)
S. glauca 300 (160) 5.7 (0.58)
Populus balsamifera 150 (80) 9.0 (1.52)
Total 11 750

Adjusted
Browse Browsing carrying
available! intensities capacity? Utilization?
(kg/ha) (%+SE) (M.D./ha*) (M.D./ha‘4)
31.9 68.5 (2.61) 4.8 4.4
7.3 31.7 (5.41) 1.1 0.5
39.2 56.8 (3.14) 59) 4.9
85.4 59.1 (2.56) 12.8 10.1
6.2 55.0 (5.05) 0.9 0.7
1.0 16.0 (6.79) 0.2 0
92.6 58.0 (2.45) 13.9 10.8
101.7 50.6 (3.13) 15.3 10.3
6.2 34.6 (7.86) 0.9 0.4
1.0 23.0 (8.05) 0.2 0
1.8 0 0.3 0
110.7 48.1 (2.94) 16.7 10.7

'Salix alaxensis 0.64 g/twig, S. arbusculoides 0.55 g/twig, S. glauca 0.60 g/twig, P. balsamifera 1.32 g/twig.

2Adjusted carrying capacity:
5 kg:‘Moose -d
Browsing intensity X total browse available
‘ 5 kg:Moose 'd '
4M.D./ha: Moose days per hectare.

3Utilization:

75% of browse available (Wolff and Zasada 1979).
Let ON ENISS e

(Gasaway and Coady 1974).

88 THE CANADIAN FIELD-NATURALIST

a greater number of species and individual plants from
which to select good quality forage (Oldemeyer et al.

_ 1977) than they do in smaller and more dense single-
species stands. In small, patchily distributed stands it
is probably easier to consume large quantities of for-
age before moving to another patch.

The relatively high browsing intensities recorded
suggest that the population is approaching or at the
carrying capacity as determined by available food.
Subjective observations made during this study sug-
gest browsing intensities were also high in previous
years, but relatively high twinning and natality rates in
1979 indicate the population might still be below the
actual carrying capacity. If this is the case, the Moose
population is currently being controlled by some
means other than low forage availability. The twin-
ning rate was lower in 1979 than in previous years,
however, and this suggests that forage quality or
quantity may be becoming a limiting factor in Moose
numbers. Small sample sizes prevent drawing any
firm conclusions.

The drop incalf:cow ratios recorded from spring to
fall suggest high summer mortality, likely the result of
predation by Gray Wolves (Canis /upus) or, to a lesser
extent, Grizzly Bears (Ursus arctos) (Haber 1977;
Gasaway et al. 1977; S. Buskirk, National Park Ser-
vice, Mount McKinley National Park, Alaska; and
W. Ballard, Alaska Department of Fish and Game,
Glenn Allen, Alaska, unpublished report). Even if
predation were reduced, the Moose population might
increase by only 10-15% before reaching the current
carrying capacity of the habitat.

Most of the tall willow shrub communities in
upland sites in the park are ina climax state with low
current annual growth. The seral communities along
the major river systems are self-perpetuating with
yearly alluvial deposits providing a substrate for
primary succession of willow communities. An
increase in seral habitat resulting from fire in uplands
or continued alluvial deposition along rivers would
improve Moose habitat in Mount McKinley National

Vol. 95

Park. Species composition and size of stands should
be taken into consideration when evaluating and
managing Moose habitat.

Acknowledgments

We thank William C. Gasaway, and two anony-
mous reviewers for helpful comments on this manu-
script. J. Dalle-Molle provided unpublished data and
logistic support. This study was financed by coopera-
tive aid agreement 48-PNW-78 between the Univer-
sity of California, Berkeley, and the Institute of
Northern Forestry, Pacific Northwest Forest and
Range Experiment Station, Fairbanks, Alaska.

Literature Cited

Gasaway, W. C.,and J. W. Coady. 1974. Review of energy
requirements and rumen fermentation in moose and other
ruminants. Naturaliste canadien. 101: 227-262.

Gasaway, W.C., D. Haggstrom, and B.E. Burris.
1977. Preliminary observations on the timing and causes
of calf mortality in interior Alaskan moose populations.
Proceedings of the 13th North American Moose Confer-
ence and Workshop. pp. 54-70.

Haber, G. C. 1977. Socio-ecological dynamics of wolves
and prey ina subarctic ecosystem. Ph.D. thesis, University
of British Columbia. 817 pp.

Oldemeyer, J. L., A. W. Franzmann, A. L. Brundage, P. D.
Arneson, and A. Flynn. 1977. Browse quality and the
Kenai moose population. Journal of Wildlife Manage-
ment. 41: 533-542.

Shafer, E. L., Jr. 1963. The twig-count method for measur-
ing hardwood deer browse. Journal of Wildlife Manage-
ment. 27: 428-437.

Wolff, J. O. 1978. Burning and browsing effects on willow
growth in interior Alaska. Journal of Wildlife Manage-
ment. 42: 135-140.

Wolff, J. O., and J. C. Zasada. 1979. Moose habitat and
forest succession on the Tanana river floodplain and
Yukon-Tanana upland. Proceedings of the 15th North
American Moose. Conference and Workshop. pp.
213-244.

Received 7 January 1980
Accepted 23 May 1980

The Sedge Carex loliacea in Eastern North America

A. A. REZNICEK! and P. W. BALL?

'University of Michigan Herbarium, North University Building, Ann Arbor, Michigan 48109
2Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario L5L 1C6

Reznicek, A. A., and P. W. Ball. 1981. The Sedge Carex /oliacea in eastern North America. Canadian Field-Naturalist

95(1): 89-92.

Carex loliacea, a circumboreal sedge found mostly in northwestern North America, has been known since 1954 from the
northern boreal forest of Ontario. Recent collections have considerably extended its range eastward and southward and it is
now known from within the area covered in Gray’s Manual. A key is provided separating C. /oliacea from other similar species
in eastern North America, and complete citations of all known collections east of Manitoba are given.

Key Words: Carex loliacea, flora, Ontario, geographical distribution, records.

Carex loliacea (section Heleonastes) is an essen-
tially circumboreal species that is widespread in
northern Asia, northern Europe, and northwestern
North America (Hultén 1968). Recently it has been
reported froma few scattered localities in the northern
boreal forest of Ontario (Dutilly et al. 1954; Baldwin
1958; Moir 1958). These published records are outside
the area covered by the northeastern North American
floras (Fernald 1950; Gleason 1952; Gleason and
Cronquist 1963), and the species does not extend suf-
ficiently north to be included in eastern arctic floras
(e.g. Porsild 1964). Recent collections (cited below)
have greatly extended its known range both eastward
and southward so that it is now known from within
the area covered by Fernald (1950) and closely
approaches the area of Gleason (1952) and Gleason
and Cronquist (1963).

In this paper we report the distribution of C. /olia-
cea in Ontario and discuss ways in which it can be
distinguished from similar species. We hope that bot-
anists will become more familiar with the species so its
distribution and ecology in eastern North America
will become more fully known.

Description of Carex loliacea

Carex loliacea has a loosely cespitose habit with an
infructescence of few separated spikes bearing a few
narrowly elliptical spreading or reflexed perigynia. In
these characters it closely resembles a number of
widespread and common eastern North American
species of Carex subgenus Vignea, particularly C.
interior of section Stellulatae, C. sy/vicola (C. rosea
auct.) of section Bracteosae, and C. disperma of sec-
tion Heleonastes, and to a lesser extent several other
members of sections Heleonastes and Stellulatae. This
convergence in appearance is a remarkable feature
and has several other parallels in Carex, for example,
in the west, there is great similarity between C. i//ota of
section Ovales, C. hoodii of section Bracteosae, C.

89

vernacula of section Foetidae, and C. jonesii of sec-
tion Vulpinea. Carex /oliacea can be most readily
recognized by the complete absence of a beak on the
perigynium, although in a number of other similar
species the beak may be extremely short (0.2 mm
long). In this regard C. disperma is the species most
likely to be confused with C. /oliacea because it occurs
in similar habitats, may have virtually no beak on the
perigynium, and has only one or two staminate flow-
ers in the terminal spike which can be easily over-
looked. Although the position of the staminate flow-
ers is the only certain means of distinguishing these
species, there are several other features which may be
useful. For example, the perigynia of C. disperma are
either erect or spreading at an angle of less than 90° to
the main axis, tend to taper rather abruptly at the
apex, and often have nine or fewer rather thin nerves
on each surface. The perigynia of C. /oliacea often
spread at an angle of 90° or greater to the main axis,
taper very gradually from the middle to the apex, and
often have nine or more rather stout nerves on each
surface. Two representative infructescences of C. /oli-
acea are illustrated in Figure |.

Identification Key

We have provided the following short key to facili-
tate identification. This key includes all northeastern
North American species of Carex subgenus Vignea
that are cespitose and which have infructescences
composed of several simple spikes. Species other than
C. loliacea are not keyed in detail.

| Terminal spike with staminate flowers at apex or
entirely staminate.
Section Bracteosae, Carex disperma,

Ce ee er

C. sterilis

| Terminal spike with staminate flowers at base or
entirely pistillate.

90 THE CANADIAN FIELD-NATURALIST

53MM _,

FIGURE |. Two representative infructescences of Carex
loliacea.

2 Perigynia lanceolate to broadly ovate, widest
well below middle; spongy layer at base of
perigynium thick, occupying basal 4 to '4 of
perigynium body, usually marked by an ex-
ternally differentiated zone (except in C.
AN CLO BSR rote Maca ia ere cae Silye bs weal aicaniorotnereee onan

2 Perigynia narrowly to broadly elliptic, widest
near middle; spongy layer at base of perigy-
nium thin or absent, not marked by an exter-
nally differentiated zone.

3. Lower two spikes of infructescence over-
lapping, the apex of the lowest reaching to
or beyond the base of the next spike.

SMU yaaa Carex glareosa, C. lachenalii,
C. heleonastes, C. tenuiflora, C. amblyo-
rhyncha, C. mackenzei

3 Lower two spikes of infructescence separ-
ate, the apex of the lowest not reaching to
the base of the next spike.

Vol. 95

4 Perigynia tapering uniformily to the
apex, not contracted to a beak
EN see Seas ai es Carex loliacea

4 Perigynia abruptly contracted to a
short but definite beak 0.2-0.9 mm long
a SE eS ey ae Carex canescens, C.
brunnescens, G. ‘tripsenmaamaGe
mackenzei

Distribution and Ecology of Carex loliacea

The known distribution of Carex loliacea east of
Manitoba now extends to within 130km of the
Quebec boundary (Figure 2). It is likely that searches
will disclose this species in Quebec. In Ontario, the
species is rare in the northern boreal forest and occurs
disjunctly near the north shore of Lake Superior (Fig-
ure 3), paralleling a number of other northern species
(Soper and Maycock 1963). This species should be
looked for in the Lake Superior area of Minnesota
and Michigan as well.

Virtually no information is available about the
ecology of the species in eastern Canada. It evidently
grows in Sphagnum bogs shaded by conifers and in
rich, wet sites that are mossy but often with little or no
Sphagnum. Most of the collections are along river
banks or associated with small streamlets. At the
Moose River site (TRTE 5348) the species occurred

FIGURE2. Distribution of Carex loliacea in North America.

1981

REZNICEK AND BALL: CAREX LOLICEA IN EASTERN NORTH AMERICA 91

FiGuRE 3. Distribution of Carex loliacea in Ontario. @ Specimens examined. Literature records.

with Carex media and other fine-leaved sedges on the
sides of small, mossy hummocks in a network of
streamlets through Picea glauca (White Spruce)
~Thuja occidentalis (White Cedar) woods just above
the bank of the Moose River.

Ontario Records of C. loliacea

Kenora District
Fawn River between Big Trout Lake and Otter
River, 8 July 1952, Moir 357. Deep Sphagnum bog
(CAN 259793).
Attawapiskat River 53°08’N, 83°18’W. 17 July
1957. Porsild et al. 20253. Dry river bank (CAN
787751).*
Attawapiskat River, 60 miles (100 km) upstream
from town. 7 July 1977. Riley 6699. Closed spruce
forest by cool seepage streamlet (TRT 202024) (sub
C. brunnescens).
Albany R. au rapide Frenchman. 51°22’N,
87°48’W. 22 August 1952. Dutilly, Lepage &
Duman 30574. Bois de coniféres (n.v.).

* A number of other specimens from this locality collected in
the same year (e.g., Porsild et al. 20096, 20205) were distrib-
uted as C. loliacea. These specimens are C. disperma.

Thunder Bay District
Port Arthur, east of Hodder Avenue. 2 July 1950. Garton
1156. In muskeg under alders. (DAO 172285).
Brodeur Island, Lake Superior. Bay at north end of
island, 48°33’N, 88°18’W. 8 July 1973. Soper & Given
13151. In Sphagnum under Balsam-Spruce. (CAN
367847) (sub C. interior).

Algoma District
Kapuskasing; 26 miles (42 km) southwest in Shanly Tp.,
on Kapuskasing River. 30 June 1953. Baldwin 4963.
Road through old black spruce forest. (CAN 225843).

Cochrane District
Canfield Tp. Grey Goose Island; 54 miles (9 km) south
west of Moose River railway crossing. 50° 46’N, 81° 24’W.
15 July 1974. Reznicek & Carlton. Wet mesic Black
Spruce —- White Cedar forest at river’s edge. Infrequent
on hummocks associated with runnels of water (TRTE
5348).

Acknowledgements

Our thanks to the curators of the herbaria cited for
making their material available to us and to G. W.
Argus and D. J. White for permission to publish the
maps. This work was partially financed by Natural
Sciences and Engineering Research Council of Can-
ada grant A6494.

92 THE CANADIAN FIELD-NATURALIST

Literature Cited

Baldwin, W. K. W. 1958. Plants of the Clay Belt of north-
ern Ontario and Quebec. National Museum of Canada
Bulletin 156. 324 pp.

Ball, P. W. 1980. Cyperaceae. Atlas of the Rare Plants of
Ontario. Edited by G. W. Argus and D. J. White. Sy//o-
geus. In press.

Dutilly, A., E. Lepage,and M. Duman. 1954. Contribution
a la flore du versant occidental de la baie James, Ontario.
Contributions of the Arctic Institute of the Catholic Uni-
versity of America. SF. 144 pp.

Fernald, M. L. 1950. Gray’s Manual of Botany. 8th edi-
tion. American Book Company, New York. 632 pp.

Gleason, H. A. 1952. The New Britton and Brown illus-
trated flora of the northeastern United States and adjacent
Canada. New York Botanical Garden, New York. Vol. 1.
482 pp.

Vol. 95

Gleason, H. A., and A. Cronquist. 1963. Manual of vascu-
lar plants of northeastern United States and adjacent Can-
ada. Van Nostrand, Princeton. 810 pp.

Hultén, E. 1968. Flora of Alaska and neighboring territo-
ries. Stanford University Press, Stanford. 1008 pp.

Moir, D. R. 1958. A floristic survey of the Severn River
drainage basin, northwestern Ontario. Ph.D. thesis, Uni-
versity of Minnesota, St. Paul. 261 pp.

Porsild, A. E. 1964. Illustrated flora of Canadian Arctic
Archipelago. 2nd edition. National Museum of Canada
Bulletin 146. 218 pp.

Soper, J. H., and P. F. Maycock. 1963. A community of
arctic—alpine plants on the east shore of Lake Superior.
Canadian Journal of Botany 41: 183-198.

Received 6 March 1980
Accepted 17 June 1980

Notes

First Records of the Tundra Shrew (Sorex tundrensis)
in British Columbia

DAVID W. NAGORSEN! and DONALD M. JONES?

!Department of Mammalogy, Royal Ontario Museum, Toronto, Ontario M5S 2C6
2Department of Biology, University of Waterloo, Waterloo, Ontario N2L 3G1

Nagorsen, David W., and Donald M. Jones. 1981. First records of the Tundra Shrew (Sorex tundrensis) in British
Columbia. Canadian Field-Naturalist 95(1): 93-94.

Five Tundra Shrews (Sorex tundrensis) were collected near Haines Road in northwestern British Columbia. These are the
first records of the Tundra Shrew outside the boundaries of the Beringian Refugium, and they extend the range of this shrew
about 500 km south of its known limits.

Key Words: Tundra Shrew, Sorex tundrensis; new records, British Columbia, Beringian Refugium.

The Arctic Shrew (Sorex arcticus)! and the Tundra
Shrew (Sorex tundrensis)'! are boreal and tundra
forms of the Sorex arcticus group described by Jack-
son (1928). The two taxa are allopatric in distribution
(Figure 1). The Arctic Shrew inhabits the southern
Yukon, southwestern Northwest Territories, central
and eastern Canada, and north central United States.
The Tundra Shrew is restricted to Alaska, northern
Yukon, and Northwest Territories. As there are few
records from the Yukon and Northwest Territories,
the limits of the geographic ranges of these two shrews
are poorly known.

During field studies in 1978 and 1979, one of us(D.
Jones) collected five specimens of the S. arcticus
group near Haines Road, Cassiar District, British
Columbia. These specimens are deposited in the col-
lections of the Department of Mammalogy, Royal
Ontario Museum (ROM Nos. 82833, 82834, 83471,
83497, 83498). Based on cranial measurements from
the five specimens (Table 1) and pelage of a skin
prepared from 83471, we identified these shrews as
Tundra Shrews. These represent the first records for FiGure |. Distributions of the Tundra Shrew (light gray)

this shrew in British Columbia. and Arctic Shrew (darker gray) in northwestern

Five Tundra Shrews were trapped at three sites. In North America (modified from Hall and Kelson 1959
August 1978 two specimens were taken in an alpine and Youngman 1975). Diamonds are peripheral
willow community (59°41’N, 136°34’W) at an eleva- records for the Arctic Shrew, dots are peripheral

records for the Tundra Shrew, and the star indicates
the Tundra Shrew records from northwestern British
Columbia.

tion of 1037 m adjacent to a stream. The community

‘Although the Tundra Shrew has a smaller skull, shorter tail,

and lighter ventral pelage than the Arctic Shrew, the taxon- . . sae
omy has been problematic and the Tundra Shrew has been tallvand various species|of bryophytes and Gramin

regarded as a subspecies of the Arctic Shrew, Sorex arcticus eae. Other plants included horsetails (Equisetum sp.),
tundrensis (Bee and Hall 1956; Banfield 1974) and as a Sedges (Carex sp.), yarrow (Achillea sp.), Burnet
distinct species, Sorex tundrensis (Jackson 1928; Youngman (Sanquisorba stipulata), Larkspur (Delphinium glau-
1975; C. G. van Zyll de Jong, personal communication). cum), Wormwood (Artemisia arctica), Indian Paint-

was dominated by willows (Salix sp.), about 1.5 m

3

94 THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 1—Comparison of means (ranges) in millimetres of selected cranial measurements of the five Tundra Shrews from
British Columbia with Tundra Shrews from Alaska and Arctic Shrews from the Northwest Territories and Yukon.

Sample Condylobasal Cranial Palatal Maxillary
size length breadth length toothrow length
Tundra Shrew
Haines Road,
British Columbia 5 18.1 (17.8-18.3) 8.9 (8.7-9.0) 7.3 (7.1-7.4) 6.3 (6.2-6.4)
Northeastern Alaska* 13 17.9 (17.2-18.5) 9.1 (8.7-9.4) 7.3 (7.0-7.5) 6.6 (6.4-6.8)
Arctic Shrew
Fort Norman, Northwest
Territories* 4 18.9 (18.7-19.2) 9.4 (9.1-9.5) 7.8 (7.6-8.1) 7.0 (7.0-7.0)
Yukon Crossing,
Yukon* l 20.3 10.4 8.6 7.8
* Measurements denoted by an asterisk are from Youngman (1975).
brush (Castilleja unalaschcensis), Bluebell (Mertensia Acknowledgments

paniculata), and Northern Grass-of Parnassus (Par-
nassia palustris).

Two specimens were collected in August 1979 ina
subalpine willow community (58° 48’N, 136°37’W) at
885 m. This community also bordered a stream, and
vegetation was similar to that of the alpine willow
community described above. Another Tundra Shrew
specimen was trapped in June 1979 in a subalpine
grass community (59° 49’N, 136° 36’W) at 885 m. This
habitat was dominated by willows, Wormwood,
sedges, Dwarf Birch (Betula glandulosa), and various
species of Gramineae and Compositae. Small mam-
mals trapped at Tundra Shrew sites included the
Dusky Shrew (Sorex monticolus), Long-tailed Vole
(Microtus longicaudus), Yundra Vole (Microtus
oeconomus), Meadow Vole (Microtus pennsylvani-
cus), Heather Vole (Phenacomys intermedius), and
Deer Mouse (Peromyscus maniculatus).

There are no records of the Tundra Shrew from
southeastern Alaska or southern Yukon (Hall and
Kelson 1959; Youngman 1975), and our specimens
extend the range of this species about 500 km south of
its known limits (Figure 1). Other populations of
Tundra Shrews may occur in alpine habitats in the
Coast Mountains of southern Alaska and northwest-
ern British Columbia. Rand (1954), Hoffman and
Peterson (1967), and Youngman (1975) hypothesized
that the Tundra Shrew was isolated in an unglaciated
region of Alaska and the Yukon (Beringian Refu-
gium) during the Wisconsin glaciation, and Young-
man (1975) noted that the present distribution of this
species is completely within the boundaries of Berin-
gia. Our records from northwestern British Columbia,
however, are 450-550 km south of the limits of the
Beringian Refugium (as mapped by Prest et al. 1968).
Presumably this population in British Columbia
resulted from postglacial dispersion.

We especially thank Stan van Zyll de Jong,
National Museum of Natural Sciences, Ottawa, for
verifying our identifications of the Tundra Shrews
and for providing us with his unpublished morpho-
metric data on the Sorex arcticus-tundrensis group.
Fieldwork was part of an M.Sc. research by Donald
Jones and was supported by research grants from the
University of Waterloo and from NSERC to John
Theberge. Adriana Jones is acknowledged for her
capable field assistance. We thank J. R. Tamsitt and
R. L. Peterson for critically reviewing the manuscript.

Literature Cited

Banfield, A. W. F. 1974. The mammals of Canada. Univer-
sity of Toronto Press, Toronto. 438 pp.

Bee, J. W., and E. R. Hall. 1956. Mammals of northern
Alaska on the arctic slope. University of Kansas, Publica-
tions in Natural History 8. 309 pp.

Hall, E.R., and K.R. Kelson. 1959. The mammals of
North America. Vol. 1. The Ronald Press Co., New York.
546 pp.

Hoffman, R. S.,and R. S. Peterson. 1967. Systematics and
zoogeography of Sorex in the Bering Strait area. System-
atic Zoology 16(2):127-136.

Jackson, H. H. T. 1928. A taxonomic review of the Ameri-
can long-tailed Shrews (genera Sorex and Microsorex).
United States Department of Agriculture Biological Sur-
vey. North American Fauna 51. 238 pp.

Prest, V. K., D. R. Grant, and V. N. Rampton. 1968. Gla-
cial map of Canada. Geological Survey of Canada, Map
1253 A.

Rand, A. L. 1954. The Ice Age and mammalian speciation
in North America. Arctic 7(1): 31-35.

Youngman, P. M. 1975. Mammals of the Yukon Territory.
National Museums of Canada. Publications in Zoology
10. 192 pp.

Received 26 May 1980
Accepted 16 July 1980

1981]

NOTES

95

Great Blue Heron (Ardea herodias) Colony
in the Peace-Athabasca Delta, Alberta

JOHN KRISTENSEN

LGL Limited, environmental research associates, 10110— 124 Street, Edmonton, Alberta T5N 1P6

Kristensen, John. 1981. Great Blue Heron (Ardea herodias) colony in the Peace-Athabasca Delta, Alberta. Canadian

Field-Naturalist 95(1): 95-96.

The northernmost colony of Great Blue Herons (Ardea herodias) was recorded in the Peace-Athabasca Delta, Alberta, in

1977.

Key Words: Great Blue Heron, Ardea herodias, bird colony, Peace- Athabasca Delta, Wood Buffalo National Park, Alberta.

On 24 August 1977, while conducting research on
Goldeye (Hiodon alosoides), 1 found an active Great
Blue Heron (Ardea herodias) colony in the Wood
Buffalo National Park section of the Peace-Atha-
basca Delta in northeastern Alberta, following infor-
mation given me by E. Courtorielle, a local trapper.
This represents the northernmost known breeding
locality of the species.

The heronry was situated on a small point of land
on the left bank of the Birch River (58°27’N,
112°17’W; Figure 1) about 100m from the river.
Dominant tree cover consisted of Balsam Poplar
(Populus balsamifera) and the understory was domi-

NORTHWEST
kame

NATIONAL

km 10 O 10 20 30 40

/

-_ ALBERTA

\,
\ |

e |
Calgary
ae SES)

Chenal des

Bey Quatre Fourches—=_
| y Vsileg
| | Ka Vane, i

| | ony z
| ae GREAT BLUE
( — | HERON COLONY ‘
@
ae Edmonton |
a |

nated by Red Osier Dogwood (Cornus stolonifera),
Low-bush Cranberry (Viburnum edule), and River
Alder (Alnus tenuifolia). Ground cover consisted
primarily of horsetails (Equisetum spp.) and grasses.

The heronry consisted of 12 nests, all located
12-18 m above ground in living Balsam Poplar trees.
Four trees contained two nests each; nests in the same
tree were as little as 4 m apart. The other nests were
constructed one toatree. All of the nests were located
withina circular area of about 30 m radius. One fallen
nest was slightly less than | m in diameter, con-
structed almost entirely of Balsam Poplar branches,
and lined with Balsam Poplar leaves. A dead Great

Fort Smith
—-— eas

~

bs , y
Fort Chipewyan

SO
—s —

|

}

| -

Po NWMaHOLVYSVS

FIGURE |. Location of the Great Blue Heron colony along the Birch River.

96 THE CANADIAN FIELD-NATURALIST

Blue Heron nestling was found on the ground beneath
another nest. Egg shell fragments were present
beneath several nests, as were abundant remains
(scales, caudal fins, vertebrae, and skull fragments) of
Goldeye, Northern Pike (Esox /ucius), and Walleye
(Stizostedion vitreum).

This colony, as far as is known (Godfrey 1966;
W. E. Godfrey, National Museum of Natural Scien-
ces, personal communication), is the northernmost
breeding locality of Great Blue Herons in Canada.
Jung (1930) was told (presumably by residents of Fort
Chipewyan) that Great Blue Herons nested along the
south shore of Lake Athabasca, but no heronry was
found at that time. The previous northernmost defi-
nite record of a heronry in Alberta was at Pelican
Lake at approximately 55° 50’N (Vermeer 1969). Most
Great Blue Heron colonies in Alberta have been
located in the southern half of the province (Vermeer
1969; Salt and Salt 1976). Colonies in other provinces
were also south of the colony described in this note
(Vermeer 1973; Mark 1976; Gray et al. 1980).

Residents of Fort Chipewyan have only recently
observed Great Blue Herons regularly in the Peace-
Athabasca Delta (personal communications with
local trappers and fishermen). The Great Blue Heron
was listed as an “accidental visitant” in Wood Buffalo
National Park by J. D. Soper (1935. Mammals and
birds of Wood Buffalo Park, northern Alberta and
District of Mackenzie. Canadian Wildlife Service
unpublished report. 365 pp.) and as a “rare vagrant” in
the Peace- Athabasca Delta by Hohn (1973). A single
bird was observed along the Chenal des Quatre
Fourches by Jung (1930). Great Blue Herons have
been observed in the Birch River area from 1975 to
1979, and as far north as the Salt River (59° 52’N,
112°03’W) (E. Kuyt, Canadian Wildlife Service, per-
sonal communication). I observed numerous single
Great Blue Herons along the Chenal des Quatre
Fourches during 1975 and 1980, and several single
birds along the Prairie River in 1975 and 1976.

On the basis of the suggestion by Vermeer (1969)
that herons’ excrement contributes to the death of
trees that they nest in, I concluded that the Birch River
heronry, composed of nests in live trees, is probably
relatively recent. Although the Balsam Poplar trees
and understory were covered with excrement, they
appeared to be healthy and there was no evidence of
nettles, etc., plants typically found where the soil has
been enriched with excrement.

Vol. 95

Great Blue Herons and their nesting colonies are
conspicuous and vulnerable to disturbance by
humans. Industrial and lakeshore residential expan-
sion may have diminished both the number and size of
Great Blue Heron colonies in Alberta (Salt and Salt
1976), and Vermeer (1969) indicated that some Great
Blue Herons in Alberta have moved to new locations
as a result of human disturbance. The northern nest-
ing of Great Blue Herons in Alberta, shown by this
record may be a result of increasing human develop-
ment and disturbance in more settled areas, coupled
with ideal feeding habitat for Great Blue Herons in the
Peace-Athabasca Delta. It is also possible that the
apparent recent northward expansion of Great Blue
Heron colonies in Alberta and other provinces is an
artifact of increasing northern field work.

Thanks are extended to A. Courtorielle for assisting
me in locating the heronry; to W. E. Godfrey, E. Kuyt,
W. J. Richardson, R. E. Salter, S. G. Sealy, and C. E.
Tull for reviewing an earlier draft of this note; and to
K. Bruce for preparing the figure.

The heronry was located while the author was
under contract with LGL Limited to Fisheries and
Environment Canada.

Literature Cited

Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203. 428 pp.

Gray, P.A., J. W. Grier, G.D. Hamilton, and D. P.
Edwards. 1980. Great Blue Heron colonies in northwest-
ern Ontario. Canadian Field-Naturalist 94: 182-184.

Hohn, E. O. 1973. The birds of the Peace- Athabasca Delta
and of the Lake Athabasca region. Appendix C. /n Peace-
Athabasca Delta Project. Ecological Investigations. Vol.
2. Technical Appendices. 32 pp.

Jung, C. S. 1930. Notes on birds of the delta region of the
Peace and Athabasca rivers. Auk 47: 533-541.

Mark, D.M. 1976. An inventory of Great Blue Heron
(Ardea herodias) nesting colonies in British Columbia.
Northwest Science 50: 32-41.

Salt, W. R., and J. R. Salt. 1976. The birds of Alberta.
Hurtig Publishers, Edmonton. 498 pp.

Vermeer, K. 1969. Great Blue Heron colonies in Alberta.
Canadian Field-Naturalist 83: 237-242.

Vermeer, K. 1973. Great Blue Heron and Double-crested
Cormorant colonies in the prairie provinces. Canadian
Field-Naturalist 87: 427-432.

Received 13 March 1980
Accepted 15 August 1980

1981

NOTES

97

An Unusually Small American Eel (Anguilla rostrata)

from the Lake Superior Drainage!

PHILIP A. COCHRAN

Department of Entomology, Fisheries and Wildlife, University of Minnesota, Saint Paul, Minnesota 55108
Present address: Laboratory of Limnology, University of Wisconsin, Madison, Wisconsin 53706

Cochran, Philip A. 1981. Anunusually small American Eel (Anguilla rostrata) from the Lake Superior drainage. Canadian

Field-Naturalist 95(1): 97-98.

An unusually small specimen (total length 157 mm) of the American Eel (Anguilla rostrata) was collected from the Lake
Superior drainage. At least some eels in Lake Superior must migrate directly into the lake at a small size. The specimen in
question is much smaller than the size previously inferred for eels migrating into the Great Lakes at Lake Ontario.

Key Words: Anguilla rostrata, American Eel; Lake Superior, Great Lakes, Anguillidae, geographic distribution, body size,

migrations.

A specimen of the American Eel, Anguilla rostrata,
was collected on 11 February 1978 from the Black-
hoof River, a tributary of the Nemadji River in the
Lake Superior basin (Carlton County, Minnesota,
46° 33’N, 92°28’W). The stream at the site of capture
was ~ 0.5 m deep and 5-6 m wide, with open water at
the center and shelf ice along the banks. Total length
of the specimen was 157 mm. It was givento James C.
Underhill to be deposited in the University of Minne-
sota collection.

American Eels have been present in the Upper
Great Lakes in small numbers for some time (Hubbs
and Lagler 1964), including Lake Superior (Moore
and Braem 1965; Eddy and Underhill 1974). It is
generally presumed that they have gained access to the
Upper Great Lakes through the St. Lawrence Seaway
(Eddy and Underhill 1974) or the Welland Canal
(Eales 1968; Scott and Crossman i973). The possibil-
ity of eels also gaining access to the Great Lakes from
the Mississippi drainage via the Chicago Drainage
Canal has apparently been overlooked by recent
authors, although Hubbs and Lagler (1964) stated
that several fish species may have used it to enter Lake
Michigan. As early as 1815, Clinton (see Webster
1980) discounted the possibility of young eels migrat-
ing upstream against the rapid currents of the Missis-
sippi and Illinois rivers to take advantage of the then
seasonal connection between the Lake Michigan and
Mississippi drainages. Nevertheless, eels penetrate the
upper Mississippi River and its tributaries as far north
as Minnesota (Eddy and Underhill 1974), and speci-
mens have been collected from the Illinois River as
recently as 1974 (Sparks and Starrett 1975). Sparks
and Starrett (1975) commented onan improvement of
water quality in the upper Illinois River and Chicago

'Paper Number 11123, Scientific Journal Series, Minnesota
Agricultural Experiment Station, St. Paul, Minnesota 55108.

Canal resulting from aeration and improved waste
treatment. This would conceivably facilitate fish
movements.

The significance of the Blackhoof River record is
that it apparently represents by far the smallest Amer-
ican Eel yet taken in the Lake Superior basin. The
next smallest specimens that could be located on
record measured 373 mm (J. C. Underhill, personal
communication) and 406 mm (Moore and Braem
1965). The Blackhoof specimen is evidence that at
least some American Eels migrate directly into Lake
Superior and reach the western limit of their Great
Lakes distribution at a small size, much smaller than
the 200- to 300-mm length inferred by Hurley (1972)
for eels entering Lake Ontario. It might previously
have been supposed that the few large eels captured in
Lake Superior had merely wandered into the lake
after attaining their size elsewhere, because American
Eels are capable of long movements (e.g., Viladykov
1956) and have been known to make movements
apparently not associated with sexual maturity or
reproduction (Smith and Saunders 1955).

Capture of the Blackhoof specimen was fortuitous.
It was collected with electrofishing gear during the
marking run of a mark-and-recapture population
estimate for Brown Trout (Sa/mo trutta) and Rain-
bow Trout (Sa/mo gairdneri). In the water it was
mistaken for a larval lamprey, which were being
stunned in some numbers and which it resembled in
general form and coloration. It was set aside for later
scrutiny, the only “lamprey” so collected. After its true
identity was revealed, however, close attention was
paid to the lampreys stunned during the recapture run
over the same stretch of stream the next day, but no
additional eels were observed. Fisheries personnel
working along Lake Superior tributaries should
nevertheless be aware of the possibility of securing
additional specimens of small American Eels which

98 THE CANADIAN FIELD-NATURALIST

might easily be overlooked because of a superficial
resemblance to lampreys.

George C. Becker (Department of Biology, Univer-
sity of Wisconsin—Stevens Point), E. J. Crossman
(Royal Ontario Museum), and James C. Underhill
(Department of Zoology, University of Minnesota,
Minneapolis) kindly made available their records of
American Eels from Lake Superior. James C. Under-
hill and Thomas F. Waters read earlier drafts of the
manuscript.

Literature Cited

Clinton, D. 1815. Some remarks on the fishes of the west-
ern waters of the State of New York, in a letter to S. L.
Mitchell. Transactions of the Literary and Philosophical
Society of New York. 1: 493-501. (Not seen, reprinted with
interspersed commentary in Webster (1980).)

Eales, J.C. 1968. The eel fisheries of eastern Canada.
Fisheries Research Board of Canada Bulletin 166. 79 pp.

Eddy, S., and J. C. Underhill. 1974. Northern fishes. Uni-
versity of Minnesota Press, Minneapolis. 414 pp.

Hubbs, C. L., and K. F. Lagler. 1964. Fishes of the Great
Lakes region. University of Michigan Press. 213 pp.

Vol. 95

Hurley, D. A. 1972. The American Eel (Anguilla rostrata)
in eastern Lake Ontario. Journal of the Fisheries Research
Board of Canada 29: 535-543.

Moore, H.H., and R.A. Braem. 1965. Distribution of
fishes in United States streams tributary to Lake Superior.
United States Fish and Wildlife Service, Special Scientific
Report Fisheries Number 516. 61 pp.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada Bulletin
184. 966 pp.

Smith, M. W., and J. W. Saunders. 1955. The American
Eel in certain freshwaters of the maritime provinces of
Canada. Journal of the Fisheries Research Board of Can-
ada 12: 238-269.

Sparks, R. E., and W. C. Starrett. 1975. An electrofishing
survey of the Illinois River 1959-1974. Illinois Natural
History Survey Bulletin 31: 317-380.

Vladykov, V. D. 1956. Fish tags and tagging in Quebec
waters. Transactions of the American Fisheries Society 86:
345-349.

Webster, D. A. 1980. De Witt Clinton’s “. . . Fishes of the
western waters of the State of New York” reexamined.
Fisheries 5(2): 5-12.

Received 5 March 1980
Accepted 27 May 1980

Panicled Knapweed (Centaurea paniculata: Compositae)

New to Eastern Canada

DANIEL F. BRUNTON! and PAUL D. PRATT?

'Southwick Drive, R.R. 3, Manotick,Ontario KOA 2NO

2Windsor Department of Parks and Recreation, 2450 McDougall Street, Windsor, Ontario N8X 3N6

Brunton, Daniel F., and Paul D. Pratt. 1981. Panicled Knapweed (Centaurea paniculata: Compositae) new to eastern

Canada. Canadian Field-Naturalist 95(1): 98-99.

The discovery of a small station of Panicled Knapweed, Centaurea paniculata, in Essex County, Ontario, constitutes the first
locality for this species in eastern Canada and the second for North America. This introduced species is very rare in Ontario.

Key Words: Centaurea paniculata, Compositae, Ontario, rare flora, new records, eastern Canada.

The Panicled Knapweed (Centaurea paniculata) is
a rare introduction in Canada and was previously
known only from Vancouver Island, British Colum-
bia, before the recent discovery of a station in Essex
County, Ontario. It is a native of Europe, ranging
across the Mediterranean region from the Iberian
Peninsula to Italy and Switzerland. Although first
collected in Victoria in 1893 by John Macoun and
noted occasionally thereafter on southern Vancouver
Island, it has not become well established nor has it
been found elsewhere in British Columbia (Moore
and Frankton 1974). It is not listed in Montgomery
(1957), Boivin (1967), Scoggan (1979), Gleason
(1952), Hitchcock et al. (1955), or Fernald (1950),

indicating that this species is largely unknown as an
element of the North American flora and is apparently
unknown in the United States. A specimen search in
the DAO, CAN, and MICH herbaria revealed no
additional records, nor did an examination of the
large number of Essex County specimens in the per-
sonal herbarium of W. Botham.

The C. paniculata — C. maculosa group of Centau-
rea is complex and its species have been combined
under various specific and subspecific rankings. The
Essex County material keys out to C. paniculata ssp.
esterellensis (Burnat) Dostal (Dostal 1976), a taxon of
southeastern France. The validity and relationships of
this taxon are not clear.

1981

The Ontario station is located on disturbed ground
near a railway spur line. No additional plants were
noted in the area; however, a thorough search was not
conducted. Its means of introduction is unknown,
although the railway line is an obvious possibility. It
seems quite unlikely that these plants originated from
the Victoria population, given the species’ inability to
become well established even there. It is more likely
that the Essex County station represents an independ-
ent introduction from Europe. The species is also
known in Great Britain (Fitter and Fitter 1974) where
it was probably introduced. It seems that the Ontario
station may have originated from any of a number of
areas.

The label data for the new collection are as follows:
1.75 km NE of Amherstberg (sic), Anderdon Twp.,
Essex Co., Ontario. 42°07’N, 78° 05’W. On overgrown
road across bare limestone in old quarry, growing in
open with Melilotus spp. 2 September 1979. D. F.
Brunton & P. D. Pratt 1933 (DAO).

On the basis of the three or four plants noted at the
Essex County station, C. paniculata would seem to be
very rare in eastern Canada.

We greatly appreciate the assistance of C. Frankton
for his critical review of the determination, his com-
ments on Centaurea in general, and for his review of
an earlier draft of the manuscript. Our thanks also go
to the curators of DAO and CAN for their assistance.
A. A. Reznicek was kind enough to examine the Uni-

NOTES 99

versity of Michigan collection for us and his assistance
is also appreciated.

Literature Cited

Boivin, B. 1967. Enumération des plantes du Canada. Pro-
vancheria Numero 6, Université Laval, Québec.

Dostal, J. 1976. Centaurea L. In Flora Europaea, Volume
4. Edited by T. G. Tutlin, V. H. Heywood, N. A. Burges,
D. M. Moore, D. H. Valentine, S. M. Watters, and D. A.
Webb. Cambridge University Press, Cambridge. pp.
254-301.

Fernald, M. L. 1950. Gray’s manual of botany (eighth edi-
tion). American Book Company, New York.

Fitter, R., and A. Fitter. 1974. The wildflowers of Britain
and Northern Europe. William Collins Sons & Company
Ltd., London.

Gleason, H. A. 1952. The new Britton and Brown illus-
trated flora of the northeastern United States and adjacent
Canada. New York Botanical Garden, New York.

Hitchcock, C. L., A. Cronquist, M. Ownbey, and J. W.
Thompson. 1955. Flora of the Pacific Northwest. Uni-
versity of Washington Press, Seattle.

Montgomery, F. H. 1957. The introduced plants of Onta-
rio growing outside of cultivation (Part II). Transactions
of the Royal Canadian Institute 32(1): 1-34.

Moore, R. J., and C. Frankton. 1974. The thistles of Can-
ada. Monograph 10, Agriculture Canada, Ottawa.

Scoggan, H. J. 1979. The flora of Canada, Part 4. National
Museum of Canada, Ottawa, Ontario.

Received 15 April 1980
Accepted 21 July 1980

Ruby-crowned Kinglets (Regulus calendula)
Feeding a Brown-headed Cowbird (Molothrus ater)

PETER C. BOXALL

Department of Biology, University of Calgary, Calgary, Alberta
Present address: Provincial Museum of Alberta, 12845-102 Avenue, Edmonton, Alberta TSN 0M6

Boxall, Peter C. 1981. Ruby-crowned Kinglets (Regulus calendula) feeding a Brown-headed Cowbird (Molothrus ater).

Canadian Field-Naturalist 95(1): 99-100.

A pair of Ruby-crowned Kinglets (Regulus calendula) were observed feeding a fledged Brown-headed Cowbird (Molothrus
ater). This is the third report of successful brood parasitism upon Ruby-crowned Kinglets, despite a number of observations

of cowbird eggs in kinglet clutches.

Key Words: Regulus calendula, Molothrus ater, brood parasitism, British Columbia.

On | August 1979, I observed a young Brown-
headed Cowbird (Molothrus ater) being fed by a pair
of Ruby-crowned Kinglets (Regulus calendula) at the
south end of Bridge Lake (51° 29’N, 120° 42’W), about
80 km N of Kamloops, British Columbia. I was
attracted to the scene by the loud begging calls of the

cowbird. Although cowbirds are usually noisy and
aggressive towards their foster parents (Lavers 1974),
this particular bird seemed more vocal than cowbirds
I have seen attended by Yellow Warblers (Dendroica
petechia) or Tennessee Warblers (Vermivora pere-
grina). |continued to watch the trio to see whether the

100

much smaller kinglets were having problems supply-
ing enough food to the cowbird. From 16:50 to 17:30
(PDT), I observed four instances of both kinglets
feeding the cowbird virtually simultaneously, and 14
feedings by a single kinglet. The cowbird begged per-
sistently and chased its fosterers for 1-2 m several
times. I returned to the area a number of times over
the next 3 d, and I found one kinglet each time but did
not find or hear a begging fledgling.

This observation is the third report of successful
parasitism by the Brown-headed Cowbird upon the
Ruby-crowned Kinglet. The latter were initially con-
sidered very rare victims of cowbirds owing to the
disparity in size between the two species (Friedmann
1929). Friedmann et al. (1977), however, reported 14
cases of cowbird parasitism on the Ruby-crowned
Kinglet, and suggested that it may bea regularif nota
favorite host. Seven records involved the western race
Regulus calendula cineraceus; six from California and
one from British Columbia (Grant 1965). Apparently,
12 of the 14 cases involved cowbird eggs seen in kinglet
nests. Only Friedmann (1963) and Mans and Peyton
(1962) describe Ruby-crowned Kinglets feeding
fledged cowbirds, and Lavers (1974) observed
Golden-crowned Kinglets (Regu/us satrapa) feeding a
cowbird.

Kinglet clutches commonly contain 5-I1 eggs.
Since five kinglet eggs are about 150% of the volume
of a cowbird egg, and egg volume is an approximation
of the energy needs of nestlings (Murton and West-
wood 1977), five or more kinglet nestlings require
more energy than one cowbird. This, and the observa-
tions of kinglets feeding cowbirds, suggest that kin-
glets may not have problems meeting the energy
requirements of a cowbird. Considering the numerous

THE CANADIAN FIELD-NATURALIST

Vol. 95

cases of cowbird parasitism known for other host
species, however, the 15 records of parasitism do not
suggest that kinglets are important hosts. Factors
such as well-concealed spherical nests high above the
ground (Bent 1949) may give cowbirds few opportuni-
ties to parasitize kinglets.

I thank Dan Busby, Gordon Holton, and M. Ross
Lein for commenting on early drafts of this manus-
cript. I gratefully acknowledge Stephen Rothstein for
providing me with an important reference, and also
for his valuable comments.

Literature Cited

Bent, A.C. 1949. Life histories of North American
thrushes, kinglets, and their allies. United States National
Museum Bulletin Number 196. 452 pp.

Friedmann, H. 1929. The cowbirds: a study in the biology
of social parasitism. C. C. Thomas, Springfield, Illinois.
Friedmann, H. 1963. Host relations of the parasitic cow-
birds. United States National Museum Bulletin Number

233.

Friedmann, H., L. F. Kiff, and S. I. Rothstein. 1977. A
further contribution to knowledge of the host relations of
the parasitic cowbirds. Smithsonian Contributions to
Zoology, Number 235.

Grant, J. 1965. /n Great Basin, Central Rocky Mountain
Region. Audubon Field Notes 20: 589.

Lavers, N. 1974. Three more cases of White-crowned Spar-
rows parasitized by Brown-headed Cowbirds. Auk 91:
829-830.

Mans, M.,andG. S. Peyton. 1962. /n Middle Pacific Coast
Region. Audubon Field Notes 16: 505-506.

Murton, R. K.,and N. J. Westwood. 1977. Avian breeding
cycles. Clarendon Press, Oxford.

Received 19 March 1980
Accepted 31 May 1980

Northern Record for Three-flowered Avens

(Geum triflorum) in Manitoba

H. L. MUNRO and D. H. MUNRO

Department of Geography, Brandon University, Brandon, Manitoba R7A 6A9

Munro, H.L., and D. H. Munro. 1981. Northern record for Three-flowered Avens (Geum triflorum) in Manitoba.

Canadian Field-Naturalist 95(1): 100-101.

We have documented the occurrence of Geum triflorum (Three-flowered Avens) at Flin Flon, 350 km N of its previously
known location in Manitoba. Plants were similar in size to those described else where.

Key Words: Geum triflorum (Three-flowered Avens), distribution, Flin Flon, Manitoba.

Geum triflorum (Three-flowered Avens, Old Man’s
Whiskers, Lion’s beard) is anerect perennial herb with

flowering stems 15—40 cm tall, pinnately compound
basal leaves 10-20 cm long, and one to five nodding

1981

flowers 12-20 mm in diameter (Scoggan 1978; Loo-
man and Best 1979). Three-flowered Avens are com-
mon spring flowers on the prairies.

On 10 June 1978, a Three-flowered Avens was
found growing on the north-facing slope of a railroad
right-of-way near Flin Flon, Manitoba (54°47’N,
101°52’W). Flowering stems were 27 cm tall, basal
leaves were 15 cm long, and flowers averaged 15 mm
across. On 23 June 1979, the site was revisited and two
specimens were located, one of which was collected
and deposited in the herbarium at Brandon
University.

Other herbs which grew along the right-of-way were
Maianthemum canadense (Wild Lily-of-the-valley),
Smilacina trifolia (YThree-leaved Solomon’s-seal),
Heuchera richardsonii (Alumroot), and Carex spp.
(sedges). Intermittent clumps of Cornus stolonifera
(Red Osier Dogwood) and Rosa acicularis (Prickly
Rose) were also present. The right-of-way was bor-
dered by Picea mariana (Black Spruce) forest.

The previous northernmost occurrence of Three-
flowered Avens in Manitoba was recorded along
roadsides and in clearings in the Duck Mountains
(approximately 51°30’N, 100° W), 350 km S of Flin
Flon (Scoggan 1957). The species has also been
recorded in prairie communities at Prince Albert
National Park, Saskatchewan (53°45’N, 106°03’W)
and in roadside meadows at Wood Buffalo Park,
Alberta (59° 57’N, 112° 17’ W) (J. M. Gillett, National
Museum of Natural Sciences, personal communica-
tion) as well as on shallow residual soils in limestone
outcrop areas beside the Enterprise- MacKenzie River
Highway, Northwest Territories (approximately
61°N, 117° W) (Porsild and Cody 1968).

NOTES 101

Presence of Three-flowered Avens in the Flin Flon
area does not necessarily represent a natural range
extension. The vegetation along the right-of-way is
characteristic of moist parkland or woodland margins
(Looman and Best 1979). Although Three-flowered
Avens seeds are probably capable of long-distance
wind dispersal, the absence of other species character-
istic of prairie communities suggests that Three-
flowered Avens was introduced to the area by train.
The disclimax characteristics of the limestone gravel
and open environment of the railway allowance pro-
vided suitable habitat for the establishment of this
non-boreal species.

Our measurements indicate that Three-flowered
Avens does not decrease in vigor in northerly lati-
tudes. It is also able to overwinter and maintain vital-
ity in the boreal zone provided that suitable habitat is
available.

We express thanks to R. C. Rounds of Brandon
University for reviewing the manuscript.

Literature Cited

Looman, J., and K. F. Best. 1979. Budd’s flora of the Can-
adian Prairie Provinces. Research Branch, Agriculture
Canada, Ottawa. 863 pp.

Porsild, A. E., and W. J. Cody. 1968. Checklist of the vas-
cular plants of continental Northwest Territories, Canada.
Canada Department of Agriculture, Ottawa. 102 pp.

Scoggan, H. J. 1957. Flora of Manitoba. National
Museum of Canada, Ottawa. 619 pp.

Scoggan, H.J. 1978. The flora of Canada (Part 3).
National Museum of Natural Sciences, Ottawa. 1115 pp.

Received 15 May 1980
Accepted 14 August 1980

FARRELL E. BANIM, O.M.I., 1902-1979

On8 April 1979, with the death of Father Farrell E.
Banim, O.M.I., Professor Emeritus of Biology,
Carleton University, The Ottawa Field-Naturalists’
Club lost one of its oldest and most faithful members.
As far back as 1957, in his Christmas letter, Father
Banim described himself as “one of the few remaining
real old-timers.”

He was born in Dublin, Ireland, on 17 October
1902, the son of Farrell P. and Mary J. Banim, and
thus a descendant of John and Michael Banim, the
19th-century Irish novelists (a connection which
meant a great deal to him). After his early schooling in
Dublin, he was graduated in 1924 from the National
University of Ireland. During this period he entered
the Novitiate of the teaching and missionary order,
the Oblates of Mary Immaculate, and was ordained in
Rome in 1927. In Rome, he pursued his studies in
theology and philosophy and, after gaining his Licen-
tiate, proceeded to Cambridge University in 1928 to
earn his B.Sc. and M.Sc. degrees. For many years
thereafter, and often facing great difficulties, he con-
tinued postgraduate work at the Catholic University
of America (Washington, D.C.), and at Fordham and
Columbia universities. In 1957 and 1958 he did
research in Palaeozoology at the University of Paris,
and during the summer of 1957, he worked at the
Mediterranean Biological Station.

After his graduation from Cambridge in 1931, he
was sent to what was then Ceylon (now Sri Lanka), to
teach at St. Joseph’s College, Colombo.

In the meantime, in Ottawa, the Oblate Fathers,
with more faith than foresight, had founded St.
Patrick’s College, planned to provide a liberal arts
education from first-form high school to university
graduation for English-speaking students. The college
opened its doors on the eve of the stock market crash
of 1929 and the years of the Great Depression. Witha
commitment of $660 000, a vast sum in those days, it
was imperative that a staff of unpaid teachers be re-
cruited from the members of the Order. Among the
gifted and dedicated young men to staff the college in
those early years was Father Banim, who came to St.
Patrick’s in 1933 as head of the Science Department,
and who was appointed Dean of Arts the following
year, forbidding tasks for a man barely 30 years of age.

It was typical of Father Banim that he lost no time
in marshalling all available resources. At the college
on Echo Drive he built, with his own hands, the work
benches, shelves, and cupboards in the first science
laboratory, as well as assisted with the installation of
the old Bunsen burners and laboratory plumbing. In
the same year, he discovered The Ottawa Field-
Naturalists’ Club, which he immediately joined. The

writer, who was, at the time, a member of the Excur-
sions Committee, well remembers a hurried call from
the late W. H. Lanceley. “I’ve a note here to call a
Father Banim. I don’t know how you address a priest
so I’m giving him your number.” “Neither do I,” an-
swered a very scared committee member. (Despite the
extensive work of Frére Marie-Victorin, Frére Alex-
andre, Father (later Bishop) Vachon, and others, it
was still accepted locally that priests and science did
not mix.) The call came through, however, with a
warm, genial “Hello, could you tell me something
about the field-naturalists’ club?” He turned up at the
next outing and thus began an association with the
club and a friendship with both W. H. Lanceley and
myself that lasted unbroken through the years.
Nobody knew quite what to expect, but we who had
visualized a stern, black-robed cleric were totally
unprepared for the unorthodox figure riding his bicy-
cle, in old clothes and a battered felt hat.

In 1936 he was first elected to Council and, with the
exception of his years in Africa, served until the end of
1966. When pressed to continue, he answered “No. It
is time for the young to take over. We have had our
day.” He had been Second Vice-President in 1941 and
1942, First Vice-President in 1943 and 1944, and Pres-
ident in 1945 and 1946. He carried out his duties with
the enthusiasm that characterized all that he did. In
December 1971, he was elected an Honorary Member.

In 1936 and 1937, and again in 1940 and 1941, he
took part in the Christmas Bird Census, an activity
curtailed only by increasing work at the college.
Beginning in 1936, the club held regular winter lec-
tures at St. Patrick’s, a program that lasted for the
following nine years. For many years the college was
almost a headquarters for the club, and council meet-
ings, with few exceptions, were held in the Common
Room from 1946 until 1957. At the symposium on the
“Natural history of McKay Lake based on field excur-
sions,” held on 16 November 1943, there was passed a
special vote of thanks to the college “for the use of
rooms and equipment.” Father Banim’s personal con-
tribution to the program was contained in his “Review
of the freshwater biology of McKay Lake.” Indeed it
could be said that Father Banim and St. Patrick’s
College were almost synonymous. At this time he was
especially interested in the algae, and using the bel-
lows of an old-folding kodak, photographed a
number of local species. One wonders what became of
those pictures.

Science teaching during those times of depression
and the war years that followed was a vastly different
situation from the well-equipped laboratories and
libraries of present-day schools and colleges. Micro-

102

1981

OBITUARY: FARRELL E. BANIM 103

Farrell E. Banim, O.M.I.

scopes were costly and difficult to come by, sources of
basic information, particularly of local biology, were
much less extensive than those now available. Stras-
burger, Tansley, Godwin, Bower, Seward, Borradaile,
and Ward and Whipple held pride of place on the
shelves, and Father Banim’s own books were always
available to student and club member alike. There are
still members who will recall his patched laboratory
coat (“I doall my own mending.”). They were the days
when straight razors and carrots were used to cut
sections, and tubs of dogfish preserved in formalde-
hyde emitted odors that drifted through the corridors
to the disgust of the nonscientific academics. The
writer still recalls the shocked and disapproving looks
of Ottawa’s elite as Father Banim waited on the front

steps of the stately Chateau Laurier Hotel one lovely
June day, burdened with the skull, ribs, and leg-bones
of a cow which he had found in an old pasture near
Pink Lake. They were the days of the club’s bus excur-
sions, and a member, most likely W. H. Lanceley, had
agreed to transport Father Banim and his treasures
back to the college. Those bones were part of the
collection for many years.

The war years were times of great sadness for him as
he watched his young students leave the predomi-
nantly male college for service overseas, some never to
return.

In 1936, 1937, and again in 1941, he addressed the
club on “Mosses, lichens and algae” (illustrated with
his own slides), “How to know the flowers,” and “The

104

cultural value of nature study.” There are still a few of
us who remember his reading the passages on
“Mosses” and “Lichens” from Ruskin’s Frondes
Agrestes, his reciting of Padraic Pearse’s hauntingly
lovely The beauty of the world hath made me sad, and
the lines from Wordsworth quoted in his tribute to the
late Rowley C. Frith in The Canadian Field-
Naturalist Volume 89(4), 1975,

For I have learned to look on nature

Not as in the days of thoughtless youth,

But hearing oft’ times

The still sad music of humanity.

In 1939 he organized study groups which met at the
college on alternate Saturdays during the winter. The
first of these was the bird group which he led with
J. M. Robinson. In 1943 he worked with the moss
group, at the same time developing a new interest in
arachnology.

Five years later he was appointed Chairman of the
Constitution and By-laws Committee, to carry out a
long-overdue revision. This was completed and
approved by Council the following year and appeared
as “The Ottawa Field-Naturalists’ Club By-laws —
Revised 1949” in The Canadian Field- Naturalist
Volume 64(2), 1950. When the club celebrated a
delayed 75th anniversary in 1955, he served as Chair-
man of the Council of Former Presidents to arrange
the anniversary banquet held on 8 November at Lans-

downe Park.
The greater part of 1957 and 1958 was spent at the

University of Paris and at the Mediterranean Biologi-
cal Station. He described his experience ina letter toa
friend as “a wonderful year in France,” and on his
return to Ottawa in December, spoke of his work at
the club’s annual dinner. Four years later, he was sent
by his Order to Roma, Basutoland, to take over the
administration of St. Pius XII University College. It
was a difficult assignment, demanding organizing and
teaching ability as well as the skills of a diplomat. He
returned to Canada in 1964, his health impaired by the
burden of work and conflict imposed upon him. Dur-
ing his African years, he met Mary and Louis Leakey
and spent his brief vacations working at the famous
site at Olduvai Gorge, Tanzania. It is not certain
whether he had met Teilhard de Chardin, but certainly
in his later years his teaching was influenced by the
thought of the great Jesuit scientist.

Back in Ottawa he resumed his work at St. Patrick’s
College and later at Carleton University when St.
Patrick’s joined Carleton in 1967. In 1973 he was
appointed Professor Emeritus of Biology, and con-
tinued to lecture on physical anthropology until
within a few weeks of his death. Unfortunately, the
pressure of his life left little time for writing, and apart
from detailed reports to his superiors and an extensive
personal correspondence, he seldom recorded his

THE CANADIAN FIELD-NATURALIST

Voltes

work and observations. His review of Pieter W. Fos-
burgh’s The natural thing in The Canadian Field-
Naturalist Volume 83(3), 1969, was typical of his free,
easy style, even to the characteristic comment at the
end — “the price, $4.75, is not a bit too much.”

He was first and foremost a teacher, patient, under-
standing but demanding, and this applied to his study
groups as wellas to his students. Perhaps the influence
of his Banim forbear, John, who was artist and draw-
ing teacher as wellas writer, had something to do with
his insistence upon all work being accompanied by
careful and accurate drawings. A master of the Eng-
lish language, both written and spoken, he refused to
tolerate careless or sloppy notes. (It might be added
here that he was also fluent in his native Gaelic,
French, and Spanish, and toward the end of his life
had hoped to retire to the Oblate Mission at Lima,
Peru, where he had already spent some time, with the
intention of perfecting his knowledge of Spanish).

But his students came to know another side of him
as well. He had a habit of beginning class with one of
his endless array of jokes, showing his sharp Irish wit.
He also loved to tell stories about personal experien-
ces particularly in relation to his archaeological digs.
What his students will probably remember most
about his classes, however, is his longtime friend
“Mrs. Ples,” short for Plesianthropus, one of the pre-
historic skull models used in the anthropology labs.
This nickname was an example of his efforts to bring
familiarity to a discipline that can be very complex. By
his own words “She will live on in the minds of my
students long after 1am gone.” Among his friends and
students there will long survive a legion of “Banim
stories.” One such concerns an incident in the class-
room when he was being needled by a student to
express his views on the relationship of religion and
the theory of evolution, which was quite irrelevant to
the subject of his lecture. In exasperation he removed
his clerical collar, flung it across the room, and
exclaimed, “Here I am a professor of biology. If you
want to discuss religion, come to my office after
class!” And there was the winter when he agreed to
board in the laboratory, ina large cage constructed by
himself, a young crow found on a club excursion to
Carlsbad Springs, raised by a member in Westboro,
but still too young to release. The results were disas-
trous in terms of the decibel level of the bird’s daily
“hymn to the dawn.” That year he painted a delightful
Christmas card showing the crow, and bearing the
limerick,

I’m a sleek bit of mischief, Jim Crow,
Whose mind is supposed to be slow,
So they sent me to college
To brush up my knowledge
But my heart is in Westboro.
The unremitting financial struggles of the college

1981 OBITUARY: FARRELL E. BANIM 105

and the inevitable changes, along with the disap- epitaph for him than Stevenson’s immortal lines . . .
pointments of his African experience, all took their To go on forever and fail and go on again,

toll but failed to daunt him. What to many would have And be mauled to the earth and arise,

been obstacles, to him were challenges. St. Patrick’s And contend for the shade of a word

ceased to exist at the end of the academic year gud shine oes: cawidaytiieseyes:

Sia : : : With the half of a broken hope for a pillow at night
1978-1979 and with it, Father Banim. A final painful Thai somaeingnn dae mabe 16 dhe mabe

illness forced him to relinquish his course in physical And the smooth shall bloom from the rough: . . .
anthropology only a few weeks before the end of term
and the closing of the college to which he had devoted PEGGY WHITEHURST KURATA

so many years of his life. There could be no better 478 Edison Avenue, Ottawa, Ontario K2A 1T9

News and Comment

Experimental Oil Spills off Baffin Island Approved

Approval for experimental oil spills to take place in
the nearshore waters of Baffin Island during the
summers of 1980 and 1981 was announced in June
1980. The spills are part of a 4-yr study to assess the
impact of oil spills on arctic coasts and to test cleanup
measures. The study is to be conducted near the
northern tip of Baffin Island on Cape Hatt, 70 km
from Pond Inlet, the nearest settlement. All partici-
pants in the study are satisfied that the environmental
effects of the oil spills will be confined to a very small
area.

During the summer of 1980 until freeze-up in
October, scientists were to gather baseline data and
also to release a small amount of oil on a limited
section of shoreline to begin studying the long-term
fate of oil on arctic beaches. The major part of the
study, however, will take place in the summer of 1981.
Up to 275 barrels of oil will be discharged in the
nearshore waters of several small bays. About 100
barrels of crude oil will be spilled into one bay, and a
similar volume of crude oil with a dispersant, which
breaks up oil and mixes it into the water, will be
discharged into another bay. A third bay will remain
uncontaminated as a control.

Prairie Wildlife Interpretation Centre

“The Prairie Wildlife Interpretation Centre is
designed to provide the travelling public an opportun-
ity to experience the uniqueness of the prairies,” said
federal Environment Minister, John Roberts. The
center is located on the Trans-Canada Highway near
Webb, Saskatchewan.

With summer students interpreting the region’s
ecology and the history of man’s use of the grasslands,
visitors will be shown the full range of grassland char-
acteristics on trails that lead through a wheat field,
creek valley, tree grove, and marsh. Some 45 species of
mammals, more than 450 species of plants, and up to
270 species of birds have been identified on the 1 100-
acre (~ 445-ha) site by the Canadian Wildlife Service.

In addition to an abundance of natural life, the site

In an intensive sampling and analysis program that
will continue through 1983, scientists will study the
fate of oilin the water and sediments and determine its
effects on bottom-dwelling organisms such as seaweed
and clams. One aim will be to determine whether
ecosystems are damaged more by contamination with
an oil-dispersant mixture than by untreated oil.
Fishes, birds, or mammals will not be purposely
exposed to the oil spills. Bird-scaring devices will be
used and project personnel will attempt to ensure that
no sea or land mammals enter the shoreline test areas.

The Baffin Island Oil Spill (BIOS) project, initiated
by the federal government’s Arctic Oil Spill Program,
is the culmination of several years of planning. The
$4-million project is managed by an international
committee composed of representatives from the
Department of the Environment, the Department of
Indian and Northern Affairs, the Department of
Fisheries and Oceans, the Canadian oil industry, the
Norwegian government, and the United States
Oceanic and Atmospheric Administration. Northern
residents have been consulted throughout the plan-
ning of BIOS and support the aims of the project; in
fact, the test site selected was suggested by the Pond
Inlet Council.

offers a glimpse of prairie history. Visitors may view
sections of the original Canadian Pacific railbed, as
well as teepee rings left by pre-European plainsmen.

The Prairie Wildlife Interpretation Centre is the
most recent addition to a series of Environment Can-
ada centers designed to provide opportunities for
direct contact with natural phenomena, typical of a
specific region. The existing four centers interpret (1)
the wet interior British Columbia forest, a large
freshwater marsh, and the valley environment at Cres-
ton, British Columbia; (2) the hardwood forest region
and Wye marsh at Midland, Ontario; (3) the migra-
tion and population management of Snow Geese at
Cap Tourmente, Quebec; and (4) the Atlantic coastal
region at Percé, Quebec.

106

Book Reviews

ZOOLOGY

Conservation of Marine Birds of Northern North America

Edited by James C. Bartonek and David N. Nettleship. 1979.
Wildlife Research Report 11. United States Department
of the Interior, Fish and Wildlife Service, Washington.
ix + 319 pp. Free.

James C. Bartonek and David N. Nettleship
brought together a representative selection of the pap-
ers presented at the International Symposium on the
Conservation of Marine Birds of Northern North
America, held in May 1975, in Seattle, Washington.
This symposium, which was sponsored by the Natural
Resources Council of America, the National
Audubon Society, the National Wildlife Federation,
and the United States Department of the Interior,
Fish and Wildlife Service, was “convened because of a
growing awareness that not all was well with our
marine birds.” Participants included scientists,
administrators, conservationists, and many interested
lay people. The objectives of the symposium were, in
chairman H. K. Nelson’s words, “to identify problems
and the needed information and programs necessary
for the conservation of marine birds of northern
North America.” From the 22 communications
included in this volume, and from I. C. T. Nisbet’s
summary, it is clear that the symposium succeeded in
fulfilling its objectives.

The following subject areas are covered in the pap-
ers: (1) the marine environment of birds, (2) the status
of marine bird populations, (3) the biology and ecol-
ogy of marine birds in the north, (4) conflicts between
the conservation of marine birds and uses of other
resources, (5) programs and authorities related to
marine bird conservation, and (6) conservation of
marine birds in other lands.

In two papers the environment of marine birds is
introduced. In Long term climatic and oceanographic
cycles regulating seabird distributions and numbers,
M. T. Myers of the University of Calgary discusses
short-term and long-term influences on seabirds, and
the necessity of continuous research. He emphasizes
the fact that “seabird ornithologists must not rely on
government programs to provide continuous data
Over a period of years.” Instead, he suggests that biol-
Ogists, governments, and amateurs should combine
their resources “so that any one of them can continue
the work [of censusing marine birds] if any other
element should be incapacitated.” In Sea ice as a
factor in seabird distribution and ecology in the Beau-
fort, Chukchi, and Bering seas, G. J. Divoky discusses
the general effects of sea ice on birds and the specific
effects of ice on birds in the western Arctic. He also
points out the difference between arctic and antarctic

pack-ice systems.

Part 2, on the status of marine bird populations,
contains various papers on birds breeding along the
arctic seacoasts, in the Aleutian Islands, the Gulf of
Alaska, and in British Columbia and Washington
State.

Section 3 includes papers on the Trophic relations
of seabirds in the northeastern Pacific Ocean and
Bering Sea, on Population dynamics in northern
marine birds, and on Time-energy use and life history
strategies of northern seabirds. M. D. F. Udvardy, in
a brief article, describes the Zoogeography and taxo-
nomic relationships of seabirds in northern North
America. Their distribution in the northwest is pres-
ented in a well-organized table. ;

The social and economic values of marine birds,
resource development along the coast, marine birds
and commercial fishing, and seabirds and introduced
animals are some of the topics covered in the next
section entitled “Conflicts between the conservation
of marine birds and uses of other resources.” J. G.
King and G. A. Sanger provide an interesting and
timely “oil vulnerability index” for marine-oriented
birds; this is especially useful at a time when oiled
birds and their cleaning and rehabilitation are an
increasing concern for ornithologists not only along
the coast, but also along inland waterways.

Programs and authorities related to conservation,
in Washington State and in British Columbia, and the
petroleum industry’s role in marine bird conservation,
are the subjects of the next section, while the last part
of the book provides an interesting look at conserva-
tion in such widely separated areas as Norway, the
Danish monarchy (including the Faroe Islands), and
New Zealand.

lan C. T. Nisbet, in hissummary of the symposium,
stresses that there are many important topics that
have not been treated in the papers presented. He calls
for more emphasis on the surveying of medium-sized
seabird colonies, and for more information on the
factors influencing the winter range of seabirds.
Nisbet believes that in the future “highest research
priority should be given to obtaining basic informa-
tion on reproductive success and life table data for
some representative species” of seabirds. He further
underlines the need for long-term studies of a few
carefully selected species, so “that we can trace the
effect of environmental fluctuation on their perfor-
mance for a long period.” Threats to marine bird
populations constitute another line of research,
because only threats by introduced predators and by

107

108 THE CANADIAN FIELD-NATURALIST

oil pollution are well documented. Potential threats,
such as the drowning of diving birds in fish nets, the
dumping of toxic substances, mineral development
(both onshore and offshore), and human disturbance
resulting from increased industrialization and even
tourism (particularly the prospective influx of natural
history tours), all of which can have major adverse
effects, have to be carefully studied.

Most of the articles are accompanied by well-
prepared, informative maps and tables. There are also
some useful diagrams. There are only two photo-
graphs, of rather poor reproduction, but this, in no
way, detracts from the merits of the book. It is regret-
table, however, that many other papers presented at

The Insects and Arachnids of Canada

Vol. 95

the symposium could not be included. Of particular
interest would have been R. E. Le Resche’s communi-
cation dealing with the institutional problems
involved in protecting and managing seabirds on an
interregional and international basis.

I would recommend this book to anyone interested
in marine birds, their environment, and their conser-
vation. Although almost five years have elapsed
between the symposium and the publication of this
volume, it was well worth waiting for.

MARIANNE GOSZTONYI AINLEY

4828 Wilson Avenue, Montreal, Quebec H3X 3P2

Part 1. Collecting, Preparing and Preserving Insects, Mites and Spiders
Part 3. The Aradidae of Canada (Hemiptera: Aradidae)
Part 4. The Anthocoridae of Canada and Alaska (Heteroptera: Anthocoridae)

By J. E. H. Martin, Ryuichi Matsuda, and Leonard A.
Kelton (respectively). 1977, 1977, 1978. Supply and Ser-
vices Canada, Hull. 182 pp., illus. $3.50 in Canada; $4.20
elsewhere. 116 pp., illus. $4 in Canada; $4.80 elsewhere.
101 pp., illus. $4 in Canada; $4.80 elsewhere.

The first part to appear in this long needed series
was Part 2, The bark beetles of Canada and Alaska, by
Donald Bright, Jr., in 1976, covering the beetle family
Scolytidae. Since the appearance of Part 4 and the
time of this writing, Parts 5 and 6 have been published
on The crab spiders of Canada and Alaska (Araneae:
Philodromidae and Thomisidae), by C. D. Dondale
and J. H. Redner, and The mosquitoes of Canada
(Diptera: Culicidae), by D. M. Wood, P. T. Dang,
and R.A. Ellis. It is to be hoped that the series will
continue to appear at least at its present rate if not
more rapidly, for there are a lot of insects and arach-
nids yet to be covered.

Part | appears to be a greatly expanded and
updated version of Beirne’s earlier compilation, with
the addition of photographs as well as more and
improved drawings. In the foreword, D. F. Hardwick,
Director of the Biosystematics Research Institute,
states the two objectives of this publication: “to
acquaint amateur entomologists or potential ama-
teurs with the basic methods of obtaining insect and
arthropod material and preparing a collection; and to
serve as a guide to both amateur and professional
entomologists on how to prepare material before
submitting it to the National Identification Service of
the Biosystematics Research Institute.”

The first section, 68 pages in length, is concerned
with equipment and methods for collecting, followed

by a large section on equipment and methods for
preserving and mounting, including microscopical
preparations as well. This is a section on general tech-
niques. Storage and care of collections is dealt with
next. Most of the remainder of the book treats the
insects and arachnids by order, discussing special col-
lecting and preservation techniques pertaining to each
order. Three pages are devoted to formulas for preser-
vatives, mounting media, stains, etc., and nearly three
pages to references.

This is a very useful book, one of the best of its kind
available, which I heartily recommend to beginning
amateurs and professionals alike.

Part 3 deals with the Aradidae or flat bugs, also
known as fungus bugs, for all but one of the 49 Cana-
dian species feed upon fungi. They are usually found
under the bark of dead trees.

The introduction is very brief and is followed by a
discussion of structure and terminology. There is a
key separating the two subfamilies represented in
Canada, each containing one genus and a key foreach
of the two genera. The key to Aradus separates the
species occurring in Canada and the adjacent states of
the USA. Aneurus is represented in Canada by only
two species.

The species accounts consist of a description, dis-
tribution, a list of associated plants (for most species),
and remarks that aid in identification. There is also a
spot map for most species. Supplementing the keys,
but located at the end of the book, are 15 figures
consisting of line drawings, and 15 plates in the form
of black-and-white photographs. Reference is made
to both figures and plates in the keys. The glossary is

1981

three pages long, followed by two and one-half pages
of references. There is no index. I noticed only one
apparent error, on page 45, where “Vermont” should
probably read “Vernon.”

This book will be useful to those with a special
interest in the Hemiptera, or true bugs, as will the
following publication.

Part 4 is concerned with the Anthocoridae, or
flower bugs, of Canada and Alaska. These are also
known as minute pirate bugs, for their small size
(2-5 mm) and their predacious habits. Of approxi-
mately 85 species known to occur in North America
north of Mexico, 41, representing 14 genera, are
known from Canada and Alaska.

The book begins with an introduction followed by
discussions on collecting and preserving specimens,
classification, and morphology, and a short glossary.
There is a key to the three subfamilies represented in
the region covered by the book and keys to the tribes,

The Birds and Birders of Beaverhills Lake

By Robert Lister. 1979. Edmonton Bird Club, Box 4441,
Edmonton, Alberta. 264 pp., illus. $9.50.

Beaverhills Lake, deep in the Aspen Parkland,
75 km E of Edmonton, is Alberta’s Point Pelee for
water-loving birds of every size and description:
common or accidental, migrant or summer resident. It
is one of the few places where, in spring, summer,
autumn, or, of course, winter, if you pick your day
with reasonable care, you will see no other bird-
watchers, and in fact probably not even any other
people! At the-same time you may be completely
mesmerized by the multitudes and diversity of the
shore- and water-birds. In early spring you may wit-
ness hundreds and often thousands of waterfowl - real
wild ones, not their Stanley Park or Toronto Island
subspecies — flying to and from their feeding areas,
clearing leafless treetops by inches. A month later the
shorebirds will have their turn to impress you with
their numbers and variety.

Scientific importance of Beaverhills Lake as a
breeding area and a migratory stopover was first rec-
ognized early in this century by Professor William
Rowan. Asa result, Beaverhills Lake is probably the
best known area, ornithologically, in Alberta. The
birds and birders of Beaverhills Lake is a popular
(narrative) account, a local ornithological history, of
this famous lake and the observations recorded on its
shores from the time of Rowan to the present. The
author, Robert Lister, was Rowan’s field technician
for over 30 years and is today a pillar of the Edmonton
Bird Club.

BOOK REVIEWS

109

genera, and species, where required. A total of 123
figures illustrate this publication, consisting of 46 line
drawings, 40 scanning electron micrographs, and 37
drawings of the bugs. The species accounts consist of
descriptions of each sex, remarks aiding in identifica-
tion, habitat, anda spot map. At the end of the book is
a list of scientific and common names for plants
referred to in the text, three and one-half pages of
references, and an index. As I mentioned earlier, this
book will be of special interest to those with an interest
in the true bugs, and they are an interesting group of
insects.

Not only is this series useful to the professional
entomologists, but it will also serve as an important
stimulus to the amateurs and beginning amateurs.

WILLIAM B. PRESTON

Manitoba Museum of Manand Nature, 190 Rupert Avenue,
Winnipeg, Manitoba R3B0N2

Much of Lister’s story is taken from Professor
Rowan’s field notes. Rowan’s interests were many,
from his world-renowned studies on the effects of
photoperiod manipulation on migration to his lesser
known studies on loon embryology and Alberta wad-
ers on the British list. Regardless of his interests, they
were always carried out at Beaverhills Lake — captur-
ing juncos or crows, collecting loons, waders et al., or
just plain waterfowl hunting, which seems to have
been freely intermingled with everything. Over the
course of 30 or more years, the exploits and daily
undertakings of this scientific and ornithological pio-
neer are described and quoted.

Thus we have many of the diverse exploits of
Rowan from the early 1920s to the late 1950s. Some of
these exploits were carried out in the finest detail ina
pioneer setting, others failed miserably, but there was
always humor in them, for example, the day Rowan
backed his car over his shotgun, bending both barrels.
“Nothing daunted he simply turned the gun over and
backed over it again to straighten them.”

In Chapter 7, Lister has concluded the history of
Beaverhills Lake as seen through the eyes of Professor
Rowan and turns to the records of the Edmonton Bird
Club (EBC). In his own words, Lister states that this
brought to Beaverhills Lake “. .. a new kind of out-
door enthusiast who was neither interested in collect-
ing birds nor shooting them for sport.”

The book’s contribution is decidedly historical; its
scientific ornithological contents are few. Dates, exact
locations, and observers are provided for only some of

110

the more outstanding records. A checklist of the birds
of Beaverhills Lake area appears as an appendix;
however, it contains at least one obvious error and a
questionable call. Cinnamon Teal, which are omitted,
have been recorded from Amisk Creek, just south of
the lake. They are also mentioned on p. 231 in the text!
Also a single Surfbird and at least one Sharp-tailed
Sandpiper have appeared at Beaverhills Lake, and
have been photographed and accepted by the Alberta
Ornithological Records Committee. Is it anything
other than coincidence that the one seen on an outing
is fully accepted while the one not seen on an EBC
outing is only hypothetical!

On concluding the book, as one who once regularly
visited Beaverhills Lake, I was left with many mixed
feelings. I was disturbed at much of the wholesale
slaughter that went on, of Peregine Falcons being so
abundant that“... one settled on Harold’s back when
he was laying camouflaged for geese. He rolled over
and put it up and adroitly dropped it.” I was embar-
rassed by Rowan’s frequent racial slurs, and the print-

THE CANADIAN FIELD-NATURALIST

Vol. 95

ing of them is totally inexcusable. I was amazed by the
scenes that have not changed with time, e.g., the diffi-
culty in getting around the lake. I was disappointed in
the almost total lack of reference to colonial water-
birds which have abounded on the lake for years
(cormorants, pelicans, gulls, terns, and herons).
Finally I was pleased that except for Greater Prairie
Chickens and Whooping Cranes, which are admit-
tedly a large exception, the variety of birdlife at Bea-
verhills Lake has not changed all that much in 50
years.

To conclude, if you know or have known birding on
Beaverhills Lake, the book is a must; it will greatly
increase your appreciation of the lake. If you don’t
know the lake you can perhaps find better things to
read or at the very least be ready for a boring series of
hunting stories.

D. V. CHIP WESELOH

Canadian Wildlife Service, P.O Box 5050, Burlington, Onta-
rio L7R 4A6

The Peregrine Falcon in Greenland: observing an endangered species

By James T. Harris. 1979 University of Missouri Press,
Columbia. 255 pp., illus. U.S. $15.95.

The search for Peregrine Falcon eyries and the
detailed observation of one during a summer in
Greenland provide the basis for a discussion of pere-
grines as an endangered species, primarily in North
America, from both scientific and humanistic per-
spectives. In 1972, the author was part of a research
team investigating the breeding status of peregrines in
western Greenland. The narrative of this book con-
centrates on his search for eyries and detailed observa-
tion of one eyrie during incubation, feeding of young,
and to a limited extent the interactions of these pere-
grines with other birds. Throughout this narrative a
variety of aspects of peregrine biology, such as the
effect of DDT on populations, nest site selection, and
prey utilization, are included to detail the life of pere-
grines in general rather than just the Greenland
population.

The book is enhanced by some accurate and concise
summaries of biological concepts, but it also suffers
from contradictions in fact from section to section,
inclusion of many extraneous details, and some poor
summaries of biological concepts. Very understand-
able treatments of the effects of DDT, Mayr’s views of
species versus subspecies, and peregrine courtship
flights give one an impression that the book ade-
quately covers peregrine biology. Unfortunately,
either selective reading or an ignorance of some
important peregrine research leaves contradictions

throughout the book. The author wants peregrines to
be symbols of wild areas and hence at one point
emphasizes that they nest in places inaccessible to
humans (p. 6), while he later discusses peregrines nest-
ing in cities such as Montreal. Coverage of historic
peregrine numbers and their subsequent decline is
hampered because it is not dealt with as a unit but is
split by a discussion of the effects of pesticides. Also,
the 1975 North American Peregrine Falcon Survey
(Fyfe et al. 1976, Canadian Field-Naturalist
90: 228-273) is ignored resulting in a biased represen-
tation. The inclusion of many mealtime conversations
concerning weather, who prefers coffee to tea, and an
attempt to feed cheese to a spider often slow the book
to a snail’s pace and distract the reader from the main
theme. The suggestions that mammals are scarce in
Greenland because they have few places to hide, that
Caribou have antlers to protect themselves from
wolves, and that foxes see people as ‘idle, furless and
itchless’ reduce the credibility of the reporting.

The idea of using observations of one population of
Peregrine Falcons to popularize the life of this falcon
is very good. The execution as shown by the above
examples is mediocre, certainly not as the dust jacket
suggests, ‘a model of scientific accuracy,’ or reminis-
cent of Peter Mathiesson.

DAVID MCCORQUODALE

758 Ridgewood Aveune, Ottawa, Ontario K1V 6N1

1981

Some Adaptations of Marsh-Nesting Blackbirds

By Gordon H. Orians. 1980. Princeton University Press,
Princeton, NJ. 295 pp., illus. Cloth U.S. $18; paper U.S.
$7.95.

This is the 14th book in Princeton’s Monographs in
Population Biology series. It is a compilation of 20
years of research on marsh-nesting blackbirds with an
aim to testing and expanding ecological theory
through measurements and observations in the field.

The book is well organized into eight chapters. The
first is an introduction to the biology of the Red-
winged Blackbird (Agelaius phoenicus), the Yellow-
headed Blackbird (Xanthocephalus xanthocephalus),
and Brewer’s Blackbird (Euphagus cyanocephalus),
which are the subjects for most of the studies. A brief
outline of the characteristics of the marshes and of the
materials and methods complete the chapter.

Next, the marsh is considered in detail as a source of
food, cover, and nest sites. This provides background
for discussion of the adaptations of blackbirds to the
resources available, starting with habitat, territory,
and mate selection and finishing with a detailed look
at foraging behavior. From this basis of interactions
between individual birds and the marshes, a jump 1s
made to population and community patterns that
result from these interactions. In the next chapter
variability in resource use between populations and
species is considered, which leads to a discussion of
the implications of this variability in competition and

BOOK REVIEWS Leta

community structure, thus concluding the section on
the marsh-nesting blackbirds of western North
America.

In the following chapter, the author compares the
generalizations molded by north temperate expe-
rience with south temperate examples from Argen-
tina. The marshes and the blackbirds are briefly intro-
duced and then foods, foraging, and competition are
compared.

The final chapter is a discussion of the adaptations
of blackbirds in marshes ona global scale. The signifi-
cance of the structure of marsh vegetation, avian
social systems, island biogeography, and the effect of
blackbirds on the phenotypes of odonates are dealt
with in turn.

This is not an easy reading book full of novel ideas
and evolutionary insights. It consists of ecological
theory and subsequent tests in the field, and the prob-
lems of collecting data relevant to testing hypotheses
are emphasized. The result is a cautious interpretation
of voluminous data, accepting some generalities and
suggesting alternate tests for others. The complement
between field data and theory is the strength of the
book. It will be of value to all students of avian
ecology.

DAVID MCCORQUODALE

758 Ridgewood Avenue, Ottawa, Ontario KIV 6NI

The First Ten Years of the Co-operative Breeding Bird Survey in Canada

By Anthony J. Erskine. 1978. Canadian Wildlife Service
Report Series 42. Supply and Services Canada, Ottawa. 61
pp., illus. $3.75 in Canada; $4.50 elsewhere.

The Co-operative Breeding Bird Survey (BBS), a
project designed to measure annual population
changes in birds (especially small landbirds), was
initiated in 1966 by the United States Fish and Wild-
life Service, with support in Canada from the Cana-
dian Wildlife Service. Annual summaries of the Can-
adian results have been issued since 1970 (Canadian
Wildlife Service Progress Notes, Numbers 15, 21, 26,
32, 38, 45, 60, 74, 84). About 1800 BBS routes (250 of
these in Canada) are surveyed each year in North
America, mostly by amateur volunteers. Each route
consists of 50 “stops” at 0.5-mi (0.8-km) intervals,
where all birds seen and heard in 3 min are recorded.
Each is surveyed by car, once a year (in June or early
July) under standardized conditions. This review is as
much about the results of the project itself as about
Erskine’s report of it.

Of the routes surveyed in both of two consecutive
years, about 80% are considered by Erskine as “com-
parable” (same observer, no major changes in survey
date, weather conditions, etc.), and only these are used
in determining population trends. For analytical pur-
poses, Erskine divides settled Canada into six ecogeo-
graphic regions. (Note: these six regions are not the
same as those shown on the report’s cover!) Since
1966, BBS data have shown significant population
changes in several bird species. These include an
increase in Red-winged Blackbirds over much of
southern Canada, probably because of increased corn
acreage; population changes in several forest-bird
species resulting from spruce budworm epidemics,
and from spraying programs to control the latter; and
short-term declines of some insectivorous birds in
eastern Canada because of cold spring weather.

Nonetheless, | am skeptical of some population
trends suggested by these analyses. For most of the six
regions, the number of comparable surveys (20-30) in

112 THE CANADIAN FIELD-NATURALIST

successive years barely permits meaningful statistical
treatment; furthermore, only 20 or 30 routes per
region may not bea truly representative sample of that
region. When analyzing results for all of North Amer-
ica, the United States Fish and Wildlife Service
groups BBS routes into only three regions (eastern,
central, western). With hundreds of routes in each
region, population trend analyses obviously stand on
much firmer ground than those for the six Canadian
regions used by Erskine.

My most serious criticism of this report is of the
graphs of population trends (Figures 4a through 12),
which occupy part or all of 14 pages of this 61-page
bulletin. Most of these graphs attempt to show sim-
ilarities in population trends among several species
within a region. Although most of these data are also
presented in Tables 3 through 8, they do seem worth
graphing. Unfortunately, most of these graphs lack
any scale to indicate the actual population index
values. Even those graphs which do have a scale, e.g.,
Figure 10a and b) are often cut off well above the zero
mark, making the population changes look larger
than they really are. To bea little facetious, someone
writing an article entitled “How to lie with statistics”

_ The Dragonflies of British Columbia

By Robert A. Cannings and Kathleen M. Stuart. 1977.
British Columbia Provincial Handbook Number 35.
British Columbia Provincial Museum, Victoria. 256 pp.
$2.00.

This is the first of the excellent British Columbia
Provincial Museum Handbook series to be devoted to
insects, and what better insect group to begin with
than the Odonata. Not only are they familiar to all of
us but, as the authors point out, they are “. . .among
the most ancient of insects... .” Although the title of
this handbook is The dragonflies of British Columbia,
it includes the damselflies as well.

The introduction contains a number of sections,
first introducing the order Odonata. This is followed
by a brief discussion on the dragonfly in Indian cul-
ture. In the section on the vanishing marshland the
authors refer to the phenomenal destruction of rich
aquatic habitats in the Okanagan Valley and point out
that “. . . the sizes of dragonfly populations that
impressed collectors around Penticton in the 1930's
no longer exist.” It is to be sincerely hoped that we can
learn to avoid this kind of destruction before it 1s too
late.

Also included in the introduction are sections on
basic structure, biology, and zoogeography of Odo-

Vol. 95

could easily pick examples from this report!

A major potential use of BBS data is zoogeogra-
phic, i.e., for mapping a species’ relative abundance in
different parts of its range. In American publications
on the BBS (e.g., Robbins and Van Velzen, United
States Bureau of Sport Fisheries and Wildlife, Special
Scientific Reports (Wildlife) Number 102 (1967) and
Number 124(1969)), many such maps, using different
kinds of shading for different levels of abundance,
have been published. I was disappointed that Erskine
did not include any range maps in his report.

Despite the above criticisms, the BBS has proved to
be a worthwhile and popular project, and Erskine’s
report provides a detailed and thought-provoking
summary of the first 10 years’ results. | hope these
surveys can be continued for many years in the future,
because the amount of information one can extract
from the results seems to increase almost geometri-
cally with the number of years of coverage.

WAYNE C. WEBER

303-9153 Saturna Drive, Burnaby, British Columbia V3J
TKI

nata. The authors list seven additional species to
watch for in the province, and they discuss observing,
photography, and collecting of dragonflies, and
finally, how to use the book.

The body of the book is made up of the keys and
species accounts. There are keys to the larvae as well
as to the adults, and the keys are well supplemented
with excellently clear line drawings by Robert Can-
nings. Asa further aid to identification there are eight
halftone plates by Kathleen Stuart. The species
accounts include a synonymy, descriptions, range,
distribution in British Columbia, field notes, and a
spot map.

The authors admit that their book is based “to a
large degree” upon existing publications. They have
made this information more readily available, how-
ever, and with the addition of their own research have
produced an excellent guide to the 80 species of dra-
gonflies known to occur in British Columbia. Along
with the British Columbia Provincial Museum, they
are to be commended.

WILLIAM B. PRESTON

Manitoba Museum of Manand Nature, 190 Rupert Avenue,
Winnipeg, Manitoba R3B 0N2

1981]

BOOK REVIEWS

113

Annotated List of Workers on Systematics and Faunistics of Canadian Insects and
Certain Related Groups. Pilot Study for a Biological Survey of the Insects of Canada

By the Secretariat. Entomological Society of Canada,
Ottawa. 1977. 107 pp.

This useful little publication, especially to entomol-
ogists, consists mainly of an alphabetic list of workers
in entomology, giving their addresses, disciplines, and
specialties. Included also are current projects of each
worker and an indication of his intended involvement
in the Biological Survey of the Insects of Canada. A
total of 436 workers are listed, and in a supplement
published in 1978, thirteen more names were added.

The introduction outlines the origin and purpose of
the publication and the method of data acquisition.
As well, there is a section on how to use this list. There

are three indices, one to locations, one to taxa, and
one to ecological groups. The user can easily locate
workers ina particular locality, with a certain specialty
or with an interest in certain ecological areas, for
example, insects of alkaline lakes. ;

What more can one say? This is a useful publication
and an excellent beginning for a biological survey of
the insects of Canada.

WILLIAM B. PRESTON

Manitoba Museum of Manand Nature, 190 Rubert Avenue,
Winnipeg, Manitoba R3B 0N2

Wolves of Minong — their vital role in a wildlife community

By Durward L. Allen. 1979, Houghton Mifflin Company
Boston. 544 pp. U.S. $16.95.

Minong is the Ojibway name for Isle Royale. This
544-km2 island has become well known for its Gray
Wolf/ Moose studies, now extending into the 22nd
year. Durward Allen’s book is the third monograph
on the subject. Others were written by L. D. Mech,
The wolves of Isle Royale and by R. O. Peterson,
Wolf ecology and prey relationships on Isle Royale.
Allen has the longest continuous involvement with
Isle Royale. He began planning the wolf studies in
1951 and in 1958 initiated field work with L. D. Mech
as his first graduate student. Mech was followed by
five other principal field workers (graduate and post-
graduate students) all under Durward Allen’s direc-
tion. It is therefore appropriate that this book,
designed for biologists, naturalists, environmental-
ists, game administrators, visitors to the park, and
outdoor enthusiasts should be written by the master
himself.

The title of the book is misleading as the subject is
less about just wolves and their role ina vital commun-
ity as it is about the total Isle Royale ecosystem. That
includes such items as redpolls feeding on birch seeds,
nesting behavior of birds of prey, and the many nuan-
ces in the weather patterns.

The story begins with a speculative account of how
the first wolves could have wandered over an ice
bridge onto the island. The rest is a detailed descrip-
tive account about the principal subjects, namely
human investigators, Gray Wolves, Moose, and the
many other components in the ecosystem. In an elo-
quent style his writing guides the reader through the
park. Allen always maintains a perspective on the
total scene (the whole of nature as he calls it), never

giving up an opportunity to share with us this keen
sense of observation. I found a great deal of satisfac-
tion in reading about those things which field workers
often observe but which are rarely seen in print. Des-
criptions of the existence of “moose jumps” (“elk
jumps” in our studies), evidence of the spurting of
arterial blood on vegetation at kill sites, and the habits
of ravens utilizing food remains in wolf scats are but a
few examples. Photographs are well chosen and bring
the Isle Royale scene into the living room of the
reader.

For those interested in purely technical information
this book may be disappointing. Durward Allen’s
treatise does not shy away from technical informa-
tion, but more importantly his work has a philosophi-
cal theme to it. | was impressed by the author’s broad
visions and the insights that these provided. Because
there is an emphasis on the broader picture some
biological details have fallen between the cracks. The
concept of surplus killing, for example, was used
incorrectly; an error frequently made by wolf
researchers. References are heavily oriented towards
classical studies of the past, with less emphasis on
more recent findings. There are numerous typogra-
phical errors and one annoying misspelling of an
author’s name repeats itself; however, these are minor
shortcomings.

We can learn a lot from the Isle Royale studies.
First and foremost is the need for long-term research.
Allen, on page 369, states “a weakness of ecological
research in its first 50 years has been the short term
approach.” Had the Isle Royale studies ended after 10
years the results and conclusions would have been
different than was the case after the second 10 years,
or alternatively what would have been the conclusion

114

if studies had begun 10 years later and then ended?
Secondly we find out (page 90) that wolf/ prey systems
are in rough equilibria — “in the operation of ecosys-
tems we see repeatedly that calamitous extremes of
almost any kind are countered by compensations. The
high-to-low fluctuations tend to even out ina pattern
that spells survival over long time periods for all kinds
of life that fits the local organization.” Thirdly, givena
lack of human interference, predation on ungulates is
a vital biological process. The scene is ever changing

Mammals of the Eastern United States

By William J. Hamilton, Jr., and John O. Whitaker, Jr.
1979. Cornell University Press, Ithaca, New York. 346 pp.,
illus. U.S. $19.95.

This revision of Hamilton’s 1943 edition treats 105
species of land mammals which occur east of the
Mississippi River. The preface states that species nat-
urally inhabiting this area are described, but three
introduced species of Lepus and the nutria are also
included. About 65% of these 105 species also occur in
eastern Canada.

In the first edition, 253 species and subspecies were
considered with each receiving a separate description
and range map. In the current edition, the authors
state that an excessive number of eastern subspecies
has been described and follow the proposal of Whi-
taker (The Biologist 52: 12-15. 1970) that a biological
definition be used for subspecies as well as for species.
The omission of subspecies treatments in the present
work allows emphasis on habits rather than on the
taxonomy of mammals and increases the value of the
work for naturalists and amateur ecologists.

The species accounts contain the following: (1) a
verbal description including measurements and usu-
ally a drawing or photograph, (2) a statement on
distribution and a range map, (3) a discussion of the
species habits. Keys to identification, which are
included throughout the text, often use skull and den-
tal criteria, but skull characteristics are not included in
the species descriptions. The lack of a glossary or any
general treatment of the mammal skull may make the
keys difficult to use by those unfamiliar with mamma-
lian anatomy.

This is an attractive volume; the species accounts
are interesting to read, and the maps, photos, and
drawings are clear. Much information in the accounts
is inadequately documented, however. For example,
it is stated that recent studies indicate Vulpes fulva to
be the same species as the old world Vulpes vulpes, but
no citation is provided. Blarina brevicauda and B.
carolinensis are treated tentatively as separate species
without citing pertinent studies such as Ellis et al.

THE CANADIAN FIELD-NATURALIST

Vol. 95

and ultimately habitat and winter severity so far have
determined the course of events. Anyone reading this
book will understand that there never will be a final
outcome. In the meantime this book stands as an
important reference on a fascinating subject.

L. CARBYN

Canadian Wildlife Service, 9942-108 Street, Edmonton,
Alberta TSK 2J5

(Journal of Mammalogy 59: 305-311. 1978). The
unsupported statement that Clethrionomys gapperiis
an agile climber differs from the results of Getz and
Ginsberg (Animal Behavior 16: 418-424. 1968) in
which an absence of significant arboreal behavior in
this species was reported. Much of the literature cited
is old; 52% of the 83 references predate 1940. The guide
to further reading for Rodentia includes 41% before
1940 and only 7% (half of which are papers by Whi-
taker) since 1970. Whitaker’s accounts of Cryptotis
parva and Zapus hudsonius are the only citations
from the Mammalian Species series, although many
others exist for eastern species. Further, Vaughan’s
1972 edition of Mammalogy and Walker’s 1968 edi-
tion of Mammals of the World are listed although
both occur in new editions (1978 and 1975).

The authors strongly emphasize that much remains
to be learned about many eastern mammals and even
state that we have added little to the knowledge of
many species since Audubon and Bachman’s Vivipar-
ous quadrupeds of North America; however, the text
suffers by not including reference to many important
eastern studies. Choate’s paper (Journal of Mammal-
ogy 54: 41-49. 1973) on Peromyscus in New England
is not cited nor are any of the many pertinent studies
by Getz (for example, Ecology 49: 276-286. 1968).
Microtus breweri, a probable semispecies found only
on Muskeget Island, Massachusetts, and reported on
extensively by Tamarin in the 1970s, could have at
least been mentioned under Microtus pennsylvanicus,
but it is not discussed at all.

Many of these deficiencies will not be obvious or
important to the amateur reader who wishes to
become familiar with the mammals of the eastern
United States and Canada. This readable book should
increase the public awareness of eastern mammals and
may stimulate the further study of these forms.

DAVID A. LOVEJOY

Westfield State College, Westfield, Massachusetts 01086.

1981

BOOK REVIEWS

115

Responses of Peary Caribou and Muskoxen to Helicopter Harassment

By Frank L. Millerand Anne Gunn. 1979. Occasional Paper
Number 40. Canadian Wildlife Service, Edmonton, 90 pp.
Free.

Miller and Gunn have produced a fine example of
applied biological research. This publication isa tool.
In the face of increasing oil and gas exploration and
development in the Canadian Arctic, this volume pro-
vides resource managers with a means of developing
site-specific management guidelines for the operation
of helicopters in areas frequented by large ungulates.
In short, their goal was to determine the effects of
various types of helicopter activity likely to occur
during the construction and maintanence of a
pipeline.

The studies were conducted during the summers of
1976 and 1977 on Prince of Wales and Russell islands,
Northwest Territories. The authors examined the
overt behavioral responses of Peary Caribou (Ran-
gifer tarandus pearyi) and Muskoxen (Ovibos
moschatus) to helicopter harassment. The 1976 field
season objectives were to develop techniques for
observation and to describe the responses of harassed
animals from ground observation sites and from the
air. In 1977 more specific objectives led to the exami-
nation of the effect of the aircraft’s altitude on animal
response, animal habituation to overflights, delayed
responses to harassment, and animal response to
helicopter landings and ground-based work parties.

Of major importance was the determination of an
inverse relationship between the response level of the
animals and the altitude of the overflights, and of the
horizontal distance between the animals and a landing
helicopter. Many of the recommendations are based
upon this relationship. The statistical rigor employed
to analyze the data is impressive. In completing a
literature review on the topic, I found no other study
that details the responses of free-ranging ungulates to
aircraft harassment to the extent found here. Gener-
ally, Millerand Gunn found that the responsiveness of
cows and calves of both species and of solitary bull
Muskoxen, the group size and social organization, the
number of calves in a group,the position of the sun
and the direction of the wind relative to the flight, the
previous activity of the animals, and the terrainare all
contributing factors to the level of response exhibited
by the harassed animals.

To minimize harassment the authors recommend
that minimum flying heights be restricted to 300 magl
(above ground level) from December through April,
and that to ensure minimal disturbance during calving
and post-calving periods (May—August) aircrafts be
restricted to altitudes no lower than 600 m agl. In
addition, it is recommended that the 600 m agl min-

imum flying height be in effect from August through
November to prevent harassment during the rut; too,
itis during this time that the young animals are build-
ing up crucial winter fat reserves. To limit ground
harassment they recommend that crews and vehicles
not approach the animals and maintain a buffer dis-
tance of 1000 m. Camp dogs should not be permitted
to interfere with Caribou or Muskoxen, and construc-
tion materials that might hamper movement or be
hazardous to the animals should not be left on the site.
Juvenile Caribou are known to be inquisitive, which
makes them extremely vulnerable to harassment and
hunting. The important point here, as Miller and
Gunn rightly mention, is that “. . . the investigative
behavior of young caribou should not be regarded as
evidence that the animals will not be subsequently
stressed as a result of the encounter with ground crews
and vehicles.”

The recommendations should be taken seriously,
especially in view of the fact that Peary Caribou and
Muskoxen are inhabiting areas near the northern lim-
its of their range. The authors report that reproduc-
tion Is inconsistent and the general physical condition
of the animals over | yr of age is poor. Consequently,
recruitment is low and natural mortality is high. To
inflict additional forms of potential mortality-causing
agents on a population whose status is at best precar-
lous 1s simply playing with fire.

The physiological effects of helicopter harassment
are not considered. It is known that a harassing stimu-
lus precipitates excitement that is not always detecta-
ble. The authors allude to the problem of not knowing
“.. . whether animals that did not overtly respond
were in fact responding at a physiological level.” This
is often referred to as active inhibition.

V. Geist (1975. University of Calgary, unpublished
report) states that “... intense harassment or frequent
harassment which imposes a burden on the energy and
nutrient supply of the animal can become the so-called
alarm reaction.” Physiological trauma can be detri-
mental to an animal’s quest for survival. The need for
a physiological approach, in conjunction with a
behavioral one, is apparent. Telemetric techniques
have not, however, been successfully refined to a level
that will permit monitoring of physiological responses
(i.e., heart and respiratory rate) to harassment of
remote populations of wild, free-ranging animals. In
the interim, biologists will have to continue to develop
sound baseline data with the available techniques.

“Need for further study” is an important section
because it outlines the limitations of this study. Miller
and Gunn call for increased research into the physio-
logical aspects of harassment, long-term monitoring

116

programs, and harassment studies during the fall and
winter months. It is not known how Peary Caribou
and Muskoxen respond to harassment from Sep-
tember to June. The late winter months (March to
May) are especially critical, for it is during this period
when the animals are in their poorest physical condi-
tion and most vulnerable to harassment.

This volume is an important contribution to

BOTANY

The Rare Vascular Plants of Saskatchewan

By Robert V. Maher, George W. Argus, Vernon L. Harms,
and John H. Hudson. 1979. Syllogeus #20. National
Museums of Canada, Ottawa. 55 pp. English and 57 pp.
French. Free.

In Southern Saskatchewan the land is mostly flat,
fertile, and free of rocks, allowing easy and almost
complete exploitation by cultivation. There has been
no long-term policy to preserve the diversity of vegeta-
tion adapted to our climatic and soil conditions since
the retreat of the last great glacier. Now, belatedly,
agriculturalists wish that there were natural areas,
bench marks, against which change could be mea-
sured and the performance of other ecosystems could
be judged. Botanists are belatedly trying to take inven-
tories of all the species (the total genetic diversity)
existing in the province so that man’s impact on envi-
ronment can be measured.

In the book The rare vascular plants of Saskatche-
wan, after an eight-page introduction 1s a list of 368
species (in 60 families) which have been reported as
rare. The brief note on each species includes the refer-
ences which identify that plant as rare, lists the herba-
ria where voucher specimens are located, and com-
ments on habitat, total range, distribution within the
province, and status of protection. The list (about 30
pages) is followed by a bibliography (5 pages) and a
list of excluded species (unconfirmed, misidentified,
or considered too common for inclusion). The text and
appendices are in both English and French and
between these two sections there are 300 small maps of
Saskatchewan (25 pages) clearly showing locations of
those 300 species. The list and the maps of species are
in alphabetic arrangement but there is a summary of
the species by family in taxonomic sequence.

The authors admit that the information about
many of the plants in the list is incomplete. They also
admit that only preliminary verification of the speci-
mens has been undertaken and that there 1s real need
for field verification of the current status of the popu-
lations. Following the Preface the authors make a plea
for information, “additions, deletions, comments and

THE CANADIAN FIELD-NATURALIST

Vol. 95

resource management and is a must for professionals
who are faced with the task of developing sound land-
use management strategies in Canada’s north.

PAUL A. GRAY

Wildlife Service, Government of the Northwest Territories,
Yellowknife, Northwest Territories X1A 2L9

corrections,” with “as complete documentation as
possible.”

This reviewer, in spite of a lifetime in Saskatchewan
and considerable interest in plants, has never seen
many of the species listed in the book. Certain species
do occur over a fairly wide range but apparently they
are not common and | do not have any evidence to
back the suggestion that they are too common for
inclusion in this book. Rather than suggesting dele-
tions my first inclination on reading the list was to
suggest certain species for possible addition to the list.
There are some species which I have always consi-
dered rare that are not in this list. I will be considering
possible evidence to support such recommendations.
The book is valuable because it makes us realize how
little we know of the world around us and how rare
and precious so many of the species are. The book
encourages further study and research and there will
be revisions and modifications as our knowledge and
understanding grow. Future lists will surely distin-
guish some species as endangered, and perhaps some
as extinct; they will help to locate sites which must be
set aside as ecological reserves for the purpose of
protecting certain plant species from extinction.

There is no legal protection for any plant species in
Saskatchewan, but there are some areas which receive
token protection. It is hoped, however, that Saskat-
chewan’s ecological reserves act will soon be
operative.

The authors are to be congratulated. The book
brings together a large body of information on rare
plants of Saskatchewan. It will encourage much addi-
tional study. It is essential reading for anyone inter-
ested in preserving biological diversity, man’s impact
on environment, or the establishment and manage-
ment of ecological reserves to protect habitats essen-
tial for the existence of certain rare plants.

GEORGE F. LEDINGHAM

Vascular Plant Herbarium, University of Regina, Regina,
Saskatchewan S4S 0A2

—e.

1981

BOOK REVIEWS

il 7/

Identifying Grasses: data, methods and illustrations

By H.T. Clifford and L. Watson. 1977. University of
Queensland Press, St. Lucia, Queensland, Australia. U.S.
distributor Prentice Hall. 146 pp., illus. U.S. $35.

This book presents a fascinating mixture of ideas
and information, but anyone looking for an identifi-
cation manual will be disappointed. It is about
grasses, but many of the ideas in it should appeal even
to those taxonomists who (to their shame) avoid
grasses. The first chapter starts with a discussion of
the difference between classification and identifica-
tion and contains one of the more readable and com-
plete accounts of different types of keys (conven-
tional, directory, and punched card) that I have come
across. The use of computers in plant identification is
the subject of the second chapter. In both these chap-
ters it is evident that the authors have considerable
experience with their subject. They admit that “the
potential flexibility of ... computerized key making...
seems to us very attractive” yet state very clearly that
under certain circumstances, e.g., if only a single key is
to be produced, computer key generation is not worth
the effort necessary to obtain the data in the appro-
priate format. Together the two chapters provide a
very useful overview and assessment of different con-
cepts of identification keys.

The next two chapters are on a totally different
topic: the morphological variation and vegetative
anatomy of grasses. The authors make an interesting
distinction between racemose spikelets, in which the
rachilla is usually prolonged beyond the uppermost
female fertile floret, and cymose spikelets in which it is
not. “Experience in using our data to generate keys
has shown that ‘rachilla prolonged beyond the
uppermost female fertile floret/ not’ produces a sym-
metrical first division for many samples of grass gen-
era.” It would be interesting to determine whether the
same would be true for samples of Canadian grass
genera.

Those who have struggled with grass keys that
expect the beginner to recognize as “sterile lemmas”
minute hairy appendages or to know intuitively
whether a spikelet lacks a glume ora sterile lemma will
take pleasure in the statement that “Keys ought not to
demand levels of comparative morphological and
taxonomic knowledge that most would-be users can-
not reasonably be expected to possess.”

The discussion of the vegetative anatomy of grasses
is particularly interesting, for the authors draw atten-
tion to a number of characters not previously consi-
dered to be of much significance such as the number of
vascular bundles in the midrib (often three or more in
panicoid and andropogonoid grasses) as well as indi-
cating that other characteristics often thought to be

constant within a genus may in fact be quite variable.
They found, forexample, that it was not worth record-
ing more than seven types of silica body whereas
Metcalfe recognized twenty.

Most of the book is taken up with photographs of
epidermal and cross-sectional preparations of differ-
ent grasses, approximately 120 epidermal prepara-
tions and 60 cross sections. To my prejudiced eye (I
like grasses) these are beautiful and I am delighted to
have such a large array of species represented in one
place. The photographs are also used to explain the
terminology used. It was interesting to me that the
Australian members of the Stipeae seem to be as
variable as the North American members.

The last part of the book is taken up by generic
descriptions. Their style is best illustrated by a quota-
tion:

“Agropyron 1/3/5,6/8/10/12/14/16/18/20/22/
23/25,26/ 28/29, etc.” The numbers refer to attributes
which are described at the start of the section. For
instance, | /3 means the upper leaves have a ligule (1),
and the 3 means that the ligules are membranous. It is
obviously nota style that makes for easy reading but it
does pack a large amount of information into a rela-
tively small space.

The last chapter presents a summary of the author’s
classification of Australian grasses. It is an informal
summary with some groups given formal taxonomic
names (e.g., Stipeae) and others informal recognition
(e.g., danthonioids). Although I was glad to see their
classification, I found their introductory paragraphs,
in which they indicate those areas that are “exasperat-
ing” (the arundinoid grasses), and attributes that they
suspect may be of greater taxonomic import than has
so far been recognized, of even greater interest. The
groups were derived with the aid of a divisive poly-
thetic computer program, a fact that is not belabored.
The groups formed correspond, so far as I can tell,
reasonably well to those accepted by Gould for North
American genera.

I recommend /dentifying grasses primarily to two
groups of people: those interested in methods of iden-
tification and those interested in the variation that
occurs within the grass family. It contains a wealth of
ideas for both groups. Those looking for an identifica-
tion manual for Australian grasses must turn
elsewhere.

MARY E. BARKWORTH

Director, Intermountain Herbarium, Utah State University,
Logan, Utah 84322

118 THE CANADIAN FIELD-NATURALIST

Vol. 95

Native Trees and Shrubs of Newfoundland and Labrador

By A. Glen Ryan. 1978. Park Interpretation Publication
Number 14. Parks Division, Department of Tourism,
Government of Newfoundland and Labrador, St. John’s.
120 pp., illus. Free.

I have mixed feelings about this booklet. On one
hand, it is a very useful handy little reference for
amateurs; indeed, as far as park interpretative mate-
rial in Canada and elsewhere 1s concerned, it is better
than most. On the other hand, it has many weaknesses
and inconsistencies which detract from the overall
value of the booklet.

In justifying this booklet the author states “In com-
piling this book frequent reference was made to
already published material. But this was inadequate.
A considerable amount of original field work and
examination of herbarium specimens had to be done
to make the study complete.” I must confess that,
despite the ‘inadequacies’ of his 44 references, few of
which are quoted in the text, he provides no new
information either taxonomically or phytosociologi-
cally; on the contrary, his treatment falls far short of
his references. In addition, he has failed to consult a
wealth of well-known local literature and data eman-
ating from phytosociological studies on the flora of
Newfoundland and Labrador, which would have
given a broader and more realistic insight into the
taxonomy, ecology, and distribution of trees and
shrubs in the region.

On the premise that they are too difficult to identify,
species of the genera Salix, Amelanchier, Rubus, and
Cratageous were avoided. Yet in Appendix A he lists
no less than 33 Salix, 6 Amelanchier, 4 Cratageous,
and 8 Rubus — most of which have been well illus-
trated in several of the references. I would have pre-
ferred to have seen distinctly recognizable species
included to give these genera some representation.

In the short superficial key, from which it is
assumed that the reader knows the habit of the plant,
the plants are divided into trees and shrubs. Beyond
this the reader is on his/her own because there are no
guiding symbols to let one know where each group of
plants begins. Ericaceae, Caprifoliaceae, and Ros-
aceae, for example, occur sporadically throughout,
depending upon whether it is a creeping or upright
shrub, so that thé order of the plants appears a con-
fused medley. This makes comparisons between spe-
cies tedious.

The emphasis of the publication is on the illustra-
tions. The habit drawings are of excellent quality. But
it is a pity that only a token effort was given to various
features of the plants (twigs, leaves, flower structures,
etc.). Many are erroneous, others poorly labeled, and
yet others are so obscure as to be quite useless for

diagnostic purposes. Occasionally the text does not
match the illustrations. Comparatively, the scale of
some of the illustrations is misleading and often, espe-
cially with regard to seeds, size is not given in the text.
Some of the more obvious errors are these: failure to
note the decussate leaves of Mitchella repens; the
structure of the male catkin in Betula michauxii, seeds
of Pinus resinosa, Betula pumila, B. cordifolia, and
the flower of Diapensia lapponica are all incorrect.
The illustration labeled fruit in Juniperus horizonta-
lis, should be labeled seed (the text described “fruit are
bluish, berrylike cones . . .”). Similarly, the illustra-
tions labeled ‘fruit’ in Potentilla fruticosa does not
show the fruit at all.

Despite the author’s intent to use as much collo-
quial language as possible the terminology is anything
but non-technical; indeed, for any technical subject it
would be presumptuous to assume otherwise for a
booklet of this sort. Much critical inconsistency arises
in the sequence of the descriptive characteristics, and
ambiguity persists throughout the text. In comparing
like species, ambiguous and erratic treatment 1s
tedious to follow, especially when no diagnostic keys
or underlining of differentiating characters are
provided.

The habitat and distribution notes need revision.
The geographical designations are much too vague
even for amateurs. It would have been useful to pro-
videa map of the province outlining the main regions.
Since it is a park interpretation publication perhaps a
map showing park locations and administrative
regions would have served the purpose. Many of the
habitat notes, though not erroneous, do not give a true
representation.

Park interpretative literature quite often does not
measure up to acceptable standards and if my critique
serves any useful purpose, it is perhaps to focus atten-
tion on common faults which occur in so-called col-
loquial park literature. Park naturalists must realize
that it is not simply enough to provide pleasant illus-
trations but also precise and coherent texts. Wherever
there is doubt that may lead to ambiguity, the desire
for colloquialism should be constrained. Ryan, on the
whole, has done a commendable job. He has not
underrated the skill and intelligence of the serious
amateur, and his book is certainly one that most ama-
teurs will enjoy and find useful.

ALEXANDER ROBERTSON

Newfoundland Forest Research Station Herbarium, P.O.
Box 6028, St. John’s, Newfoundland AIC 5X8

1981

BOOK REVIEWS

119

Vascular Flora of the Southeastern United States: Volume 1, Asteraceae

By Arthur Cronquist. 1980. University of North Carolina
Press, Chapel Hill. xv + 261 pp. U.S. $25.00.

Before I opened the cover of this book, I had some-
how anticipated the format of the Jntermountain flora
and Vascular plants of the Pacific Northwest, other
floras with which Arthur Cronquist has been asso-
ciated. This flora, however, does not contain the same
detailed synonomy and citation of types that is found
in the Intermountain and Pacific Northwest floras,
but follows the format laid down by A.E. Radford et
al. in Contributors’ guide for the vascular flora of the
southeastern United States, which is repeated in part,
in the preface.

The presentation is essentially that of a classical
flora with dichotomous keys and detailed descrip-
tions, habitat, and known geographic distribution
within the southeastern states and the adjacent states.
Chromosome numbers are given, but there is no indi-
cation as to whether these counts were based on mate-
rial from within the area treated by the flora.

Synonomy is given where names used differ from
the four manuals which cover parts of the region

Ferns of North-western Himalayas

By K. K. Dhir. 1980. In Bibliotheca Pteridologica, Band 1. J.
Cramer, Braunschweig, West Germany. 158 pp., illus.
DM40.

This book will have little utility for either amateurs
or professionals in North America. For the amateur,
he is told that there are 264 species in the region under
consideration, which is in the states of Jammu and
Kashmir. The region is said to be “not rich in variety
of ferns” but has 3.3 times as many species (80) as
given by Scoggan for Canada. There are no keys, no
descriptions, and no illustrations to distinguish these
264 species, so obviously other works will have to be
consulted. A Canadian will be glad to see some famil-
iar friends such as Ophioglossum vulgatum, Osmunda
claytoniana, O. regalis, Adiantum pedatum, Crypto-
gramma_ crispa, Pteridium aquilinum, Woodsia
alpina (rare), Polystichum lonchitis, Cystopteris fragi-
lis, C. montana (rare as usual), Athyrium filix-femina,
Phegopteris polypodioides, Gymnocarpium dryopte-
ris, Thelypteris palustris, Asplenium viride, A. trich-
omanes, etc., but these will be insufficient to extrapo-
late to all the new genera and species in this region.
The professional will note that the system of classifica-
tion is not discussed, but follows a scheme of Mehra
presented in 1961. A large amount of space is taken up
with a literature citation for the names used, followed

treated: Small 1933; Fernald 1950; Gleason and
Cronquist 1963; and Radford et al. 1968. Literature
citations are provided for recent monographs.

The only awkward feature of the book is the use of
abbreviations for the physiographic regions, with
which the user must familiarize himself.

The area covered is that part of the forested United
States lying east of the prairie and extending north to
the southernmost terminal moraines. The aim is to
include all of the vascular plants growing outside of
cultivation within this region.

The need for a new treatment to replace Small’s
Manual of the southeastern flora has been evident for
many years. This first volume of the new Vascular

flora of the southeastern United States by Cronquist is

an excellent beginning for the series, and it is to be
hoped that the authors for the remaining families will
follow his fine example.

WILLIAM J. CODY

Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6

by references to floristic works concerning the species
in India. The other contribution is to give some eco-
logical notes about each species, e.g., Osmunda clay-
toniana, “very commonat high elevations,” and some
locality data for the collections.

The only light reading is in the first 23 pages, where
there is a brief sketch of the climate, soils, vegetation,
epiphytes, climbers, forest floor species, ravine ferns,
thicket species, lithophytes, water ferns, etc. The wri-
ter uses a colorful style which is not often encountered
in works such as this, e.g., “The water holding capac-
ity of rocky soils along the roadside is desperately
poor’; regarding evergreen ferns, “perpetuating dur-
ing the snowy period”; regarding epiphytes, “in the
forests with lax crown,” “their root systems are more
prolific and wiry thus anchoring them to the substra-
tum strongly so as to withstand the wind velocity,”
“leafy liverworts which retain lot of moisture,” etc.

The most enjoyable portion of this book is the 20
figures (actually high-quality photographs) of a few
species in their natural habitat although this reviewer
is not sympathetic to the words “infested with” when
viewing a luxuriant growth of a Po/ypodium ona tree
trunk.

The 12 pages of bibliography 1s a good cross section
of the world’s literature on ferns as it pertains to the
ferns of India. These references might be useful to a

120

worker studying individual species, but to a casual
reader they add little because there is no elaboration
regarding their significance, and no discussion of their
merits for the list of species.

The number of typographical errors is far too large
for a science such as taxonomy which prides itself in

ENVIRONMENT

THE CANADIAN FIELD-NATURALIST

Vol. 95

being painstakingly accurate about minute details.
DONALD M. BRITTON

Department of Botany and Genetics, University of Guelph,
Guelph, Ontario NIG 2W1

Order in Living Organisms: a systems analysis of evolution

By Rupert Ried]. 1979. Wiley, New York. 313 pp., illus. US
$37.50.

This is a curious book. It reexamines nature from
an original point of view. The structure of life from
molecules to man is studied to detect to what extent
order is found in the constituent blocks. The
approach, although similar in some respects to syste-
matics, produces quite different results because its
goal is not producing phylogenies and classifications,
but an understanding of order. The diverse multidis-
ciplinary approach gives a refreshing view of life.

Life, Ried] proposes, is the result of interaction
between accident (mutation) and necessity (inheri-
tance). The degree of complexity that evolves can be
measured. Information theory /= i/ P, where / is the
amount of information and P is the probability of
occurrence of the event /, is applied. From this he
develops a measure of order, the determinacy content.
The determinacy content for small bacterium, about
10!! bits, exceeds the more complicated machines such
as the American telephone system by a factor of 5
thousand or 10 thousand. The content of a man is
about 2 X 1028 whichexceeds the content of all librar-
ies on earth. It would be interesting to compare the
content of a child with that of a highly educated
person. He does explain the paradoxical difference of
14 orders between the content of our germ cells and
those of an adult.

Order theory must involve complex and simple
structures like the spinal cord, the tail of a sperm, and
chemical processes. As examples of the latter he cites
the turning on and off of processes by chemical
“switches” including synchronous switches, regulator-
repressors, and sequential switches. Complexes of
these he calls order-on-order. He believes that there is
little chance of complex systems being dismantled or
seldom if ever being turned off. However, I can think
of one, at least, the loss of eyes in cave fishes. But he
does reemphasize the importance of the new building
on the old, or transforming the old, in evolution, or
more simply, building blocks.

One chapter is devoted to standard part patterns of
order. These are structures which agree so well that
there is no doubt left of identical determinative laws.
Examples range from elementary physical particles
through cells and organisms to societies.

In another chapter arrangement of hierarchies of
order, their constancy, and variation are examined.
As an example of constancy he cites the posterior
rectus muscle, little changed in 400 million years of
evolution of man from fish. He points out that the
burden (complexity), and constancy (duration) of
hierarchical blocks are positively correlated, that is,
complex structures are likely to persist.

Part of the delight of the book is examining old
facts in a new light, the linkage of warm-bloodedness
and fur (or feathers), as an example of traditive inher-
itance in man, “A fish fin is used for playing the piano,
an archaic olfactory brain has been adopted to pro-
duce logic; and an original torpedo design was manip-
ulated in tetrapods ina bridge design and finally come
to be balanced on two legs, these being the behind
pillars of the bridge.” Enjoyment is also provided by
the extent of the author’s presentation using examples
from history, embryology, and anatomy. The last fea-
ture I enjoyed was the philosophical approach. He
contradicts, rightly I think those authors who distin-
guish between rules and laws of nature, for example.
Rules to Riedl are imprecisely recognized laws where
either accident reigns or further laws of an unknown
kind are at work.

Some readers may be unable to cope with the
mathematical formulae or German quotations. But
there is enough left from which they will still benefit.

Ried] has refreshingly recaptured much old wine in
new bottles. This is a book to be recommended to the
philosophically inclined or to those who wish to
enlarge their horizons.

DON E. MCALLISTER

National Museum of Natural Sciences, Ottawa, Ontario
KIA 0OM8

1981

BOOK REVIEWS 12)

The Environmental Effects of Forestry Operations in Alberta: report and recommendations

By the Environmental Council of Alberta, Edmonton. 181
pp., illus. Free.

This report is the published result of a fact-finding
mission designed to examine the environmental
effects of logging operations in Alberta. It isan erudite
synopsis of a series of public hearings held throughout
the province, and although broad in scope, it deals
with many of the problems confronting resource
managers who are responsible for the utilization of
public lands.

Major themes include the development of intensive
forest management practices which are integrated to
represent other resources and interests, improving the
administration of public lands, initiation of research
(e.g., on multiple-use and integrated land-planning
procedures) and improved data collection programs,
improvement of planning and construction tech-
niques, and the development of dynamic land-use
practices that best serve the natural resources of
Alberta.

The volume is divided into four sections. Section
one introduces the reader to forestry in Alberta by
reviewing forest policy, history, administration, man-
agement, and annual operating plans. Section two Is
an overview of the public hearings. Public concern
ranging from soil erosion to fish and wildlife and
recreation is presented.

The recommendations, comprising section three,
are perceptive, and in most cases useful and well-
grounded; this section justifies the existence of the
publication. There are 140 recommendations to
improve forestry operations in the province. At the
risk of failing to give the positive recommendations
ample review, and in hope that the following com-
ments will be viewed as constructive criticism, I will
dwell upon a few of the weak points in this section.

Recommendation 12 is particularly bothersome.
The recommendation states that “. . . the present
general regulation of minimum regeneration height of
6 feet or 20 years between cuts seems to be an unwel-
come compromise. Conditions between those
favoured by industrial foresters and wildlife biologists
have been adopted as a province-wide uniform rule
when site-specific compromises would solve more
problems. However, until fish biologists, game biolo-
gists, and foresters can agree on a more appropriate
set of figures these should be retained ... The Council
recommends that portions of each forest (area to be
determined by fish biologists, game biologists, and
foresters) be handled under different rules to suit the
relevant prime uses .. .” Prime use means land-use

management through the development of zones, a
concept that does not allow for the equal expansion of
other resources. Sound management is to develop a
guideline system flexible enough to change with the
natural dynamics of the forest systems. This requires
site-specific evaluation on behalf of all of the interest
groups, culminating in comprehensive guidelines gov-
erned bya time schedule that dictates the implementa-
tion of appropriate action.

The prime-use concept assumes a two-cut cycle,
which is unacceptable to me, as a biologist, especially
in view of our knowledge of wildlife habitat require-
ments and modified cutting procedures. Current prac-
tices involve clear-cutting 50% of the mercantile
timber within watersheds in cutblocks on the first cut.
The second or residual cut 20 years later (or after 6 to
8 ft (1 ft = 0.3048 m) regeneration on the initial cutb-
locks) is not conducive to climax forest-dwelling spe-
cies (e.g., Pine Martin, Woodland Caribou, and toa
certain degree, Wolverine and Grizzly Bear). Gener-
ally, 20-30 ft is the minimum height required for win-
ter cover for some large ungulates. Not only will the
loss of old growth habitat eliminate the species re-
stricted to mature and over-mature stands, it reduces
the numbers of species which utilize a broad spectrum
of habitats.

Recommendation 45 states that Forest Manage-
ment Agreements (FMAs) associated with new mills
should be undersized to promote the establishment of
local markets for mill-site wastes. The concept of util-
izing mill-site wastes is sound, but to undersize FMA
areas can be extremely detrimental to wildlife popula-
tions. Icontend that to manage effectively for wildlife,
longer rotation rates, involving 3- to 4-cut cycles, are
necessary. In addition, strategically placed buffer
strips of ample area are required to promote and
maintain ungulate and furbearer populations. To
reduce the FMA areas means to harvest more inten-
sively at the expense of critical wildlife habitat.

Section four contains a forest management map, a
map of the fish and wildlife zones and recreation
areas, an Outline of active research programs, and a
glossary.

The book is well worth the reading and is recom-
mended to professional resource managers in all
fields.

PAUL A. GRAY

Wildlife Service, Government of the Northwest Territories,
Yellowknife, Northwest Territories -X1A 2L9

122

Soil Organisms as Components of Ecosystems

Edited by U. Lohmand T. Persson. 1977. Proceedings of the
4th International Soil Zoology Colloquium, Uppsala,
Sweden, June 1976. Ecological Bulletins Volume 25.
NFR, Stockholm, Sweden. 614 pp., illus. 140 SwCr.

Considering that the soil of North America Is one of
its greatest assets, and, within normal time spans, a
nonrenewable resource, one can only continue to be
amazed at the small number of North American
scientists studying the biology of soil organisms. This
weakness is reflected in the authorship of the articles
in this book, for only 11 of this book’s 158 contribu-
tors are from North America. Twenty of 94 contribu-
tions are in languages other than English (usually
French).

The editors have divided the book into five sections
roughly corresponding to the session order of the
Colloquium, the first four of which deal with the
following themes: (1) Community structure and niche
separation, (2) Soil organisms in nutrient cycling, (3)
Plant roots in the soil system, and (4) Simulation
models of soil organisms. The fifth section gives sum-
maries of presentations in the Poster Session. There is
some overlap between sections, and, at times, a lack of
focus among articles within the theme sections.

The first section begins with a description of the
Organization, distribution, habitats, and food of a
wide variety of soil animals, in a well-organized and
succinct article by J. M. Anderson. The rest of the
section attempts to illustrate aspects of community
structure using a wide variety of soil animals
(amphipods to proturans).

The second section is an effort to define the role of
soil animals in soil quality. Reichle’s summary of
Research problem areas on pp. 152-153 is quite
cogent. Biologists and ecologists outside of soil zool-
ogy are frequently unaware of the importance of soil
fauna as soil modifiers allowing more rapid coloniza-
tion of soil by microbes, and as regulators of microbial
biomass and respiration.

Section 3 concerning plant roots in the soil was a
revelation. Soil biologists often overlook the point
that from one-half to two-thirds of the organic matter
in soil is due to the mortality of plant roots rather than
detrital fall onto the soil surface. Probably because of
the relatively narrow terms of reference, this section
had the best focus.

No recent ecology conference could be considered
complete without a section on ecological models, and
one is included. Although I would agree with the

THE CANADIAN FIELD-NATURALIST

Vol. 95

remarks of P. Berthet that models “oblige us to clarify
our ideas,” I often feel that the models presented at
ecology symposia are divorced from reality and not
tested rigorously (1.e., the models are limited by the
constraints of the data used to develop them, and asa
result are inapplicable to real situations). These com-
ments probably apply to this section, too. The funda-
mental weakness in attempting ecological models in
soil biology is the yawning abyss between what little
we know, but need to know, to construct sound
models.

The final section contains a mixed bag of 41 short
papers. For example, there are brief but excellent
articles by Dindal and Levitan on The soil inverte-
brate community of composting toilets, Hagvar and
Abrahamsen on The effect of artificial acid rain on
soil organisms, and Perfect et al. on The effect of DDT
on soil organisms in Nigeria.

I found several of the discussions following papers
particularly useful in exposing weaknesses in meth-
odology and experimental design. The student could
read them with considerable profit.

Obviously soil biology is still at the stage where
descriptive studies still predominate probably because
relatively little is known about even which species
comprise a given soil community. Until descriptive
problems are substantially solved, experimental stu-
dies on the functions of soil animals and communities
will continue to lag.

This book is only available as a not very sturdy
paperback. It will appeal mostly to students and
workers in soil bioiogy, but I can recommend selected
readings, including the summary papers of Ghilarov
and Satchell, to nonspecialists, and ecologists who
look upon the soil as a “black box” in their model
diagrams.

This book provides a reasonable reflection of the
quality and direction of current studies of soil ecosys-
tems. Even casual reading makes obvious the vast
diversity of soil organisms and their myriad interac-
tions. One also becomes aware of the enormous diffi-
culties under which soil biologists labor in sampling
the soil environment to construct some idea of what
goes on in that nether world under our feet.

A. D. TOMLIN

Research Institute, Agriculture Canada, London, Ontario.
N6A 5B7

1981

Exploring Nature with Your Child

By D. Shuttlesworth 1977. Abrams, New York (Canadian
distributor Prentice Hall, Toronto). 240 pp., illus., U.S.
$18.50.

I have experienced the joys and frustrations of
exploring nature with children. I had expected to
learn from Dorothy Shuttlesworth’s volume but have
not. The title Exploring nature with your child
implied to me that the theme revolved around an
active participation between parent and child. How-
ever, the book is, at best, a passive treatment of this
theme. The author does not directly address the prob-
lems confronted by the parent with children of differ-
ent ages, abilities, and interests. Only a few examples
of activities that can be undertaken by the parent and
child are provided. Most of the book is a superficial
survey of the plant and animal kingdom, and the
physical or abiotic component of nature is completely
ignored.

The author has taken the standpoint that education
of the parent is the key to exploring nature with a child
successfully. However, information alone is insuffi-
cient. Children have an uncanny knack of asking
questions that often require complicated answers. The
author has provided examples of questions that “your
young explorer” might ask but in most cases these
questions could even be asked by an uninformed par-
ent. The answers to example questions are not
adapted to the special problems of communicating
with a child.

The book begins with a discussion of exploring
nature in a changing world and following chapters
deal with birds, mammals, marine life, fish, reptiles
and amphibians, zoo animals, pets, spiders, insects,
and plants. The author did not include a preface and
concluding remarks are lacking. An index is provided
at the back of the book and is convenient to use.
Chapters on fish, zoo animals, and pets are interesting
and are more appropriate to the theme of exploring
nature withachild. The book as a whole, however, has
several downfalls that result from poor format, mis-
leading or unclear statements, and inadequate
explanations.

Although subsections conveniently divide the text,
the titles of the subsections do not always reflect the
content. For example, the subsection entitled “Expert
excavators—moles and prairie dogs” contained discus-

BOOK REVIEWS

13

sions of these as well as the range of marmots, the
folklore of groundhogs, and the defence mechanism
of the skunk.

The book is profusely illustrated with stunning pho-
tographs and artwork that complement and streng-
then the discussions in the text. However, the captions
accompanying the illustrations are often repetitive of
the text and the illustrations are not referenced in the
text. This reduces the readability of the book and
makes it inconvenient to use.

Misleading or unclear statements occur throughout
the book. “The beaver is a builder that wastes no
material” (p. 55) is a misleading statement because
beavers certainly do not utilize all that they harvest. I
think the author intended to say that the beaver is
efficient in its utilization of tree parts. Other confusing
statements result from poor grammar, suchas, “When
Spaniards sailed back to Europe taking along some
little ‘pigs’ they had discovered they frequently madea
stop at Guinea on the western coast of Africa” (p.167).
The directions to build a birdhouse on page 38 are not
easily understood and the accompanying photograph
of a birdhouse is not of the same design. An illustra-
tion of different life stages of the frog is confusing
because it is not properly labeled and the drawings of
each stage are not proportionate. Text from at least
three paragraphs on pages 86 and 87 is missing.

The author has attempted to illustrate the diversity
of nature and in doing so neglects to provide detailed
discussions. This limits the usefulness of the book
because children often ask questions that require full
but simple explanations. For example, the discussion
of how a tadpole breathes (p. 132) is not clear and
lacks the detail required to explain this very important
and basic function.

Although the book is handsomely illustrated and
does contain some interesting sections. I would not
recommend it to a person that already has a basic
understanding of natural history. Parents and inter-
pretive naturalists who are looking for novel ways to
explore nature with a child will be disappointed with
the text.

J. DAVIE

451 Marsh Road NE, Calgary, Alberta T2E 5B2

124 THE CANADIAN FIELD-NATURALIST

Island Forest Year: Elk Island National Park

By Deirdre Griffiths. 1979. University of Alberta Press,
Edmonton. x + 257 pp., illus. $12.50.

Elk Island National Park, 40 km east of Edmonton,
Alberta, lies in a southern island of mixed boreal
forest. Because of past fires and clearing, this “island
forest” is now a subclimax poplar parkland. As the
author states, “At first glance Elk Island seems
monotonous, ...” but it does have “. . . an intriguing
diversity...” Itis refreshing that Elk Island maintains
this natural diversity on 195 km? of land so close toa
major metropolis, and while almost surrounded by
extensive agricultural lands.

Griffiths states that “. . . for the writer to evolve
beyond plain description a place must be experienced
day to day . . . Factual observation and recording
transcend to empathy when the human becomes an
integral element in countless interacting events, many
of them fleeting, and fragmentary, and minor, but all
facets of a unity.” According to the publisher’s promo-
tional pamphlet, the book is “. . . to be enjoyed and
reread by anyone from the mid-teens on, who has even
the slightest interest in the outdoors ... and the
experienced naturalist.”

The book has a hard cover with a handsome dust
cover depicting some of the wildlife, plants, and habi-
tats of the park. No serious topographical errors were
found. The 232 pages of text are indented 9 cm on
every page and only 11% of this space is used for
illustrations. This is a disconcerting waste of paper
considering high book prices and the intended readers
who are probably sensitive to waste of resources. The
book contains 60 of the author’s drawings. These are
attractive but undistinguished.

The first chapter is an outline of the park’s history
and environmental setting. It describes the dominant
natural features and provides information of use to
most naturalists. Although published in 1979, the
author uses only British measurement units without
metric equivalents.

The rest of the text contains chapters for each
month from December through to the following
December. Each chapter describes the events expe-
rienced by the authorin the field. This design is highly
reminiscent of Aldo Leopold’s classic, A sand county
almanac (1949, 1966). Griffiths fails, where Leopold
did not, in presenting the unity of natural events.

Vol. 95

Griffiths does not manage to rise above the fleeting,
fragmentary, and minor events she experiences.
Repeatedly, there are lists of species and events. She
shows little selectivity in the events she records in
terms of their significance to the whole. Often she
mentions an encounter with an animal or its sign (e.g.,
a Moose track) without any description of its appear-
ance or why it might be seen, and where and when it
was seen. The layman with an interest in ecology
probably will not find enough information to develop
an understanding of ecological interrelationships. The
reader cannot develop an empathy for the natural
features described by the author.

Griffith’s writing style is tedious, nonstimulative,
and difficult. The frequent listing will tire many read-
ers. The poetic style is not suitable for the prose form
the author has selected.

The book includes annotated checklists of the
mammals, birds, reptiles, amphibians, fish, inverte-
brates, and plants of Elk Island. This is by far the most
useful section of the book. However, her usage of
scientific names from J. D. Soper’s The mammals of
Alberta (1964) is somewhat outdated with terms such
as Citellus spp. and Euarctos americanus. The bird
checklist is updated to 1978.

There is also a glossary, a bibliography, and an
index. The glossary and bibliography will be useful to
the layman, although they are not extensive. The
index is not complete. Among a list of birds given in
the text, one or more may not be indexed. Other
omissions can also be found.

Elk Island National Park is a very interesting and
accessible area for natural history study and observa-
tion. However, there has beena lack of popular litera-
ture about this park. It is unfortunate that /s/and

forest year has not filled this gap in a readable and

informative way. This book would be suitable for
public library shelves to be borrowed and browsed. At
$12.50 a copy, I cannot recommend this text to natu-
ralists or others as a useful addition to their personal
libraries.

MICHAEL A. D. FERGUSON

Department of Wildlife Ecology, University of Wisconsin,
1630 Linden Drive, Madison, Wisconsin 53706.

1981

MISCELLANEOUS

Photography and the Art of Seeing

By Freeman Patterson. 1979. Van Nostrand Reinhold Ltd.,
Toronto, Canada. 156 pp., illus. $12.95.

Freeman Patterson is developing a large reputation
as an extraordinary photographer and writer. Images
such as those seen in Canada: a year of the land have
brought Patterson many prestigious awards such as
The Hon. EFIAP, and much recognition as a photo-
grapher. Patterson’s writing ability is clear in the
numerous articles published in Photolife, Photo Can-
ada, and other popular photographic periodicals.
However, it was not until the publication of his recent
book, Photography for the joy of it, that the public
became aware of his combined talents as photo-
grapher and author.

Patterson’s most recent book is equally as informa-
tive and enjoyable as the last. The topic of the present
work, that of composition in photography, is abstract
and often elusive to describe because of the difficulty
in expressing verbally how one feels about a photo-
graphic image. Patterson, with an obvious artistic
awareness and much technical experience, has devel-
oped the ability to bridge the gap between impres-
sionism and technical know-how. The result is a sim-
plistic treatise of an inherently difficult photographic
topic.

The text is divided into two parts. In the first half,
Patterson introduces the reader to many enlighting
concepts of photographic composition. One such
concept is what Patterson describes as thinking side-
ways. This is defined as the process of abandonment
of normal photographic standards in search of origi-
nal composition.

One method of thinking sideways is to make a list of
three or four items accepted as essential elements to
photography. One such item would be to keep the
subject always in focus. The photographer then pro-
ceeds to break these rules. Patterson admits that on
many occasions the resultant image may not be

BOOK REVIEWS 125

appropriate. But experimentation is integral to the
thinking sideways scheme and is essential to the
development of new composition technique.

The second section of the text relates the authors’
concepts of sound mental impressionism in composi-
tion to the technical limitations of the camera. The
knowledge that the camera is not a magic box that
captures exactly what the eye sees is germane to
improving composition in a photograph. Patterson
explains that seeing in photography includes knowing
what the camera sees. He describes what the camera
actually perceives and how the photographer can use
this knowledge to his or her advantage.

A detailed discussion on visual design completes the
second portion of the text. Visual design is necessary
to make a good image because it is concerned with
expressing one’s subject in the most effective manner.
This discussion is, therefore, very important to all
photographers.

This book is applicable to those making wilderness
photographs. Nature photographs, especially those of
animals and plants, have previously simply attempted
to capture the subject in full frame. However, at pres-:
ent these types of shots must be original and even
impressionistic to enthuse the modern nature photo-
critic. Therefore, the ability to see photographically
can certainly improve nature photography through
introducing original theme and design into the
images.

This book is very well presented and therefore
highly recommended. The image reproduction is
excellent, which is a credit to the publisher; and the
text is understandable, which is a credit to the author.

ALAN J. KENNEDY

Canadian Wildlife Service, #1000, 9942 - 108 Street, Edmon-
ton, Alberta TSK 2J5

126 THE CANADIAN FIELD-NATURALIST

NEW TITLES

Zoology

Arctic summer: birds in north Norway. 1979. By Richard
Vaughan. Buteo, Vermillion, South Dakota. 152 pp., illus.
U.S. $17.50.

Birds of the Labrador peninsula and adjacent areas: a dis-
tribution list. 1980. By W. E. Clyde Todd. Reissue of 1963
edition with corrections and addendum. Buteo Books,
Vermillion, South Dakota. 822 pp., illus. U.S. $70. (New
pages available separately U.S. $1 for handling.)

Bright waters, bright fish. 1980. By Roderick Haig-
Brown. Douglas and McIntyre, Vancouver. 144 pp., illus.
$19.95.

Butterflies and moths of Newfoundland and Labrador: the
Macrolepidoptera. 1980. By Ray F. Morris. Report 1691.
Agriculture Canada, Ottawa. 407 pp., illus. $15 in Canada;
$18 elsewhere.

Catalog of Hymenoptera in America north of Mexico.
1979. Prepared under the direction of K. V. Krombein,
P. D. Hurd, Jr., D. R. Smith, and B. D. Burks. Superin-
tendent of Documents, Washington. 3 volumes. 2736 pp.
U.S. $78.

Cat behavior: the predatory and social behavior of domes-
tic and wild cats. 1979. By Paul Leyhausen. Translated by
B. A. Tonkin. Garland STPM Press, New York. xvi + 340
pp., illus. U.S. $24.50.

*Charrs: salmonid fishes of the genus Salvelinus. 1980. By

Eugene K. Balon. Perpectives in Vertebrate Science
Volume |. Junk, The Hague. ix + 928 pp., illus. U.S.
$210.60.

A country-lover’s guide to wildlife: mammals, amphibians
and reptiles of the northeastern United States. 1979. By
Kenneth A. Chambers. Johns Hopkins University Press,
Baltimore. xviii + 230 pp., illus. U.S. $14.95.

Ecology of Great Horned Owls and Red-tailed Hawks in
southeastern Wisconsin. 1979. By L. Petersen. Technical
Bulletin Number I11. Wisconsin Department of Natural
Resources, Madison. 63 pp. No price given.

Evolutionary history of the primates. 1979. By Frederick
S. Szalay and Eric Delson. Academic Press, New York. 608
pp., illus. U.S. $43.

The evolution of culture in animals. 1980. By John Tyler
Bonner. Princeton University Press, Princeton. x + 218 pp.,
illus. U.S. $14.50.

The George Reserve deer herd: population ecology of a
K-selected species. 1979. By Dale R. McCullough. Uni-
versity of Michigan Press, Ann Arbor. xiv + 272 pp., illus.
U.S. $16.

Vol. 95

Greenshanks. 1979. By Desmond and Maimie Nethersole-
Thompson. Buteo, Vermillion, South Dakota. 272 pp.,
illus. U.S. $27.50.

+Handbuch der Vogel Mitteleuropas, Band 9: Columbi-
formes bis Piciformes. 1980. Akademische Verlagsgesell-
schaft, Wiesbaden. 1152 pp., illus. DM208.

+The Hawaiian Goose: an experiment in conserva-
tion. 1980. By Janet Kearand A. J. Berger. Buteo, Vermil-
lion, South Dakota. 154 pp., illus. U.S. $30.

Herbivores: their interaction with secondary plant metabol-
ites. 1979. Edited by Gerald A. Rosenthal and Daniel H.
Janzen. Academic Press, New York. 744 pp. U.S. $59.50.

+Hydromedusae of British Columbia and Puget
Sound. 1980. by M.N. Arai and A. Brinckmann-Voss.
Canadian Bulletin of Fisheries and Aquatic Sciences 204.
Supply and Services Canada, Hull. viii + 192 pp., illus. $13
in Canada; $15.60 elsewhere.

tInnovative teaching in aquatic entomology. 1979. Edited
by V. H. Resh and D. M. Rosenberg. Canadian Special
Publication of Fisheries and Aquatic Sciences Number 43.
Supply and Services Canada, Hull. 114 pp., illus. $4 in
Canada; $4.80 elsewhere.

Mesozoic mammals: the first two-thirds of mammalian
history. 1980. Edited by J. A. Lillegraven, Z. Kielan-
Jaworowska, and W. A. Clemens. University of California
Press, Berkeley. x + 312 pp., illus. Cloth U.S. $35; paper
WESH SOA:

Museum studies and wildlife management: selected
papers. 1979. By R.C. Banks. Smithsonian Institution
Press, Washington. Available from Museum of Natural
History, University of Kansas, Lawrence. 297 pp. U.S. $2 in
North America; U.S. $3.25 elsewhere.

+ Ornithological and ecological studies in S. W. Greenland
(59° 46’-62° 27’ N.Lat.). 1979. By Finn Salomonsen. Nyt
Nordisk Forlag Arnold Busck, Kobenhavn. 214 pp., illus. +
3 plates. Paper 160 Danish Crowns.

Pollution ecology of estuarine invertebrates. 1979. Edited
by C. W. Hart, Jr., and S. L. H. Fuller. Academic Press,
New York. 432 pp. U.S. $28.50.

Predator-prey systems in fisheries management. 1979. Ed-
ited by Henry Clepper. Papers froma symposium, Atlanta,
July, 1978. Sport Fishing Institute, Washington. vii + 504
pp., illus. U.S. $20.

Proceedings of management of north central and north-
eastern forests for nongame birds. 1979. Compiled by
R. M. DeGraaf and K. W. Evans. U.S. Department of

1981

Agriculture General Technical Publication NC—SI. North
Central Experimental Station, St. Paul, Minnesota. 268 pp.
No price given.

Reindeer and caribou hunters: an archaeological study.
1980. By Arthur E. Spiesse. Academic Press, New York.
xiv + 314 pp., illus. U.S. $25.

Reproduction of marine invertebrates. Volume 5, molluscs:
pelecypods and lesser classes. 1979. Edited by Arthur C.
Giese and John S. Pearse. Academic Press, New York. 392
pp. U.S. $45.50.

Songbirds in your garden. 1980. By John K. Terres. 3rd
edition. Clarke Irwin, Toronto. 289 pp. $7.75.

Splendid isolation: the curious history of South American
mammals. 1980. By George Gaylord Simpson. Yale Uni-
versity Press, New Haven. x + 266 pp., illus. U.S. $17.50.

+Toxicity tests for freshwater organisms. 1979. Edited by
E. Scherer. Canadian Special Publication of Fisheries and
Aquatic Sciences 44. Department of Fisheries and Oceans,
Winnipeg. 194 pp. Free.

Vertebrate ecology in the northern neotropics. 1980.
Edited by John F. Eisenberg. Smithsonian Institute Press,
Washington. 272 pp., illus. Cloth U.S. $17.50; paper U.S.
$8.95.

Wildlife and America: contributions to an understanding of
American wildlife and its conservation. 1978. Edited by
Howard P. Brokaw. Superintendent of Documents,
Washington. x + 532 pp., illus. U.S. $8.50 plus 25% foreign
handling.

Wildlife habitats in managed forests, the Blue Mountains of
Oregon and Washington. 1979. Edited by J. W. Thomas.
Agricultural Handbook Number 553. U.S. Forest Service,
Washington. 511 pp. U.S. $14.

Botany

Algal assays and monitoring eutrophication. 1979. Edited
by P. Marvan, S. Pribil, and O. Lhotsky. Schweizer-
bart’sche, Stuttgart. vill + 252 pp., illus. DM49.

The biology of Canadian weeds: contributions
1-32. 1979. Edited by Gerald A. Mulligan. Publication
1693. Agriculture Canada, Ottawa. 380 pp. Free.

*Common and botanical names of weeds in Cana-
da. 1980. By Jack F. Alex, Richard Cayouette, and Gerald
A. Mulligan. Agriculture Canada Publication 1397. Supply
and Services Canada, Hull. 132 pp. $5.75 in Canada; $6.90
else where.

Diatoms in New Zealand, the north island. 1979. By N.
Foged. Bibliotheca Phycologica 47. Cramer, Braunsch-
weig. 226 pp., illus. DM60.

Diatoms in Oland, Sweden. 1980. By N. Foged. Bibli-

BOOK REVIEWS

127)

otheca Phycologica 49. Cramer, Braunschweig. 194 pp.,
illus. DMSO.

*The flora of Canada: part 4, Dicotyledoneae (Loasaceae to
Compositae). 1979. By H. J. Scoggan. National Museums
of Canada, Ottawa. Sold only as 4 volume set (previous
volumes published 1978). xx + 1711 pp. $131.

A flora of New Mexico. 1980. By W. C. Martinand C. R.
Hutchins. Cramer, Braunschweig. c3000 pp., illus. cU.S.
$80.

*Flore des champignons au Québec. 1980. Par René
Pomerleau. Les Editions La Presse, Montreal. 652 pp..,
illus. $65.

*Grasses of Ontario. 1980. By William G. Dore and J.
McNeill. Agriculture Canada Monograph 26. Supply and
Services Canada, Hull. 566 pp. $12 in Canada; $14.40
elsewhere.

+Légumes sauvage du Canada. 1980. Par Adam F. Szcza-
winskiand Nancy J. Turner. Plantes Sauvages comestibles
du Canada, 4. Musee national des sciences naturelles,
Ottawa. 179 pp., illus. $9.95.

The orchids of the high mountains of New Gui-
nea. 1980. By P. van Royen. Cramer, Braunschweig. p.
51-834, illus. DM 200.

Physiological ecology of the alpine timberline: tree exis-
tence at high altitudes with special reference to the Euro-
pean alps. 1979. By W. Tranquillini. Translated from
German. Springer-Verlag, New York. xii + 140 pp., illus.
U.S. $29.70.

Rare and endangered vascular plants and vertebrates of
Wyoming. 1979. Edited by T. W. Clark and R. D. Dorn.
Committee for Rare and Endangered Plants and Animals
of Wyoming, Jackson. 78 pp. U.S. $10 contribution
suggested.

Responses of plants to environmental stresses. 2nd edition.
Volume 1, chilling, freezing, and high temperature stresses.
1980. By J. Levitt. Academic Press, New York. 512 pp. U.S.
$29.50.

Survey of Canadian herbaria. 1980. By Bernad Boivin.
Provancheria Number 10. Available from B. Boivin, Herb-
ier Louis-Marie, Université Laval, Québec. 187 pp. $10.

Threatened and endangered plants in selected areas of the
Bim Fortymile Planning Unit, Alaska. 1979. By A. R.
Batten, D. F. Murray, and J. C. Davie. Alaska Technical
Report 3. Bureau of Land Management, Anchorage. 127
pp. No price given.

Topics in plant population biology. 1979. Edited by O. T.
Solbrig, S. Jain, G. B. Johnson, and P. H. Raven. Papers
from a conference, Ithaca, New York, June, 1977. Colum-
bia University Press, New York. xvili + 590 pp., illus. U.S.
$25.

128

Environment

An all-season guide to easy hiking around Vancou-
ver. 1980. By Jean Cousins and Heather Robinson. Dou-
glas and MacIntyre, Vancouver. $5.95.

The Arun: a natural history of the world’s deepest val-
ley. 1979. By Edward W. Cronin, Jr. Houghton Mifflin,
Boston. xil + 236 pp., illus. U.S. $10.95.

Bioengineering for land reclamation and _ conserva-
tion. 1980. By Hugo Schiechtl. Translation and editing
coordinated by Nick Horstmann. University of Alberta
Press, Edmonton. 400 pp., illus. $30.

Canoe routes British Columbia. 1980. By Richard and
Rochelle Wright. Douglas and MacIntyre, Vancouver. 176
pp. Paper $6.95.

Canoe routes Yukon Territory. 1980. By Richard and
Rochelle Wright. Douglas and MacIntyre, Vancouver. 112
pp. Paper $6.95.

*Circulation models of lakes and inland seas. 1980. By T. J.
Simons. Canadian Bulletin of Fisheries and Aquatic Scien-
ces 203. Supply and Services Canada, Hull. 146 pp., illus.
$12 in Canada; $14.40 elsewhere.

*Environmental effects of dams and impoundments in
Canada. 1980. By R. M. Baxter and P. Glaude. Canadian
Bulletin of Fisheries and Aquatic Science 205. Supply and
Services Canada, Hull. 34 pp. $2.50 in Canada; $3
elsewhere.

+ Estimation of density from line transect sampling of biolog-
ical populations. 1980. By Kenneth P. Burnham, David R.
Anderson, and Jeffrey L. Laake. Wildlife Monographs
Number 72. The Wildlife Society, Washington. 202 pp.,
illus. $4.

The heyday of natural history 1820-1870. 1980. By Lynn
Barber. Jonathan Cape (Canadian distributor Clarke
Irwin, Toronto). 320 pp. $27.95.

Home is the high country. 1980. By Mona Anderson.
Reed (Canadian distributor Douglas and MacIntyre, Van-
couver). 72 pp. illus. $11.95.

Human impacts on life in fresh waters. 1979. Edited by J.
Salanki and P. Biro. Papers from a symposium, Tihany,
Hungary, September, 1977. Akademiai Kiado, Budapest
(distributed by Heyden, Philadelphia). 224 pp., illus. U.S.
$22.

Land/wildlife integration. 1979. Edited by D. G. Taylor.
Proceedings of a technical workshop, Saskatoon, 1-2 May
1979. Ecological Land Classification Series, II. Environ-
ment Canada, Ottawa. 160 pp. Free.

The natural selection of populations and communi-
ties. 1980. By David Sloan Wilson. Benjamin/Cummings,
Menlo Park, California. xviii + 186 pp., illus. U.S. $12.95.

THE CANADIAN FIELD-NATURALIST

Vol. 95

+A nature guide to Alberta. 1980. Edited by David A. E.
Spalding. Hurtig, Edmonton. 368 pp., illus. $14.95.

Population dynamics: alternative models. 1979. By Ber-
tram G. Murray, Jr. Academic Press, New York. 224 pp.
U.S. $21.

Riparian and wetland habitats of the Great Plains.
1979. By the Great Plains Agricultural Council. Publica-
tion Number 91. Rocky Mountain Forest and Range
Experimental Station, Fort Collins. 88 pp. No price given.

Serengeti: dynamics of an ecosystem. 1980. Edited by
A. R.E. Sinclair and M. Norton-Griffiths. University of
Chicago Press, Chicago. x11 + 390 pp., illus. U.S. $28.50.

Miscellaneous

Bird, beast and flower. 1980. Watercolours by Marie
Angel. Poems chosen by Ilan Parsons. Clarke Irwin,
Toronto. 64 pp., illus. $14.95.

Biochemical and photosynthetic aspects of energy produc-
tion. 1980. Edited by Anthony San Pietro. Academic
Press, New York. 256 pp. U.S. $24.50.

Directory of Canadian environmental experts. 1980. By
Environment Canada. Government Publishing Centre,
Hull. $12.50 in Canada; $15 elsewhere.

The earth: its origin, structure and evolution. 1979. Edited
by M. S. McElhinny. Academic Press, New York. xvit+ 598
pp., illus. U.S. $74.50.

The enduring Great Lakes. 1979. Edited by John Rous-
maniere. Norton, New York. xiv + 356 pp. U.S. $12.95.

Energy future. Report of the Energy Project at the Harvard
Business School. 1979. Edited by Robert Stobaugh and
Daniel Yergin. Random House, New York. xii + 356 pp.
WES Si128952

Energy in America’s future: the choices before
us. 1979. By S. H. Schurr, J. Darmstadter, H. Perry, W.
Ramsay, and M. Russell. Published by Resources for the
Future by Johns Hopkins University Press, Baltimore.
xxvill + 556 pp., illus. Cloth U.S. $30; paper U.S. $10.95.

A fossil-hunter’s notebook: my life with dinosaurs and
other friends.. 1980. By Edwin H. Colbert. Dutton Cana-
dian distributor Clarke Irwin, Toronto. 288 pp., $19.95.

Gathering what the great nature provided: food traditions
of the Gitksan. 1980. Edited by the Book builders of
*Ksan. Douglas and McIntyre, Vancouver. 128 pp., illus.
$18.95.

Water resources assessment: methodology and technology
sourcebook. 1979. By Larry W. Canter. Ann Arbor
Science, Ann Arbor. xxii + 530 pp. U.S. $29.50.

* Assigned for review
+ Available for review

i=

pa eae j iin}
Bi &

Instructions to Contributors

Content

The Canadian Field- Naturalist is a medium for the publi-
cation of scientific papers by amateurand professional natu-
ralists or field-biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All
readers and other potential contributors are invited to sub-
mit for consideration their manuscripts meeting these crite-
ria. 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 significance.

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 sub-
mitted. 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 biblio-
graphic 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. Under-
line only words meant to appear in italics. The names of
authors of scientific names should be omitted except in taxo-
nomic manuscripts or other papers involving nomenclatural
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 Serial Sources for the BIOSIS Data Base,
published yearly by BioSciences Information Service, Phila-
delphia, Pennsylvania 19103. 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 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. Webs-
ter’s New International Dictionary and le Grand Larousse
Encyclopédique are the authonties 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
$50 for each page in excess of six journal pages, plus $5 for
each illustration (any size up toa full page), and up to $50 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 $50 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.

Reviewing Policy of The Canadian Field-Naturalist

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, whois 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

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)
Notes (continued)

An usually small American Eel (Anguilla rostrata) from the Lake Superior drainage
PHILIP A. COCHRAN of

Panicled Knapweed (Centaurea paniculata: Compositae) new to eastern Canada
DANIEL F. BRUNTON and PAUL D. PRATT 98

Ruby-crowned Kinglets (Regulus calendula) feeding a Brown-headed Cowbird (Molothrus ater)
PETER C. BOXALL 99

Northern record for Three-flowered Avens (Geum triflorum) in Manitoba
H. L. MUNRO and D. H. MUNRO 100

Obituary
Farrell E. Banim, O.M.I., 1902-1979 PEGGY WHITEHURST KURATA 102
News and Comment 106

Book Reviews

Zoology: Conservation of marine birds of northern North America — The insects and arachnids of 107
Canada. Part 1. Collecting, preparing and preserving insects, mites and spiders. Part 3. The
Aradidae of Canada (Hemiptera: Aradidae). Part 4. The Anthocoridae of Canada and Alaska
(Heteroptera: Anthocoridae) — The birds and birders of Beaverhills Lake — The Peregrine
Falcon in Greenland — Some adaptations of marsh-nesting blackbirds — The first ten years of
the Co-operative Breeding Bird Survey in Canada — The dragonflies of British Columbia
—Annotated list of workers on systematics and faunistics of Canadian insects and certain
related groups — Wolves of Minong — Mammals of the eastern United States — Responses of
Peary Caribou and Muskoxen to helicopter harassment

Botany: The rare vascular plants of Saskatchewan — Identifying grasses — Native trees and shrubs 116
of Newfoundland and Labrador — Vascular flora of the southeastern United States: Volume 1,
Asteraceae — Ferns of north-western Himalayas

Environment: Order in living organisms — The environmental effects of forestry operations in 120
Alberta — Soil organisms as components of ecosystems — Exploring nature with your child
—Island forest year: Elk Island National Park

Miscellaneous: Photography and the art of seeing 125
New Titles 126

Mailing date of previous issue 5 December 1980

1980 Council — The Ottawa Field-Naturalists’ Club

President: Roger Taylor Ron Bedford Bill Gummer
Vice-President: Loney Dickson Frank Bell Peter Hall
Treasurer: Barry Henson Bill Cody Don Lafontaine
Recording Secretary: Dan Brunton Ellaine Dickson Diana Laubitz
Corresponding Secretary: Frank Pope Roger Foxall Hue MacKenzie

Courtney Gilhiatt Ken Strang

Fran Goodspeed Ken Taylor

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 95, Number 1

The Ottawa Field-Naturalists’ Club Centennial Symposium
100 years of natural history in Canada

Introduction IRWIN M. BRODO
200 years of ornithology in Canada FRED COOKE
Mammalogy in Canada: a historical overview DONALD A. SMITH
Canadian entomology of the last century J. R. VOCKEROTH
Geology, 1879-1979 ROBERT F. LEGGET

Federal wildlife conservation work in Canada in the past 100 years
VICTOR E. F. SOLMAN

Viewpoint
A Canadian paradox — private land, public wildlife: can it be resolved?
J. P. RYDER and D. A. BOAG

Articles

Dynamics of Moose populations near Rochester, Alberta, 1975-1978
WILLIAM R. MYTTON and LLOYD B. KEITH

Régime alimentaire des Orignaux du sud-ouest Québécois pour les mois d’avril a
octobre
MICHEL CRETE et PETER A. JORDAN

Caribou (Rangifer tarandus) encounters with pipelines in northern Alaska
WAYNE C. HANSON

Analysis of weight lost by eggs of eleven species of birds during incubation
T. H. MANNING

Sex differential in use of the physical environment by Bighorn Sheep (Ovis canadensis)
LUIGI E. MORGANTINI and ROBERT J. HUDSON

Terrestrial molluscs of central Alberta JAMES VAN ES and D. A. BOAG

Deer mortality in the Pohénégamook wintering area, Quebec
FRANCOIS POTVIN, JEAN HUOT, and FRANCOIS DUCHESNEAU

Moose browse utilization in Mount McKinley National Park, Alaska
JERRY O. WOLFF and JOANNE COWLING

The sedge Carex /oliacea in eastern North America A. A. REZNICEK and P. W. BALL

Notes

First records of the Tundra Shrew (Sorex tundrensis) in British Columbia
DAVID W. NAGORSEN and DONALD M. JONES

Great Blue Heron (Ardea herodias) colony in the Peace-Athabasca Delta, Alberta
JOHN KRISTENSEN

1981

35

38)

50

oY,

63

69
75

81

85
89

93

O5

concluded on inside back cover

ISSN 0008-3550

0 —
0. &
aig ©
; ee
gl 2
Ran! —
© (J —
J

The CANADIAN
FIELD-NATURALIS

-

cANW
pS 27

‘

, Canada

Ottawa

CLUB,

Published by THE OTTAWA FIELD-NATURALISTS’

“~~

Se

?
.

Ze

“s “ a
SON as «
IN be

Oe
. NEE Gaal & \
Cw

April-June 1981

Volume 95, Number 2

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

C. D. 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: Patricia A. Lalla Chairman, Publications Committee: R. E. Bedford
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.

Production Manager: Pauline A. Smith, R.R. 3, Wakefield, Quebec JOX 3G0
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 iJO

Coordinator, The Biological Flora of Canada: Dr. George H. La Roi, Department of Botany, University of Alberta,
Edmonton, Alberta T6G 2E9

Associate Editor (Ornithology): Dr. A.J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick E0A 3C0
Editor: Dr. Lorraine C. Smith, R.R. 3, Stittsville, Ontario KOA 3G0

All manuscripts intended for publication should be addressed to the new Editor:
The Canadian Field-Naturalist, The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C,
Ottawa, Ontario Canada KI Y 4J5

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: Mountain Plover photographed at Onefour, Alberta, on 5 June 1979 by C. R. Wershler. For information on the
nesting and status of the species in Canada see Article on page 133.

The Canadian Field-Naturalist

Volume 95, Number 2

April-June 1981

Tiger Salamanders (Ambystoma tigrinum) and
Stocked Rainbow Trout (Sa/mo gairdneri):
Potential Competitors for Food in Manitoba Prairie Pothole Lakes

ROBERTA J. OLENICK! and JOHN H. GEE

Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
'Present address: Institute of Animal Resource Ecology, University of British Columbia, Vancouver, British Columbia

V6T 1W5

Olenick, Roberta J., and John H. Gee. 1981. Tiger Salamanders (Ambystoma tigrinum) and stocked Rainbow Trout
(Salmo gairdneri): potential competitors for food in Manitoba prairie pothole lakes. Canadian Field-Naturalist 95(2):

129-132.

Tiger Salamanders (Ambystoma tigrinum) were aquatic and coexisted with Rainbow Trout (Sa/mo gairdneri) from May until
early August, the first half of the trout growing season, in two Manitoba pothole lakes. Salamanders were benthic and ate
Gammarus almost exclusively. Trout were mainly limnetic and in May-June consumed mainly Chaoborus larvae and
Gammarus, switching almost entirely to Gammarus in July and August. The considerable overlap in diet and only partial
segregation in time (season) and space indicate a potential for competition for food. This is probably minimized by differences

in mouth size and the plasticity of feeding by trout.

Key Words: Ambystoma tigrinum, Salmo gairdneri, diet, spatial and temporal distributions, interspecific competition.

While salamanders have minimal effect on fish pro-
ductivity in large lakes, their influence in smaller
water bodies, where they are abundant, may be con-
siderable (Efford and Tsumura 1973). Densities of
Tiger Salamanders, Ambystoma tigrinum diaboli, in
prairie pothole lakes range from 100 to 364 salaman-
derssha | (Tavarutmaneegul 1978; Meyers 1973).
While these lakes do not contain indigenous, com-
mercially valuable fishes, some are currently used for
aquaculture of Rainbow Trout, Salmo gairdneri
(Lawler et al. 1974). Efford and Tsumura (1973) state
that, because salamanders can be important competi-
tors for food in fish ponds, the role of salamanders
should be considered in the development of aquacul-
ture programs. Thus, the present study was under-
taken to describe the diet of Tiger Salamanders in
stocked and unstocked pothole lakes, and to assess
dietary similarity between salamanders and trout in
stocked lakes. Data on mouth size and vertical distri-
bution provided further information on the potential
for competition for food between these two species.

Materials and Methods

This study was conducted in 1977 on Lakes 156 and
675 of an aquaculture study area near Erickson,
Manitoba (Sunde and Barica 1975). Lake 156 is circu-

lar and has a surface area of 6.9 ha,a maximum depth
of 1.8 m, anda substrate of soft mud. Aquatic vegeta-
tion includes dense mats of Aphanizomenon flos-
aquae, Chara, and emergent Scirpus and Phragmites.
It contained no trout. Lake 675 is rectangular and has
a surface area of 9.7 ha, a mean depth of 2.6 m, and a
substrate of mud. Vegetation includes Cladophora,
Myriophyllum, Ceratophyllum, Chara, and emergent
Scirpus and Typha. A woven polyethylene sea cur-
tain, constructed in late May 1977, divided Lake 675
into approximately equal north and south portions.
These were stocked with about 3500 and 10 000
trout:ha |, respectively.

Salamanders and trout were collected during the
spring and summer (Table |) as specimens for analysis
of vertical distribution, mouth size, and diet. Gill nets
(3 m deep) varying from 13 to 5I-mm stretch mesh
were normally used, but in Lake 675 gillnetting was
restricted at certain times to accomodate other
research programs. Here, some samples for stomach
analysis were obtained by seining. Gill nets were
examined every 4 h and time of capture and vertical
position were recorded. Specimens were preserved in
20% formalin and later transferred to 75% isopropyl
alcohol. Mouth width (inside distance between
corners of mouth) and mouth length (distance from

129

130 THE CANADIAN FIELD-NATURALIST

TABLE |—Number and dates of capture of salamanders and
trout collected by gill net or seine. Trout were not stocked in
Lake 156

Lake 156 Lake 675
Numbers 12 May- 1J8July- 18 May 20 July
captured 22 June 10 Aug. 30June 12 Aug.
Salamanders
Larvae 27 119 23 20
Adults 15 0 70 2
Trout — — 91 29

corner to anterior tip of premaxilla) were measured
with vernier calipers (+0.1 mm).

Food items were identified to genus wherever pos-
sible and numbers in each taxon were recorded. The
weight of each taxon was estimated by multiplying
numbers eaten by the average weight of an undigested
item. The proportion of individuals that had eaten
each taxon (frequency of occurrence) was also deter-
mined. Parasites, vegetation, and detritus were not
included in the analysis. Some taxa were grouped for
within- and between-species comparisons of the per-
centages of number and percentages of weight of food
categories eaten. Comparisons were made using Mor-
isita’s index, CA, of overlap (complete dissimilar-
ity = 0; complete similarity = 1.0) as modified by Horn
(1966).

Results and Discussion

Trout were planted in pothole lakes in late May and
harvested in October and November. Adult sala-
manders migrated into lakes in late April and early
May before breeding and remained until late July and
early August. Larvae were first observed on 13 June
and 22 June in Lakes 675 and 156, respectively. Newly
transformed salamanders were seen on land on 15
August. Salamanders were aquatic during the first
half of the growing season for trout and, therefore,
coexistence occurred at a time when the rate of growth

Vol. 95

of trout was greatest (J. A. Mathias, Freshwater Insti-
tute, Winnipeg, Manitoba, personal communication).

Salamanders and trout were caught most often at
night with trout mainly taken in mid to upper waters
while salamanders were benthic (Table 2). The ratio of
the number of salamander larvae in upper and lower
portions of gill nets differed significantly (P < 0.005,
chi-square test with Yates’ correction for continuity)
between day and night as some larvae moved upwards
in darkness. The ratio of salamander larvae in upper
and lower portions differed significantly (P < 0.005)
from that of trout for both day and night samples as
did salamander adults and trout with day and night
combined.

Gammarus were found in all salamander stomachs
and made up over 80% of the number and weight of
items eaten. Comparisons between lakes, sampling
periods, and adults and larvae revealed high similarity
in both numbers and weight of items eaten
(CA > 0.97). Because there were no intraspecific dif-
ferences between north and south portions of Lake
675 in the diet of either trout or salamanders, samples
from the two portions were combined. In Lake 675 in
spring, 73% of the items consumed by trout were
Chaoborus larvae, but in terms of weight, Chaoborus
larvae and Gammarus were of similar proportion and
both items occurred in over 70% of the stomachs
(Table 3). The diets of salamanders and trout in Lake
675 in spring were dissimilar. In summer Cladocera
and Gammarus made up similar proportions of the
number of items eaten by trout, but Gammarus con-
tributed 69% of the weight of the food and occurred in
all stomachs. The diets of salamanders and trout, in
terms of weight of items eaten, were very similar dur-
ing summer (Table 3). The vertical distribution of
Gammarus in the water column is primarily benthic
(J. A. Mathias, personal communication).

Mouths of trout and larval salamanders from Lake
675 in June were similar in width but were longer in
trout (Table 4). In July mouth length was similar but
mouths of larvae were wider than trout. Adult sala-
manders had much larger mouths than trout. Mouth

TABLE 2—Comparison of depth of capture in gill nets of salamanders in Lake 156 and salamanders and trout in Lake 675.
Upper is top two-thirds, lower is bottom third. No larvae were captured in gill nets in Lake 675 because of restrictions on mesh

sizes.
Lake 156 Lake 675
Salamander Galamander
Larvae Adults adults Trout
Upper Lower Upper Lower Upper Lower Upper Lower

Day 3 33 | 6 12 13 10 7
Night 142 186 13 22, 25 56 51 19

Total 145 219 14 28 37/ 69 61 26

1981

TABLE 3—Comparison of diet of salamanders (S) and trout
(T) in spring (18 May - 30 June) and summer (20 July — 12
August) in Lake 675. Adult and larval salamanders are com-
bined. | = larvae; p = pupae

Spring
Number, % Weight, % Occurrence, %
S T S T S TT
Hirudinea 0.1 O 10 0O 3.2 0
Cladocera O6 7 - 0 0.6 Dp) 27.5
Gammarus 85.9 10.9 87.1 38.2 100.0 73.6
Hyalella 04 #05 O1 08
Corixidae SON 24 04 2457, 4.4
Dytiscidae (1) 12 O02 65 33 20 14.3
Chaoborus (1) 0.4 73.5 04 496 15.1 87.9
Chironomidae
(I, p) I® S37 O02 309 ies 36.3
Other 5-5 oli8' 25, Bk —
Cr 0.15 0.59
Summer
Hirudinea 0.3 O05 7.6 20.6 15.0 34.4
Cladocera 0 424 O 1.3 0 62.0
Gammarus 80.3 38.9 87.2 68.9 100.0 100.0
Hyalella 0 48 0 2.8
Corixidae 14 04 07 O02 40.0 44.8
Dytiscidae (1) 0.1 0.2 06 1.0 5.0 13.8
Chironomidae
(1, p) 16.8 9.2 3.7 3.0 30.0 62.1
Other 10) 3:6), 074 2.2 — —
Chr 0.65 0.96

size limits maximum size of prey taken by salamand-
ers (Dodson and Dodson 1971) and by trout (Hart-
man 1958). The larger mouth size of salamanders in
July, when both species fed on Gammarus exten-
sively, may have allowed salamanders to eat larger
individuals. While mouth size differences may have
been important in Lake 675 in 1977, in general relative
mouth sizes of co-occurring salamander larvae and

OLENICK AND GEE: SALAMANDER-T ROUT FOOD COMPETITION, MANITOBA

131

trout depend on time of trout stocking and growth
rates of the two species.

The diet of salamanders described here is almost
identical with that shown by A. K. Maclem (Fresh-
water Institute, Winnipeg, Manitoba, unpublished
data). He examined 103 adults from 14 lakes in June
and 97 larvae from 8 lakes in July in the Erickson area
and found 82% of the number of.items consumed were
Gammarus. The extensive utilization of Gammarus
by trout indicates a potential for competition should
Gammarus be limited in supply. That stomachs of
nearly all individuals of both species were full suggests
an abundance of food in Lake 675 in 1977.

The severity of competition for food can be reduced
if species feed at different times or in different places.
Weare uncertain of time of feeding of the two species,
but there are differences in places of feeding. Adult
and larval salamanders were observed to feed while on
the bottom when held in aquaria (A. K. Maclem,
unpublished data). Their high frequency of capture
along the bottom of gill nets, and the occurrence of
vegetation and detritus in 33% of stomachs support
this observation. Trout are known to consume benthic
prey but they also feed in mid to surface waters as
indicated by their position of capture in gill nets,
consumption of limnetic invertebrates, and infrequent
occurrence (8%) of incidental vegetation and detritus
in their stomachs.

Salamanders specialize in eating Gammarus on the
bottom, and their niche, in terms of food and space, is
very narrow. That of trout is much broader and over-
laps the niche of salamanders. However, trout are
opportunistic, nonselective feeders, and although they
may prefer Gammarus when available, they are capa-
ble of exploiting alternate foods from several places
(Johannes and Larkin 1961). In Lake 200, for exam-
ple, the abundant Brook Stickleback (Culaea incon-
stans) is the major food of trout (Tavarutmaneegul
1978). Nevertheless, there is potential for competition
for food between salamanders and trout and this

TABLE 4—Mean dimensions (+99% confidence limits) in millimetres of trout and salamanders from Lake 675. Numbers

examined in parentheses

Salamanders

Month Dimension Trout Larvae Adults
June Body length! 112.9424 .33 41.1+22.46 109.4+2.48
Mouth length 9.1+ 0.65 (85) 7.0+ 0.84 (23) 15.00.43 (70)
Mouth width 7.8 0.65 7.9+ 0.95 18.1+0.61
July Body length 144.8+68.67 80.5+16.31
Mouth length 11.04 0.69 (13) 11.0 0.51 (20)
Mouth width 8.7+ 1.20 4a 7ee ESS

'Body length = fork length for trout and snout-vent length for salamanders.

132 THE CANADIAN FIELD-NATURALIST

could cause trout to shift in the food and space dimen-
sions of their niche. The impact of sucha niche shift on
the production of trout can only be assessed by
manipulating numbers of both species within re-
stricted areas and recording their growth. Differences
in mouth size and partial segregation along the space,
food, and time (season) dimensions of the niche sug-
gest that such an impact on trout may not be great.

Acknowledgments

We thank C. H. Johnston for her assistance in the
field; J. Mathias and M. Pabst for kindly providing
space, facilities, and equipment at the Erickson aqua-
culture study area along with many useful suggestions
during the field study; T. Galloway, I. Lubinsky, and
A. Salki for verifying the identification of stomach
contents; J. Mathias and A. Maclem for providing
their unpublished data; D. I. MacKenzie and D.
Marmorek for their helpful suggestions for data anal-
ysis; J. Mathias, B. Ayles, W. E. Neill, and A. Copsey
for providing constructive comments on the manu-
script. The study was funded by a Fisheries and
Marine Service grant from the Department of Fisher-
ies and Oceans.

Literature Cited

Dodson, S.I., and V. E. Dodson. 1971. The diet of Amby-
stoma tigrinun larvae from western Colorado. Copeia
1971: 614-624.

Vol. 95

Efford, I. E., and K. Tsumura. 1973. A comparison of the
food of salamanders and fish in Marion Lake, British
Columbia. Transactions of the American Fisheries
Society 102: 33-47.

Hartman, G. F. 1958. Mouth size and food size in young
Rainbow Trout, Sa/mo gairdneri. Copeia 1958: 233-234.

Horn, H.S. 1966. Measurement of “overlap” in compara-
tive ecological studies. American Naturalist 100: 419-424.

Johannes, R. E., and P. A. Larkin. 1961. Competition for
food between Redside Shiners ( Richardsonius balteatus)
and Rainbow Trout (Sa/mo gairdneri) in two British
Columbian lakes. Journal of the Fisheries Research Board
of Canada 18: 203-220.

Lawler, G. H., L. A. Sunde, and J. Whitaker. 1974. Trout
production in prairie ponds. Journal of the Fisheries
Research Board of Canada 31: 929-936.

Myers, G. L. 1973. Prairie pothole ecology and the feasibil-
ity of growing Rainbow Trout (Sa/mo gairdneri Richard-
son) in prairie potholes. M.Sc. thesis, North Dakota State
University, Fargo, North Dakota. 91 pp.

Sunde, L. A., and J. Barica. 1975. Geography and lake
morphometry of the aquaculture study area in the Erick-
son-Elphinstone district of southwestern Manitoba.
Fisheries and Marine Service Research and Development
Technical Report 510. 35 pp.

Tavarutmaneegul, P. 1978. Production of Rainbow Trout
in small eutrophic lakes subject to periodic anoxia. M.Sc.
thesis, University of Manitoba, Winnipeg, Manitoba. 170

Pp.

Received | May 1980
Accepted 27 August 1980

Status and Breeding of Mountain Plovers (Charadrius montanus)

in Canada

CLIFFORD A. WALLIS and CLEVE R. WERSHLER

615 Deer Croft Way, S.E., Calgary, Alberta T2J SY4

Wallis, Clifford A., and Cleve R. Wershler. 1981. Status and breeding of Mountain Plovers (Charadrius montanus) in

Canada. Canadian Field-Naturalist 95(2): 133-136.

The first Canadian nesting records of Mountain Plovers (Charadrius montanus) are documented, with details of habitat, nest
construction, and behavior. All known Canadian records are listed, and recommendations are given for further research and

management with respect to this species.

Key Words: Mountain Plover, Charadrius montanus, ecology, nesting, behavior, management, distribution, Canada.

Until 1979, there had been few reports of Mountain
Plovers (Charadrius montanus) in Canada, although
Coues (1878) indicated that Mountain Plovers were
relatively common along the Canada — United States
boundary adjoining southeastern Alberta and
southwestern Saskatchewan. Mountain Plovers are
inconspicuous and their remote habitat has been sel-
dom visited by naturalists. In this paper we summarize
existing records, including the first recorded nestings
in Canada. On the basis of our observations of their
ecology, we suggest that special land management
may be required if Mountain Plovers are to continue
breeding in Canada.

Summary of Canadian Observations

While accompanying the United States Northern
Boundary Commission in 1874, Elliott Coues (1878)
noted Mountain Plovers:

“in the Milk River Country, along the parallel of
49° [N], where it was breeding in considerable
numbers... the northernmost points at which the
species has thus far been observed .. . it was first
seen July | (where the Milk River joins the Mis-
sourl), and it was traced thence across the country
nearly to the Sweetgrass Hills, beyond which it
was lost. Its centre of abundance in this region
was the vicinity of the Frenchman’s River, where
many specimens, both adult and young, together
with a set of three eggs, were secured during the
first and second weeks of July.”

The exact location of many of those specimens is
unclear, but all were collected within 50 km of the
Canada - United States border and some along the
boundary itself. The only precise location cited is the
crossing of the Milk River (Figure 1), an area very
close to the border, where on 23 July 1874 Coues
collected one adult.

Soper (1941) saw one bird 5 June 1939 near
Bracken, Saskatchewan. He also found four Moun-
tain Plovers near Wildhorse, Alberta, on 22 June

133

1941. He collected two males there and suspected that
they were breeding. That area is slightly rolling with
short and mid-grasses as well as scattered sagebrush
(Artemisia cana) and sparsely vegetated alkaline
“blow-outs.”

There were no further reports until 12 June 1966,
when one of us (CRW) observed two at Elkwater in
the Cypress Hills, Alberta. They were not thought to
be breeding in this unsuitable habitat. Salt and Salt
(1976) noted a report of two Mountain Plovers on 5
July 1971, a few kilometres north of the international
boundary about 30 km west of Wildhorse, Alberta.
On 22 September 1977, eight Mountain Plovers were
seen near Val Marie, Saskatchewan (Peart and
Woods 1980). On 30 September 1977, W. Smith and
C. Wallis located five Mountain Plovers northeast of
Onefour, Alberta. Three of the birds had some buffy
coloration on the breast while the other two had clear
white breasts, possibly indicating a family group. That
site was vegetated with short grasses and had alkaline
“blow-outs.” It was ona slightly rolling plateau along
the edge of an eroding slope, about 100 m above the
plains to the south where Soper’s Alberta record and
our later nesting records were obtained.

Nesting in Canada was first documented 25 May
1979 (Wallis and Loewen 1980). Later, additional
nesting Mountain Plovers were found in the vicinity
(Table 1). Anadult with three eggs was photographed
on 5 June 1979 (Figure 2). No birds were observed
after 14 July although surveys were made 15 July, 29
July, and 14 August. All three sites are in a remote
area between Onefour and Wildhorse, Alberta. Site A
lies 100 m west of site B, while site C lies 300 m north
of site B. The exact location is not described because
we do not want to jeopardize the nesting success of the
birds by attracting casual observers.

Habitat and Nest Descriptions
The general habitat around the nests at sites A and
B was heavily grazed winter pasture witha recent burn

134

ALBERTA

THE CANADIAN FIELD-NATURALIST

Vol. 95

SASKATCHEWAN

ELKWATER

probable

limit in Canada

~ _BRACKEN #

FiGuRE |. Location of place names mentioned in text. Broken line indicates probable breeding range of Mountain Plovers in

Canada.

characterized by short green growth. There was taller
denser vegetation in the adjacent unburned areas and
in more moist depressions within the burn. It is on
moderately rolling terrain near a drainage. Vegetation
cover at ground level was estimated on 2 X 2 m plots
centered on each nest, 14 June 1979. Nomenclature
for the vascular plants follows Moss (1959).

Site A is on a slight incline facing south. The
vegetation cover was as follows: Bouteloua gracilis
(Blue Grama) 30% and Carex filifolia (Thread-leaved
Sedge) 10% with minor amounts of Eurotia lanata
(Winter Fat), Carex eleocharis (Low Sedge), Cym-
poterus acaulis (Plains Cymopterus), Stipa comata
(Spear Grass), Plantago purshii (Pursh’s Plantain),

TABLE |—1979 observations of Mountain Plovers near Onefour, Alberta
Number of
Date Site adults eggs young’ Observers”
25 May A ] 3 = VL, CB,NW, WN, CW, NK, MF
B | — —
4-5 June A l 3 = CRW
B 3 —
Between A and B 2 _ —
9-10 June A 1 3 a RW
B l 3 —
Near C 1 — In
14 June A ] = 3n CW, CRW
C 1,1 — 3n,3n
11 July A ] = 2u CW, CRW
Cc D — 3u
C (from B) =o =
14 July Cc 3 = = WS

“n, newly hatched young; u, unfledged young.

°CB, C. Bradley; MF, M. Fairbarns; NK, N. Kondla; VL, V. Loewen; WN, W. Nordstrom; WS, W. Smith; CW, C. Wallis;

RW, R. Wershler; CRW, C. Wershler; NW, N. Woerns.

198]

WALLIS AND WERSHLER: MOUNTAIN PLOVER STATUS AND BREEDING

185

FiGureE 2. Mountain Plover at nest, site B, 5 June 1979. Note the cow manure pile on the left.

and the lichens Parmelia chlorochroa, Caloplaca sp.,
and Candelariella sp., with 55% bare sandy loam soil.
Other plants nearby included: A//ium textile (Prairie
Onion), Atriplex nuttallii (Salt Sage), Astragalus pec-
tinatus (Narrow-leaved Milk Vetch), and Poa secunda
(Sandberg’s Bluegrass).

Site B is on a gentle northwest slope. It was charac-
terized by: Bouteloua gracilis 25%, Carex filifolia
15%, and Poa secunda 5% with minor amounts of
Carex eleocharis, Cymopterus acaulis, Eurotia
lanata, Allium textile, and the lichens Physconia mus-
cigena, Parmelia chlorochroa, Fulgensia sp., Clado-
nia sp., Caloplaca sp.,and Candelariella sp., with 45%
bare sandy loam soil.

Both nests were located within the burned area and
less than 10 m from its edge. At site C there were more
weedy species and a higher proportion of mid-grasses
(Stipa comata, Koeleria cristata (June Grass), Poa
secunda). Detailed vegetation analysis was not under-
taken here as the exact location of the nests could not
be determined.

The nest at site A was located beside two large heaps
of dried cow manure. The shallow nest depression was
lined mostly with pieces of dry cow manure and very
small twigs, including broken stems of Eurotia lanata.
There were also Cymopterus acaulis seeds, Selaginella
densa (Little Club-moss) cuttings, and pieces of the
lichens Physconia muscigena, Cladonia sp., and Ful-
gensia sp. Another lined nest depression, identical in

material, shape, and size, was located a few metres
east of the site A nest. It was assumed to bea nest from
a previous year; although Mountain Plovers construct
several scrapes before nesting, only the actual nest is
lined (Graul 1975).

The nest at site B was located next to a group of dry
cow chips and a coyote scat. It was lined mostly with
pieces of dry cow manure and some old lagomorph
(probably Lepus townsendii) droppings.

Vocalizations

Several vocalizations similar to those described by
Graul (1974) were noted. One, a low guttural sound,
was commonly given by the adults when near young.
It was a ventriloqual descending call that may be
Graul’s brood call. However, unlike Graul’s verbal
description, this call was sufficiently loud to attract us
to adults from a considerable distance.

Discussion

Four Mountain Plovers nested at the study sites in
1979. It is possible that these represented two pairs
because the male and female frequently incubate
separate clutches of eggs simultaneously (Graul 197.,
At least nine young were hatched from sites A and C.
The fate of the eggs at site B was uncertain.

All observers have noted that the best habitat for
Mountain Plovers is extensive unbroken grassland
with very short native grass (<8 cm) at the start of the

Vol. 95

nesting season. This is usually Blue Grama (Bouteloua
gracilis) or Buffalo Grass (Buchloe dactyloides), but
some areas have a mixture of short grasses, sedges,
mid-grasses, and some shrubs. Most good habitat is
subject to heavy summer grazing. In the Colorado
studies 90% of nests were on slopes less than 2%,
usually on large flat areas (Graul 1976).

The Canadian nesting habitat appears to be gener-
ally similar to that in the United States, although
Buffalo Grass does not occur within the presumed
breeding range of Mountain Plovers in Canada. Nest
materials used in Canada also were similar to those in
Colorado (Graul 1975). Extrapolating from Graul’s
(1975) and our data, the Alberta birds arrive about the
2nd or 3rd week of April and begin laying in the 2nd or
3rd week of May. The old lined nest depression near
site A indicates that the birds may return to a site if
habitat conditions remain suitable, as found also by
Graul (1973).

We and a few others have travelled extensively in
extreme southeastern Alberta on foot and by car,
looking specifically for Mountain Plovers and other
unusual plants and animals in the region. Unless one is
close to a pair of Mountain Plovers, their presence is
seldom obvious, and other nesting populations may
have been overlooked. There is also a large and little
studied area from the Alberta border to Val Marie,
Saskatchewan. From the historical records and recent
observations, the most likely breeding areas are within
15-20 km of the Canada- United States border
between the points where the Frenchman and Milk
rivers cross the international boundary, including the
proposed Grassland National Park area.

The Mountain Plover is now absent or rare in many
areas where previously it was numerous. In Wyoming,
there are only isolated populations, and Skaar (1975)
documents only five recent breeding locations in
Montana. The stronghold of the species is in north-
eastern Colorado where there is still prime habitat on
the Pawnee National Grassland (Graul 1976).

The recent nesting records in Canada probably do
not represent freak occurrences. Although the species
is at the extreme edge of its range, the Mountain
Plover may have always been a characteristic species
of this region. It should be added to the list of Cana-
dian endangered species and afforded protective
management because of apparent declines in its total
world population. The appropriate management of
extensive blocks of native grassland, such as those at
the Milk River Canyon and at Val Marie-Killdeer,
may be crucial to the long-term survival of Canadian
populations of the Mountain Plover and other grass-
land species.

Mountain Plover habitat can probably be des-
troyed by inadequate grazing. To simulate the range
of conditions that must have existed when bison

THE CANADIAN FIELD-NATURALIST 136

grazed the plains, a balance is needed between heavily
grazed, lightly grazed, and ungrazed areas. There may
be danger in current trends to uniform grasslands
caused by long-term overgrazing,.reduction in pasture
size, even distribution of cattle use by fencing, water-
ing, and salting, or by totally protecting large areas
from all grazing. The key to the preservation of the
Mountain Plover probably lies in maintaining sub-
stantial patches of short grass in extensive native
mixed grassland areas. Locally prescribed heavy
summer or late winter grazing or the use of fire may be
beneficial, judging from the use of winter pasture and
recent burns in the Canadian sites. More research is
needed to determine: (1) the extent of the Canadian
range and approximate population size; (2) whether
habitat is the major limiting factor and whether pre-
scribed burning or various grazing prescriptions could
improve habitat; (3) whether climatic changes alter
Mountain Plover habitat; (4) more details of their
breeding biology in Canada; and (5) wintering areas
and how conditions there affect Canadian pop-
ulations.

Acknowledgments

We express our appreciation to W. Smith and R.
Wershler for their field notes and their dedication to
field surveys; N. Kondla for review of the manuscript;
and B. Peart for details of his recent Saskatchewan
observations.

Literature Cited
Coues, E. 1878. Field notes on birds observed in Dakota and
Montana along the forty-ninth parallel during the seasons
_ of 1873 and 1874. Article XXV. /n Bulletin of the United
States Geological and Geographical Survey Volume IV.
Government Printing Office, Washington. pp. 545-661.
Graul, W. 1973. Adaptive aspects of the Mountain Plover
social system. Living Bird 12: 69-94.
Graul, W. 1974. Vocalizations of the Mountain Plover. Wil-
son Bulletin 86: 221-229.
Graul, W. 1975. Breeding biology of the Mountain Plover.
Wilson Bulletin 87: 6-31.
Graul, W. 1976. Breeding status of the Mountain Plover.
Condor 78: 265-267.
Moss, E. H. 1959. Flora of Alberta. University of Toronto
Press, Toronto.
Peart, B.,and J. G. Woods. 1980. Mountain Plover observa-
tion near Val Marie, Saskatchewan. Blue Jay 38: 41.
Salt, W.R., and J. R. Salt. 1976. The birds of Alberta.
Hurtig, Edmonton.
Skaar, P. D. 1975. Montana bird distribution. P.D. Skaar,
. Bozeman. Privately printed.
Soper, J. D. 1941. The Mountain Plover in western Canada.
Canadian Field-Naturalist 55: 137.
Wallis, C., and V. Loewen. 1980. First nesting record of the
Mountain Plover in Canada. Alberta Naturalist 10: 63-64.

Received 23 May 1980.
Accepted 20 September 1980.

Estimates of the Standing Stocks of Fishes in Four Small

Precambrian Shield Lakes!

J. M. FRASER

Research Section, Fisheries Branch, Ontario Ministry of Natural Resources, Maple, Ontario LOJ 1E0

Fraser, J. M. 1981. Estimates of the standing stocks of fishes in four small Precambrian Shield lakes. Canadian Field-

Naturalist 95(2): 137-143.

Estimates of the standing stocks of fishes in four small Precambrian Shield lakes ranged from 35.2 to 50.8 kg/ha at the time of
reclamation with a fish toxicant. In three lakes with similar species composition, White Suckers (Catostomus commersoni)
comprised 50% (49-62), Golden Shiners (Notemigonus crysoleucas) 18% (13-25), and planted Brook Trout (Sa/velinus
fontinalis) 14% (11-17) of total standing stocks. In the fourth lake, Pumpkinseeds (Lepomis gibbosus), Brown Bullheads
(Ictalurus nebulosus), and planted Brook Trout contributed 58, 19, and 13%, respectively, of the total standing stock of 50.7
kg/ha. The relatively rigorous proportionality among species, evident for both communities, and the similarity of their
standing stocks of fishes suggest that the carrying capacities of the lakes had been reached.

Key Words: biomass, Precambrian Shield, reclamation, standing stocks, toxicant.

Although there is considerable information con-
cerning the yields of fishes from lakes on the Precam-
brian Shield (Ryder 1965; Fraser 1972; Adams and

Olver 1977; Martin and Fry 1972), much lessisknown -

about the size and composition of the standing stocks
of fishes in these lakes. This information is important
to fish managers because it provides a measure of the
productivity and carrying capacity of such waters.
When a lake reclamation project was conducted in
conjunction with a research program, it provided an
Opportunity to estimate the population of fishes in
four small lakes on the Precambrian Shield. Three
lakes, Sawlog, Pine, and Lonesome (45.35°N,
75.05° W), form the headwaters of a small watershed
that empties into the Madawaska River, a tributary of
the Ottawa River. The lakes were not fished until a
hydro power transmission line provided access in
1950. Initial angling attempts were unsuccessful and
according to two highly experienced fishing guides the
lakes were devoid of fish, even minnows, before 1950.
Preliminary gillnet surveys in 1950 caught nothing,
corroborating these observations, at least for the
larger fish. A complete survey of the lakes was not,
however, conducted at that time. Annual plantings of
hatchery-reared Brook Trout (Sa/velinus fontinalis)
began in 1950 and during the following 5-7 years
provided reportedly excellent angling. The quality of
the angling declined thereafter and in 1961 when the
lakes were selected for special study, small Brook
Trout were being caught on a put-take basis at the
rate of 0.2 fish per hour. Surveys conducted in 1961
showed that all lakes supported the White Sucker

‘Contribution No. 79-17 of the Fisheries Branch, Ontario
Ministry of Natural Resources, Maple, Ontario

(Catostomus commersoni), five cyprinid species, and
the Brook Stickleback (Culaea inconstans). An excep-
tion was Sawlog Lake, the headwater, in which
suckers did not appear until 1962. The above three
lakes were treated with a fish toxicant in July 1968.
The fourth lake, Redpine, is the headwater of an
adjacent watershed, and it was treated in July 1969. In
this paper the standing stocks of fishes at the time of
reclamation are described.

The study lakes range in area from 3.6 to 16.4 ha
and in maximum depth from 6.1 to 11.9 m (Table 1).
Their waters are slightly acidic, poorly buffered, and
relatively infertile. Total dissolved solids ranged from
31 to 40 mg/L. Sawlog, Pine, and Lonesome lakes
were thermally stratified at the time of poisoning, with
thermoclines located at 3-6 m in depth. Dissolved
oxygen content of the bottom waters was 0-2 mg/L.
In contrast, only a weak temperature gradient existed

TABLE !—Physical and chemical characteristics of study
lakes. Water sampled in 1961-1967

Sawlog Pine Lonesome Redpine

Area (ha) 6.3 16.4 3.6 8.3
Maximum depth

(m) 8.5 11.9 6.4 6.1
Mean depth (m) 4.3 46 2.8 2.4
Secchi disc

visibility (m) 4.3 3.4 Dell 4.1
Total dissolved

solids (mg/L) 31 40 IS 39
Total alkalinity

(mg/L) 9 11 SI 13
pH 6.9 6.8 6.5 6.9

“Data from 23 May 1967 — R. A. Ryder data files.

137

138 THE CANADIAN FIELD-NATURALIST

in Redpine Lake and the bottom waters were moder-
ately warm (14°C) and nearly saturated with oxygen.
Fish species in the three connected lakes were identical
and are listed with their scientific names in Table 2.
Redpine Lake differed from the others in species
composition (Table 2).

Methods and Materials

Several weeks before reclamation, fish were cap-
tured, marked, and released in the study lakes. The
trap nets used were 1.2 m deep and were constructed
of nylon 25-mm mesh (bar measure) in the crib and
32-mm mesh in the leader. The 1.8-m-long Winder-
mere traps were constructed of 6.3-mm mesh hard-
ware cloth attached to three 0.8-m steel hoops. Cap-
tured fish were anesthetized with tricaine metha-
nesulfonate (MS222), finclipped, and released. White
Suckers, Pumpkinseeds, and Brown Bullheads were
measured (fork length) to the nearest 2.5 mm (0.1 in.)
before being released.

Marked yearling Brook Trout had been planted in
all lakes 1-4 d before poisoning to serve as an index to
the recovery of earlier yearling plantings. These
“index” trout were not included in the standing stock
estimates.

The fish toxicant containing 2.5% rotenone, 2.5%
synergist, emulsifier, and a water-miscible solvent was

Vol. 95

distributed chiefly by slow discharge into the prop
wash of two outboard motor boats which slowly criss-
crossed each lake. In the two deeper lakes, about 25%
of the toxicant was pumped through a weighted hose
reaching 6 m beneath the surface. Several small
streams and bog areas were sprayed by hand pump.
Sawlog, Pine, and Lonesome lakes were treated witha
1.0 mg/L concentration on 3, 9, and 11 July 1968,
respectively. Redpine Lake received a 2.0 mg/L con-
centration on 7 July 1969. Although the lakes were
treated separately, the method of treatment and the
subsequent collection of fish was similar for all the
lakes.

The application of the toxicant was completed by
noon and the collection of fish, which began within 30
min of the beginning of treatment, continued
throughout that day and the following day. Five or six
two-man crews working from small boats or canoes
dipnetted dead and dying fish from the surface of the
lake and along the shore. The collection was discon-
tinued after 2 d in Sawlog, Pine, and Lonesome lakes
because few fish surfaced thereafter. Fish continued to
surface in Redpine Lake and were collected over a 5-d
period. Small fish that had drifted to shore were col-
lected with difficulty in areas of aquatic vegetation
and overhanging shoreline shrubs.

The fish collected were sorted by species, examined

TABLE 2—Numbers and weights of various species of fish collected from four lakes following treatment rotenone

Sawlog

Species

Brook Trout®

(Salvelinus fontinalis) 215 12.9 301

White Suckers

(Catostomus commersoni) 474 35.8
Golden Shiners

(Notemigonus crysoleucas) 1239 15.9
Creek Chub

(Semotilus atromaculatus) 167 2.9 136

Pearl Dace

(Semotilus margarita) 240 De 111

Redbelly Dace

(Chrosomus €0s) P P

Fathead Minnows
(Pimephales promelas) 251 0.9
Brown Bullheads
(Ictalurus nebulosus)
Pumpkinseeds
(Lepomis gibbosus)
Brook Sticklebacks

(Culaea inconstans) 80 0.1
Minnows and sticklebacks 684 1.9
All species 3350 72.6
“Planted.

et a present but not collected.

4092

6480

Pine Lonesome Redpine

Number Weight (kg) Number Weight (kg) Number Weight (kg) Number Weight (kg)

16.7 90 4.9 SN 37.6
40.6 426 Zed
17.1 636 2.6
52 10 0.3 Pp?
1.0 33 0.4 P
360 0.9 P
a= 21 0.1 P
1316 28.9
5641 20.9
407 0.3
5.5 0.6
86.1 1983 27.8 7474 87.4

1981 FRASER: FISH STANDING STOCKS, PRECAMBRIAN SHIELD LAKES 139

individually for finclips, counted, and weighed in the
aggregate. Fork lengths of Brook Trout, White
Sucker, Brown Bullhead, and Pumpkinseed were
measured individually to the nearest 2.5 mm. Small
fish, which formed the residue left after collections
had been sorted, deteriorated rapidly and were
weighed in the aggregate and described “minnows and
sticklebacks.”

SCUBA divers swam six to eight transects along a
measured, weighted line laid on the lake bottom in the
afternoon of the day following rotenone treatment.
The transect lines, stretching from shore to shore,
were perpendicular to the long axis of the lake, and
divers, swimming in pairs, collected fish from a 3.7-m-
wide strip of bottom. In this manner, 5.8, 2.6, 9.0, and
6.0% of the bottom of Sawlog, Pine, Lonesome, and
Redpine lakes, respectively, were searched for dead
fish. Bright sunshine aided the divers, and good vis-
ibility to a depth of 9 m was reported. Visibility was
poor at greater depths and consequently the deeper
sector (2.3 ha) of Pine Lake could not be searched. An
estimate of the numbers of fish on the entire lake

bottom was made by expanding the transect data by
direct proportionality. No fish were caught in gill nets
(38-, 51-, 64-, 76-mm mesh-stretched measure) set in
the treated lakes 5-7 wk after the treatment.

Ratios of marked to unmarked fish in the total
collection were used to calculate Petersen-type esti-
mates of the Brook Trout, White Sucker, and Golden
Shiner populations. Confidence limits (95%) were cal-
culated using formula 1.13 of DeLury (1951). Esti-
mates of standing stocks were calculated by multiply-
ing the mean weights of each species by their
respective population estimates.

Results
Sawlog Lake

A total of 3350 fish weighing 72.6 kg was collected
from Sawlog Lake following rotenone treatment
(Table 2). The collection was made up of Brook Trout,
White Suckers, Brook Sticklebacks, and five cyprinid
species.

Estimates of the numbers and weights of the three
chief species were made and are presented in Table 3.

TABLE 3—Estimates of the standing stocks of fishes in four study lakes at the time of rotenone treatment

Number marked

Number collected

PopulationConfidence Mean _ SHERI SHO

and released Unmarked Marked estimate levels(95%) weight(g) kg/ha %
Sawlog Lake
Brook Trout 150 178 37 872 671-1245 60.0 8.3 17
White Sucker (© 10.0 cm) 207 432 45 2194 1708-3066 75.5 26.3 53
Golden Shiner (> 6.4 cm) 817 905 334 3030 2784-3314 12.8 6.2 13
Minnows and stickleback 920 8.7 8.5 17
Total standing stock 49.3
Pine Lake
Brook Trout 250 240 57 1302 1052-1711 55.5 4.4 13
White Sucker (© 10.0 cm) 124 1493 17 11014 7430-21277 25.5 17.1 49
Golden Shiner 3388 3227 350 34625 31437-38534 4.2 8.9 25
Minnows and stickleback 4.8 14
Total standing stock 35.2
Lonesome Lake
Brook Trout 99 66 24 368 275-571 54.5 5.6 11
White Sucker 84 413 13 2753 1780-6065 41.5 Sod 62
Golden Shiner 845 358 46 7421 5813-10283 4.2 8.7 17
Minnows and stickleback 4.8 10
Total standing stock 50.8
Redpine Lake
Brook Trout 200 379 138 749 699-808 72.8 6.6 13
Brown Bullhead
(8.0 cm) 392 1059 140 3357 2898-3990 D35) 9.5 19
Pumpkinseed 1153 5543 98 66368 53307-82960 3.7 29.6 58
Minnows and darters 5.0 10
Total standing stock 50.7

140

The most numerous fish was the Golden Shiner, but
White Suckers contributed the most weight (53%) to
the total standing stock of 310 kg or 49.3 kg/ha. The
Golden Shiner population consisted of two distinct
size groups, large (>6.4 cm) and small (<3.8 cm).
Only the larger Golden Shiners could be captured,
marked, and released, and this was the size group
most commonly collected after poisoning (Figure 1).
Mean weights of marked and unmarked Golden Shin-
ers in the collection were 11.8 and 13.2 g, respectively,
indicating that the population estimate of 3030 applies
to the larger Golden Shiners which made up 13% of
the lake’s total standing stock.

An estimated 872 Brook Trout were extant in Saw-
log Lake at the time of poisoning. Ratios of the differ-
ent finclips indicate that 66% had been planted in
May, 6 wk before poisoning and the remainder in the
preceding autumn.

It was not logistically practical to estimate the pop-
ulations of small Golden Shiners, four other cyprinid
species, and the Brook Stickleback by mark and rec-
overy methods. However, an approximate estimate
was made by assuming that the collection of small fish
represented about 15% of the standing stock of small
fish. Because the mean recovery after poisoning of
Brook Trout, White Suckers, and Golden Shiners was
27, 18, and 19%, respectively (Table 4), the assumed
15% recovery of small fish conservatively allows for
the reduced effectiveness in collecting them.

SAWLOG

BROOK TROUT
— (215)

WHITE SUCKER
— (468)
—-——(207)

PERCENTAGE OF TOTAL

GOLDEN SHINER
(77)

5 10 IS 20 25 5 10

FORK

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 4—The percentage recovery of marked fish following
treatment with rotenone. Duration of pickup was 2 d

Brook White Golden Brown Pumpkin
Lake Trout Suckers Shiners Bullheads seeds
Sawlog 25 22 41
Pine 23 15 10
Lonesome 24 17 5)
Redpine BOR 19° 8
Mean 27 18 19

“These percentages were increased to 69 for Brook Trout and
36 for Brown Bullheads when the duration of pickup in this
lake was extended to 5 d.

Pine Lake

The total standing stock of Pine Lake was estimated
to be 35.2 kg/ha made up of 49% White Suckers, 25%
Golden Shiners, 13% Brook Trout, and 14% cyprinids
and Brook Sticklebacks (Table 3), very similar to the
proportionate representation in Sawlog Lake. Most
of the Brook Trout were recently planted yearlings.

The estimate of 11 014 White Suckers is an approx-
imation at best because only 124 suckers were marked
and released into the population. Apparently, the
traps caught the few large suckers in the lake but were
ineffective in catching the small ones which comprised
90% of the population (Figure 1). The scarcity of large
suckers is likely the result of annual gillnet sampling in

PINE LONESOME

15 20

LENGTH

BROOK TROUT
— (299)

WHITE SUCKER
—— (1805)
---— (124)

GOLDEN SHINER
(11)

25 30
(cm)

BROOK TROUT
— (90)

WHITE SUCKER
— (426)

GOLDEN SHINER
— (404)

20 25

FIGURE I. Length distribution (solid lines) of Brook Trout, White Suckers, and Golden Shiners collected following the
reclamation of Sawlog, Pine, and Lonesome lakes. Broken lines represent fish marked and released before poisoning.

Numbers of fish measured are in parentheses.

1981

1963-1967. The estimated population of 34 625
Golden Shiners represented a standing stock of 8.9
kg/ha (Table 3). The collection of minnows (other
than Golden Shiners) and sticklebacks weighed 11.7
kg and assuming they represent 15% of the small fish
population, their standing stock was 4.8 kg/ha.

Lonesome Lake

Golden Shiners and White Suckers were also the
predominant species in the collection of fish from
Lonesome Lake. Golden Shiners were most numerous
but constituted only 17% of standing stock owing to
their small size (mean, 4.2 g). The White Sucker popu-
lation was bimodal in length distribution with peaks
occurring at 8.9 and 16.5 cm (Figure 1). It contributed
62% of the lake’s total standing stock of 5.8 kg/ha
(Table 3). The remainder consisted of recently planted
yearling Brook Trout, small minnows, and stickle-
backs.

Redpine Lake

Following treatment of Redpine Lake on 7 July
1969, Pumpkinseeds, Brown Bullheads, and Brook
Trout were collected daily to 11 July 1969 (Table 2).
Five cyprinid species and the lowa Darter were noted
floating on the surface, but were not collected. The
standing stocks of the three chief species were esti-
mated to total 45.7 kg/ha and allowing 5.0 kg/ha for
cyprinids and darters, the lakes standing stock of
fishes amounted to 50.7 kg/ha (Table 3).

Pumpkinseeds, which constituted 58% of the stand-
ing stock, ranged from 2 to 11 cm with the majority
falling in the 4- to 5S-cm length-class (Figure 2) and
weighing 3.7 g on the average. Pumpkinseeds, marked
and released from the traps, were larger than those
collected after poisoning (Figure 2), but this probably
did not affect the population estimates because all
sizes floated on the surface after dying and were col-
lected with equal efficiency. Brown Bullheads (>8.0
cm) and yearling Brook Trout made up much of the
remainder of the lake’s standing stock (Table 3).

Population Estimates Based on SCUBA Transects

On the day following treatment on each lake,
SCUBA divers swam six to eight transects along the
lake bottom searching for dead fish. The numbers of
fish picked up by the divers were extrapolated to the
total substrate area to provide estimates of the total
number of fish on the lake bottom. This estimate was
combined with the number of fish collected on the
surface to provide an estimate of the population
before poisoning of the lake. Estimates were calcu-
lated in this manner for Brown Bullheads in one lake,
White Suckers in three lakes, and Brook Trout in all
four lakes (Table 5). Petersen estimates of the same
populations derived from mark and recovery data are
entered in Table 5 for comparison.

FRASER: FISH STANDING STOCKS, PRECAMBRIAN SHIELD LAKES

141

BROOK TROUT
me (C8)

4

=

° BULLHEAD
iz —— (1316)
m 20 =——— (392)
[o)

lJ

oO

a 10

Ss

za

WW

oO

a

W

a PUMPKINSEED

el (OI)

30 el (III)

20

T
5 20 25 30
FORK LENGTH (cm)

FIGURE 2. Length distribution (solid lines) of Brook Trout,
Brown Bullheads, and Pumpkinseeds collected following the
reclamation of Redpine Lake. Broken lines represent fish
marked and released before poisoning. Numbers of fish mea-
sured are in parentheses.

In all instances, the estimates based on underwater
transects were significantly lower than the Petersen
estimates. A probable explanation is that some dying
fish entered the soft organic substrate and died there,
out of sight. Estimates based only on fish seen and
collected from the lake bottom would, therefore, be
biased on the low side.

Divers observed fish with only tails protruding
from the lake bottom and suggested that there were
probably others completely hidden in the organic
ooze. Rupp and DeRoche (1965) reported that dying
bullheads apparently had burrowed into the bottom
silt during rotenone treatment of Shagg Lake, Maine.
Divers in our study reported seeing Brook Trout and
suckers, as well as bullheads, with only their tails
protruding from the lake bottom.

Discussion

The procedure of marking and releasing fish before
lake poisoning, as recommended by Carlander and
Lewis (1948), provides for an estimate of percentage
recovery in the collection of fish. Appleget (1949) used

142 THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 5—Population estimates of Brook Trout, White Suckers, and Brown Bullheads based on extrapolated bottom

collections added to surface collections

Sawlog Pine Lonesome Redpine

Number of transects 6 8 6 6
Total length of transects (m) 1003 1155 885 1372
Area of lake bottom searched (m2) 3671 4227 3239 $017
Percent of lake searched 5.8 2.8 9.0 6.0
Brook Trout

Number estimated on bottom 534 577 100 356

Estimated population

— bottom estimate + surface collection 718 863 181 599

— mark and recovery’ 872 1302 368 749
White Sucker

Number estimated on bottom 672 2692 522

Estimated population

— bottom estimate + surface collection 1107 4428 901

— mark and recovery 2194 11014 2753
Brown Bullheads

Number estimated on bottom 1683

Estimated population

— bottom estimate + surface collection on

days 1, 2 2279
— mark and recovery 3357

“Mark and recovery estimates from Table 3.

this method and estimated the standing stock of fish in
a 4.5-ha softwater lake in Minnesota to be about 88
kg/ha. The procedure was especially useful in the
present study because accurate enumeration of dead
fishes by direct count was not possible. Shoreline
counts were hampered by vegetation and because of
poor visibility, counts of fish on the lake bottom were
restricted to depths less than 9 m. In addition, some
distressed fish apparently entered the bottom ooze
and died there, out of sight.

Several of the population estimates are of question-
able accuracy mainly because of the difficulty in cap-
turing a sufficient number of fish for marking and
release. Hence, the confidence intervals for the esti-
mates of White Suckers in Pine and Lonesome lakes
are rather wide. For the same reason estimates of
cyprinids (other than Golden Shiners) and stickle-
backs could not be made, and it was necessary to use
the percentage recovery of other species to make con-
servative approximations of the standing stocks of
minnows. These difficulties could undoubtedly be
overcome in future projects by an intensive trapping
program using appropriate gear.

Estimates of the standing stocks of fishes in our
study lakes ranged from 35.2 to 50.8 kg/ha, somewhat
higher than the 14.6 kg/ha determined for Red Deer
Lake, 90 km distant (Chadwick 1976). According to
Carlander (1955), standing stocks of this order are
representative of trout lakes and indicate a low pro-

ductive capacity compared to warmwater lakes and
reservoirs. The poor supply of nutrients to Precam-
brian Shield lakes is reflected in the low values of total
alkalinity (5-13 mg/L) and total dissolved solids
(15-40 mg/ L) in these waters. Ryder (1965) has shown
that a low level of fish production can be expected
from lakes of similar fertility. The study lakes,
although physically and chemically suitable for Brook
Trout, lack adequate spawning areas and conse-
quently cannot support reproducing Brook Trout
populations.

The predominance of the White Sucker and rather
rigorous proportionality among the three major spe-
cies in Sawlog, Pine and Lonesome lakes (Table 3)
indicates a moderately stable community structure.
Adams and Olver (1977) have noted that percid spe-
cies tend to constitute 30% by weight of the fish yield
obtained from communities in which they occur, and
they proposed this relation to bean emergent property
of that type of community. Our study lakes may
represent another type of community and a commun-
ity conservatism (Kerr 1977) that places the White
Sucker in a predominant role, comprising 49-62% of
the total standing stock, followed by the Golden
Shiner comprising 13-25%. In Redpine Lake, interest-
ingly, although the species composition was different,
a corresponding proportionality among species
occurred and the total standing stock was approxi-
mately equivalent to that of the other three lakes.

1981

Native Pumpkinseed and Brown Bullheads were pre-
dominant contributing 58 and 19%, respectively, of
the lake’s total standing stock of 50.7 kg/ha.

The lakes considered in this study are relatively
infertile, of simple morphometry, and support small
standing stocks of fish. Because two different fish
communities produced standing stocks of similar size,
the carrying capacities of the lakes had apparently
been reached and intrusion by planted Brook Trout

would have been severely limited. This probably »

accounts for the poor survival of Brook Trout planted
in these and similar lakes on the Precambrian Shield
(Fraser 1972).

Acknowledgments

I am indebted to K. Buckingham, D. Cucin, J.
Murdock, B. Kukhta, and a host of student assistants
and volunteers for their assistance in the treatment of
the lakes and the collection, examination, and enu-
meration of the kill of fishes; to R. Ryder, J. Cassel-
man, and C. Olver for reviewing the manuscript.

Literature Cited.

Adams, G. F., and C. H. Olver. 1977. Yield properties and
structure of boreal percid communities in Ontario. Jour-
nal of the Fisheries Research Board of Canada 34:
1613-1625.

Appleget, J. C. 1949. Elimination of a rough fish population
from Surprise Lake, Itasca County, Minnesota, and an
estimate of the total population. Minnesota Department
of Conservation, Fisheries Research Investigation Report

FRASER: FISH STANDING STOCKS, PRECAMBRIAN SHIELD LAKES

143

89.5 pp.

Carlander, K. D. 1955. The standing crop of fish in lakes.
Journal of the Fisheries Research Board of Canada 12:
543-570.

Carlander, K. D. and W. M. Lewis. 1948. Some precautions
in estimating fish populations. Progressive Fish-Culturist
10(3): 135-137.

Chadwick, E. M. 1976. Ecological fish production ina small
Precambrian Shield lake. Environmental Biology of Fish
1: 13-60.

DeLury, D. B. 1951. On the planning of experiments for the
estimation of fish populations. Journal of the Fisheries
Research Board of Canada 8: 281-307.

Fraser, J. M. 1972. Recovery of planted brook trout, splake,
and rainbow trout from selected Ontario lakes. Journal of
the Fisheries Research Board of Canada 29: 129-142.

Kerr, S. R. 1977. Structure and transformation of fish pro-
duction systems. Journal of the Fisheries Research Board
of Canada 34: 1989-1993.

Martin, N. V.,and F. E. J. Fry. 1972. Lake Opeongo: effects
of exploitation and introductions of the salmonid com-
munity. Journal of the Fisheries Research Board of Can-
ada 29: 795-805.

Rupp, R.S., and S. E. DeRoche. 1965. Standing crops of
fishes in three small lakes compared with C'4 estimates of
primary productivity. Transactions of the American
Fisheries Society 94: 9-25.

Ryder, R. A. 1965. A method of estimating the potential fish
production of north-temperate lakes. Transactions of the
American Fisheries Society 94: 214-218.

Received 26 June 1980
Accepted 23 September 1980

A Test of the Peninsular Effect on Species Diversity

ROBERT J. TAYLOR! and LEE A. PFANNMULLER?

Department of Ecology & Behavioral Biology, University of Minnesota, Minneapolis, Minnesota 55455
'Present address: Department of Zoology, Clemson University, Clemson, South Carolina 29631.
2Present address: 909 West Sixth Street, Duluth, Minnesota 55806.

Taylor, Robert J., and Lee A. Pfannmuller. 1981. A test of the peninsular effect on species diversity. Canadian Field-

Naturalist. 95(2): 144-148.

The hypothesis tested was that species diversity declines along peninsulas because of the direct effects of peninsular geometry
upon rates of colonization and extinction of local populations. Communities of carabid beetles and small mammals (rodents
and insectivores) were sampled at three sites along Red Deer Point in Lake Winnipegosis, Manitoba; this was chosen as a
research site from a map of North America solely on the basis of its size and shape. From the samples taken from this 39-km
peninsula, we conclude that carabids and small mammals fail to exhibit an identifiable peninsular effect.

Key Words: peninsular effect, Manitoba, small mammals, carabid beetles, species diversity.

Simpson (1964) observed, while studying the geo-
graphic distributions of Recent mammals, that species
number decreases toward the tips of all major North
American peninsulas. This “peninsular effect” has
subsequently been discovered in other vertebrate
groups as well, e.g., birds (MacArthur and Wilson
1967; Cook 1969), and amphibians and reptiles (Kies-
ter 1971). Simpson suggested that this was not a for-
tuitous pattern, that the geometry of peninsulas pre-
disposed them to exhibit such a gradient in diversity.
The mechanism for this was an equilibrium between
local extinctions of species and the colonization of
empty habitats, a mechanism familiar to most ecolo-
gists by its application to islands. Peninsular faunas
living in an approximately linear chain of communi-
ties would be relatively more vulnerable to the loss of
species near peninsular tips where recolonization was
limited and extinction possibly more likely.

The logic and evidence behind this hypothesis were
explored in another paper by one of us (Taylor and
Regal 1978). The primary conclusion of that paper
was that Simpson’s reasoning was acceptable; a simu-
lation model of the processes of colonization and
extinction did produce appropriate equilibrium pat-
terns, although some rigid constraints were necessary
upon both the model’s structure and the values of its
parameters. Recently Gilpin (in press) has suggested
that the model may produce appropriate peninsular
patterns under less severe constraints.

Mathematical models yield only suggestions and
possibilities, of course. Although a good hypothesis
should certainly be internally consistent, logic does
not, by itself, constitute a test. Testing of the validity
of the peninsular effect is certainly necessary because
the second major conclusion of Taylor’s and Regal’s
paper was that the species diversity patterns of verte-
brates on all North American peninsulas, with one

exception, can be explained more parsimoniously as
responses to vegetational change. That exception is
Baja California, a puzzling location requiring more
attention.

Empirical approaches to the examination of bio-
geographic hypotheses require either direct manipula-
tion of biotas (e.g., Simberloff and Wilson 1969), or,
more traditionally, indirect analysis of natural exper-
iments. Inasmuch as direct manipulation is rarely
practical, biogeographic research has relied heavily
upon the indirect test of noting the presence or
absence of predicted patterns in previously uninvesti-
gated locations. In this paper we report the results of
sucha test of the peninsular effect. We chose an unex-
amined peninsula froma map of North America using
as criteria only that it be long, narrow, and small
enough that it could be sampled.

Study Site and Methods

Red Deer Point, 39 km long, extends into Lake
Winnipegosis in central Manitoba (Figure |). Because
Red Deer Point is substantially smaller than Florida,
Baja California, Yucatan, and the other major North
American peninsulas, we tried to compensate for the
effects of scaling down. To do this we looked at two
quite different animal groups, small rodents and
insectivores which are relatively uniform in dispersal
ability, and ground beetles (Coleoptera: Carabidae), a
more variable group with some very good and some
very poor dispersers. The reason for using an inverte-
brate group was the success enjoyed by Simberloff in
scaling down investigations of island biogeography by
looking at insects on Florida mangrove keys (e.g.,
Simberloff 1976).

The width of Red Deer point ranges from | to
5.8 km. It is quite flat, probably no greater than 8 m
above the lake at its highest point. The chief topogra-

144

1981

on |
0 10 20 30 40 50
km

Lake Winnipegosis

FIGURE 1. Location of the peninsula studied, Red Deer
Point in Lake Winnipegosis, Manitoba.

phic feature is a I- to 3-m-high levee along the lake-
shore. Long-term data on the lake’s water level are not
available; however, local residents, several of whom
have been in the area since the 1920s, claimed that the
water level at the time of the study was as high as it
ever had been.

The vegetation consists of a mosaic of sedge and
grass meadows interspersed with second-growth
forest. Trembling Aspen (Populus tremuloides) dom-
inates the forest, but one finds large numbers of Bur
Oak (Quercus macrocarpa), White Elm (U/mus amer-
icana), Red Ash (Fraxinus pennsylvanica), and a few
White Spruce (Picea glauca). The soil is thin, averag-
ing less than 10 cm. Meadows and woods occur in
about equal proportions in the southern half of the
peninsula; in the northern half, woods predominate.
Nearly all the forest understory is grazed by cattle,
although grazing is heavier in the southern half.
About one-third of the meadows are fenced and used
for hay. The even-aged stands of aspen in the northern
third of the point have grown up after intensive log-
ging in the 1920s and 1930s and a major fire in 1949.

We sampled small mammals and ground beetles at
three locations along the point in late June and early

TAYLOR AND PFANNMULLER: PENINSULAR EFFECT 145

July of 1975 (Figure 2). The primary study areas were
at Glory Harbor and Kristjanson’s Farm. At both
places small mammals were collected from meadows
and aspen woods at each of two sites. Only one wood-
land site was trapped at Robinson’s Bay. Ground
beetles were collected at all sites but only in the woods.
At Glory Harbor beetle traps were placed in two
patches of woods, one of which was nearly pure aspen,
the other a mixture of aspen and ash. The dominant
understory plant at both sites was Aralia nudicaulis,
Wild Sarsaparilla. We tried to set traps at both sites in
transects spanning the transition from the edge to the
interior of the woods. At the time Forest Tent Cater-
pillars (Malacosoma disstria) were defoliating the
pure aspen stands. Beetle traps at Kristjanson’s Farm
were placed in two patches of woods, one of which
consisted of a fairly open aspen canopy with a dense
ungrazed understory of ash. The second location con-

‘Kristjanson’s Farm

RED DEER POINT)...

FIGURE2. Red Deer Point, showing the three sampling sites.

146 THE CANADIAN FIELD-NATURALIST

tained taller aspens, a more closed canopy, and an
open, grazed understory dominated by hazel (Corylus
sp.). The physical environment varied more among
the sites at Kristjanson’s Farm than among the Glory
Harbor sites, being on the average a bit drier and
brighter. The site at Robinson’s Bay was intermediate
in structure and physical environment between the
other two locations. Its ungrazed understory of Aralia
and Corylus was quite dense. The point of sampling at
various sites at each location was to get a qualitative
feeling for the differences among sites at one location
on the peninsula for comparison to the variation
between locations. If the former overshadows the lat-
ter, then an apparent peninsular effect could merely
reflect random variation among sites.

We captured small mammals with both Museum
Special snap traps and Sherman live traps. Specimens
were deposited in the collection of the Bell Museum of
Natural History, the University of Minnesota.
Ground beetles and shrews were captured in pitfall
traps containing a small quantity of propylene glycol.

The sampling scheme requires explanation. In the
absence of knowledge of the local fauna, the approp-
riate procedure in a study of this sort would be to
sample equally intensively at the base of the peninsula
and at the tip. We followed this procedure with pitfall
trapping for carabid beetles. When the local fauna is
known, however, equal sampling is not the best
procedure. One’s goal becomes the demonstration of
the absence of known species from the tip. This is a
difficult task to achieve by the sampling of acommun-
ity of species of unknown population size. We had
some information on the small mammal fauna based
upon a pilot trapping project in September 1974. The
sampling effort then although small (318 trap nights
with Sherman live traps and 72 pitfall trap nights)
yielded the following species: Peromyscus manicula-
tus (Deer Mice), Clethrionomys gapperi (Southern
Red-Backed Voles), Microtus pennsylvanicus (Mea-
dow Voles), Zapus hudsonius (Meadow Jumping
Mice), Sorex cinereus (Masked Shrews), and Micro-
sorex hoyi (Pygmy Shrews). This collection contained
almost all the small mammals one could expect to
capture in Sherman and pitfall traps in meadows and
aspen woods of this part of Manitoba (Hall and Kel-
son 1959) with the exception of several uncommon
shrews.

Given this information, we devised a sampling
scheme for small mammals in which pitfall trapping
for the relatively unknown shrew community would
be equally intense at Glory Harbor and Kristjanson’s
Farm. We trapped, with Sherman live traps and snap
traps, at Glory Harbor as intensively as possible, and
at Kristjanson’s Farm only until all the species known
to be present were caught.

Vol. 95

Results

In Table | the small mammals are listed. The most
interesting feature of these data is the absence of Cle-
thrionomys gapperi and Sorex monticolus (Dusky
Shrews), at Glory Harbor. Clethrionomys, an easily
trapped woodland vole, was about as common at the
other two locations as Peromyscus maniculatus.
Sorex monticolus, although not common, occurred at
both Kristjanson’s Farm and Robinson’s Bay.

The other differences among sites do not merit
serious consideration. The taking of one chipmunk,
Eutamias minimus, reflects only the setting of a few
traps near a stand of hardwoods on a levee at Glory
Harbor. We set no traps in meadows at Robinson’s
Bay and, as expected, caught neither of the two mea-
dow mice, Microtus pennsylvanicus and Zapus hud-
sonius, there.

The one obvious feature of the ground beetle collec-
tions (Table 2) was that Glory Harbor had nearly as
many species as Kristjanson’s Farm. What difference
does exist pales by comparison to the variability
between the two sites within each of the two locations.

The Carabidae, a diverse group, are noted for their
affinity for particular physical microclimates. The
lack of vegetational identity among the three sites
leads to concern over possible bias toward or against a
peninsular effect. We do not think this a cause for
concern for several reasons. The vegetation is a highly
variable mosaic at its most uniform and probably
provides suitable habitat for any species living in
aspen woods at any point along the length of the
peninsula; moreover, we are not certain if the habitat
differences we perceive are meaningful to carabid bee-
tles living in the leaf litter. Of the 17 species at Krist-
janson’s Farm and 15 at Glory Harbor, 9 were held in
comon, including 6 of the 8 most abundant species. Of
the remaining 14 species not held in common, 8 are

TABLE 1—Comparison of small rodents and insectivores
captured at three sites on Red Deer Point, Lake Winnipego-
sis, Manitoba, in June and July of 1975

Kristjanson’s Robinson’s Glory

Farm Bay — Harbor
Sorex cinereus l 9 16
monticolus 2 l
Microsorex hoyi 3 19 7
Peromyscus maniculatus 2 11 4
Clethrionomys gapperi 3 8
Microtus pennsylvanicus 9 11
Eutamias minimus I
Zapus hudsonius l 3
Total species 7 5 6
Trap nights — 700 101 1200

1981

TAYLOR AND PFANNMULLER: PENINSULAR EFFECT 147

TABLE 2—Comparison of carabid beetle fauna of Red Deer Point in June and July of 1975

Kristjanson’s
Farm

Species Site I
Agonum cupreum*
decentis l
gratiosum l
trigeminum
(complex) 20
Amara cupreolata
Badister obtusus 2
Bembidion (a) 3
Bembidion (b)
Bembidion (c) I
Calathus ingratus* 103
Calosoma frigidum 18
Carabus maeander*
Chlaenius pensylvanicus |
Cymindis cribricollis* D,
Harpalus fulvilabrus 5
pleuriticus
Loricera pilicornis
Metabletus americanus 2
Notiophilus semistriatus*
Patrobus foveocollis*
Pterostichus adstrictus* 31
Sphaeroderus lecontei* 7
Synuchus impunctatus* 26
Total numbers 223
Total species 15
Species per location 17
Total trap days 400

*These species have poorly developed or vestigial wings.

represented by only one specimen. Therefore, of the
original 23 species occurring either at Kristjanson’s
Farm or Glory Harbor, only 6 significant members of
either community could be considered potential habi-
tat specialists.

The choice to collect ground beetles arose from our
impression that as a group they would be most likely
to meet Simpson’s assumption that colonization of
empty habitats is more difficult near the tips of penin-
sulas. As a group, carabid beetles are not known as
good dispersers, although the family is quite variable
in this respect. Calosoma frigidum has large wings and
flies well; it could probably disperse directly to the tip
of Red Deer Point across the lake. By contrast,
Sphaeroderus lecontei with no wings and fused elytra
must walk wherever it goes. The species designated by
asterisks in Table 2 are those that in our collections
had either vestigial or only poorly developed wings.
Some carabid species are dimorphic with respect to
the presence of wings, but we found no winged
morphs in our collections of these species. This is a

Glory
Robinson’s LEO
Site II Bay Site I Site II
]
) 3 21
l
I
9 2 6
]
3 l 1
1 l
l
3 29 6 12
2 | |
3 12
|
|
1
1 2
8 l 9 13
| 3 l 2
l 21
20 67 32 al
9 7 14 10
15
150 225 180 338

conservative list of poor dispersers because some spe-
cies with wings, such as Amara cupreolata, are not
caught in light traps and probably cannot fly (Lin-
droth 1968). Even within this group we think it note-
worthy that Kristjanson’s Farm with six species and
Glory Harbor with five had just about the same
number. The proportion of the species assumed capa-
ble of flight is 53% at Glory Harbor (8 of 15) and 53%
at Kristjanson’s Farm (9 of 17). One would expect the
flying species to comprise a greater percentage of the
fauna at Glory Harbor if there were a peninsular
effect.

Discussion

One could argue that one year’s sampling effort is
inadequate to support or refute the proposed peninsu-
lar effect. This would certainly be true if the hypothe-
sized pattern were weak and fluctuating. If strong and
stable, however, it should be revealed with one sam-
pling effort. Under this assumption, we conclude that
the carabid beetle community offers little support for

148 THE CANADIAN FIELD-NATURALIST

a peninsular effect, inasmuch as the differences
between sites at one location on the peninsula exceed
the differences among locations.

The only data for which we find no reasonable
alternative explanation is the absence from the tip of
Clethrionomys. V oles fluctuate dramatically, and one
could argue that Clethrionomys is merely at a popula-
tion low point at Glory Harbor. Although not an
unreasonable argument, this requires that popula-
tions separated at most by 17 km fluctuate out of
phase. We suggest that this itself would be evidence of
a limitation of dispersal through no other mechanism
than peninsular geometry.

Reserving judgment on Clethrionomys, we con-
clude that Red Deer Point does not exhibit an identif-
iable peninsular effect in either the small mammal or
ground beetle communities. Inasmuch as Taylor and
Regal (1978) have shown that the peninsular effect is
logically possible, we can surmise only that the condi-
tions for its unambiguous demonstration were not
met at this site.

Rates of extinction are extraordinarily difficult to
measure; any comments regarding them that we might
make would be mere speculation. Rates of dispersal
are more approachable, inasmuch as they are inti-
mately linked with the morphologies and habits of the
dispersing species. The possibility of overwater dis-
persal for small mammals in Lake Winnipegosis is
minimal, although some species might be expected to
disperse the length of Red Deer Point over land in as
few as 3 or 4 yr. The point may be too small relative to
the dispersal powers of mammals to allow extinctions
to play an important role in determining distributions.
Clethrionomys may be the exception to this.

The distributions of carabid beetles require differ-
ent explanations. The species known to be capable of
flight might colonize the peninsula quite rapidly over
land or might be blown across the 5-8 km of water
separating the peninsula from the mainland; the
remaining species must either walk or drift on floating
debris. Dispersal rates for several carabid beetles have
been measured by den Boer (1971), as the length of
time required to catch one beetle 100 m from the
population using a standard set of pitfall traps. For
Calathus erratus, a ground beetle of average size
(8-11 mm), approximately 6 mo was required to trap
one individual 100 m from the population. The first
individual of Carabus problematicus, one of the larg-
est ground beetles (21-28 mm), was trapped in
2-3 mo. Although these are conservatively high esti-
mates for both species, they do provide a general idea
of the dispersal power of flightless carabid beetles over
land. If we assume that each generation can expand its
range 100 m, the fact that these beetles are univoltine
means that complete colonization of Red Deer Point

Vol. 95

would take about four centuries. This would seem to
be ample time for the extinction of local populations
to plananimportant role. The absence of a peninsular
decrease among the flightless species points to over-
water colonization as an important factor.

Some recent theoretical work (Taylor, unpublished
material) extends the original model of Taylor and
Regal to include different orientations of peninsulas
to the surrounding mainland. An unsurprising con-
clusion of that work is that of all possible orientations,
a peninsula close to the mainland on both sides 1s least
likely to show a decrease in species diversity near the
tip in taxa capable of overwater dispersal.

We suggest, in conclusion, that the conditions for
expression of a peninsular effect must be quite strict
and that this is one biogeographic pattern that is likely
to be rare.

Acknowledgments

We thank H. Kulman for advice and assistance and
R. Armstrong and S. Taylor for help in the field. We
are grateful in particular for the hospitality of Mr. and
Mrs. T. Kristjanson, R. Kristjanson, and A. Kristjan-
son, without whose generosity the work would have
been impossible to complete. Financial support was
provided by the Graduate School of the University of
Minnesota.

Literature Cited

Cook, R. E. 1969. Variation in species density of North
American birds. Systematic Zoology 18: 63-84.

den Boer, P. J. 1971. On the dispersal power of carabid
beetles and its possible significance. /n Dispersal and dis-
persal power of carabid beetles. Miscellaneous Papers
Landbouwhogeschool, Wageningen 8: 119-137.

Gilpin, M. E. Peninsular diversity patterns. American Natu-
ralist. Jn press.

Hall, E. R., and K. R. Kelson. 1959. The mammals of North
America. Ronald, New York.

Kiester, A. R. 1971. Species density of North American
amphibians and reptiles. Systematic Zoology 20: 127-137.

Lindroth, C. H. 1968. The ground-beetles of Canada and
Alaska. Opuscula Entomologica Supplement 23.

MacArthur, R. H., and E. O. Wilson. 1967. The theory of
island biogeography. Princeton University Press, Prin-
ceton, 203 pp.

Simberloff, D.S. 1976. Experimental zoogeography of
islands: effects of island size. Ecology 57: 629-648.

Simberloff, D.S., and E. O. Wilson. 1969. Experimental
zoogeography of islands. The colonization of empty
islands. Ecology 50: 278-296.

Simpson, G. G. 1964. Species density of North American
Recent Mammals. Systematic Zoology 13: 57-73.

Taylor, R. J.,and P. J. Regal. 1978. The peninsular effect on
species diversity and the biogeography of Baja California.
American Naturalist 112: 583-593.

Received 21 June 1980
Accepted 5 October 1980

Population Characteristics and Movements of Striped Skunks
(Mephitis mephitis) in Central Alberta

RONALD R. BJORGE,' JOHN R. GUNSON,! and WILLIAM M. SAMUEL?

'Alberta Fish and Wildlife Division, 6909-116 Street, Edmonton, Alberta T6H 4P2
2Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9

Bjorge, Ronald R., John R. Gunson, and William M. Samuel. 1981. Population characteristics and movements of Striped
Skunks (Mephitis mephitis) in central Alberta. Canadian Field-Naturalist 95(2): 149-155.

Population characteristics and movements of Striped Skunks (Mephitis mephitis) were studied on a 130-km? area in central
Alberta during 1971-1974. Estimates of a minimum density were amongst the lowest recorded; 0.7-1.2/km?. Following the
long winter of 1973-1974, numbers of young in natal dens and numbers of skunks on the study area were less than in 1973. Sex
ratio of neonates was 1:1 but that of adults favored females. Adult male skunks moved greater distances than adult females;
independent juveniles moved greater distances than adults. Dispersal was greatest during July and August when juveniles
were about 3 mo of age; maximum dispersal was 22 km. The extensive dispersal of juvenile skunks, and lowered litter size and
reduced density following a severe winter have not been reported from more southerly populations.

Key Words: Striped Skunk, Mephitis mephitis; population dynamics, movements, dispersal.

The Striped Skunk (Mephitis mephitis), common
throughout much of North America (Halland Kelson
1959), is of considerable public and animal health
significance, particularly as a vector of rabies and
leptospirosis (Ferris and Andrews 1966; Sikes 1970;
Parker 1975). Studies of wild populations, including
population characteristics and movements, have been
limited to the eastern and central United States (Allen
1939: Allen and Shapton 1942; Verts 1967; Upham
1967; Houseknecht 1971; Bailey 1971; Storm 1972).
The purpose of this paper is to summarize population
trends, reproduction, sex and age ratios, and move-
ments of adults and juveniles in a population of
Striped Skunks in central Alberta, near the northern
limit of agriculture, and to compare them with skunks
of more southerly areas.

Study Area

The 130-km?2 Tofield study area (approximately
53°21’N, 112°43’W) was gridded by gravel roads
every 1.6 km N/S and every 3.2 km E/ W. The area is
typical of agriculturally modified Trembling Aspen
(Populus tremuloides) parkland which covers much
of eastcentral Alberta (Bird and Bird 1967). The area
contained eight aspen stands ranging in size from 30 to
65 ha, and many smaller patches of trees and shrubs in
low or hilly areas, along fences, roads, and in farm-
yards. Most farms were about 200 ha; raising of cereal
and forage crops and livestock predominated. Of 112
farmyards, 72 were occupied by humans while 40 were
abandoned.

The climate is continental; characterized by warm
summers and cold winters (Government of Alberta
and University of Alberta 1969). The mean tempera-
ture from May through September is 13°C and from

149

November through March -9°C. The frost-free period
is about 100 d and the mean annual precipitation is 45
cm with 30% falling as snow. Landscape is undulating
to gently rolling (0-9% slope) and natural drainage is
good (Bowser et al. 1973).

Methods

The study was carried out during 1971 through 1974
in conjunction with studies of skunk rabies in eastern
Alberta and Saskatchewan (Gunson et al. 1978).
Skunks were captured by livetrapping (wooden and
metal box traps), night-lighting, and excavation of
dens. During April through October, 50 to 60 live
traps were placed throughout the study area at loca-
tions ~ 800 m apart and subjectively determined to be
good areas to trap skunks. Such locations commonly
included fence rows, abandoned buildings, woodlot
fringes, and culverts. Traps were checked daily and
baited about once per week, usually with chicken
entrails. Skunks were detected during night-lighting
froma pickup truck equipped with two aircraft land-
ing lights. These skunks were pursued on foot, netted,
and handled similarly to live-trapped skunks.

Skunks were captured or counted at natal dens
located beneath buildings by removing floor boards
or digging under the foundation during June and July.
Young skunks were returned to the den within | h.
Buildings and diggings were restored before skunks
were returned. In December 1974, at the end of the
study, skunks were gassed with carbon monoxide
while in winter dens under buildings. Uteri were
removed and prepared for counts of placental scars
(Orsini 1962).

Skunks were handled as described by Jacobson et
al. (1970). Eachear was tagged with numbered finger-

150 THE CANADIAN FIELD-NATURALIST

ling metal tags. Fourteen skunks, including nine adult
females and five juveniles were equipped with radio
collars. Transmitters were similar to those utilized by
Brand et al. (1975) and constructed by the Depart-
ment of Electrical Engineering, University of Alberta.
Locations were determined by triangulation with a
loop antenna and a portable receiving unit. Most
telemetry locations were determined during daytime
when skunks were resting.

Juveniles were distinguished from adults by body
weight, size, and general appearance. As juveniles
were similar in appearance to adults after October,
and most were captured in natal dens during May
through July, only skunks captured during August
through October were used in calculating age ratios.

Estimates of annual variation in numbers were
derived from livetrapping success during August and
September because comparable data were available
from all years of the study only during these months;
in 1971 trapping began in late July and in 1972 trap-
ping was conducted only in spring and fall.

Greatest distance between the most extreme cap-
tures during a specified period (Brant 1972; Sander-
son 1966; Bailey 1971) was utilized as an index of
home range and dispersal. Because of the relatively
wide spacing of traps, some greater movements likely
were not recorded. Information derived from teleme-
try was utilized to calculate home ranges (Mohr 1947)
and minimum daily movements.

Weather records were taken from Environment
Canada, Annual Meteorological Summaries 1970-
1974. Temperature records were from Camrose,
Alberta, about 30 km south of the study area. Precipi-
tation records were available only from Edmonton,
Alberta, about 70 km to the west.

Results and Discussion
Reproduction

During 1973 and 1974, 42 of 48 (88%) adult females
were either lactating or had suckled young (32 of 35 in

Vol. 95

1973, 10 of 13 in 1974). Such high pregnancy rates
have been noted elsewhere; Verts (1967) found 72 of
75 (96%) females in Illinois were pregnant, while Gun-
son and Schowalter (unpublished data) observed that
99 of 112 (88%) females from eastern Alberta and
Saskatchewan were pregnant or had given birth to
young.

Mean litter size + SE (counts of young in natal
dens) was larger in 1973 (5.2 + 0.8 of nine litters) than
in 1974 (2.5 + 0.4 of six litters) (t-test, P< 0.05). Lit-
ter sizes of wild skunks, including embryos, neonates,
and placental scars, range from 5.8 to 7.3 (Allan 1939;
Hamilton 1963; Verts 1967; Bailey 1971; Gunson and
Schowalter, unpublished data), suggesting our 1974
skunk litters were considerably smaller than those
reported elsewhere.

Difference in litter size between 1973 and 1974 was
likely related to snowfall and temperature during win-
ter denning (Table 1). Snowfall was about 1.7 times
greater during winter 1973-1974, with more snow fal-
ling earlier and staying longer. Temperatures were
also lower during November 1973 and March 1974,
usually the first and last months of winter denning
(Gunson and Bjorge 1979), than | year earlier. Analy-
sis of age data, as follows, suggested that reproduction
and/or survival of juveniles was low following the
relatively heavy snowfall and long winter of
1970-1971, as well. Of the skunks captured during
August through October, 18 of 37 (49%) were juve-
niles in 1971 compared to 23 of 33 (70%) in 1972, 52 of
75 (69%) in 1973, and 35 of 60 (58%) in 1974.

In northern parts of their geographic range, such as
Alberta, Striped Skunks spend more time within the
winter den during long winters than short winters
(Gunson and Bjorge 1979). As they do not feed during
winter denning at northern latitudes (Mutch and
Aleksiuk 1977), their body condition would likely be
poorer after long, cold winters. Skunks in Minnesota
lost a mean of 49% body weight during winter 1968
followed by significantly greater mean losses of 58%

TABLE |—Weather records during winter denning, parturition (May), and lactation (May-June) of Striped Skunks at

Tofield, Alberta®

Variable Month 1970-71 1971-72 1972-73 1973-74
Snowfall (cm) October—April 100 167 130 219
Snow on ground (cm) 30 November 25 3 5 20
31 March 56 10 0 74
Rainfall (cm) April-June 20 17 32 16
Mean monthly temperature (°C) November 9.3 ~14.5 3.8 -14.1
December 17.6 16.8 15.8 -12.9
January 18.9 19.3 -12.5 -15.3
February 9.6 -16.0 -10.1 -I1.1
March 7.8 -6.1 -3.1 -12.1

“Most important are the snow on ground 30 November and 31 March and temperatures during November and March.

1981

during the longer winter of 1969 (Sunquist 1974).
Breeding occurs primarily during March (Wright
1931; Hamilton 1963; Bjorge 1977) near the end of
winter denning. It follows that reproduction and sur-
vival of juveniles may be lowered as a result of a long
winter. Lower reproduction and survival of young
during periods of decreased food supply have been
documented for several carnivores including Coyotes
(Canis latrans) (Clark 1972) and Lynx (Lynx cana-
densis) (Brand et al. 1976).

Numbers of placental scars = SE from 15 females
with scars killed during December 1974 averaged
6.0 + 0.5; 2.4 times as great as the mean number of
young counted in dens that year. Previous studies
documented resorption of embryos (Verts 1967) and
death of small juveniles within a few days of birth
(Rakowski 1972). Increased resorption of embryos
and neonatal mortality after a long winter likely
accounted for the small litters observed in 1974. As
well, the cooler temperatures recorded during May
1974 could have hampered survival of neonates
because natal dens of Striped Skunks are often in
relatively exposed (granaries and other structures
without foundations) sites in Alberta (Bjorge 1977).
Heavy rainfall occurred during May and June 1973
when litters were largest, suggesting that rain during
the first 2 mo after birth was not a negative factor.

Numbers, Age, and Sex

Livetrapping success during August and September
increased from 1971 through 1973, and then decreased
in 1974, suggesting two- to three-fold changes in popu-
lation densities (Figure 1). Total numbers captured
during May through October in 1974 (93) were lower
(chi-square test, P< 0.001) than in 1973 (155) despite
increases in trapping and night-lighting efforts in 1974
(4757 trap-nights versus 4082 in 1973; 448 km versus
121 km in 1973).

Marked fluctuations in numbers of Striped Skunks
have been reported (Allan and Shapton 1972; Verts
1967). Verts (1967) summarized success of skunk
trappers during 25 seasons in Illinois; twofold changes
in trapper success during successive years likely
reflected comparable changes in density. Allan and
Shapton (1942) attributed population variations to
“disease” while Verts (1967) provided no explanation.
Tests for rabies virus and antibodies to leptospirosis,
tularemia, and western equine encephalitis from 47
skunks collected at the termination of this study were
negative (J. Iverson, Western College of Veterinary
Medicine, Saskatoon, and D. Schowalter, Alberta
Fish and Wildlife Division, Edmonton). It is unlikely
that disease was a significant factor in population
regulation of skunks in this study.

Decreased recruitment of young in 1974 was not
entirely responsible for lowered populations that year.

BJORGE ET AL.: STRIPED SKUNKS, CENTRAL ALBERTA 151

200

2

175

150

August
September

125

100 700

HTN TRSNNN TELE

75

CAPTURES/1000 TRAP-NIGHTS

tA WH)

50

25

1971 1972 1973 1974
YEAR
FIGURE 1. Success of Striped Skunk captures at Tofield,

Alberta, during August and September, 1971 through
1974. Numerals indicate trap-nights.

Fewer adults were located in 1974 (37) than 1973 (51)
despite greater capture efforts in 1974. Sunquist
(1974) thought that winter-related mortality of skunks
was important in Minnesota. Limited data from this
study support that; we found I! dead skunks under
buildings following the severe winter compared to 3
following the mild winter.

Our observations of capture success, litter size, and
age structures indicate that populations of Striped
Skunks in northern areas are importantly influenced
by severity of weather during the preceding winter,
especially the length of winter. Numbers of skunks
decrease following long winters and increase follow-
ing short winters. Particularly important is accumu-
lated snow during the first and last months of winter,
usually November and March. In northern areas, win-
ters likely have greater potential regulation of
numbers of Striped Skunks. Densities of skunks in
more southerly habitats with less rigorous winters are
more apt to be regulated by other factors.

Minimum density estimates in our study, based on
total captures for 1973 and 1974, were 1.2/ km? and

152 THE CANADIAN FIELD-NATURALIST

0.7/km?*, respectively. Other estimates, also deter-
mined from total numbers of skunks captured within
a given area, varied from 0.4—1.4/ km? in Iowa (Scott
and Selko 1939) to 11.5—-27.0/km2 in Illinois (Ferris
and Andrews 1966). Additional estimates of density,
determined from a variety of techniques, varied from
1.9-3.1/km? in North Dakota (Upham 1967) to
4.4-4.6/km2? in Ohio (Bailey 1971).

The sex ratio of captured adults was significantly
unbalanced in favor of females (chi-square test,
P< 0.01) (Table 2). This pattern was comparable in
both adult skunks captured by livetrapping (36%
males of 120) and night-lighting (35% males of 23).
When sex ratios for all years were combined by
month, more females than males were captured in all
months except April when 7 of 13 skunks were males.
All evidence indicated that more adult females than
males were present as reported elsewhere (Bailey 1971;
Storm 1972). Verts (1967) observed a change from a
ratio favoring males during the first 3 yr to one favor-
ing females during the last 2 yr of his 5-yr study. Most
collections or captures of skunks during winter and
early spring (Bennitt and Nagel 1937; Hamilton 1937;
Casey and Webster 1975) have favored males, likely
because males are more active than females during
cold months (Gunson and Byjorge 1979).

The apparent change in sex ratio from one favoring
males among juveniles (Table 2) to one favoring
females among adults suggests a greater mortality rate
for adult males. Adult males are intolerant of each
other (Allan and Shapton 1942), more active during
cold months, often den alone during winter, may
compete with other males for aggregations of females
(Gunson and Bjorge 1979), and travel extensively in
early spring during mating (Allan and Shapton 1942;
Ferris and Andrews 1966). Males are probably subject
to higher mortality from accidents, fighting, preda-
tion, and human causes, because of the above factors.

The sex ratio of juveniles after leaving the natal den
favored males [84 of 138 (61%)] (chi-square test,
P< 0.01) (Table 2). That of neonates in dens also
favored males [35 of 62 (56%)], but the difference was

TABLE 2—Percentage (N) male Striped Skunks captured at
Tofield, Alberta, during April through October, 1971-74.

% males (N)
Year Adults Juveniles®
1971 37(19) 61(18)
1972 48(33) 65(23)
1973 29(51) 54(56)
1974 38(37) 68(41)
Mean 37(140) 61(138)

“Includes juveniles captured July through October (excludes

juveniles captured in natal dens).

Vol. 95

not significant; a finding not uncommon for neonates
and embryos of wild skunks (Hamilton 1963: Verts
1967). Monthly sex ratios of independent juveniles (all
4 years combined) varied from 67 to 69% males during
July, August, and September, but dropped to 53%
males during October. Possible reasons for this
change would include differential trapability, mortal-
ity, or dispersal.

Previous studies indicate varying sex ratios among
independent juveniles. Verts (1967) and Storm (1972)
documented 44 and 45% males among 84 and 42
livetrapped juveniles in Illinois. In Saskatchewan and
eastern Alberta, Gunson and Schowalter (unpub-
lished data) found 56% of 364 juveniles were males.

Movements of Adult Skunks

Comparison of greatest linear distance between
captures of adult skunks indicated larger home ranges
for males than females (Table 3). Upham (1967) and
Storm (1972) also recorded more extensive move-
ments for adult male than adult female Striped
Skunks in North Dakota and Illinois, respectively,
but Houseknecht (1971) observed the opposite in
Minnesota.

TABLE 3—Mean = SE greatest linear distance and ranges
between captures of Striped Skunks at Tofield, Alberta,
during May through October, 1973 and 1974

Range®

Number of Mean
Cohort skunks distance (m) (m)
Adult males 17 2556+ 231 1200-4 050
Adult females 30 1970 +170 700-4 200
Total 47 2197+ 143 700-4 200
Juvenile males 29 3310 + 410 850-10 100
Juvenile females 19 4150+ 1191 650-21 700
Total 48 3640 + 640 650-21 700

“Measured to the nearest 50 m.

Seasonal variation in home range, as indexed by
greatest linear distance between captures, was
obvious. In general adult females were most sedentary
during parturition and lactation (May and June), and
travelled more following weaning (July and August)
and less during predenning (September and October)
(Table 4). Adult males moved increasingly greater
distances from May and June through September and
October. Greater movement of adult males during fall
was also reported by Storm (1972). During winter,
communes of females generally den with one male,
usually an adult (Gunson and Byjorge 1979). Adult
males appear to search for groups of females during
fall (Gunson and Bjorge 1979), leading to greater
mobility during that period.

1981

TABLE 4— Mean + SE (JN) greatest distance movements (m)
of adult Striped Skunks between capture at Tofield, Alberta,
during 2—mo periods between | May and 31 October, 1973
and 1974

September-—
Cohort May-June July-August October
Adult
male 1782+234(4) 2108+462(11) 2849+369(8)
Adult
female 510+214(11) 22074£331(12) 1260£269(8)

Mean home range calculated according to Mohr
(1947) for six adult females radiotracked for 45-105 d
(mean of 44 fixes, range 20-62) was 208 ha (range
110-370 ha). Storm (1972) reported mean home
ranges of five adult females that were radiotracked for
more than 15 d during summer at 362 ha.

Minimum daily movement (distance between dens
used on consecutive days) for five radio-collared adult
females using 41 dens was 1.5 km (range 0.1-2.8 km).
One adult female was extremely mobile, moving a
mean of 2.3 km between dens during 8 d; however, all
these movements were within one home range of 370
ha. The greatest distance between daily locations for
this skunk during 97 d of radiotracking (51 fixes) was
3.7 km. This suggests that adult skunks are capable of
moving over a large portion of their range withina day
Or so.

Movements of Juveniles

Juvenile skunks moved significantly greater distan-
ces (t-test, P< 0.05) between captures than adults
(Table 3). At least 4 of 62 juveniles marked in natal
dens during June 1973 and 1974 dispersed, moving a
mean distance of 11.1 km (dispersal is defined as any
straight-line movement greater than 4.8 km, because
no adults moved more than this distance). Of the four
juveniles, only one was captured on the study area
beyond August, despite intensive livetrapping and
night-lighting during 1973 and 1974 and excavation of
winter dens in December 1974. This skunk, a male,
was present on the study area about 8 km from his
natal den, during September of the year following
marking. Of 196 juveniles captured (including those
captured in natal dens), 148 were caught only once. Of
the 48 recaptures, 11 (23%) had dispersed (Table 5).

These data suggest that most juveniles born on the
study area during 1973 and 1974 dispersed. Occa-
sional observations at six natal dens following excava-
tion indicated that families of skunks usually did not
vacate the den after marking. Death resulting from
capturing and marking neonates was not considered a
significant mortality factor. Survival of juveniles after

BJORGE ET AL.: STRIPED SKUNKS, CENTRAL ALBERTA 1593}

TABLE 5—Sex, dates, and distances of dispersing juvenile
Striped Skunks at Tofield, Alberta, during 1973 and 1974.
Includes all skunks known to have travelled straight-line
distance % 4.8 km

Date of first and last capture Distance’ (km)

Males ;
7 July — 17 August 10.1
26 June — 29 August 8.4
21 August — 28 August 8.1
24 July — 14 August D2
1 August — 21 August 48
15 August — 27 August 5.1
28 August — 30 August 8)
Females
4 July — 31 August Ned)
17 July — 30 November 13.7
1 October — 30 October 5.8
1 September — 22 October 5.6

“Measured the nearest 50 m.

marking was demonstrated at 7 of 15 dens while no
data were available from the other dens. Maximum
recorded distance of dispersal was 22 km. Dispersing
juveniles travelled relatively great distances within a
short time; a radio-instrumented male travelled
8.0 km within 6d, a juvenile male moved 5.8 km
within 2 d,and a juvenile female moved 3.7 kmin 1 d.

Most dispersal occurred between weaning in early
July (Bjorge 1977) and | September. Recapture rates
of juveniles first captured in natal dens during June
and July were lower than recapture rates of adults
captured during the same period (chi-square test,
June, P< 0.005; July, P< 0.05) (Table 6). Juveniles

TABLE 6— Occurrence of recapture and mean number of days
(+ SE) between first and last capture during the same year
for adult and juvenile Striped Skunks captured at Tofield,
Alberta, between May and October, 1973 and 1974.

Recapture Time

Month Age N (%) (d)
May Adult 16 81 76+£16
June Adult 21 62 88+31
Juvenile* 62 8 91+16
July Adult 18 61 T1+£16
Juvenile 17 24 19+ 4
August Adult 13 69 60+14
Juvenile 31 48 36+ 8
September Adult 12 75 46+13
Juvenile 31 »y 44+ 7
October Adult 5 100 44+13
Juvenile 17 47 29+ 6

“Includes juveniles captured in natal dens.

154 THE CANADIAN FIELD-NATURALIST

captured during July and August were known to be
alive on the study area for fewer days (based on days
between first and last capture) than adults captured
during these months (July, P< 0.005; August,
P<0.05). The known periods of dispersal (Table 5)
and the high rate of disappearance of marked juven-
iles from the study area indicate that dispersal was
greatest when juveniles were 3 to 4 mo of age.

Juvenile skunks in Illinois, monitored during late
summer and fall by telemetry and livetrapping, moved
up to 2.2 km (Verts 1967). Storm (1972) concluded
that the home ranges of radio-marked juveniles in
Illinois were smaller than those of adults, and Bailey
(1971), in Ohio, determined that the average straight-
line distance between live captures was less for juven-
iles than adults. The most extensive movement of a
juvenile in his study was 2.1 km over 27 d. Upham
(1967) noted that juveniles in his study in North
Dakota moved only slightly greater mean distances
than adults; one juvenile travelled nearly 9 kin.

These observations suggest that juvenile skunks in
Alberta disperse greater distances than those in more
southerly areas. Gunson and Bjorge (1979) suggested
that the observed mobility of juveniles, particularly
females, could be related to a search for a suitable and
available winter den. Although female skunks in cen-
tral Alberta den in winter communes, juvenile females
may be excluded from some dens (Gunson and Byjorge
1979). Winter dens are likely more widely dispersed in
central Alberta than in more southerly areas. Exten-
sive dispersal of juveniles is of major significance to
management of diseases transmitted by skunks. The
potential for spread of disease by highly mobile juven-
ile skunks is recognized.

Acknowledgments

This study was supported by the Veterinary Servi-
ces Division, Alberta Agriculture, the Fish and Wild-
life Division, Alberta Energy and Natural Resources,
the National Research Council of Canada (operating
grant A-6603 to WMS), and the University of Alberta.
The interest and support of H. Vance of the Veteri-
nary Services Division is gratefully acknowledged. P.
Andersen, E. Ewaschuk, W. Johnson, L. Pecharsky,
D. Pipella, and W. Wynnyk capably assisted in the
field. D. Schowalter and L. B. Keith reviewed drafts
of this paper.

Literature Cited

Allen, D. L. 1939. Winter habitats of Michigan skunks.
Journal of Wildlife Management 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: 59-68.

Bailey, T. N. 1971. Biology of Striped Skunks on a south-
western Lake Erie marsh. American Midland Naturalist
85: 196-207.

Vol. 95

Bennitt, R., and W. O. Nagel. 1937. A survey of the resi-
dent game and furbearers of Missouri. University Mis-
souri Studies 12: 1-215. (Not seen, cited by Verts 1967.)

Bird, C. D., and R. D. Bird. 1967. The aspen parkland.
In Alberta—a natural history. Edited by W. D. Hardy.
Hurtig Publishers, Edmonton. 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, Alberta. 96 pp.

Bowser, W. E., A. A. Kjearsgaard, T. W. Peters, and R. E.
Wells. 1973. Soil survey of Edmonton (sheet 83H).
Alberta Soil Survey Report No. 21. University of Alberta,
Edmonton.

Brand, C. J., R. H. Vowles, and L. B. Keith. 1975. Snow-
shoe hare mortality monitored by telemetry. Journal of
Wildlife Management 39: 741-747.

Brand, C. J., L. B. Keith, and C. A. Fischer. 1976. Lynx
responses to changing Snowshoe Hare densities in central
Alberta. Journal of Wildlife Management 40: 416-428.

Brant, D. 1962. Measures of the movements and popula-
tion densities of small rodents. University California Pub-
lication Zoology 62: 105-184.

Casey, D.A., and W. A. Webster. 1975. Age and sex
determination of Striped Skunks (Mephitis mephitis)
from Ontario, Manitoba and Quebec. Canadian Journal
of Zoology 53: 223-226.

Clark, F. W. 1972. Influence of jackrabbit density on
Coyote population change. Journal of Wildlife Manage-
ment 36: 343-356.

Ferris, D. H., and R. D. Andrews. 1966. Parameters of a
natural focus of Leptospira pomona in skunks and oppos-
sums. Bulletin of the Wildlife Disease Association 3: 2-I1.

Government of Alberta and the University of Alberta.
1969. Atlas of Alberta. University of Alberta Press,
Edmonton, Alberta. 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.

Gunson, J. R., and R. R. Bjorge. 1979. Winter denning of
the Striped Skunk in Alberta. Canadian Field-Naturalist
93: 252-258.

Hall, E.R., and K.R. Kelson. 1959. The mammals of
North America (Vol. I and II). Ronald Press Company,
New York. 1083 pp.

Hamilton, W. J., Jr. 1937. Winter activity of the skunk.
Ecology 18: 326-327.

Hamilton, W. J., Jr. 1963. Reproduction of the Striped
Skunk in New York. Journal of Mammalogy 44: 123-124.

Houseknecht, C. R. 1971. Movements, activity patterns
and denning habits of Striped Skunk (Mephitis mephitis)
and the exposure potential for disease. Ph.D. thesis, Uni-
versity of Minnesota, Minneapolis. 46 pp.

Houseknecht, C. R., and J. R. Tester. 1978. Denning hab-
its of Striped Skunks. American Midland Naturalist 100:
424-430.

Jacobson, J.O., E.C. Meslow, and M.F. Andrews.
1970. An improved technique for handling Striped
Skunks in disease investigations. Journal of Wildlife Dis-
eases 6: 510-512.

Mohr, C. O. 1947. Table of equivalent populations of an
endangered species. American Midland Naturalist 37:
223-249.

1981

Mutch, G. P.,and M. Aleksiuk. 1977. Ecological aspects of
winter dormancy in the Striped Skunk (Mephitis mephi-
tis). Canadian Journal of Zoology 55: 607-615.

Orsini, M. W. 1962. Technique of preparation, study and
photography of benzyl-benzoate cleaned material for
embryological studies. Journal of Reproduction and Fer-
tility 3: 283-287.

Parker, R. L. 1975. Rabies in Skunks. /n Natural History
of Rabies. Edited by G. M. Baer, Academic Press, New
York. pp. 41-51.

Rakowski, P. W. 1972. Studies on the Striped Skunk in
southeastern North Dakota. M.Sc. thesis, North Dakota
State University, Fargo. 62 pp.

Sanderson, G. C. 1966. The study of mammal movements
—a review. Journal of Wildlife Management 30: 215-233.

Scott, T. G.,and L. F. Selko. 1939. A census of Red Foxes
and Striped Skunks in Clay and Boone counties, Iowa.
Journal of Wildlife Management 3: 92-98.

Sikes, R. K. 1970. Rabies. /n Infectious diseases of wild
mammals. Edited by J. N. Davis, L. N. Karstad, and

BJORGE ET AL.: STRIPED SKUNKS, CENTRAL ALBERTA 155

D. W. Trainer. lowa State University Press, Ames. pp.
3-19.

Storm, G. L. 1972. Daytime retreats and movements of
skunks on farmlands in Illinois. Journal of Wildlife Man-
agement 36: 31-45.

Sunquist, M. E. 1974. Winter activities of Striped Skunk
Mephitis mephitis in east-central Minnesota. American
Midland Naturalist 92: 434-446.

Upham, L. L. 1967. Density, dispersal, and dispersion of
the Striped Skunk (Mephitis mephitis) in southeastern
North Dakota. M.Sc. thesis, North Dakota State Univer-
sity, Fargo. 61 pp.

Verts, B. J. 1967. The biology of the Striped Skunk. Uni-
versity of Illinois Press, Urbana, Illinois. 218 pp.

Wright, H. M. 1931. Reproduction in the eastern skunk
(Mephitis mephitis nigra). Journal of Mammalogy 12:
42-47.

Received 25 March 1980
Accepted 6 October 1980

Natural History of the Ebony Spleenwort,
Asplenium platyneuron (Aspleniaceae), in the Great Lakes Area

WARREN H. WAGNER, JR., and DAVID M. JOHNSON

Department of Botany and Herbarium, The University of Michigan, Ann Arbor, Michigan 48109

Wagner, Warren H., Jr.,and David M. Johnson. 1981. Natural history of the Ebony Spleenwort, Asplenium platyneuron
(Aspleniaceae), in the Great Lakes area. Canadian Field-Naturalist 95(2): 156-166.

Once very rare in the Great Lakes region, the Ebony Spleenwort, Asp/enium platyneuron, appears to be spreading rapidly in
many localities where it was previously unknown. The species had been either very sporadic or absent in this region. The major
habitats are dry forest and forest edges, shaded steep road banks, pine plantations, and second-growth old fields, pastures,
and orchards. The fronds average 14-16 per plant, the number of low, short, sterile leaves usually twice as many as upright,
tall, fertile leaves. Much of the starch storage occurs in the thickened petiole bases which may be retained fora number of years
after the rest of the frond has died. Vegetative proliferation by tiny plantlets produced on the lowest pinnae takes place in
about | of 20 plants growing in dry upland sites and | of 5 in moist lowland sites. Variations with strongly incised pinnae are
rare, but in at least one population | of 8 plants showed this condition. Intermediates between incised and nonincised forms
are also found. We hypothesize that the invasive success of A. platyneuron is due to its ability to grow terrestrially in
competition with other plants, its tall sporophylls, the capacity of storing considerable starch in the petiole bases, and the
ability to propagate vegetatively under certain conditions. Factors such as increased area of second-growth habitats and
warming climatic conditions may have also played a role. We believe that none of the so-called “varieties” of this species merit
that taxonomic designation; they should be treated instead as mere trivial forms without special nomenclature.

Key Words: Ebony Spleenwort, Asp/enium platyneuron; range expansion, leaf dimorphy, starch storage, vegetative prolifer-
ation, incised variations, taxonomy.

Ebony Spleenwort, Asplenium platyneuron, is some of which were cleared in 5% aqueous NaOH,
unusual among temperate North American spleen- stained with safranin, and drawn under a micropro-
worts in being terrestrial as well as saxicolous. The _jector for examination of pinna and venation varia-
species occurs from Quebec and Ontario south to tion. We examined foliar storage organs using hand
Florida and Arizona, becoming rare and sporadic sections and the iodine test for starch.
near the periphery of its range. In the Great Lakes
region it was considered a very rare fern until the Distribution of Ebony Spleenwort

1960s (Billington 1952), but during the past 20 years it A northward range expansion has taken place in the
has been invading this region, spreading rapidly and past 20 years (see Figure | where the dots represent
forming large populations. older records and the stars more recent ones). This

Throughout its range the species is highly variable, trend is supported by statements from numerous field

and six infraspecific taxa have been recognized, five of | workers based on their experiences over the last two
which have been found to occur in the Great Lakes decades. In Kalamazoo County, Michigan, the species
area. The biological significance of these taxa has not was once unknown (Hanes and Hanes 1947), but Har-
been studied adequately, and the purpose of this study vey Ballard of Western Michigan University (personal

was to elucidate the factors allowing Ebony Spleen- communication 1979) informed us that now “It
wort to increase its range, and to determine the nature appears to be an aggressive and ‘weedy’ species, com-
and extent of this variability. mon in many places in the county.” Thomas Carlson
of The University of Michigan (personal communica-

Materials and Methods tion 1979) wrote that in the Three Rivers Game
To assess the expanding patterns in the range of A. Reserve in St. Joseph County, Michigan, “the original
platyneuron, we used our field observations in Michi- ten populations (in 1971) have now expanded, while
gan, published reports, data from herbarium speci- many new ones have become established, particularly
mens, and information provided by various individu- on disturbed ground along roads, trails, and corn
als for Michigan and elsewhere. We recorded plant fields, and in pine stands.” In Ontario, A. A. Reznicek
species growing within several metres of spleenwort of The University of Michigan (personal communica-

plants in a number of habitats in Michigan. We also tion 1980) reported that this species, once confined to
studied frond structure and variation by collecting the Kingston and Georgian Bay areas, “has undergone
random samples of plants from large populations, a tremendous explosion in the past decade... The

156

1981

WAGNER AND JOHNSON: EBONY SPLEENWORT, GREAT LAKES AREA

SY

FiGURE |. The distribution of Ebony Spleenwort in the Great Lakes region. Dots represent records from the following
sources: Illinois, Mohlenbrock (1967); Indiana, Deam (1940); lowa, Cooperrider (1959); Michigan, Billington (1952);
Minnesota, Tryon (1954); Ohio, Adams (1958); Ontario before 1965, A. A. Reznicek (personal communication 1980);
and Wisconsin, Tryon et al. (1953). Stars indicate records since these respective dates supplied by individuals (see
Acknowledgments), who examined specimens in herbaria in the state or province with which they are identified. Large
stars serve to emphasize peripheral stations, and the irregular line through Iowa, Indiana, Illinois, Ohio, Pennsylvania,
and New York represents the glacial boundary. The inset shows the entire range of the Ebony Spleenwort.

plant can now barely be considered rare in Ontario.”
Charles Sheviak of New York State Museum, Albany,
(personal communication 1980) considered A. platy-
neuron “now a common plant throughout Illinois,”
where Pepoon (1927) called it “the rarest of our ferns”
in the Chicago region.

The best documented instance of increase in indi-
vidual population size was provided by the work of
Arthur J. Cooper of North Carolina State University,
who found and mapped the localities of six plants at
five sites in a 1958 study at The University of Michi-

gan’s Edwin S. George Reserve in Livingston County,
Michigan. He was impressed that he never found
more than one plant (except for two plants at one
locality) at any one place, and therefore collected no
vouchers to avoid eradicating any of the populations.
He provided us, however, with a minutely detailed
map of the 1958 localities which enabled us to revisit
each site in 1979 and find out what had happened to
the five populations in the intervening 21 years. At
four of the five sites where plants could still be located,
the number of plants had increased from 6 to 34. In

158

addition, we found 122 plants at three additional sites
within the Reserve, making it fairly common where in
1958 it had been rare and local.

The Ebony Spleenwort is clearly extending its range
northward, being known now from Chippewa County
in the Upper Peninsula of Michigan (Hagenah 1955),
Manitoulin Island, Ontario (J. Morton, University of
Waterloo, personal communication 1980), and from
the vicinity of Ottawa (D. Britton, University of
Guelph, personal communication 1980). Even consid-
ering the amount of field work that has been carried
out by botanists in this region since 1950, this increase
in range exceeds normal expectations. If this popula-
tion explosion continues, néw localities for this spe-
cies should be discovered at the northern and western
peripheries of its range where it was never before
anticipated.

Most ferns of the genus Asp/enium in temperate
North America occur in crevices, on rocks, or on
talus, and are unable to grow in competition with
other vascular plants. In contrast, Ebony Spleenwort
usually grows directly upon ground soil interspersed
with many species of other vascular plants. In south-
ern Michigan we have found it principally in dry forest
and forest edges, on shaded steep road banks, in pine
plantations, and in second-growth old fields, pastures,
and orchards. These habitats are discussed below in
order of increasing frequency of the ferns.

In all habitats, including the forest, growth of the
spleenwort appears to be fostered by disturbance. In
the forest this is provided by natural and artificial light
gaps and clearings and by erosion on steep slopes. The
typical forest occurrences are in dry upland woods of
Red Maple (Acer rubrum), Serviceberry (Amelanch-
ier arborea), Pignut Hickory (Carya glabra), Black
Walnut (Jug/ans nigra), Black Cherry (Prunus sero-
tina), White Oak (Quercus alba), Red Oak (Q. rubra),
and Black Oak (Q. velutina) which have a well-
developed oak litter, and associated shrubs including
Black Huckleberry (Gaylussacia baccata), Fall
Witchhazel (Hamamelis virginiana), Common
Juniper (Juniperus communis), Prickly Gooseberry
(Ribes cynosbati), and Black Raspberry (Rubus occi-
dentalis). Ebony Spleenwort also occurs on steep road
banks in such forested areas, often growing with Fra-
gile Fern (Cystopteris fragilis var. mackayit).

Plantations of Scots Pine (Pinus sylvestris) or
White Pine (P. strobus) in southern Michigan usually
support small Ebony Spleenwort populations. Here
the plants grow in the thick needle layer with other
plantation adventives such as Manitoba Maple (Acer
negundo), Tatar Honeysuckle (Lonicera tatarica),
White Mulberry (Morus alba), Black Cherry, Prickly
Gooseberry, Swamp Nightshade (Solanum dulcam-
ara), Black Raspberry, and Red-berried Elder (Sam-
bucus pubens).

THE CANADIAN FIELD-NATURALIST

Vol. 95

Second-growth old fields, pastures, and orchards in
later stages of succession (Figure 2) provide the most
productive habitats for the Ebony Spleenwort, which
in these sites may build up populations of thousands
of individuals. The plants usually grow in the shade of
scattered trees such as Red Maple, Black Walnut,
Eastern Red Cedar (Juniperus virginiana), Trembling
Aspen (Populus tremuloides), Black Cherry, Sassa-
fras (Sassafras albidum), and White Elm (U/mus
americana), but occasionally appear in grassy areas in
full sunlight. The plants are often concentrated in
thickets of Gray Dogwood (Cornus racemosa),
Common Juniper, or Rubus spp. Other associates in
these habitats include saplings of Pignut Hickory,
White Ash (Fraxinus americana), Scarlet Oak (Quer-
cus coccinea), Red Oak, and Black Oak, the shrubs
New Jersey Tea (Ceanothus americanus), Autumn
Olive (Elaeagnus umbellata), Chokecherry (Prunus
virginiana), Tatar Honeysuckle, and Rosa spp., and
the vines Virginia Creeper ( Parthenocissus quinquefo-
lia) and Riverbank Grape (Vitis riparia). Many of the
herb associates are of Eurasian origin. The clubmoss
Lycopodium digitatum (formerly treated as Lycopo-
dium complanatum var. flabelliforme) is also com-
monly present. Four species of grape ferns (Botry-
chium) are known to grow with the spleenworts, but
they do not form weedy populations.

A number of observers have commented on the
intrusive nature of the Ebony Spleenwort. In Ontario,
A. A. Reznicek (personal communication) reported
Ebony Spleenwort from rock outcrops, forested
dunes, and beach ridges, and pointed out that “Ebony
Spleenwort does not seem to care whether the rock is
acidic or basic and a wide variety of sandy soils also
seem suitable. It seems to be drought-tolerant and
happy in both sun and shade.” In Illinois Charles
Sheviak (personal communication) attributed its
dramatic increase in abundance to “the species coloni-
zation in old fields, abandoned pastures, and similar
disturbed sites.” The success of A. platyneuron in
becoming an aggressive and weedy plant cannot be
explained, however, entirely in terms of its coloniza-
tion of successional habitats, as evidenced by the non-
weedy nature of Botrychium species. Other character-
istics which might preadapt it for making an invasion
are best illustrated by comparing it with near relatives
such as A. montanum, A. resiliens, A. septentrionale,
and A. trichomanes which have never been reported
to be aggressive invaders.

Morphology of Ebony Spleenwort

As background we expand here the descriptions of
frond structure to include features not previously
emphasized by other authors. The horizontal rhizome
of the Ebony Spleenwort is very short so that the
leaves are crowded together and the plants appear

FIGURE 2. The second-growth old field habitat of Asplenium platyneuron in southeastern Michigan.

tufted. The leaves reach two extremes in morphology
— stiff, tall, erect, brittle, easily broken, fertile fronds
which tend to die back in the fall, and pliable, spread-
ing or prostrate, short, sterile fronds which persist
through the winter — but a spectrum of intermediates
occurs. Plants of dry upland habitats occasionally
have fertile fronds which resemble the sterile ones. On
a given plant the number of sterile leaves greatly
exceeds the number of fertile and intermediate leaves;
20 upland plants from Lyndon Township, Washtenaw
County, Michigan, gave an average of 16 (range 7-36)
fronds per plant, of which 5 were fertile, | was inter-
mediate, and 10 were sterile, and 20 lowland individu-
als from the same locality averaged 14 (range 6-34)
fronds per plant, of which 4 were fertile, | interme-
diate, and 9 sterile.

Starch storage in petiole bases is a phenomenon
which has not been described for this species and is
probably more widespread in the ferns than is realized
because it is best recognized in live plants and may not
be evident unless sections are made. Herbarium spec-
imens do not show the storage areas well because of
the drying and collapse of tissues.

Most fertile fronds die in the Fall while the sterile
fronds persist through the winter. All of the fronds
appear to remain alive at their bases for several years
(Figure 3). We discovered rhizomes estimated, from
the number of frond bases present, to be 5-7 years old
on which even most of the oldest frond bases were still
alive and contained masses of storage parenchyma.
The firm yellowish-white storage tissue contains
numerous small starch grains and is protected by the
dark occluded outer cortical and epidermal cells of the
petiole base. The extent of the storage on a given
petiole can be readily determined by pinching the
petiole base. The storage region feels hard and thick-
ened, while the dead petiolar area distal is dry and
collapses easily. Many petioles that are several years
old have disintegrated completely down to the point
where the storage base terminates.

Food storage in A. platyneuron is substantially fol-
iar rather than cauline (Figure 3), possibly partly asa
result of the size of the dictyostele in the true stem. The
sclerotized densely overlapping petiole bases proba-
bly also function in protecting the delicate rhizome
and bud tissues. The tissues of the storage petiole

160 THE CANADIAN FIELD-NATURALIST

eon /ETETATTT TULA TTT

Vol. 95

2

FIGURE 3. Starch storage in petiole bases of Asplenium platyneuron. Top left, longitudinal section of rhizome and petioles;
top right, cross section through rhizome and petioles; lower left, lateral view of intact rhizome and petioles; lower right,

ventral view of rhizome and petioles.

bases are also capable of dedifferentiation and growth
— petiole bases on damp sphagnum or sand produced
new plants.

In other fern genera with storage petiole bases, such
as Onoclea, Matteuccia, Athyrium, and Dryopteris,
there is a tendency toward abortion of the entire frond
except for the storage base. This tendency reaches its
best development in the Southern Fragile Fern (Cys-
fopteris protrusa) which has cataphyll-like storage
bases (Wagner et al. 1970). Other North American
species of Asplenium have storage areas which are
weakly or not developed.

The production of foliar buds is known in many
fern families and genera and has been discussed by
DeBenedictus (1969); who recognizes three types, all
of which occur in the Aspleniaceae: (1) buds on leaf
blades and margins, (2) axillary or rachis buds, and (3)
leaf tip proliferations. Faden (1973) showed that
rachis buds are most common among the gemmifer-
ous spleenworts of tropical East Africa. The ability of
the Ebony Spleenwort to form foliar proliferations

has long been known, having been reported from
plants cultivated in England in 1816 (Weatherby
1924). Plants exhibiting these proliferations were rec-
ognized taxonomically by Eaton (1879) as Asplenium
platyneuron var. proliferum, later reduced to the sta-
tus of form by Tanger (1933). Some writers have
regarded the proliferation process as the result of
abnormal conditions. Marshall (1923) reported find-
ing that 25 out of 75 plants growing ina basement had
buds, and then only on sterile fronds. Floyd (1924)
believed that the proliferations were a form of repro-
duction which resulted when the plants were under
stress.

In the populations of A. platyneuron we examined
from the Great Lakes region, we found proliferous
plants at practically every site. Most buds were but-
tonlike, pale-green, or whitish masses 1.0-1.5 mm in
diameter on the adaxial side of the lowest pinna (Fig-
ure 4). Rarely there was also a bud on the second
pinna. These buds were composed of a shoot apex, a
closely arched leaf primordium, sometimes accom-

1981

WAGNER AND JOHNSON: EBONY SPLEENWORT, GREAT LAKES AREA : 161

FIGURE 4. Buds at bases of basal pinnae of Asplenium platyneuron.

panied by a second, much smaller primordium, and
masses of narrow clathrate scales around the apex and
primordia. Roots are found on the more fully devel-
oped proliferations on which the leaf primordia have
expanded. Similar proliferations have been found in
the southern Appalachian populations of A. mona-
thes, especially those grown in culture (Wagner,
unpublished data).

Unless the buds have formed leaves they are diffi-
cult to detect, as they occur at the bases of the lowest
pinnae and usually on only a single frond per plant
which is buried in a mass of 10-20 other fronds.
Among 250 plants at three localities which were stu-
died intensively, proliferations were found on | out of
6 plants. The buds were found on both fertile and
sterile fronds, but were almost always borne singly on
one leaf of the plant.

Most of these buds probably die back with the blade
and upper portion of the stipe, but under certain
conditions they come in contact with the soil at the
base of the mother plant and produce new, separate
plants by decay of the connection between the bud and
the mother leaf. This accounts for the clusters of

plants which occur at certain sites, particularly in
shaded, moist, woodland habitats.

Mickel (1976) wrote that in Asplenium exiguum,
which normally grows in dry habitats, the frequency
of proliferation increases enormously if the plants are
grown under 100% humidity. To see if available mois-
ture also stimulated the formation of leaf buds in A.
platyneuron, we sampled three populations in Michi-
gan from dry, grassy hillsides and from nearby
shaded, moist bottoms at the edges of woods. Whole
plants were collected and brought back to be
inspected for buds. On the drier more open sites, only
4% (4/108) of the individuals bore proliferations,
while in the more moist, more shaded sites, 27%
(37/138) had them. Budding, while apparently a
genetic possibility for all Ebony Spleenwort plants,
appears to be influenced by the environment. We tried
unsuccessfully to force nonproliferous old fronds to
produce buds by laying them on wet sphagnum or wet
sand under high humidity. This suggests that bud
formation is initiated during the early ontogenetic
stages of the fronds in the spring and that by fall the
tissues have matured and will not dedifferentiate.

162 THE CANADIAN FIELD-NATURALIST

Occurrence of Incised Variations

Plants which have the pinnae of both the fertile and
sterile leaves deeply incised, giving the plants a ruffled
appearance (Figure 5), have been given the name
Asplenium platyneuron var. incisum. Taylor et al.
(1976), who reviewed the nomenclature and identifi-
cation of the infraspecific taxa of the Ebony Spleen-
wort, characterize var. incisum as having pinnae
which are “doubly serrate to deeply incised; all or
nearly all of the pinnae cut less than 4/5 of the way to
the midrib.”

The geographical and taxonomic significance of
this variety relative to typical A. p/atyneuron has been
variously interpreted. Some authors, such as Shaver
(1954), have considered it to be a coastal race of A.
platyneuron, while others, such as Correll (1956),
have considered it to be only a shade-induced envi-
ronmental form. Distribution maps (Shaver 1954;
Steyermark 1963; Taylor and Demaree 1979) show it
to be scattered throughout the range of A.
platyneuron.

a
Pps te
My ee

Vol. 95

An Ebony Spleenwort population in Lyndon
Township, Washtenaw County, Michigan, which
contained plants conforming to var. incisum, was stu-
died to determine whether this taxon has an environ-
mental or genetic basis. Thirteen samples were taken
from a 20 X 20 m area, each composed of the largest
frond from every plant within a plot of 2.3 m2.
Voucher specimens from this site are Wagner 79419
(MICH). The samples were uniformly spaced and
included plants both low and high on the hillside and
in both sun and shade. The collected fronds were
scored for six characters thought to be correlated with
the incised condition: greater frond length and width,
greater maximum pinna length and width, greater
number of pinnae, and non-confluent sori. In addi-
tion, pinnae from typical A. platyneuron and from
var. incisum were Cleared and stained for comparison
of their venation and soral characters.

Of 204 plants sampled, 27 were assignable to var.
incisum, 165 to the typical variety, and 12 were more
or less intermediate (Figure 6, c and e). These identifi-

FIGURE 5. Asplenium platyneuron var. incisum, Washtenaw County, Michigan, showing ruffled appearance caused by

overlapping pinna lobes.

1981

5mm +———

WAGNER AND JOHNSON: EBONY SPLEENWORT, GREAT LAKES AREA 163

FIGURE 6. Variation in pinna form in Asplenium platyneuron: a and b, var. platyneuron;f, g, and h, var. incisum, ¢, d,ande,

intermediate.

cations were based solely on pinna incision, as there
was no significant correlation found between the
other characters examined and the incised condition.
The plants of var. incisum were scattered at random,
making up 0-20% of the plants in each plot. There was
no obvious frequency gradient from plot to plot in the
sample area.

Extremely well-developed plants of var. incisum
differed from typical plants in several morphological
features having to do with the increase in lobing and
surface area of the pinnae (cf. Figure 6, a and b withf,
g, and h). Greater dissection of the pinnae is accom-
panied by an increase in the number of quaternary and
higher-order veins, an increase in the number of ulti-
mate veinlet endings in each vein complex, the occur-
rence of several sori per lobe on ultimate higher-order
veins, the change froma dichotomous branching plan
of the lateral veinlet system to a sympodial one, and
the overlapping of pinnule lobes. This last character is
visible on pressed specimens, although the ruffling of
the pinna lamina found in the living state in incisum is
not.

As a further investigation on the nature of var.
incisum, plants of it and of the typical variety were dug
up and grown indoors under uniform conditions for
several months. Both varieties retained their respec-
tive morphologies in the new leaves they produced
during this period.

Discussion

We have documented the range expansion of the
Ebony Spleenwort, but it is still not clear why this fern
did not undergo its spread before the 1960s, as dis-
turbed habitats and old farmlands and fields have
been available for over a century. Perhaps there has
been a much greater area of second growth available
than in the more distant past. Climatic warming
trends may have also played a role (cf. Britton 1977).
We may be witnessing an entirely statistical pheno-
menon of population growth, and the species may
have existed at a low level for a much longer period
and over a much wider range than we have thought.
As the populations built up, the spores produced
increased geometrically, and what was a trivial spore

164

shower in the Great Lakes region in 1900 is now a
massive fallout. The leading edge of the range is
clearly heading northward, and the advent of this
species in new areas of Canada, north of Lake Supe-
rior and Lake Huron, is expected.

In comparison with other native Asp/enium species,
A. platyneuron has a number of characteristics that
enable it to compete more aggressively with other
plants. Its ability to grow on the ground in addition to
rock surfaces allows it to invade areas where outcrops
are absent, as in much of southern Michigan and
Ontario. The tall, erect sporophylls enable the plant
not only to produce large numbers of spores but to
release them efficiently into the airstream. The condi-
tion of dimorphy is unusual in asplenioid ferns and its
presence in this species may be a strong factor in its
success (cf. Wagner and Wagner 1977, p. 260, Tables |
and 2). The starch storage in the petiole bases provides
the plant with a ready supply of food for quick devel-
opment in the spring and for the production of fronds
large enough to overtop competing vegetation. In this
respect Ebony Spleenwort is like species of a number
of common terrestrial fern genera such as Onoclea,
Dryopteris, and Athyrium. Colonization of available
habitats, particularly those in shaded, moist bottom-
lands, is facilitated by the production of new plants
from the foliar proliferations. Finally, the origins of
many populations are probably traceable to single
spores, as intragametophytic mating has been shown
to be possible in a large number of ferns (Klekowski
1971, 1972); this may explain why many of the reports
of Ebony Spleenwort in the 1950s and 1960s were only
of single plants. As shown by our studies in the George
Reserve, such plants are capable of giving rise to local
populations, even in forests.

As a result of examining plants which could be
referred’ to the infraspecific taxa forma proliferum
and variety incisum, we conclude that neither should
be treated as a true taxonomic variety (cf. Wagner
1960). By extrapolation of these studies to the other

THE CANADIAN FIELD-NATURALIST

t

Vol. 95

infraspecific taxa of A. platyneuron recognized by
Taylor et al. in 1976 (f. furcatum, var. platyneuron, f.
hortonae, var. bacculum-rubrum), we conclude that
none of them merit the designation variety, and ques-
tion the value of giving them taxonomic names at all.
Proliferous leaves have been found on plants “other-
wise referable to var. platyneuron, var. incisum, or
var. bacculum-rubrum” (Taylor et al. 1976), and our
observations suggest that bud formation is a normal
potentiality of the Ebony Spleenwort. The only way to
tella plant of forma proliferum is to find a bud on one
of its leaves, as usually all of the other leaves lack it, a
trivial basis for the recognition of a formal taxon.
Plants of the “variety” incisum occur throughout the
range of the Ebony Spleenwort at low frequencies and
grade into the typical “variety,” criteria for recogni-
tion of a weak form at best (cf. Wagner 1960).

Of the remaining four taxa, all but forma hortonae,
a rare bipinnate-leaved, entirely sterile form, have
been found in the Great Lakes area. The remaining
three are the typical “variety” (var. platyneuron),
forma furcatum, and “variety” bacculum-rubrum.
Forma furcatum is the name given to plants in which
the rachis forks repeatedly; such crested forms, caused
by the repeated abortions of the apical cell, are well
known in ferns, having been designated with formal
nomenclature for at least 11 North American taxa.
“Variety” bacculum-rubrum, characterized by Taylor
et al. (1976) as a southern “variety” with fertile fronds
up to 70 cm long, up to 70 pairs of pinnae per frond,
and with pinnae frequently serrate-incised and over
3.5 cm long, has been collected in Cass County, Mich-
igan (Wagner 73431, MICH). So far as we can deter-
mine, the taxon bacculum-rubrum embraces only the
largest plants from the southern part of the range of A.
platyneuron, and these, like “variety” incisum, occur
at low frequencies in collections and presumably in
populations; of 55 A. platyneuron specimens from the
coastal plain from Virginia to Texas at MICH, only 5
could be identified as var. bacculum-rubrum, while

TABLE !—Comparison of infraspecific taxa of Asplenium platyneuron

incisum bacculum-rubrum proliferum hortonae furcatum
Distribution Sporadic Clinical north to Sporadic Sporadic Sporadic
south
W. Great Lakes Rare Rare Common Unknown Unknown
incidence
Transition to Gradual Gradual Gradual Abrupt Gradual
typical form
Major character Incised Pinnae linear, Presence of Bipinnate Forked midrib
pinnae more than 3 cm long buds
Control Genetic or Modification Modification Genetic Genetic or

modification

modification

1981

most specimens were indistinguishable from typical
plants from New England and Ontario. There is no
real break in the variation pattern, and the characters
are primarily those of size alone. Accordingly, we
believe that if this taxon must be recognized taxonom-
ically, it should be as a form. When specimens are
discovered in the Great Lakes area that conform to
“bacculum-rubrum,” they do not constitute range
extensions, but only extreme variations of local
populations.

The characteristics of the described nontypical
infraspecific taxa of A. platyneuron are summarized
in Table |. In all cases we appear to be dealing with
potentialities—for forking fronds, incised pinnae,
giant fronds with long pinnae, and foliar
proliferation—which reflect different aspects of the
adaptations and genetic constitution of the Ebony
Spleenwort. Similar situations in the animal world,
such as the color phases of hawks, the budding of
hydras, and the regeneration of new tails by lizards,
have not been saddled with formal nomenclature, and
we see no reason why it is appropriate here. It should
be sufficient to recognize that the Ebony Spleenwort is
a dynamic and variable species, keeping in mind that
the recognition of these taxa was historically impor-
tant in pointing out its capabilities.

Acknowledgments

The information presented in this paper represents
the collaboration of many workers, especially H. E.
Ballard, Jr., J. M. Beitel, D. J. Bay, D. M. Britton
(Ontario), T. J. Carlson, A. J. Cooper, T. Crovello
(Indiana), T. F. Daniel, G. J. Gastony (Indiana), the
late D. J. Hagenah, Matt Heumann, J. Morton
(Ontario), J. H. Peck (lowa, Minnesota, and Wiscon-
sin), R. A. Pippen, A. A. Reznicek (Ontario), C. J.
Sheviak (Illinois), W. C. Taylor (Wisconsin), R. L.
Stuckey (Ohio), Olive Thompson (Wisconsin), F. S.
Wagner, and J. R. Wells. Those persons who supplied
data from outside Michigan are so indicated. Michael
K. Hansen aided us in the design of data collection,
and the authorities of Park Lyndon, of Washtenaw
County, Michigan, and the E. S. George Reserve of
The University of Michigan, kindly permitted us to
make collections. R. J. Ayotte contributed much of
the work on the incidence of proliferations.

Literature Cited

Adams, William. 1958. A preliminary list of Ohio pterido-
phytes. Distributed by the Ohio Flora Commiteee. 22 pp.

Billington, Cecil. 1952. Ferns of Michigan. Cranbrook
Institute of Science Bulletin 32. 240 pp.

Britton, Donald M. 1977. The Fern Woodsia obtusa
(Spreng.) Torrey in Ontario. Canadian Field-Naturalist
91: 84-85.

WAGNER AND JOHNSON: EBONY SPLEENWORT, GREAT LAKES AREA 165

Cooperrider, T. S. 1959. The ferns and other pteridophytes
of lowa. State University of lowa Studies in Natural His-
tory 20(1): 1-66.

Correll, D. S. 1956. Ferns and fern allies of Texas. Contri-
butions of the Texas Research Foundation, 2. Texas
Research Foundation, Renner, Texas. 188 pp.

Deam, Charles C. 1940. Flora of Indiana. Department of
Conservation, Division of Forestry, Indianapolis. 1236
Pp.

DeBenedictus, Virginia M. M. 1969. Apomixis in ferns.
Ph.D. thesis, University of Michigan, Ann Arbor. 193 pp.

Eaton, D. C. 1879. New or little-known ferns of the United
States. Bulletin of the Torrey Botanical Club 6: 306-307.

Faden, R. B. 1973. Some notes on the gemmiferous species
of Asplenium in tropical East Africa. American Fern
Journal 63: 85-90.

Floyd, F.G. 1924. Proliferation of Asplenium platyneu-
ron. American Fern Journal 14(1): 13-17.

Hagenah, Dale J. 1955. Notes on Michigan Pteridophytes.
I. New county records in Osmundaceae and Polypodia-
ceae. American Fern Journal 45: 65-80.

Hanes, Clarence R., and Florence N. Hanes. 1947. Flora of
Kalamazoo County, Michigan. Vascular Plants.
Anthoensen Press, Portland, Maine. 295 pp.

Klekowski, E. J., Jr. 1971. Ferns and genetics. BioScience
317-322.

Klekowski, E. J., Jr. 1972. Genetical features of ferns as
contrasted to seed plants. Annals of the Missouri Botani-
cal Garden 59: 138-151.

Marshall, M.A. 1923. Proliferous
American Fern Journal 13(1): 7-13.

Mickel, John T. 1976. Vegetative propagation in Asple-
nium exiguum. American Fern Journal 66: 81-82.

Mohlenbrock, Robert H. 1967. The illustrated flora of Illi-
nois: ferns. Southern Illinois University Press, Carbon-
dale. 191 pp.

Pepoon, H.S. 1927. Flora of the Chicago Region. Chicago
Academy of Sciences, Chicago. 554 pp.

Shaver, Jesse M. 1954. Ferns of Tennessee. George Pea-
body College for Teachers, Nashville, Tennessee. 502 pp.

Steyermark, Julian A. 1963. Flora of Missouri. lowa State
University Press, Ames, Iowa. 1728 pp.

Tanger, Louise F. A. 1933. Fernsand fernallies of Lancas-
ter County, Pennsylvania, 1931 and 1932. American Fern
Journal 23(1): 13-18.

Taylor, W. Carl, and Delzie Demaree. 1979. Annotated list
of the ferns and fern allies of Arkansas. Rhodora 81:
503-548.

Taylor, W. Carl, Robert H. Mohlenbrock, and Fredda J.
Burton. 1976. Variation in North American Asplenium
platyneuron. American Fern Journal 66(2): 63-68.

Tryon, R. M., Jr. 1954. The ferns and fern allies of Minne-
sota. University of Minnesota Press, Minneapolis. 166 pp.

Tryon, R. M., N.C. Fassett, D. W. Dunlop, and M. E.
Diemer. 1953. The ferns and fern allies of Wisconsin.
Department of Botany, University of Wisconsin, Madi-
son. 158 pp.

Wagner, W.H., Jr. 1960. Evergreen grapeferns and the
meanings of infraspecific categories as used in North
American pteridophytes. American Fern Journal 50(1):
32-45.

Ebony Spleenwort.

166 THE CANADIAN FIELD-NATURALIST Vol. 95

Wagner, W. H., Jr., Donald R. Farrar, and Bruce W. McAI- Weatherby, C. A. 1924. The earliest record of proliferous
pin. 1970. Pteridology of the Highlands Biological Sta- Ebony Spleenwort. American Fern Journal 14(3): 95-96.
tion area, southern Applachians. Journal of the Elisha
Mitchell Scientific Society 86(1): 1-27.

Wagner, W. H., Jr.,and F. S. Wagner. 1977. Fertile—sterile Received 15 July 1980
leaf dimorphy in ferns. Gardens’ Bulletin 30: 251-267. Accepted 6 October 1980

Distributional History of Juncus compressus (Juncaceae)
in North America

RONALD L. STUCKEY
Department of Botany, College of Biological Sciences, The Ohio State University, Columbus, Ohio 43210

Stuckey, Ronald L. 1981. Distributional history of Juncus compressus (Juncaceae) in North America. Canadian Field-
Naturalist 95(2): 167-171.

Juncus compressus Jacq., native to Europe and portions of western Asia, is a poorly known member of the North American
wetland flora. Although sometimes misidentified as J. gerardii, verified records of its occurrence date from 1881 to the early
1900s. These early records are froma coastal locality in Maine and in Newfoundland, along the St. Lawrence River in Quebec,
the Finger Lakes region of New York, and Salt Lake City, Utah. Within the past 30 years, J. compressus has become more
widespread, and records have been obtained from in and near Toronto, Ontario, from southern Manitoba, and from five
states, namely Colorado, Minnesota, Montana, Wisconsin, and Wyoming, all west of the Great Lakes. The plants are
perennial, and they invade and inhabit salt marshes, man-created saline habitats, ditches along highways and railroads,
limestone quarries and calcareous shores, and banks of canals and roadsides. Its recent discovery from six widely spaced
localities in Wyoming suggests that the species may be established in other areas of western United States, even though it is not
included in the most recent floristic treatments of that region.

Key Words: Juncaceae, Juncus compresus, J. gerardii, rush, wetland flora, non-indigenous, introduced, distributional

history, halophyte, North America.

During a study of the migration and establishment
of Juncus gerardii (Juncaceae) in the interior of North
America (Stuckey 1980), it became apparent that a
morphologically similar species, J. compressus (Fig-
uré 1), has sometimes been confused with and misi-
dentified as J. gerardii. This paper presents documen-
tation for the known distributional history of J.
compressus, Flattened Rush, an apparently non-
indigenous member of the North American wetland
flora. Native throughout most of Europe and portions
of western Asia, J. compressus was first reported for
North America by Bartlett (1906). This record was
based on specimens obtained in 1904 by Arthur S.
Pease (6312, 6312A, 6313, GH), who discovered the
plants as abundant on the Plains of Abraham near the
city of Quebec. Bartlett pointed out the morphologi-
cal differences between J. compressus and J. gerardit,
and one or more of these differences (Table 1) have
since been used as key characters in several manuals,
for example, Fernald (1950), Gleason (1952), Her-
mann (1975), Scoggan (1978), and Wiegand and
Eames (1926). Like J. gerardii, J. compressus invades
and inhabits salt marshes, man-created saline habi-
tats, and ditches along highways and railroads, lime-
stone quarries and calcareous shores, and banks of
canals and roadsides. Marie-Victorin (1929) believed
that the plants may have been introduced at an early
date with military forage.

Examination of herbarium specimens now reveals
that J. compressus was present in North America
prior to Bartlett’s report as verified from two speci-
mens. One specimen was obtained on 30 July 1881 by

Agnes Saunders (DAO) from a “bog” near shore at
Great Chebeag, Casco Bay, Maine. This specimen,
mounted on the same sheet with a collection of J.
gerardii that apparently was also obtained at the same
locality, represents a record that she would have
secured while ona vacation trip, rather than one from
her home area near London, Ontario, where most of
her plants were collected (Dore 1970). Another speci-
men from Murray Bay, Quebec, obtained on 20
August 1902 by W. W. Eggleston (2974, GH, NY),
represents the voucher for the notation by Robinson
and Fernald (1909), who considered the species as
“possibly naturalized from Eu[rope].” The report of J.
compressus from Coney Island, New York, by Smith
(1945) is based ona plant of J. gerardii (CU) collected
in 1879 by an unidentified collector.

Elsewhere in the eastern United States, J. compres-
sus has been reported from the Finger Lakes region of
New York. Wiegand and Eames (1926) considered the
species “introduced with sand and other material
brought to a glass factory formerly existing near the
salt works” at Ithaca, Tompkins County. Wiegand
and Eames considered its arrival shortly before 1895,
when the plants were first detected. The species had
not been reported earlier from the area (Dudley 1886).

The following records have been verified as J. compressus
from Tompkins County: marsh near D. L. & W. coal dump,
Ithaca, 22 July 1895, K. M. Wiegand s.n. (CU); marsh W of
Glass Works near D. L. & W. dump, 22 July 1895, K. M.
Wiegand s.n. (CU); at edge of marsh in soft mucky soil, near
Ithaca Salt Works, Ithaca, 7 July 1913, E. L. Palmer 269
(CU, GH, TRT); brackish pond near salt works, Ithaca, 21
July 1916, E. P. Metcalf6/39 (CU); salt works, flats, Ithaca,

167

168

[}

FIGURE |. Juncus compressus, showing upper portion of
plant taken from specimen obtained at Oshkosh,
Winnebago County, Wisconsin, 11 July 1974 (Har-
riman 10,186, OS).

30 July 1919, K. M. Wiegand 11731 (CU); E bank of inlet,
Ithaca, 31 July 1919, K. M. Wiegand 11732 (CU, MO).

Wiegand and Eames (1926) also reported the spe-
cies about the railroad station at Freeville, but their
specimen (//733, CU) is J. gerardii. The record of J.
compressus from nearby Onondaga County is based
on plants (CU) obtained in 1912 froma salt marsh at
Syracuse (Smith 1945). The species was also listed as a
member of the flora of Onondaga County (Faust
1961; Bye and Oettinger 1969) and the Cayuga Quad-
rangle (Clausen 1949).

The spread of J. compressus has been more exten-
sive in Canada than in the United States. Following its
early discovery at Murray Bay (La Malbaie) in 1902
and near the city of Quebec in 1904, the species has
been located at several sites along the St. Lawrence
River.

The following additional specimens from Quebec
arranged chronologically have been examined: dry soil,

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE !|—Morphological differences between Juncus com-
pressus and J. gerardii based on an examination of plants by
the author and statements by Fernald (1950), Gleason
(1952), Hermann (1975), Scoggan (1978), and Wiegand and

Eames (1926)

Stamen length

Anther length

Perianth color

Style length

Capsule length

Capsule shape

Primary
involucral
bract

J. compressus

About half as long
as the perianth
segments

About as long or
twice as long as
the filaments

Light brown with
greenish center
and membrana-
ceous margin

Shorter than the
ovary

About one and
one-half times
as long as the
perianth

Subglobose or
globose-
obovoid

Usually longer
than the
inflorescence

J. gerardii

About equal in
length to the
perianth
segments

About three to four
times as long as
the filaments

Dark brown to
blackish with
green central
stripe

Equal to or longer
than the ovary

Shorter than or
barely exceeding
the perianth

Subellipsoid or
ellipsoid-ovoid

Usually shorter
than the
inflorescence

Heights of Abraham, 5 August 1910, C. S. Williamson 1112
(NY); berges ducanal de Chambly, Ile Ste. Thérése, St. Jean
County, 14 juillet 1932, M.-Victorin and R.-Germain 49134
(GH, TRT); narrow channel of Richelieu River, Chambly to
St. Jean, Chambly County, 13 September 1932, M.-Victorin
and K. M. Wiegand s.n.(CU, MO, NY); abondant au revers
du remblai du canal, Ile Ste. Thérése, prés de St. Jean, St.
Jean County, 20 septembre 1932, M.-Victorin and R.-
Germain 44667 (DAO, GH); introduit le long du canal en
gazon abondant, Ile Ste. Thérése, St. Jean County, 25 aoat
1933, M. Raymond 3498 (TRT); trés rare récolté prés du
pont qui mene a I’ile, Ile Ste. Thérése, 28 aoait 1933, M.
Raymond 126731 (CAN); dans une prairie saumatre au bord
de la mer, Les Emoulements, Charlevoix County, 16 juillet
1935, M.-Victorin, R.-Germain, and R. Meilleur 43615 (CU,
DAO); sur la gréve, Pointe-au-Pic, Charlevoix County, 9
aout 1937, C. Morin 897 (DAO); berges du canal Chambly
St. Jean, 7 aott 1943, M.-Victorin, R.-Germain, E. Rouleau,
and A. Blain 2158 (DAO, GH, MICH); sommet de la gréve,
St. Joachim, Cap Tourmente, 25 aoat 1950, Y. Desmarais
1237 (CAN, DAO); talus du cété de St. Jean, Ile Ste.
Thérése, St. Jean County, 2 aotit 1953, L. Cing- Mars and M.
Raymond s.n. (DAO); dans une prairie humide 4 mi
[0.93 km] au sud du Cap, St. Joachim, Montmorency
County, 6 juillet 1960, G. Lemieux 67084 (DAO); anse-aux-
foins, dans un petit misseau, St. Fulgence, cté Chicoutimi, 18
juillet 1973, J. Cayouette 73-516 (TRT); open alkaline road-
sides of highway, N side of Hwy 20, 2 mi [3.7 km] E of

1981

Pointe-Claire, just E of Boul des Sources, Dorval, Montreal
County, 10 June 1976, P. M. Catling, A. A. Reznicek, S. M.
McKay, and K. M. Lindsay 81 (CAN, DAO, TRT); salt
marsh, near Gaspé, S side of Gaspé Bay, Gaspé County, 10
August 1978, P. M. Catling and F. H. Catling s.n. (RT).

In Ontario, J. compressus was apparently first
noted by Fassett (1933) based ona specimen obtained
in 1931 from a small stream near a highway along the
Mississippi River near Ottawa, Carleton County
(Fassett 13280, CAN, GH). Since then the species has
been obtained from several localities in Ontario par-
ticularly along railroads and in saline wetlands in
areas near Ottawa, Toronto, Hamilton, and as far
north as Cochrane.

The following additional specimens from Ontario have
been identified, arranged by county: Carleton County: in
shallow soil over shale, Rockliffe Park, along Ottawa river
near boathouse below “lookout,” 4 September 1947, J. A.
Calder 1641 (DAO); a small clump in alluvial, perhaps
saline, pasture, at Mississippi River (about 5 mi[9.3 km] SE
of Arnprior, 27 mi [50 km] W of Ottawa), 19 June 1959,
W. G. Dore and W. J. Cody 17426 (DAO, TRT).[Cochrane
District]: le long de la voie ferré, Cochrane, 20 August 1946,
E. Lepage s.n. (DAO). Halton County: lining ditch in a
distinctive dark-green band in Nelson Quarry, near Burling-
ton, 16 July 1975, G. W. Dore 25444 (DAO, TRT). Ontario
County: floodplains and flanking slopes, from confluence of
Little Rouge and Rouge Rivers S to Lake Ontario, 4 July
1973, J. L. Riley s.n. (TRT); roadside, along Highway 69
near Rathburn, Mara Township, !5 July 1979, P. M. Catling
s.n. (OS, TRT). Peel County: one clone seen, disturbed clay
soil on floodplain, Credit River just N of Erindale College
campus, 16 June 1977, A. A. Reznicek and S. A. White s.n.
(TRT). Simcoe County: locally dense, wettest section of
roadside ditch, Conc. XIII, lot 1, Vespra Township, 22 July
1972, R. S. W. Bobbette 2851 (TRT); moist ground by end of
road along drainage canal, West Gwillimbury Township,
4.5 mi. [8.3 km] N NE of Bradford, 15 July 1973, R. S. W.
Bobbette 3830 (DAO); solitary dense patch, NW side of Hwy
27,5 mi{9.3 km] S of Bondhead, Conc. I, Lot 23, Tecumseth
Township, 29 June 1975, A. A. Reznicek 14541 (TRT).
Wentworth County: marsh, banks of old creek, Royal
Botanical Gardens, Hamilton, 17 July 1954, Tamsalu 1419
(HAM); in pools, edge of Queen Elizabeth Highway, 2 km
NW of Fifty Road near Stoney Creek, Saltfleet Township, 6
October, P. M. Catling and K. L. Catling s.n. (OS, TRT).
York County: moist sandy soil in open location, Toronto
Island, 9 August 1972, P. M. Catling and S. M. Mc Kay s.n.
(DAO); on landfill ca. 5 years old, wet ground at foot of
Leslie Street, Toronto, September 1972, P. M. Catling s.n.
(CAN, TRT); additional specimens from the Leslie Street
locality have been obtained by Catling and McKay on! July
1973 (TRT), 10 November 1974 (TRT), July 1975 (TRT),
October 1975 (TRT), | August 1976 (DAO, TRT), 8 August
1976 (CAN, TRT); sandy lake shore, Hanlan’s Point,
Toronto Island, 25 July 1974, D. Hoy, H. Saifi, and N.
Purcell s.n. (TRT); in snow dumping area opposite Don
Valley Brickyard, Don Valley, Toronto, 19 October 1974,
P. M. Catling and S. M. McKay s.n. (DAO); in wet depres-
sion receiving runoff from rock salt storage depot opposite
Don Valley Brickyard, Don Valley, Toronto, 20 October

STUCKEY: DISTRIBUTION OF JUNCUS COMPRESSUS

169

1974, P. M. Catling and S. M. McKay s.n. (CAN, TRT);
open alkaline soil by railway yard at Bathurst Street under-
pass, just N of Front Street, just E of Old Fort York,
Toronto, 21 August 1977, P. M. Catlingand K. L. McIntosh
s.n. (TRT); alkaline ditch receiving salty runoff, edges of
Don Valley Parkway, near Bloor Street, Toronto, 13
October 1979, P. M. Catling s.n. (OS, TRT).

In the maritime provinces, J. compressus was first
discovered in Newfoundland, in boggy meadows and
wet grassy slopes at Birchy Cove, Curling, 7 Sep-
tember 1910 by Fernald and Wiegand (2986, CAN,
CU, GH, NY). Two records, one from Inverness
County and one from Richmond County, have been
cited for Nova Scotia (Erskine 1951), but specimens
verifying these localities have not been found. The
earliest report for Nova Scotia (Roland 1945) is /.
gerardii based on a specimen obtained in 1930 from
Guysborough by J. Rousseau (35358, CAN). Fur-
thermore, the report of J. compressus for Prince
Edward Island by Erskine (1960) is also /. gerardii,
having been based on specimens collected in 1912
from Charlottetown by Fernald, Long, and St. John
(7140, CAN, GH, MICH, MO, NY).

West of the Great Lakes region, J. compressus was
first obtained from Manitoba in 1950 at Brandon
(Stevenson 1957), and subsequently at Duck Bay,
Lake Winnipegosis, and northeast of Duck Mountain
(Scoggan 1957). Later, Love and Bernard (1959)
noted it from a roadside south of Kleefeld in the
Otterburne area.

Specimens seen from Manitoba are wet flats W of Exhibi-
tion grounds, Brandon, 16 July 1950, G. Stevenson 201
(DAO); locally common around Brandon, 3 August 1954, G.
Stevenson 882 (CAN); Fosse, coin nord-ouest du terrain
d’Exposition, Brandon, 3 juillet 1959, B. Boivin, G. Steven-
son, and E. Laishley 13162 (DAO, GH, NY, TRT).

In the United States west of the Great Lakes region,
J. compressus was first obtained in 1909 from Salt
Lake City, Utah, based on a specimen of M. E. Jones
(s.n., NY) which had been identified as J. gerardii. In
the /ntermountain Flora, Cronquist (1977) did not
include J. compressus.

Verified records of J. compressus are known from five
other states, all since 1950 as follows: Montana: Beaverhead
County: marshy roadside ditch, 9 mi[16.7 km] S of Wisdom,
1955 (Hermann, 1975). Minnesota: Kittson County: road-
side at Caribou, 1962 (J. W. Moore 26083, TRT, mapped
erroneously as J. gerardii by McGregor and Barkley (1977).
Colorado: Larimer County: open wet soil adjoining bait
minnow ponds, Colorado State University foothills campus,
Fort Collins, 1967; sedge meadow south of Spring Canyon,
5 mi [9.3 km] SW of Fort Collins, 1971 (Hermann 1975).
Wisconsin: Winnebago County: along grassy edge beneath
U.S. 41 overpass at state route 21, on the S side of route 21
only, and onto the lawn at the Howard Johnson’s, Oshkosh,
1974 (Harriman and Redmond 1976). Wyoming: Carbon
County: around pool in floodplain of Medicine Bow River,
Y% mi [0.93 km] N of Medicine Bow, 1973 (S. Stephens

170 THE CANADIAN FIELD-NATURALIST

FiGuRE 2. The distribution of Juncus compressus in North America based on herbarium specimens examined and reliable
literature citations. Dots represent localities where plants have been recorded before about 1950, and squares represent
localities where plants have been obtained since about 1950.

70504, KANU, NY); same locality, 31 July 1979, B. Nelson
and R. Hartman 4215 (RM); abundant in wet sandy soil
around pool, Seminoe Reservoir, ca. 16 air km SSE of
Seminoe Dam (T24N R84W S14, 24), ca. 1950 m, 11 July
1979, R. Hartman 992] (RM). Converse County: North
Platte River, ca. 10.4 air km E of Glenrock (T33N R74W S39),
ca. 1524 m,9 August 1979, K. Dueholm 8816 (RM). Johnson
County: ca. 27.2 air km SE of Buffalo (T48N R81 W S24), ca.
1370 m, 29 June 1979, K. Dueholm 7447 (RM). Natrona
County: North Platte River, ca. 1.6 air km E of Pathfinder
Reservoir Dam(T29N R83W S19), ca. 1740 m, 11 July 1979,
R. Hartman 9958, 9959 (RM); oil field, ca. 6.8 air km N of
Waltman (T36N R86W S6 and R87W S11), ca. 1900 m, 12
July 1979, R. Hartman 10037 (RM).

The recent discovery of J. compressus from six
widely spaced localities in Wyoming suggests that the
species may be established in other areas of western
United States even though it is not included in two of
the most recent floristic treatments of the region
(Cronquist 1977; Dorn 1977).

The known distribution of J. compressus in North
America is mapped based on the above cited records
(Figure 2). The map shows the range of the species
before about 1950 to be confined to Newfoundland,
Nova Scotia, Quebec, eastern Ontario, Maine, New
York, and Utah. Since then J. compressus has been
obtained from additional localities in Quebec, and has

appeared in the Toronto region in Ontario, southern
Manitoba, Colorado, Minnesota, Montana, Wiscon-
sin,and Wyoming. The plants are perennial, and once
they invade an area, they persist, as has been verified
by the recent records from the St. Lawrence River
valley and the Toronto area. Current records, how-
ever, have not been seen from the early reported local-
ities in Maine, New York, Utah, and Newfoundland.

Acknowledgments

My thanks are extended to Paul Catling, who pro-
vided the records and identifications of the specimens
at TRT; to Ronald Hartman, who provided the
records and identifications of the specimens at RM; to
James S. Pringle, who provided data on the single
specimen at HAM; and toA. E. Spreitzer, for prepar-
ing the photograph. My thanks are also extended to
the curators of the herbaria who made specimens
available. Those herbaria in which I have examined
specimens are CAN, CU, DAO, GH, MICH, MO,,.
NY, OS.

Literature Cited
Bartlett, H. H. 1906. Juncus compressus in the province of
Quebec. Rhodora 8: 233.

1981

Bye, Robert A., and Frederick W. Oettinger. 1969. Vascu-
lar flora of Onondaga County, New York: an annotated
list. Department of Forest Botany and Pathology, State
University College of Forestry, Syracuse University, Syra-
cuse. 248 pp.

Clausen, Robert T. 1949. Checklist of the vascular plants of
the Cayuga Quadrangle 42°-43° N., 76°-77° W. Cornell
University Agricultural Experiment Station Memoir 291.
87 pp.

Cronquist, Arthur. 1977. Juncaceae, pp. 47-67. /n Inter-
mountain flora: vascular plants of the Intermountain
West, U.S.A. By Arthur Cronquist, Arthur H. Holmgren,
Noel H. Holmgren, James L. Reveal, and Patricia K.
Holmgren. Vol. Six. The Monocotyledons. Colombia
University Press, New York. 584 pp.

Dore, William G. 1970. The Saunders collection. Green-
house-Garden-Grass 9: 11-20.

Dorn, Robert D. 1977. Manual of the vascular plants of
Wyoming. Garland Publishing Company, New York. 2
Volumes. 1498 pp.

Dudley, William R. 1886. The Cayuga flora. Part I: a cata-
logue of the phaenogamia growing without cultivation in
the Cayuga Lake basin. Bulletin Cornell University 2:
I-XXX, 1-132+ additions and corrections, index to
orders and genera, I-v.

Erskine, D. S. 1951. Species newly or rarely reported from
Nova Scotia and Cape Breton Island. Rhodora 53:
264-271.

Erskine, David S. 1960. The plants of Prince Edward
Island. Plant Research Institute, Research Branch, Can-
ada Department of Agricultural Publication 1088. 270 pp.

Fassett, Norman C. 1933. Notes from the herbarium of the
University of Wisconsin-X. Rhodora 35: 387-391.

Faust, Mildred E. 1961. Checklist of the vascular plants of
Onondaga County, New York. Bulletin of the Syracuse
Museum of Natural Science No. 9. 83 pp.

Fernald, Merritt Lyndon. 1950. Gray’s Manual of botany.
Eighth Edition. American Book Company, Incorporated.
Ixiv + 1632 pp. (Reprinted by D. Van Nostrand Com-
pany, New York. 1970).

Gleason, Henry A. 1952. The new Britton and Brown illus-
trated flora of the northeastern United States and adjacent
Canada. Volume I. New York Botanical Garden, New
York. 482 pp.

STUCKEY: DISTRIBUTION OF JUNCUS COMPRESSUS 171

Harriman, Neil A., and Arrell Redmond. 1976. Somatic
chromosome numbers for some North American species
of Juncus L. Rhodora 78: 727-738.

Hermann, Frederick J. 1975. Manual of the rushes (Juncus
spp.) of the Rocky Mountains and Colorado Basin. Uni-
ted States Department of Agriculture Forest Service Gen-
eral Technical Report RM-18. 107 pp.

Love, Doris, and Jean-Paul Bernard. 1959. Flora and vege-
tation of the Otterburne area, Manitoba, Canada. Svensk
Botanisk Tidskrift 53: 335-461.

Marie-Victorin, Frére. 1929. Les Luiluflores du Québec
[Liliacees, Pontederiacees, Iridacees, Joncacees]. Contri-
butions du Laboratoire de Botanique de |’Univerisé de
Montréal 14: I-1 92.

McGregor, R. L.,and T. M. Barkley (£ditors). 1977. Atlas
of the Flora of the Great Plains. lowa State University
Press, Ames. 600 pp.

Robinson, B. L.,and M. R. Fernald. 1909. Emendations of

the seventh edition of Gray’s Manual, — I. Rhodora 11:
33-61.
Roland, A. E. 1945[1947]. The flora of Nova Scotia. Pro-

ceedings and Transactions of the Nova Scotian Institute of
Science 21: 95-642. (Reprinted Truro Printing and Pub-
lishing Company, Truro. 552 pp., 127 figures. 1948.)

Scoggan, H. J. 1957. Flora of Manitoba. National Muse-
um of Canada Bulletin 140. 619 pp.

Scoggan, H. J. 1978. The flora of Canada. Part 2: Pterido-
phyta, Gymnospermae, Monocotyledoneae. National
Museum of Canada, Ottawa. 545 pp.

Smith, Stanley Jay. 1945. Contributions to the flora of cen-
tral New York-I. New York State Museum Bulletin. 338.
74 pp.

Stevenson, George A. 1957. Additions to the flora of Mani-
toba. Blue Jay 15: 74.

Stuckey, Ronald L. 1980. The migration and establishment
of Juncus gerardii (Juncaceae) in the interior of North
America. Sida 8(4): 334-347.

Wiegand, Karl M.,and Arthur J. Eames. 1926. The flora of
the Cayuga Lake basin, New York. Cornell University
Agricultural Experiment Station Memoir 92. 491 pp.

Received 9 July 1980
Accepted 14 October 1980

Le régime alimentaire du Coyote (Canis latrans)
et du chien errant (C. familiaris) dans le sud du Québec

JEAN-MARIE BERGERON! et PIERRE DEMERS?

'Département de biologie, Faculté des sciences, Université de Sherbrooke, Sherbrooke, Québec JIK 2R1
Service de l Aménagement et de Exploitation de la Faune, Ministére du Loisir, de la Chasse et de la Péche, 85 rue Holmes,
Sherbrooke, Québec JIE 1S1

Bergeron, Jean-Marie, et Pierre Demers. 1981. Le régime alimentaire du Coyote (Canis /atrans) et du chien errant (C.
familiaris) dans le sud du Québec. Canadian Field-Naturalist 95(2): 172-177.

L’analyse de contenus stomacaux de 84 Coyotes (Canis latrans) et 58 chiens errants (C. familiaris) échantillonnés dans un
milieu agro-forestier nous a permis d’identifier les éléments importants de leur alimentation. Beaucoup d’indices portent a
croire que les Coyotes de la région se nourissent de charogne puisqu’une grande partie des contenus digestifs est composée de
restes d’animaux domestiques comme les bovins (Bos taurus) et le porc (Sus scrofa). Par contre, les mammiféres sauvages
constituent un élément moins important du régime alimentaire de ces prédateurs. Pour le Coyote, viennent par ordre
d'importance, le Li¢vre d’ Amérique (Lepus americanus), les petits mammifeéres, le Cerf de Virginie (Odocoileus virginianus),
la Marmotte Commune (Marmota monax) et le Raton Laveur ( Procyon /otor). Cependant, des restes de cerfs ont été analysés
dans plus du tiers des chiens examinés en hiver, ce qui en fait un animal trés nuisible face a la survie hivernale des cerfs.
Certains éléments de végétation et les oiseaux completent le régime alimentaire de ces prédateurs.

Mots clés: Coyote, chien errant, prédateur, régime alimentaire.

Bergeron, Jean-Marie, et Pierre Demers. 1981. Le régime alimentaire du Coyote (Canis /latrans) et du chien errant (C.
familiaris) dans le sud du Québec. Canadian Field-Naturalist 95(2): 000-000.

We identified the components of their diet from the stomach analyses of 84 Coyotes (Canis latrans) and 58 domestic dogs (C.
familiaris) trapped in the forested agricultural land of southeastern Quebec. Domestic animal carrion, especially cattle (Bos
taurus) and hogs (Sus scrofa), was very important for Coyotes. Wild mammals were not an important food source for Coyotes
but Snowshoe Hares (Lepus americanus), small mammals, White-tailed deer (Odocoileus virginianus), Woodchucks (Mar-
mota monax), and Raccoons (Procyon /otor) regularly occurred in that order in their diet. Deer remains, however, were
identified in more than a third of the dogs examined in winter. Some avian and vegetal components were classified as
secondary food sources for both species.

Key Words: Coyote, wild dog, predators, diet.

Depuis la derniére décennie, les populations de
Coyotes (Canis /atrans) et de chiens errants (C. famili-
aris) ont tellement augmente dans les Cantons de l'Est
au Québec qu’ils entrent maintenant en conflit avec
plusieurs secteurs d’activités humaines. Par exemple,
dans les milieux agro-forestiers, leur présence souleve
une polémique sérieuse et met en évidence l’opportu-
nisme de ces prédateurs. En effet, du printemps a
lautomne, ces prédateurs s’attaquent fréquemment
aux animaux domestiques, jeunes et adultes, tandis
qu’en hiver, ils doivent se nourrir d’animaux sauvages.
Cette situation mest pas particuliére a notre régioneta
été vérifiée a maints endroits en Amérique du Nord ou
les pratiques agricoles de petite et moyenne envergure
sont effectuées dans un encadrement forestier.

Depuis 1935, de nombreuses analyses du régime
alimentaire de ces prédateurs ont été effectuées aux
Etats-Unis. Cependant, peu d’études canadiennes
autres que celles de Murie (1935), Pastuck (1974) et
Nellis et Keith (1976) ont été entreprises dans ce
domaine. Au Québec, les seules études que nous con-

naissons sont celles de Jean-Paul Blais,Ministere du
Tourisme, de la Chasse et de la Peche (MTCP), Sha-
winigan, rapport d’étape, 1974 et de Messier et
Barrette (1979). Dans la premieére, l’examen macros-
copique des contenus stomacaux de 81 Coyotes cap-
turés dans la région de Shawinigan a permis de con-
stater dans 70% des cas un bol alimentaire constitué de
diverses espéces de mammiféres dont les plus impor-
tantes sont représentées par les bovins domestiques
(13%) et le Liévre d’Amérique (Lepus americanus)
(8%). Dans la seconde étude, les auteurs ont étudié le
régime alimentaire de Coyotes provenant de régions
forestiéres en se basant sur les crottins et les restes
laissés le long de leurs pistes sur la neige. Ils ont trouvé
que le Cerf de Virginie (Odocoileus virginianus) et le
Liévre d’Amérique constituaient les proies les plus
recherchées avec des fréquences d’abondance respec-
tives de 59 et 38%. Ces auteurs ont également noté
d’importantes variations de ces deux espéces dans le
régime alimentaire au cours de l’année et selon la taille
des groupes de Coyotes.

2

1981

De janvier 1975 a juin 1977, nous avons recueilli les
carcasses de Coyotes et de chiens errants provenant de

trappeurs des Cantons de !’Est afin d’analyser leur”

régime alimentaire. Nous voulions par cette compa-
raison vérifier les changements saisonniers dans I’ali-
mentation des prédateurs et analyser importance rel-
ative des sources de nourriture de provenance
domestique ou sauvage de facon a jeter les bases d’un
aménagement de ces prédateurs par les organismes
concernés.

La région étudiée

Les Cantons de l'Est (Figure |) sont limités au sud
par la frontiére canado—américaine (45°00’N) et au
nord par le piémont appalachien (46°30’N). L’enca-
drement est et ouest se fait a 70°30’ et 73°00’ de
longitude ouest. Le territoire couvre environ 20 000 km?
et comprend un relief peu accidenté dans |’ensemble,
ce qui permet un bon drainage de sols sédimentaires a
fort potentiel agricole et de podzols offrant de bonnes
aptitudes forestiéres.

Les versants fertiles et bien drainés favorisent la
croissance de certains feuillus comme !’Erable a Sucre
(Acer saccharum), VErable Rouge (A. rubrum), le
Merisier (Betula alleghaniensis) et de peuplements
conifériens comme !|’Epinette Blanche ( Picea glauca),

Zone agricole

fees: Zone moyennement
agricole

Zone forestiere

A Chiens errants

@ Coyotes

BERGERON ET DEMERS: COYOTE ET CHIEN ERRANT REGIME ALIMENTAIRE, QUEBEC

3)

le Sapin Baumier (Abies balsamea) et la Pruche
(Tsuga canadensis).

Le développement du réseau routier a permis l’ex-
pansion de l’agriculture dans la portion occidentale.
La section centrale posséde une vocation agricole et
forestiére, tandis que l’est demeure un important sec-
teur forestier. La région présente donc une mosaique
agro-foresti¢re dont lensemble des composantes
passe par une gamme complexe d’exploitations par-
tagées entre l’agriculture et la foresterie.

Matériel et méthodes

L’analyse des contenus stomacaux de Coyotes et de
chiens s’est faite selon la technique décrite par
Korschgen (1969). Les carcasses nous parvenaient (1)
de trappeurs travaillant a leur propre compte dans la
région; (2) de trappeurs professionnels employes par
le Ministére de l’ Agriculture, des Pécheries et de l’Al-
imentation du Québec (MAPAQ), dont la fonction est
de trapper les animaux ayant causé des pertes de
bestiaux; et (3) d’Agents de Conservation exergant un
controle hivernal et printannier prés des quartiers
d’hiver de Cerfs de Virginie. Les carcasses étaient
acheminées au laboratoire d’écologie animale de
l'Université de Sherbrooke, munies d’une étiquette
indiquant la date et le lieu de capture, le mode de

FIGURE |: Répartition des captures de Coyotes et de chiens errants en fonction de l'utilisation du sol des Cantons de l'Est
(Districts électoraux. Ministére des Terres et Foréts, 1973).

174 THE CANADIAN FIELD-NATURALIST

trappage, l’appat utilisé, le nom du trappeur et l’habi-
tat dans lequel l’animal était capturé. Celles-ci étaient
ensuite 4gées et séparées en Coyotes, chiens ou
hybrides selon les techniques de Lawrence et Bossert
(1967).

Les analyses stomacales ont été effectuées princi-
palement a l’aide d’exemples cuticulaires de poils de
mammiféres. Nous avons préparé une collection de
400 photographies de référence regroupant 39 espéces
de mammiferes et d’oiseaux sauvages ou domestiques.
Les échantillons de poils prélevés dans les contenus
stomacaux étaient montés sur lames pour analyse
microscopique, a la fagon d’Adorjan et Kolenosky
(1969) et comparés a ceux de la collection. Les autres
éléments du contenu stomacal étaient identifiés et
classés en groupes de fagon a exprimer leur fréquence.
Nous n’avons pas effectué de mesures volumétriques
de composantes étant donné que les structures avaient
s€journé plus de 5 h dans l’estomac (Gier 1968).

Résultats et discussion

La distribution géographique des Coyotes et des
chiens errants qui nous ont servi pour l’étude apparait
a la figure |. L’effort de cueillette des carcasses n’est
pas réparti uniformément dans les Cantons de l'Est.
Les petites concentrations de captures peuvent s’ex-
pliquer par emplacement des quartiers d’hiver de
Cerfs de Virginie, la provenance des plaintes de pertes
d’animaux de ferme, le lieu de résidence des trappeurs
et leur habileté respective a capturer Coyotes et
chiens. La figure montre que la section sud-est forme
un bloc forestier contrastant avec la région sud-ouest
de la plaine du Saint-Laurent dont la vocation est
fonciérement agricole. La moitié de la portion cen-
trale présente des terres non défrichées (MACQ 1965).
Ainsi, la majeure partie de notre récolte provient de
secteurs forestiers et partiellement agricoles.

La distribution de notre échantillonnage dans le
temps est directement fonction de notre approvision-
nement en carcasses. Les trappeurs professionnels
engagés pour le contréle des prédateurs des animaux
de ferme ont fourni au printempset a l’été 46% (38/84)
des carcasses. La participation des autres trappeurs
constitue 39% (33/84) de l’échantillonnage dont les
animaux ont surtout été piégés l’automne et Phiver. Le
reste de la cueillette provient du contréle des préda-
teurs a proximité des quartiers d’hiver de Cerfs de
Virginie par les Agents de Conservation. Ce controle
nous a également procuré 55% (32/58) des chiens de
notre échantillon, alors que la différence, en majorité
des captures d’automne, nous est parvenue de trap-
peurs répartis dans la région.

Bien que la récolte soit assez bien répartie sur 12
mois, il faut tenir compte de la compartimentation
saisonniere faite en fonction des objectifs du piégeage
ou du contréle de prédateurs. Cet état de fait rend la

Vol. 95

comparaison des régimes alimentaires difficile a effec-
tuer puisque beaucoup de chiens ont été pi¢gés prés
des aires hivernales de Cerfs de Virginie tandis que les
Coyotes proviennent de différentes sources.

Les mammiferes représentent sans contredit l’éle-
ment le plus important du régime alimentaire du
Coyote. Nous avons trouvé des restes de mammifeéres
domestiques dans 30% des Coyotes et 28% des chiens
analysés (Tableau |). La présence d’animaux domes-
tiques dans les contenus digestifs du Coyote est par-
ticulierement importante en hiver et au printemps,
probablement a cause de la libre disposition d’ani-
maux morts a proximité des fermes. Les bovins (Bos
taurus) constituent lélément le plus fréquemment
rencontré chez ces prédateurs, totalisant 18% chez le
Coyote et 24% chez le chien. Nos résultats se compar-
ent assez bien avec ceux du Québec (Jean-Paul Blais,
MTCP, rapport d’étape, 1974), du Manitoba (Pas-
tuck 1974) et des Etats-Unis (Ozaga 1963; Mathwig
1973; Andrews et Boggess 1978; Berg et Chesness
1978). L’élevage du porc (Sus scrofa) dans nos régions
se fait dans des conditions de rétention trés contrdlées.
Le fort pourcentage d’indices de porcs dans les conte-
nus digestifs peut s’expliquer soit par Pabondance des
carcasses pres des fermes, soit par l’utilisation des
carcasses aux dépotoirs municipaux.

La faible abondance (2%) du mouton (Ovis aries)
dans les prédateurs échantillonnés est un peu surpre-
nante. Malgré l’attention particuliére apportée a la
détection d’ indices (laine), nous n’avons pu confirmer
par l’analyse des contenus digestifs limpact réel
qu’exerce le Coyote sur les moutons de la région. Bien
que nos résultats se comparent avec ceux de Pastuck
(1974, 3%), Andrews et Boggess (1978, 5%), et Berg et
Chesness (1978, 2%), nous ne pouvons nier l’impact
qu’ont les Coyotes et les chiens errants sur le cheptel
ovin de la région. Les nombreuses plaintes (P.
Demers, chargé du dossier des prédateurs au MLCP
(Ministére du Loisir, de la Chasse et de la Péche) de
Sherbrooke) enregistrant la perte d’animaux ou du
moins la présence de Coyotes et de chiens errants a
proximité des paturages temoigne du probleme de
certains éleveurs. L’impact des chiens errants sur les
animaux de la ferme est indirect, c’est-a-dire qu'il
s’agit le plus souvent de harcélement que de prédation
réelle.

Mathwig (1973) affirme que 20% des pertes en
bétail étaient causées par des chiens et Pastuck (1974)
précise que les moutons étaient la plupart du temps
tués par les chiens. La présence plus fréquente d’indi-
ces de bovins chez les chiens plut6t que chez les
Coyotes s’explique par la promiscuité avec le bétail
ainsi que la proximité des dépotoirs de ferme et des
carcasses d’animaux facilement accessibles.

Parmi les mammiféres sauvages faisant partie du
régime alimentaire des Coyotes nous n’avons pas

1981 BERGERON ET DEMERS: COYOTE ET CHIEN ERRANT REGIME ALIMENTAIRE, QUEBEC 175

TABLEAU 1—Fréquence (pourcentage) des principaux éléments rencontrés dans les analyses stomacales de 84 Coyotes et 58

chiens errants récoltés en différentes saisons

Printemps Eté Automne Hiver Total
Coyotes, Chiens, Coyotes, Chiens, Coyotes, Chiens Coyotes, Chiens, Coyotes, Chiens,
Eléments 22 12 17 3 23 19 22 24 84 58
Mammiféres domestiques 40.9 50.0 5.9 0 26.1 15.8 40.9 29.2 29.8 Dales)
Bovin domestique
(Bos taurus) 13.6 41.7 — — 26.1 10.5 Dies 29.2 17.6 24.1
Pore domestique (Sus scrofa) DD) — 5.9 — 4.3 DS 13.6 = 11.9 lod
Autres! 9.1 16.7 — 4.5 _ 3.6 3.4
Mammifeéres sauvages 31.8 25.0 29.4 33.3 43.5 10.5 40.9 25.0 36.9 20.7
Liévre d’ Amérique
(Lepus americanus) 13.6 — 5.9 4.5 a 6.0 --
Marmotte Commune
(Marmota monax) 9.1 4.3 3.6 —
Cerf de Virginie
(Odocoileus virginianus) — 8.3 — 33.3 4.3 10.5 13.6 20.8 4.8 5.5)
Raton Laveur (Procyon lotor) 9.1 — 2.4 —
Porc-épic (Erethizon
dorsatum) — 16.7 — — 4.3 — 4.5 4.2 2.4 Sy
Campagnol des Champs
(Microtus pennsylvanicus) 4.5 _ 58) — 4.3 — 4.5 — 4.8 _—
Souris Sylvestre (Peromyscus
maniculatus) 4.5 — 5.9 aaa 8.7 — 4.5 — 6.0 —
Autres _— _ 11.8 — 17.4 — 4.5 _— 8.3 _
Mammnifeéres indéterminés 54.5 66.7 70.6 33.3 34.8 15.8 36.4 Bye) 47.6 36.2
Oiseaux 9.1 8.3 5.9 — 4.3 5.3 Oe — 10.7 3.4
Poulet domestique (Gallus
gallus) 91 _ 52) — 4.3 3)3) 18.2 — 9.5 1.7
Autres _— 8.3 4.5 — 12 Ila
Végétaux 86.4 83.3 76.5 ‘100.0 95.7 78.9 V2 70.8 83.3 77.6
Pommier nain (Malus pumila) 4.5 13.0 — Ded 4.2 10.7 1.7
Graminées cultivées PORT 58.3 11.8 33.3 17.4 26.3 13.6 37.5 16.7 37/8)
Débris végétaux divers V3 50.0 64.7. ‘100.0 95.7 Wel) 63.6 62.5 76.2 65.5
Litiére forestiére 50.0 8.3 35.3 — 43.5 31.6 Del) 8.3 38.1 S25)
Autres éléments? 40.9 100.0 29.4 66.7 17.4 36.8 54.5 50.0 3507 56.9
Fragments osseux 31.8 50.0 11.8 — 13.0 31.6 22) 33.3 20.2 34.5
Autres structures animales Delf 50.0 11.8 _— 13.0 10.5 13.6 8.3 15.5 17.2
Nourriture commerciale pour
chiens — 25.0 — 66.7 8.3 — 12.1
Déchets domestiques — 50.0 11.8 _ _— 15.8 9.1 313},33 4.8 29.3
Matiére indéterminée 4.5 8.3 10.5 Deg) 8.3 8.3 8.6

'Le mouton (Ovis aries), la chévre (Capra hircus), et le cheval (Equus caballus).

2Regroupement des éléments importants; les fragments osseux de méme que les autres structures animales servaient a
l'identification des proies; les déchets domestiques incluaient souvent des restes de table et des éléments non-comestibles. La
matiére indéterminée provenait d’animaux ou de déchets domestiques.

remarqué de prédominance d’une espéce en particu-
lier. Le Liévre d’Amérique n’a été identifié que chez
6% des Coyotes. Des restes de Cerfs de Virginie, de
Marmottes Communes (Marmota monax) et de Ra-
tons Laveurs (Procyon lotor) apparaissent chacun

dans moins de 5% des Coyotes. Représentant un faible
pourcentage chez le Coyote, le Cerf de Virginie est
apparu chez 16% des chiens analysés. En se basant sur
les résultats obtenus, il ne semble pas que les Coyotes
aient un effet sérieux sur les populations de cerfs de la

176 THE CANADIAN FIELD-NATURALIST

région tout au long de l’hiver. De plus, les premiéres
captures pres des quartiers d’hiver ne se font qu’a
partir de février (Laurent Cloutier, Agent de Conser-
vation, communication personnelle, 1979). I] faut se
rappeler qu’a cette date, 20% des Cerfs peuvent périr
dans un hiver moyenet que cette quantité peut s’élever
a 40% au cours d’un hiver difficile (Potvin et Huot
1975):

Par contre, l’impact des chiens errants sur les Cerfs
de Virginie en période de confinement est trés impor-
tant. A la fin de l’hiver les conditions de neige et de
disette alimentaire rendent pénibles les déplacements
des cerfs, en particulier ceux des femelles gravides. La
libre circulation de groupes de chiens parmi les cerfs
occasionne des poursuites et des dépenses d’énergie a
la suite desquelles, les victimes survivent difficilement.
L’analyse des contenus stomacaux de ces chiens a
révélé la présence d’indices frais chez la plupart d’entre
eux, ce que nous n’avons pas observe chez le Coyote.
De leur cété, Perry et Giles (1971), Scott et Causey
(1973), Denney (1974), Lachance (1975) et Thériault
(1978) ont tous noté les effets négatifs de la présence de
chiens dans les quartiers d’hiver des cerfs. De plus,
maints témoins d’agression par des chiens errants sur
des cerfs nous ont décrit les poursuites et les pertes
infligées aux cheptels locaux. Cependant, il faut
retenir que le pi¢geage des chiens s’est fait a Pintérieur
des aires hivernales tandis que celui des Coyotes s’est
effectué en périphérie, ce qui peut expliquer partielle-
ment la différence des résultats.

Les petits mammiféres dans leur ensemble ont été
identifiés dans plus de 19% des Coyotes examinés, ce
qui représente un élément assez stable de leur alimen-
tation tout au long de l'année. Contrairement a d’au-
tres études qui ne leur donnaient quwune valeur sai-
sonniére, nos résultats indiquent que le Campagnol
des Champs (Microtus pennsylvanicus) et la Souris
Sylvestre (Peromyscus maniculatus) sont des
éléments stables de leur régime alimentaire, alors
qu’apparaissent plus sporadiquement d’autres espéces
de campagnols et de souris sauteuses.

La plupart des descriptions de contenus digestifs
montrent que les oiseaux ne sont pas importants dans
lalimentation du Coyote. Des restes de poulet domes-
tique (Gallus gallus) abandonnés dans des dépotoirs
ou autour des fermes sont utilisés, particulierement
Vhiver, par 11% des Coyotes analysés. Des conclu-
sions semblables ont d’ailleurs été suggérées par
Murie (1935), Fichter et al. (1955), Gier (1968), Tie-
meier (1975), Nellis et Keith (1976). Des indices de
végétation ont été analysés dans plus de 83% des
Coyotes et 78% des chiens examinés. I] s’agit le plus
souvent de débris végétaux comme des aiguilles de
coniféres, des feuilles d’herbacées, d’écorce et de
ramilles de plantes ligneuses, ce qui peut étre une
cons€quence directe de la méthode de piégeage uti-

Vol. 95

lisée. Leur présence est importante chez les Coyotes
capturés au printemps et a l’automne tandis qu'elle est
plus uniforme tout au long de année pour le chien.
Gier et al. (1978) et Niebauer et Rongstad (1977) ont
décrit les qualités purgatives, vermicides, ou vitami-
niques de certaines plantes. La présence de vergers
abandonnés et de pommiers sauvages procure aux
Coyotes de la région une source alimentaire surtout
utilisée 4 ’'automne et a l’hiver. Certaines substances
plus ou moins comestibles ont été identifiées dans les
analyses stomacales de 36% des Coyotes et 57% des
chiens examinés. Parmi les déchets domestiques
prélevés, nous avons trouvé du papier d’emballage,
des pelures de fruits et de l¢gumes, des restes de tables,
des tissus et des objets métalliques ou de plastique.

Malgré le maximum d’objectivité accordé a lana-
lyse des résultats, nous demeurons conscients des
limites de l’interprétation d’une partie de ceux-ci. La
récolte de carcasses dépendait des sites de piégeage des
trappeurs travaillant a leur solde, ainsi que de l’en-
droit ot s’effectuaient les contrdles de prédateurs a
proximité d’aires hivernales de Cerfs de Virginie et de
fermes d’élevage. De plus, il est possible que dans
certains cas, l'utilisation de carcasses d’animaux en
guise d’appats ait pt influencer les résultats. Notons
cependant que plusieurs trappeurs utilisaient des
appats odoriférants et que d’autres nous ont menti-
onné la présence d’animaux déja morts sur le site de
trappage. Ces éléments, lorsque retrouvés dans les
contenus digestifs, étaient enregistrés mais omis de
Yanalyse.

Il faut également tenir compte du fait qu’une bonne
partie des carcasses provenant des programmes de
répression de prédateurs a puamener un biais dans les
résultats a cause d’animaux “problémes.” Nous pen-
sons tout de suite a la sur-représentation des restes
d’animaux domestiques provenant de Coyotes trap-
pés pres des fermes et des dépotoirs et de la sur-
représentation des restes de Cerfs de Virginie trouvés
chez les chiens capturés ou abattus a proximité des
quartiers d’hiver de cerfs.

Nous ne pouvons cependant nier deux grandes
caractéristiques se dégageant de ce travail et venant
appuyer les résultats de plusieurs auteurs du Canada
et du nord-est des Etats-Unis dont les travaux se sont
faits dans un contexte similaire au notre. II s’agit de
lopportunisme du Coyote face aux sources de nourri-
ture qui l’entourent et de l’impact des chiens errants
sur les Cerfs de Virginie en période de confinement.

Remerciements

Cette étude a été réalisable grace a un contrat de
recherches et a des bourses d’études accordées par le
Service Canadien de la Faune. Nous désirons remer-
cier M. Serge Gonthier, chef de Service a l Aménage-
ment et a l’Exploitation de la Faune du Ministere du

1981

Loisir, de la Chasse et de la Péche, pour avoir coor-
donné la récolte de spécimens provenant du pro-
gramme de contr6le des prédateurs.

Références

Adorjan, A. S.,et G. B. Kolenosky. 1969. A manual for the
identification of hairs of selected Ontario mammals.
Research Report No. 90, Department of Lands and
Forest, Ont. 64 pp.

Andrews, R.D., et E.K. Boggess. 1978. Ecology of
Coyotes in Iowa. Dans Coyotes: biology, behavior and
management. Edité par M. Bekoff. Academic Press Inc.,
New York. pp. 249-265.

Berg, W. E.et R. A. Chesness. 1978. Ecology of Coyotes in
northern Minnesota. Dans Coyotes: biology, behavior
and management. Edité par M. Bekoff. Academic Press
Inc., New York. pp. 229-247.

Denney, R.N. 1974. The impact of uncontrolled dogs in
wildlife and livestock. Transactions of North American
Wildlife Natural Research Conference 39: 257-291.

Fichter, E., G. Schildmanet J. H.Sather. 1955. Some feed-
ing patterns of Coyotes in Nebraska. Ecological Mono-
graphs 25: 1-37.

Gier, H. T. 1968. Coyotes in Kansas. Kansas Agricultural
Experimental Station Bulletin 393. 118 pp.

Gier, H. T.,S. M. Kruckenberg et R. J. Marler. 1978. Par-
asites and disease of Coyotes. Dans Coyotes: biology,
behavior and management. Edité par M. Bekoff. Aca-
demic Press Inc., New York. pp. 37-71.

Korschgen, L. J. 1969. Procedures for food habits ana-
lyses. Dans Wildlife management techniques. Rédigé par
R. J. Giles. The Wildlife Society, Washington, D.C. pp.
233-250.

Lachance, J. R. 1975. Les chiens errants tuent 100 chev-
reuils dans le Parc de la Gatineau. Québec, Chasse et
Péche 4: 38-89.

Lawrence, B., et W. H. Bossert. 1967. Multiple character
analysis of Canis lupus, latrans, and familiaris with a
discussion of the relationships of Canis niger. American
Zoologist 7: 223-232.

Mathwig, H. J. 1973. Food and population characteristics
of lowa Coyotes. lowa State Journal Research 47:
167-189.

Messier, F., et C. Barrette. 1979. Etude de la prédation du
Cerf de Virginie par le Coyote dans le ravage d’Armstrong,

BERGERON ET DEMERS: COYOTE ET CHIEN ERRANT REGIME ALIMENTAIRE, QUEBEC 177

Beauce sud. Rapport remis au Ministere du Loisir, de la
Chasse et de la Péche, Direction de la recherche faunique,
RRF 40, mai 1979. 167 pp.

Ministére de l’Agriculture et de la Colonisation du Qué-
bec. 1965. L’agriculture du Québec jusqu’en 1961. Cata-
logue de cartes. 102 pp.

Murie, O. H. 1935. Food habits of the Coyote in Jackson
Hole, Wyoming. United States Department of Agriculture
Circular 362: 1-24.

Nellis, C. H., et L. B. Keith. 1976. Population dynamics of
Coyotes in Central Alberta, 1964-1968. Journal of Wild-
life Management 40: 389-399.

Niebauer, T. J.,et O. J. Rongstad. 1977. Coyote food hab-
its in northwestern Wisconsin. Dans Proceedings of the
1975 Predator Symposium. Edité par R. L. Phillips et C.
Jonkel. Montana Forest Conservation Experimental Sta-
tion, University of Montana. pp. 237-251.

Ozaga, J.J. 1963. An ecological study of the Coyote on
Beaver Island, Lake Michigan. M.Sc. thesis, Michigan
State University. 122 pp.

Pastuck, R. D. 1974. Some aspects of the ecology of the
Coyote (Canis latrans Say) in southwestern Manitoba.
M.Sc. thesis, University of Manitoba, Winnipeg. 159 pp.

Perry, M.C. et R.H. Giles, Jr. 1971. Studies of deer-
related dog activity in Virginia. Proceedings Annual Con-
ference, Southeast Association Game Fish Commission
1970, 24: 64-73.

Potvin, F., et J. Huot. 1975. Mortalité d’hiver du Cerf de
Virginie dans le ravage Pohénégamook, 1971-1975. Rap-
port d’étape, Service de la recherche biologique, Ministere
du Tourisme, de la Chasse et de la Péche. 21 pp.

Scott, M. D. et K. Causey. 1973. Ecology of feral dogs in
Alabama. Journal of Wildlife Management 37: 253-265.

Thériault, R. 1978. The impact of uncontrolled domestic
dogs on Gatineau Park Resources. Section de Conserva-
tion des Ressources, Service des Parcs, Commission de la
Capitale nationale. 122 pp.

Tiemeier, J. W. 1975. Winter foods of Kansas Coyotes.
Transactions of the Kansas Academy of Science 58:
196-207.

Recu le 1 décembre 1979
Accepté le 20 octobre 1980

Variation in Frequencies of Pelvic Phenotypes of the Brook
Stickleback, Culaea inconstans, in Redwater Drainage, Alberta!

JAMES D. REIST

Department of Zoology, The University of Alberta, Edmonton, Alberta T6G 2E9
Present address: Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6

Reist, James D. 1981. Variation in frequencies of pelvic phenotypes of the Brook Stickleback, Cu/aea inconstans, in
Redwater drainage, Alberta. Canadian Field-Naturalist 95(2): 178-182.

In Brook Stickleback, Cu/laea inconstans, geographical variation in relative frequencies of pelvic phenotypes was noted
especially between lotic and lentic environments. In the primary study area, Wakomao Lake incentral Alberta, no association
of any pelvic phenotype with grossly defined habitats was evident. The relative frequencies of all pelvic phenotypes were stable
from 1971 to 1977; however, the frequency of the form without the pelvis and spines decreased over winter. Frequencies of the
subsequent generation returned to prewinter values, thus suggesting some cyclical seasonal variation. Causative agents for the
maintenance of the frequencies were sought. The observed frequencies are not due to sampling biases, rather the most likely
cause is some selective agent(s). Mass die-off of sticklebacks during midsummer oxygen depletion was not differential. Other
studies implicate predation by a variety of piscivores as a selective agent.

Key Words: Culaea inconstans, Brook Stickleback; pelvic variation, spatial and temporal frequency.

Brook Stickleback, Culaea inconstans, show clines
in lengths of pelvic and dorsal spines (Nelson 1969).
Also, a high proportion of individuals lack the pelvic
skeleton and associated pelvic spines in numerous
populations in midcentral Alberta and Saskatchewan
(Nelson 1969, 1977; Nelson and Atton 1971; Nelson
and Paetz 1974).

Four pelvic states are observed in variable popula-
tions of Culaea: complete presence — with(s), com-
plete absence —without(s), intermediate with a single
spine — spined, and intermediate with only an inter-
nal bony plate —spineless (see figures in Nelson 1969,
1977). This variation has at least a partial genetic basis
(Nelson 1977).

Extent and range of variation of the phenomenon
has been described (Nelson 1977; Nelson and Atton
1971), but adequate, in-depth examination is lacking
for the variation over space and time, both within
single sites and between sites in close spatial proxim-
ity. In this study I have described geographical, habi-
tat, and temporal variation of frequencies of pelvic
phenotypes in the Redwater drainage of Alberta, as
background to investigations into the evolutionary
significance of the phenomenon (Reist 1980 a, b).

Additionally, some aspects of the explanation for
such phenotypic variation are investigated. Such
explanations may be grouped into three classes: (1)
variation might be due to methodological problems
—sampling biases or errors of assignment to pheno-

1Based upona thesis submitted to the University of Alberta
in partial fulfillment of the requirements for the degree of
Master of Science.

178

typic classes; (2) historical canalization of phenotypic
frequencies might result in the observed variation,
that is, the present situation might be due to stochastic
influences such as drift or immigration, or it may bea
transient situation proceeding to future fixation of
one phenotype; (3) finally, variation might be due to
selective force(s) qualitatively or quantitatively vary-
ing Over space and time, resulting in a balanced sys-
tem. Evidence is presented which seems to refute the
first two of these explanations. One aspect of the third
explanation is tested in additional studies (Reist
1980a, b) which investigated predation as a potential
selective force in more detail.

Study Area

The Redwater river system (Figure 1), a collection
of small lakes and rivers about 80 km north of
Edmonton, Alberta, is a tributary to the North Sas-
katchewan River. The lakes tend to be shallow and
eutrophic, and the creeks are narrow and meandering.

Land surrounding the bodies of water is flat, and
extensive marshes, of either aquatic macrophytes or
drowned bogs of Black Spruce (Picea mariana) — wil-
low (Salix sp.), are typical. Extensive beaver activity
and large numbers of deadfall blockages on the creeks
likely restrict fish movements, especially during win-
ter or the drier periods of the year.

The major study area, Wakomao Lake (54°09’N,
113°32’W), is a small (about 325 ha) lake most proxi-
mate to the North Saskatchewan River. Maximum
depth is 2.5 m. Most of the shoreline is marshy with
extensive beds of emergent macrophytes, but the
windward southeast corner is sandy beach. Large

1981

113 30

2 18 16 8
S {|

REIST: PELVIC FREQUENCIES OF STICKLEBACKS

179

FiGurRE |. The study area, Redwater River system, Alberta. Numbers refer to collection sites (see Table 1 for site names); and
( 4), areas sampled but where no sticklebacks were found. The major study area was Wakomao Lake (Number 10).

amounts of submerged and floating vegetation are
present throughout the lake during the summer.
Wakomao Lake is subject to light recreational pres-
sure — angling and boating. It was also the subject of
an abortive fisheries management program in 1955.
Four species of fish were found in Wakomao Lake
during the study: Culaea inconstans, Pimephales
promelas (Fathead Minnow), Catostomus commer-
soni(White Sucker), and Esox /ucius (Northern Pike).

Materials and Methods

All collections of fishes were made in water depths
of 0-1.5 m by the most convenient method, usually
dip net or bag-seine. Sampling was conducted at each
location by methods which would lessen potential
biased collection of phenotypes. The samples were
immediately preserved in 10% formalin.

Pelvic phenotypes are easily scored by running a
sharp probe over the belly, although some individuals
of the spineless phenotype may incorrectly be classed
as withouts; however, the pelvic remnant of spineless
individuals is usually large enough to permit unam-
biguous identification. Pelvic development occurs
between 13 and 2! mm standard length (SL, the dis-
tance from tip of the snout to end of the fleshy lobe of

the caudal peduncle). The first structures to appear
are the pelvic spines (Nelsonand Atton 1971). All fish
less than 21 mm SL were examined under a low-
power dissecting microscope. Fish less than 15 mm
SL were considered unassignable and left out of statis-
tical considerations.

To investigate variation in phenotypic frequencies,
I designated eight habitats in Wakomao Lake using
predominant vegetation type, time of year, and spatial
position. Association of any phenotype with these was
tested using multiple samples obtained from each.

Samples taken at the appropriate times were com-
pared to establish change of frequency with season,
age, and longer periods of time. In the 1977 cohort,
age-classes were defined using standard length as the
criterion.

Collection froma 5 X 10 m quadrat of the beach of
Wakomao Lake permitted investigation of collection
biases. This area was sequentially collected by bag-
seine, dip net, and electrofisher (Smith-Root Inc.
Type VII). Baited minnow traps set in other areas of
the lake constituted a fourth collection method.

Collection at a single site (No. 8, Figure 1) in early
May fortuitously resulted in a sample of “naturally”
dead fish and a sample of “live” fish thus permitting

180 THE CANADIAN FIELD-NATURALIST

the test of a selective death hypothesis. Cause of mor-
tality was not identified, but the fish were subject to
extreme crowding in a small creek so that oxygen
depletion may have been a factor.

Many sites were visited several times; thus, the
sample sizes represent composite samples. All multi-
ple samples were tested for heterogeneity (Sokal and
Rohlf 1969) and then combined, because none were
heterogeneous. Statistical comparison of samples was
by row by column chi-square tests.

Results

Moderate amounts of variation in phenotypic fre-
quencies are evident among the 15 sites (Table 1). The
greatest deviations from expected frequencies are sites
6,8, 13, 15,and 16. Frequencies in remaining areas are
relatively consistent at 70% with, 5% intermediates
(spined and spineless), and 25% without.

The average frequencies for collections from creeks
(sites 4,5, 6, 8,9, 12, 14, 15, 17, 18) are 72.8% with,
5.0% intermediates, and 22.2% withouts. Average fre-
quencies for collections from lake environments (sites
7, 10, 11, 13, 16) are 70.6% with, 4.4% intermediate,
and 25.0% without. The differences are statistically
significant (P< 0.05).

Association of any phenotype witha particular hab-
itat is not evident (Table 2, P > 0.05). No sticklebacks
were visually sighted or taken in surface minnow traps
set for 270 trap-hours. The gross nature of habitat
definition and collection does not preclude microhab-
itat association.

Vol. 95

Collections from Wakomao Lake in 1971 show
phenotypic frequencies (71.0% with, 4.4% interme-
diate, and 24.6% without, N = 284) similar to those of
1977.

A significant decrease in frequency of intermediate
and without phenotypes with a concomitant increase
in frequency of withs was observed over winter (Table
3, P<0.025). Relative frequencies tended to be re-
stored to prewinter values in the next sample from
1977 which contained scoreable young from that year
(Table 3). Within the year-class trom 1977, there was a
tendency for increase in frequency of withs and inter-
mediates and a decrease in without frequencies,
although this is not significant (Table 4, P > 0.05).

On the collected quadrat, no method of collection
preferentially selected for any particular phenotype
(Table 5, P = 0.8). Minnow trap samples from other
areas of the lake also did not preferentially select fora
phenotype (P = 0.4). The phenotypic frequency in the
sample of fish presumed to have died of oxygen star-
vation did not differ from that of the sample still living
when collected (Table 6, P = 0.6), implying nonselec-
tive death of pelvic phenotypes.

Discussion

Sites that contribute to significant geographical
variation in frequencies differ in physicochemical and
biotic features not quantified in this study. For exam-
ple, site 6 is a spring-fed creek with water temperatures
consistently below those from other sites, and site 8
contains little or no aquatic vegetation. These findings

TABLE 1—Frequencies of pelvic phenotypes of Brook Stickleback in the Redwater River system. Map numbers refer to
locations on Figure |. Samples with less than 50 individuals are not included. The intermediate phenotypes were combined for

calculating the chi-square statistic (P< 0.05)

Number of phenotypes (%)

Map Number
number Site name With Spined Spineless Without N of samples
4 Outflow from Halfmoon Lake 93(70.5) 4(3.0) 4(3.0) 31(23.5) 132 l
5 Redwater River 402(69.8) 16(2.8) 18(3.1) 140(24.3) 576 4
6 Fairydell Creek 72(68.6) 4(3.8) 6(5.5) 23(21.9) 105 l
7 Overflow from unnamed lake 51(79.7) 1(1.6) 1(1.6) 11(17.2) 64 l
8 Beaver Pond Creek 617(76.5) 13(1.6) 21(2.6) 155(19.2) 806 I
9 Creek south of Wakomao
Lake 190(70.1) 9(3.3) 8(2.9) 64(23.6) 271 l
10 Wakomao Lake 1526(70.6) 73(3.4) 31(1.4) 531(24.6) 2161 15
11 Slough south of Wakomao
Lake 89(70.1) 1(0.8) 4(3.1) 33(26.0) 127 l
12 Halfway Creek 105/72.4) 4(2.7) 3(2.1) 33(22.7) 145 |
13 Halfway Lake 71(63.4) 1(0.9) 1(0.9) 39(34.8) 112 I
14 Wakomao Creek 322(70.9) 9(2.0) 10(2.1) 106(23.7) 447 3
15 Wakomao Creek 247(73.3) 1(0.3) 6(1.8) 83(24.6) 337 3
16 West Bridges Lake 205(71.2) 4(1.4) 4(1.4) 75(26.0) 288 3
17 Bridges Creek 205(72.4) 9(3.2) 7(2.5) 62(21.9) 283 2
18 Duggans Creek 226(74.1) 11(3.6) 7(2.3) 61(20.0) 305 y,

1981

REIST: PELVIC FREQUENCIES OF STICKLEBACKS

TABLE 2—Phenotypic frequency of Brook Stickleback by habitat within Wakomao Lake

Number of phenotypes (%)

Habitat type With Spined
1. Burreed-7) pha shoreline 111(65.3) 7(4.1)
2. Scirpus shoreline 432(74.6) 19(3.3)
3. Myriophyllum-Lemna: washed on

shore submerged 325(69.1) 18(3.8)
4. Myriophyllum: rooted on shore

submerged 207(70.2) 8(2.7)
5. Open sand beach less than 5% cover

vegetation, Culaea here May-

June only 236(68.2) 10(2.9)
6. Submerged algal mats (Cladophora

sp. — May-June only) 83(67.5) 6(4.9)
7. Offshore rooted Myriophyllum—

Potamogeton 132(74.1) 5(2.8)
8. Open, unvegetated water’ 0(00.0) 0(0.0)
Pooled 1526(70.6) 73(3.4)

“Not included in computation of chi-square statistic.

agree with a general correlation of variation in pheno-
typic frequencies with characteristics of the water
body. Nelson and Atton (1971) suggested that with-
outs tend to occur in lakes lacking outlets and those
subjected to management practices.

Nelson (1969) suggested lack of choice of prey items
available to a predator, thus inevitable predation des-
pite deterrents such as spines may make it favorable
for the loss of minimally useful structures such as the
pelvic spines. The result would be a more streamlined

TABLE 3—Change in phenotypic frequency of Brook
Stickleback overwinter (1976 juveniles to 1977 adults) in
Wakomao Lake

Number of Phenotypes (%)

Time With Intermediate Without N
June 1976 495(68.6) 70(9.8) 161(22.2) 726
May 1977 144(77.1) 7(3.7) 36(19.1) 188
July 1977 563(72.5) 43(5.5) 170(21.9) 776

TABLE 4—Change in phenotypic frequency of Brook
Stickleback with age in Wakamao Lake (four collections,
one each from July, August, September, October 1977)

Standard length Number of phenotypes (%)

(= age) group With Intermediate Without N
15-29.9 mm 315(75.0) 17(4.1) 88(20.9) 420
30-39.9 mm 324(72.5) 31(6.9) 92(20.6) 447
40+ mm 106(77.9) 12(8.8) 18(13.2) 136

181
Number
Spineless Without N of samples
7(4.1) 45(26.5) 170 2
9(1.5) 119(20.5) 579 4
3(0.6) 124(26.4) 470 2
2(0.7) 78(26.4) 295 2
8(2.3) 92(26.6) 346 2
2(1.6) 32(26.0) 123 l
0(0.0) 41(23.0) 178 2)
0(0.0) 0(00.0) 0 2
31(1.4) 531(24.6) 2161

and lighter animal, perhaps capable of quicker accel-
eration away from predators (Nelson 1969). If truly an
advantage such streamlining should also be favorable
in flowing water presumably as a result of decreased
hydrodynamic drag (assuming the predator/ prey
situation is equivalent). The significantly lower fre-
quency of withouts in creeks mitigates against sucha
difference; however, care is warranted in interpreting

TABLE 5—Phenotypic frequencies of Brook Stickleback
from different collecting methods in Wakomao Lake

Number of phenotypes (%)

Method With Intermediate Without N
Bag-seine 251(74.0) 17(5.0) 71(21.0) 339
Dip net 152(68.2) _ 14(6.3) Si(25)5) 23
Electrofisher 160(74.8) 12(5.6) 42(19.6) 214
Total 563(72.5) 43(5.5) 170(22.0) 776
Minnow trap 70(68.0) 9(8.7) 24(23.3) 103

TABLE 6—Phenotypic frequency of Brook Stickleback in
collection Number 8, Beaver Pond Creek, between “live” and
“dead” fish

Number of phenotypes (%)

State of

fish With Intermediate Without N
Live 607(75.6) 43(5.3) 153(19.1) 803
Dead 231(78.6) 14(4.8) 49(16.6) 294

182 THE CANADIAN FIELD-NATURALIST

these data because the creeks generally are not rapidly
flowing.

Lack of association of any phenotypic class with a
particular habitat indicates that differential usage or
partitioning of the environment does not occur, at
least at the gross level considered. Predators such as
Northern Pike commonly associate with cover vegeta-
tion which is not too dense (Christiansen 1976). If
association was habitat specific, such predators would
not be attempting to prey upon a frequency of pelvic
phenotypes different from that in the entire lake.
Therefore the discrepancies found by Reist (1980a),
between phenotypic frequencies in the lake and in pike
stomachs, are not due to differential availability of the
pelvic phenotypes.

The nonsignificant change in frequencies between
1971 and 1977 suggests the frequencies are stable thus
mitigating against a transient evolutionary event;
however, the short duration precludes a firm conclu-
sion. Variation in frequency of withouts with season
suggests that a seasonal, environmental parameter
may be a cause of selective elimination of the pheno-
type. Such cyclical selection has been advanced as a
potential factor maintaining polymorphisms (May-
nard Smith 1970). Similarly, the tendency for decrease
in frequency of withouts with age suggests they may
have some disadvantage relative to withs.

Methodological biases or scoring are not responsi-
ble for observed variation in phenotypic frequencies;
thus, a biological explanation for temporal and geo-
graphic variation is most likely. Few data are availa-
ble on historical influences, such as genetic drift or
gene flow, which may significantly affect frequencies
prior to the action of selective forces. Accordingly, the
most productive area to investigate for factors contri-
buting to the maintenance of the observed phenotypic
frequencies is one involving selective explanations.
Indication of a selective disadvantage of withouts has
been presented, that is, decrease in frequency over
winter and with age, but a specific selective agent has
not been identified. The lack of selectivity for any
phenotype in the sample of “naturally” dead fish indi-
cates not all environmental agents are selective forces
involved in maintaining this variation.

Other studies have investigated one potential aspect
of selection, predation by vertebrate and invertebrate
piscivores (Reist 1980a, b). These studies suggest a
relative disadvantage for the withs during predation
by large Northern Pike and by Dytiscus larvae (Dytis-
cidae, water beetles). This disadvantage seems to be
the result of behavioral rather than morphological
differences between the with and without phenotypes.
This disadvantage of the with phenotype could poten-
tially be balanced by the aforementioned disadvan-
tage of the without phenotype. Thus the frequency of
phenotypes at any one point in time or space probably

Vol. 95

represents a balance between opposing selective forces
that tend to eliminate the least fit phenotype. Qualita-
tive or quantitative variation of the selective regime,
primarily predators, would result in different stable
equilibria of the frequencies thus explaining the diver-
sity of frequencies observed (Nelson 1969, 1977; this
study).

These studies are insufficient to explain all aspects
of this phenotypic variation, thus other predators or
other selective forces such as differential reproductive
success (for example, fecundity, assortative mating)
must be considered.

Acknowledgments

I thank E. J. Crossman, B. G. Naylor, and J. S.
Nelson for valuable comment on early versions of the
manuscript. J. S. Nelson, my thesis supervisor, pro-
vided welcome guidance and encouragement through-
out the study. R. Foote, L. Reist, and T. van Meer
assisted in the field collections. Funding for the study
was provided by National Research Council of Can-
ada operating grant A5457 (to JSN) and a National
Research Council of Canada postgraduate
scholarship.

Literature Cited

Christiansen, D. G. 1976. Feeding and behavior of North-
ern Pike (Esox /ucius Linnaeus). M.Sc. thesis, Depart-
ment of Zoology, University of Alberta. 302 pp.

Maynard Smith, J. 1970. The causes of polymorphism.
Symposia Zoological Society of London 26: 371-383.

Nelson, J. S. 1969. Geographic variation in the Brook
Stickleback, Cu/aea inconstans, and notes on nomencla-
ture and distribution. Journal of Fisheries Research Board
of Canada 26: 2431-2447.

Nelson, J. S. 1977. Evidence of a genetic basis for absence
of the pelvic skeleton in the Brook Stickleback, Culaea
inconstans, and notes on the geographical distribution and
origin of the loss. Journal of the Fisheries Research Board
of Canada 34: 1314-1320.

Nelson, J.S., and F.M. Atton. 1971. Geographic and
morphological variation in the presence and absence of the
pelvic skeleton in the Brook Stickleback, Culaea incon-
stans (Kirtland), in Alberta and Saskatchewan. Canadian
Journal of Zoology 49: 343-352.

Nelson, J. S., and M. J. Paetz. 1974. Evidence of under-
ground movement of fishes in Wood Buffalo Park Can-
ada, with notes on recent collections made in the park.
Canadian Field-Naturalist 88: 157-162.

Reist, J. D. 1980a. Selective predation upon pelvic pheno-
types of Brook Stickleback, Cu/aea inconstans, by Nerth-
ern Pike, Esox Jucius. Canadian Journal of Zoology 58:
1245-1252.

Reist, J. D. 1980b. Predation upon pelvic phenotypes of
Brook Stickleback, Culaea inconstans, by selected inver-
tebrates. Canadian Journal of Zoology 58: 1253-1258.

Sokal, R.R., and F. J. Rohlf. 1969. Biometry. W. H.
Freeman and Co., San Francisco. 776 pp.

Received 28 June 1980
Accepted 21 October 1980

4

Feeding and Social Behavior of Some Migrant Shorebirds in Southern
Manitoba

RICHARD A. WISHART,' PATRICK J. CALDWELL,2 and SPENCER G. SEALY!

!Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2
2Ducks Unlimited (Canada), 1190 Waverley Street, Winnipeg, Manitoba R3T 2E2

Wishart, Richard A., Patrick J. Caldwell, and Spencer G. Sealy. 1981. Feeding and social behavior of some migrant
shorebirds in southern Manitoba. Canadian Field-Naturalist 95(2): 183-185.

The foraging and social behavior of spring migrant shorebirds were observed as they fed on earthworms ina ploughed field in
southern Manitoba. Black-bellied Plovers (Pluvialis squatarola) and American Golden Plovers (P. dominica) foraged by
visually locating prey, interspersed with bursts of walking and probing. A greater proportion of feeding attempts by
Black-bellied Plovers were successful compared to those of American Golden Plovers, but the latter made more attempts and
hence obtained about the same number of prey items per unit time of foraging. Ruddy Turnstones (Arenaria interpres) had a
lower percentage of successful feeding attempts than plovers but obtained more than twice the number of food items per
minute. Social interactions in the flocks, particularly involving Black-bellied Plovers, led to a dominance hierarchy that was
related directly to body size. Bill and leg lengths may also have affected the ability of birds to capture prey and influenced their
feeding success rates and strategies.

Key Words: Black-bellied Plover (Pluvialis squatarola), American Golden Plover (P. dominica), Ruddy Turnstone (Arenaria
interpres), foraging, dominance, migration, Manitoba.

Shorebirds often associate in mixed-species flocks length of the bird, and counted the number of pauses
while feeding. Such assemblages during migration between feeding attempts. Intra- and inter-specific
offer opportunities to compare foraging strategiesand interactions were also described.
possible competition among closely related species We measured earthworm (Oligochaeta) densities
that may not come into contact on breeding or winter- on 27 May, using Raw’s (1959) formaldehyde expel-
ing grounds (Recher and Recher 1969; Goss-Custard _ lant techique. Quadrats, each 0.8 m2, were sampled in
1970; Baker and Baker 1973; Burger et al. 1979). In _ the field used by birds and inan adjacent unused field.
this study, we compared the feeding behavior of | The pH ofasoilsample from the top 4 cm ineach field
Black-bellied Plovers (blackbellies) (Pluvialis squa- was measured, and the soil oven-dried to estimate its
tarola), American Golden Plovers (goldens) (P. moisture content.
dominica), and Ruddy Turnstones (Arenaria inter-
pres) that fed in a mixed flock in a ploughed field in Results
southern Manitoba during spring migration. We doc- Shorebirds were seen at the field from 25 May until
umented social interactions within the flock which 29 May, but counts were made only on 27 and 28 May
also included Ring-billed Gulls (Larus delawarensis) | (Table 1). Soil from this field had a moisture content

and Least Sandpipers (Cal/idris minutilla). of 31.1% compared to 8.7% for the unused field. The
pH of these fields was similar (6.7 and 6.9, respec-
Study Area and Methods tively), but earthworm (Aporectodea sp.) density dif-

The birds were concentrated on a wet ploughed fered markedly (77.1 worms/m? and zero, respec-
field 0.6 km2, 16km north of Portage la Prairie, tively). Other soil invertebrates sampled (e.g.,
Manitoba (50° 06’N, 98° 16’W). Other such fields, used
for growing cereal grains, surrounded the area. al

We made 4 h of detailed observations on 27 and 28 Se Bee as offbirds; counted onstudy.areaoni jane
Mavaloy/ ss usingsa 20-45) timesispottingascope froma. (iusnalnieenhnliiseen ales Co eee
stationany vellicle. Birds, weres30—100 matron thes "5 seit; Sli nmin mille pinta angie
observers. Before each observation period we counted

the number of each species present. Behavior of a 27May = A.M. 0 125 61 3 0
feeding individual was watched for as long as possible Fons y 25) 20 2 36
(1.5-14 min), and feeding attempts and their success 28 May A.M. 0 0 2) 0 0
were described into a tape recorder. We could see _—SsP-M. 20 16892300

birds pulling up and mandibulating prey when suc- *RB, Ring-billed Gull; BB, Black-bellied Plover, GP, Ameri-
cessful. We estimated the distance plovers moved after can Golden Plover; RT, Ruddy Turnstone; LS, Least

successive feeding attempts in relation to the known Sandpiper.

183

184

Araneae, Acarina, and Chilopoda) were not common
in either field.

Six goldens were observed for 44.5 min (range of
4-12 min each), five blackbellies for 43 min
(5-14 min), and two turnstones for 7.5 min
(1.5-6 min). The remaining 145 min of observation
were spent watching the flock as a whole and recording
social interactions.

Birds moved about the field probing in the soil or
pecking at its surface for food. Most food items
obtained were earthworms, which we could see being
pulled from their burrows oras they dangled from the
birds’ bills.

Blackbellies and goldens typically moved with head
erect in rapid bursts of walking, interspersed with
pauses when they often looked down and probed or
pecked once or several times for food. Both species
moved similar distances (with means + SD for black-
bellies = 1.3 + 1.0 m,n = 52 vs. goldens= 1.1 + 1.1 m,
n= 105) and paused a similar number of times
between feeding attempts (blackbellies = 3.3 + 2.3,
n= 52 vs. goldens = 3.2 + 2.3,n = 105) whether aftera
successful or unsuccessful feeding attempt (P > 0.05,
t-test) (thus data above are pooled).

Blackbellies made slightly fewer feeding attempts
per minute of foraging than goldens (2.9 + 1.l,n=5
birds vs. 3.8 + 1.3, n = 6) (P= 0.17, Mann-Whitney U
test), but a greater proportion of these were successful
(40.4+9.1% vs. 23.94 15.4%) (P< 0.05). Thus
blackbellies obtained slightly more earthworms per
minute of foraging (1.1 + 0.3 vs. 0.8 = 0.3) (P = 0.06).

In contrast, two Ruddy Turnstones we watched
averaged 42.7 feeding attempts/min (range
25.3-60.0), although only 7.4% (5.6—-9.2) of these were
successful. They secured an average of 2.8 worms/ min
(2.3-3.3), more than double the rate for either plover
(P< 0.05). Unlike the plovers, turnstones moved in
rapid bursts with tail up and bill down, almost con-
stantly pecking and probing in the soil for food.

Although none of the birds in the flock appeared to
be paired and none was courting, aggressive interac-
tions between individuals often occurred (Table 2). In
43 min five blackbellies were involved in 20 encoun-
ters (0.47/ min) whereas in 44.5 min six goldens were
in 10 encounters (0.22/ min).

Overall, 57.3% (n= 110) of aggressive encounters
were between conspecifics. Fifty-five percent of all
encounters resulted in one bird chasing another, usu-
ally for less than 3 m. The remainder involved one
bird supplanting another without pursuit. No fighting
was observed but many intraspecific encounters
involved displays. Both blackbellies and turnstones
gave chattering calls during some encounters.

An interspecific dominance hierarchy, related
directly to body size, existed in the flock (Table 2).

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 2— Aggressive encounters among birds’, arranged by
size and position in the dominance hierarchy

Winner
Loser RB BB GP RF LS Total
RB 0 0 0 0 0 0
BB 15 S7/ 0 l 0 73
GP 0 17 3 0 0 20
RT 4 4 D) 3 0 13
LS 0 3 l 0 0 4
Total 19 81 6 4 0 110

“RB, Ring-billed Gull; BB, Black-bellied Plover; GP, Ameri-
can Golden Plover; RT, Ruddy Turnstone; LS, Least
Sandpiper.

Ninety-nine percent of interspecific encounters were
won by a larger species. Only once did a turnstone
(smaller) supplant a blackbelly (larger).

Several encounters may have been attempts at
pirating food. One turnstone grabbed an earthworm
as it dangled from the bill of a blackbelly. Ring-billed
Gulls occasionally chased blackbellies up to 90 m in
flight, presumably trying to steal their prey (see Payne
and Howe 1976; Kallander 1977). However, no suc-
cessful kleptoparasitism was observed. Gulls were
successful in securing earthworms independently, but
their individual feeding rates were not timed.

Discussion

The birds observed in this study exploited a locally
abundant food source. Others have also noted
migrant plovers using agricultural fields and feeding
on earthworms (Bent 1929; Burton 1974; Payne and
Howe 1976), a nutritious food (Kallander 1977). We
do not know if earthworms are available at this field
every spring, nor if it is traditionally used by shore-
birds, as are certain coastal areas and inland fields on
both the breeding and wintering grounds (Bengtsonet
al. 1978; Fuller and Youngman 1979).

The plovers fed similarly but the blackbellies
obtained more earthworms/ min. Blackbellies were
dominant over goldens, and may have been able to
usurp superior feeding sites. Goldens, by avoiding
encounters with blackbellies (which they always lost),
may have spent less of their feeding time in ‘superior’
areas. Like the American Robin (Turdus migratorius)
(Heppner 1965), most land-foraging shorebirds prob-
ably rely on sight to detect earthworms. Blackbellies
have slightly longer legs (4.7%) and much longer bills
(25%) (Godfrey 1966) than goldens, and so may be
able to observe potential prey at a greater distance,
move faster, and be more successful at capturing it.

Foraging by turnstones seemed to be energetically
more costly (faster rate of pecking and movement)

1981

than that used by plovers, although they obtained
more food items per foraging minute. Turnstones
have much shorter legs than either plover (40%) (God-
frey 1966) and may less often observe prey from a
distance. Turnstones seemed to take observable prey
when available but supplemented this by exposing
food that would have been missed by relying on visual
cues only (plover method). The latter part of their diet
is, at least momentarily, unavailable to plovers, and
this may allow turnstones to reduce competition with
the more dominant species. Although food intake
rates varied, without measuring time budgets and prey
sizes taken, it is not yet possible to compare net energy
gains among the species.

Most social interactions were intraspecific, as
found by Recher and Recher (1969) and Burger et al.
(1979). Dominance in the hierarchy was related
directly to body size, as noted also by Bailey and Batt
(1974), Morse (1974), and Burger et al. (1979). Thus,
larger birds have an advantage in competing, particu-
larly if resources are limited or patchy. However, if
they are relying on the same food, larger individuals
require more items and so must expend more effort in
meeting their nutrient needs.

Stinson (1977) saw little intraspecific aggression in
a wintering flock of blackbellies, and believed that
avoidance was responsible for the spaced-out distri-
bution that he observed. Strauch and Abele (1979)
observed a similar relationship for other wintering
plovers. However, in our study the birds spent much
time in aggressive encounters, perhaps owing to dif-
ferences in food distribution or flock density. Where
food is aggregated or bird numbers are large in rela-
tion to the available feeding space, encounter rate and
thus competition increase (see Stinson 1980). Burger
et al. (1979) and Mallory and Schneider (1979) found
similar relationships between rates of aggressive
encounters and habitat conditions with other shore-
bird species. Unlike wintering shorebirds those we
observed were presumably depositing fat to meet the
demands of migration and reproduction. This is likely
another reason why we saw more competition than
did other observers.

Acknowledgments

We thank C. M. Savage and T. G. Neraasen for
helping to collect data and J. W. Reynolds for identi-
fying the earthworms. While involved in this and
other studies we were funded by the Delta Waterfowl
Research Station, Canadian Wildlife Service, Univer-
sities of Manitoba and Kansas State, and grant A9556
(to S.G.S.) of the Natural Science and Engineering
Research Council of Canada. G. W. Page and another
reviewer offered helpful advice.

WISHART ET AL.: MIGRANT SHOREBIRD FEEDING BEHAVIOR

185

Literature Cited

Bailey, R. O., and B. D. J. Batt. 1974. Hierarchy of water-
fowl feeding with Whistling Swans. Auk 91: 488-493.

Baker, M. C., and A. E. M. Baker. 1973. Niche relation-
ships among six species of shorebirds on their wintering
and breeding ranges. Ecological Monographs 43: 193-212.

Bengtson,S.-A., A. Nilsson, and S. Nordstrom. 1978. Selec-
tive predation on lumbricids by Golden Plover, Pluvialis
apricaria. Oikos 31: 164-168.

Bent, A. C. 1929. Life histories of North American shore
birds II. United States National Museum Bulletin Number
146. 412 pp.

Burger, J.. D.C. Hahn, and J. Chase. 1979. Aggressive
interactions in mixed-species flocks of migrating shore-
birds. Animal Behavior 27: 459-469.

Burton, P. J. K. 1974. Feeding and the feeding apparatus
in waders: a study of anatomy and adaptations in the
Charadrii. British Museum of Natural History Publica-
tion Number 719. 150 pp.

Fuller, R. J..and R. E. Youngman. 1979. The utilisation of
farmland by Golden Plovers wintering in southern Eng-
land. Bird Study 26: 37-46.

Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203. 428 pp.

Goss-Custard, J.D. 1970. Feeding dispersion in some
over-wintering wading birds. /m Social Behaviour in birds
and mammals. Edited by J. H. Crook. Academic Press,
London. pp. 3-35.

Heppner, F. 1965. Sensory mechanisms and environmental
clues used by the American Robin in locating earthworms.
Condor 67: 247-256.

Kallander, H. 1977. Piracy by Black-headed Gulls on
Lapwings. Bird Study 24: 186-194.

Mallory, E.P., and D.C. Schneider. 1979. Agonistic
behavior in Short-billed Dowitchers feeding on a patchy
resource. Wilson Bulletin 91: 271-278.

Morse, D. H. 1974. Niche breadth as a function of social
dominance. American Naturalist 108: 818-830.

Payne, R. B., and H. F. Howe. 1976. Cleptoparasitism by
gulls of migrating shorebirds. Wilson Bulletin 88: 349-351.

Raw, F. 1959. Estimating earthworm populations by using
formalin. Nature 184: 1661-1662.

Recher, H. F.,and J. A. Recher. 1969. Some aspects of the
ecology of migrant shorebirds. II]. Aggression. Wilson Bul-
letin 81: 140-154.

Stinson, C. H. 1977. The spatial distribution of wintering
Black-bellied Plovers. Wilson Bulletin 89: 470-472.

Stinson, C. H. 1980. Flocking and predator avoidance:
models of flocking and observations on the spatial disper-
sion of foraging winter shorebirds (Charadrii). Oikos 34:
35-43.

Strauch, J. G., Jr., and L. G. Abele. 1979. Feeding ecology
of three species of plovers wintering on the Bay of
Panama, Central America. Studies in Avian Biology 2:
217-230.

Received 26 May 1980
Accepted 6 October 1980

Distribution, Growth, and Foods of Arctic Cod
(Boreogadus saida) in the Bering, Chukchi, and Beaufort Seas

LLOYD F. LOWRY AND KATHRYN J. FROST

Alaska Department of Fish and Game, 1300 College Road, Fairbanks, Alaska 99701

Lowry, Lloyd F., and Kathryn J. Frost. 1981. Distribution, growth, and foods of Arctic Cod (Boreogadus saida) in the
Bering, Chukchi, and Beaufort seas. Canadian Field-Naturalist 95(2): 186-191.

Arctic Cod (Boreogadus saida) are abundant and ecologically important in arctic and subarctic marine ecosystems. Based on
otter trawl collections made in the northern Bering and northeastern Chukchi and Beaufort seas, Arctic Cod were most
abundant in the northeastern Chukchi and western Beaufort seas. There was no relationship between water depth and
abundance; however, in the Chukchiand Beaufort seas small fishes were more common in water less than 100 mdeep. Annual
growth increments appear to be quite variable which complicates interpretation of size at age data. Arctic Cod less than 4 yr
were larger at a given age in the Bering Sea. In the Bering Sea gammarid amphipods were the main food with mysids, shrimps,
and hyperiid amphipods also eaten. In the northeastern Chukchi and Beaufort seas, calanoid copepods were by far the
predominant prey with other crustaceans eaten much less frequently. Arctic Cod appear to bea very adaptable species which
may explain their overwhelming success in Arctic marine waters.

Key Words: Boreogadus saida, Arctic Cod, distribution, foods, growth, abundance.

Three species of gadid fishes are abundant and eco-
logically important in marine waters of western and
northern Alaska. Walleye Pollock (Theragra chalco-
gramma) is the most abundant finfish in southern and
central Bering Sea (Bakkala, R. G., and G. B. Smith
1978. Demersal fish resources of the eastern Bering
Sea: spring 1976. Northwest and Alaska Fisheries
Center Processed Report. 233 pp.). Saffron Cod (Ele-
ginus gracilis) is the most abundant species during
summer months in the northern Bering and southern
Chukchi seas (Wolotira, R. J.. J. M. Sample, and M.
Morin, Jr. 1977. Demersal fish and shellfish resources
of Norton Sound, the southeastern Chukchi Sea and
adjacent waters in the baseline year 1976. Northwest
and Alaska Fisheries Center Processed Report. 292
pp.). In truly arctic waters and in winter months in the
northern Bering and southern Chukchi seas, Arctic
Cod (Boreogadus saida) is by far the most abundant
forage fish species (Moskalenko 1964; Ponomarenko
1968). In combination these species provide a large
portion of the annual nutrition of most species of
marine mammals in these areas (Klumov 1937;
Tomilin 1957; Johnson et al. 1966; Lowry etal. 1979).
Generally, a more diverse assemblage of forage fish
species are available and utilized in the Bering than in
the Chukchi and Beaufort seas. In the Beaufort Sea,
Arctic Cod is virtually the only widely distributed and
abundant forage fish species.

The biology of Walleye Pollock has been studied in
detail because of its importance in commercial bot-
tomfish catches. Less is known of the biology of Arctic
and Saffron Cods although there have been several
significant studies of Arctic Cod in the Barents Sea
and in Siberian waters (Moskalenko 1964; Baranen-

186

kova et al. 1966; Hognestad 1968; Ponomarenko
1968; Rass 1968). In light of their ecological impor-
tance and pending developments which may affect
marine systems, detailed information on the biology
of Arctic and Saffron Cod in Alaskan waters Is greatly
needed. In this paper we present observations on the
biology of Arctic Cod caught by bottom trawls in the
Bering, Chukchi, and Beaufort seas.

Methods and Materials

Tows were made using a 5.8-m (19-ft) headrope
semiballoon otter trawl with 3.2-cm (1.25-in.) stretch
mesh webbing in the body and 0.6-cm (0.25-in.)
stretch mesh liner in the cod end. In the northeastern
Chukchi and Beaufort seas, 33 successful tows
(10 min bottom time) were made during the period 2
August to 3 September 1977. This set of tows was
broken down by three subareas: northeastern Chuk-
chi Sea (west of Point Barrow), 10 tows; western
Beaufort Sea (Point Barrow to Prudhoe Bay), 8 tows;
central Beaufort Sea (east of Prudhoe Bay), 15 tows.
In the northern Bering Sea 32 tows (20 min bottom
time) were made between 27 May and 10 June 1978
(Figure |). Arctic Cod were sorted from other compo-
nents of the trawl catches. Fishes caught in the Bering
Sea tows were immediately enumerated, weighed, and
measured (fork length to the nearest millimetre), and
otoliths and stomachs were removed. Stomachs were
split open and the contents were washed into a petri
dish where prey items were identified and separated by
taxon. Components of the stomach contents were
ranked according to their relative abundance, 1.e., the
most abundant prey was givena rank of I, the second
most abundant a rank of 2, and so on. Arctic Cod

—

198]

160

CHUKCHI
SEA

Saint
Lawrence
Island

LOWRY AND FROST: BIOLOGY OF ARCTIC COD 187

140°

BEAUFORT SEA

CANADA

ALASKA

FIGURE |. Map of Alaska showing the two regions (broken lines) where Arctic Cod were collected.

caught in the Chukchi and Beaufort seas were treated
similarly except that fishes were preserved by injection
of 10% formalin into the abdominal cavity and
immersion in a 10% formalin-seawater solution.
Fishes were weighed and measured and stomach con-
tents were examined 3-6 wk later. For presentation of
results, lengths of preserved fishes were corrected for
shrinkage resulting from preservation by increasing
measured length by 2.1% (Lowry and Frost, unpub-
lished data based on 25 Arctic Cod measured before
and after 3 wk preservation in 10% formalin). Ages of
fishes were determined using annular rings in otoliths.
The medial surface of each otolith was ground ona
fine stone. The otoliths were then immersed in xylene
and the rings counted using a binocular microscope
and reflected light. In some otoliths, rings were more
easily observed by examining a cross section. In such
cases the otolith was broken transversely through the
middle and the broken surface was polished, then
examined under the microscope.

Results and Discussion

The overall distribution of Arctic Cod generally
coincides with that of seasonal and multiyear sea ice
(Andriyashev 1954; Ponomarenko 1968). With the
exception of three tows in the Bering Sea, all our tows
were made in or very near pack ice. In the Bering Sea,
Arctic Cod were caught in three tows south of the ice;
the most southerly occurrence was at 61°33’0’N,
174° 30'48” W, about 300 km from the ice on 27 May
1978. This area had been ice-covered 3 wk previously.
Arctic Cod are still widely distributed although not
particularly abundant in the northern Bering Sea in
late summer, long after the ice has gone (Wolotira et
al. 1977, op. cit.).

In our samples, Arctic Cod were most widespread
and abundant in the northeastern Chukchi and west-
ern Beaufort seas and least abundant in the northern
Bering and central Beaufort seas (Table 1). This is
perhaps due to many Arctic Cod from the northern
Bering and southern Chukchi seas moving north with

188 THE CANADIAN FIELD-NATURALIST

TABLE 1—Occurrence and relative abundance of Arctic Cod
in the study areas

Tows with No. of
Number Arctic Arctic Cod
of Cod per tow,
Area tows (%) mean (range)
Northern
Bering Sea! 32 56.2 3.3 (0-37)
Northeastern
Chukchi Sea? 10 100.0 10.3 (1-26)
Western
Beaufort Sea? 8 100.0 7.8 (2-24)
Central
Beaufort Sea? 15 66.7 1.9 (0-11)

!'Bottom time 20 min.
2Bottom time 10 min.

the receding ice edge, and suggests that they do not
penetrate far to the east in the Beaufort Sea. There was
no obvious relationship between depth of water and
catches of Arctic Cod. In the northeastern Chukchi
and Beaufort seas, 26 tows were made in water
40-100 mdeep while 8 tows were in waters 101-400 m
deep. The mean number of Arctic Cod caught was 5.7
in shallow tows (range 0-26) and 5.5 in deeper tows
(range 0-24). In the Bering Sea most tows were in

30

2s

20

1S

=
o

NUMBER IN SIZE CLASS

CHUKCHI AND BEAUFORT SEAS

Vol. 95

waters less than 50 m deep and again there was no
obvious relationship between abundance and water
depth.

We casually examined the gonads of the larger Arc-
tic Cod caught; in both sexes these were small, ranging
in weight from 0.1 to 0.9 g, usually less than 3% of the
total body weight of the fish. That is consistent with
previous observations that gonadal development be-
gins in September in preparation for spawning during
winter months (Moskalenko 1964).

Fishes caught in the northeastern Chukchi and
Beaufort seas were generally smaller than those
caught in the northern Bering Sea (Figure 2). In the
former area mean fork length was 8.8 cm (range
4.6-18.4) while in the latter area mean length was
14.7 cm (range 7.2—22.2). This difference is attributa-
ble to several factors. First, the fishes caught in the
Bering Sea were generally older than those from the
more northern area. In the northeastern Chukchi and
Beaufort seas l-yr-old Arctic Cod made up 81% of the
catch, but in the Bering Sea they comprised only 27%
of the catch. Arctic Cod older than 4 yr were caught
only in the Bering Sea. Second, as indicated in Figure
2, Arctic Cod less than 4 yr old were longer at a given
age in the Bering Sea than in the Chukchi and Beau-
fort seas. This difference was largest for l-yr-old fishes
and was progressively less in older age-classes; 4-yr-

FORK LENGTH-cm

FIGURE 2. Size distribution of Arctic Cod caught in the two regions sampled. Mean (triangle) and range (horizontal bar) in
size for each age-class in each region are shown above the size distributions.

1981

old Arctic Cod were of similar length in both areas.
Third, in the northeastern Chukchi and Beaufort seas
small fishes (<10.5 cm) were proportionately more
abundant in waters 100 m or less deep (Table 2). The
size distribution of Arctic Cod from that area (Figure
2) is therefore influenced by the depth distribution of
tows. Twenty-four of 33 tows were made in water
100 m or less in depth. Tows in northern Bering Sea
spanned a much smaller depth range (29-80 m) and
no relationship between fish size and water depth was
evident.

Growth rates of Arctic Cod appear to be quite
variable, especially during the first 2 yr of life (Figure
3). The transition from larvae to juvenile is reported to
occur in August at a length of 3-5 cm (Rass 1968).
Should we assume a mean length at transition of
4.0 cm, Arctic Cod we examined from northern Ber-
ing Sea had grown 6.1 cmin 10 mo while those from
the Chukchi and Beaufort seas grew only 3.4 cm in
12 mo. First-year growth rates reported for Arctic
Cod in the Barents Sea (Hognestad 1968; Gjosaeter
1973) fall between those values. Our data for the Ber-
ing Sea and that reported by Gjosaeter for the Barents
Sea show the expected sequential decrease in annual
length increment with age. However, in the Chukchi
and Beaufort seas and in the samples from the Barents
Sea examined by Hognestad, growth in length was
greater in the second year of life than during the first.
Hognestad found differences in postlarval growth
during the first summer of life both between areas in
the same year and between years at the same area.
Interestingly, he found the greatest annual growth

increment in the second year of life even when data for -

10 sampling years was combined (Figure 3). This
strongly suggests that greatest growth in length during
the 2nd year of life may be common or characteristic
in Arctic Cod. Reported sizes attained at the end of
the Ist year of adult life vary considerably: Bering Sea,
10.1 cm; Chukchi and Beaufort seas, 7.1 cm; Barents
Sea, 9.3 cm (Hognestad 1968; Gjosaeter 1973). In our
samples, 4-yr-old Arctic Cod were of similar mean
sizes in the Bering (17.4 cm) and the Chukchi and
Beaufort seas (17.5 cm), but were smaller than that
reported from the Barents Sea (x = 19.8 cm (Hognes-

TABLE 2—Relationship between water depth and sizes of
Arctic Cod caught in the northeastern Chukchiand Beaufort
Seas

% in length class
Fork length

Depth pels 10.5-

range(m) Mean(range) <10.5cm 14.0cm 414.0 cm
40-100 8.2 (4.6-15.9) 89.0 10.3 0.7
101-400 _—‘10.7 (6.8-18.4) 59.6 5) 14.9

LOWRY AND FROST: BIOLOGY OF ARCTIC COD

189

GROWTH IN PREVIOUS YEAR -cm

AGE CLASS

FiGURE 3. Annual growth increments reported for Arctic
Cod in this study (solid lines) and other studies
(broked lines).

1. Northern Bering Sea, May-June 1978.

2. Northeastern Chukchiand Beaufort seas, August—
September 1977.

3. Barents Sea, August-September 1956-1966 (Hog-
nestad 1968).

4. Barents Sea, July-September 1970-1972 (Gjosae-
ter 1973).

tad 1968) and 19.1 cm(Gjosaeter 1973)). Moskalenko
(1964) reviewed Soviet data on growth of Arctic Cod
in the Kara, Laptev, and Barents seas. He suggested
two forms of Arctic Cod: a slow-growing form from
coastal regions of the Kara and Laptev seas, and a
fast-growing form from the Barents and Kara seas,
which might be more common in the open sea.
Although we cannot comment on the size and mor-
phometric differences discussed by Moskalenko, the
size at age differences observed by us and others seem
adequately explained by variations in growth rates

190 THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 3—Food items in stomachs of Arctic Cod collected in the northern Bering Sea, May-June 1978 and northeastern

Chukchi and Beaufort seas, August-September 1977

Northern Bering Sea, = 73

Northeastern Chukchi and
Beaufort seas, n = 157

Rank in stomach

Frequency of

Rank in stomach Frequency of

Prey item l 2 3 4 occurrence (%) l 2 3 4 occurrence (%)
Gammarid amphipod 38 1] | — 68.4 47 40 10 — 61.8
Copepod - - a 86 22 2 — 70.1
Shrimp 17 3 _ 27.4 l 2 — = 1.9
Hyperiid amphipod 7 — l — 10.9 2 7 4 3 10.2
Mysid 6 5 2 l 19.2 4 9 l _ 8.9
Euphausiid — 2 2 l _ 37)
Chaetognath — 4 3 — — 4.5
Larval capelin 4 — — — S65) - - =
Polychaete worm — 2 — — Del — —
Cumacean I — — — 1.4 —
Medusae _ l — = — 0.6

induced by annual variations in food availability
(primary production), without invoking discrete
forms in a species with a circumpolar distribution
(Ponomarenko 1968).

We examined stomachs of 288 Arctic Cod. Recog-
nizable food remains were found in 73 of 101 stom-
achs from the northern Bering Sea and 157 of 187
stomachs from the northeastern Chukchi and Beau-
fort seas (Table 3). Major prey in the northern Bering
Sea, gammarid amphipods (mostly Ampelisca
macrocephala), shrimps (mostly Eualus fabricii and
E. gaimardii), and mysids (Neomysis rayii), were
primarily benthic forms. Prey found in stomachs col-
lected in the northeastern Chukchi and Beaufort seas
indicate feeding primarily on_pelagic/ planktonic
organisms; copepods (mostly Calanus hyperboreus,
C. glacialis, Euchaeta glacialis, and few Metridia
longa and C. cristatus) and Apherusa glacialis, a
pelagic gammarid amphipod (Barnard 1959), were by
far the predominant prey. The species of mysid eaten
in this area were Pseudoma truncata and Mysis litora-
lis in contrast to Neomysis rayii in northern Bering
Sea. The widely distributed hyperiid amphipod
Parathemisto libellula was quite commonly eaten in
both areas.

We examined size-related differences in foods of
Arctic Cod (Table 4). In both areas, hyperiid amphip-
ods were more commonly eaten by larger fishes, and
chaetognaths were found only in smaller fishes. Over-
all, a greater size-related difference in foods was seen
in the northern Bering Sea where the primary prey of
larger cod, gammarid amphipods, and shrimps were
much less commonly eaten by smaller fishes.

Similar results have been reported in other studies
of foods of Arctic Cod. In the eastern Canadian Arc-

tic, Bohn and McElroy (1976) reported copepods as
the main food with pelagic amphipods and other crus-
taceans eaten in much smaller quantities. They found
that copepods were more important in the diet of
small fishes while large cod ate more shrimp. Hognes-
tad (1968) reported that copepods were the major
food of Arctic Cod collected in the eastern Barents
Sea in September. Euphausiids, hyperiid amphipods,
mysids, fish and shrimp eggs, phytoplankton, and
small fishes have all been reported as foods of Arctic
Cod in the Kara Sea (Klumov 1937; Moskalenko
1964). Similar foods have been reported from other
portions of the Siberian Arctic (Moskalenko 1964).

TABLE 4—Size-related differences in foods of Arctic Cod
collected in the northern Bering and the northeastern Chuk-
chi and Beaufort seas. Only prey that occurred in more than
two stomachs are included

Frequency of occurrence (%)

Northeastern
Northern Chukchi and
Bering Sea Beaufort seas
Prey item <12.0 cm 212.0 cm S10.0 cm >10.0 cm
Gammarid
amphipod 15.4 82.9 57.8 70.8
Copepod _ _ 74.2 62.5
Shrimp 15.4 46.3 2.3 4.2
Hyperiid
amphipod _ 19.5 7.0 20.8
Mysid 38.5 39.0 7.0 12.5
Euphausiid — = 1.6 4.2
Chaetognath 30.8 — B29 —
Larval capelin doll ies — =

1981

Our results and data available in the literature indi-
cate considerable variability in a number of the bio-
logical characteristics of Arctic Cod. Their abundance
and food habits vary both geographically and season-
ally. Growth rates vary by area and by year. Abun-
dance, feeding, and growth rates are undoubtedly
interrelated in such a way that Arctic Cod optimally
utilize available resources each year. Such adaptabil-
ity is essential in a system with great annular fluctua-
tions in physical (ice cover) and biological (primary

productivity) characteristics and probably explains
the overwhelming success of the species in Arctic
waters. More data are needed before a rigorous exam-
ination of functional aspects of Arctic Cod productiv-
ity will be possible. Further studies of the biology of
Arctic Cod are urgently needed to consider the ecolog-
ical importance of Arctic Cod both as a potential
competitor with species such as Ringed Seals (Phoca
hispida) and Bowhead Whales (Balaena mysticetus)
(Lowry and Burns 1980) and as a food source for
many arctic marine mammals, birds, and fishes
(Klumov 1937).

Acknowledgments

We thank the officers and crew members of the
NOAA ship SURVEYOR and the USCG cutter
GLACIER for their essential support of our collec-
tions. P. Wagner and K. Coyle aided in the identifica-
tion of copepods and amphipods. Project support was
provided by the United States Bureau of Land Man-
agement Outer Continental Shelf Environmental
Assessment Program and Federal Aid in Wildlife Res-
toration Project W-17-9.

Literature Cited

Andriyashey, A. P. 1954. Fishes of the northern seas of the
U.S.S.R. Keys to the fauna of the U.S.S.R. No. 53. (Trans-
lated from Russian by Israel Program for Scientific Trans-
lations, 1964).

Baranenkova, A. S., V. P. Ponomarenko, and N. S. Khokh-
lina. 1966. The distribution, size, and growth of the lar-
vae and fry of Boreogadus saida (Lep.) in the Barents Sea.
Voprosy Ikhtiologii 6: 498-518. (Translated from Rus-
sian, available as Fisheries and Marine Service Transla-
tion Series Number 4025, 1977. 37 pp).

Barnard, J. L. 1959. Epipelagic and under-ice amphipods
of the central Arctic Basin. Geophysical Research Papers
63(1): 115-152.

LOWRY AND FROST: BIOLOGY OF ARCTIC COD IS)!

Bohn, A., and R. O. McElroy. 1976. Trace metals (As, Cd,
Cu, Fe, and Zn) in Arctic Cod, Boreogadus saida, and
selected zooplankton from Strathcona Sound, Northern
Baffin Island. Journal of the Fisheries Research Board of
Canada 33(12): 2836-2840.

Gjosaeter, J. 1973. Preliminary results of Norwegian polar
cod investigations 1970-1972. International Council for
the Exploration of the Sea Report, 1973. 23 pp.

Hognestad, P. T. 1968. Observations on polar cod in the
Barents Sea. /n Symposium on the ecology of pelagic fish
species in arctic waters and adjacent seas. Editedby R. W.
Blacker. International Council for the Exploration of the
Sea Report, Volume 158. pp. 126-130.

Johnson, M. L., C. H. Fiscus, B. T. Ostenson, and M. L.
Barbour. 1966. Marine mammals. /n Environment of the
Cape Thompson Region, Alaska. Edited by N. J. Wil-
imovsky and J. N. Wolfe. United States Atomic Energy
Commission, Oak Ridge, Tennessee. pp. 897-924.

Klumoy, S. K. 1937. Polar cod and their importance for
certain life processes in the Arctic. Izvestiya Akademiya
Nauk SSSR (Biol.), No. 1 (in Russian).

Lowry, L.F., and J.J. Burns. 1980. Foods utilized by
Bowhead Whales near Barter Island, autumn 1979.
Marine Fisheries Review. /n press.

Lowry, L. F., K. J. Frost, and J. J. Burns. 1979. Potential
resource competition in the southeastern Bering Sea:
fisheries and phocid seals. Proceedings of the 29th Alaska
Science Conference, Fairbanks, 15-17 August 1978. pp.
287-296.

Moskalenko, B. F. 1964. On the biology of Arctic Cod,
Boreogadus saida (Lepechin). Voprosy Ikhtiologii 4(3):
32, 433-443.

Ponomarenko, V. P. 1968. Some data on the distribution
and migrations of polar cod in the seas of the Soviet arctic.
Jn Symposium on the ecology of pelagic fish species in
arctic waters and adjacent seas. Edited by R. W. Blacker.
International Council for the Exploration of the Sea
Report, Volume 158. pp. 131-134.

Rass, T.S. 1968. Spawning and development of polar cod.
In Symposium on the ecology of pelagic fish species in
arctic waters and adjacent seas. Edited by R. W. Blacker.
International Council for the Exploration of the Sea
Report, Volume 158. pp. 135-137.

Tomilin, A. G. 1957. Cetacea. Jn Mammals of the USSR
and adjacent countries. Edited by V. G. Heptner. (Trans-
lated from Russian by Israel Program for Scientific Trans-
lations, 1967).

Received 25 June 1980
Accepted 30 October 1980

Polymorphism in Colonies of the Land Snail Cepaea nemoralis at
London, Ontario: Changes Over Three Decades

S. M. SINGH

Department of Zoology, The University of Western Ontario, London, Ontario N6A 5B7

Singh, S. M. 1981. Polymorphism in colonies of the land snail Cepaea nemoralis at London, Ontario: changes over three

decades. Canadian Field-Naturalist 95(2): 192-197.

Genetically determined variation for band pattern in the land snail Cepaea nemoralis introduced into North America,
provides an opportunity to study evolutionary processes. Judd reported different band patterns in two London colonies in
1952. In 1979, I located seven additional well-established colonies in London. The degree of polymorphism in 1979 colonies
(75-87%) was comparable to values obtained for the 1952 colonies. Colony A of 1952 formed a separate lineage in the
“similarity” dendrogram whereas colony B grouped with three 1979 colonies. Four other 1979 colonies formed their own
grouping, suggesting differentiation during the process of colonization.

Key Words: Cepaea, land snail; polymorphism, evolution, genetics.

The land snail Cepaea nemoralis is a western Euro-
pean species. It is common to a wide variety of habi-
tats at different altitudes ranging from dunes to culti-
vated fields. Since the time of Linnaeus it has been
known to be one of the most polymorphic members of
the European fauna. From the examination of fossil
specimens, Diver (1929) showed that this species has
been polymorphic for its banding patterns since the
Neolithic period and commented that it has remained
approximately constant since then. Various aspects of
the ecology of Cepaea have been extensively studied
and information is available on its biologically impor-
tant ecological characteristics (Jones et al. 1977). The
genetics of its visible polymorphism (band patterns) is
fairly well understood (Cain et al. 1960; Cain and
Sheppard 1957; Cook 1967; Lamotte 1951; Wolda
1969: Murray 1975). The band pattern polymorphism
is extensive in most colonies and is easy to score.
About 10 000 populations have been studied in their
natural habitat. In every colony a number of different
morphs are observed and various colonies are distin-
guished by ratios of these phenotypes (Murray and
Clarke 1978).There is a wide range of opinion among
biologists as to the evolutionary mechanisms at work
in causing and maintaining this polymorphism in nat-
ural populations. Among others, Cain and Sheppard
(1950, 1954) and Clarke (1960, 1962) attributed the
differences in the frequency of various morphs
between colonies to selection by predators against
most conspicuous morph(s); others, however, attrib-
uted this difference to random drift and founder effect
(Lamotte 1959; Goodhart 1963), “cryptic” environ-
mental factors (Cain and Currey 1963), and climatic
selection (Arnold 1968, 1971; Jones 1973). The rela-
tive importance of these forces stays an unresolved
problem (Jones et al. 1977).

A long-term follow-up of the changes in the fre-
quency of different morphs along with the dispersal
and development of new colonies under different eco-
logical circumstances is essential in evaluating and
understanding evolutionary forces. Unfortunately,
well-documented studies of morph (gene) frequencies
over long periods of time are not common. The color
polymorphism in the Peppered Moth, Biston betula-
ria (Kettlewell 1965; Ford 1971), probably provides
the best example of this kind. Murray and Clarke
(1978) reported gene frequency changes over a 50-year
period in Cepaea inhabiting the coastal sand dunes at
Berrow, Somerset, England, an area which has stayed
fairly stable. In this paper, I report on the changes in
morph frequencies and the development of new colo-
nies of C. nemoralis in London, Ontario, located
within the city limit in gardens, parks, backyards, and
river banks. The area has undergone extensive urban
development and thus has subjected the existing fauna
to differential selection pressures.

Banding Patterns of the London Colonies
in 1952

Cepaea nemoralis was introduced to North Amer-
ica around 1857 (Reed 1964). Pilsbry (1928, 1939)
reported colonies of introduced Cepaea at various
locations in the USA and Canada while the occur-
rence of C. nemoralis in the London, Ontario, area
was reported at different times by Pilsbry (1928),
Latchford (1930), and Oughton (1948). The species is
now fairly abundant in southern Ontario. Judd (1953,
1955) was the first to report on the band pattern
frequencies in the two colonies he observed in 1952 in
London.

The five bands on the shell (Figure |) are numbered
from the top down: I, 2, 3, 4, 5, and a pattern of five

192

1981

SINGH: POLYMORPHISM IN LAND SNAILS 193

FIGURE |. Three views of a five-banded Cepaea nemoralis shell. Bands are numbered.

separate bands is designated as | 2 3 45. A missing
band is designated by ‘0.’ A complete fusion of two
bands is designated by a round bracket and an incom-
plete fusion by square brackets around the bands
involved. Presence or absence of any combination of
five bands along with fusion of any two or more bands
provides a large number of possible phenotypes in any
colony. In the first colony (A) which was located near
the western city limits on grassy plots, Judd (1953)

found that over 90% of the shells expressed all five.

bands. Of these 35% were 1 2345, 43% were | 23[4 5],
and 10% were | 2 3 (4 5). All other 27 recognized
patterns were present in much lower frequencies. Ina
sample size of 1000, 13 of these patterns were repre-
sented by single individuals and 5 by only two individ-
uals. For others the frequencies were less than 2%
each. Thus characteristically this population was pre-
dominantly | 2 345 type with some fusion of bands 4
and 5. Furthermore, there were no 0 0 3 - - (00 3 with

presence or absence of bands 4 and/or 5) individuals

in this colony. In a second colony (B) which was
located in a well-maintained garden within the city
limit 5 km NE of the first colony, Judd (1955) noted
that the frequencies of banding patterns were signifi-
cantly different from the first colony. About. 25% of
the snails were 0 0 - - -(absence of bands | and 2) as
compared to 1/1000 in the first colony. The fusion of
bands | and 2 was pronounced in the second colony
(41%) as compared to the first colony (2%). Frurther-
more, the band pattern 0 0 3 00 was present only in the
second colony. This study was undertaken to see if
more colonies of C. nemoralis have developed since
1952 and how they compare with the findings of Judd
(1953, 1955) in this area.

Materials and Methods

During the summer of 1979, 10 additional colonies
of C. nemoralis were located within the city boundar-
ies of London, Ontario. At least 200 mature individu-
als were available for scoring from seven of these
colonies. The description of the habitat of these colo-
nies is as follows:

Colony 1. Located in the middle of residential area
at the low-lying edge of a heavily used playing field in
a park. The ground is covered with various grasses
and goldenrod (Solidago sp.) and shaded by large
trees. It leads to a drainage area having a marshy
appearance and faces a high bank behind housing.
Total area approximately 15 X 15 m.

Colony 2. Located about 200 m SE of Colony | in
the same park area separated by a road. It is more
elevated with very little ground vegetation and densely
shaded by a large number of small trees. Total area
approximately 15 X 15 m.

Colony 3. Located ina backyard of an old residen-
tial area about 10 city blocks to the north of Colony 1.
It has a history of gardening with grasses and small
hedges. Approximate area 5 X 5 m.

Colony 4. Located on the river bank about 3.5 km N
of Colony |. The area is covered with dense vegetation
including goldenrod, wild grape (Vitis sp.), and small
bushes. Approximate area 10 X 30 m.

Colony 5. Located ina small patch(5 X 5 m) of gold-
enrod bordering dense coniferous trees in a heavily
used campground about 12 km NE of Colony 1.

194

Colony 6. Located onthe bank of the river ina small
shady area (5 X 5 m) under two low trees with pre-
dominantly grass covering approximately 3 km S of
Colony |.

Colony 7. Located in a low-lying grassy area on the
bank of a drainage channel under high trees about
200 m from Colony 6. Approximate area 5 X 7 m.

Collected individuals were scored for their band
pattern following the usual method. Animals were
returned to their colonies after scoring. An attempt
will be made to follow these and other London colo-
nies at yearly intervals to obtain long-term data. This
will be a further attempt to understand processes of
colonization, selection, and population differentia-
tion in Cepaea in a changing habitat.

Data I obtained were used to compare band pat-
terns of 1979 and 1952 colonies, to establish a degree
of similarity statistically between seven 1979 and two
1952 colonies, and to assess a degree of polymor-
phism, an index of variability, in these colonies.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Results and Discussion

Table | records the banding pattern of 1598 live
adult snails collected from seven colonies in 1979, plus
2000 from two colonies in 1952. Yellow morphs pre-
dominated in all these colonies. It is evident that the
two colonies of 1952 are very different for the fre-
quency of band patterns (P < 0.95). Bands | and 2 are
absent in 25% of the individuals in colony B while only
one individual in a sample size of 1000 shows this
pattern in colony A. Also the fusion of bands 12
and/or 45 is more pronounced in colony B. The 1979
colonies are similar to 1952 colonies in many respects.
Most of the major morphs were present in both
groups. The differences observed in 1952 between the
A and B colonies were also present between colonies
in 1979. Similar to colony A, colonies 2 and 5 have
zero frequency for 0 0 3 0 0 morphand its frequency is
comparable to the B colony in colonies |, 3, and 4.
Colonies 6 and 7 have very high frequency of this
morph (76/302 and 74/272, respectively). Other
morphs (where bands | and 2 are absent), namely,
00345,00 3 [4 5], and 0 0 3 (4 5) are present in all

TABLE |. Comparison of the banding pattern in London colonies of Cepaea studied in 1952 (A, 27 April; B, 5 September) and

1979 (August and September)

Colonies in 1979

Pattern | 2 3 4
12345 4 28 52) 60
123[45] 8 6 8 4
123(45) 16 28 24 24
(12)345 l 0 2 l
[12]3(45) 24 8 6 7
(12)3(45) 66 47 45 18
[12]345 l 0 0 l
[12]3[45] 0 0 | 2
[123](45) 22 1] 3 7
(123)(45) 0 4 0 4
[12345] 0 Dp) 0 0
12345 0 0 0 0
123[45] 0 0 0 0
12045 l 3 l 0
(12)0(45) 4 22 l 8
120[45] 0 l l 0
0234 0 0 0 0
023(45) 0 0 0 0
003[45] | 4 12 0
003(45) 62 18 16 3
00345 14 y) 52 13
00300 2 0 12 3
Others 4 11 3 9
Total 230 195 210 191
Polymorphism

index 0.815 0.872 0.867 0.871

*Modified from Judd (1955).

Colonies* in 1952

5 6 7 A B
68 122 110 353 104
8 22 8 429 20
35 36 16 102 183
2 I 2 0 4
12 12 19 19 31

20 14 0 266

l 2 0 0 4
0 0 | 0 10
8 0 0 0 24
3 0 0 0 59
0 0 0 15 2
0 0 0 13 0
0 0 0 19 0
l 0 4 7 0
4 0 l 0 4
| 0 2 4 0
0 0 0 8 0
0 0 0 ) 0
3 0 0 ] 10
21 0 4 0 139
8 8 14 0 50
0 76 74 0 49
5 3 3 25 4]
198 302 272 1000 1000
0.835 0.765 0.757 0.680 0.856

1981

colonies of 1979 but were found only inthe Bcolony in
1952. The degree of fusion between two or more bands
when all five bands are present, however, varies from
colony to colony in 1979 as in 1952. The frequency of
fused bands 3 and 4, when bands | and 2 are absent,
was quite variable and ranged from 0% in colony 6 to
27% in colony 1. Others, like colonies 2, 3, and 5, had
intermediate frequencies around 10%. A number of
rare morphs that are lumped together as “others”
varied from colony to colony and were represented by
very few individuals in each sample.

Judd (1955) postulated that the bandless pattern
“would probably soon become prevalent when once
established” owing to its dominant nature over the
banded forms (Stelfox 1918); but the frequency of a
given form is not dependent on whether it is dominant
or recessive. Genetic dominance is totally uncon-
nected with gene frequency, as shown by the Hardy-
Weinberg equilibrium law. The frequency of a given
morph or rate of its increase is dependent on its rela-
tive Darwinian fitness ina Mendelian population (see
Spiess 1977 for details).

The similarity among seven 1979 plus two 1952
colonies for their banding pattern was evaluated by
obtaining minimum spanning distance based on two
main principal components (Figure 2). Although the
two principal components account for only 58% of the

PRINCIPAL COMPONENT 1

SINGH: POLYMORPHISM IN LAND SNAILS 195

variance, the relative position of different colonies in
this network is comparable to the dendrogram (Figure
3) based on similarity indexes between pairs of colo-
nies which utilized all the information. The CLUS-
TAN !1C computer program (Wishart 1975) was used
for obtaining the patterns in both figures. Colony 4 of
1979 and B of 1952 are most similar for their band
patterns. Furthermore all 1979 colonies are closer to
the B colony as compared to the A colony of 1952,
although the patterns that were specific to the A col-
ony in 1952 are observed in different proportions in all
colonies of 1979. This suggests differential mixing of
existing colonies in the development of new colonies
and/or some form of selection imposed by factors
associated with urbanization of the study area.
Samples from the colonies of 1952 and 1979 were
also compared using a diversity (polymorphism)
index (Simpson 1979). Here the probability of finding
individuals of different morphs when two are sampled
at random from an infinite population = (1 - y p,).
is |
where pis the frequency of the ‘th morph and ms the
number of morphs. An unbiased estimate of ¥
m G>l|

is given by © nv, i=") where v7, is the number of the
VOVEN)

= a
= P;

(ic=

PRINCIPAL COMPONENT II

FIGURE 2, Minimum spanning tree based on two principal components of band pattern variation showing relationship
between London colonies studied in 1979 (1-7) and 1952 (A and B).

196 THE CANADIAN FIELD-NATURALIST Vol. 95
15-053 +
NBs. “ar
IDo2I
10-802 ‘i
9.365 +
7-967 ar
6-550 +
Sd S3h = ot
3-716 + ie
20289 >
eo

- 1 2

3 A

FIGURE 3. Dendrogram showing relationship between 1979 (1-7) and 1952(A and B) colonies of London based on similarity
in their banding patterns. ( Y axis numbers — Ward’s coefficient of dissimilarity).

ith morph and Nis the sample size. Fora monomor-
phic sample the value for this index is zero and the
maximum value will be (m — 1)/m for a population
with equally frequent m morphs. It is obvious that all
populations are fairly polymorphic (bottom row,
Table 1), although there are differences between the
frequency of various morphs in these colonies. In the
city of London the ecological characteristics
obviously have changed since 1952 with increase in
population density, and new colonies of land snails
have been founded during this period. However, the
diversity (polymorphism) index is comparable
between the colonies observed in 1952 and 1979. This
may reflect on the inherent property of the species and
may be explained by the reproductive biology and
colonizing property of Cepaea. It is monoecious, an
obligate outcrosser, has multiple matings, and stores
sperm from many of them (Murray 1964). One mature
individual may well represent several individual
genomes in terms of its reproductive potential. The
genetic effect of each colonizing individual, therefore,
is greater than in most other species.

The extensive body of information on Cepaea

polymorphism has led to an increase in the complexity
of our explanations of genetic variation, rather than
to the emergence of a single unifying explanation.
Jones et al. (1977) suggested that this polymorphism
cannot be explained by simplistic models which
depend entirely on random processes, entirely on one
type of selection, or even on a few types of selection.
At least eight evolutionary forces are known to affect
shell polymorphism in Cepaea and, in most popula-
tions, several of these in fact may act simultaneously
to control morph frequency (see Jones et al. 1977 for
review). Unique combination of these evolutionary
explanations is needed for almost every Cepaea popu-
lation. Furthermore it is not possible to produce gen-
eral rules from limited studies on few populations.
The pattern of variation in North American colo-
nies may reflect circumstances involved in their intro-
duction and founder effect rather than deterministic
processes like millenia of selection in local environ-
ments. Selection may be the reason for the extinction
of some known colonies that were not found even
after diligent searches. Additional studies over long
periods are necessary to assess the effects of stochastic

1981

processes and natural selection in North American
populations of C. nemoralis, an excellent species for
ecological/evolutionary genetic studies.

Acknowledgments

I express my thanks to W. W. Judd and D. M.
Scott for helpful advice on colonies of Cepaea in
London and to Roger Green for statistical advice.
Financial support was provided bya Natural Sciences
and Engineering Research Council research grant.

Literature Cited

Arnold, R. W. 1968. Studies on Cepaea VII. Climatic selec-
tion in Cepaea in the Pyrenees. Philosophical Transac-
tions of Royal Society of London 253: 549-593.

Arnold, R. W. 1971. Cepaea nemoralis on the East Sussex
south Downs, and the nature of area effects. Heredity 26:
277-298.

Cain, A. J., and J. D. Currey. 1963. Differences in interac-
tions between selective forces acting in the wild on certain
pleiotropic genes in Cepaea. Nature 197: 411-412.

Cain, A. J.,J. M. B. King, and P. M.Sheppard. 1960. New
data on the genetics of polymorphism in the snail Cepaea
nemoralis L. Genetics 45: 393-411.

Cain, A. J., and P.M. Sheppard. 1950. Selection in the
polymorphic land snail Cepaea nemoralis. Heredity 4:
275-294.

Cain, A. J.,.and P. M.Sheppard. 1954. Natural selection in
Cepaea. Genetics 39: 89-116.

Cain, A. J., and P.M. Sheppard. 1957. Some breeding
experiments with Cepaea nemoralis. Journal of Genetics
55: 195-199.

Clarke, B. 1960. Divergent effects of natural selection on
two closely related polymorphic snails. Heredity 14:
423-443.

Clarke, B. 1962. Natural selection in mixed populations of
two polymorphic snails. Heredity 17: 319-345.

Cook, L.M. 1967. The genetics of Cepaea nemoralis.
Heredity 22: 397-410.

Diver, C. 1929. Fossil records of Mendelian mutants.
Nature 124: 183.

Ford, E. B. 1971. Ecological genetics (3rd ed.). Chapman
and Hall, London.

Goodhart, C. B. 1963. “Area effects” and nonadaptive
variation between populations of Cepaea (Mollusca).
Heredity 18: 459-465.

Jones, J.S. 1973. Ecological genetics and natural selection
in molluscs. Science 182: 546-552.

Jones, J. S., B. H. Leith, and P. Rawlings. 1977. Polymor-
phism in Cepaea: a problem with too many solutions.
Annual Review of Ecology and Systematics 8: 109-143.

SINGH: POLYMORPHISM IN LAND SNAILS

197

Judd, W. W. 1953. A colony of the land snail Cepaea
nemoralis (L.)(Helicidae) in the vicinity of London, Onta-
rio. Canadian Field-Naturalist 67: 87-89.

Judd, W. W. 1955. Observations ona second colony of the
land snail Cepaea nemoralis (L.) at London, Ontario, with
a consideration of the banding patterns in the two colo-
nies. Canadian Field-Naturalist 69: 148-150.

Kettlewell, H. B. D. 1965. A twelve-year survey of the fre-
quency of Biston betularia (L.) (Lep.) and its melanic

forms in Great Britain. Entomologist’s Record 77:
195-218.
Lamotte, M. 1951. Recherches sur la structure génétique

des populations naturelles des Cepaea nemoralis (L.). Bul-
letin Biologique de la France et de la Belgique-supplément
35: 1-239.

Lamotte, M. 1959. Polymorphism of natural populations
of Cepaea nemoralis. Cold Spring Harbor Symposium on
Quantitative Biology 24: 65-84.

Latchford, F. R. 1930. Some introduced mollusks. Cana-
dian Field-Naturalist 44: 33-34.

Murray, J.J. 1975. The genetics of the Mollusca. Jn Hand- —
book of Genetics. Volume 3. Edited by R. C. King. Ple-
num, New York. pp. 3-31.

Murray, J. J. 1964. Multiple mating and effective popula-
tion size in Cepaea nemoralis. Evolution 18: 283-291.
Murray, J. J., and B. Clarke. 1978. Changes -of gene fre-
quency in Cepaea nemoralis over fifty years. Malacologia

17: 317-330.

Oughton, J. 1948. A zoogeographical study of the land
snails of Ontario. University of Toronto Studies, Biologi-
cal Series, No. 57.

Pilsbry, H. A. 1928. Helix nemoralis L. in Ontario. Nauti-
lus 42: 42.

Pilsbry, H. A. 1939. Land Mollusca of North America
(north of Mexico). Academy of Natural Sciences, Phila-
delphia, Monograph No. 3, Vol. I.

Reed, C.F. 1964. Cepaea nemoralis (Linn.) in eastern
North America. Sterkiana 16: 11-18.

Simpson, E.H. 1949. The measurement
Nature 163: 668.

Spiess, E. B. 1977. Genes in populations. John Wiley and
Sons, New York. pp. 441-496.

Stelfox, A. W. 1918. Researches into the hereditary charac-
ters of some of our British Mollusca. Part 2. Helix aspersa
Mull. and H. nemoralis L. Journal of Conchology 15:
268-275.

Wishart, D. 1975. CLUSTAN IC, Users manual. Univer-
sity College, London.

Wolda, H. 1969. Genetics of polymorphism in the land
snail, Cepaea nemoralis. Genetica 40: 475-502.

of diversity.

Received 28 April 1980
Accepted 10 November 1980

Scheduling Censuses of Breeding White Pelicans (Pelecanus
erythrorhynchos) in Northern Alberta

RIcK D. BEAVER! and VICTOR LEWIN

Department of Zoology, University of Alberta, Edmonton, Alberta T6G 2E9
\Present address: 318—109 Street, Saskatoon, Saskatchewan S7N IRS

Beaver, Rick D., and Victor Lewin. 1981. Scheduling censuses of breeding White Pelicans (Pelecanus erythrorhynchos) in
northern Alberta. Canadian Field-Naturalist 95(2): 198-201.

In May 1976 and June 1977 aerial surveys were flown over Birch Lake in northeastern Alberta to determine the numbers of
White Pelicans (Pelecanus erythrorhynchos) occupying nests. Ground counts conducted concurrently with the 1977 aerial
surveys revealed that the aerial counts included some pelicans not occupying nests. Pelicans that were present but not
occupying nests were loafing birds, either non-breeders, mates off the nest, or both. Evening rather than morning census
flights are preferable, because the number of such loafing pelicans appeared then to be lower. Censuses to record numbers of
incubating pelicans in northern Alberta should be conducted in late May or early June during the peak of the incubation

period.

Key Words: White Pelican, Pelecanus erythrorhynchos; aerial census, breeding colonies, Alberta.

During 1976 and 1977 we studied breeding White
Pelicans (Pelecanus erythrorhynchos) at Birch Lake
(local name; 57°14’N, 112°44’W) in northeastern
Alberta. Several biases that we found in the census
techniques used to estimate the breeding population
are reported here. These may be applicable in other
areas in Canada as well (Trottier et al. 1980).

White Pelicans nest on remote islands in permanent
or intermittent water bodies, and they frequently
occupy sites which are not heavily forested (Behle
1958: Vermeer 1970; Strait 1973; Beaver and Ballan-
tyne, unpublished data”). Such colonies, which may
be composed of several discrete subcolonies, usually
offer unobstructed viewing from the ground or from
the air. Determination of the number of nests by visits
on the ground can cause abandonment of the site by
the pelicans (Farley 1919; Carson 1966; Sanderson
1966; Vermeer 1969; Johnson and Sloan 1976), and
the practice should be discouraged. Gull predation,
and heating, chilling, or trampling of nest contents
also may result from activities that force adults to
abandon their nests (Johnson and Sloan 1976). Nest
censuses should therefore entail minimal disturbance,
particularly if subsequent productivity (fledgling)
surveys are to be reliably free from the bias introduced
by research activities.

Study Area

The Birch Lake colony, a small island consisting of
sand, clay, and scattered rock, is essentially free of
vegetation and hence offers an unobstructed view of
the pelicans. The island, about 2 m above lowest water
levels (observed in 1977), is subject to periodic flood-

2Canadian Wildlife Service AOSERP (Alberta Oil Sands
Environmental Research Program) 1979. Report 82. 93 pp.

ing caused by Beaver (Castor canadensis) dams on the
outflow drainage of Birch Lake. Birch Lake covers an
area of about 1.7 km?. The island was almost com-
pletely covered by rising water levels in the summer of
1976, but in 1977, because of periodic removal of the
beaver dams, water levels were lower and the island
measured 90 X 23 m.

In 1976 and 1977, 15 and 25 pairs of California
Gulls (Larus californicus), respectively, were nesting
associates of the pelicans. The gulls, although they
had arrived earlier in the spring, were subsequently
displaced to nesting locations peripheral to the pelican
subcolonies.

Methods

Data from literature on other colonies in Canada
were used to select the 27 May census date in 1976.
The 8 June 1977 survey date was selected after prior
observations that year revealed a peak in the number
of pelicans incubating at that time. A Cessna 185
aircraft was flown directly over the colony at eleva-
tions of 600, 450, and 300 m at 09:00 on both dates.
Photographs of the colony were taken at each eleva-
tion using 55- or 200-m lenses on a hand-held 35-mm
single-lens reflex camera with Kodak ektachrome 160,
kodachrome 64, and Plus-X Pan film. Black-and-
white enlargements and projected color slides were
used when counting the pelicans photographed on the
island from the air.

In 1977 we made 16 counts of the numbers of loaf-
ing birds (birds not occupying nests) as well as incu-
bating birds from 8 to 13 June. An observation plat-
form, constructed for the purpose, was situated | m
above water, about 300 m from the island. We con-
ducted the counts with the aid of a 15-60 power spot-
ting scope over the course of several days at different

198

1981]

times of the day. The presence of pelicans loafing on
the island obscured our view of some incubating birds.
Birds thus imperfectly seen were assigned to the two
categories (loafing or incubating) by assuming that
the same proportions of nesting or loafing birds
existed among those not clearly visible as among those
that were clearly either on or off nests. Although
loafing birds regularly congregated at certain areas of
the island, they did not do so exclusively. Birds
observed at these areas, which were always located
apart from established subcolonies, were considered
to be loafing.

Results and Discussion

The chronology of events seems to be similar at all
colonies in Canada (Table 1) although there is some
variability (Vermeer 1970).

The Birch Lake pelicans established nests over a
4-wk period in 1976 and a 2-wk period in 1977. Seven
individual subcolonies (two in 1976, five in 1977) were
established in a mean of 9.3 d (range: 7-14 d) (Table
2). The length of the period when most incubating
birds would be counted may be expected to vary.
Factors suchas infertile eggs, weather, disturbance, or
fluctuating water levels can cause abandonment of
nests, and less reliable census data would result from
later censuses because the evidence (Schaller 1974)
indicates that White Pelicans do not renest. If surveys
had been made after | June in 1976, a number of
incubating birds would not have been counted. Sub-
colonies were established over a 4-wk period and some
birds in the first subcolony formed were nearing the
completion of incubation by | June in 1976. In 1977,
ground observations before 8 June were indispensable

BEAVER AND LEWIN: WHITE PELICAN CENSUSES, ALBERTA 199

in determining the best time for censusing. All birds
were incubating by 4 June with earliest nesting birds
about 19 d into incubation. An aerial census for incu-
bating birds was then scheduled for 8 June to ensure a
maximum count.

Schaller (1964) and Knopf (1975) stated that nest
reliefs for incubating White Pelicans occurred around
midday. Aerial censuses conducted at times other
than midday should have avoided counting birds that
were not on nests (i.e. loafing birds) as only one
member of the pair is present except during nest reliefs
(Schaller 1974; Knopf 1975). Boeker (1972) employed
a morning aerial census to determine the number of
occupied nests during the peak incubation period and
multiplied this figure by 2. We also used this tech-
nique; however, ground observations at Birch Lake
revealed a potential bias due to the presence of loafing
birds on the island throughout the day. These birds
would have been indistinguishable from incubating
birds in the photographs.

Projections of color transparencies (Kodak ekta-
chrome 160), taken at 300 m above ground level with
the 200-mm lens, provided the best image. Larger
format cameras might have further improved resolu-
tion. Using the projections, we tallied 140 pelicans in
1976 and 70 in 1977.

Sixteen ground counts from 8 to 13 June in 1977
(Table 3) revealed a mean of 70.1 (range 59-75; SD
3.9) birds on nests. The 1977 aerial count was thus
accurate, but this was fortuitous, as ground checks
revealed.

The proportion of pelicans not clearly visible in
ground counts varied from 0 to 21% of the total
numbers present on the island. Adjustment as out-

TABLE |—Timing of annual cycles at several Canadian White Pelican colonies

Cessation of

Spring Autumn
Colony Year arrival Incubation brooding departure Source
Stum Lake, Mid-May- Campbell and
British Columbia 1964 early June Frost 1969
Birch Lake, Alberta 1976 Late April May-June June-July September This report
1977 Late April- May-June July September-__ This report
late May October
Beaverhill Lake, Alberta 1975 3-31 May Did not Early Weseloh et al. 1975
breed September
Lavallee Lake, 1975 Late April- May-June June-July August- Trottier et al., 1980
Saskatchewan May September
Dog Lake, Manitoba 1965 May-June Anderson and
Bartonek 1967
Pelican Lake, Manitoba 1964 Late April- May-June June-July Anderson and
May Bartonek 1967
East Shoal Lake, Manitoba 1964, May June-July Hosford 1965
1967,
1971 May June-July Evans 1972
Lake of the Woods, Ontario June-July Lies and Behle

1966

Vol. 95

THE CANADIAN FIELD-NATURALIST

200

‘poled sw) SuLNp spew sjunod jo 1aquinu = Ny,

(€1-1) 09 (€L-$9) O'0L 9 C 00:7c—-00:91
(0r-0) $6 (€L-6S) 8°89 ¢ € 00:91—-00:01
(St OTH (SL-89) PIL S Z 00:01-00:90
(adue1) UvauI (asue1) ueaw WN poyxseyo shep potod owiy
‘sloyeoy Jo Jaquinyy ‘s10}BQNdUI JO JaquinN jo Jaquiny

LL61 UNE E] 03 g Woy Aep ayy Jo sinoy
JUIIOJJIP 1B PaONpuod sjunod Zulinp Auojoo axe] Yosrg ay} Ie suKdTTad 9314 MQ SUNeQnoU! puke Zulyeo] Jo SioquinN—¢ aTEV_L

“LL61 Ul ABT] YOM 12 sJUsAa JO UOTVAIASGO [ene Aq Pajepl[eA J19M S9JBUIISA IS9Y | ‘P OE INOGe S}se] UOTeQndUT yey} UOTIBAIOSGO (¢/6]|) SJdouy pue
uedaq Apnjs uaym payoiey Apeadye (p96] J9][B4OS) SuNO< Jo sonstiajoe1eYo [eJUaWIdo[aAap Woy SUNePYIeG WOIJ POALJap So1eUII]Sa UO pase 1k 9/6] Ul s$31eq ,
‘Sp1lq Suljeqnoul Jo sasnsuad [ellae WOIJ Pautwsa}0q ,

“reak 0} 1e9A WI stun
Auojoogns Jo Aytadajut ayy A[duat Ay1essadau Jou Op puke spiig Bulpaaig jo sdnoi3 payesedas Aj[e1odui9} 10 / pure Ayjeneds 0} pauisse 219M suoleusisap Auojoogns,

A[N¢ 97-S VN A[ng p-oune p VN oune p-AeW 8Z a AN 8 VN q

AIne €7-Z A[n¢ ¢{-ounr ¢z Ajng [-oung | Ajng [-oung | aung |-Aey 97 aune | -Aey SZ Il v d

A[nf 97-S GN Aine p-oune p dN oun p-AeW CZ dN €l €l 5)

Ainge /Z-oune 97 GN oune ¢Z7—AeW 97 dN ABW 97-81 dN Ol €¢ a

Ajng ¢{]-oung 7Z ~=—s oun ¢Z-p aune [Z7-AeW ZZ aune g-Aey ABW 7-11 Aew p-udy 1z 8Z OL Vv

LL61 ,9L61 LL61 ,9L61 LL6I ,9L61 LL61 9161 Auojoogns
sulpooig uoneqnouy diysjinog qo}S9u JO JOQUINN]

Byep ou ‘CN ‘o[qeordde jou ‘WN *LZ6] Puke 9/6] Ul Oye] Yosg 1e Busou suvoyag aIYA\ JO AZO[ouoIYO pue ozis AuOjooqng—Z alaV _L

1981

lined in the Methods gave a mean of 11% loafing birds
(range 0-40%; SD 12%) from 8 to 13 June in 1977. The
proportion of loafing birds appeared to decline
throughout the day and was least variable in late
afternoon/ evening based on 16 counts (Table 3).

Estimates of numbers of nesting White Pelicans
derived from aerial photographs taken during the
peak incubation period thus appeared to be poten-
tially variable owing to the presence of up to 40%
loafing pelicans. A ground count is thus deemed more
accurate than an aerial census of nesting pelicans. The
former, if conducted by distant observation where the
nesting areas are clearly visible, would eliminate dis-
ruptive intrusions in the colonies.

In summary, from this study we conclude that aerial
censuses to count incubating pelicans should be con-
ducted during late May or early June, the peak incu-
bation period in Canada. Further we suggest that
evening census flights might be more accurate than
morning flights, as the proportions of loafing birds
appear to be lower in the evening. Concurrent ground
counts and census flights at other locations, however,
are desirable to validate the potential biases suggested
here.

Acknowledgments

We gratefully acknowledge financial assistance by
the Alberta Oil Sands Environmental Research Pro-
gram for this study. We thank D. Ealey, K. Zurfluh,
A. Richards, D. Richards, and M. Ballantyne for
assistance with the field work. C. Boyle and D. Hadler
(field operations staff, Alberta Oil Sands Environ-
mental Research Program) provided valuable logistic
assistance. Our appreciation is conveyed to E. Kuyt,
L. Allan, and G. Trottier of the Canadian Wildlife
Service for their helpful criticism of earlier drafts of
this manuscript.

Literature Cited

Anderson, D. W., and J. C. Bartonek. 1967. Additional
observations on the status of North American White Peli-
cans. Condor 69: 311-313.

Behle, W. H. 1958. The bird life of Great Salt Lake. Uni-
versity of Utah Press, Salt Lake City. 203 pp.

BEAVER AND LEWIN: WHITE PELICAN CENSUSES, ALBERTA

201

Boeker, E. L. 1972. A survey of White Pelican nesting col-
onies in 1972. American Birds 26: 24, 125.

Campbell, R. W., and D. L. Frost. 1969. Additional notes
on the White Pelican Colony at Stum Lake, British
Columbia. Condor 71: 73.

Carson, R. D. 1966. Destruction of colonial birds on an
island on Suggi Lake. Blue Jay 24: 96-97.

Evans, R. M. 1972. Some effects of water level on the
reproductive success of the White Pelican at East Shoal
Lake, Manitoba. Canadian Field-Naturalist 86: 151-153.

Farley, F. L. 1919. The White Pelicanin Alberta. Canadian
Field-Naturalist 33: 38-39.

Hosford, H. 1965. Breeding success of the WhitePelican in
two colonies in Manitoba in 1964. Blue Jay 23: 21-24.
Johnson, R.F., and N.F. Sloan. 1976. The effects of
human disturbance on the White Pelican colony at Chase
Lake National Wildlife Refuge, North Dakota. Inland

Bird Banding News 48: 163-170.

Knopf, F. L. 1975. Spatial and temporal aspects of colonial
nesting of the White Pelican, Pelecanus erythrorhynchos.
Ph.D. thesis, Utah State University, Logan, Utah. 76 pp.

Lies, M. F., and W. H. Behle. 1966. Status of the White
Pelican in the United States and Canada through 1964.
Condor 68: 279-292.

Sanderson, R. M. 1966. The colonial birds at Suggi Lake,
Saskatchewan, in 1966. Blue Jay 24: 121-123.

Schaller, G. M. 1964. Breeding behavior of the White Peli-
can at Yellowstone Lake, Wyoming. Condor 66: 3-23.
Strait, L. E. 1973. Population dynamics of a White Pelican
population, Chase Lake Wildlife Refuge, North Dakota.
M.Sc. thesis, Michigan Technological University, Hough-

ton, Michigan. 76 pp.

Trottier, G.C., R.J. Breneman, and N.A. Young.
1980. Status and foraging distribution of White Pelicans,
Prince Albert National Park, Saskatchewan. Canadian
Field-Naturalist 94(4): 383-390.

Vermeer, K. 1969. Colonies of Double-crested Cormorants
and White Pelicans in Alberta. Canadian Field-Naturalist
83: 36-39.

Vermeer, K. 1970. Distribution and size of colonies of
White Pelicans, Pelecanus erythrorhynchos, in Canada.
Canadian Journal of Zoology 48: 1029-1032.

Weseloh, D. V., D. Dekker, S. Brechtel, and R. Burns.
1975. Notes on the Double-crested Cormorants, White
Pelicans and Great Blue Herons of Beaverhill Lake,
summer, 1975. Alberta Naturalist 5: 132-137.

Received 10 March 1980
Accepted 11 November 1980

Notes

Nesting Northern Gannets (Morus bassanus) Killed by
Rock Falls at Great Bird Rock, Quebec

IAN R. KIRKHAM

Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland AIB 3X9

Kirkham, Ian R. 1981. Nesting Northern Gannets (Morus bassanus) killed by rock falls at Great Bird Rock, Quebec.

Canadian Field-Naturalist 95(2): 202-203.

Nine Northern Gannet (Morus bassanus) adults and six eggs were crushed by a falling rock at Great Bird Rock. Falling rocks
and related seabird mortality are probably common at this colony as the islands are slowly eroding away.

Key Words: rock falls, Northern Gannets (Morus bassanus), Great Bird Rock, erosion, accidental mortality.

During my study of Northern Gannets (Morus bas-
sanus) on Great Bird Rock (47° 50’N, 61°09’W), Mag-
dalen Islands, Quebec, in May 1979, I saw two dead
adult gannets floating in the water at the base of the
eastern cliffs. As I climbed down the cliff to collect the
dead birds, I saw that a large rock (0.3 X 1.5 X 2.5 m)
had broken away, falling about 3 m and coming to
rest on a group of incubating gannets. Four dead
adults were visible on their nests. Given the local
nesting density, it is likely that at least two more nests
were destroyed beneath the rock. An estimated nine
adults were killed and six eggs destroyed, including
the two birds on the water and a dead adult on the
shore below the rock fall. Lightkeepers gave me
accounts of other rock falls which killed gannets. In
1973, 24 dead gannets were in the water near the
northeast point of the island where part of the cliff had
collapsed on a group of nests (D. McLean, Entry
Island, Magdalen Islands, P.Q. GOB 1C0, personal
communication). In May 1978, W. Dixon (Entry
Island, Magdalen Islands, P.Q. GOB 1CO, personal
communication) saw a large number ( > 30) of dead
gannets along the northern shore of the island and in
the water beneath a nesting area where a large over-
hanging section of the top shelf had given way. Along
the northern end of the island, rock slides are frequent
(D. McLean, personal communication) and during
my 26-d stay a slide there destroyed about 20 nests.

The Bird Rocks were possibly once the largest gan-
netry in the world, estimated at about 112 000 pairs in
1833 (Gurney 1913). Recently this colony was esti-
mated at 5331 pairs (Nettleship 1976). The two
obvious reasons for the population decline are (1) the
erection of a lighthouse in 1869, which discouraged
possibly 50 000 pairs from nesting during the next 20
years (Lucas 1888), and (2) the reduction of breeding
sites through erosion of the islands. Ina short period

of time, Little Bird Rock has apparently eroded
greatly. A third island, located on the shoal between
the two Bird Rocks, was seen by Cartier in 1534 but
disappeared before the 19th century (Lucas 1888). The
Bird Rocks are remnants of a once extensive rock
formation and consist primarily of red sandstone
which is fine-grained and poorly consolidated. They
have a weathered appearance due to the small
amounts of calcite which have mostly eroded away
and left a porosity of close to 45% (Sanschagrin 1964).

Most of the mortality due to rock slides at breeding
sites would be expected to occur during spring thaw
which causes multiple fractures and disintegration of
rock. Reports of deaths related to rock falls among
gannets appear to be confined to the Bird Rocks,
except for a small colony on Grimsey, Iceland, that
disappeared owing to falling cliff material above nest-
ing sites (Gudmundsson 1953). The long-term effect of
the erosion and rock slides at Bird Rocks will be a
decline in the number of breeding birds, owing to a
reduction in the number of safe nesting sites, and
eventually one of the oldest and once the largest gan-
netry in the world will disappear.

I thank the lightkeepers of Great Bird Rock for
providing information about the gannets, Chris-
topher Lynas for reviewing the geological reports, and
William Montevecchi for reviewing the manuscript.
Support was provided by a Canadian Wildlife Service
student scholarship and a NSERC grant A0687 to
W. A. Montevecchi.

Literature Cited

Gudmundsson, F. 1953. Islenzkin fuglar VII Sula (Sula
bassana (L.)). Natturufraedingurinn 23: 170-177.

Gurney, J. H. 1913. The gannet. Witherby and Company,
London. 567 pp.

202

1981

Lucas, F. A. 1888. The Bird Rocks of the Gulf of St. Law-
rence in 1887. Auk 5: 129-135.

Nettleship, D. N. 1976. Gannetsin North America: present
numbers and recent population changes. Wilson Bulletin
88: 300-313.

Sanschagrin, R.

NOTES 203

1964. Geological Report 106, Magdalen
Islands. Department of Natural Resources, Quebec. 58 pp.

Received 21 May 1980
Accepted 27 August 1980

Postglacial Fossil Fishes from Coppermine River,

Northwest Territories, Canada

DON E. MCALLISTER! and DENIS ST-ONGE?

\Ichthyology Section, National Museum of Natural Sciences, Ottawa, Ontario KIA 0M8
2Département de Géographie, Université d’Ottawa, Ottawa, Ontario KIN 6NS5

McAllister, Don E. and Denis St-Onge. 1981. Postglacial fossil fishes from Coppermine River, Northwest Territories,
Canada. Canadian Field-Naturalist 95(2): 203-205.

Fossils remains referred to Arctic Charr (Sa/velinus alpinus), Arctic Grayling (Thymallus cf. arcticus), Northern Pike (Esox
cf. Jucius), and Slimy Sculpin (Cottus cognatus), were collected from freshwater, sandy-silt deposits at 66° 50’N, 115°55’W.
These may provide the earliest yet known Northwest Territories records of fishes originating from the Mississippian or
Beringian refugia approximately between 8400 and 9000 years BP.

Key Words: fossil fishes, Coppermine River, Northwest Territories, Sa/velinus alpinus, Thymallus arcticus, Esox lucius,
Cottus cognatus, Pleistocene.

McAllister, Don E. and Denis St-Onge. 1981. Postglacial fossil fishes from Coppermine River, Northwest Territories,
Canada. Canadian Field-Naturalist 95(2): 000-000.

Des débris fossils d’?Omble Arctique (Sa/velinus alpinus), d’ Ombre Arctique (Thymallus cf. arcticus),de Grand Brochet (Esox
cf. Jucius) et de Chabot Visqueux (Cottus cognatus) furent trouvés dans un dépét de limon sableux a 66° 50’N, 115°55’W. Ces
restes représentent peut-étre les témoins les plus anciens de poissons en provenance des refuges Mississipien ou Beringien des
Territoires du Nord-Ouest approximativement entre 8400 et 9000 ans AP.

Mots clefs: Poissons fossils, Riviere Coppermine, Les Territoires du Nord-Ouest, Sa/velinus alpinus, Thymallus arcticus,

Esox lucius, Cottus cognatus, Pli€istocéne.

In this note we report on post-Wisconsin fish
remains found in the valley of the Coppermine River
at 66°50’N, 115°55’W, north-central District of
Mackenzie, Northwest Territories. The Quaternary
fish fauna of Canada is poorly known and Harington
(1978) reports only two fossil fish collections for the
Northwest Territories, and fishes from one of these
have not been identified.

The fish remains occurred in an organic silty-sand
unit, | m thick, at the top of a section which forms a
61-m-high bluff on the north side of the Coppermine
River. Most of the outcrop is composed of outwash
(sand, gravel, and drop stones) resting onan irregular
till surface (boulders in sand-gravel matrix). These
sediments are part of a hummocky terrain within the
valley that are capped, in part at least, by the fossilif-
erous silty sand. A perched delta, 300 m upstream,

marks the level of a major glacial lake which formerly
occupied the valley (Glacial lake Coppermine, St-
Onge, in press). The l-m-thick organic silty sand in
which the bones were found appears to be river flood
deposits which occur at an altitude similar to that of
the delta.

Another major section, 30 m high, is exposed along
a small stream (Quicksand Creek, 66°49’20’N,
116°21’W), 15 km downstream from the high bluff. It
shows three major units: bouldery till at the base;
22-25 m of varved sandy silts and clays covering the
bouldery till; cross-bedded deltaic sands, 5-8 m thick
which overlie the varves. In the deltaic sands is found a
former channel filled with silty sand, rich in organic
debris, and capped by a dense peat bed. The channel
sequence is about 4 m thick. A piece of willow (Salix
sp.) at the base of the channel fill has yielded an age of

204

8400 + 80 (GSC-2959) while the peat at the top of the
fill gave an age of 3210 + 60 (GSC-2998).

National Topographic Series maps recently pub-
lished (1979) by the Canadian Department of Energy,
Mines, and Resources ata scale of 1:50 000 show that
the top of both sections is at approximately 320 masl.
What then is the relationship between the fossil fish
bed and the dated units in the Quicksand Creek sec-
tion? It is difficult to imagine water being present at
the top of the hummocky terrain unless it was part of
Glacial Lake Coppermine. Because the fossiliferous
bed occurs at an elevation corresponding to the high-
est level of the lake, it is reasonable to assume that the
fish were alive during that period of maximum lake
elevation, probably 8400-9000 years BP. This deduc-
tion has considerable paleoecological implications
and will require substantiation by radiocarbon dating
of organic material from the fossiliferous beds.

The fish remains studied consisted of numerous
bones and scales, little eroded, in fine silty sand. All
the bones and scales apparently originated from small
specimens about 120 mm in standard length.

Species of Fossil Fishes

Four taxa of fishes were determined. From the
bones identified, the relative frequency of taxa, with
the most frequent first, seemed to be Thymallus, Esox,
Salvelinus, and Cottus; the taxonomic position of
many other bones difficult to determine were not iden-
tified. Catalogue numbers are those of the Paleobiol-
ogy collection, National Museum of Natural Sciences,
Ottawa.

Salvelinus alpinus, Arctic Charr

A single dentary (NMC 36122), 8.0 mm long, with
the tips of the larger teeth hooked and pointing poste-
riorly, was identified as Arctic Charr. Jaw and tooth
form distinguish this fossil from the Lake Charr
(= Lake Trout, Sa/velinus namaycush).

Harington (1978) did not list Sal/velinus alpinus
amongst the known Quaternary fishes of Canada and
Alaska, so this specimen provides the first fossil
record for this species in North America.

Thymallus c.f. arcticus, Arctic Grayling

Several small scales (NMC 36123) about 1.8 mm in
diameter from juveniles of this species with their char-
acteristic crenulate border (about five crenulations)
were found. Among Canadian arctic fishes the North-
ern Pike (Esox /ucius) is the only other species with
crenulate scales, but the scales of pike have grooves
running posteriorly from the crenulations, a feature
lacking in these specimens.

We refer them to 7. arcticus, rather than one of the
Palearctic species, solely on the basis of provenance.
Pleistocene 7. arcticus have been reported from the
Yukon Territory by McAllister and Harington (1968).

THE CANADIAN FIELD-NATURALIST

Vol. 95

Esox c.f. lucius, Northern Pike

This species is represented by a few small dentaries
(NMC 36124) about 11 mm long with teeth character-
istic of the genus.

The fossils are identified as E. /ucius instead of E.
niger, E. americanus, or E. masquinongy, because it is
unlikely that these three species would be found in the
cool northern waters so soon after deglaciation. Evi-
dently the Coppermine was deglaciated about 9500
BP. Esox lucius has been reported from late Pleisto-
cene deposits in the Yukon Territory (Crossman and
Harington 1970).

Cottus cognatus, Slimy Sculpin

The species was represented by a single almost
complete preoperculum (NMC 36125). It is identified
as Cottus cognatus rather than Cottus ricei because
the spine forms a V with the body of the bone instead
of a U. The bone is about 6.4 mm long, the spine
1.2 mm long. S. L. Cumbaa, D. E. McAllister, and
R. E. Morlan have discovered a vertebral centrum
attributable to Co/tus from Pleistocene deposits in the
northern Yukon Territory.

Discussion

The fossils bear significantly on the timing of dis-
persal of these species. All four had refugia in both the
Beringia and the Mississippi system. Pike probably
entered the upper Mackenzie from the Mississippi
(McPhail and Lindsey 1970), hence the Coppermine
population probably reached the area from the same
source. These fossil pike thus fix the earliest known
date for fish of Mississippian origin to have arrived in
arctic Canada.

Slimy Sculpins presently in the Coppermine area
appear to be intergrades between populations which
survived glaciation in Beringian and Mississippian
refugia (McAllister and Lindsey 1961), although
closer in anal ray ‘branching’ to the form that survived
in Beringia. Unfortunately the preoperculum does not
have characters known to assign it to one of those
forms. However, one can say that very shortly after
the glacier had left the Coppermine area, Cottus cog-
natus from one of these refugia had already entered
the region. Arctic Charr in the Coppermine area today
display a mixture of forms which survived in Beringia
and the Atlantic coastal plain (McPhail and Lindsey
1970).

The bones are all derived from small individuals
thus suggesting shallow water. Possibly the fish were
caught in a drying sandy pool near the margin of a
river or lake, or an eroding sandy river bank collapsed
into shallow waters. The four species are freshwater
dwellers; none except for the Arctic Charr are
anadromous.

The fine state of preservation of the bones and their

1981

high density in the matrix suggest that further
research on the fauna would be worthwhile. Addi-
tional species of, for example, Salmonidae, Catosto-
midae, or Gadidae may be represented. Occurrence of
adult scales or operculas may reveal growth patterns
and life cycles.

L. D. Farley-Gill, Quaternary Paleoecology Lab-
oratories, Geological Survey of Canada, kindly iden-
tified a piece of wood as willow. We thank the
National Museum of Natural Sciences scientists Ste-
phen L. Cumbaa for confirming the identification of
the Arctic Charr dentary and C. Richard Harington
for criticizing the manuscript.

Literature Cited
Crossman, E. J., and C. R. Harington. 1970. Pleistocene
pike, Esox /ucius and Esox sp. from the Yukon Territory

NOTES

205

and Ontario. Canadian Journal of Earth Sciences 7:
1130-1138.

Harington, C. R. 1978. Quaternary vertebrate faunas of
Canada and Alaska and their suggested chronological
sequence. Syllogeus, Ottawa 15: 1-105.

McAllister, D. E., and C. R. Harington. 1969. Pleistocene
Grayling, Thymallus, from Yukon, Canada. Canadian
Journal of Earth Sciences 6: 1185-1190.

McAllister, D. E., and C. C. Lindsey. 1961. Systematics of
the freshwater sculpins (Cottus) of British Columbia.
National Museums of Canada Bulletin 172: 66-89.

McPhail, J. D.,and C. C. Lindsey. 1970. Freshwater fishes
of northwestern Canada and Alaska. Fisheries Research
Board of Canada Bulletin 173: 1-381. 3

St-Onge, D. A. 1980. Glacial Lake Coppermine, north-
central District of Mackenzie, Northwest Territories.
Canadian Journal of Earth Sciences 17: 1310-1315.

Received 30 June 1980
Accepted 12 September 1980

Gulls Robbing Prey from Great Blue Herons (Ardea herodias)

T. E. QUINNEY,! B. N. MILLER, and K. R. S. QUINNEY

Department of Biology, Acadia University, Wolfville, Nova Scotia BOP 1X0
'Present address: Department of Zoology, University of Western Ontario, London, Ontario N6A 5B7

Quinney, T. E., B. N. Miller, and K. R. S. Quinney. 1981. Gulls robbing prey from Great Blue Herons (Ardea herodias).
Canadian Field-Naturalist 95(2): 205-206.

Three of five attempts by gulls to steal prey from adult Great Blue Herons (Ardea herodias) during 80 h of observation were
successful. Over half of the aggressive actions involving adult herons were directed at gulls. Prey stolen were always longer
than 140 mm. The handling times of these items indicates that herons have difficulty swallowing such large prey and thus
become susceptible to piracy.

Key Words: Great Blue Heron (Ardea herodias), Great Black-backed Gull (Larus marinus), kleptoparasitism, handling time.

Kleptoparasitism, or piracy, is well documented in
birds (Brockman and Barnard 1979). Gulls are com-
monly described as pirates (Lockley 1953; Meinertz-
hagen 1959; Hatch 1970, 1975; Morrison 1978), but
Great Blue Herons (Ardea herodias) have seldom
been described as victims of piracy (Carlsson 1979).

Gull-heron interactions were recorded during a
study of the foraging behavior of Great Blue Herons
(Quinney and Smith 1980). Herons on their foraging
grounds in three estuaries of the Minas Basin, Nova
Scotia (45° 08’N, 65° 16’W), were watched for 80 h 24
July - 17 August 1978. Up to 17 adult and newly
fledged juvenile herons foraged concurrently in the
numerous tidal pools and streams exposed during low
tide in these estuaries. Great Black-backed Gulls
(Larus marinus) and Herring Gulls (L. argentatus)

were usually present and spent considerable time
apparently loafing along the stream banks or on mud
bars.

Adult herons performed inter- and intra-specific
aggressive actions on 21 occasions (not involving cap-
tured prey). Eleven such events resulted when Great
Black-backed Gulls attempted to approach too
closely to feeding herons. Black-backs stole prey, in all
cases Flounder (Liopsetta putnami) longer than
140 mm, from adult herons on three occasions. This
size approximates the length of a male heron’s bill
(Godfrey 1966, p. 36).

The handling time of large (over 105 mm) com-
pared with smaller prey indicates that adult herons
have difficulty swallowing large prey (large: 52s,
n= 12, vs. smaller: 7 s,n = 114). Only 5% (13/279) of

206

all prey captured by adult herons were large. When a
heron captured a large prey item and did not swallow
it immediately, one or more gulls often quickly flew in.
If the gulls came within 2 m, the heron partially
extended its wings and lunged at the gulls, still holding
the captured prey inits bill. The gulls retreated imme-
diately about 5 m but continued to watch. If the prey
was dropped during mandibulation, a gull often flew
in, stole the prey, and flew away followed by other
gulls. Conflict over the stolen prey often ensued. After
a loss, the heron vocalized and lunged toward the thief
for a few steps, but never flew in pursuit. An adult
‘Herring Gull attempted twice to steal a large fish from
a heron in flight, but failed.

The relatively low frequency of interspecific aggres-
sive interactions and prey robbing involving gulls and
herons indicates that the effect upon heron foraging is
minor. Prey robbing by gulls appears limited to large
prey that are captured infrequently and handled with
difficulty by Great Blue Herons.

We thank P. C. Smith, J. A. Kushlan, R. D. Bayer,
and J.-L. Des Granges for their comments on the
manuscript. This study was supported by National
Research Council of Canada grant A0236 to P. C.
Smith, Canadian Wildlife Service, University

THE CANADIAN FIELD-NATURALIST

Vol. 95

Research Support Fund, and Federal Summer Job
Corps project No. 16-01-003.

Literature Cited

Brockman, H. J., and C. J. Barnard. 1979. Kleptoparasit-
ism in birds. Animal Behaviour 27: 487-514.

Carlsson, Le. 1979. Trutar som naringsparasit pa hager.
Vaar Fagelvarld 38: 50.

Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203. 428 pp.

Hatch, J. J. 1970. Predationand piracy by gulls at a ternery
in Maine. Auk 87: 244-254.

Hatch, J.J. 1975. Piracy by Laughing Gulls (Larus atri-
cilla): an example of the selfish group. Ibis 117: 357-365.

Lockley, R. M. 1953. Puffins. J. M. Dent and Sons Ltd.,
London. 186 pp.

Meinertzhagen, R. 1959. Pirates and predators. Oliver and
Boyd, Edinburgh. 230 pp.

Morrison, R. I. G. 1978. Herring Gulls stealing prey from
Parasitic Jaegers. Wilson Bulletin 90: 649-650.

Quinney, T. E.,and P. C.Smith. 1980. Comparative forag-
ing behaviour and efficiency of adult and juvenile Great
Blue Herons. Canadian Journal of Zoology 58: 1168-
1173.

Received 6 March 1980
Accepted 8 September 1980

First Record for Canada of the Bat Mite Spinturnix globosus

and a New Host, Myotis lucifugus'

HUGH C. SMITH

Provincial Museum of Alberta, 12845— 102 Avenue, Edmonton, Alberta T5N 0M6

Smith, Hugh C. 1981. First record for Canada of the bat mite Spinturnix globosus and a new host, Myotis lucifugus.

Canadian Field-Naturalist 95(2): 206-207.

The first Canadian record for the bat mite Spinturnix globosus is documented from Little Brown Myotis, Myotis lucifugus, at
four Alberta nursery colonies. This is the first report of the mite’s occurrence on Myotis lucifugus.

Key Words: Spinturnix globosus, Myotis lucifugus, first record, host.

During the preparation of a series of Little Brown
Myotis, Myotis lucifugus, as study skins, mites were
noted in and around the anal orifice of several indi-
viduals, Five mites were collected and forwarded to
the Biosystematics Research Institute, Agriculture
Canada, Ottawa, for identification. There, Evert E.
Lindquist identified these mites as adult female Spin-

‘Provincial Museum of Alberta Natural History Contribu-
tion 59.

turnix globosus. They represent the first record for
this mite in Canada and the first time that Myotis
lucifugus has been reported as a host (Whitaker and
Nixon 1974). Other locality records for this mite are
from Tennessee, Indiana, Utah, Arizona, Oklahoma
(Rudnick 1960), Kansas (Ubelaker 1966), and Oregon
(J. O. Whitaker, Jr., Indiana State University, per-
sonal communication). This mite has been reported
on various species of bats: Myotis sodalis (Indiana
Myotis), Myotis velifer (Cave Myotis), and Myotis

1981

grisescens (Gray Myotis) by Rudnick (1960), and
Myotis volans (Long-legged Myotis) by Whitaker
(personal communication).

Spinturnix globosus was originally described by
Rudnick (1960) and was placed in the genus Paraspin-
turnix. Domrow (1972) subsequently synonomized it
into its present genus Spinturnix. Spinturnix globo-
sus, in common with all other members of the family
Spinturnicidae, is exclusively parasitic on bats (Rud-
nick 1960). Rudnick (1960) noted that only the adult
female of S. globosus has been recovered, and only in
the anal orifice. But Ubelaker (1966) reported that this
mite was obtained from the penis of a male and from
the vagina of females of M. grisescens.

Spinturnix globosus were recovered from bats that
were collected at four nursery colonies in Alberta in
July 1978; and no attempt was made to obtain any
specific age or sex cohort. No systematic search for
parasites was made on all bats collected. The number
of mites per bat ranged from one to three (Table 1).

TABLE 1—Incidence of infection with the mite Spinturnix
globosus at four nursery colonies of Little Brown Myotis
Myotis lucifugus, in Alberta (All female bats were adults; the
male bat was a juvenile)

Number of bats

Location Examined Infected Mites/bat
52° 29’N-112°11’W 48 1Q ]

8Q 2

59 3
54°01’N-110°54’W 21 19 !
54° 10’N-110° 56’W 8 19 l

1Q 2
54° 11’N-110°48’W 33 1d I

1Q 1

1Q 2

NOTES

207

Ubelaker (1966) reported that he found no more than
three mites per host.

Bat ectoparasites, especially in Canada, are rela-
tively poorly known. Rudnick (1960) suggested that
the role played by blood-sucking ectoparasites should
be included in the study of bat diseases. Recently in
Alberta, as part of an overall rabies control program
(see Dorward et al. 1977 re-rabies in Alberta), there
has been a concerted effort to study the life history of
bats. To my knowledge no studies of ectoparasites of
these bats have been carried out.

I thank D. Schowalter for collecting the bats, Lorne
Duncan and Anne Allen for preparing the specimens
as study skins and for observing and collecting the
mites, Evert E. Lindquist for identifying the mites and
supplying requested literature references, and John
Whitaker for bringing several pertinent references to
my attention.

Literature Cited

Domrow, R. 1972. Acari Spinturnicidae from Australia
and New Guinea. Acarologia 13: 552-584.

Dorward, W.J., D. B. Schowalter, and J. R. Gunson.
1977. Preliminary studies of bat rabies in Alberta. Can-
adian Veterinary Journal 18: 341-348.

Rudnick, A. 1960. A revision of the mites of the family
Spinturnicidae (Acarina). University of California Publi-
cations in Entomology 17: 157-284.

Ubelaker, J. E. 1966. Parasites of the Gray Bat, Myotis
grisescens, in Kansas. American Midland Naturalist 75:
199-204.

Whitaker, J. O., Jr., and N. Wilson. 1974. Host and distri-
bution lists of mites (Acari), parasitic and phoretic, in the
hair of wild mammals of North America, north of Mexico.
American Midland Naturalist 91: 1-67.

Received 24 June 1980
Accepted 3 November 1980

208 THE CANADIAN FIELD-NATURALIST

Vol. 95

Estimating Winter Defecation Rates for Moose, Alces alces

JOHN L. OLDEMEYER! and ALBERT W. FRANZMANN?2

'Denver Wildlife Research Center, U.S. Fish and Wildlife Service, 1300 Blue Spruce Drive, Fort Collins, Colorado 80524
2Alaska Department of Fish and Game, Soldotna, Alaska 99669

Oldemeyer, John L.,and Albert W. Franzmann. 1981. Estimating winter defecation rates for Moose, Alces alces. Canadian

Field-Naturalist 95(2): 208-209.

The pellet-group count technique was tested for 2 yr ina 1.6-km? enclosure with known numbers of Moose (Alces alces) on the
Kenai National Moose Range, Alaska. Pellet-groups were cleared from one hundred and sixty 17.8-m? quadrats each fall and
counted and cleared from those same quadrats in the spring. Clearing of permanent pellet-group plots in fall is necessary for
an accurate estimate of Moose numbers-and winter use. In winter 1976-1977, seven Moose inhabited the enclosure and
deposited a daily average of 16.2 pellet-groups per Moose. The next winter, six Moose deposited a daily average of 17.2

pellet-groups per Moose.

Key Words: Moose, Alces alces, Alaska, pellet-group counts, population size.

The pellet-group count technique (Neff 1968) has
been used for estimating Moose (A/ces alces) numbers
and determining habitat preferences since the estab-
lishment of the Kenai Moose Research Center
(MRC), Alaska, in 1968 (LeResche and Davis 1973;
Franzmann et al. 1976b). They reported, however,
that May counts of permanent plots in a 1.6-km?
enclosure at MRC with known numbers of Moose
resulted in unusually high daily defecation rates (20.2
groups per Moose in 1970-1971 to 28.7 groups per
Moose in 1973-1974). To lend support to the apparent
bias in the plot counts, Franzmann et al. (1976a)
followed eight individual Moose during winter 1975,
when snow cover allowed accurate counts of daily
defecations, and found the daily defecation rate (and
range) for adult cows was 14.6 (10-22) groups, that for
adult bulls was 19.6 (14-25) groups, and rates were
significantly different (P < 0.01) between sexes.

We assumed that the daily defecation rates from the
winter-trailed Moose was a more accurate estimate of
Moose defecation rates, even though the plot counts
had been consistent over the years in their random
distribution of groups within vegetative types and in
relative Moose use among types. In the plot counts,
the only variable not taken into consideration was
summer-deposited pellet-groups. The objective of this
study was to determine if the daily defecation rates
obtained when plots were cleared in fall (to eliminate
summer-deposited pellet-groups) and counted and
cleared in spring were similar to the rates obtained by
winter-trailing Moose.

Methods

In 1969, 160 permanent 17.8-m? (8 X 24 ft) plots
were randomly located in the seven important upland
vegetation types in the 241-ha Enclosure 1 at MRC.
The plots were used for estimating plant utilization
and for pellet-group surveys (R. Bishop. 1969. Moose

Report. Alaska Department of Fish and Game 153
pp. (mimeo.)). Eighteen to 26 plots were located in
each vegetation type without regard to type size.
These 160 plots represented about 0.1% of the enclo-
sure. Through spring 1976, pellet-groups were
counted and removed from the plots in early May.

All plots were cleared of defecations in September
1976 before we began this study. Plots were counted
and cleared in May 1977, cleared in September 1977,
and counted and cleared in May 1978.

The counting procedure followed was as follows: A
7.3-m (24-ft) rope was strung between the two stakes
placed at midpoints of the end of each plot. A 1.2-m
(4-ft) width was walked oneach side of the rope and in
each direction so that the plot was counted twice. All
pellet-groups in which the group center, or at least half
of the pellets fell within the plot, were counted and
removed from the plot.

As part of other studies at MRC, the number of
Moose in the enclosures was closely monitored. In
Enclosure |, there were seven Moose providing 1722
Moose days between October 1976 and May 1977 and
six Moose providing 1308 Moose days from October
1977 to May 1978.

Analysis of the data followed Bowden et al. (1969)
in which the data were first tested for random deposi-
tion with the Poisson distribution.

Results and Discussion

The distribution of our data fit the Poisson distribu-
tioneach year forall vegetation types except one. This
implies random deposition of defecations within the
enclosure, which we would expect in this area because
snow is not deep enough to force yarding; topographic
differences are not distinct enough to result in prefer-
ential use; and even in dense populations Moose tend
to be solitary animals.

Analysis of variance of \/x+0.5 transformed

1981

counts (to equalize variance) comparing mean
number of pellet-groups per plot among types failed
to show significant differences; therefore, we used a
pooled total estimate of pellet-groups in the enclosure.
There were 1722 d of use by seven Moose during the
1976-1977 winter and they deposited an average
(+ 95% confidence interval) of 16.2 +5.78 pellet-
groups per Moose day. The next winter, there were
1308 Moose days and an average deposition rate of
17.2 + 7.76 pellet-groups per Moose day. These rates
are higher than most reported rates for Moose (Tim-
merman 1974) or for other ungulates (Neff 1968). The
results for the two winters, however, are within the
range (14.6-19.6) of that found by Franzmann et al.
(1976a) when individual Moose were trailed in snow
and represent the most controlled study of the tech-
nique for Moose defecation rates.

These rates are more realistic for Moose than those
reported earlier (Franzmannet al. 1976b) and demon-
strate the need to clear permanent pellet-group plots
in fall if estimating Moose winter use or numbers. Neff

(1968) has remarked on the lack of reliability of esti- —

mating age of pellet-groups, which reinforces the need
to clear plots before the season of interest.

As an additional test, we estimated the number of
pellet-groups Moose would have deposited in the
enclosure over winter by multiplying the known
number of male and female Moose days by the
number of defecations per day.

Based on the data of Franzmann et al. (1976a), we
calculated in winter 1976-1979 that there would have
been 25 232 groups deposited by three adult males and
three adult females. This value would leave 2791 pellet
groups to be deposited by one calf resulting ina rate of
11.4 groups per day. In winter 1977-1978 there would
have been 19 184 groups deposited by two males and
three females leaving 3310 groups to be deposited by
the one male yearling. The defecation rate for that
animal would have been 15.2 groups per day to reach
our sample estimated total of 22 494.

In summary, we have had an ideal opportunity to
test pellet-group surveys for estimating Moose popu-

NOTES 209

lations and habitat use. The estimated defecation rates
of 16.2 and 17.2 groups per Moose day were well
within the range observed for winter-trailed Moose
and appear to be precise estimates of winter defeca-
tion rates.

In addition when comparing our results with earlier
studies at MRC, we have demonstrated the need for
clearing pellet-group plots in both fall and spring if
one 1s to estimate winter use by Moose accurately. We
would expect this practice to be a requirement for
other western ungulates.

Acknowledgments

We thank C. J. Breckenridge. W. L. Regelin, and
C.C. Schwartz for their field assistance. Daniel
Welsh, Canadian Wildlife Service, provided construc-
tive criticism of the manuscript. The project was
funded in part by Federal Aid in Wildlife Restoration
Project W-17-R.

Literature Cited

Bowden, D.C., A. E. Anderson, and D. E. Medin.
1969. Frequency distributions of Mule Deer fecal group
counts. Journal of Wildlife Management 33: 895-905.

Franzmann, A. W., P.D. Arneson, and J. L. Olde-
meyer. 1976a. Daily winter pellet groups and beds of
Alaskan moose. Journal of Wildlife Management 40:
374-375.

Franzmann, A. W., J. L. Oldemeyer, P. D. Arneson, and
R. K. Seemel. 1976b. Pellet-group count evaluation for
census and habitat use of Alaskan moose. Twelfth North
American Moose Conference and Workshop. St. John’s,
Newfoundland. 16 pp.

LeResche, R.E., and J. L. Davis. 1973. Importance of
nonbrowse foods to Moose on the Kenai Peninsula,
Alaska. Journal of Wildlife Management 37: 279-287.

Neff, D. J. 1968. The pellet-group count technique for big
game trend, census, and distributions: a review. Journal of
Wildlife Management 32: 597-614.

Timmerman, H.R. 1974. Moose inventory methods: a
review. Naturaliste Canadien 101: 615-629.

Received 22 August 1980
Accepted 27 November 1980

210 THE CANADIAN FIELD-NATURALIST Vol. 95

First Canadian Records of the Ghost Shiner (Notropis buchanani)
and the Orangespotted Sunfish (Lepomis humilis)

ERLING HOLM and GEORGE A. COKER

Department of Ichthyology and Herpetology, Royal Ontario Museum, Toronto, Ontario M5S 2C6

Holm, Erling, and George A. Coker. 1981. First Canadian records of the Ghost Shiner (Notropis buchanani) and the
Orangespotted Sunfish (Lepomis humilis). Canadian Field-Naturalist 95(2): 210-211.

Ghost Shiners (Notropis buchanani) and Orangespotted Sunfish (Lepomis humilis), previously unknown from Canada, are
apparently well established in the Thames River and Cedar Creek, respectively, in southwestern Ontario. The record for
Ghost Shiners extends the known range into the Great Lakes drainage and 270 km north of the nearest record. The record for

Orangespotted Sunfish extends the known range 44 km north of the nearest record.

Key Words: Ghost Shiner, Notropis buchanani; Orangespotted Sunfish, Lepomis humilis, range extension.

Notropis buchanani, the Ghost Shiner, and Lepo-
mis humilis, the Orangespotted Sunfish, not pre-
viously reported from Canada, were captured in 1979
and 1980 in southwestern Ontario.

We caught nine N. buchanani on 3 June 1979 in
quiet and turbid backwaters of the Thames River,
near the mouth of Dolsen Creek, Kent County, Dover
Township (42°20'11”N, 082°23’40”W). The shiners
were seined from water less than | m in depth over a
bottom of clay, silt, and detritus. The specimens, six
females and three males, 36-43 mm TL, have been
added to the collection of the Royal Ontario Museum
(ROM 35794). Identification of these specimens has
been confirmed by R. M. Bailey, University of Michi-
gan Museum of Zoology (UMMZ).

Notropis buchanani is apparently common in the
lower Thames. On 12 June 1980, we captured 86 fish
at five of the six stations sampled from the Thames
River below Kent Bridge (42° 30’48”N, 082° 04’15” W).
At three locations, N. buchanani was more abundant
than any other species. These specimens are cata-
logued as ROM 36436-36440. An additional 15 spec-
imens were donated to UMMZ (207676).

The maximum size attained by N. buchanani,
according to Trautman (1957), is 64mm TL. The
largest individual we captured was a female, 51 mm
TL, full of eggs. Most of the males, the largest 43 mm
TL, were tuberculate (see Cross 1967 for best descrip-
tion of tuberculation).

The habitat in the Thames River was similar to that
described by Smith (1979) for this species, i.e., large
rivers with turbid water and a soft bottom of silt and
detritus. But Trautman (1957) stated that NV. bucha-
nani sought clear, quiet water with clean sand and a
gravel bottom, and that it was apparently intolerant of
turbidity and current.

Notropis buchanani is easily confused with the
Mimic Shiner, Notropis volucellus, but differs in that
N. buchanani rarely possesses an infraorbital canal,

has more highly elevated anterior lateral line scales,
and differs in pigmentation (see Bailey 1956 for best
key).

Notropis buchanani was previously known to occur
only in the Mississippi drainage. The new record is 270
km north of the nearest record in Ohio (Trautman
1957). Smith (1979) recorded the Ghost Shiner in the
upper Illinois River near Morris, Illinois. The species
has access to the Great Lakes via the Chicago Ship
Canal, but there is apparently no record of this species
in the Great Lakes system of Illinois or adjacent states
(see Gilbert 1980). The Illinois record is 510 km west
of the mouth of the Thames.

When one considers its abundance in the lower
Thames, it is surprising that this shiner has been
missed in past surveys. Because of its size, however, it
would have eluded large-mesh nets, and according to
ROM and the Ontario Ministry of Natural Resources
records, there have been few small fish collections
from the lower Thames. Stream characteristics may
have discouraged seining in the past. Notropis bucha-
nani may also have been regularly confused with N.
volucellus, although a check of past ROM collections
of N. volucellus from the upper Thames River did not
turn up any N. buchanani.

We captured L. humilis on 11 June 1980 at two
locations in Cedar Creek, Essex County, Colchester
South Township (42°01'53”N, 082°49’53”W and
42°01’38”N, 082°49’27”W). Thirty-eight specimens,
36-97 mm TL, catalogued as ROM 36441 and 36442,
were seined from shallow muddy water witha bottom
of silt, clay, and detritus.

Lepimis humilis was the most abundant species
comprising almost half of the total catch of several
species. Eight specimens (51-58 mm TL) were females
full of eggs. The two largest individuals (83 and 97 mm
TL) were older males with enlarged ear flaps, and the
remainder was smaller males and juveniles. One of the
collections included four hybrids between the Pump-

1981

kinseed and the Orangespotted Sunfish, L.
gibbosus X L. humilis, 49-69 mm TL (ROM 36443),
identified by intermediacy of diagnostic characters.

Lepomis humilis is distinguished from other Lepo-
mis by a flexible opercle, greatly elongated preopercu-
lar sensory pores, and conspicuous orange (male) or
brown (female) spots on the cheeks and sides (see
Trautman 1957). Number of dorsal spines (10-12),
dorsal soft rays (10-11), and anal soft rays (8-10) of
the Cedar Creek individuals averaged higher than
Kansas specimens (Cross 1967), the only specimens
for which these values are given in the literature.

According to Trautman (1957), L. humilis was first
recorded from the Lake Erie drainage in 1929 in Lake
St. Mary’s, Ohio, and spread rapidly northeastward.
In 1952 the first specimens were captured on South
Bass Island, Lake Erie, 44 km south of the present
record. Specimens of L. humilis should be looked for
in future surveys in the drainages of Lake Erie and
Lake St. Clair because the range of this silt-tolerant
species will likely continue to expand.

Acknowledgments
We thank E. J. Crossman for valuable help in the
preparation of this note, R. M. Bailey for confirming

NOTES Dial

the identity of the Ghost Shiners, and G. A. Good-
child for providing Ontario Ministry of Natural
Resources fish survey records of Kent and Essex
Counties.

Literature Cited

Bailey, R. M. 1956. A revised list of the fishes of lowa, with
keys for identification. Jn lowa Fish and Fishing. Edited
by J. R. Harlan and E. B. Speaker. Iowa State Conserva-
tion Commission. pp. 325-377.

Cross, F. B. 1967. Handbook of fishes of Kansas. Univer-
sity of Kansas Museum of Natural History. Miscellaneous
Publication 45. 357 pp.

Gilbert, C.R. 1980. Notropis buchanani Meek, Ghost
Shiner. p. 243 Jn Atlas of North American freshwater
fishes. By D. S. Lee, C. R. Gilbert, C. H. Hocutt, R. E.
Jenkins, D. E. McAllister, and J. A. Stouffer, Jr. North
Carolina State Museum of Natural History, Raleigh,
North Carolina. 854 pp.

Smith, P. W. 1979. The fishes of Illinois. University of IIhi-
nois Press. 314 pp.

Trautman, M. B. 1957. The fishes of Ohio. Ohio State Uni-
versity Press. 683 pp.

Received 28 August 1980
Accepted 8 December 1980

Reports of Significant Range Extensions

Range Extensions for 15 Teleost Fishes
in the Hudson Bay Lowlands, Ontario

Osteoglossiformes: Hiodontidae:

Salmoniformes: Osmeridae:
Cypriniformes: Cyprinidae:

Perciformes: Percidae:

Cottidae:

Hiodon alosoides, Goldeye

H. tergisus, Mooneye

Mallotus villosus, Capelin

Phoxinus eos, Northern Redbelly Dace

Couesius plumbeus, Lake Chub

Notropis atherinoides, Emerald Shiner

N. cornutus, Common Shiner

N. heterolepis, Blacknose Shiner

N. hudsonius, Spottail Shiner

Pimephales promelas, Fathead Minnow

Semotilus margarita, Pearl Dace

Perca flavescens, Yellow Perch

Etheostoma exile, lowa Darter

E. nigrum, Johnny Darter

Myoxocephalus quadricornis, Four-horned
Sculpin

As part of its Aquatic Habitat Inventory Program,
the Ontario Ministry of Natural Resources (OMNR)
has recently been systematically evaluating the
resources of the Hudson Bay lowlands of Ontario.
Between 1975 and 1979 surveys of the fish fauna have
revealed range extensions for the 15 species of fishes
listed. Sources used to define known ranges were

Scott and Crossman (1973), Lee et al. (1980), and
Ryder et al. (1964), as well as personal communica-
tions with scientists at the Royal Ontario Museum
(ROM) and the National Museum of Natural Scien-
ces (NMNS).

The sites where the fish were collected are shown in
Figure |. Details of the capture methods and habitats

TABLE |—Range extension records for |5 species of fishes collected in the Hudson Bay lowlands of Ontario. Specimens were
deposited at the Royal Ontario Museum (ROM) or National Museum of Natural Sciences (NMNS)

Location No. Fork Catalogue Extension
on of length, or accession of range,
map fishes mm Date collected no. km
Goldeye, Hiodon alosoides
(17) | 65 8 September 1976 NMNS 76-611 150 NE”
Mooneye, Hiodon tergisus
(16) 325 8 August 1976 NMNS 608 abe
Capelin, Mallotus villosus
(9) I 118 3 July 1979 NMNS 79-966 160 SE*
Northern Redbelly Dace, Phoxinus eos
(8) 5 36-53 2 October 1978 ROM 3877 335 NNW°
(8) 1, 2 June 1979 ROM 3957

PD

1981

TABLE I — Concluded

REPORTS OF SIGNIFICANT RANGE EXTENSIONS

Location No. Fork
on of length,
map fishes mm Date collected
Lake Chub, Couesius plumbeus
(17) 4 37-S7 17 June 1976
(18) 3 52-73 7 June 1976
Emerald Shiner, Notropis atherinoides
(17) | 36 12 September 1976
(19) l 85 5 June 1976
Common Shiner, Notropis cornutus
(8) 44 20-94 2, 9 October 1978
(8) 12 June 1979
Blacknose Shiner, Notropis heterolepis
(8) 6 35-42 2 October 1978
(8) 2 June 1978
Spottail Shiner, Notropis hudsonius
(3) l 58 14 August 1976
(11) 21 26-98 5, 6 June 1978
(12) 13 28-94 5 June 1978
(2) | 87 26 June 1978
(13) 7 33-85 5 June 1978
(4) 5 30-58 23, 24 July 1976
(10) 10 19-101 18 July 1976
Fathead Minnow, Pimephales promelas
(8) 25 23-57 2 October 1978
(8) 1, 2 June 1979
Pearl Dace, Semotilus margarita
(6) 39 29-67 11, 19 June 1977
(8) 19 38-69 2 October 1978
(8) 1, 2 June 1979
Yellow Perch, Perca flavescens
(7) l 250 15 August 1979
(11) 3 46-57 5, 8 June 1978
(12) 5) 41-57 5 June 1978
(13) yy 44-79 3, 5 June 1978
(1) 34 40-47 21, 22 August 1977
(10) 29 34-93 15, 16 July 1976
lowa Darter, Etheostoma exile
(3) | 50 27 June 1978
(5) 5 26-31 16 August 1976
(2) 2 31-38 21, 26 August 1977
(13) l 35 5 June 1978
(8) I 31 2 June 1978
Johnny Darter, Etheostoma nigrum
(17) 2 38-54 17, 23 June 1976
(14) | 28 30 October 1976
(15) 19 26-60 August-October 1975
(8) 6 29-63 2 October 1978
(8) 1, 2 June 1979
Four-horned Sculpin, Myoxocephalus quadricornis
(18) l 44 12 September 1976
(14) | 180 5 October 1976
(8) I 150 2 October 1978

“Southernmost record in James Bay.
Not previously recorded within lowlands in this watershed.

“Previous record from this location disputed (Dymond 1937).

“Caught in area that at times is within tidal salt water influence.

“Not previously recorded in this watershed. Isolated by salt water from previous record.
‘Not previously recorded in Hudson Bay drainage in Ontario.

Catalogue
or accession
no.

ROM 3129
ROM 3129

ROM 3260 |
ROM 3129

ROM 35739
ROM 36385

ROM 35743
ROM 36387

ROM 3268
ROM 3695
ROM 35635
ROM 35897
ROM 35634
ROM 3263
ROM 3263

ROM 35740
ROM 36401

ROM 35089
ROM 35742
ROM 36386

NMNS 79-965
ROM 3695
ROM 3695
ROM 3695
ROM 3566
ROM 3846

ROM 3263
ROM 3846
ROM 35898
ROM 3695
ROM 36384

ROM 3129
ROM 3264
ROM 2437
ROM 3877
ROM 36389

ROM 3263
NMNS 76-612
ROM 35738

PNB)

Extension
of range,
km

170 NE¢*
165 NE¢*

260 N*
165 N°

250 N'
300 NE

240 NE
135 NW
130 NW°
22 50NIEx
130 NW
230 NE
180 WNW

335 NNW

140 ESE®
335 NNW°

435 E°

380 ESE*
175 NNW
175 NNW
275 NE‘
200 WNW”

720 NNW
700 NWW*
720 NWW
330 NW°

340 NNW’

180 SE"

ISS3SSEx
140 SSE
70 SSE

MTS See
2350S En
160 SE*

214

if

Uff Vi

| ONTARIO

51° -—— oat
104
f° 8

8 6°

At!

FIGURE |. Sites where fishes were captured within the Hud-
son Bay lowlands of Ontario. Inset delimits extent of
the lowlands in the province. Collection sites were as
follows: 1, Shamattawa Lake (55° 10’N, 58°41’W); 2,
North Washagami Lake (54°31’N, 85°03’W); 3, Haw-
ley Lake (54°31’N, 84°38’W); 4, Sutton Lake
(54° 16’N, 84°42’W): 5, Kinusheo Lake (54° 16’N,
84°11’W); 6, Lakitusaki River, (54°22’N, 82°29’W);
7, Albany River (52°13’N, 81°36’W); 8, Wapiskau
River (51°51’N, 80°47’W): 9, James Bay (51°53’N,
80°45’W); 10, Trilsbeck Lake (50° 48’N, 84°02’W); 11,
Jaab Lake (51°09’N, 82°58’W): 12, North Sandbank
Lake (51°08’N, 82°42’W) — not listed in Gazetteer of
Canada (Ontario); 13, Sandbank Lake (51°06’N,
82°42’W); 14, Moose River (51° 14’N, 80°38’W); 15,
Partridge River (51°19’N, 80°19’W):; 16, Abitibi
River (51°04’N, 80°55’W): 17, Harricanaw River
(51° 10’N, 79°47’W); 18, Kesagami River (51°08’N,
79°47’W): 19, Kattawagami River (51°04’N,
79°44’W).

THE CANADIAN FIELD-NATURALIST

Vol. 95

are available from the Ontario Ministry of Natural
Resources (Box 190, Moosonee, Ontario POL IYO).
We have summarized information on the 15 species in
Table | including the approximate distance and direc-
tion of our new records from the closest previously
recorded localities. The range extensions are from 70
to 720 km.

Individuals who assisted us in our collections or
from whose collections the records are drawn are T.
Archibald, T. Bellhouse, T. Edwards, D. Fletcher, F.
Johnson, R. Stitt, G. Sudbury, and J. P. Prevett who
also assisted with the preparation of the manuscript.
Specimens were identified by D. E. McAllister of
NMNS or one of the following OMNR staff: G. C.
Gillespie, E. Holm, F. Johnson, and O. C. O'Neil:
Collections were funded by OMNR with assistance by
a grant from the Federal Industrial Development
Agency.

Literature Cited

Dymond, J.R. 1937. New
Copeia 1937(1): 59.

Lee, D. S., C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E.
McAllister, and J. R. Stouffer, Jr. 1980. Atlas of North
American freshwater fishes. North Carolina State
Museum of Natural History, Raleigh, North Carolina. 854

records of Ontario fishes.

Pp.

Ryder, R.A., W. B. Scott and E. J. Crossman. 1964.
Fishes of northern Ontario, north of the Albany River.
Royal Ontario Museum Life Sciences Contribution 60. 30
Pp.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada Bulletin
184. 966 pp.

B. R. ZALEWSKI and J. B. WEIR

Ontario Ministry of Natural Resources, Box 190, Moosonee,
Ontario POL 1Y0

Received 14 April 1980
Accepted 18 November 1980

News and Comment

Editor’s Report for 1980

During 1980 the manuscript flow for The Canadian
Field-Naturalist was similar to that of the previous
year. Table | summarizes the numbers of manuscripts
accepted for publication with respect to the numbers

-submitted in any given year. Note that the figures for
accepted manuscripts for recent years are not final as
some manuscripts will still be revised and published.

TABLE |—Statistics on manuscripts submitted and accepted
to the end of 1980

Number of manuscripts

Year

submitted Submitted Accepted
1973 153 117
1974 152 116
1975 167 122
1976 147 93
1977 137 88
1978 149 91
1979 148° 90
1980 137° 47

*Includes four manuscripts not formally refereed: one View-
point and three papers from The Ottawa Field-Naturalists’
Club Centennial Symposium.

Volume 94 (1980) of The Canadian Field- Naturalist
contained 93 scientific papers (48 Articles and 45
Notes). Table 2 gives the breakdown according to
subject matter; this is similar to those of previous
years. I hope that the publication of the first of the

“Reports of Significant Range Extensions” will
encourage others to contribute to this annotated list.
Although I originally proposed to publish them as an
annual list, these range extensions will now, until

circumstances change, appear in the next available
issue of the journal.

TABLE 2—Number of manuscripts published in The Cana-
dian Field-Naturalist in 1980 with respect to field of study.

Number of manuscripts

Subject Total (Articles + Notes)
Birds 31 (18 + 13)
Mammals 27 (12 + 15°)
‘Plants 19 (10 + 9)
Fishes 5) Gar)
Amphibians and Reptiles 5 (3 + 2)
Invertebrates 4 (0+ 4)
Other subjects 2» (2° + 0)

“Includes one Note under the heading Reports of Significant
Range Extensions.
"Includes a Viewpoint.

The Business Manager, W. J. Cody, has supplied
some figures on the current mailing list of The Cana-
dian Field- Naturalist, and these are included as an
item of general interest in Table 3. The data indicate
that the journal receives a wide national and interna-
tional distribution both to individuals.and institutions
(including libraries). Besides addresses in Canada and

TABLE 3—Current (1980) geographic distribution of recipients of The Canadian Field- Naturalist

Club Individual Institutional All recipients

Area Memberships" subscribers subscribers Number (%)
Canada

Maritime provinces 40 30 36

Quebec 68 18 43

Ontario 832 105 128 1735 (77.3)

Prairie provinces

and territories 135 114 85

British Columbia 46 32 23
United States 103 66 272 44] (19.6)
Other countries 7 9 54 70 (@3ab)
Total number (%) 1231° (54.8) 374° (16.7) 641° (28.5) 2246 (100)

“Includes Family Memberships.

Includes copies to 15 Honorary Members and to the club’s patrons.

“Includes seven copies for journal personnel.

“Includes 13 copies to depository libraries and abstracting and indexing services.

2S

216 THE CANADIAN FIELD-NATURALIST

the United States, the journal is mailed to the follow-
ing countries: Australia, Belgium, Brazil, Czechoslo-
vakia, Denmark, Finland, France, West Germany,
Iceland, Jamaica, Japan, Mexico, Netherlands, New
Zealand, Norway, Panama, Peoples Republic of
China, Poland, St. Pierre and Miquelon, South
Africa, Sweden, Switzerland, United Kingdom, and
USSR.

This year we especially express our appreciation to
Marilyn Dadswell for her seven years of dedicated

Voleo5

copy-editing. At the same time we welcome Patricia
Lalla to fill the void created by Marilyn’s retirement.
My personal thanks are also extended to my editorial
colleagues, to the Production Manager, to the Busi-
ness Manager, and to our many willing referees, all of
whomassist substantially in maintaining standards or
otherwise help with the publication of interesting pa-
pers pertinent to the natural history of Canada.

LORRAINE C. SMITH
Editor

A New National Wildlife Area — Long Point in Lake Erie

The donation of the major part of Long Point in
Lake Erie, the largest unspoiled sand dune - marsh
land complex in the Great Lakes, was made to the
people of Canada by the Long Point Company and
the Nature Conservancy (U.S.). Except for small
pieces of the point, the land had belonged to the Long
Point Company since 1866. Now under the manage-
ment of the Canadian Wildlife Service of Environ-
ment Canada, Long Point becomes one of 39 National
Wildlife areas under the Canada Wildlife Act.

Long Point, a 32-km-long sandy spit, juts into Lake
Erie, south of Simcoe, Ontario. As one of Lake Erie’s
three major points (the others being Point Pelee
National Park and Rondeau Provincial Park), it
serves as a resting or nesting stop for 150 species of
birds and is considered vital for maintaining certain
bird populations at healthy levels. A significant por-
tion of the world’s population of Redhead and Can-

vasback ducks and Whistling Swans pass through
Long Point each year. In addition to migrating birds,
Long Point is important for bats and butterflies that
traverse Lake Erie. They funnel through the area
because it is the first point of land encountered on
northward spring migration and the most southerly
point for their fall movement.

The vegetation on Long Point is unusual because it
is of a type usually associated with the middle-eastern
United States. Also the ridges and valleys of the sand
dunes create an intriguing juxtaposition of different
plant communities. The older ridges contain mature
forests while savannas predominate near the younger
tip of the point. At least five rare or threatened species
of amphibians and reptiles are found on Long Point.

Because of the care and foresight of the Long Point
Company, Long Point exists today in its wild state as
valuable wildlife habitat.

Canadian Wolf Status and Conservation Strategy Workshop

A workshop on wolf conservation problems in
Canada will be held 12-14 May 1981 in Edmonton,
Alberta. The purpose of the workshop is to review the
status of the species in Canada, and to map out a
conservation strategy based on sound technical
information. This will lay out the groundwork for the
species’ survival into the 21st century.

Emphasis of the workshop will be on the guidelines
provided by the International Union for Conservation
of Nature and Natural Resource (IUCN) on World
Conservation Strategy. The conservation strategy
focusses on the preservation of genetic diversity and it

endeavors to ensure sustainable utilization of species
and ecosystems.

The participants already confirmed will deliver
papers on the following topics: IUCN perspective,
wolves and ungulates, wolves and livestock, wolf tax-
onomy, and general papers including ones on wolf
management.

The proceedings of the workshop will be published.

For further information please contact Dr. L.N.
Carbyn, Canadian Wildlife Service, 9942-108 Street,
Edmonton, Alberta T5K 2JS.

1981

NEWS AND COMMENT S Dla

The Ottawa Field-Naturalists’ Club

Special Publications

1. Autobiography of John Macoun

A reprint of the 1922 edition of the fascinating life story of one of Canada’s outstanding early
naturalists, with a new introduction by Richard Glover and bibliographical essay, footnotes,
and index by William A. Waiser, plus three maps of John Macoun’s western travels.

Individuals
Libraries

$12.50 plus $2 postage and handling
$15.00 plus $2 postage and handling

2. Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index.

Compiled by John M. Gillett

A complete author, title, and subject index to the predecessors of The Canadian Field-
Naturalist, the first thirty-nine volumes of the publications of The Ottawa Field-Naturalists’

Club.

$25 plus $2 postage and handling

Centennial Bird Record

Songs of the Seasons

More than fifty eastern North American birds and amphibians are presented in full stereo-
phonic sound as recorded in the wild by wildlife recording expert F. Montgomery Brigham.

$9.11 (postage and handling included but
Ontario residents must add 7% sales tax

Please send orders to:

The Ottawa Field-Naturalists’ Club
Box 3264 Postal Station C
Ottawa, Ontario, Canada

KIY 4J5

The Ontario Field Biologist

The Ontario Field Biologist isa semiannual natural
science journal of the Toronto Field Naturalists,
which has been published regularly for over 25 years.
Articles and notes dealing with amateur and profes-
sional research in all aspects of natural history are
published in it. It fills a role between the senior scien-
tific journals and club publications, and hence pro-
vides a vehicle particularly for amateur papers of
merit. Emphasis is on Ontario subject matter, and all
papers are subject to review. Topics in the last two

years include a monitoring survey of winter birds in
Wellington County, Ontario, Ontario’s pondweeds, a
review of the status of the Greater Black-backed Gull
on the Great Lakes, Tremblay’s Salamander in
Algonquin Park, and notes on the flora of the Cana-
dian portion of the Niagara frontier. Subscriptions
are available ($4.00 a year) from the Toronto Field
Naturalists, 195 Glengarry Avenue, Toronto, Ontario
MSM IEI.

Submitted by Clive Goodwin

The Wilsori Ornithological Society Annual Meeting

The 61st annual meeting of The Wilson Ornitholog-
ical Society will be held at Sackville, New Brunswick,
Canada, on 4-7 June 1981. The meeting will be hosted
by the Canadian Wildlife Service — Atlantic Region,
the Biology Department of Mount Allison University,
and the Chignecto Naturalists’ Club. Information
concerning accommodations, transportation, excur-

sions, and related matters, and a call for papers will be
mailed to the membership in early March 1981. The
deadline for submission of abstracts will be 15 April
1981. Chairman of the local arrangements committee
is Dr. Anthony J. Erskine, Canadian Wildlife Service,
P.O. Box 1590, Sackville, New Brunswick, Canada
E0A 3C0.

Book Reviews

ZOOLOGY

A Bibliography of British Columbia Ornithology, Volume I

By R. Wayne Campbell, Harry R. Carter, Christopher D.
Shepard, and Charles J. Guiguet. 1979. British Columbia
Provincial Museum Heritage Record Number 7, Victoria.
185 pp., illus. $4.00.

The compilation of a bibliography is among the
most time-consuming and frustrating tasks facing any
scientist. The literature is usually scattered widely and
contains many references published in obscure places,
making the compilation of a complete bibliography
an unrealistic goal. Yet, even a poor bibliography is
useful to others. This volume is an excellent beginning
on a bibliography to British Columbia birds.
Although not stated anywhere in the book, the desig-
nation as Volume | in the title indicates that at least a
second volume is planned.

The book consists of a short introduction, a list of
sources searched, an explanation of format, acknowl-
edgments, a bibliography of 2100 references, and indi-
ces to species, geographic areas of the province, and
authors. Although the authors state that books and
notes on single observations are not included, they
presumably mean non-bird books with some bird data
or large regional works, which include British Colum-
bia, as several bird books are included. Similarly,
many notes documenting single sightings out of range
are included, and the authors apparently mean that
they have excluded short notes on common birds inan
area, such as may occur in local newsletters. Many
theses and government reports are included, a feature
of particular value to researches, as these are the sort
of reports that are especially difficult to track down.

The list of sources searched is extensive, although

Population Ecology of Raptors

By lan Newton. 1979. Buteo Books, Vermillion, South
Dakota.:399 pp., illus. U.S. $35.

At last a book on raptors which is not simply a
rehash of previously written books on this subject!
This latest book by Ian Newton (his first was on
finches in 1972) will surely have great impact on the
thought processes of not only today’s raptor enthusi-
asts, but wildlife ecologists in general. On page 13, he
states his objectives which are to organize and inter-
pret critically existing knowledge on population regu-
lation of raptors and to present this synthesis in a
simple informative manner for maximum benefit to
professional and amateur alike. Newton succeeds ina
most admirable way and leaves very few stones
unturned, indeed.

some obvious journals, such as Audubon Field Notes
— American Birds, covering North America as a
whole, and others, suchas Western Birds, covering the
west coast, including British Columbia, specifically
have not yet been included. Presumably these have
been reserved for Volume 2. Some papers, of which
the senior author has a copy, are excluded, presuma-
bly until the next volume.

The references are listed in the “order received,” and
not in any particularly useful sequence. However, the
three useful indices give easy access to the papers by
species, author, and area. Distinct races, until recently
considered full species, are not listed separately in the
species index, nor are hybrids. A few references per-
haps should have included brief annotations to indi-
cate their relevance to British Columbia where titles
seem to indicate otherwise (e.g. “First Alberta record
of the Glaucous-winged Gull”). The book is remarka-
bly free of errors — I detected only “Porchard” for
Pochard (p. 135) — and is produced well with attrac-
tive drawings, by an anonymous artist, interspersed
throughout. Ornithologists in British Columbia and
the entire west coast of North America will find this a
very valuable document. When Volume 2 is complete,
few will need to do more thanconsult these books and
search subsequent literature when compiling available
data on an ornithology subject within British
Columbia.

MARTIN K. MCNICHOLL

128 Silvergrove Hill N. W., Calgary, Alberta T3B 4Z5

The presentation of the information contained in
the book is best described in the author’s own words
on page 16: “The first two chapters .. . deal with the
social organisation and spacing behaviour of raptors,
and the next two with density regulation. There are —
then five chapters on various aspects of breeding, two
On movements, One on mortality, and three on the
effects on raptor populations of human persecution,
organo-chlorine chemicals, and other pollutants,
respectively. Lastly, there are two chapters dealing
with conservation management and captive breed-
ing.” The book deals virtually with the diurnal raptors
or the Falconiformes.

I personally found the book extremely useful in two
ways. First, by accumulating and synthesizing the vast

218

1981

wealth of raptor literature, including his own intensive
research on British Sparrow Hawks and Peregrine
Falcons, the author has been able to draw certain
conclusions about population regulation of raptors.
Previously, most of these generalizations were no
more than mere gut feelings on the part of scientists,
but never substantially proven. Secondly, Newton
offers much food for thought in his book by providing
his own theories and speculations in this field. Most
are based on no more than good common sense, but
nevertheless are just begging to be studied. For exam-
ple, on page 177, the author points out the scarcity of
long-term studies involving marked birds.

With regard to the layout, I found the presentation
of the tables (68 of them) at the back of the book
instead of interspersed throughout the text rather
bothersome, but admittedly, the author did a fine job
of summarizing them in the written text. I especially
liked the short summary presented at the end of each
chapter.

The ink drawings by Jim Gammie introducing and

Love Affair with a Cougar

By L. Hancock. 1978. Doubleday Canada Ltd., Toronto.
252 pp., illus. $9.95.

Love Affair with a Cougar is a detailed and tiring
account of Lyn Hancock’s experience raising or-
phaned cougars in southwestern British Columbia.
The story follows the development and fate of four
cougar kittens. Interspersed throughout the story are
discussions of care and handling of cougars, public
image of cougars, human-cougar interactions, and
the social—political problems of keeping wild
animals.

The pace of the book is very sluggish. Descriptions
of the author’s relationship with the litter, her re-
unions with one of the cougars after it had been placed
in a local zoo, and the interactions between the cou-
gars and the author’s students are redundant and
laborious to read.

Camel Quest
By Anne Innis Dagg. 1978. York, Toronto. 150 pp., illus.

Anne Dagg recounts a journey that she and a col-
league, Hilde (whose last name is never revealed),
embarked upon to study camels in Mauritania, a
small Islamic Republic bounded in the west by the
Atlantic Ocean, in the south by Senegal, in the east by
Mali, and in the north by Algeria. The southern part

BOOK REVIEWS 219

occasionally closing each chapter are superb. The
black and white photographs selected from various
photographers add to the book’s appeal, as they were
obviously selected with quality in mind. Also clearly
presented throughout the text in an informative sup-
portive style are 50 graphs. All in all, the drawings,
photographs, figures, lists of tables and figures, bibli-
ography, index, and general lack of typographical
errors indicate that this book was written and pub-
lished with professionalism in mind.

This book is an absolute must for any conscien-
tious, self-respecting, wildlife, population ecologist,
particularly one who shares an affinity for birds of
prey.

DAVID M. BIRD

Macdonald Raptor Research Centre, Macdonald Campus
of McGill University, Ste. Anne de Bellevue, Quebec H9X
1C0

I disagree with the author’s use of her animals as
representatives of the wild cougar population because
they were heavily conditioned to the human environ-
ment. “Only by constant association and a buildup of
mutual trust could we hope to maintain the friendship
between ourselves and the cougars as they continued
to develop.” p. 145. The discussions of the domestic
relationship between humans and cougars offered lit-
tle convincing evidence of a wild cougar’s behavior.

The downfalls of the book could have been elimi-
nated by strict editing. The interesting parts of the
book would then have been easier to appreciate.

JOHN DAVIE

Alberta Department of Energy and Natural Resources, Fish
and Wildlife Division, Calgary, Alberta T2G 419

of the country is savanna grassland and the northern
part, Sahara Desert. About 80% of the | million
inhabitants are Moors, a people of mixed Berber and
Arab blood.

It all began in Calgary, Alberta, in 1971, when she
attended a lecture that Hilde presented at a conference
on ungulate behavior. Dagg, because of her interest in
African mammals, found herself intrigued by Hilde’s

220 THE CANADIAN FIELD-NATURALIST

lecture which focused on the biology of camels. She
had earlier carried out studies for her doctorate, which
concentrated on the locomotion of giraffes, in eastern
Transvaal of South Africa. Subsequently, she wanted
to study the camel, the only other large mammal that
could survive on thorny vegetation for extended peri-
ods without drinking water, but to her dismay lacked
the funds.

Hilde, a French biologist, who had spent time in
Mauritania studying camel physiology, wanted to
return there, and hence a trip was planned. Dagg went
to study camel locomotion, and Hilde the vegetation
surrounding wells utilized by the camels and their
nomadic owners. At the time of their visit, Mauritania
was being ravaged by drought. Many of the free-
roaming camels were dying off, and many of the nom-
ads had been forced to migrate to the Spanish Sahara
where they could obtain food and water. Unfortu-
nately, their camels were left in the desert to fend.
usually unsuccessfully, for themselves.

In addition to their biological work, Dagg and
Hilde decided to seek out and study the Reguibat“...
the greatest camel men and perhaps the last truly free
people of the world who had once reigned in the
western desert as far south as Timbuctoo and as far
north as Spain.” Should this way of life disappear

Charrs: salmonid fishes of the genus Sa/velinus

Edited by Eugene K. Balon. Dr. W. Junk bv Publishers, The
Hague, The Netherlands. 928 pp., illus. U.S. $210.

This monograph provides a broad summary of our
knowledge of the charrs, by 40 contributors from
Europe, Asia, and North America. It is a landmark in
the history of charr research. The 30 chapters are
grouped into four parts: I. Review of the main forms;
II. Taxonomy, biogeography, phylogeny, and mor-
phology; III. Ontogeny, ethology, physiology, and
ecology; and IV. Utilization of charrs by man and the
creation of a new taxon.

Lionel Johnson, inan excellent chapter in Part I on
the Arctic Charr, conveys the impression of the
immense variability of this circumpolar species, for
example, the frequent occurrence in Europe of dwarf
and normal-sized populations, of spring and fall
spawners, river and shallow or deep lake spawning,
possession of several patterns of life cycles with vary-
ing stays and sequences in sea, lakes, and rivers. The
chapter by R. H. Armstrong and J. Morrow shows
Dolly Varden Charr have equally variable life history
patterns. These authors also inform us that J. D.
McPhail’s western Arctic-Bering Sea form of Arctic
Charr is in reality a northern form of Dolly Varden.

Vol. 95

forever, Hilde felt it important that it be documented
for the archives.

In this volume are tidbits of information about the
camel, but emphasis is placed more on Dagg’s expe-
riences. It is an informal analysis of the nomadic way
of life, a description of the various people encountered
throughout their journey, and a review of the climate
and terrain of the country. The book is of a socio-
anthropological slant. If you are interested in Mauri-
tania, then the volume will serve as introductory
reading.

The book is tainted in some sections by certain
remarks that only serve to detract from the story. For
example, “... I was associated as a researcher with the
Department of Biology at the University of Waterloo,
Canada, but none of the other biologists, all men,
cared to discuss the ecology of animals in Africa.”
Undoubtedly, too, there are many women biologists
in this country who would not be interested in African
animal ecology.

PAUL A. GRAY

Wildlife Service, Government of the Northwest Territories,

Yellowknife, NWT XIA 2L9

This redefinition of the Dolly Varden is based on
discriminant function analysis (DFA) of gill raker,
vertebral, and pyloric cecal counts. Unfortunately the
calculated weights for the characters are not given, so
other workers cannot use their results for identifica-
tion. However, assuming samples are adequate, the
graphs convince one that there are three groups: Arc-
tic Charr, northern and southern Dolly Varden, and
that multivariate analysis is a useful approach in this
difficult group.

The chapter on the Brook Charr emphasizes physi-
ology as well as life history. Valuable information is
given on the effects of pH on the life cycle of Brook
Charr. Distressingly, the range of Brook Charr is
decreasing in parts of the United States because of
developmental pressures. It is also suggested that
angling has selected for short life span and small size
in the eastern United States.

Ksenia Savvaitova provides an overview of charr
taxonomy in the Palearctic, valuable because of the
number of charrs that have been described there, often
in journals not generally available. Several charrs des-
cribed from the Taimyer, USSR, are synonymized
following unclear reasoning. She does not seem to

1981

consider occurrence of sympatric reproductively iso-
lated populations as species and presents much theory
but little actual data on individual taxa. Mednikov et
al., applying the technique of DNA hybridization,
concluded that all Eurasian charrs should be regarded
as S. alpinus except S. leucomaenis. Other papers in
this volume affirm and provide evidence for the dis-
tinctiveness of §. ma/ma and S. alpinus, but Savvai-
tova relegates these to ‘malmoid,’ ‘alpinoid,’ and ‘high
arctic’ forms. This approach should be contrasted
with Cavender’s, who discovers, clearly describes, and
illustrates new morphological characters for charr
phylogeny (comparative morphology lives), and Mor-
row’s multivariate analysis.

Robert Behnke was given the difficult task of dis-
cussing taxonomy of the genus Sa/velinus. This genus
he divides into three subgenera. In the subgenus Sa/ve-
linus, he recognizes S. alpinus, S. malma, S. confluen-
tus, S. profundus (Bodensee, Europe), S. taimyricus
(Taimyr, USSR), the unnamed stone charr (Kam-
chatka), and S. leucomaensis (western Pacific). He
considers Morrow’s S. anaktuvukensis to represent
only northern Dolly Varden populations in small
stream habitat in the Alaskan Brooks Range.
Cavender, unlike Behnke, recognizes S. pluvius
(Japan) as a distinct species. Only one species is rec-
ognized in the subgenus Cristivomer, S. namaycush.
Two species are recognized in Baione, the Brook
Charr, S. fontinalis, and the Silver Charr, S. agassizi
(Dublin Pond, New Hampshire). The latter, unrecog-
nized by the American Fisheries Society list of fishes
(in press), is nevertheless well distinguished by its low
gill-raker count (14-17), the raker’s knob-like form,
and the high number of pyloric ceca (43-58). The
Silver Charr is believed extinct. A comparative table
of characteristics of each species would not have been
difficult to compile and would have been very useful.

The Evolution of Culture in Animals

By John Tyler Bonner. 1980. Princeton University Press,
Princeton. 216 pp. $17.00.

Sociobiology is a hot topic and Bonner is both an
erudite biologist and an articulate author. It is there-
fore no surprise that a book on this topic by this
author is a pleasure to read. In eight chapters he
surveys the biological background for cultural evolu-
tion. The work is intended as a personal document
examining issues between the biological and social
sciences, while avoiding political issues, for a broad
audience. The central thesis is that the ability for
culture, taken as the transfer of information by behav-
ioral means, is an evolutionary product. After due

BOOK REVIEWS

22)

Balon’s five chapters on ontogeny of charrs wisely
includes provenance and descriptions of parents. No
evidence is provided for the suggestion in the title of
Part IV that a new species of charr has been created
(by hybridization of the Brook and Lake Charrs).

Many of the color photos of fish are excellent, but
rather small. Some illustrations are poor, the symbols
on the Arctic Charr distribution map are unreadable,
the American range of the Lake Charr is omitted on
the Lake Charr map. Otherwise the graphs, photos of
embryos, histological sections, and osteological fig-
ures are generally of high quality.

The authors and editor have had a formidable task
in just assembling the material, let alone digesting and
drawing generalizations from it. But, aside from
Behnke’s paper, there is little cross referencing
between chapters. There is no overview of how species
differ in response to their environments, nor on com-
parative adaptations of planktivores and predators to
their ways of life. No advantage was taken of the
opportunity to use the assembled information at
another level. Nevertheless, the editor and the con-
tributors are to be congratulated for the general excel-
lence of the book. There are few typographical errors.

Charrs provides a good summary of new and pre-
viously available but scattered information. The book
will be essential for researchers specializing in the
genus Sa/velinus and a useful resource book for those
having occasion to manage or investigate one or more
species. Invaluable although it is, the price puts the
volume beyond the budgets of most individuals and
some institutions.

DON E. MCALLISTER

Ichthyology Section, National Museum of Natural Sciences,
National Museums of Canada, Ottawa, Ontario KIAOMS8.

attention to problems such as nature versus nurture,
reductionism versus holism, and anthropomorphism,
Bonner contrasts genetical and cultural evolution and
examines ‘the resulting interactions. The functioning
of the brain, the eventual agent of culture, and of
genes, especially in behavior genetics, are discussed,
along with the time and size scales for different pro-
cesses. The biological steps toward culture are pre-
sented: the evolution of motility, of societies (bacte-
rial, insect, and vertebrate), of communication
systems, of learning and teaching, and of behavioral
flexibility in higher vertebrates. The discussion
includes the differences in the evolution of sociality
and of culture, and the advantages of both various

jpipe

sensory modalities for complex communicatory sig-
nals and sociality under what Bonner terms commu-
nal task selection, by which he means selection for
cooperation to perform tasks unachievable by indi-
viduals. In the final chapter, instances of nonhuman
and human cultures, the adaptiveness of cultures, and
the evolutionary trend to flexibility are surveyed. The
cultural passage of memes, or ideas, between individ-
uals is analogized with the genetic transmission of
genes across generations. As in so much of contem-
porary evolutionary thinking, the importance of the
contributions by W. D. Hamilton on kin selection and
J. Maynard Smith on evolutionarily stable strategies
is very obvious throughout the discussion. In keeping
with his uniformitarian stance, Bonner has provided
an excellent perspective of the long biological pro-
gression underlying cultural evolution.

With a very few minor blemishes, the book is well
composed and illustrated and in these respects
appropriately provides the impression of being a ligh-
ter version of E. O. Wilson’s hefty “Sociobiology.”
The explanatory footnotes are helpful, although a few
more (e.g. on Fisher’s Fundamental Theorem) would
assist “the well-read layman” of the intended
audience. The calm and friendly style also enhance the

BOTANY

The Whole Fungus: the sexual—asexual synthesis

Edited by Bryce Kendrick. 1979. Co-published by National
Museum of Natural Sciences, National Museums of Cana-
da, and the Kananaskis Foundation. 2 volumes, 793 pp.,
illus. Paper bound; available by mail from W. B. Ken-
drick, Department of Botany, University of Waterloo,
Waterloo, Canada N2L 3G1.

The covers and spines lack the subtitle, and the
half-title pages of both volumes bear a different subti-
tle — ‘Kananaskis II.’ The 27 chapters represent
reports on papers either presented at the conference
held in Kananaskis, Alberta, in 1977, or submitted for
publication at the editor’s request. Dialogues by par-
ticipants follow each presentation. A postcript, a list
of about 2000 references, and an index completes
Volume 2.

The aim of the book is to present ‘the sexual—asex-
ual synthesis,’ and ‘to unite the different phenotypes,
at least conceptually, and to use all of the data thus
made available in the elaboration and refinement of
our classificatory schemes.’ For over a century mycol-
ogists have been concerned with the fact that fungi can
be pleomorphic: Ascomycetes, Basidiomycetes of var-
ious kinds, and some so-called ‘Phycomycetes’ pro-

THE CANADIAN FIELD-NATURALIST

Vol. 95

attractiveness of the volume. The avoidance of politi-
cal issues doubtless contributes to this calm, but it is
nonetheless strange that Bonner should regard a dis-
cussion of the stability of recent Western sex role
changes as a topic beyond the scope of the book. The
need for the lengthy review of motility in lower forms,
long one of Bonner’s research interests, is doubtful,
even allowing for the deep perspective he wishes to
illustrate. On the other hand, given that, as Bonner
argues, brains have succeeded in bypassing genes, it
seems worthwhile to speculate on the possibility of
computers bypassing brains. Similarly, a couple of
mentions of “the evolutionary scale” and the lack of
discussion of Lamarckism are perhaps surprising,
although Bonner’s Princeton predecessor, Baldwin,
and his effect are mentioned. But such questions of
emphasis are rarely decidable and the criticisms are
minor. Overall this is an excellent volume as an intro-
duction to the topic.

PATRICK COLGAN

Department of Biology, Queen’s University, Kingston,
Ontario K7L 3N6

duce a sexual state and one or more kinds of asexual
states. Independent names have been applied to such
various states and mycologists have concerned them-
selves with connecting one state with the other(s). A
chapter on ‘Pleomorphism of fungi as treated in the
history of mycology and nomenclature’ is an excellent
introduction to the problems involved. Three useful
new terms — anamorph for imperfect (asexual) state,
teleomorph for perfect (sexual) state, and holomorph
for the whole fungus are used: the terms are explained
in a chapter reprinted from Mycotaxon (6: 207-211.
1977). Five chapters are devoted wholly or in part to
problems associated with the nomenclature primarily
of pleomorphic ascomycetous fungi.

The main thrust of the book is found in several
well-illustrated and documented chapters on
anamorph-teleomorph connections in specific groups
of fungi — Plectomycetes, nectrioid fungi, some
freshwater Hyphomycetes, some Coelomycetes, in
ascomycetous and so-called basidiomycetous yeasts,
Agaricales and other Basidiomycetes (excluding
Uredinales and Ustilaginales), and in Zygomycetes.
There is a ‘Classification of Ascomycetes partly based

1981

on their anamorphs’ and a chapter on ‘Deuteromy-
cetes and their relationships’ which includes a classif1-
cation of Deuteromycetes.

The largest chapter is a compilation, from the litera-
ture, of teleomorph-anamorph connections in unitun-
icate and bitunicate Ascomycetes which are listed
alphabetically by generic name of the teleomorph.
The records are then listed under the orders and fami-
lies of the teleomorph. Unfortunately the extent of the
literaiure search is not mentioned.

Records of teleomorph-anamorph connections
vary in their reliability for several reasons. Apart from
the basis for claiming a connection, a subject which is
discussed in the book, there are also problems of the
generic dispositions of teleomorph and anamorph(s).
For instance, in Me/anomma three species are listed
with apparently related pycnidial anamorphs whereas
a fourth species has an entirely different hyphomyce-
tous anamorph. The four connections were estab-
lished by ascospore isolations. If we accept the identi-
fications and connections as correct, then it is likely
that the teleomorph of the fourthentry, M. subdisper-

sum, is nota Melanomma.

The chapter on an appraisal of the taxonomic sig-
nificance of some modes of conidium ontogeny, and
discussions on the same topic elsewhere, serve to
remind us that so much remains to be done in the
classification of anamorphs. But it is evident that the
refinement of schemes of classification of both
morphs must proceed together.

In a brief review it is impossible to cover the whole
book composed for the most part of different

Grasses of Ontario

By W. G. Dore and J. McNeill. 1980. Agriculture Canada
Research Branch Monograph 28. Supply and Services
Canada, Ottawa. 566 pp. $12 in Canada; $14.40 elsewhere.

Dore and McNeill’s Grasses of Ontario is likely to
become the standard taxonomic treatment of central
North American grasses and to remain so for some
time. There are several reasons for this. Firstly, some
270 species in 78 genera are treated, representing the
large majority of grasses native to the Great Lakes
area; secondly, the taxonomic scholarship of the
authors renders the work highly reliable. In this
respect, the treatment of Panicum, for instance,
represents an entrée into a difficult genus that pos-
sesses numerous species southwards, many very sim-
ilar to Ontario species. To have the former eliminated
instead of attempting to identify Ontario Panicum
with a key designed for the United States is obviously
very helpful.

BOOK REVIEWS

223

approaches to essentially the same problems and
goals. But this is its strength.

A few chapters, such as ‘the classification and
nomenclature of fossil fungi,’ are lucidly written but
are, I believe, somewhat on the fringes of the stated
aim of the book.

The checking of several citations amongst the com-
pilation of ‘Teleomorph-anamorph connections in
Ascomycetes’ of particular interest to the reviewer has
revealed several errors. For example, ‘Torulopsis’ (pp.
380 and 387) should be ’?Torulopsiella.’ ‘Capnocybe,’
in the reference(s) given, was not included as an ana-
morph of Limacinia moniliformia and L. quinquesep-
tata (p. 374), or of Ophiocapnocoma sp. (p. 380). The
entry of ‘Phoma herbarum’ as an anamorph of Pleos-
pora herbarum, with the notation ‘2.3.1’ to indicate
considerable evidence of affiliation, is not substan-
tiated in the references given, including ‘Wehmeyer
(1961) which is not found in the references. On page
362 it is stated that ‘Spegazzini never illustrated his
papers’ yet his ‘Fungi Chilenses’ alone has 130 text
figures.

The book brings together a considerable amount of
information and should be available particularly to
those involved in the taxonomy, classification, and
nomenclature of Ascomycetes and Fungi imperfecti,
as well as of Agaricales and most other Basidiomy-
cetes in pure culture.

S. J. HUGHES

Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6

With the reservations noted below, Dore and
McNeill’s Grasses of Ontario is sensibly produced.
The introductory chapter explaining the layout of the
book, the sources used, and directing the reader else-
where for morphological interpretations necessary for
grass identification is sensibly brief. The taxonomic
part includes keys to genera, species, and sometimes
infraspecific taxa. Distribution maps of most species,
line drawings of a few, numerous photographs of
spikelets and caryopses as well as anecdotal informa-
tion about distribution and sometimes identificatory
matters, especially where dispute has existed or there
are real identification difficulties, are all of great
value. As is customary in regional floras, no species
descriptions as such are given.

The book will startle many by its novel format, but
because it will probably be used in conjunction with
other works providing species descriptions and mod-

224

ern tribal affiliations, this shortcoming is not great.
Nevertheless it is surprising that the authors have not
used modern tribal affiliations, and the implication in
their introduction that this might cause difficulties is
not well founded. Minor typographical errors are
quite numerous but will not confuse the careful
reader. The maps, particularly, are models of clarity
and will be very useful. Somewhat surprising is the
virtually equivalent treatment given to essentially
transient taxa (e.g. Pennisetum) and extremely rare
introductions (e.g. Nardus) with treatments of com-
mon taxa, but perhaps all have equal reason for being
properly identified. If space is no limitation then
nothing is really lost.

Thus it is essentially at the local level, i.e., critical

ENVIRONMENT

THE CANADIAN FIELD-NATURALIST

Vol. 95

species distinctions, good keys, and excellent maps,
that Dore and McNeill will be invaluable. The book
fails to be, but is not intended to be, a handbook of
Gramineae taxonomy. Dore’s lifework finally sees the
light of day, and his great firsthand knowledge of
Ontario grasses is at last presented for all to see. One
admires the depth of knowledge indicated. McNeill’s
organizational and taxonomic skills, one assumes,
have seen the work into print. He too 1s to be congrat-
ulated with the first author.

J. B. PHIPPS

Department of Plant Sciences, University of Western Ontar-
io, London, Ontario N6A 5B7

The Natural Selection of Populations and Communities

By David Sloan Wilson. 1980. Benjamin/Cummings
(Canadian distributor Addison-Wesley, Don Mills,
Ontario). xv + 186 pp., illus. U.S. $12.95.

At last there is a book in which the theory of group
selection has been given the attention it has deserved
for over a decade. Someone has finally provided a text
witha plausible mechanism for the operation of group
selection in nature without using the inconsistent
altruistic pathway. In fact, Wilson’s models would be
nullified if strong altruism were prevalent in nature.
Therefore, past arguments rejecting group selection
via rejection of the altruistic pathway cannot be used
against his thesis.

Wilson stresses models without using conventional
constant fitness values. He feels that fitness is spatially
and temporally heterogeneous and therefore constant
homogeneous population fitness values cannot be
used to model natural selection. In short, he feels that
individual selection’s predictions about relative fitness
are artifacts of the homogeneity assumption. Wilson
shows the reader that his feelings are indeed plausible
with convincing ridicule of conventional population
genetics theory. If there is spatial and temporal genetic
variation ina population, as we strongly suspect, Wil-
son’s models most likely are true. He shows that the
differential productivity of groups (group selection) is
an inevitable consequence of relaxing the homoge-
neity assumption. Conventional models of homoge-
neity only fit when applied to tiny populations.

Wilson believes that demes in nature are made up of
somewhat philopatric small groups of individuals
who ecologically interact more often with their imme-
diate neighbors and microclimatic conditions and

therefore are likely to experience similar selective
pressures regardless of genetic relationships. Because
of these similar selective pressures, these small groups
are to some extent similarly genetically structured and
are termed trait groups. Wilson terms demes of this
type structured demes. A population of a species is
envisioned to be made up of many structured demes
each subdivided into still smaller trait groups. It is the
genetic variation within and between trait groups on
which Wilson is concerned. The increase of genetic
variation among trait groups is always associated with
the regulation of population density at optimum size.

Selfish behavior of individuals causes the differen-
tial fitness of populations and communities through
what is termed indirect effects. For example, bark
beetles create galleries under tree bark while feeding.
This creates a habitat for many insects that can inhabit
trees only where beetles are present. The theory of
structured demes predicts that bark beetle predators
come from “outside” the community the bark beetle
creates. The data support this theory. Wilson shows
that conventional coevolutionary theory cannot
account for this observation and indeed predicts the
opposite trend (cf. p. 119-126). Interesting analogies
of indirect effects in human communities are
provided.

According to the theories presented here, it
becomes altruistic to compete with (or exclude)
another individual from gaining resources when the
population size is below optimum. After all, if an
animal wastes time competing instead of reproducing
when resources are abundant, he is reducing his
potential fitness. Dominant individuals chase lowly

1981

subordinates from their area while the rest of the deme
(intermediates) enjoy reduced interference, improved
conditions, and possibly increased productivity with-
out sharing the costs. In this case, Wilson points out
that lowly subordinates are selected against, but dom-
inance is neutral at best. The entire deme is rid of
subordinates which is a group advantage. Regardless
of the underlying causes of this event, the group has
benefitted through the indirect effects provided by the
direct beneficial effect the dominant individual was
trying to obtain. In short, a large portion of an indi-
vidual’s fitness is indirectly influenced by the activities
of other members in the community, and these influ-
ences are not accounted for in conventional models.
Throughout the text, mathematical models rein-
force the verbal arguments, but are not used to make
them. Models show how a lower fecundity is favored
by selection in one generation. Examples show where
niche differentiation between stages (e.g., larva vs.
adult insects) cannot be explained by individual selec-
tion (cf. p. 57-64). Models show that in some cases
individual fitness is maximized only if the individual
performs in the best interests of the community. Mod-
els for multilevel evolution in the final chapter (6)
provide ample mathematical evidence supporting all
contentions presented throughout the text. In short,

Circulation Models of Lakes and Inland Seas

By T. J. Simons. 1980. Canadian Bulletin of Fisheries and
Aquatic Sciences, Bulletin 203. Supply and Services Can-
ada, Hull, Quebec. 146 pp., illus. $12.00 in Canada; $14.40
elsewhere.

As Simons points out in his introduction, there has
been increasing activity in the field of modeling both
because of demands from many disciplines needing
information on water movement and asa result of the
increasing capabilities of modern computers. This
book comes conveniently at a time when some of the
basic modeling methods involving depth averaged
currents and transports are sorting themselves out
into standard procedures, and more complex models
with vertical variation of current and density are pro-
liferating. Thus the book is a valuable summary of
models to date and an indication of present trends and
limitations.

The title suggests a restriction to circulation in
enclosed small bodies of water, and in fact the open-
boundary problem and tides are not considered. In
addition the modeled area is taken to be small com-
pared to the weather systems that influence it. There is
still, however, a considerable amount of material
applicable to ocean modeling. There is some examina-
tion of surface response since it can be a useful tool in

BOOK REVIEWS

Das

these show that structured deme theory causes
increased community productivity until an equili-
brium at an adaptive peak is reached.

Perhaps some will criticize that Wilson is repeti-
tious in stating his theories. I feel, however, that he
keeps reminding the reader of them because his ideas
are so novel that one will forget their components
unless reminded. Also, a few passages are difficult to
grasp while conventional evolutionary theory is
engrained in the reader’s mind because of his earlier
training. Repetition of these passages makes one
understand the significant contribution to evolution-
ary science that Wilson makes. Overall, most discus-
sions are written in a way that allows the careful
reader to understand Wilson’s theories as though they
were common sense.

No serious naturalist should proceed into the future
without reading this book, perhaps the most signifi-
cant work of natural selection advanced within 15
years. Skeptics need only read pages 13-20; if they are
still entirely skeptical, they needn’t read further.

RICHARD M. ZAMMUTO

Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7

circulation model verification and in theoretical
examinations. There are many examples for ocean
modeling and many analogies and references to the
atmospheric sciences from which ocean and lake
modeling traces much of its development.

The book gives a good background. Terms such as
Rossby waves and Ekman number are developed and
defined in context, instead of just being dropped.
Illustrations are used to good advantage often in the
context of theoretical models of simple geometry lead-
ing to more complex models of real lakes and basins.
Simons uses many examples, frequently from his own
extensive work, and supplements these with a wealth
of references. Most of the book deals with the analyti-
cal bases of the models, with only one chapter devoted
to numerical procedures. If the book only had an
index and possibly a symbol table, it would be an ideal
reference book.

In his preface Simons says he is aiming the book for
“investigators of water quality problems and other
ecological concerns.” For these people it is a very
useful book, provided they already have a good foun-
dation in university level mathematics and perhaps
some ideas on fluid mechanics. A field naturalist
without this background trying to evaluate what this

226 THE CANADIAN FIELD-NATURALIST

technical field has to offer his science would have
difficulty extracting that information. In any case, the
low cost of this book makes it a bargain compared to
most of the modern literature available.

NEW TITLES

Zoology

Animals as monitors of environmental pollutants. 1979.
By the National Academy of Sciences, Washington.
xilit+ 421 pp. U.S. $20.

America’s endangered birds: programs and people working
to save them. 1979. By Robert M. McClung. Morrow,
New York. 160 pp., illus. U.S. $7.95.

Assessing toxic effects of environmental pollutants.
1979. Edited by S. D. Lee and J. R. Mudd. Ann Arbor
Science, Ann Arbor. 306 pp. U.S. $30.

+ Atlas of North American freshwater fishes. 1980. ByD. S.
Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E.
McAllister, and J. R. Stauffer, Jr. North Carolina State
Museum of Natural History, Raleigh. 825 pp., illus. U.S.
$20 plus U.S. $3 postage. (Canadian price $22.50 plus $3
postage).

Avian community structure of six forest stands in La Mau-
ricie National Park, Quebec. 1980. By J.-L. DesGranges.
Canadian Wildlife Service Occasional Paper, 41. Environ-
ment Canada, Ottawa. 34 pp., illus. Free.

The biology and physiology of the living coelacanth.
1979. Edited by John E. McCosker and Michael D.
Lagios. Papers from a symposium, June, 1977. California
Academy of Sciences, San Francisco. vi+ 176 pp., illus.
U.S. $10.

+Bird island in Antarctic waters. 1980. University of Min-
nesota Press, Minneapolis. xi+ 140 pp., illus. U.S. $18.95.

Birds of prey. 1979. By Gareth Parry and Rory Putman.
Simon and Schuster, New York. 120 pp., illus. U.S. $29.95.

*Birds of prey of the world. 1980. By Friedhelm Weick, in
collaboration with Leslie H. Brown. Parey (U.S. distribu-
tor IR Unlimited, New York). 159 pp., illus. U.S. $48.

*Birds of Regina. 1980. By Margaret Belcher. Revised edi-
tion. Special Publication Number 12. Saskatchewan Natu-
ral History Society, Regina. 151 pp., illus. $5 plus 50¢
postage.

*Birds of the Qu’Appelle, 1857-1979. 1980. By E. Manley
Callin. Special Publication Number 13. Saskatchewan
Natural History Society, Regina. 180 pp., illus. $7 plus 50¢
postage.

Vol. 95

DAVID A. GREENBERG

Coastal Oceanography, Bedford Institute of Oceanography,
P.O. Box 1006, Dartmouth, Nova Scotia B2Y 4A2

+Character variation and evolution of sibling species in the
Empidonax difficilis-flavescens complex (Aves: Tyranni-
dae). 1980. By Ned K. Johnson. Publications in Zoology,
Volume 112. University of California Press, Berkeley.
x+ 151 pp., illus. + plates. U.S. $9.50.

+A cold look at warm-blooded dinosaurs. 1980. Edited by
Roger D. K. Thomas and Everett C. Olson. Papers froma
symposium, Washington, February, 1980. American Asso-
ciation forthe Advancement of Science Symposium Series,
28. Westview Press, Boulder, Colorado. xxx + 514 pp.,
illus. U.S. $28.50

Comparative studies of the courtship and mating behavior
of tropical araneid spiders. 1980. By Michael H. Robin-
son and Barbara Robinson. Pacific Insects Monograph 36.
Bishop Museum Press, Honolulu. 218 pp., illus. U.S.
$22.50.

+Completely outfoxed. 1980. By Miles Smeeton. Van Nos-
trand Reinhold, Toronto. xi+ 148 pp., illus. $9.95.

Conservation and management of whales. 1980. By K. R.
Allen. University of Washington Press, Seattle. 120 pp.
U.S. $12.50.

tEcological investigation of a population of Dall
sheep. 1979. By Manfred Hoefs and lan McTaggart
Cowan. Syesis Volume 12, Supplement |. British Columbia
Provincial Museum, Victoria. 81 pp., illus. $7.

Ecology of Africanmammals. 1979. By M. J. Delany and
D. C. D. Happold. Longman, London, England. £25.

Ecology of small mammals. 1979. Edited by D. Michael
Stoddart. Chapmanand Hall, London. xiv + 386 pp., illus.
U.S. $44.95.

+A field guide to the birds of Australia. 1980. By Graham
Pizzey. Princeton University Press, Princeton. 460 pp.,
illus. U.S. $27.50.

A field guide to western birds’ nests. 1979. By Hal H.
Harrison. Peterson Field Guide Series, 25. Houghton Mif-
flin, Boston. xxxii + 280 pp., illus. + plates. U.S. $11.95.

First aid and care of wild birds. 1979. Edited by J. E.
Cooperand J. T. Eley. David and Charles, North Pomfret,
Vermont. 288 pp. U.S. $27.50.

1981

Fisheries ecology of floodplain rivers. 1979. By Robin L.
Welcomme. Longman, New York. x + 318 pp., illus. U.S.
$45.

The fishes of the islands in the South China Sea. 1979. By
the South China Sea Fisheries Institute and Others. Science
Press, Beijing, China. 613 pp., illus. No price given.

Fishes of the north Pacific. 1980. By Robert Browning.
Revised edition. Alaska Northwest, Edmonds, Washing-
ton. 432 pp., illus. U.S. $29.95.

+The fool hen: the Spruce Grouse on the Yellow Dog
Plain. 1980. By William L. Robinson. University of Wis-
consin Press, Madison. xviit+ 221 pp., illus. U.S. $18.50.

*Great Grey Owl: phantom of the northern forest. 1980. By
Robert Nero. Smithsonian Institution Press, Washington.
167 pp., illus. U.S. $17.50.

A guide to the behavior of common birds. 1979. By
Donald W. Stokes. Little, Brown: Boston. x + 336 pp.,
illus. U.S. $9.95.

A guide to the marine flora and fauna of the Bay of Fundy:
Annelida: Polychaeta. 1980. By Teresa D. Appy et al.
Technical Report 920. Fisheries and Oceans Canada,
Ottawa. 124 pp., illus. Free.

+Hawk lady: the story of a woman who opened her home to
care for wild birds of prey. 1980. By Stellanie Ure. Dou-
bleday, Toronto. 215 pp., illus. $14.95.

Investigating animal abundance: capture-recapture for
biologists. 1979. By Michael Begon. University Park
Press, Baltimore. vit+ 98 pp., illus. U.S. $9.95.

+Mammal collectors’ manual: a guide for collecting, docu-
menting, and preparing mammal specimens for scientific
research. 1980. By D. W. Nagorsen and R. L. Peterson.
Life Sciences Miscellaneous Publications. Royal Ontario
Museum, Toronto. 79 pp., illus. $6.50.

Mammals: their Latin names explained: a guide to animal
classification. 1979. By A. F. Gotch. Sterling, New York.
DANG p WES a o3-95-

Manual for bird watching inthe Americas. 1979. By D. S.
Heintzelman. Universe Books, New York. 255 pp., illus.
WES Sl7295:

Nearctic sawflies IV, Allantinae: adults and larvae (Hyme-
noptera: Tenthredinidae). 1979. By David R. Smith. Uni-
ted States Department of Agriculture Technical Bulletin
Number 1595. Superintendent of Documents, Washington.
iv+ 172 pp., illus. + plates. U.S. $4.25 plus 25% foreign
handling.

+Observations of wildlife. 1980. By Peter Scott. Cornell
Univeristy Press, Ithaca, New York. 112 pp., illus. U.S.
$19.95.

NEW TITLES 27

Osteology for the archaeologist. Number 3, the woolly
mammoth; number 4, American birds: skulls and mandi-
bles; number 5, North American birds; postcranial skele-
tons. 1979. By Stanley J. Olsen. Papers of the Peabody
Museum of Archaeology and Ethnology, Harvard Univer-
sity, volumes 3, 4, and 5. Peabody Museum, Cambridge,
Massachusetts. x + 186 pp., illus. U.S. $15.

Pacific coast subtidal marine invertebrates: a fishwatcher’s
guide. 1979. By Daniel W. Gotshall and Laurence L. Lau-
rent. Sea Challengers, Los Osos, California. 107 pp., illus.
Cloth U.S. $12.50; paper U.S. $9.50.

Penguins. 1979. By Roger Tory Peterson. Houghton Mif-
flin, Boston. xvit 238 pp., illus. U.S. $25.

*The population ecology of cycles in small mam-
mals. 1980. By James Patrick Finerty. Yale University
Press, New Haven. 260 pp., illus. U.S. $18.50.

Sea otter: core of conflict: loved or loathed. 1979. By Jane
H. Bailey. El Moro, Morro Bay, California. 167 pp., illus.
U.S. $5.50.

*The squirrels of Canada. 1980. By Shirley E. Woods, Jr.
National Museum of Natural Sciences, Ottawa. 208 pp.,
illus. $29.95.

The struggle for survival: the elephant problem. 1979. By
J. Hanks. Mayflower Books, New York. 176 pp. U.S.
$14.95.

+Systematics, osteology, and phylogenetic relationships of
fishes of the ostariophysan subfamily Anostominae (Cha-
racoidei, Anostomidae). 1980. By Richard Winterbot-
tom. Life Sciences Contribution 123. Royal Ontario
Museum, Toronto. 112 pp., illus. $7.50.

+Upper Cambrian to lower Ordovician conodont biostrati-
graphy and biofacies, Rabbit kettle Formation, District of
Mackenzie. 1980. By Ed Landing, Rolf Ludvigsen, and
Peter H. vonBitter. Life Sciences Contributions 126. Royal
Ontario Museum, Toronto. 42 pp., illus. $3.

Watching birds: an introduction to ornithology. 1980. By
R. F. Pasquier. Houghton Mifflin, Boston. 320 pp., illus.
WESaOS 052

Wildfowl 30. 1979. Edited by G. V.T. Matthews and
M. A. Ogilvie. Wildfowl Trust, Slimbridge, England. 176
pp. U.S. $10.

Wildlife of the oceans. 1979. By A. C. Jensen. Chanticleer
Press, New York. 231 pp., illus. U.S. $16.95.

Wildlife survival. 1979. Edited by R. Lockwood. Wild
Canid Survival and Research Centre, St. Louis. 303 pp. No
price given.

+A world list of mammalianspecies. 1980. By G. B. Corbet

and J. E. Hill. Cornell University Press, Ithaca. 226 pp.
WES3>:

228

+The world of the tent-makers: a natural history of the

eastern tent caterpillar. 1980. By Vincent G. Dethier. Uni-
versity of Massachusetts Press, Amherst. 148 pp., illus.
Cloth U.S. $12.50; paper U.S. $5.95.

Botany

Aquatic plants, lake management, and ecosystem conse-
quences of lake harvesting. 1979. Edited by James E.
Breck, Richard T. Prentki, and Orie L. Loucks. Proceed-
ings of aconference, Madison, Wisconsin, February, 1979.
University of Wisconsin Institute for Environmental Stu-
dies, Madison. 436 pp., illus. No price given.

The Audubon Society field guide to North American wild-
flowers: eastern region. 1979. By William A. Niering and
Nancy C. Olmstead. Visual key by Susan Rayfield and
Carol Nehring. Knopf, New York. 863 pp., illus. U.S.
$12.50.

The Audubon Society field guide to North American wild-
flowers: western region. 1979. By Richard Spellenberg.
Visual key by Susan Rayfield and Carol Nehring. Knopf,
New York. 862 pp., illus. U.S. $12.50.

Checklist of the mosses of Canada. 1980. By the Canadian
Botanical Association Checklist Committee, Robert R. Ire-
land, Chairman. Publications in Botany, 8. National
Museum of Natural Sciences, Ottawa. 75 pp. Free.

Flowering plants: willows to mustards. 1980. By Robert
H. Mohlenbrock. Volume 8, the Illustrated Flora of IIli-
nois. Southern Illinois University Press, Carbondale.
xii+ 286 pp., illus. U.S. $18.

*The fossil hunters: in search of ancient plants. 1980. By
Henry N. Andrews. Cornell University Press, Ithaca. 421
pp., illus. U.S. $28.50.

Kosciusko alpine flora. 1979. By A. B. Costin, M. Gray,
C. J. Totterdell, and D. J. Winbush. Collins, Sydney, Aus-
tralia. 408 pp., illus. A. $25.

Toxic plants. 1979. Edited by A. Douglas Kinghorn.
Columbia University Press, New York. ix + 195 pp., illus.
U.S. $20.

*The vascular plants of the continental Northwest Territo-
ries, Canada. 1980. By A. Erling Porsild and William J.
Cody. National Museums of Canada, Ottawa. 676 pp.,
illus. $80.

Wetland functions and values: the state of our understan-
ding. 1979. Edited by P. E. Greeson, J. R Clark, and
J. E. Clark. American Water Resources Association, Min-
neapolis. 700 pp. U.S. $49.

+Wild green vegetables of Canada. 1980. By Adam F.
Szcezawinski and Nancy J. Turner. National Museum of
Natural Sciences, Ottawa. 179 pp., illus. $9.95.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Environment

Acid rain: the North American forecast. 1980. By Ross
Howard and Michael Perley. Anansi, Toronto. 208 pp.,
illus. $16.95.

Advances in marine biology. 1979. Edited by FrederickS.
Russell and Maurice Yonge. Volume 16. Academic Press,
New York. xiit+ 426 pp., illus. U.S. $75.

The Aleutians. 1980. By Lael Morgan. Alaska Northwest,
Edmonds, Washington. 224 pp., illus. + map. U.S. $17.95
plus $1 postage. Separate maps $3.50 plus 75¢ postage.

Aquatic ecosystems: an operational research approach.
1980. By J. E. Beyer. University of Washington Press,
Seattle. 351 pp. U.S. $20.

*Bibliography on the natural history of Newfoundland and
Labrador. 1980. By M. Laird. Academic Press, London.
378 pp. $57.50.

Biochromy: natural coloration of living things. 1979. By
Denis L. Fox. University of California Press, Berkeley.
xiv+ 248 pp., illus. U.S. $24.50.

+The ecological approach to visual perception. 1979. By
James J. Gibson. Houghton Mifflin, Boston. xvi + 332 pp.,
illus. U.S. $18.95.

*Ecology of a subarctic mire. 1980. Edited by M. Sones-
son. Ecological Bulletins Volume 30. NFR, Stockholm. 313
pp., illus. SwCr 125.

Environmental contaminants in food. 1979. By the Office
of Technological Assessment. Superintendent of Docu-
ments, Washington. x + 230 pp., illus. U.S. $5.50 (25%
surcharge for foreign handling).

Energy and environmental issues: the making and imple-
menting of public policy. 1979. Edited by Michael Stein-
man. Lexington Heath, Lexington, Massachusetts.
xii+ 206 pp. U.S. $18.95.

*Environmental impact analysis: a new dimension in deci-
sion making. 1981. By R. K. Jain, L. V. Urban, and G. S.
Stacey. 2nd edition. Van Nostrand Reinhold, New York.
xv + 393 pp. U.S. $27.50.

The functioning of freshwater ecosystems. 1980. Edited
by E. D. LeCrenand R. H. Lowe-McConnell. Cambridge
University Press, New York. 470 pp. U.S. $75.

Handbook of variables for environmental impact assess-
ment. 1979. By Larry W. Canter and Loren G. Hill. Ann
Arbor Science, Ann Arbor. x + 204 pp., illus. U.S. $25.

Kluane: pinnacle of the Yukon. 1980. Edited by John
Theberge. Doubleday, Toronto. 192 pp., illus. $35.

+Public participation in environmental decision making:
strategies for change. 1980. By the Environment Council
of Alberta, Edmonton. Free.

1981]

People, peregrines, and arctic pipelines: the critical battle to
build Canada’s northern gas pipelines. 1980. By D. Pea-
cock. University of Washington Press, Seattle. 224 pp. U.S.
$5.95.

{Selected thematic maps of man’s activities in Canada’s
watersheds. 1980. By B. W. Mitchell. Statistics Canada,
Ottawa. 88 pp., illus. No price given.

A Sierra Club Naturalists’ guide to the Sierra Neva-
da. 1979. By Stephen Whitney. Sierra Club, San Fran-
cisco. xiil+ 526 pp., illus. Cloth U.S. $14.95; paper U.S.
$8.95.

{To whom the wilderness speaks. 1980. By Louise de Kir-
iline Lawrence. McGraw-Hill Ryerson, Scarborough. viii?
180 pp., illus. $14.95.

Track of the grizzly. 1979. By F. C. Craighead, Jr. Sierra
Club Books, San Francisco. 261 pp. U.S. $10.95.

Water conservation alternatives for the North. 1980. By
James J. Cameron and Bryan C. Armstrong. EPS 3-WP-
80-2. Environment Canada, Ottawa. 45 pp.., illus. Free.

NEW TITLES 229

Wilderness economics and policy. 1979. By Lloyd C.
Irland. Lexington Heath, Lexington, Massachusetts.
xx + 228 pp. U.S. $18.95.

Miscellaneous

World dictionary of national parks and other protected
areas. 1979. By the International Union for Conservation
of Nature and Natural Resources, Morges, Switzerland. 2
volumes in looseleaf binder. U.S. $85.

{Nature Saskatoon, an account of the Saskatoon Natural
History Society 1955-1980. 1980. By C. Stuart Houston.
Saskatoon Natural History Society, Saskatoon. 48 pp.
$2.50 plus 50¢ postage.

Exploring the deep frontier: the adventures of man in the
sea. 1980. By Sylvia A. Earle and Al Giddings. National
Geographic Society, Washington. 296 pp., illus. U.S.
$17.20 (Canadian $20.98) includes postage.

* Assigned for review
+ Available for review

Instructions to Contributors

Content

The Canadian Field- Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natu-
ralists or field-biologists reporting observations and results
of investigations in any field of natural history provided that
they are original, significant, and relevant to Canada. All
readers and other potential contributors are invited to sub-
mit for consideration their manuscripts meeting these crite-
ria. 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 significance.

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 sub-
mitted. 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 biblio-
graphic 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. Under-
line only words meant to appear in italics. The names of
authors of scientific names should be omitted except in taxo-
nomic manuscripts or other papers involving nomenclatural
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 Serial Sources for the BIOSIS Data Base,
published yearly by BioSciences Information Service, Phila-
delphia, Pennsylvania 19103. 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 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
K1A 082. 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. Webs-
ter’s New International Dictionary and le Grand Larousse
Encyclopédique are the authonities for spelling.

Illustrations— 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
$50 for each page in excess of six journal pages, plus $5 for
each illustration (any size up to a full page), and up to $50 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 $50 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.

Reviewing Policy of The Canadian Field-Naturalist

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 isa 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

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)
Notes (continued)
Estimating winter defecation rates for Moose, Alces alces
JOHN L. OLDEMEYER and ALBERT W. FRANZMANN

First Canadian records of the Ghost Shiner (Notropis buchanani) and the Orangespotted
Sunfish (Lepomis humilis) ERLING HOLM and GEORGE A. COKER

Reports of Significant Range Extensions

Range extensions for 15 teleost fishes in the Hudson Bay lowlands, Ontario
B. R. ZALEWSKI and J. B. WEIR

News and Comment

Book Reviews

Zoology: A bibliography of British Columbia ornithology, Volume | — Population ecology of
raptors — Love affair with a cougar — Camel quest — Charrs: salmonid fishes of the genus
Salvelinus — The evolution of culture in animals

Botany: The whole fungus: the sexual-asexual synthesis — Grasses of Ontario

Environment: The natural selection of populations and communities — Circulation models
of lakes and inland seas

New Titles

Mailing date of previous issue 20 January 1981

The Ottawa Field-Naturalists’ Club

Honorary Members

208

210

Ze
215

218

222
224

226

C.H. Douglas Clarke
William J. Cody
William G. Dore

R. Yorke Edwards
Clarence Frankton

W. Earl Godfrey
George H. McGee
Hugh M. Raup
Loris S. Russell
Douglas B. O. Savile

Pauline Snure

J. Dewey Soper
Charles M. Sternberg
Mary E. Stuart
Robie W. Tufts

1981 Council

President: R. Taylor R.E. Bedford C.S. Gilliatt
F.H. Bell F.E. Goodspeed
Vice-President: H.L. Dickson D.R. Bewley J.A. Jackson
: D.F. Brunton D. Laubitz
Recording Secretary: E.F. Pope P.M. Catling eo can
Corresponding Secretary: W.K. Gummer W.J. Cody R.C. Montgomery
S. Darbyshire J.K. Strang
Treasurer: B.C. Henson E. Dickson K. Taylor

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 95, Number 2

1981

—— eT

Articles

Tiger Salamanders (Ambystoma tigrinum) and stocked Rainbow Trout (Sa/mo
gairdneri): potential competitors for food in Manitoba prairie pothole lakes
ROBERTA J. OLENICK and JOHN H. GEE

Status and breeding of Mountain Plovers (Charadrius montanus) in Canada
CLIFFORD A. WALLIS and CLEVE R. WERSHLER

Estimates of the standing stocks of fishes in four small Precambrian Shield lakes
J. M. FRASER

A test of the peninsular effect on species diversity
ROBERT J. TAYLOR and LEE A. PFANNMULLER

Population characteristics and movements of Striped Skunks (Mephitis
mephitis) in central Alberta
RONALD R. BJORGE, JOHN R. GUNSON, and WILLIAM M. SAMUEL

Natural history of the Ebony Spleenwort, Asp/enium platyneuron (Aspleniaceae),
in the Great Lakes area WARREN H. WAGNER, JR, and DAVID M. JOHNSON

Distributional history of Juncus compressus (Juncaceae) in North America
RONALD L. STUCKEY

Le régime alimentaire du Coyote (Canis /atrans) et du chien errant (C. familiaris)
dans le sud du Québec JEAN-MARIE BERGERON et PIERRE DEMERS

Variation in frequencies of pelvic phenotypes of the Brook Stickleback,
Culaea inconstans, in Redwater drainage, Alberta JAMES D. REIST

Feeding and social behavior of some migrant shorebirds in southern Manitoba
RICHARD A. WISHART, PATRICK J. CALDWELL, and SPENCER G. SEALY

Distribution, growth, and foods of Arctic Cod (Boreogadus saida) in the Bering,

Chukchi, and Beaufort seas LLOYD F. LOWRY and KATHRYN J. FROST
Polymorphism in colonies of the land snail Cepaea nemoralis at London,

Ontario: changes over three decades S. M. SINGH
Scheduling censuses of breeding White Pelicans (Pelecanus erythrorhynchos)

in northern Alberta RICK D. BEAVER and VICTOR LEWIN
Notes

Nesting Northern Gannets (Morus bassanus) killed by rock falls at Great Bird Rock, Quebec
IAN R. KIRKHAM

Postglacial fossil fishes from Coppermine River, Northwest Territories, Canada
DON E. MCALLISTER and DENIS ST-ONGE

Gulls robbing prey from Great Blue Herons (Ardea herodias),
T. E. QUINNEY, B. N. MILLER, and K. R. S. QUINNEY

First record for Canada of the bat mite Spinturnix globosus and a new host,
Myotis lucifugus HUGH C. SMITH

129

133

137

144

149

156

167

We

178

183

186

192

198

202

203

205

206

concluded on inside back cover

ISSN 0008-3550

ahs /Ze4 “US. Comp, ZOOL.

The CANADIAN * *%

SRVARD

FIELD-NATURALIST

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

ce

Volume 95, Number 3 July-September 1981

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

C2 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: Patricia A. Lalla Chairman, Publications Committee: R. E. Bedford
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.

Production Manager: Pauline A. Smith, R.R. 3, Wakefield, Quebec JOX 3G0
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

Associate Editor (Ornithology): Dr. A.J. Erskine, Canadian Wildlife Service, Box 1590, Sackville, New Brunswick E0A 3C0
Editor: Dr. Lorraine C. Smith, R.R. 3, Stittsville, Ontario KOA 3G0

All manuscripts intended for publication should be addressed to the new Editor:
The Canadian Field-Naturalist, The Ottawa Field-Naturalists’ Club, Box 3264, Postal Station C,
Ottawa, Ontario Canada KIY 4J5

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 (613-996-1665).

Cover: Deer Mouse, Peromyscus maniculatus, photographed in mixed forest near Burritts Rapids, Ontario, on 13 August
1971 by Donald A. Smith. See Article on page 319.

Volume 95, Number 3

The Canadian Field-Naturalist

Editorship of The Canadian Field-Naturalist

It is with regret that The Ottawa Field-Naturalists’ Club announces the decision of
Dr. Lorraine C. Smith not to accept reappointment as Editor of The Canadian
Field-Naturalist. Dr. Smith has edited the journal efficiently, effectively, and with
great dedication for nine years, a period almost double that of her recent predecessors.
In two decades The Canadian Field- Naturalist has grown extensively in both size (200
pages annually in 1960 to 500 pages annually now) and stature. The former is largely
dependent upon our authors; the latter is due in no small measure to Lorraine. During
her tenure she successfully guided The Canadian Field- Naturalist to its present role as
an important medium for recording the results of research and field observations in
natural history. With a full-time professional appointment of her own, Lorraine’s
work as Editor had to be pursued in her “spare time,” of which in consequence she had
little to devote to other interests. The Ottawa Field-Naturalists’ Club and, we are sure,
all readers of The Canadian Field- Naturalist thank Lorraine sincerely for her long and
devoted service as Editor and wish her success and fulfillment in her future endeavors.

ROGER TAYLOR

President, The Ottawa Field-Naturalists’ Club
R. E. BEDFORD
Chairman, Publications Committee

231

July-September 1981

Editor’s Thanks

As Editor of The Canadian Field- Naturalist, |had the great fortune and pleasure of
‘meeting’ many interesting and nice people. Moreover, the rapport that developed
between many authors and referees and me was welcome and very gratifying. I have
enjoyed particularly happy relationships with the Associate Editors and I sincerely
appreciate the fine job they have done in evaluating the scientific worth of manus-
cripts. During my editorship I received much valued expert advice from Earl Godfrey,
Pete Peterson, Bob Wrigley, Stan Rowe, Herb Lawler, Ernie Brodo, Tony Keith, and
Bob Risebrough (all retired); Ed Bousfield and Dave Scott (retiring with me); and
Charley Bird, Francis Cook, Tony Erskine, Charles Jonkel, Charley Krebs, George La
Roi, Bill Pruitt, and Steve Smith (all continuing to serve the journal). My sincere
thanks to them all for helping me maintain the scientific quality of the journal.

I also put on record here my heartfelt thanks to all those other persons who have
helped me with the journal but have received little formal recognition for their
contributions. Marilyn Dadswell, the journal’s first Copy Editor, served for almost
eight years (1973-1980) and did a fine job in helping upgrade its technical quality.
Patricia Lalla kindly agreed to carry on as Copy Editor during the remainder of my
editorship. When Joe Dafoe offered to lend a much needed helping hand in 1975, I
appointed him the journal’s first Production Manager. Late the following year Joe left
for Australia, so | asked Pauline Smith (no relation) to carry on this important work.
She will now be leaving the journal after conscientiously seeing five years of issues into
print: I have always been deeply appreciative of her devoted and competent service.

Diligent and careful work on the useful annual indices during my editorship was
completed by Stan Teeple, Emerson Whiting, and Harvey Beck. The book review
section, under Iola Price, Anne Dagg, and since July 1975 under the enthusiastic and
dependable Wilson Eedy, has become an important component of the journal. I am
grateful too to Business Manager Bill Cody who has contributed significantly not only
to his particular parts of the operation but to other aspects as well, and to Emil Holst
who has served as our very affable contact at M.O.M. Printing.

And finally I give my very special thanks to my husband Don whoas Assistant to the
Editor has given me continuing support through his advice and help in scientific,
editorial, cartographic, and clerical matters and also total backing in controversial
and emotional issues. ]

I am indeed indebted to my many associates for what they have done and I thank
them all for their help, time, and for the pleasure and knowledge I have gained in being
associated with them.

LORRAINE C. SMITH
Editor (1972-1981)

2B

Editor’s Farewell Musings

For historical and general interest reasons, I decided to document what the editorship of The Canadian
Field- Naturalist has entailed for me, the changes initiated, major decisions made, actions taken, and what I
consider my important accomplishments. !

Normally a scientific editor’s role is to evaluate, with the help of referees who are chosen because they are
specialists in certain fields of study, the scientific content of submitted manuscripts and to accept for publication
material that significantly advances scientific knowledge within the scope of the journal. Most editors take
special pride in the results of their endeavors and I certainly feel that the issues of The Canadian Field- Naturalist
produced under my editorship have been of high quality. Moreover, as the person responsible for the content of
the journal, I tried to ensure not only that papers conformed to certain standards (i.e., that they met the criteria
for acceptance) but that their presentation was as understandable as possible to the average reader.

The attention an editor gives to each submitted paper differs from journal to journal and from editor to editor
of the same journal. Some editors rely completely on the chosen referees (and advice if needed from members of
their editorial boards) and hence never actually read the submitted papers. Because scientific editors are
sometimes appointed for only short time spans and must carry on as well the work associated with their regular
permanent positions, many don’t take extra time with each paper. Although The Canadian Field- Naturalist isa
general journal and no one person can be knowledgeable in all the areas it covers, I have always read each
submitted paper whatever the subject. It is usually possible to evaluate generally the research or observations
upon which a paper is based, to discern whether the contribution is placed in the proper perspective concerning
the state of knowledge of the subject, to determine if it is clear and concise, and if its organization is logical and
geared to the stated purpose. Moreover, I think that to control the quality of the accepted papers, an editor
should try to ascertain that the results are valid, accurate, and important, and that the language and presenta-
tion are comprehensible. Although to me the quality of writing is important, others have argued that if the new
information is communicated to readers, other ‘improvements’ are unnecessary. Nevertheless I believe the final
published products have been better because of my editorial assistance to authors. It is obvious to many of them
too that I cared.

When I somewhat reluctantly agreed to take on the editorship of The Canadian Field- Naturalist in 1972, in
addition to the scientific editing and processing of manuscripts, I assumed other unexpected responsibilities. I
was also the office staff (filing, keeping records, and typing), production staff (preparing a mockup of the
journal, correcting proofs, and seeing the operation through to the final production stage), and copy editor
(marking manuscripts for the printer; correcting the spelling, punctuation, grammar, and English and French
composition; and reading the galley proofs). Unfortunately, there has been a decline in general competence in
writing in the English and French languages and authors who lack good communications skills benefit
particularly from the skills and experience of editors and copy editors. Some of my first chores as Editor also
included starting a much-needed referee file and preparing standard letters, cards, and other forms for routine
correspondence.

Because there were so many extra duties to perform, the position was a great deal more time-consuming than I
had imagined. Moreover, the location and maintenance of the editorial office in our private home put
considerable constraints on obtaining clerical, typing, and other assistance. Unfortunately, because it has been
difficult for others to imagine the extent of the enormous work load, it has also been difficult to obtain help,
needed support, or approval for funds to pay assistants. Eventually the publisher approved a token remunera-
tion for a Copy Editor and I was able to obtain the services of an experienced person in 1973. By 1975, I had
arranged for a volunteer Production Manager. Even so, the time I continued to expend editing The Canadian
Field- Naturalist has been so extensive (more than 800 hours per year) that I have had to forgo most other
activities.

Throughout my editorship, I have strived to improve the format and style of the journal; some of these
innovations are recorded in my annual published “Editor’s Reports.” In 1977 I published an editorial detailing
current editorial policy. A new series entitled “The Biological Flora of Canada” was launched in 1979 and a new
method of reporting some range extensions under the heading “Reports of Significant Range Extensions” was
started in 1980. Several manuscripts for these sections are in process and others continue to be submitted; |
expect the series will soon be well-established components of the journal. In late 1973 I submitted the first

1Additional detailed notes are on file with The Ottawa Field-Naturalists’ Club.

233

234 THE CANADIAN FIELD-NATURALIST Vol. 95

material supplementary to one of our published journal papers to the Depository of Unpublished Data (located
at Canada Institute for Scientific and Technical Information, National Research Council of Canada, Ottawa).
Other relevant material has been deposited there from time to time and is always available to any interested
person requesting it.

The Canadian Field- Naturalist was at last selected for inclusion in Current Contents/ Agriculture, Biology,
and Environmental Sciences starting in 1977, and it was included as a “source publication” in the Science
Citation Index starting in 1979. These last two actions signify formal acceptance of The Canadian Field-
Naturalist as a primary scientific journal by the North American scientific community.

Although well established and now widely recognized as a scientific journal, The Canadian Field- Naturalist is
in the rather anomalous position of being published, not by a scientific society, but by The Ottawa Field-
Naturalists’ Club, a local natural history club. In the past many professional biologists have served on the club’s
Council but hardly any do so anymore. This change has serious implications for The Canadian Field- Naturalist
in the long run because the Council appoints (and can dismiss) editors and otherwise can have considerable
influence on the journal. For example, the Ad Hoc Committee on Club Publications, approved in principle by
the Council in November 1978 as a result of unspecific rumors of “unease” within the club about The Canadian
Field-Naturalist, and set up in November 1979, had a narrow and restricted data base for its deliberations
because opinions of editors, managers, subscribers, readers, and authors were not solicited. This committee
presented its report to Councilin November 1980. Eventually its recommendations, perhaps ina modified form,
will be approved by the Council as club policy. Some of the recommendations are blatantly contrary to current
editorial policy and practices, some are real infringements on editorial freedom, some cast doubt on the
capabilities of some Associate Editors (based it seems on mistaken hearsay evidence), and others have the
appearance of being new when, in fact, they represent the status quo. And yet the club’s policy on this journal
(and other publications) will be decided by a Council most of whose members and executive have little
knowledge, understanding, or appreciation of what primary biological publication is all about. Will this be
acceptable to those the journal now serves?

Because of my reluctance to continue the tremendous work load I have carried in my “spare time” for nine
years of journal issues and because of what I have perceived as the lack of trust, appreciation, and support by the
club, I informed the Council that took office in January (I had already alerted the Publications Committee
Chairman in mid-1980 about this possibility) that my term of office was completed and I would decline a
reappointment. I did, however, agree to put out the present issue; this will thus be the last appearing under my
editorship. I wish readers to know, however, that it was with considerable regret that I took this action not only
because the journal has become an important part of my life but also because feedback from the scientific
community has assured me that the high quality and standards I instituted and my personal attention to
improving manuscripts have been deeply appreciated.

Through expenditure of considerable time and effort during my term of office, | have applied my knowledge
and ability to upgrade the scientific and technical quality of The Canadian Field- Naturalist. One of my first and
probably the most important of my major policy changes was to adopt a consistent and rigorous peer review
system by commencing to send each submitted manuscript to at /east two referees. Previous editors had either
not done this or had done so only occasionally; most manuscripts submitted to them were accepted either after
evaluation by only an Associate Editor or surprisingly often with no external review at all. Another important
policy change was to limit the contents of the journal to information that is original (i.e., confirmatory material
although of importance to have on record was directed elsewhere), scientifically sound, and of importance at the
national level. Moreover, I have always helped authors to convey their messages including interpretations of
their observations as clearly, concisely, and intelligibly as possible.

It is particularly important to me that my successors should try to maintain the scientific and technical quality
I have built up. Of course, the cornerstones of a successful primary scientific journal are a good editor and a
functional peer review system. That is why I strongly recommended that the new editor should be an
experienced and competent biologist (preferably with a Ph.D.) who is highly respected by other scientists and
who is responsible, well organized, efficient, dependable, dedicated to the reporting of good science, dedicated
to the expenditure of considerable time for the sake of communicating good science via the journal, tactful,
diplomatic, and perhaps what is most important of all, able to exercise sound judgment. This latter quality is
especially needed to determine if there is new knowledge worth reporting and if its presentation meets high
scientific and technical standards. Of course, an editor should have good communication skills to maintain
credibility. Also, for a generalist journal, an editor should have broad interests and be widely experienced in
science especially for choosing the right referees. Nevertheless, | was informed recently by two members of the

1981 SMITH: EDITORIAL 235

current Search Committee (neither I as retiring Editor nor any other scientist with specialized training in
biology were asked to serve on this committee) that because of expediency almost the sole criteria for the
choosing of a new Editor became availability, willingness to serve, release time by the employer to carry out
editorial duties during office hours, and the promise of secretarial help from the employer. Surely if new editors
are to be chosen on a hit-or-miss basis by laymen-naturalists using or emphasizing such criteria, however
important they may be, rather than those listed earlier, the journal could sooner or later run into serious
problems of quality and credibility.

Although there has been a long association of The Ottawa Field-Naturalists’ Club and The Canadian
Field- Naturalist, 1 strongly urge that concerned readers and authors consider the possibility of an alternative
publisher, a science-based one that will vigorously support a journal reporting observations and results of
investigations in all fields of natural history provided that they are original, significant, and relevant to Canada.
Of course such a hypothetical new publisher should be financially sound and otherwise stable as well. Perhaps,
too, another publisher would give more support to the Editor in delegating to him or her the responsibility for
the journal. At least the time-consuming local club politics and misdirected unjust criticisms, which presumably
resulted from the restricted interests and lack of knowledge by many members of the club’s Council regarding
primary publication of biological information and which necessitated large unproductive portions of our
valuable time to be taken up with defensive actions, might no longer exist. Of course, the club has gained in
stature from its role as publisher of this journal and its predecessors (since 1880) and it may not wish to
relinquish control. Nevertheless, the burden of being accountable to the scientific natural history community for
a national journal and especially the serious responsibility of choosing new editors may now be more that this
local club should retain. Therefore, in the future if the journal’s reputation is not maintained, and there is no
move fora change in publisher, I challenge those who want a top-quality Canadian natural history journal, such
as The Canadian Field- Naturalist is now, to initiate a new journal.

Because I have dedicated myself to its scientific and technical editing for so long and because I have a vested
interest in the future of The Canadian Field- Naturalist, | sincerely hope that this journal or its replacement will
continue to fill an important role communicating information of scientific worth in natural history and that it
will do so ina readily comprehensible and retrievable manner. Moreover, I hope that rational procedures can be
instituted to ensure that the current scope, quality, and standards of the journal, and its consequent support by
authors and recognition by the scientific community will be maintained.

There is no doubt that I will miss the journal and the enjoyable contacts I have had with authors, referees,
Associate Editors, and my other coworkers on The Canadian Field- Naturalist. Many thanks to those who have
helped and supported me during my editorship.

LORRAINE C. SMITH
Editor (1972-1981)

Is the Impact Factor a Meaningful Index for the Ranking of

Scientific Research Journals?

ROGER TAYLOR

Physics Division, National Research Council of Canada, Ottawa, Ontario KIA 0R6

Taylor, Roger. 1981. Is the Impact Factor a meaningful index for the ranking of scientific research journals? Canadian

Field-Naturalist 95(3): 236-240.

In a given year the Impact Factor for a journal is defined as the number of citations to papers published in the previous two
years in that journal divided by the total number of these papers. This is often used as a measure of a journal’s effectiveness. I
have shown that, as defined, the Impact Factor can give a distorted view of the worth of some journals. In particular,
field-biology journals such as The Canadian Field-Naturalist seem to be badly misrepresented by this index. 1 propose an
alternative definition of the Impact Factor which is a more realistic measure of the importance of a journal to the scientific
community. The two éorrections that I suggest take account of the fact that (a) some disciplines, notably field biology, takea
much longer time from conception to completion of a project and (b) the average length of a paper varies considerably from

journal to journal.

Key words: Impact Factor, citation, field-biology, journal ranking.

The Scientific Citation Index, published annually
since 1963 by the Institute for Scientific Information
in Philadelphia, has become over the intervening
years a widely used tool for evaluating the usefulness
of published papers to the scientific community. It
analyses all citations contained within papers pub-
lished in a set of “source journals” (currently about
3000) and groups them in various ways. Perhaps the
best-known and most commonly utilized grouping is
by author of cited articles. Despite some very obvious
criticisms, the ranking of scientists by number of ci-
tations is clearly more meaningful than ranking by
number of articles published, and this procedure is
becoming widely accepted. Hence it should come as
no surprise that similar ranking procedures are
becoming adopted for journals. Starting with 1975 the
Institute for Scientific Information has included in the
Science Citation Index a volume entitled Journal Ci-
tation Reports (Garfield 1976a) which contains ana-
lyses of scientific journals and their impact on the
scientific community. Among other things, the data
contained in this publication make it possible to rank
scientific journals in two ways. One is by the total
number of citations received and the other is by
Impact Factor. The Impact Factor is basically a ratio
between citations and citable items published. It is
computed for any particular journal for the year 1979,
say, in the following way. The total number of ci-
tations, in all journals in 1979, of 1977 and 1978
papers published in the particular journal ‘X’ is
divided by the total number of these same 1977 and
1978 papers to give the Impact Factor. For example, if
X published 100 articles in 1977 and 1978 and if in
1979 these articles were cited 48 times, the 1979
Impact Factor for X would be 0.48. There are many

other possible choices of Impact Factor and some of
these are discussed by Garfield (1972), but this index
clearly provides a more meaningful basis for journal
comparison than a simple listing of numbers of jour-
nal citations. To put things in perspective, an Impact
Factor of 1.00 is considered to be good.

In discussing relative assessments of scientific jour-
nals Garfield (1972, 1976a, b) emphasizes that caution
should be used and that statistics should not be taken
out of context. The very nature of a journal can appre-
ciably affect its citation frequency (e.g., compare
Scientific American with Journal of Molecular Biol-
ogy) and different fields are not always comparable.
Nevertheless the Impact Factor is an easily accessible
ranking parameter, and there is a temptation to use it
without due consideration of circumstances. Recog-
nizing this fact it is clearly desirable to construct an
Impact Factor which takes into account, in some way,
fundamental differences between the journals that are
being studied. This is particularly important when
making comparisons such as those in The Press of
Knowledge (Royal Society of Canada 1978).

The purpose of this article is to point out that, as _
presently defined by the Science Citation Index, the
Impact Factor badly misrepresents some areas of
science such as field biology and to suggest an alterna-
tive definition that seems to allow for a much more
meaningful comparison of journals from different
fields. This is particularly important for journals such
as The Canadian Field- Naturalist.

The Press of Knowledge, commissioned by the
National Research Council of Canada and prepared
by the Royal Society of Canada, is the report of a
committee that undertook a study of policies and
practices for publication of research journals in Can-

236

1981

ada. The committee noted that there were 67 primary
scientific, medical, and engineering journals pub-
lished in Canada and that 38 of them were included
among the 2630 source journals analyzed in the 1975
issue of Journal Citation Reports (Garfield 1976a). In
Table II of The Press of Knowledge these 38 journals
are ranked according to total number of citations in
1974 and to 1974 Impact Factors. The latter ranges
from a high of 1.671 for the Canadian Journal of
Biochemistry to a low of 0.051 for the Canadian
Aeronautical Space Journal. By implication those
other 29 Canadian journals that are not included in
this list (e.g., The Canadian Field- Naturalist) should
be considered to have impact factors even lower than
0.051. In fact, there is no further mention of any of the
29 non-source journals in The Press of Knowledge
and it is hard not to draw that inference of inferior
quality, even if it was not intended by the authors of
the report. This is simply one example of a situation
where journals from vastly different fields have been
compared using Impact Factor, and the points | will
make in the following paragraphs are first that the
currently defined Impact Factor is a poor basis for
such a comparison and second that if such compari-
sons must be made, a much better index can be used.
If we focus our attention on The Canadian Field-
Naturalist, we find that we have to turn to the year
1978 to obtain an Impact Factor because it does not
appear to be reported in earlier years. In the journal-
ranking package of the 1978 issue of Journal Citation
Reports (Garfield 1979), the value 0.127 is given for
this quantity, marginally better than 0.051 but cer-
tainly not an impressive figure. However, we must
keep in mind that because The Canadian Field-
Naturalist is not a source journal, this figure does not
contain self-citations, so that it is clearly low. To
generate the 1978 Impact Factor the total number of
citations of papers published in 1976.and 1977 should
be divided by the total number of papers (referred to
as source items) published in the same two years, but
for some unexplained reason only the last two issues
of 1976 were included in the figures. This again is a
misrepresentation since 1976 articles are more likely
- to becited in 1978 than are 1977 articles. Therefore all
four 1976 issues should have been included. To correct
these deficiencies in the published Impact Factor, I
have determined the number of self-citations in 1978
of 1976 and 1977 papers and doubled the number of
citations for 1976 articles. The results are summarized
in Table | which shows that the corrected Impact
Factor, hereafter referred to as Jo, is equal ‘to
44/209 = 0.211. I shall take this to be the Impact
Factor as generated by the Science Citation Index
prescription.

Now if we look carefully at The Canadian Field-

TAYLOR: VIEWPOINT

23)

TABLE 1|—Number of citations in 1978 to papers published in
The Canadian Field- Naturalist

Citations S
ource
Year Self Other Total items”
1976 7 30° 37 110
1977 3 4 7 99
1976-77 10 34 44 209

“The total number of scientific papers (excluding editorials,
book reviews, news and comment etc.) published by The
Canadian Field- Naturalist in the indicated year(s).

’Because the number of citations for 1976 in the 1978
Science Citation Index includes only two issues, this
number has been doubled to represent the entire year.

Naturalist, it readily becomes apparent that there are
two factors which work to produce a low Impact
Factor. The first of these is the unusually high number
of notes making up the content of the journal. Of the
110 Source Items in volume 90 (1976 year) of the
journal, only 41 are full-length articles and 69 are
notes. The articles cover 310 pages whereas the notes
cover only 131 pages. The average article in that
volume, therefore, is about seven and a half pages long
and the average note only about two pages. This ratio
of articles to notes is certainly not typical of a scientific
journal. Generally speaking, within a given journal,
the probability of an article or note being cited will be
roughly proportional to its length. Of course some
short papers can have major impact and be cited
heavily whilst other verbose papers on obscure topics
can be entirely ignored, but, on average, if ajournal is
carefully edited, a long paper will contain more citable
information and refer to a broader field than a short
one. Therefore, each individual note will draw fewer
citations than an article although it is reasonable to
assume that any group of four notes is likely to draw
about the same number of citations as an article.
Hence the presence of an unusually large number of
notes in The Canadian Field- Naturalist will result ina
disproportionately small Impact Factor because each
note is treated as a source item despite the lower
citation probability, which has nothing to do with
scientific content.

The second factor working to the detriment of The
Canadian Field- Naturalist Impact Factor is the very
nature of the research that is published therein. It is
exclusively field biology which frequently requires a
much longer time between conception and completion
of a project than does laboratory work. Inreferring to
a chronological distribution of cited papers Garfield
(1972) stated, “An analysis of this distribution has
shown that the typical cited article is most heavily
cited during the 2 years after its year of publication.”
One has only to peruse the citation pages of any issue

238

of The Canadian Field- Naturalist to appreciate that
this is certainly not true for that journal and it is
obviously because of the nature of the research. Gar-
field’s comment clearly refers to laboratory work. To
confirm this difference between laboratory and field
work, I plotted histograms of the chronological dis-
tribution of citations occurring in the 1978 issues of
The Canadian Field- Naturalist and of the February,
May, August, and November 1978 issues of the Cana-
dian Journal of Physics. These are shown in Figure |
(A and B). Consistent with Garfield’s comment, the
Canadian Journal of Physics distribution of citations
shows that the peak years are 1976 and 1977 whereas
for The Canadian Field- Naturalist they are 1973 and
1974. As a test of my hypothesis, I also plotted a
histogram of the corresponding distribution for the
February, July, and November 1978 issues of the
Canadian Journal of Zoology which contains a mix of
laboratory and field work (Figure IC). There a
double-peaked distribution seems to be present, with
the peaks centered on 1975-1976 and 1972-1973. This
distribution appears to be intermediate between the
two foregoing ones and this I confirmed by isolating a
subgroup consisting of those papers which reported

exclusively field work. This subgroup of the Canadian ,

Journal of Zoology papers gave a distribution, peaked
in 1973, which was very similar to that of The Cana-
dian Field- Naturalist (Figure 1A). The remainder, by
the way, consisting of laboratory or laboratory plus
field work, gave rise to a distribution quite similar to
that of the Canadian Journal of Physics but with the
peak years in 1976 and 1977 and with an appreciably
smaller peak in 1973. Clearly then, to base an Impact
Factor for 1978 solely on citations for 1976 and 1977
articles does not do justice to journals with a signifi-
cant amount of research requiring long periods
between conception and completion.

I can suggest two corrections that can be applied to
the Impact Factor Jo to make it more meaningful to
compare journals of different disciplines. To correct
for the problem caused by unusually large numbers of
notes, we could, first of all, define a standard source
item as containing, say, 7500 words. Then we could
compute the number of standard source items from
the number of words per page and the number of
pages for the years under consideration. Using the
number of standard source items to compute the
Impact Factor gives an effective Impact Factor,
hereafter referred to as /;. I estimate the number of
words per page for The Canadian Field- Naturalist to
be 750, the number of pages in 1976-1977 to be 797
(This excludes editorials, book reviews, etc., which are
not included in the Science Citation Index) and there-
fore the number of standard source items to be 79.7).
Hence /; = 4/79.7 = 0.552, a much more respectable
looking number than /. It is interesting to compare Jo

THE CANADIAN FIELD-NATURALIST

Vol. 95

A

Lora The Canadian Field -Naturalist

25 | B

Canadian Journal of Physics

Canadian Journal of Zoology

1978
=|
1969 +
1964-4
1959 +
1954 4-
1940-1947
1930-1939
before 1930

FIGURE |. The chronological distribution of citations con-
tained in papers published in (A) The Canadian Fiela-
Naturalist in 1978 (all four issues), (B) the Canadian
Journal of Physics in 1978 (February, May, August,
and November issues), (C) the Canadian Journal of
Zoology in 1978 (February, July, and November
issues).

1981

and /; for 1978 for eight of the Canadian Journals of
Research published by the National Research Council
of Canada. The results are displayed in Table 2. They
are based on anestimated 900 words per page for these
journals. As one might expect, in most cases, the
change from J) to /; is not very large because the
numbers of notes are relatively small. The most note-
worthy change, relatively speaking, is for the Cana-
dian Journal of Earth Sciences where Io = 0.980 and
I, = 0.697. Apparently earth scientists write longer
articles than do those in other disciplines.

TABLE 2—1978 Impact Factors for eight of the Canadian
Journals of Research. J) is computed using the SCI definition
and /, is computed using the number of standard source
items (see text).

Journal Ih /,
Canadian Journal of Physics 0.982 0.971
Canadian Journal of Chemistry 1.516 1.765
Canadian Journal of Earth Sciences 0.980 0.697
Canadian Journal of Botany 0.986 0.897
Canadian Journal of Zoology 0.879 0.890
Canadian Journal of Physiology

and Pharmacology 1.308 1.469
Canadian Journal of Biochemistry 1.543 1.874
Canadian Journal of Microbiology 1.073 1.296

To correct for the problem of the different chrono-
logical distributions I suggest that for each journal the
percentage of all citations occurring for 1976-1977 be
computed and used as a basis for renormalizing the
1978 Impact Factor to give a new index called /2. This
can of course be done for any year. For The Canadian
Field- Naturalist this percentage is 8.6%, for the Can-
adian Journal of Zoology it is 13.5%, and for the
Canadian Journal of Physics it is 19.4%. If we choose
arbitrarily to say that 19.4% is what we would expect
for an “average” journal, then we compute /, for The
Canadian Field-Naturalist by multiplying /, by
0.194/0.086. The results for J), /,, and /, for all three
journals are displayed in Table 3 where we see that for
The Canadian Field- Naturalist, I, is avery respectable
1.240, putting this journal roughly on a par with the

TABLE 3—1978 Impact Factors for three journals including a
correction for the chronological distribution of citations (/,)

Journal hy /, I,
Canadian Journal of Physics 0.982 0.971 0.971
Canadian Journal of Zoology 0.879 0.890 1.279
The Canadian

Field-Naturalist 0.211 0.552 1.240

TAYLOR: VIEWPOINT

239

two Canadian Journals of Research. In other words
the two suggested corrections actually increase the
Impact Factor for The Canadian Field- Naturalist by a
factor of 6, and by a factor of about 1.5 for the
Canadian Journal of Zoology whilst making no
change for the Canadian Journal of Physics.

Implicit in the correction leading to /, is the
assumption that the chronological distribution of ci-
tations for articles published in journal X will be
about the same as the distribution of citations within
the same journal. This is not an unreasonable assump-
tion because both distributions reflect activity within
virtually the same field of scientific research. Any
further corrections to take into account the small
differences between the two distributions would be
much more complicated and would be unlikely to
produce significant alterations to /,.

The foregoing arguments are not a thinly disguised
attempt to make The Canadian Field- Naturalist look
good. They should be regarded as some suggestions
for correcting what is clearly a poor procedure for
comparing journals of different disciplines. The
example of The Canadian Field- Naturalist shows how
poor this can be. The chief proponent of the Impact
Factor concept, Garfield, would not recommend mak-
ing such comparisons, judging by many of his remarks
(Garfield 1972, 1976b). Nevertheless, as long as the
data are there, these comparisons will be made.
Although, in the report of the Royal Society of Can-
ada (1978), the committee warns against using the
Impact Factor as the sole basis of judgment, the very
fact that they rank the journals in this way in tabular
form and dismiss those journals for which no data
were available make it all too easy for others to come
to poorly conceived conclusions. By using the renor-
malized effective Impact Factor /,, the data can be
presented in a much more equitable fashion.

Asa final comment, a glance at the journal-ranking
package contained in Journal Citation Reports (Gar-
field 1979) reveals that the top-rated journals are of a
review nature with Impact Factors greater than 20. If
the above procedures were used, these Impact Factors
would be cut down to a size which would more truly
reflect their importance to science. I very much doubt
that any scientist when asked to name the three most
important journals for his research would include a
review journal in the list.

Literature Cited

Garfield, E. 1972. Citation analysis as a tool in journal
evaluation. Science 178: 471-479.

Garfield, E. 1976a. Journal Citation Reports, a bibliomet-
ric analysis of reference processed for the 1974 Science
Citation Index. Science Citation Index 1975 Annual 9, I.

Garfield, E. 1976b. Significant journals of science. Nature
264: 609-615.

240 THE CANADIAN FIELD-NATURALIST Vol. 95

Garfield, E. 1979. Journal Citation Reports, a bibliometric Practices for Research Journals and Other Periodicals in
analysis of science journals in the ISI data base. Science Science and Engineering in Canada. 59 pp.
Citation Index 1978 Annual 13.

Royal Society of Canada. 1978. The Press of Knowledge: a Received | December 1980
report by the Committee on Publication Policies and Accepted 7 March 1981

Vegetation with Atlantic Coastal Plain Affinities in
Axe Lake, near Georgian Bay, Ontario

P. A. KEDDY

Department of Botany and Genetics, University of Guelph, Guelph, Ontario NIG 2W1

Keddy, P. A. 1981. Vegetation with Atlantic coastal plain affinities in Axe Lake, near Georgian Bay, Ontario. Canadian
Field-Naturalist 95(3): 241-248.

Axe Lake is a small, sandy lake about 25 km NW of Huntsville, Muskoka District, Ontario. Many species with eastern or
Atlantic coastal plain affinities occur in the lake, including Muhlenbergia uniflora (One-flowered Dropseed), Juncus militaris
(Bayonet Rush), Nymphoides cordata (Floating-heart), Rhexia virginica (Meadow-beauty), Rhynchospora capitellata
(beak-rush), Woodwardia virginica (Virginian Chain-fern), Utricularia purpurea (Purple Bladderwort), and Xyris carolini-
ana (Yellow-eyed Grass). The lake occurs on the edge of a sand plain which marks the former high waterline of postglacial
Lake Algonquin. The vegetation of this lake may thus be of a relict shoreline type which was formerly much more widespread
in the sandy bays of postglacial Lake Algonquin. Extensive beds of emergents and shallow water aquatics occur on the gently
sloping shores. During July 1979, quantitative data on species composition were collected, using 25 transects run from 0.5 m
above to 0.5 m below the waterline. These transects represented much of the variation in species composition and substrate
types occurring in Axe Lake (except for floating bog mats and rock outcrops). The transects were divided into three categories
based on the substrate type of the shoreline at 0.5 m above the waterline: sand (mean = 0.9% organic matter), transition
(mean = 3.7% organic matter), and peat (mean = 16.4% organic matter). Species composition varied markedly among these
categories. Many coastal plain species (such as R. virginica, X. caroliniana, and M. uniflora) reached their maximum
abundance in the transition areas. Few occurred in Sphagnum bogs elsewhere on the lakeshore. Axe Lake appears to be a
small remnant of a larger open water area, most of which is now Sphagnum bog. Thus, it appears that bog formation is
gradually eliminating a relict type of shoreline vegetation.

Key Words: aquatic plants, coastal plain flora, Georgian Bay, Lake Algonquin, phytogeography, rare species, shoreline

vegetation, substrate type.

It has long been known that areas about the Great
Lakes, particularly the southern end of Lake Michi-
gan and the sand barrens of northwestern Wisconsin,
have floristic affinities with the Atlantic coastal plain
of the eastern United States (Peattie 1922; McLaugh-
lin 1932). The Atlantic coastal plain is a relatively flat
marginal area of eastern North America which
extends as a narrow band up the eastern coast between
the Applachian Mountains and the Atlantic Ocean; it
ranges in width from a few kilometres on Cape Cod to
up to 250 km in Georgia. It is often poorly drained,
with extensive wetlands; as well, acid soils are fre-
quent (Fernald 1942). Fernald (1942) was critical of
the use of the term “Atlantic coastal plain species” to
describe many of the Great Lakes disjuncts, noting
that the term was overused, and that “the true
members of the Atlantic coastal plain flora are rarely
found off the coastal plain.” Although some of the
species listed by Peattie (1922) and McLaughlin
(1932) are questionable, there are still many species
known to occur along the Atlantic coastal plain, with
extensions northwards into eastern Canada, and
inland to the Great Lakes. In spite of Fernald’s admo-
nition, the use of the term “coastal plain species”
empahsizes the geographical affinities of these species,

and thus seems useful to retain in its broader sense
(e.g., in Roland and Smith 1969; Voss 1972).

In recent years, there has been growing evidence of
another concentration of coastal plain species about
the Great Lakes, around Georgian Bay in Ontario.
Some of the more notable examples include Rhexia
virginica (Meadow-beauty) (Soper 1956), Potamo-
geton bicupulatus (pond weed) (Reznicek and
Bobbette 1976), Bartonia paniculata (Screw-stem)
(Reznicek and Whiting 1976), and Panicum spretum
(panic-grass) (Catling et al. 1977). Reznicek and Whit-
ing (1976) note that other eastern species such as
Listera australis (Southern Twayblade), Juncus mil-
itaris (Bayonet Rush), Gratiola aurea (Golden-pert),
and Linum striatum (Flax) have similar disjunct dis-
tributions. R. C. Simpson and H. Simpson (1973. The
biology of Balckstone Harbour-Moon Island Provin-
cial Park Reserve, Parry Sound District, Ontario.
Resource Inventory Report, Ontario Ministry of
Natural Resources, Toronto, 153 pp.) found many
shoreline species with eastern affinities including R.
virginica, P. bicupulatus, and Nymphoides cordata
(Floating-heart) on a portion of the Georgian Bay
shoreline in Parry Sound District.

Field work on shoreline vegetation in the Muskoka

241

242 THE CANADIAN FIELD-NATURALIST

and Parry Sound areas has revealed another locality
for some of these species. This locality, Axe Lake, 1s
further north than previous reports, and is also inland
from Georgian Bay. The purpose of this paper is to
describe and discuss the vegetation at this locality with
reference to the substrate type of the shoreline and the
geological history of the area.

Study Area — Axe Lake

Axe Lake occurs on the boundary between Mus-
koka and Parry Sound districts, about 25 km NW of
Huntsville, Ontario (Figure 1). This region has a
growing season of between 180 and 190 d. The frost-
free period is between 100 and 120 d, with the last frost
occurring around 25 May and the first occurring
around 20 September. There are between 160 and
180d per annum with precipitation, for a total of
about 100 cm (Department of Energy, Mines and
Resources 1974).

Figure 2 shows Axe Lake and the adjacent

Vol. 95

wetlands. The lake is about 0.5 X 3 kmand has asand
bottom with occasional outcrops of Precambrian
granitic rocks. The southern third of the lake is largely
surrounded by floating bog mats, although in places a
sand bottom is exposed during low water periods. The
northern two-thirds has scattered boggy areas, and
shorelines dominated by shrubs. In deeper water on
the lake side of the shrubs, emergent and floating-
leaved aquatics are well developed. Most of the lake is
shallow enough to support floating-leaved aquatics,
producing extensive beds of Nymphaea odorata (Fra-
grant Water-lily), Brasenia schreberi (Water-shield),
and Utricularia purpurea (Purple Bladderwort). Only
one area in the northern portion of the lake is deep
enough to be free of floating-leaved aquatics. Small
streams draining peat bogs enter at both the north and
south end; the lake initially drains eastward into Buck
Lake and then Lake Vernon, eventually emptying
westward into Georgian Bay via the Moon River.
Although no long term data are available on lake

79°
+ 46°

PARRY SOUND

Georgian Bay

45°

Sprucedale @

45°

@
Gravenhurst

79°

FicureE |. Location of Axe Lake relative to existing Georgian Bay shoreline and postglacial Lake Algonquin shoreline.
Extensive sand plains parallel the old shoreline (after Chapman 1975, Figure 6).

1981

FIGURE 2. Axe Lake and immediate vicinity. Note the exten-
sive areas of peat bog, and the location of the study
area.

levels, there is considerable variation. The water level
in July 1979 was about 30 cm lower than it had been
the preceding autumn. The water level fell another
5 cm from 13 to 21 July. A subsequent visit in Sep-
tember 1979 revealed the water level 20 cm above the
July level, approaching that of the previous autumn.
While 20- to 30-cm fluctuations may seem minimal,
the extremely gentle slopes (as little as 1:250) means
that a few centimetres drop may expose more than a
metre-wide strip of lake bottom.

Sampling Techniques

Floristic data were gathered over a period of three
years: September 1978, July 1979, September 1979,
and August 1980. Voucher specimens were taken for
most taxa. The problematic taxa (Potamogeton and
Carex in particular) were confirmed or determined by
P. W. Ball (Herbarium, Erindale College, University
of Toronto) or A. A. Reznicek (Herbarium, Univer-

KEDDY: RELICT LAKESHORE VEGETATION

243

sity of Michigan). These vouchers are on file in the
author’s herbarium with duplicates, where available,
in TRT, CAN, and the National Water Research
Institute Herbarium at the Canada Centre for Inland
Waters, Burlington.

The vegetation data were gathered during 13 to 21
July 1979. A 600-m section of shoreline was surveyed
along the northeastern end of the lake (Figure 2); this
encompassed much of the range of variation seen in
the lake as a whole, from open sand beach to mature
bog shoreline (but excluding floating bog mats and
rock outcrops). Along this perimeter I located 25 tran-
sects chosen by means of random numbers subject to
the criterion that no two transects could be closer
together than 10 m. This ensured that each transect
was relatively independent. The transects were
sampled in the order that the random numbers were
drawn to ensure that variation resulting from changes
in observer expertise, working conditions, and/or
plant maturity would be randomly assigned along the
shoreline.

Each transect contained a belt of vegetation 50 cm
wide and was surveyed into 5-cm-height increments
using an automatic level. The waterline on 13 July was
used as the reference point, and for each transect
increments were marked out ranging from 50 cm
above to 50 cm below this water level, a total of 20
height increments (or quadrats) per transect. Because
the water level fell as the study progressed, daily cor-
rections were made for falling water levels. An
observer started at the top of each transect and
recorded the presence of each species in each height
increment. The final set of data then consisted of lists
of species occurrences in 20 height increments for 25
transects.

These randomly chosen transects were assigned to
three shoreline types: sand (6 transects), transition (12
transects), and peat (7 transects). For each of these
categories frequency of occurrence for each species
was calculated by summing all occurrences over all
transects and dividing by the total number of height
increments examined. For example, 120 height
increments (6 transects X 20 height increments) were
examined on sand. The proportion of these which
contained a given species is a measure of that species’
frequency.

Soils data presented are based on samples collected
toa 10-cm depth at 0.25 mabove the waterline in each
transect. Organic content was estimated by percent-
age loss on ignition at 400°C.

Nomenclature follows Fernald (1950) except for the
use of Triadenum fraseri (Marsh St. John’s Wort),
instead of Hypericum virginicum. Most common
names, where available, generally also follow Fernald
(1950).

244

Results and Discussion
Floristics

Peattie (1922) provided a summary table of coastal
plain species known to occur around the Great Lakes.
One of Fernald’s (1942) main criticisms was that too
many botanists thought of the coastal plain flora in
ecological rather than phytogeographic terms, thus
lumping many species of acid sands and bogs into the
coastal plain category. The following discussion of the
flora of Axe Lake attempts to separate these
components.

Two species from Axe Lake, R. virignica and J.
militaris, are widely disjunct from the Atlantic coastal
plain. Their Ontario distribution is largely restricted
to the Georgian Bay region, although J. militaris
occurs north to Temagami District (Fernald 1950)
and east to the Ottawa River (M. 1. Moore. 1972.
Vascular Plants of the Middle Ottawa Valley and
Northeastern Algonquin Park. Petawawa Forest
Experiment Station Information Report PS-X-34,
Environment Canada. 48 pp.). In eastern Canada R.
virginica reaches southwestern Nova Scotia (Roland
and Smith 1969) and J. militaris reaches Newfound-
land (Fernald 1918).

Other species from Axe Lake are also prominent on
the coastal plain and form a strong component of
vegetation in the vicinity of coastal plain disjuncts,
although there are scattered inland stations elsewhere
in North America. Such species include Eleocharis
olivacea (Olive-brown Spike-rush), WN. cordata,
Rhynchospora capitellata (beak-rush), U. purpurea,
U. resupinata (bladderwort), Woodwardia virginica
(Virginian Chain-Fern), and Xyris caroliniana
(Yellow-eyed Grass) as well as J. militaris and
R. virginica.

Many more widespread and northern species now
occur primarily on glaciated areas off the Atlantic
coastal plain, but appear to have survived on the
coastal plain during the last glaciation. Cladium
mariscoides (Twig-rush), Drosera intermedia
(Narrow-leaved Sundew), Juncus pelocarpus (Brown-
fruited Rush), Myriophyllum tenellum (Water-
milfoil), Muhlenbergia uniflora, Potamogeton con-
fervoides, P. oakesianus, Utricularia cornuta (Horned
Bladderwort), and Viola lanceolata (Lance-leaved
Violet) are frequently conspicuous in sites supporting
coastal plain disjuncts.

Lastly, there are species considered by Peattie
(1922) and/or McLaughlin (1932) as coastal plain
species, but which are much more widespread,
although ecologically they may be a prominent com-
ponent of the flora on lakes such as Axe Lake. Species
such as Calamagrostis canadensis (Blue-joint), Duli-
chium arundinaceum (Three-way Sedge), Eriocaulon
septangulare (Pipewort), Lycopodium inundatum

THE CANADIAN FIELD-NATURALIST

Vol. 95

(Bog Club-moss), Lycopus uniflorus (Water-
horehound), Pontederia cordata (Pickerel weed),
Rhynchospora fusca (beak-rush), Scirpus torreyi(bul-
rush), and Utricularia gibba (Humped Bladderwort)
are probably best classified simply as species of acid
sands and bogs.

The restricted distribution of the following seven
species results in their being classed as “rare” in Onta-
rio (Argus and White 1977): E. olivacea, J. militaris,
P. oakesianus, R. virginica, R. capitellata, S. torreyi,
X. caroliniana.

Miller (1977) used floristic and water chemistry
data to classify 56 lakes of southern Ontario into five
“types,” based on floating and submerged macro-
phytes. Axe Lake was included in his study and classi-
fied as type A, “. . . very soft, low conductivity lakes
that are dominated by Isoetids and Utricularids...”.
These lakes generally have alkalinity values of 0-7 mg
CaCO,'L' and conductivity values from 20-42 uS
[umhos]:cm '. Miller reported that Axe Lake had an
alkalinity of 3, conductivity of 29, and a pH value of 6.
Miller did not provide the raw data matrix used in his
floristic analysis, but based on his tables of “character-
istic species,” type A lakes are characterized by species
such as Lobelia dortmanna (Water-lobelia). E. sep-
tangulare, M. tenellum, U. purpurea, U. resup-
inata, J. pelocarpus, J. militaris, and Elatine minima
(Waterwort). Only the latter species has not been
observed in Axe Lake. Crowder et al. (1977) studied
16 lakes in southeastern Ontario and similarly con-
cluded that lakes with conductivities of below
60 us:cm ' contained species not found in other lakes,
notably E. septangulare, L. dortmanna, and M. tenel-
lum. Swindale and Curtis (1957) also found that spe-
cies such as M. tenellum, E. septangulare, J. pelocar-
pus, and E. minima occurred together in low
conductivity lakes in Wisconsin.

Vegetation

The most botanically interesting area of the lake is
the extreme north end (Figure 2). The substrate ranges
from the gently sloping sand shown in Figure 3
through to peat. This small area at the north end
appears to encompass much of the variation in the
shoreline types of Axe Lake.

Figure 4 shows shoreline profiles from the north
end of Axe Lake in three different substrate types: (1)
sand, (2) transition, and (3) peat. These categories are
based on the organic content of the transects at 0.25 m
above the waterline, as determined by weight loss on »
ignition to 400°C (mean values for each category are:
sand = 0.9%, transition = 3.7%, peat = 16.4%). These
changes in organic content are related to the physiog-
nomy of the shoreline (Figure 4) and species composi-
tion (Table 1). These categories could be distinguished
in the field. “Sand,” as shown in Figure 3, referred to

1981

KEDDY: RELICT LAKESHORE VEGETATION 245

FIGURE 3. The shoreline vegetation at the north end of Axe Lake in a sand shoreline type. Note the gently sloping sand
exposed by falling summer water levels. The vegetation types from left to right (above to below waterline) are (1) Betula
papyrifera (White Birch) and Pinus strobus (White Pine) forest, (2) a band of shrubs, principally Myrica gale, (3) open
sand with Rhynchospora fusca, Lobelia dortmanna, Eriocaulon septangulare, Drosera intermedia, and Utricularia
cornuta (at peak of flower — see lower left of picture), (4) emergents such as Scirpus torreyi mixed with E.
septangulare and L. dortmanna, and (5) open water supporting Scirpus subterminalis, Nympoides cordata, E.
septangulare, and L. dortmanna.

shorelines with extensive areas of open sand and only
sparse plant cover. “Transition” shores were densely
vegetated, but scattered openings in the vegetation
revealed a sandy substrate. “Peat” shores were densely
vegetated and had a peat layer covering the sand.
Sand was the least common of the three shoreline
types, and occurred only on the north and east sides of
the lake. (The latter has been disturbed by recent
campsites, and possibly by a now-abandoned farm, so
it was excluded from the study.) Peat shores were the
most common of all. Transition areas intermediate in
organic content were also common. These three cate-
gories are arbitrary divisions in a continuum, so a
more accurate assessment of relative abundances of
these shoreline types was not attempted. These cate-
gories are based only on above-water characteristics.

The below-water organic content was consistently low
(mean = 0.78%), yet, as Table 1 shows, the aquatic
flora differed among the three shoreline types.

Species suchas N. cordata, E. septangulare, and M.
tenellum were abundant on areas of sand. All of the
species in column | (except for Juncus filiformis)
(rush) also occurred on more organic shorelines. In
general, sand shorelines supported few species.

The transition shorelines supported many more
species. Rhexia virginica, X. caroliniana, M. uniflora,
and V. lanceolata attained their peak abundance on
these shores. The dominant species included Cladium
mariscoides, E. septangulare, and N. cordata.

The peat shorelines were dominated by 7. fraseri,
Myrica gale (Sweet Gale), C. mariscoides, Scirpus
subterminalis (Swaying Rush), E. septangulare, and

246

T-13, TRANSITION

RELATIVE HEIGHT (m)

T-25, PEAT

= [he StS { -
0 10 20 30 40 50 60
DISTANCE (m)

FiGureE 4. Diagrammatic profiles through transects repre-
senting the three main shoreline types studied. Note
the increasing plant cover from sand through transi-
tion to peat shorelines.

N. cordata. Many floating-leaved aquatics, suchas B.
schreberi, N. ordorata. Nuphar variegatum (Yellow
Pond-lily), and Potamogeton oakesianus, reached
their peak abundance here. Myriophyllum tenellum,
L. dortmanna, U. cornuta, and J. pelocarpus all
decreased in frequency on peat shorelines.
Eleocharis olivacea did not occur in the study tran-
sects; this species and _X. caroliniana reach their peak
abundance on floating peat islands in the south end of
the lake. Woodwardia virginica and Rhynchospora
capitellata were also absent from the study area; each
occurred at a single point elsewhere on the lakeshore.

Geological History

This area emerged from beneath the Wisconsin ice
sheet roughly 12 000 years ago (Douglas 1970, fig.
XII-15). Chapman (1975) stated that the absence of
any strong moraines suggested an uneventful glacial
withdrawal from the area. The glacier left behind
sandy and stony till which is so thin that it generally
fails to cover completely the underlying rock ridges.
These ridges are composed of Precambrain granitic
rocks (Hewitt 1967). A most interesting feature, how-
ever, is the postglacial Lake Algonquin shoreline
which runs due north from Bracebridge to Lake Nip-
issing. This shoreline is now marked by extensive sand

THE CANADIAN FIELD-NATURALIST

Vol. 95

plains (Chapman 1975, map 2228). When Lake
Algonquin was present, this shoreline possessed an
intricate series of islands and bays (Figure 1). Axe
Lake is just “off shore” from this maximum water
level of Lake Algonquin. Lake Algonquin water levels
then gradually dropped (because of new outlets such
as the Fossmill outlet, as well as isostatic rebound)
leaving behind scattered lakes with rich shoreline flo-
ras. At its peak the shoreline of Lake Algonquin may
have had bays and islands with gently sloping sandy
shores not unlike those on Axe Lake today.

Peattie (1922) attempted to trace the inland migra-
tion of coastal plain species in relation to the postgla-
cial development of the Great Lakes. He concluded
that the coastal plain flora entered the Great Lakes
during the Lake Algonquin stage, when Lake Algon-
quin was drained by the Kirkfield outlet into Lake
Iroquois, which in turn drained down the Hudson
River to the Atlantic coastal plain. Speaking of this
time, he wrote “Judging from the large number of old
spits and beach ridges which have been traced out for
the glacial lakes giving rise to Lake Erie and Lake
Michigan, it is reasonable to imagine their shores to
have been an intricate series of lagoons, ridges,
strands and low dunes harbouring the newly-migrated
coastal plain types.” This appears to be an excellent
description of the shoreline of Lake Algonquin from
Bracebridge to Lake Nipissing.

McLaughlin (1932) noted that the greatest number
of coastal plain plants grow on moist, sandy lake-
shores, and that the development of Sphagnum bogs
was partially responsible for the gradual elimination
of these species. Figure 2 showed the extensive areas
of peat bog around Axe Lake. These areas were pre-
sumably open water when Lake Algonquin was at its
maximum; they have since become peat bogs, and
judging from the extensive areas of boggy shores and
floating peat islands in Axe Lake, this process is con-
tinuing. Many of the coastal plain species listed in
Table | were absent from those shores of the lake with
floating bog mats. Ona smaller scale, comparison of
column 2 (transition) with column 3 (peat) also
appears to support McLaughlin’s observation, as spe-
cies such as M. uniflora and R. virginica are absent
from peat shorelines. Aquatic coastal plain species,
however, such as P. confervoides, P. oakesianus, U.
resupinata, U. purpurea, and J. militaris reached their
peak abundance in shallow water adjacent to the peat
shores.

Recent History

Recent human activities, particularly cottage
development, may be modifying many lakes in this
area. To examine this problem, Miller and Dale
(1979) studied changes in the aquatic macrophyte
floras of eight small lakes in the Muskoka area. They

1981 KEDDY: RELICT LAKESHORE VEGETATION 247

TABLE 1—Frequency of shoreline plants on three shoreline types in Axe Lake. Columns give percentage occurrence in
“quadrats” located between 0.5 m above and 0.5 below the waterline. Species are arranged by (1) shoreline type of peak
abundance (sand, transition, peat), (2) habitat specificity (number of shoreline types occupied), and (3) abundance. Shoreline

types are shown in profile in Figure 4

Occurrence along shoreline

(%)

Sand Transition Peat

*Rare in Ontario (Argus and White 1977).
{Species with coastal plain affinities.

compared a 1953 survey of these lakes to their 1976
and 1977-surveys. Axe Lake, one of the lakes they
studied, did not appear to have lost any species of
aquatic macrophytes during this time period. Miller
and Dale (1979) record nine additional species in Axe
Lake; these were likely missed in the 1953 survey. In
addition to their total of 19 species I collected Pota-
mogeton natans (Floating Brown leaf), P. confer-
voides, Glyceria borealis (Small Floating Manna-
grass), J. militaris, J. pelocarpus, M. tenellum, and
Zizania aquatica(Wild Rice). Because the emphasis of

Occurrence along shoreline

(%)

Sand Transition Peat

Species (n= 120) (n= 240) (n= 140) — Species (n= 120) (n = 240) (7 = 140)
Sand Transition (continued)
Juncus filiformis 0.8 — — Hypericum ellipticum 3.3 7.5 4.3
Viburnum cassinoides 6.7 0.8 — Juncus canadensis 1.7 2.9 0.7
Bidens sp. 1.7 0.4 — Peat
Nymphoides cordata* 70.8 56.7 43.6 Pollarnoneiow anlanebns a ae 29
Eriocaulon septangulare 65.0 Sil 43.6 4 2
i Tris versicolor is os 1.4
Myriophyllum tenellum* 41.7 26.3 12.1
Chamaedaphne calyculata — — 0.7
Lobelia dortrganna 39.2 23.7 11.4 Vaceini
: : accinium Oxycoccus — — 0.7
Utricularia cornuta* 24.2 14.6 1.4 Sana Gey
: Dulichium arundinaceum _ 14.2 21
Hypericum boreale 10.0 9.2 2.9 5 ;
Eleocharis palustris — 3.3 21.4
Alnus rugosa 10.0 2.5 2.9 Se :
Spi Ib 33 13 14 Calamagrostis canadensis — 5.8 20.7
é LIS alse ; Brasenia schreberi — 10.4 20.0
Transition Nymphaea odorata — eS) 17.1
Rhexia virginica* + — 14.2 — Rubus hispidus = 7.9 15.7
Potamogeton natans — 2.1 _— Vaccinium macrocarpon — 9.2 15.0
Lycopodium inundatum — 1.7 — Nuphar variegatum — 2.1 14.3
Ilex verticillata _— 0.4 — Pontederia cordata = 333 13.6
Carex michauxiana a 0.4 — Carex rostrata = 4.2 12.1
Pyrus arbutifolia — 0.4 — Potamogeton oakesianus* | — 1.7 10.0
Agrostis hyemalis _ 0.4 _ Glyceria canadensis _ 1.3 10.0
Panicum lanuginosum — 0.4 — Carex vesicaria — 1.7 Dal/
Carex echinata — 0.4 _— Carex stricta — 1.7 4.3
Xyris caroliniana* ¢ 14.2 _ Utricularia intermedia — 353 3.6
Juncus brevicaudatus 1.7 4.2 — Potamogeton confervoides* — 0.4 2.9
Nemopanthus mucronata 3.3 3.3 — Carex oligosperma — 0.8 1.4
Muhlenbergia uniflora 17.5 2.1 Scirpus subterminalis 15.0 40.4 50.7
Utricularia gibba _ ES) 7.1 Triadenum fraseri 9.2 20.8 S07
Carex lasiocarpa — 1.3 0.7 Mpyrica gale 17.5 13.8 33.6
Drosera intermedia* 16.7 39.2 14.3 Lysimachia terrestris 20.8 20.8 31.4
Cladium mariscoides* 3.3 39.2 30.0 Utricularia vulgaris 3.3 13.8 17.9
Juncus pelocarpus* 22.5 27.5 2.9 Utricularia resupinata* 6.7 15.4 15.7
Viola lanceolata* 4.2 25.8 3.6 Lycopus uniflorus 10.8 6.7 13.6
Aster nemoralis 6.7 19.2 5.0 Utricularia purpurea* 1.7 8.8 12.1
Scirpus torreyit 1.7 19.2 12.1 Juncus militaris* + 1.7 3.7 5.0
Rhynchospora fusca 10.8 17.5 5.0

this study was on shoreline and shallow water plants,
additional species may yet be found. Local residents
have remarked that the lake has become more weedy,
but there is no convincing evidence of recent changes
in the flora of Axe Lake.

In conclusion, Axe Lake appears to be a relict
fragment of a once extensive sandy shoreline stretch-
ing from Bracebridge to Lake Nipissing. The flora of
Axe Lake contains some coastal plain species which
apparently migrated into the Great Lakes area near
the end of the last ice age. The vegetation of this lake

248

may be a relict of the shoreline vegetation that flour-
ished in the sandy bays of postglacial Lake Algon-
quin. Because the coastal plain flora of the Georgian
Bay area has not yet been adequately recognized in
planning parks and other reserves, it is to be hoped
that thought will be given to preserving some suitable
shores in the near future.

Acknowledgments

The assistance with summer field work by S. M.
van Walsum and K. McCulloch was much appre-
ciated. P. W. Ball (Erindale College, University of
Toronto) and A. A. Reznicek (Herbarium, University
of Michigan) kindly assisted in the annotation and
determination of voucher specimens. Thanks also to
R. Proctor for assistance with the data manipulation,
and to D. F. Brunton, C. J. Keddy, G. La Roi, and
A. A. Reznicek for constructive criticism of various
drafts of this paper. G. Burger kindly permitted us to
work on his land and use his cabin for 1980 field work.
This research was supported by University of Guelph
R.A.B. grant No. 83193 and NSERC grant A6963.

Literature cited

Argus, G. W., and D. J. White. 1977. The rare vascular
plants of Ontario. Syllogeus Series, Number 14, National
Museums of Canada, Ottawa. 63 pp.

Catling, P. M., A. A. Reznicek, and J. L. Riley. 1977. Some
new and interesting grass records from southern Ontario.
Canadian Field-Naturalist 91: 350-359.

Chapman, L. J. 1975. The physiography of the Georgian
Bay — Ottawa valley area of southern Ontario. Ontario
Division of Mines, Geoscience Report 128. 33 pp.

Crowder, A. A.,J. M. Bristow, M. R. King, andS. Vanderk-
loet. 1977. The aquatic macrophytes of some lakes in
southeastern Ontario. Le Naturaliste Canadien 104:
457-464.

Department of Energy, Mines and Resources. 1974. The
national atlas of Canada. Queen’s Printer, Ottawa. 254 pp.

Douglas, R. J. W. (Editor). 1970. Geology and economic
minerals of Canada. Department of Energy, Mines and
Resources. 838 pp.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Fernald, M. L. 1918. The contrast in the floras of eastern
and western Newfoundland. American Journal of Botany
5: 237-247.

Fernald, M. L. 1942. Misinterpretation of Atlantic coastal
plain species. Rhodora 44: 238-246.

Fernald, M. L. 1950. Gray’s manual of botany. 8th ed. D.
Van Nostrand, NY. 1623 pp.

Hewitt, D. F. 1967. Geology and mineral deposits of the
Parry Sound - Huntsville area. Ontario Department of
Mines, Geological Report 52. 65 pp.

McLaughlin, W. T. 1932. Atlantic coastal plain plants in
the sand barrens of Wisconsin. Ecological Monographs 2:
335-383.

Miller, G. E. 1977. A classification of Ontario lakes based
on their submersed and floating macrophyte flora. M.Sc.
thesis, University of Guelph, Guelph, Ontario. 95 pp.

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: 386-390.

Peattie, D. C. 1922. The Atlantic coastal plain element in
the flora of the Great Lakes. Rhodora 24: 57-70, 80-88.
Reznicek, A. A., and R. S. W. Bobbette. 1976. The taxon-
omy of Potamogeton subsection hybridi in North Amer-

ica. Rhodora 78: 650-673.

Reznicek, A. A., and R. E. Whiting. 1976. Bartonia (Gen-
tianaceae) in Ontario. Canadian Field-Naturalist 90:
67-69.

Roland, A. E., and E. C. Smith. 1969. The flora of Nova
Scotia. NS Museum, Halifax. 705 pp.

Soper, J. H. 1956. Some families of restricted range in the
Carolinian flora of Canada. Transactions of the Royal
Canadian Institute 31, Part 2: 69-90.

Swindale, D. N., and J. T. Curtis. 1957. Phytosociology of
the larger submersed plants in Wisconsin lakes. Ecology
38: 397-407.

Voss, E. G. 1972. Michigan flora. A guide to the identifica-
tion and occurrence of the native and naturalized seed-
plants of the state, Part 1. Gymnosperms and Monocots.
Cranbrook Institute Science Bulletin 55, Cranbrook Insti-
tute of Science, Bloomfield Hills, and University of Michi-
gan Herbarium. 488 pp.

Received 3 June 1980
Accepted 12 December 1980

Low DDT Residues in Piasma of Bald Eagles
(Haliaeetus leucocephalus) Wintering in Colorado and Missouri

CHARLES J. HENNY,! CURTICE R. GRIFFIN,? DALE W. STAHLECKER,?

ALAN R. HARMATA,‘4 and EUGENE CROMARTIE5

1US Fish and Wildlife Service, 480 SW Airport Road, Corvallis, Oregon 97330

2School of Forestry, Fisheries, and Wildlife, 112 Stephens Hall, University of Missouri, Columbia, Missouri 65211
3US Forest Service, Box 98, Jemez Springs, New Mexico 87025

4Department of Biology, Montana State University, Bozeman, Montana 59715

5US Fish and Wildlife Service, Patuxent Wildlife Research Center, Laurel, Maryland 20811

Henny, Charles J., Curtice R. Griffin, Dale W. Stahlecker, Alan R. Harmata, and Eugene Cromartie. 1981. Low DDT
residues in plasma of Bald Eagles (Haliaeetus leucocephalus) wintering in Colorado and Missouri. Canadian

Field-Naturalist 95(3): 249-252.

Residues of DDT and its metabolites (2 DDT) in blood plasma from Bald Eagles (Haliaeetus leucocephalus) wintering in
Colorado and Missouri were low (0.06-0.14 ug/g) in 1977 and 1978. Most of the adult wintering birds probably nest in
Canada, although a few in Missouri may be from Minnesota or the Great Lakes region. DDE residues in eggs, estimated from
those in plasma, were lower than those reported from Saskatchewan in 1969-1972 (4.5 ug/g vs. 0.67 ug/g); this suggests

reduced environmental contamination.

Key Words: Bald Eagle, Haliaeerus leucocephalus; environmental contaminants, DDT, PCB, wintering populations.

Reports of low rates of reproduction (Broley 1958;
Abbott 1967) and relatively high levels of toxic chemi-
cal contamination (Krantzet al. 1970; Wiemeyeret al.
1972) in certain Bald Eagle (Haliaeetus leucocepha-
lus) populations led to widespread concern for this
species. It is classified as “endangered” in the contig-
uous states excepting Washington, Oregon, Minne-
sota, Wisconsin, and Michigan, where it is “threat-
ened” (US Fish and Wildlife Service 1979). Sprunt
et al. (1973) documented decreased reproduction in
Florida, the northeastern states, and near the shores
of the Great Lakes. Grier (1974) noted that many
nesting birds are widely spaced in relatively inaccessi-
ble regions and thus are difficult to sample. To gain
additional insight into the pollutant contamination of
Bald Eagles, we collected blood samples for organo-
chlorine analyses from two wintering populations, the
San Luis Valley in Colorado and Swan Lake in Mis-
souri, during the winters of 1976-1977 and 1977-1978.

Henny and Meeker (1981) showed that plasma
residues could be used to predict residues in eggs for
four species of raptors and that the relationship was
species independent for the species tested. Further-
more, blood plasma residues in American Kestrels
(Falco sparverius) paralleled residues in eggs during a
recent study that evaluated the impact of an opera-
tional DDT spray project (Henny 1977a). The ability
to predict residues in eggs provides a method for
interpreting the plasma residues. Contaminants in
plasma of adults and estimated contaminants in eggs

from these two wintering segments of the Bald Eagle
population can be evaluated because some informa-
tion concerning residues in Bald Eagle eggs is
available.

Methods

During the study 14 adults (white head) and 21
subadults were trapped and blood samples collected.
Twenty were trapped in the San Luis Valley of Colo-
rado between 22 January and 7 March 1977, and I5at
Swan Lake National Wildlife Refuge (NWR), Mis-
souril, between 9 January and 16 February 1978.
About 2-3 ml of blood was collected from the brachial
vein with a 3-ml disposable syringe and a 23-gauge
needle, then placed a heparinized tube, and kept cool
until later in the day when it was centrifuged. The
plasma was drawn off and stored frozen until it was
analyzed at the Patuxent Wildlife Research Center.

The samples were mixed with anhydrous sodium
sulfate, extracted 7 h with hexane in a Soxhlet appa-
ratus, and lipids removed by Florisil column chroma-
tography as described by Cromartie et al. (1975),
except that the organochlorines were separated into
four fractions rather than three to ensure the isolation
of dieldrin or endrin, or both (Kaiser et al. 1980). The
pesticides in each fraction were quantified with a gas—
liquid chromatograph (GLC) using an electron-
capture detector and a 1.5% OV-17/1.95% QF-1
column. Average recoveries ranged from 83 to 104%
from spiked tissue and 77 to 97% from spiked Mallard

249

250

THE CANADIAN FIELD-NATURALIST

TABLE |—Pesticide residues in plasma from Bald Eagles wintering in Colorado and Missouri

Mean residues (ug/g)

Location

Year
Missouri
4 adults 1978 (Jan.—Feb.)
11 subadults 1978 (Jan.—Feb.)
Colorado}
10 adults 1977 (Jan.-March)

10 subadults 1977 (Jan.-March)

1Almost all DDE.
2Range; ND = none detected.

Vol. 95
XDDT! Dieldrin PCBs
0.06 0.03 0.24
(0.03-0.10)? (ND-0.08) (ND-0.36)
0.06 0.01 0.08
(0.01—0.14) (ND-0.06) (ND-0.27)
0.14 0.01 0.10
(0.03-0.23) (ND-0.05) (ND-0.68)
0.07 ND 0.04
(ND-0.31) (ND) (ND-0.36)

3Three contained heptachlor epoxide (0.11, 0.01, and 0.02), two contained cis-chlordane (0.02 and 0.01), and one trans-

nonachlor (0.01).

(Anas platyrhynchos) blood. Residues were not cor-
rected for percentage of recovery. The lower limit of
reportable residues for organochlorine pesticides in
the blood plasma ranged from 0.01 to 0.05 ug/g
depending upon the sample size. Residues in 5% of the
blood samples were confirmed with a Finnigan model
4000 gas chromatograph/mass spectrometer
(GC/MS) (Kaiser et al. 1980).

Results and Discussion

The XDDT in plasma (mean 0.12 wg/g) of adult
Bald Eagles was low (Table 1). Adult American Kes-
trels averaged nearly the same (X{DDT 0.11 ug/g)
before a spray project in the Pacific Northwest, while
1 year after a 3/4 lb per acre (0.84 kg/ha) treatment,
they showed plasma levels of 0.78 ug/g in the spray
area. Cooper’s Hawks (A ccipiter cooperii) and Sharp-
shinned Hawks (A. striatus) in the same spray area
averaged over 2 ug/g after spraying with some indi-
viduals showing 7-9 ug/g (Henny 1977a). The Bald
Eagles in this study showed slightly higher 2: DDT
plasma residues than found in 16 adult Golden Eagles
(Aquila chrysaetos) (X{DDT 0.02 ug/g) sampled near
Dillon, Montana, in March, April, and May of 1975

(Henny, unpublished data). Adult Bald Eagles gener-
ally showed higher residues than the younger birds
with the Missouri and Colorado populations carrying
similar pollutant burdens.

The plasma residues (X{DDT) in the Bald Eagle
samples from this study were among the lowest
encountered among adult raptors studied to date. We
also converted the plasma residues to estimated egg
residues based on the postlaying model of Henny and
Meeker (1981). Estimated XDDT residues in eggs
averaged 0.68 wg/g (Table 2). If the prelaying model
were used, the residue estimates would be slightly
lower. Polychlorinated biphenyls (PCBs) were
detected in five of the plasma samples; however, the
detection limit was higher than for XDDT. Therefore,
PCBs were probably also present at lower levels in
other eagles.

One question immediately arises: Where are the
nesting areas of the adult birds that winter in Colo-
rado and Missouri? Whitfield et al. (1974) stated that
Saskatchewan eagles must make up a large propor-
tion of the population wintering in the midwestern
United States. Recently, Gerrard et al. (1978) and
Leighton et al. (1979) estimated the population in

TABLE 2— Pesticide residues in Bald Eagle eggs (ug/g) as estimated from plasma of adults

Estimated mean residues (range)

No.
Location samples Xx DDT Dieldrin PCBs!
Swan Lake,
Missouri 4 0.36 (0.14-0.58) 0.15 (ND-0.43) 1.4 (ND-2.2)
San Luis Valley,
Colorado? 10 0.81 (0.18-1.4) 0.05 (ND-0.26) 0.67 (ND-4.8)

1Minimal estimate.

2Two also contained small amounts of heptachlor epoxide and cis-chlordane, and one trans-nonachlor. ND = none detected.

HENNY ET AL.: DDT IN BALD EAGLE PLASMA >|

(1974)
NA _ Grier (1974)

87

Gilbertson and Reynolds

Krantz et al. (1970)

Wiemeyer et al. (1972)
Wiemeyer et al. (1972)
Wiemeyer et al. (1972)
Wiemeyer et al. (1972)

Grier (1974)

0.895 This study

NA

28
Tall
De

1.1

PCB Authority
2.8

Heptachlor
epoxide
NA
NA
NA
0.02
0.07
0.08
0.02

Dieldrin
NA
0.42
1.3
1.0
0.99
0.10
0.06

DDT
NA
NA

0.16

NA
NA
0.83
0.32
1.1
1.0
0.01

DDD

4.5
14
21
4.7
9.6
2.9
0.67

DDE

9
15(10)
5(4)
7(7)
5(5)

14

No. eggs
4

(No. nests)

Year(s)
1969-1972
1967
1968-1972
1968
1969
1969-1970
1969
1970
1977-1978

5

NA = not available, ND = none detected; T = trace.
‘Geometric mean, dry weight basis, converted to fresh wet weight (divided by 5.0).

2Arithmetic mean, dry weight basis, converted to fresh wet weight (divided by 5.0).

TABLE 3—T oxic chemical residues (ug/g, fresh wet weight) in Bald Eagle eggs from Canada, Alaska, and the Great Lakes region, as compared to estimates from
3Arithmetic means computed on nest basis.

this study
‘Lake Superior site (high residues typical of Great Lakes) but not representative of Michigan eagle population at large, most of which is inland (S. Postupalsky,

personal communication).

Alaska, Admiralty Island}
S5Minimal estimate.

Nesting location
Saskatchewan!
NW Ontario?
NW Ontario
Wisconsin}
Michigané
Minnesota}
Alaska, Kodiak}
This study

NOTE:

northern Saskatchewan at over 14000 birds and
showed band recoveries which bracketed the Colo-
rado and Missouri wintering areas. Griffin et al.
(1980) reported that Bald Eagles that wintered at
Swan Lake NWR in Missouri originate in Ohio,
Michigan, Wisconsin, Minnesota, Ontario, and Sas-
katchewan, and some migrate through Manitoba and
Ontario. Dunstan (1973) reported two Minnesota-
banded birds recovered in Texas, that probably had
migrated through Missouri. Postupalsky (1976)
reported recoveries from Michigan-banded eagles
generally to the east of Missouri except for one in
Arkansas.

S. Postupalsky et al. (Department of Wildlife Ecol-
ogy, University of Wisconsin, Madison, Wisconsin,
unpublished data) pointed out that adults that breed
in Minnesota and western Ontario and points north
and northwest apparently are migratory, but few if
any that nest south of Lake Superior leave their
summer range during the winter. Thus, a subadult
found in the winter at Swan Lake could have hatched
anywhere between Saskatchewan and northern Ohio.
An adult, on the other hand, would most likely be
from Canada or possibly Minnesota.

Chemical residue levels in a few Bald Eagle eggs
from Alaska, Canada, and the Great Lakes region
have been published (Table 3). Residues in Saskat-
chewan and Alaska in 1969-1972 were among the
lowest reported. Most Bald Eagle eggs analyzed were
addled and therefore may not be representative (i.e.
eggs with higher residues may not hatch, but eggs with
extremely high residues may break). Postupalsky
(1978) recently reported improved eagle productivity
in the Great Lakes region. Bald Eagles appear to be
following the same pattern of improvement in repro-
duction with the decline of DDE residues as the
Osprey (Pandion haliaetus) (Henny 1977b; Spitzer et
al. 1978).

The improved pattern of productivity in several
Bald Eagle populations in the mid-to-late 1970s sug-
gest that our estimated egg residues in 1977-1978
(0.67 ng/g DDE) are realistic and that a residue
decline did occur in the Canadian breeding popula-
tion (cf. Saskatchewan, 4.5 ug/g DDE in 1969-1972).
Leighton et al. (1979) reported a reproductive rate of
0.95 young per breeding area in Saskatchewan in 1974
and indicated the production was well above the best
estimate of the minimum required to maintaina stable
population (0.7 young per occupied breeding area)
determined in studies of six other Bald Eagle popula-
tions (Sprunt et al. 1973). It should be noted that the
plasma-—egg relationship for }DDT in Bald Eagles has
not been studied specifically. We are assuming that
the relationship is independent of species; therefore, it
would be useful to collect a few eggs from Bald Eagles

Lys

nesting in northern Canada to determine if egg
residues agree with our predictions.

Acknowledgments

The Colorado portion of the study was partially
supported by the National Wildlife Federation and
the National Audubon Society. The Missouri portion
of the study is a contribution from the Missouri
Cooperative Wildlife Research Unit; it was funded in
part by the Unit, by a National Audubon Society
fellowship, and by the US Fish and Wildlife Service
through Agreement No. USDI 14-16-008-757,
awarded to the University of Missouri. The manu-
script was improved by the comments of Sergej Pos-
tupalsky, Thomas S. Baskett, and Lucille F. Stickel.

Literature Cited

Abbott, J. M. 1967. The Chesapeake Bald Eagles. Atlantic
Naturalist 22: 20-25.

Broley, C. L. 1958. The plight of the American Bald Eagle.
Audubon Magazine 60: 162-163.

Cromartie, E., W. L. Reichel, L. N. Locke, A. A. Belisle,
T. E. Kaiser, T.G. Lamont, B.M. Mulhern, R. M.
Prouty, and D. M. Swineford. 1975. Residues of orga-
nochlorine pesticides and polychlorinated biphenyls and
autopsy data for Bald Eagles, 1971-72. Pesticide Monitor-
ing Journal 9: 11-14.

Dunstan, T. C. 1973. Bald Eagle from Minnesota recov-
ered in Texas. Loon 45: 132.

Gerrard, J. M., D. W. A. Whitfield, P. Gerrard, P. N. Ger-
rard, and W. J.Maher. 1978. Migratory movements and
plumage of subadult Saskatchewan Bald Eagles. Cana-
dian Field-Naturalist 92: 375-382.

Gilbertson, M., and L. Reynolds. 1974. DDE and PCB
determinations in Canadian birds, 1969 to 1972. Canadian
Wildlife Service, Occasional Paper No. 19. 18 pp.

Grier, J. W. 1974. Reproduction, organochlorines, and
mercury in northwestern Ontario Bald Eagles. Canadian
Field-Naturalist 88: 469-475.

Griffin, C. R., J. M. Southern, and L. D. Frenzel. 1980.
Origins and migratory movements of Bald Eagles winter-
ing in Missouri. Journal Field Ornithology 51: 161-167.

Henny, C.J. 1977a. Birds of prey, DDT, and tussock
moths in Pacific Northwest. Transactions North Ameri-
can Wildlife and Natural Resources Conference 42:
397-411.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Henny, C. J. 1977b. Research, management, and status of
the Osprey in North America. /n Proceedings World Con-
ference on Birds of Prey. Edited by R. D. Chancellor.
International Council for Bird Preservation. pp. 199-222.

Henny, C. J., and D. L. Meeker. 1981. An evaluation of
blood plasma for monitoring DDE in birds of prey. Envi-
ronmental Pollution. /n press.

Kaiser, T. E., W. L. Reichel, L. N. Locke, E. Cromartie,
A. J. Krynitsky, T. G. Lamont, B. M. Mulhern, R. M.
Prouty, C. J. Stafford, and D. M. Swineford. 1980. Or-
ganochlorine pesticide, PCB, and PBB residues and
necropsy data for Bald Eagles from 29 states — 1975-77.
Presticide Monitoring Journal 13: 145-149.

Krantz, W. C., B. M. Mulhern, G. E. Bagley, A. Sprunt, IV,
F. J. Ligas, and W. B. Robertson, Jr. 1970. Organochlo-
rine and heavy metal residues in Bald Eagle eggs. Pesticide
Monitoring Journal 4: 136-140.

Leighton, F. A., J. M. Gerrard, P. Gerrard, D. W. A. Whit-
field, and W. J. Maher. 1979. An aerial census of Bald
Eagles in Saskatchewan. Journal Wildlife Management
43: 61-69.

Postupalsky, S. 1976. Banded northern Bald Eagles in
Florida and other southern states. Auk 93: 835-836.

Postupalsky,S. 1978. The Bald Eagles return. Natural His-
tory 87: 62-63.

Spitzer, P. R., R. W. Risebrough, W. Walker, II, R. Her-
nandez, A. Poole, D. Puleston, and I.C.T. Nisbet.
1978. Productivity of Ospreys in Connecticut-Long
Island increases as DDE residues decline. Science 202:
333-335.

Sprunt, A., IV, 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. Transac-
tions North American Wildlife and Natural Resources
Conference 38: 96-105.

US Fish and Wildlife Service. 1979. List of endangered and
threatened wildlife and plants. Federal Register 44: 3644.

Whitfield, D. W. A., J. M. Gerrard, W. J. Maher, and
D. W. Davis. 1974. Bald Eagle nesting habitat, density,
and reproduction in central Saskatchewan and Manitoba.
Canadian Field-Naturalist 88: 399-407.

Wiemeyer, S. N., B. M. Mulhern, F. J. Ligas, R. J. Hensel,
J.E. Mathisen, F.C. Robards, and S. Postupals-
ky. 1972. Residues of organochlorine presticides, poly-
chlorinated biphenyls, and mercury in Bald Eagle eggs and
changes in shell thickness — 1969 and 1970. Pesticide
Monitoring Journal 6: 50-55.

Received 10 November 1980
Accepted 14 January 1981

Changes in Small Mammal Populations Following Clear-cutting
in Upper Michigan Conifer Swamps

Louis J. VERME and JOHN J. OZOGA

Michigan Department of Natural Resources, Shingleton, Michigan 49884

Verme, Louis J.,andJohnJ.Ozoga. 1981. Changes in small mammal populations following clear-cutting in upper Michigan
conifer swamps. Canadian Field-Naturalist 95(3): 253-256.

Strip clear-cutting in an upper Michigan conifer swamp increased the relative abundance of small mammals, particularly
Deer Mice (Peromyscus maniculatus), Least Chipmunks (Eutamias minimus), and Meadow Voles (Microtus pennsylvani-
cus). Compared to a clear-cut block with slash left unburned, broadcast burning of slash resulted in a greater population of
shrews (mainly Masked Shrews, Sorex cinereus) and Deer Mice. Nevertheless, because Northern White Cedar (Thuja
occidentalis) reproduced adequately, we conclude that small rodents have little impact upon conifer swamp regeneration in

this locale.

Key Words: small mammals, Northern White Cedar, Thuja occidentalis, clear-cutting, stand regeneration.

Conifer swamps dominated by Northern White
Cedar (Thuja occidentalis); occupy over a million
acres in Michigan’s upper peninsula, or nearly half of
this forest type’s acreage within the northern Great
Lakes region (Johnston 1977). These stands are vital
to White-tailed Deer (Odocoileus virginianus)
because they congregate, or “yard,” here in winter to
gain respite from harsh weather. Efforts to restore
conifer swamp deeryards to their former high carrying
capacity involve systematic clear-cutting of timber to
produce a series of even-aged stands (Verme 1965).
Under certain conditions, broadcast burning of the
logging slash constitutes a valuable silvicultural tool
in conifer seedbed preparation (Johnston 1977).

Clear-cutting and burning of swamps conceivably
could create ideal environments for small rodents and
lead to serious depredation on conifer seeds and/or
seedlings, thereby hindering or even preventing stand
reestablishment. Surprisingly little is known about
small mammal populations inhabiting the White
Cedar forest type (Manville 1949; Verme 1958; Ozoga
and Verme 1968). The purpose of this study was to
document changes in species composition and relative
abundance of small mammals following clear-cutting
to assess their potential impact on cedar regeneration.

Study Areas and Methods

This study was conducted in the Petrel Grade deer-
yard, which is situated 5.8 km northeast of Shin-
gleton, in Alger County, Michigan (46°34’N,
86°37’W). The area lies in the cold, deep, snow belt
along the south shore of Lake Superior. The two sites
investigated were about | km apart and comprised
readily accessible portions of an irregular-shaped 100-
km? conifer swamp. Both areas, before logging, were
densely timbered chiefly in mature (100-200 yr old)
White Cedar, with Black Spruce (Picea mariana) and
Balsam Fir (Abies balsamea) as minor associates.
Logging of the first site began in the 1965-1966 winter

and consisted of clear-cutting alternate strips 33-50 m
wide aligned east-west. Cutting progressed ina south-
erly direction, with the initial series of clear-cutting
being completed in two winters. Subsequent removal
of the residual strips began in the 1975-1976 winter
and was completed in 3 years.

Small mammal populations were studied by means
of snap traps (mainly Museum Specials) baited with
peanut butter during a 2-wk period in late August
and/orearly September from 1967 to 1977, excluding
1974. Because the cutting area initially was rather
small, in 1967 we established four parallel traplines
spaced about 44 m apart. On each line two traps were
placed at either edge of a clear-cut strip, with another
two set equidistant (+15 m) apart within the cleared
portion and residual stand, respectively. Thereafter,
two 750-m-long traplines were used, spaced 110 m
apart. The traplines were shortened in 1976 and espe-
cially 1977, as cutting of the residual strips neared
completion; hence, fewer trap nights of effort resulted,
because the original spacing was retained. A 440-m
trapline established parallel to and 550 m east of the
clear-cut unit served as a control,but it was aban-
doned after the area was also logged, in 1973.

Because the number of small mammals captured
was not based on equal trap nights among units, sta-
tistical analysis employed logarithmic transforma-
tions of results to test skewed data containing zeros
(Elliot 1971). The transformed data by species by year
were then tested by analysis of variance to appraise
differences among microhabitats.

The second study area consisted of a 4-ha neck of
conifer swamp bordered on one side by northern
hardwoods and grading into a mixed forest type on
the other side. The stand was clear-cut within 2 yr,
beginning in the 1971-1972 winter. In May 1973, half
of the unit was broadcast burned to eliminate slash; all
tree limbs less that 7.5 cm in diameter were consumed
by the hot fire. A rectangular (65 X 105 m) trapline

253

254

was established in each unit in 1974 and run for 2 wk
in late summer annually through 1977. Traps were
spaced 10 m apart around the perimeter of each line.
One corner of the slash-burned trapline extended to
within 50 m of adjacent northern hardwoods; how-
ever, at least a two-fold greater buffer zone existed
between the remaining portion and its proximity to
the edge of another forest type.

A three-way contingency table (Kullback et al.
1962) was used to analyze the catch data by species
according to site treatment (1.e., slash burned or
unburned).

Results
Strip Clear-cut

Compared to the uncut control, strip clear-cutting
collectively produced twice as many (P < 0.001) small
mammals during the 7-yr study, 8.9 vs. 4.6/100 trap
nights (Table 1). Logging resulted in an eightfold
greater abundance of Deer Mice and Meadow Voles.
Relative abundance of Least Chipmunks (Eutamias
minimus) and Northern Flying Squirrels (Glaucomys
sabrinus) increased nearly fourfold following logging.
Southern Red-backed Voles (Clethrionomys gapperi)
doubled in number on clear-cuts, whereas Masked
Shrews (Sorex cinereus) were slightly more numerous
in the control site. Conifer swamps in this locale
apparently do not provide suitable habitat for Bog
Lemmings (Synaptomys cooperi) or Meadow Jump-
ing Mice (Zapus hudsonius) as few were captured.
Red Squirrels (Tamiasciurus hudsonicus) were com-
mon in the area, but proved difficult to catch in the
Museum Specials.

There was no difference (P > 0.05) in population

TABLE | —Variation in small mammal abundance in a strip
clear-cut versus an uncut conifer swamp in upper Michigan
over a 7-yr period

Strip

clear-cut Uncut
Species site site
Deer Mouse 2.9(135)° 0.4(5)
Southern Red-backed Vole 1.6(76) 0.8(11)
Meadow Vole 0.7(31) 0.1(1)
Bog Lemming 0.1(1) 0.1(1)
Meadow Jumping Mouse — 0.1(1)
Least Chipmunk 1.3(62) 0.4(5)
Northern Flying Squirrel 0.3(12) 0.1(1)
Red Squirrel 0.1(1) _
Masked Shrew 2.1(100) 2.6(37)
Short-tailed Shrew 0.1(1) 0.1(2)
Total catch 8.9(419) 4.6(64)
Total trap nights 4712 1400

“Data are means/100 trap nights; total animals snap-
trapped are shown in parentheses.

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 2—Influence of conifer swamp microhabitat on small
mammal abundance in upper Michigan

Clear-cut Clear-cut/ Uncut
strip uncut strip
Species interior edge interior
Deer Mouse 2.1(40)* 1.7(35) 3.5(68)
Southern
Red-backed Vole 0.5(10) | 1.2(24) 3.3(65)
Meadow Vole 1.3(25) 0.5(10) 0.3(5)
Bog Lemming _ 0.1(1) —
Meadow Jumping
Mouse 0.1(1) _ —
Least Chipmunk 1.4(27) 1.3(27) 0.6(12)
Northern
Flying Squirrel _ 0.2(4) 0.5(9)
Red Squirrel = 0.1(2) a
Masked Shrew 2.7(51) 2.9(59) 2.1(41)
Short-tailed Shrew — — 0.2(4)
Arctic Shrew 0.1(2) 0.1(1) —
Total catch 8.3(156) 8.1(163) 10.4(204)
Total trap nights 1883 2013 1971

“Data are means/100 trap nights; total animals snap-
trapped are shown in parentheses.

levels of small mammals captured within the residual
stands, cleared strips, or their contiguous edges. How-
ever, certain species demonstrated rather distinct hab-
itat preferences (Table 2). Their general distribution
was consistent with known living requirements for a
given species. For example, Southern Red-backed
Voles definitely preferred uncut timber, whereas
Meadow Voles were mainly found in the cleared
strips.

Block Clear-cut-burn

Broadcast burning of slash in the block clear-cut
area resulted in increased (P< 0.05) abundance of
virtually all small mammals, yielding 15.1 anim-
als/ 100 trap nights versus 9.4 where fire was not used
(Table 3). The greatest proportional increase (nearly
seven-fold) was manifested by Deer Mice; however,
some of this increase could have been due to their
influx into the burn from the adjacent northern hard-
woods stand. Meadow Vole populations evidently
were comparable on both units. Masked Shrews,
Short-tailed Shrews (Blarina brevicauda), and Arctic
Shrews (Sorex arcticus) were moderately more
abundant in the burned unit. Least Chipmunks did
not exhibit any marked preference for one site over
the other. Although the burning of slash reduced use-
ful escape cover, the prompt and rank establishment
of fire-succession vegetation served effectively in this
regard and apparently provided sufficient food for
recolonization by the various small mammals. Mea-
dow Jumping Mice, Woodland Jumping Mice

1981

TABLE 3—Abundance of small mammals following clear-
cutting with and without broadcast burning of slash in an
upper Michigan conifer swamp

Slash Slash
Species burned left
Deer Mouse 2.0(20)° 0.3(3)
Southern Red-backed Vole 0.5(5) 0.4(4)
Meadow Vole 2.8(29) 2.8(29)
Meadow Jumping Mouse 0.2(2) 0.1(1)
Woodland Jumping Mouse 0.1(1) —
Least Chipmunk 1.9(19) 1.6(16)
Masked Shrew 5.6(57) 3.0(31)
Short-tailed Shrew 0.6(6) 0.2(2)
Arctic Shrew 1.5(15) 1.0(10)
Water Shrew 0.1(1) —
Total catch 15.1(155) 9.4(96)
Total trap nights 1024 1024

“Data are means/100 trap nights; total animals snap-
trapped are shown in parentheses.

(Napaeozapus insignis), and Water Shrews (Sorex
palustris) were scarce in both areas.

Discussion

Our studies indicate that clear-cutting in White
Cedar stands increased the relative abundance of
many small mammals. The increase was somewhat
greater when the resultant slash was broadcast burned
compared to being left intact. Findings of major
increases in small mammal numbers, especially Deer
Mice, following clear-cutting with or without slash
burning have been noted in other forest types (Trous-
dell 1954; Tevis 1956; Ahlgren 1966; Krefting and
Ahlgren 1974; Hoovan and Black 1976; Martell and
Radvanyi 1977). However, Sullivan (1979a) has
argued that these increases primarily represent maxi-
mum autumn populations, and that densities at other
seasons are far less because of habitat-limiting factors.
We captured some species 1n environments where they
would not be expected to occur. Perhaps such availa-
ble habitats served as dispersal sinks for immature
(subordinate) individuals during peak autumn density
(Lidicker 1975; Sullivan 1977). Age classification of
specimens we examined, however, was equivocal in
this regard.

The natural increase or influx of seed eaters like the
Deer Mouse and Least Chipmunk and/ or plant eaters
such as the Meadow Vole and Southern Red-backed
Vole potentially could reduce or prevent regeneration
of White Cedar after logging. However, examination
of clear-cut and burned experimental plots bordering
the first study area revealed there was more than
adequate (10 000 seedlings/ha) White Cedar 5 yr
later (unpublished data). Thus, small mammal popu-
lations evidently had little effect on stand regenera-

VERME AND OZOGA: SMALL MAMMAL POPULATIONS

BS

tion. White Cedar produces seed almost every year
and a good crop every 3—Syr (Johnston 1977); hence
germination is almost a continuous process.

Small mammal populations in conifer swamps in
this locale experience dramatic fluctuations in abun-
dance from year to year (Ozoga and Verme 1968). Any
major “explosion” in rodent density from their usual
comparatively low levels doubtless would attract a
host of predators (including the ubiquitous shrews)
which would help keep the numbers in check.
Although Deer Mice inimically influence forest
regeneration in other regions, Sullivan (1979b) notes
that efforts to control their depredations have failed
because of rapid immigration into vacant habitats
from surrounding areas. Application of proper silvi-
cultural techniques quite likely is the key to adequate
regeneration of White Cedar (Johnston 1977). No
artificial measures to control prospective damage by
small mammals seem warranted in conifer swamps of
upper Michigan. The main wildlife damage to cedar
reproduction 1s due to excessive browsing by White-
tailed Deer and Snowshoe Hare (Lepus americanus)
populations (Bookhout 1965); their destructive influ-
ence can be regulated by hunting, plus sound deeryard
management practices (Verme 1965).

Acknowledgments

This paper is a contribution from Federal Aid in
Wildlife Restoration Project W-117-R, Michigan,
and the Cusino Wildlife Research Station. We thank
G. E. Burgoyne for assistance in statistical analysis;
C. S. Bienz, D. L. DeLisle, and L. J. Perry also aided
in this endeavor. W. F. Johnston and R. H. Baker
graciously reviewed the manuscript.

Literature Cited

Ahlgren, C. E. 1966. Small mammals and reforestation fol-
lowing prescribed burning. Journal of Forestry 64:
614-618.

Bookhout, T. A. 1965. The Snowshoe Hare in upper Mich-
igan: its biology and feeding coactions with White-tailed
Deer. Michigan Department of Conservation, Research
and Development Report No. 38. 191 pp.

Elliot, J. M. 1971. Some methods for the statistical analysis
of samples of benthic invertebrates. Freshwater Biological
Association Scientific Publication Number 25. 147 pp.

Hoovan, E. F., and H.C. Black. 1976. Effects of some
clearcutting practices on small mammal populations in
western Oregon. Northwest Scientist 50: 189-208.

Johnston, W. F. 1977. Manager’s handbook for Northern
White Cedar in the North Central States. USDA Forest
Service General Technical Report NC-35. North Central
Forest Experiment Station, St. Paul, Minnesota. 18 pp.

Krefting, L. W., and C. E. Ahlgren. 1974. Small mammal
and vegetation changes after fire ina mixed conifer—hard-
wood forest. Ecology 55: 1391-1398.

256

Kullback, S., M. Kupperman, and H.H. Ku. 1962. Tests
for contingency tables and Markovchains. Technometrics
4: 573-608.

Lidicker, W. Z. 1975. The role of dispersal in the demo-
graphy of small mammals. /n Small mammals: their pro-
ductivity and population dynamics. Edited by F. B. Golly,
K. Petrusewicz, and L. Ryszkowski. Cambridge Univer-
sity Press, New York and London. pp: 105-128.

Manville, R. H. 1949. A study of small mammal popula-
tions in northern Michigan. University of Michigan Mis-
cellaneous Publication Museum of Zoology Number 78.
83 pp.

Martell, A. M., and A. Radvanyi. 1977. Changes in small
mammal populations after clearcutting of northern Onta-
rio Black Spruce forest. Canadian Field-Naturalist 91:
41-46.

Ozoga, J. J., and L. J. Verme. 1968. Small mammals of
conifer swamp deeryards in northern Michigan. Papers of
the Michigan Academy of Science, Arts, and Letters 51:
37-49.

Sullivan, T. P. 1977. Demography and dispersal in island
and mainland populations of the Deer Mouse, Peromys-

THE CANADIAN FIELD-NATURALIST

Vol. 95

cus maniculatus. Ecology 58: 964-978.

Sullivan, T. P. 1979a. Demography of populations of Deer
Mice in coastal forest and clear-cut (logged) habitats.
Canadian Journal of Zoology 57: 1636-1648.

Sullivan, T. P. 1979b. Repopulation of clear-cut habitat
and conifer seed depredation by Deer Mice. Journal of
Wildlife Management 43: 861-871.

Tevis, L. 1956. Effect of a slash burn on forest mice. Jour-
nal of Wildlife Management 20: 405-409.

Trousdell, H.B. 1954. Peak populations of seed-eating
rodents and shrews occur one year after loblolly stands are
cut. USDA Forest Service Southeastern Forest Experi-
ment Station Research Note 68, Ashville, North Carolina.

Verme, L. J. 1958. Localized variation in Masked Shrew
abundance. Journal of Mammalogy 39: 149-150.

Verme, L. J. 1965. Swamp conifer deeryards in northern
Michigan: their ecology and management. Journal of
Forestry 63: 523-529.

Received 26 September 1980
Accepted 27 January 1981

Nest-tree Sharing by Herons and Cormorants in Montana

LARRY S. THOMPSON

Montana Department of Natural Resources and Conservation, 32 South Ewing, Helena, Montana 59601

Thompson, Larry S. 1981. Nest-tree sharing by herons and cormorants in Montana. Canadian Field-Naturalist 95(3):

257-260.

At two central Montana colonies where Great Blue Herons (Ardea herodias) and Double-crested Cormorants (Phalacro-
corax auritus) share nest trees, herons arrived about three weeks earlier than cormorants and selected sites in healthy trees
near the center of the colony. Cormorants often used vacant nests or recently abandoned heron nests, sometimes in the same
tree as nesting herons. No evidence was seen of vertical stratification of nest sites by species, of nest eviction, or of competition
for nest sites. Opportunistic use of old heron nests by cormorants may reduce energy expenditures in nest-building and allow

speedier onset of reproduction.

Key Words: Great Blue Heron (Ardea herodias) , Double-crested Cormorant (Phalacrocorax auritus), nesting, colonial,

Montana.

Arboreal mixed nesting colonies of colonial water
birds have been well documented in North America.
In Atlantic wetlands, five or more species may be
found nesting within a single tree. Several workers
(e.g., Jenni 1969; McCrimmon 1978) observed nest-
site stratification and inter- and intra-specific compe-
tition for nests in mixed heron colonies. There has
been little documentation of interspecific aggression
or competition for nest sites between ardeids and
water birds of other orders.

In the northern Great Plains and adjacent Rocky
Mountains, the only large colonial water birds which
commonly nest in trees are the Great Blue Heron
(Ardea herodias) and the Double-crested Cormorant
(Phalacrocorax auritus). Lewis (1929) and Gers-
bacher (1939) noted no apparent interspecific aggres-
sion or competition for nest sites in mixed heron-
cormorant colonies. The relative positions of heron
and cormorant nests vary; Gersbacher (1939) reported
cormorants nesting on the highest treetops ina mixed
colony in Tennessee, whereas in Saskatchewan the
cormorants nested lower than herons (Vermeer and
Anweiler 1970). Of the 15 known arboreal cormorant
colonies in Montana, 12 also had herons, and the two
species often nested on the same branch (Thompson
1978).

During the spring and summer in 1976-1978, I
observed mixed colonies in Montana to gain insight
on the following questions: Do herons and cormor-
ants compete for nest sites? Do mixed colonies merely
result from limited nest site availability, ordo the two
species actively seek each other out? If the latter, what
adaptive advantages are conferred by sharing of nest
trees or mixed colonies?

Study Area and Methods
Two mixed heron-cormorant colonies in Montana,
at Lake Helena and Townsend, were selected for close

study. I also made some observations in four other
mixed colonies. The Lake Helena colony (46°41’N,
112°57’W) is located near the inlet of Tenmile Creek
into Lake Helena, a 200-ha impoundment on the Mis-
souri River. Herons have nested there since about
1935, but cormorants only in 1975 (S. Martin, 1110
Wilder Avenue, Helena, Montana, personal commun-
ication) and 1976. The Townsend colony (46°21'N,
111°31’W) is on an island in the Missouri River 2 km
upstream from Canyon Ferry Reservoir. Herons and
cormorants have shared this colony for many years (S.
Martin, personal communication).

During the nesting seasons of 1976-1978, 47.6 h
were spent observing behavior of birds. Most field
time was spent at Lake Helena in 1976, when the
group of trees harboring both heron and cormorant
nests was observed with a 20X telescope for 33.5h
during March through July. Between 17 April and 15
May 1976, 7.5 h were spent recording the exact occu-
pancy of each nest and the movements and behavior
of nesting birds. Relatively little time was spent at this
colony in 1977 and 1978 because cormorants were not
nesting there. The townsend colony could not be stu-
died in as much detail owing to the inaccessibility of
the island.

Results

Nest Distribution and Habitat

Lake Helena colony. Most heron nests were situated
at heights of 12-18 min the tops of old willows (Salix
spp.). The only trees shared by herons and cormorants
were three adjacent willows, with many dead
branches, about 250 m from the main colony (Figure
1). Allnests shown, except No. 12, were present before
birds arrived in 1976. Each tree had one active heron
nest for at least part of the 1976 season. One heron and
eight cormorant nests in this group of trees success-

ZS) Hh

258

metres

THE CANADIAN FIELD-NATURALIST

Vol. 95

FIGURE |. Relative position of nests in trees occupied by both herons and cormorants at Lake Helena. In 1976, cormorants
occupied nests 2,5,6,8,9, 10, 11,and 12, and herons occupied nests | and 3. Nest 1 1 was occupied by herons early in the

season but was later taken over by cormorants.

fully fledged young in 1976, but no active nests of
either species were found here in 1977 or 1978.

Townsend colony. Both herons and cormorants
shared nest trees in the Townsend colony during all
three years of this study. In 1978, at least 43 heron and
24 cormorant nests were active. Nests were located
near the tops of Black Cottonwoods ( Populus tricho-
carpa) 14-19 mhigh. Most cormorants nested in dead
or dying trees with little foliage near the edge of the
colony, although some nested with herons in densely
leafed-out trees near the center of the colony. As at
Lake Helena, no apparent horizontal or vertical strat-
ification of nests was observed. One tree, which sup-
ported two heron and five cormorant nests in 1976,
supported nine active cormorant nests and no herons
in 1978. Some trees not used in 1976 had nests of one
or both species in 1978. At least seven trees supported
both heron and cormorant nests in 1978.

Nesting Chronology and Behavior

Lake Helena colony. During the three years of the
study, herons first began using nests in late March; by
the first week of April, most herons had arrived. The
earliest arrivals selected old nests in the center of the
main colony. In 1976, pairs were first seen on 22
March. The peak of display, courtship, and territory
establishment was around 27 March — 10 April. Incu-

bation extended through late April and early May,
and fledging occurred from mid-June until mid-July.
Herons started nesting later in that portion of the
colony later occupied by cormorants, perhaps because
the open, nearly dead trees offered less security.

In 1976, the first two cormorants were seen on 15
April. By this time, herons at the main colony were
mostly paired and well into incubation, and nests 1, 3,
and | 1 (Figure 1) were occupied by herons. Nest 6 was
claimed by a cormorant almost immediately, and
three pairs were present by 21 April. By | May, nest I 1
had been abandoned by the lone heron and claimed by
a cormorant; nests 2,6, 8,9, and 10 were also occupied
by cormorants. The peak of cormorant courtship dis-
play was between 21 Apriland | May. Between | May
and 15 May, a new nest (No. 12) was built and occu-
pied by a cormorant; nest 5 was also occupied. No
further changes occurred, and by 10 July, two young
herons were present at nest 3 and young cormorants
(two to five per nest) in each of the eight occupied
nests. Herons were last seen in nest |, which was
unsuccessful, on 12 June.

Townsend colony. Both species arrived at the Town-
send site a few days ahead of the Lake Helena arrivals.
In 1976, herons and cormorants were first seen in nest
trees on 20 March and 10 April, respectively.

1981

Territoriality and Interspecific Aggression

While cormorants exhibited much intraspecific
aggression, no displays were obviously directed at
neighboring herons. Herons, on the other hand, were
observed giving the following displays in apparent
response to nearby cormorants: forward (twice),
arched neck (once), snap (four times), and fluffed neck
(once). (Nomenclature of heron displays follows that
of Mock (1976).) No direct evidence of interspecific
supplanting or evicting of an occupant from an estab-
lished nest was obtained, although, as noted before,
the unpaired male heron at nest | 1 at Lake Helena was
replaced by a cormorant. During 1978, four heron
nests were taken over by cormorants at Townsend. I
noticed no replacement of cormorants by herons at
nest sites during a breeding season, although M.
Bishop (Polson, Montana, personal communication)
reported this at Ninepipe National Wildlife Refuge in
western Montana.

Discussion

The observed sharing of nesting colonies by herons
and cormorants may simply be a chance association
forced by shortage of suitable nesting habitat. This is
unlikely, because suitable nesting sites—tall decidu-
ous trees near large bodies of water—are abundant
throughout Montana, and yet the degree of associa-
tion is considerable. Sharing of colonies and nest trees
might therefore confer some advantage on one or both
species. Because herons arrive at colonies nearly a
month earlier and are well entrenched by the time the
cormorants arrive, the latter are likely to be both the
instigators and the main beneficiaries of the associa-
tion. Krebs (1978) discussed two possible advantages
of group nesting: protection from predation and facil-
itation of food exploitation. However, these are not
likely explanations for colony sharing by herons and
cormorants.

Cormorants are much more reactive to approach-
ing humans than are herons; they would often flush
from nests when I approached within 200 m, whereas
herons would show no response until I came within
70 m. The predator-detecting ability of herons would
do cormorants little good, especially because the most
significant potential predators in the area are other
birds. By the “information center” hypothesis of Ward
and Zahavi (1973), coloniality affords easier location
of food items through cooperative food searching or
interspecific learning of feeding areas. Herons and
cormorants, however, obtain different food items by
quite different means and seldom feed in the same
portion of a water body.

Cormorants that use vacant heron nests in an estab-
lished colony reduce the amount of time and energy
Spent in nest construction, and can reproduce more

THOMPSON: MIXED HERON-CORMORANT COLONIES, MONTANA

259

quickly. This may well provide an advantage over
cormorants constructing new nests. Although 67-93%
of nests in heron colonies in Oregon and Montana are
occupied each year (Werschkul and McMahon 1976:
Thompson 1978), the other nests are available for
immediate occupancy by cormorants arriving in late
spring. Pair formation in cormorants begins imme-
diately upon arrival at Montana colonies; this could
indicate a selective pressure for speedy reproduction
upon arrival at nesting grounds. Jenni (1969) and
McCrimmon (1978) have documented a drastic con-
traction of defended territory around heron nests as
incubation gets under way; by the time cormorants
arrive in mid-April, nests in the same tree or on the
same branch as active heron nests may be occupied
with little interspecific strife. Where the temporal sep-
aration between heron and cormorant nesting peaks is
less than in the northern Great Plains, occupancy of
nests adjacent to territorial herons would be more
difficult, and the incidence of nest-tree sharing would
be lower. This may be the case, in light of the rarity of
published accounts of nest-tree sharing.

Cormorants arriving in Montana in spring proba-
bly select arboreal nesting colony sites in response toa
number of environmental cues, including: (1) presence
of active heron colonies, which indicates relative
safety of nest sites from predators; (2) presence of
vacant nests, which allow speedier onset of nesting; (3)
presence of nearby large water areas, which indicates a
stable food source; (4) water surrounding the nest
trees or the land on which the trees are located, which
means safety of fledged young from ground predators.
Cormorants may thus let herons do part of the work
of locating and constructing nests in suitable colony
sites.

Because the first herons arriving at a colony gener-
ally select the choice sites in the centre of the colony,
most of the surplus nests are located at the colony’s
edge, often in dead or dying trees. This may explain
the apparent preference of cormorants in Montana
for nest sites in partly bare trees near the edges of
heron colonies.

Thus, there appears to be no competition between
herons and cormorants in selection of nest sites, nor
vertical stratification of nest sites in trees. This sharing
of nest trees is likely to have arisen as an opportunistic
relationship benefiting cormorants considerably and
herons slightly.

Acknowledgments

Many thanks are due to S. Martin and G. Holton,
who first alerted me to the phenomenon of nest-tree
sharing at Lake Helena and who provided continuous
advice and encouragement; and to C. Winterburn and
his wife who kindly allowed me access to the Lake

260 THE CANADIAN FIELD-NATURALIST

Helena colony. I also thank many others who shared
their knowledge of herons and cormorants in Mon-
tana. Special thanks to my wife Susan for sharing
ideas and field work and for her invaluable tactical
and moral support. D. Chitty, D. McCrimmon, D.
Mock, and K. Vermeer provided helpful comments on
earlier versions of this manuscript.

Literature Cited

Gersbacher, E. O. 1939. The heronries at Reelfoot Lake.
Journal of the Tennessee Academy of Science. 14:
162-180.

Jenni, D. A. 1969. A study of the ecology of four species of
herons during the breeding season at Lake Alice, Alachua
County, Florida. Ecological Monographs 39: 245-270.

Krebs, J. R. 1978. Colonial nesting in birds, with special
reference to the Ciconiiformes. /n Wading birds. Edited by
A. Sprunt, IV, J. C. Ogden, and S. Winckler. Research
report number 7 of the National Audubon Society.

Lewis, H. F. 1929. The natural history of the Double-

Vol. 95

crested Cormorant ( Phalacrocorax auritus). H. C. Miller,
Ottawa.

McCrimmon, D.A., Jr. 1978. Nest site characteristics
among five species of herons on the North Carolina coast.
Auk 95: 267-280.

Mock, D. W. 1976. Pair-formation displays of the Great
Blue Heron. Wilson Bulletin 88: 185-230.

Thompson, L. S. 1978. A Montana statewide heron survey
(abstract). Proceedings of the Montana Academy of
Sciences. 38: 88.

Vermeer, K., and G. Anweiler. 1970. Great Blue Heron
colonies in Saskatchewan in 1970. Blue Jay 28: 158-161.

Ward, P., and A. Zahavi. 1973. The importance of certain
assemblages of birds as “information centres” for food
finding. Ibis 115: 517-534.

Werschkul, D. F.,and E.£ McMahon. 1976. Someeffects of
human activities on the Great Blue Heron in Oregon.
Wilson Bulletin 88: 660-662.

Received 14 December 1979
Accepted 23 January 1981

Occurrence and Growth Patterns of the American Brook Lamprey,
Lethenteron lamottenii, in the Ottawa River

J. LANTEIGNE, J. M. HANSON, and S. U. QADRI

Department of Biology, University of Ottawa, Ottawa, Ontario KIN 6N5

Lanteigne, J., J. M. Hanson, and S. U. Qadri. 1981. Occurrence and growth patterns of the American Brook Lamprey,
Lethenteron lamottenii, in the Ottawa River. Canadian Field-Naturalist 95(3): 261-265.

Ammocoetes, transforming ammocoetes, and adults of the American Brook Lamprey, Lethenteron lamottenii, are reported
for the first time from the Ottawa River. Specimens were caught only from the sandy areas of three islands when the water
temperature was between 13 and 25°C. Lampreys were not present in the shallow water (< I.1 m) when water temperatures
were less than 10°C. The largest ammocoete captured was 198 mm long and weighed 10.4 g. The prebranchial and branchial
lengths (as a percentage of total length) became proportionately shorter as length increased and the trunk and tail lengths
became proportionately longer with increasing length. This showed that morphometric characters undergo changes particu-
larly during early larval life. The length-weight relationship, log WT = 2.819 log TL — 5.467, and the decrease in the values for
the Fulton’s condition factor with increasing length further demonstrate that growth was allometric.

Key Words: lamprey, ammocoetes, Lethenteron lamottenii, body proportions, condition, length-weight, pigmentation,

habitat, Ottawa River.

The American Brook Lamprey, Lethenteron
lamottenii, occurs in Canada in streams flowing into
the St. Lawrence River, Lake Champlain, in the Great
Lakes (other than the eastern portion of Lake Onta-
rio), and in the Noire, St. Anne de la Perade, and St.
Maurice Rivers of Quebec (Scott and Crossman
1973). Vladykov (1972) reported its occurrence in the
Gatineau River (a tributary of the Ottawa River)
below the Petites Chutes at Limbourg, Quebec, where
10 adults were collected on 17 May 1959. No other
collections have been made since then in the Ottawa or
Gatineau rivers.

The purpose of this study 1s to report the first occur-
rence of the American Brook Lamprey in the Ottawa
River and to describe some aspects of its growth.

Materials and Methods

American Brook Lamprey were collected with a
backpack electrofisher (type VII, Smith-Root Inc.,
Vancouver, Washington) in the shallow water of the
Ottawa River near Ottawa, Ontario. Sampling was
done along the north and south shorelines of the
Ottawa River, in all small creeks flowing into the
river, and around Kettle Island, Upper Duck Island,
and Lower Duck Island in an area from Lac
Deschenes to just east of Lower Duck Island (Figure
1). Additional sampling was done near the mouth of
the Petite Blanche River (45°30’N, 74°32’W) and
Green Creek (45°28’N, 74°33’W). The study area has
been described by Ericksson (1974) and Hanson and
Qadri (1980). Water temperatures reached 25°C in
late July and early August 1979 in the shallow water
around Upper Duck Island.

From samples collected at 2-wk intervals from 4

261

June to 7 November 1979, we kept 271 ammocoetes, 9
transforming ammocoetes, and 3 newly transformed
adults. Additional sampling was done at 2-wk inter-
vals from 5 May to 18 June 1980. Specimens were
preserved in a 5% formalin solution and transported
to the laboratory where total length (mm) and wet
weight (to the nearest 0.01 g) of specimens blotted dry
were recorded. The Fulton’s condition factor, K(TL)
(Hile 1941; Lagler 1956), was calculated for each spec-
imen and the means and standard errors for 10-mm
length-groups calculated. The length—weight relation-
ship was determined for total length (mm) and wet
weight (g) by the least squares method of linear regres-
sion (Ricker 1975). The mean values for the lengths of
the prebranchial, branchial, trunk, and tail regions (as
a percentage of total length) were calculated for 42
ammocoetes in 10-mm length-classes. The relation-
ship of these same characters to total length were
calculated by linear regression by the least squares
method. Pigmentation patterns were recorded for the
same 42 ammocoetes following the method described
by Vladykov (1950). All specimens have been depos-
ited with the National Museums of Canada Ichthy-
ological Collection (NMC 80: 958-962).

Results

American Brook Lamprey were not captured in the
Ottawa River, or its small tributaries, above the
Chaudiére Falls. Specimens were not caught along the
shorelines or from the small tributaries of the Ottawa
River below the Chaudiere Falls, but ammocoetes
were common around Kettle Island, Upper Duck
Island, and Lower Duck Island.

The shorelines of the islands differ from the shore-

262

THE CANADIAN FIELD-NATURALIST

Vol. 95

/eerue SSS

ROU RCHC EERE

¥ CHAUDIERE
FALLS

OT TAWA 1
ONTARIO

FIGURE |. Sampling sites where Lethenteron lamottenii was collected in the Ottawa River, Ottawa—Hull area.

lines of the main river channel in that the substrate of
the former is mostly sand and the substrate of the
latter is mostly clay. Ammocoetes were not collected
from the eastern tip of Kettle Island which has a hard
clay substrate nor were there any present along the
north shoreline of Kettle Island which is subjected to
pollution and dredging activities by the Canadian
International Paper Company at Gatineau, Quebec.

The large ammocoetes, transforming ammocoetes,
and newly transformed adults were usually taken
among the emergent vegetation in depths less than
0.3 m. Lampreys were not present in the shallow water
(<1.1 m) after temperatures decreased to 10°C in
October 1979, but ammocoetes were present in the
shallow water on 5 May 1980, when the water temper-
ature was 16°C.

The upper lip and suborbital areas of 42 American
Brook Lamprey ammocoetes were devoid of pigment
and the prebranchial blotch was absent. The branchial
region above the branchial openings was without
pigment for about 2 mm, leaving a whitish band
below the dorsal pigmented area. The pigmentation
on the caudal fin varied from weak to moderate; an
unpigmented margin was always present. At capture,
the newly transformed adults were yellow-brown on
the dorsal surface and the sides and the ventral surface
were silver-grey. The number of trunk myomeres
ranged from 65 to 70 (usually 68) for 42 ammocoetes.
The three adults had 69, 70, and 71 myomeres. The
values obtained for the body proportions of 42
ammocoetes divided into !0-mm_length-groups
showed the changes occurring in the different body

areas throughout larval life. The prebranchial and
branchial regions became proportionately shorter
with increasing body length, this process occurring
more rapidly in the smaller length-classes, and level-
ing off at about the 140- to 149-mm length-class (Fig-
ure 2). The relative lengths for the trunk and tail
regions increased with total length and leveled off
somewhat at about the 140- to 149-mm length-class
(Figure 3). After 160-169 mm, there seemed to be a
slight decrease in the relative length of the trunk
region and a slight increase in the tail region.

The total length of the 271 ammocoetes and 9 trans-
forming ammocoetes had a range from 60 to 198 mm
and 134 to 202 mm, respectively. The three newly
transformed adults were 146, 166, and 181 mmin total
length. The linear regression fitting the data for length
and weight of ammocoetes was log WT = 2.819 log TL
— 5.467, n= 271, r= 0.9947. As the slope of the
length-weight relationship shows that the weight is
proportionately less as length increased, the values for
Fulton’s condition factor should decrease as the
ammocoetes grow. The values for Fulton’s condition
factor decreased froma value of 0.185 for the smallest
ammocoetes toa value of 0.129 for the longest ammo-
coetes (Figure 4).

Discussion

The typical habitat of L. Jamottenii was described
by Vladykov (1949) as cold brooks and small rivers,
usually in association with Slimy Sculpin (Cottus
cognatus) and Brook Char (Sa/velinus fontinalis), and
by Morman (1979) as pool-riffle streams with low

1981

Prebranchial length

10

\ log Y = 0.337 log TL + 1.559;
9 + g ;

Deter n= 42; r=-0.9612
8 —
it Pit es
iat tol eect a

6 +
5

Branchial length

log Y= 0.260 log TL + 1.596;

n= 42; r=-0.8635

+t
~t

See =

Body lengths as percentage of total length

Dn NDA A HHA HAHAH HAHAH HH MH
OTS GB) Sy ce Ca
Des UCSF re Ties oveth eon nite Umea tient Hime es. aeindeet ert OTM ee uct J ont maT)
QC Oo & GO EC GF eC 2 Oem EG ©
@ 1S Cu Cy GY 1, se I

Total length (mm)

FIGURE 2. Mean value for the relative length of the prebran-
chial and branchial regions of Lethenteron lamottenti
for each 10-mm ammocoete length-group.

temperatures and stable bottom. In Delaware, habi-
tats occupied varied from mud to sand at the Coastal
Plain localities to clay and gravel at the Piedmont
localities with water temperatures ranging from 9.0 to
12°C (Rohde et al. 1976). The Ottawa River popula-
tion differs in that it inhabits a large river, and the
ammocoetes were only found in the sandy areas of
three large islands when water temperatures were
between 13.0 and 25°C. The ammocoetes were not
found in shallow water at temperatures below 10°C
(mid-October) when, presumably, they had moved
into deeper waters.

The patterns of pigmentation of the different body
areas of the larval American Brook Lamprey from the
Ottawa River correspond to the descriptions given by
Vladykov (1950) for specimens collected in tributaries
of the St. Lawrence River in Quebec.

Ammocoetes collected in Delaware apparently
show proportionately longer tail and_ branchial
lengths but shorter trunk lengths (Rohde et al. 1976)
than the Ottawa River population (Table 1); speci-
mens from Kentucky have similar branchial and tail
lengths (Distler 1957). Such comparisons do not,
however, show the ontogenic changes in body propor-

LANTEIGNE ET AL.: AMERICAN BROOK LAMPREY, OTTAWA RIVER

263
56 Trunk length
55 er
54 ge
53
52 A + log WT = 0.272 log TL +1170;

n= 42 r= 08199
5|

Spa ep

Body lengths as percentage of total length

Tail length
29 pease
28 elec! ‘
=
27 oe
26 dud +
25 “£ log WT = ra paee: bw
n=42; r=0.
24
yp tt tt tt tt
Cop) an Dn n Dn oO on (oz) oO a n nD
TS 9) te) Se ee) ey Sc
! 1 1 1 ! ! 1 1 1 ! ! 1 1 '
OOOO! (ONC COO OO BOmOnsn©
GS) (Se ey Se a Sy
Total length (mm)

FIGURE 3. Mean value for the relative length of the trunk and
tail regions of Lethenteron lamottenii for each 10-mm
ammocoete length-group.

log K= -O.216 log TL-0.3935
r=-08949 n= 42

—

NE

—_

—

(e)

=

oO

£

(S

2 Peers oRgenae

o —t

Cc

{e)

O

0100 a ie ene ES

i (DOD OOD NC) TO) tO) OD OD) OO) DD
Ot OD) OS No YT Oe 2 @
WORT iy ml eu Mt Wem TT Mere (PUR eee Vumert( eco. ae th vee ‘ll
j@) (~) fe) ~e) OO EO Oo OFF & © we 2
OQ NSC 1b) Mees Ga Se GS Ge!
em Se RS re eee ee ee es ey ome LC Ne
Yr Om GO £& BW & mM © te G& io @
SSeS SN SD po Bae 22

Total length (mm)

FIGURE 4. Condition factor of ammocoetes of Lethenteron
lamottenii from the Ottawa River, Ottawa- Hull area.
Means and 95% confidence limits are presented.
Sample size in each 10-mm length-group is in
parentheses.

264

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE |—Comparison of the number of myomeres and body proportions (as a percentage of total length) of American Brook
Lamprey from various drainages. Ranges, means, and number of specimens examined are presented

No. of Prebranchial Branchial Trunk Tail
TL (mm) myomeres length length length length Location Source
133-182 69-73 = 10.4-12.2 — 26.1-29.6 Greasy Creek, Distler (1957)
158.0 70.8 11.4 28.0 Kentucky
16 16 16 16
— 65-74° — _ — = Big Creek, Kott (1974)
69.1 Ontario
303
100-143 64-72” = 12.6-14.4 51.0-53.7 27.7-31.6 Delmarva Rohde et al. (1976)
121.0 68.0 13.3 52.0 29.7 Peninsula,
19 101 19 19 19 Delaware
a 63-70 _ — _— _ St. Lawrence Vladykov (1950)
67.1 River, Quebec
381
65-197 65-70 6.0-9.4 9.3-14.7 51.5-56.0 24.6-28.8 Ottawa River, Present study
130.0 67.9 V2 11.4 54.3 Dale. Ontario
42 42 42 42 42 42
“Adults only.

»Ammocoetes and adults combined.

tions relative to body length. The number of speci-
mens studied from both the Delaware and Kentucky
populations was small and only incuded a narrow
(<50 mm) range of lengths. With the exception of the
upper limit of the range for tail length for the Dela-
ware population, the ranges of values reported for the
body proportions of the Delaware and Kentucky
ammocoetes fall within the range of values observed
for the Ottawa River population. The average number
of myomeres reported by Distler (1957) is higher than
for other studies, but this is probably a sampling bias
due to the small number of specimens examined. The
range of the number of myomeres of ammocoetes
from the Ottawa River readily falls within the range
for the species (Vladykov 1950; Rohde et al. 1976).
The length-weight relationship for 139 ammo-
coetes of the American Brook Lamprey in Delaware
was log WT = 2.76 log TL — 5.31 (Rohde et al. 1976).
The relationship is very similar to that calculated for
the lampreys in the present study. Rohde et al. (1976)
did not calculate the condition factors for American
Brook Lamprey, but they did calculate the condition
factors for Least Brook Lamprey, Lampetra aepyp-
tera, which has a length-weight relationship of log
WT = 2.72 log TL — 5.24. They found a decrease in the
value of the condition factors from 0.32 for the 1 1- to
20-mm length-group to 0.20 for the largest ammo-
coetes (121-130 mm). Hardisty (1944) also found that
the value of the condition factor decreased with
increasing length for Lampetra planeri. More studies
are needed to determine if this pattern of allometric

growth observed by Hardisty (1944), Rohde et al.
(1976), and in the present study occurs with all lam-
prey ammocoetes.

Literature Cited

Distler, D. A. 1957. Some differential characteristics of
ammocoetes of two species of lampreys Lampetra aep\'p-
tera (Abbott) and Lampetra lamottenii (LeSueur). M.Sc.
thesis, University of Louisville, Kentucky. 32 pp.

Ericksson, C. 1974. A two-year study of the higher aquatic
plant community in a section of the Ottawa River, Can-
ada, with emphasis on its role in mercury uptake. M.Sc.
thesis, University of Ottawa, Ottawa. 156 pp.

Hanson, J. M.,andS. U. Qadri. 1980. Observations on the
biology of Black Crappie, Pomoxis nigromaculatus
(LeSueur), in the Ottawa River. La Naturaliste Canadien
107: 35-42.

Hardisty, M. W. 1944. The life history and growth of the
Brook Lamprey (Lampetra planeri). Journal of Animal
Ecology 13: 110-122.

Hile, R. 1941. Age and growth of the Rock Bass, Amblo-
plites rupestris (Rafinesque), in Nebish Lake, Wisconsin.
Transactions of the Wisconsin Academy of Sciences, Arts,
and Letters 33: 189-337.

Kott, E. 1974. A morphometric and meristic study of a
population of the American Brook Lamprey, Lethenteron
lamottenii (LeSueur) from Ontario. Canadian Journal of
Zoology 52(8): 1047-1055.

Lagler, K. F. 1956. Freshwater fishery biology. Wm. C.
Brown Company, Dubuque, Iowa. 421 pp.

Morman, R.H. 1979. Distribution and ecology of lam-
preys in the lower peninsula of Michigan, 1957-1975.
Great Lakes Fishery Commission Technical Report 33.
59 pp.

1981

Ricker, W. E. 1975. Computation and interpretation of
biological statistics of fish populations. Bulletin of the
Fisheries Research Board of Canada 191: 382 pp.

Rohde, F. C., R. G. Arndt, and J. C.S. Wang. 1976. Life
history of the fresh water lampreys, Okkelbergia aepyp-
tera and Lampetra lamottenii (Pisces: Petromyzonidae),
on the Delmarva peninsula (East Coast, United States).
Bulletin of the Sourthern California Academy of Sciences
75: 99-111.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada, Bulletin
184: 966 pp.

LANTEIGNE ET AL.: AMERICAN BROOK LAMPREY, OTTAWA RIVER

265

Vladykov, V.D. 1949. Quebec lampreys. List of species
and their economical importance. Department of Fisher-
ies, Quebec 26: 67 pp.

Vladykov, V. D. 1950. Larvae of eastern American lam-
preys. 1—Species with two dorsal fins. La Naturaliste
Canadien 77 (3-4): 73-95.

Vladykov, V. D. 1972. Lampreys of the Ottawa area. Trail
and Landscape 6(4): 105-132.

Received 19 September 1980
Accepted 2 February 1981.

A Time-Activity Budget for Breeding Mallards (Anas platyrhynchos)
in Manitoba

RODGER D. TITMAN

Delta Waterfowl Research Station, Portage La Prairie, Manitoba RIN 3A1
Present address: Department of Renewable Resources, Macdonald College of McGill University, Ste.-Anne-de Bellevue,
Quebec H9X 1CO

Titman, Rodger D. 1981. A time-activity budget for breeding Mallards (Anas platyrhynchos) in Manitoba. Canadian
Field-Naturalist 95(3): 266-271.

Individually marked Mallards (Anas platyrhynchos) were observed while breeding in pothole country near Minnedosa,
Manitoba, in 1970 and 1971, to determine relative amounts of time devoted to different daily activities. Breeding Mallards
spent most of their time at one or two locations within their territories. During incubation, pairs appeared to avoid openareas.
Data are presented showing relative amounts of time spent by males and females foraging, resting, loafing and alert, preening
and bathing, and in aggression. Females spent more time foraging than males before and during nesting. Males must spend
time alert and in aggressive activity to protect their reproductive investment, thus time spent foraging, and their body weights
tend to decrease while they are territorial. Although females can rely toa large extent upon endogenous energy stores, I believe
that they must still forage to complete incubation successfully.

Key Words: Mallard, Anas platyrhynchos, behavior, habitat utilization, pothole habitat, Manitoba.

In the last decade, one approach to the study of
behavioral ecology has involved the compilation of
time—activity budgets. Recent publications reporting
time—activity budgets for birds have included those
for breeding Gadwalls (Anas strepera), Northern
Shovelers (A. c/ypeata), Black Ducks (A. rubripes),
Northern Pintails (A. acuta), and Blue-winged Teals
(A. discors) (Dwyer 1975; Poston 1974; Afton 1979;
Wooley and Owen 1978; Seymour and Titman 1978;
Hickey 1980; Derrickson 1977; Miller 1976; Stewart
and Titman 1980). Dwyer (1975) and Owen (Wooley
and Owen 1978; Owen and Reinecke 1979) translated
time—activity budgets into energy budgets, and Afton
(1979) considered behavioral strategies for optimum
use of environmental resources. Time-activity
budgets help in understanding habitat utilization and
niche occupation by animal species. The aim of this
study was to determine the time allocated to different
daily activities by breeding Mallards (Anas platy-
rhynchos). Because the female assumes sole responsi-
bility for the nest and appears to invest more energy
than the male in reproduction, it was hypothesized
that the time-activity budgets of paired males and
females are different.

Study Area

The study area consisted of two 260-ha plots, with
104 and 158 potholes, respectively, amid agricultural
fields in southwestern Manitoba (50°10’N, 99°45’W),
about 30 km north of Brandon. The geological his-
tory, topography, soils, climate, and plant communi-
ties of the Minnedosa region were described by Evans
et al. (1952) and the local history, habitat conditions,

and waterfowl population trends by Kiel et al. (1972).
The ponds in the region ranged from less than 0.04 ha
to about 5.0 ha in area and from 8 cm to 2.5 m in
depth. About 52% were less than 0.2 ha in area. The
interspersion of agricultural land, cover, roads, and
water in the area where one pair was followed in 1971
is Shown in Figure 1.

Methods

Data were obtained from observations of 14
marked Mallards between early April and mid-July in
1970 and 1971. Both members of four pairs were
marked, the females only of four pairs, and only the
males in two pairs. Most of the information concerns
the members of one pair (both marked) which were
observed for over 155 h (Table 1). Collectively the
nine other pairs were observed for about 67 h.

Observations were conducted from first light until
after dark in periods lasting from 30 min to 5 h. Activ-
ity was recorded continuously instead of being
sampled at timed intervals (Altmann 1974). Periods of
observation totalling 6-8 h daily, 6 d/ wk, were stag-
gered to permit observation during each daylight hour
at least once a week. Observations aided by 7X
binoculars and 20 to 45X telescope were made at
varying distances froma tower, blinds, or an automo-
bile. In some cases where vision was obstructed by
vegetation, the birds’ location was known (e.g., Table
2), although behavior could not be recorded. Hens
were not seen as they laid or incubated eggs. Activities
were timed using a watch and stopwatch, and notes
were recorded on tape. Movements were recorded on
maps. Mallards were captured using nest traps and

266

1981 TITMAN: MALLARD TIME-ACTIVITY BUDGET 267
TABLE |—Distribution of observation time (min) for one marked pair of Mallards — Minnedosa, Manitoba, 1971
Central Renesting Renesting

Standard First laying Incubation interval Second laying Incubation interval Third laying
Time 19 April- 13 May 14-22 May 23-29May 30 May-5June 6-15 June 16-24 June 25 June — |! July
04:25-08:00 350 895 250 480 324 132 158
08:01-11:00 170 90 270 160 340 475 261
11:01-15:00 30 30 235 350 235 259 278
15:01-18:00 285 155 210 340 281 160 10
18:01-21:08 295 618 379 330 380 110 101
Totals 1130 1788 1344 1660 1560 1136 808

cannon nets (Giles 1969) and individually marked
with numbered plastic nasal saddles (Bartonek and
Dane 1964).

Activities
categories:

were classified into the following

Preening — bathing — all behavior associated with
external body maintenance and comfort (= “care of
the body surface and related activities” of McKinney
ISS)

Foraging — all behavior associated with search for
food, capturing it, and manipulating or ingesting it,
either on land or in the water.

Aggression — this ranged from simple threat and
avoidance to energetically costly chasing in pursuit
flights (McKinney 1965).

Resting — birds in repose held their heads down on
their shoulders, either sleeping with eyes closed and
bill tucked into scapular feathers or awake yet
lethargic.

Loafing — Alert — Loafing birds might either be
walking, swimming, or stationary. They held their
heads upright and were active and fully aware of their
surroundings but not performing any of the above
activities. Alert birds usually held their heads further
upstretched and watched or listened for potential
intruders, predators, or disturbance (Dwyer 1975).

Results

Location of Activity

Only two pairs sighted more than 75 times provided
sufficient information to describe the location of
activity within their range. The male of pair A spent
70% of his time on or within 60 m of one pond (about
2.0 ha) in his territory (Figure |). During the time that
the male behaved territorially, his mate was with him
48% of the time (Table 2). She spent most of the
remainder at her nest. The other pair, which was
observed for 846 min, spent 66% of its time on or
within 50 m of two ponds within its territory. These
results support Hochbaum’s (1944) observation that

breeding dabbling duck pairs concentrate their activ-
ity at one or two locations.

Pair A was kept under observation from 19 April to
8 July 1971, during which time the female made three
nest attempts. Beginning 5 May, she laid nine eggs.
After 8 d of incubation, the nest was destroyed by a
mammalian predator on 22 May. Eight eggs were laid
in a second nest that was initiated 8 d later. This was
destroyed after 8 d of incubation during the night of
15 June. Although it was not located, a third nest
probably was initiated on 23 June, and incubation
continued until 8 July when the female was last seen.

Pair A spent more time in cover and less time in the
open after the female had started to incubate, particu-
larly after beginning to incubate her second nest later
in the breeding season.

Time Allocation

The relative amounts of time spent by male and
female Mallards performing different activities at dif-
ferent times throughout the day are summarized in
Figure 2. Both sexes tended to be more active (i.e., not
resting, loafing, or alert) during the later daylight
hours. This illustration does, however, mask the well-
known momentary peaks in activity around sunrise
and sunset (Winner 1972).

Time spent flying was not plotted in the histograms
(Figures 2 and 3) because it was difficult to measure,
especially when birds were lost from sight, and
because it constituted a small proportion of total
activity. It is nonetheless important because of its high
energetic cost and should receive more careful consid-
eration in future studies.

Overall, females spent significantly more time
foraging than their mates (P< 0.01) as illustrated in
Figures 2 and 3. Although intensity was not measured
directly as Seymour and Titman (1978) did, it
appeared obvious from observation that females fed
at a greater rate.

Foraging was the predominant activity of female A
during recesses while she was incubating (Figure 3).
She also spent a lot of time preening and bathing

268 THE CANADIAN FIELD-NATURALIST Vol. 95

200 METRES
SCALE ee ——

LEGEND

X, NEST LOCATION - FIRST NEST
A MALE WITHOUT FEMALE - 20 TO 40 SIGHTINGS A >? 40 SIGHTINGS

O PAIR- 20 TO 40 SIGHTINGS @ > 40 SIGHTINGS
[-] WATER [_] POTENTIAL NESTING COVER
[_] FALLOW OR CULTIVATED FIELD BS TREES

FIGURE |. Area of activity of pair A from 19 April to 8 July 1971, near Minnedosa, Manitoba. Cultivated fields east and south
of the crossroads were seeded in grain crops (wheat, barley). In the northwest segment the field immediately west of
pond 16 was seeded with alfalfa while the one northeast was left in stubble from the previous year’s wheat.

TABLE 2— Percentages of time spent by male A on various parts of his home range (see Figure |) during daylight hours and
time spent with his mate and with other males during the breeding season

Fields near Total % %
Pond Fields near ponds Pond Pond Else- minunder time with time with

Period 18 pond 18 62 and 109 108 22 where observation female A other males
Laying Ist clutch 58.6 8.8 WS) 0.3 0.8 23.6 1130 70.3 0.5
Incubation 52.8 8.0 10.3 4.3 1.6 23.0 1808 22.6 20.8
Renesting interval 58.3 21.0 2.6 ell 37 11.8 1564 81.0 2.6
Laying 2nd clutch 62.2 23.8 3.1 0.4 4.6 5.9 1438 70.1 16.8
Incubation De] 28.1 D5) Dee) 0 Dales) 1380 Del 51.4
Renesting interval 15.5 45.3 4.1 29 13.1 0.1 1134 83.7 4.6
Laying 3rd clutch 30.8 62.1 0 1.8 Sod) 0 768 23.0 6.9

Overall (total
or mean) 45.5 24.7 4.8 7.6 Sol! 13.7 9222 48.3 16.0

1981

PREENING - BATHING

cd AGGRESSION

[_] RESTING -LOAFING- ALERT

UW,

100

FORAGING

04:25 OBO! 11:0! 15:0! 18:0!
-08:00 — 1100 —- 5:00 - 18:00 -2!'08

MALES

04:0! O8:O! 11:0! 15:0! 18:01
- 08:00 -11°00 -I5K 9 -18:00 -21:08

FEMALES

FIGURE 2. Histograms showing activity of all marked male
and female Mallards during daylight hours in the
breeding seasons of 1970 and 1971, Minnedosa,
Manitoba. Only time spent off the nest (by the female)
is shown.

LOAFING-ALERT
Ea RESTING
FORAGING

MALE A

TITMAN: MALLARD TIME-ACTIVITY BUDGET

269

during these incubation recesses, as Sowls (1955: 103)
observed, yet contrary to his impression, feeding
assumed primary importance. Her whole daily ener-
getic requirement had to be satisfied within a period
Caldwell and Cornwell (1975) found to last an average
of 78 min.

Meanwhile, her mate spent less time foraging and
more time loafing or alert, particularly while female A
was with him, which would allow their quick response
to the approach of an intruder (Figure 3). Male A
spent proportionally more time being aggressive to
intruders on their territory while his mate was laying
her first clutch. Later in the season, during the absence
of his mate at her nest, male A spent an increasing
proportion of his time with other males (Table 2). In
general, males tended to be gregarious when they left
their mates that were caring for nests and young.

Discussion

There were significant differences between the
time-activity budgets of paired male and female Mal-
lards. These differences were principally in the greater
time spent foraging by females to meet the high caloric
cost of egg production. Others have recorded similar
differences between breeding male and female Gad-
walls, Northern Shovelers, Blue-winged Teals, Black
Ducks, and Harlequin Ducks (Histrionicus histrioni-

PREENING - BATHING

faswlaletetay ei

| iil bel
WL Teale lal

1
ii
I

P =
LAY INC REN LAY INC REN LAY

FEMALE A

FIGURE 3. Histograms showing activity of pair A for seven periods through the breeding seasons, 1971. Only time off the nest

(by the female) is shown.

270

cus) (Dwyer 1975; Poston 1974; Afton 1979; Miller
1976; Stewart and Titman 1980; Seymour and Titman
1978; Bengtson 1972).

Weight losses from the beginning to the end of the
breeding season indicated a decline in energy stores
over this period by both females and males (Folk et al.
1966; Young 1977). During the breeding season, Mal-
lards do not spend sufficient time feeding to secure the
energy to balance their daily energy budgets. This
phenomenon is known in Ross’s Geese (Anser rossii),
Lesser Snow Geese (A. caerulescens), Canada Geese
(Branta canadensis), and Common Eiders (Somateria
mollissima); however, these large species can rely toa
greater extent upon endogenous reserves (Ryder 1970;
Ankney 1977; Ankney and MacInnes 1978; Raveling
1979; Milne 1976).

Eggs are costly for females to produce and then
incubate. Incubation may be no more costly than
normal daily activity, but it involves a time investment
precluding foraging. Afton (1979) argued that North-
ern Shovelers have to forage during incubation to
survive and successfully produce young. Using my
data (Titman 1973) and those of Caldwell and Corn-
well (1975), Afton calculated that Mallard females
feed for about 23 h during the 26-d incubation period
whereas Shovelers feed for 58 h over a 23-d incuba-
tion period. Although the Mallard is larger and may
rely more upon endogenous reserves, it must still for-
age to obtain sufficient energy and nutrients to sur-
vive. Incontrast, female geese and eiders may not feed
at all during incubation, and some die in the process
(Ryder 1970; Ankney and MacInnes 1978; Raveling
1979; Korschgen 1977).

A major role of the male is to ensure that his mate
can feed undisturbed while off her nest. This is
accomplished in different ways by different species.
Shovelers defend exclusive territories (Seymour
1974), whereas a Mallard territory is not as well
defined, but the female is defended from intruders
(Titman 1973). The male Northern Shoveler defends a
territory until 2 d before hatching when the female is
more attentive to her nest and spends less time off it.
Male Mallards usually abandon their incubating
mates about one-half way through incubation. The
timing may have evolved to ensure protection of the
female to a point when she could complete incubation
and survive on her reserves alone.

Males spent a lot of time loafing and alert instead of
feeding, especially during laying and incubation by
their mates. For Gadwalls, Dwyer (1975) recorded
that males spent highest proportions of time alert just
before and during laying. I believe that a male Mallard
must spend time alert as well as in aggression to pro-
tect his reproductive investment. This prevents him
from devoting sufficient time foraging to maintain
body weight (Folk et al. 1966; Young 1977). This

THE CANADIAN FIELD-NATURALIST

Vol. 95

prediction can be tested by monitoring physiological
condition and by compiling detailed time-activity
budgets for breeding males.

Acknowledgments

Iam grateful to H. A. Hochbaum, former director
of the Delta Waterfowl Research Station, and the late
Bruce S. Wright, former director of the Northeastern
Wildlife Station, for their advice and encouragement.
Robert O. Bailey gave field assistance and Hugh
Boyd, Frank McKinney, Alex Dzubin, and Norman
Seymour generously offered their comments and crit-
icisms during my study. I am grateful to L. Fredrick-
son, J. B. Gollop, and J. P. Ryder for critical review
of different drafts. Funds were granted by the North
American Wildlife Foundation through the Delta
Waterfowl Research Station and by the Department
of Biology and Faculty of Graduate Studies and
Research at the University of New Brunswick.

Literature Cited

Afton, A.D. 1979. Time budget of breeding Northern
Shovelers, Wilson Bulletin 91: 42-49.

Altmann, J. 1974. Observational study of behavior: sam-
pling methods. Behaviour 49: 227-267.

Ankney, C. D. 1977. The use of nutrient reserves by breed-
ing male Lesser Snow Geese Chen caerulescens caerules-
cens. Canadian Journal of Zoology 55: 1984-1987.

Ankney, C.D., and C.D. MaclInnes. 1978. Nutrient
reserves and reproductive performance of female Lesser
Snow Geese. Auk 95: 459-471.

Bartonek, J. C., and C. W. Dane. 1964. Numbered nasal
discs for waterfowl. Journal of Wildlife Management 28:
688-692.

Bengtson, S.-A. 1972. Breeding ecology of the Harlequin
Duck Histrionicus histrionicus (L.) in Iceland. Ornis
Scandinavica 3: 1-19.

Caldwell, P.J., and G. W. Cornwell. 1975. Incubation
behavior and temperatures of the Mallard duck. Auk 92:
706-731.

Derrickson,S. R. 1977. Aspects of breeding behavior in the
Pintail (Anas acuta). Ph.D. dissertation, Department of
Ecology and Behavioral Biology, University of Minne-
sota, Minneapolis.

Dwyer, T. J. 1975. Time budget of breeding Gadwalls. Wil-
son Bulletin 87: 335-343.

Evans, C.D., A.S. Hawkins, and W.H. Marshall.
1952. Movements of waterfowl broods in Manitoba. U.S.
Department of the Interior, Fish and Wildlife Service,
Special Scientific Report, Wildlife Number 16. 47 pp.

Folk, C., K. Hudec, and J. Toufar. 1966. The weight of the
Mallard, Anas platyrhynchos, and its changes in the
course of the year. Zoologické Listy 15: 249-260.

Giles, R. H., Jr. (Editor). 1969. Wildlife management tech-
niques. Third edition. The Wildlife Society, Washington,
DC. 633 pp.

Hickey, T. E., Jr. 1980. Activity budgets and movements of
Black Ducks (Anas rubripes) in Prince Edward Island.
M.Sc. thesis, Department of Renewable Resources,
McGill University, Montreal, Quebec. 95 pp.

1981

Hochbaum, H. A. 1944. The Canvasback on a prairie
marsh. American Wildlife Institute, Washington, D.C.
182 pp.

Kiel, W.H., Jr., A. S. Hawkins, and N.G. Perret.
1972. Waterfowl habitat trends in the Aspen parkland of
Manitoba. Canadian Wildlife Service Report Series
Number 18. 61 pp.

Korschgen, C. E. 1977. Breeding stress of female eiders in
Maine. Journal of Wildlife Management 41: 360-373.
McKinney, F. 1965. Spacing and chasing in breeding

ducks. Wildfowl Trust 16th Annual Report: 92-106.

Miller, K. J. 1976. Activity patterns, vocalizations and site
selection in nesting blue-winged Teal. Wildfowl 27: 33-43.

Milne, H. 1976. Body weights and carcass composition of
the Common Eider. Wildfowl 27: 115-122.

Owen, R.B., Jr., and K. J. Reinecke. 1979. Bioenergetics
of breeding dabbling ducks. pp. 71-102. Jn Waterfowl and
Wetlands — an integrated review. Edited by T. A. Book-
hout. Proceedings of 1977 Symposium, North Central
Section, The Wildlife Society. Madison, Wisconsin.

Poston, H. J. 1974. Home range and breeding biology of
the Shoveler. pp. 132-137. In Saskatoon Wetlands
Seminar, Canadian Wildlife Service Report Series
Number 6.

Raveling, D. G. 1979. The annual cycle of body composi-
tion of Canada Geese with special reference to the control
of reproduction. Auk 96: 234-252.

Ryder, J. P. 1970. A possible factor in the evolution of
clutch size in Ross’ Goose. Wilson Bulletin 82: 5-13.

Seymour, N.R. 1974. Territorial behaviour of wild Sho-
velers at Delta, Manitoba. Wildfowl 25: 49-55.

TITMAN: MALLARD TIME-ACTIVITY BUDGET

Pay

Seymour, N. R.,andR. D. Titman. 1978. Changes inactiv-
ity patterns, agonistic behaviour and territoriality of Black
Ducks (Anas rubripes) during the breeding season in a
Nova Scotia tidal marsh. Canadian Journal of Zoology
56: 1773-1785.

Sowls, L. K. 1955. Prairie ducks. Stackpole, Harrisburg
and Wildlife Management Institute, Washington, D.C.
193 pp.

Stewart, G. R.,and R. D. Titman. 1980. Territorial behav-
iour by prairie pothole Blue-winged Teal. Canadian Jour-
nal of Zoology 58: 639-649.

Titman, R. D. 1973. The role of the pursuit flight in the
breeding biology of the Mallard. Ph.D. thesis, Depart-
ment of Biology, University of New Brunswick, Frederic-
ton. 201 pp.

Winner, R. W. 1972. Activity of Black and Mallard Ducks
in a controlled environment. Journal of Wildlife Man-
agement 36: 187-191.

Wooley, J. B., Jr., and R. B. Owen, Jr. 1977. Metabolic
rates and heart-rate metabolism relationships in the Black
Duck (Anas rubripes). Comparative Biochemistry and
Physiology 57(3A): 363-367.

Young, D. A. 1977. Characteristics of the moults in the
male Mallard (Anas platyrhynchos). M.Sc. thesis,
Department of Zoology, University of Alberta, Edmon-
ton. 107 pp.

First submission received 10 June 1975
Revision received 30 July 1980
Accepted 10 February 1981

Invasion of a New Reservoir by Fishes: Species Composition,

Growth, and Condition

ROBIN MAHON! and MARK FERGUSON

Zoology Department, University of Guelph, Guelph, Ontario NIG 2W1
'Present address: Marine Fish Division, Bedford Institute of Oceanography, Dartmouth, Nova Scotia B2Y 4A2.

Mahon, Robin, and Mark Ferguson. 1981. Invasion of a new reservoir by fishes: species composition, growth, and

condition. Canadian Field-Naturalist 95(3): 272-275.

After the first summer of inundation, the ichthyofauna of the reservoir known as Guelph Lake was compared to that of the
Speed River before impoundment. Several species that had been rare in the river showed marked increases in relative
abundance in the reservoir. The Bluntnose Minnow ( Pimephales notatus) was most abundant among 19 species collected. For
young-of-the-year of three species, Bluntnose Minnow, Common Shiner (Notropis cornutus), and Creek Chub (Semotilus
atromaculatus), which were abundant enough for comparison, growth was faster in the reservoir than in the river. For these
species and White Sucker (Catostomus commersoni), the average condition (weight at a given length) was significantly higher

in the reservoir.

Key Words: impoundment, colonization, reproduction, abundance, river.

Although the construction of reservoirs for flood
control and low flow pollution abatement are com-
mon practice in southern Ontario, there are few stu-
dies of the biology of fishes in such reservoirs, or of the
effects of these impoundments on their rivers. In 1975,
Mahon et al. (1979) studied the distribution and pro-
ductivity of fishes in the Speed River, Ontario, to
provide a background for assessment of the develop-
ment of the ichthyofauna in the impending reservoir
(Guelph Lake) and its impact on the river.

In spring 1976 the reservoir was filled. It covers an
area of about 400 ha with the midpoint located at
43°37’'N, 80°15’W, its maximum depth is about 12 m
(D. Tregunna and K. Walker, Landscape Architec-
ture, University of Guelph, 1974, unpublished data).
The purpose of our study was to record the species of
fishes that had invaded the inshore regions of the
reservoir by the fall of 1976, and to compare the
species composition and where possible the growth
and condition of the fishes with those in the Speed
River before impoundment.

Methods

A seine (10 X 0.9 m with 6-mm circular mesh) was
used to collect fishes from 18 localities throughout the
main body of the reservoir (Figure 1) on 12 and 18
September 1976. After identification, three species for
which there were enough individuals for growth to be
compared were measured (SL) to the nearest | mm
and weighed to the nearest 0.1 g. For Common Shiner
(for scientific names see Table 1) and Creek Chub, all
individuals were measured; for Bluntnose Minnow, a
subsample of 559 individuals was taken by mixing all
the fish in a tray and separating off about one quarter
of the sample. In each case there was a distinct mode
of small fish in the length-frequency histogram. Pre-

Die

vious experience has shown that for these species this
mode is a reliable indicator of age (Mahon and Balon
1977b; Mahonetal. 1979). Growth was compared for
these young-of-the-year.

For the above three species and for White Sucker,
condition of individuals of all sizes was compared to
that of individuals of the same species in the Speed
River in 1975. This was done by comparing the
observed weight at a given length (Wo) with the
expected weight for a fish of that length before
impoundment as predicted from weight—length equa-
tions given by Mahon et al. (1979). The ratio of
observed to expected weight indicates whether the fish
is heavier or lighter than average for its length, with
heavier fish being in better condition. An average
ratio of one would indicate no difference in condition.
Direct comparison of growth and condition between
the reservoir and the river 1s possible because the
Speed River localities were sampled a year earlier on
14 and 17 September 1975.

Results

The land inundated by the reservoir was primarily
agricultural with scattered woods which were cleared
before filling. Therefore the fishes were collected from
a variety of substrate types: submerged grasses, corn
stubble, cleared woodlot, mudflat, and old river
floodplain. In contrast the Speed River is a typically
fast-flowing, warmwater (non-trout) stream with a
gravel, cobble, and rubble substrate. Some lateral
pools and backwaters did, however, provide habitat
for more lentic species (Mahon et al. 1979).

Altogether, 4130 fishes comprising 19 species were
collected from the reservoir at the end of the first
summer after filling. The relative abundance of these
species is shown in Table |. The fish community was

1981

MAHON AND FERGUSON: FISH INVASION OF NEW RESERVOIR

as

FiGureE |. Guelph Lake with sampling locations shown. For the exact location of the reservoir, refer to Figure |, Mahonet al.

(1979).

clearly dominated by the Bluntnose Minnow. Most of
the fishes collected were young-of-the-year.

In Table 2, the mean standard length for young-of-
the-year of three species is compared between the
reservoir and three localities combined in the Speed
River. These localities were a few kilometers above,
within, and just below the reservoir (QN2, QN4, and
QNS, respectively, Mahon et al. 1979). For each spe-
cies growth was faster in the reservoir than in the river.

For fishes of the above three species and the White
Sucker, the mean condition of individuals in the
reservoir for each species was significantly greater
than one (t-test, P< 0.05) (Table 3). In no case was
there any tendency for condition to be correlated with
size.

Discussion

Of the 18 species of fishes collected from the upper
Speed River watershed in 1975 (Mahonetal. 1979), 14
were collected in this study. The remaining five species
collected from the reservoir have been reported from
the Speed River watershed, but Carp and Brown Bull-
head only from below the city of Guelph wherein there
are several weirs and dams that would be expected to
restrict the upstream movement of these species.
Therefore, we did not expect to find them in the
reservoir.

There is considerable overlap between the species
lists for the reservoir and the river, but the differences

in relative abundance are marked. Table | shows the
relative abundance of the 14 species of fishes collected
in the Speed River in the immediate vicinity of the
reservoir a year before impoundment. In the river, the
Fantail Darter, Creek Chub, and Northern Hog-
sucker were dominant, whereas in the reservoir,
Bluntnose Minnow and Common Shiner were most
abundant. The dominance of the Bluntnose Minnow
young-of-the-year probably reflects their reproduc-
tive strategy which is to guard a brood of eggs at-
tached to the underside of a log, stone, or similar
object. Amongst those fishes that were common in the
river, only the Bluntnose Minnow would be expected
to reproduce equally well in the reservoir. Brook Stic-
kleback, Bluntnose Minnow, and Fathead Minnow
were the dominant species in two temporary streams
in southern Ontario (Williams and Coad 1979); this
suggests that they tend to invade newly created habi-
tats rapidly. In addition these three are not strict
‘stream species’ and are common in shallow bog lakes
and ponds. Although lacustrine populations of Creek
Chub and Common Shiner do occur, they usually
require flowing water for spawning (Scott and Cross-
man 1973). Shoal-spawning is a possibility provided
there is silt-free gravel or rubble, thus providing con-
ditions which are similar to those in a stream (Kitchell
et al. 1977); such conditions are rare in Guelph Lake.
For similar reasons neither the Smallmouth Bass nor
the Rock Bass are likely to colonize this reservoir.

274 THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE |—The relative abundance (percentage) of fishes in Guelph Lake, 1976, and in the Speed River in the vicinity of the

reservoir before impoundment (Mahon et al. 1979)

Species

Bluntnose Minnow (Pimephales notatus)
Common Shiner (Notropis cornutus)
Pumpkinseed (Lepomis gibbosus)

Fathead Minnow’ (Pimephales promelas)
Brook Stickleback (Culaea inconstans)
Redbelly Dace® (Chrosomus eos)

Brassy Minnow (Hybognathus hankinsoni)
Creek Chub (Semotilus atromaculatus)
White Sucker (Catostomus commersoni)
Carp” (Cyprinus carpio)

Johnny Darter’ (Etheostoma nigrum)
Rosyface Shiner (Notropis rubellus)
Fantail Darter (Etheostoma flabellare)
Northern Hogsucker (Hypentelium nigricans)
Brown Bullhead” (/ctalurus nebulosus)
Pearl Dace® (Semotilus margarita)
Blacknose Dace (Rhinichthys atratulus)
Rock Bass (Ambloplites rupestris)
Mottled Sculpin (Cottus bairdi)
Smallmouth Bass (Micropterus dolomieui)
Longnose Dace ( Rhinichthys cataractae)
Mudminnow (Umbra limi)

Total number of fishes collected
Total number of species

“Occurs in the Speed River above the dam.
"Occurs in the Speed River below the dam.

Guelph Speed
Lake River
60.7 4.2
13.6 5.6
8.6 0.1
6.0 —
3.4 0.1
8} —
2.0 —
1.8 16.3
0.5 2.8
0.3 —
0.3 a
0.2
0.1
0.1
0.1
0.1 —
0.1 0.2
0.1 1.3
0.1 0.1
= 3.9
— 0.9
== 0.2
4130 1897
19 14

“Previously collected only from the Eramosa River, a tributary which enters the Speed River, below the dam, in Guelph.
“Collected from the Speed River in 1951 but not found in 1976.

TABLE 2—Comparison of mean standard length (SL) of
young-of-the-year between Guelph Lake and three localities
(combined) in the Speed River before impoundment

Guelph Lake
SL+SD (n) SL (n)

36.443.63 (224) 29.4 (387)
54.7-6.30 (44) 40.9 (579)
33.35.74 (515) 27.6 (1234)

Speed River
Species

Common Shiner
Creek Chub
Bluntnose Minnow

TABLE 3— Mean condition of individuals of four species in
Guelph Lake, based on weight-length relationships for
these species in the river before impoundment

Species MeanconditiontSD (n)
Common Shiner 1.32+0.099 (84)
White Sucker 1.28+0.266 (22)
Bluntnose Minnow 1.130.177 (78)
Creek Chub 1.13£0.113 (65)

Those species that should successfully colonize a
reservoir are usually rare in fast-flowing rivers. How-
ever, this study illustrates that the small populations
inhabiting backwaters and lateral pools are sufficient
for widespread and rapid colonization. Pumpkinseed,
Brook Stickleback, Redbelly Dace, and Fathead
Minnow though rare in the river became common in
the reservoir in just a few months (Table 1). The
species composition ina vegetated lateral pool in 1975
was similar to that in the reservoir at the end of the
first summer. The Mudminnow provides an excep-
tion. Dominant in vegetated lagoons on the shore of
Lake Erie (Mahon and Balon 1977a) and abundant in
the above-mentioned pool, this species was not found
in the reservoir. The Mudminnow spawns in early
spring before the snow melts. At that time the reser-
voir was not sufficiently filled for much new suitable
habitat to have been created.

Growth and condition of those fishes for which
comparison was possible was better in the reservoir
than in the river(Tables 2 and 3). Gasaway (1970) also
found increased growth ina new reservoir but this was

1981

a temporary effect. Increased growth and better con-
dition could result from higher temperature in the
reservoir and/or from increased food abundance,
provided that food was limiting in the river. Unfortu-
nately, we are unable to separate the effects of these
factors and thereby evaluate the hypothesis that
stream fish populations are commonly limited by
density-independent rather than density-dependent
factors. We are able, however, to eliminate the possi-
bility that the difference arose through differences in
temperature between years. Monthly mean maximum
and minimum air temperatures for the 1975 and 1976
growing seasons (April—October) at a station about
20 km from the reservoir were similar (Figure 2). If
any difference between the years exists it is towards
higher temperatures in 1975, this would tend to
emphasize the differences between the fish in the
reservoir and the river.

This study shows characteristic shifts in species
composition and growth after impoundment (Jack-
son and Rogers 1976). Notable is the speed with which
many species, originally rare in the river, were able to
invade the newly created habitat. There was no
marked reduction in the number of species as is consi-
dered normal, possibly because of the presence of
leftover riverine species not yet eliminated by the
changed conditions and the explosive expansion of
those species better suited to the reservoir. In a per-
manent reservoir, this would most likely be a tempor-
ary phenomenon associated with filling (June 1976).
Guelph Lake, however, is a typical flood control

30

20

MONTHS

FIGURE 2. Mean monthly maximum and minimum air
temperatures during the 1975 and 1976 growing sea-
sons (data supplied by Department of Land Resource
Sciences, University of Guelph).

MAHON AND FERGUSON: FISH INVASION OF NEW RESERVOIR

ay

reservoir, drawn down every fall and refilled every
spring during the thaw. The extent and timing of the
drawdown depends on the amount of water required
during the summer for low flow augmentation. Under
these conditions the reservoir fauna would not be
expected to exhibit the successional sequence typical
of a more stable reservoir constructed, for example,
for hydorelectric power generation. Instead the reser-
voir may be characterized year after year by a fish
community similar to that observed in this study.

Acknowledgments

We thank Susan Mahon, who helped with the field
work and criticized the manuscript. Thanks also to the
Experience ’76 Program, Ontario, for financial assist-
ance during the sampling. The comments of an ano-
nymous referee served to improve the manuscript
greatly. This project was supported by a Natural
Sciences and Engineering Research Council operating
grant to Eugene Balon.

Literature Cited

Gasaway, C. R. 1970. Changes in the fish population in
Lake Francis Case in South Dakota in the first 16 years of
impoundment. United States Fish and Wildlife Service
Technical Paper 56. 30 pp.

Jackson, P. N. B., and K. H. Rogers. 1976. Cabora Bassa
fish populations before and during the first filling phase.
Zoologica Africana 11: 373-397.

June, F. C. 1976. Changes in young-of-the-year fish stocks
during and after filling of Lake Oahe, an upper Missouri
River storage reservoir. United States Fish and Wildlife
Service Technical Paper 87. 25 pp.

Kitchell, J. F.. M. G. Johnson, C. K. Minns, K. H. Loftus,
L. Greig, and C. H. Olver. 1977. Percid habitat: the river
analogy. Journal of the Fisheries Research Board of Can-
ada 34: 1936-1940.

Mahon, R.,and E. K. Balon, 1977a. Fishcommunity struc-
ture in lakeshore lagoons on Long Point, Lake Erie, Can-
ada. Environmental Biology of Fishes 2: 71-82.

Mahon, R., and E. K. Balon. 1977b. Ecological fish pro-
duction in Long Pond, a lakeshore lagoon on Long Point,
Lake Erie. Environmental Biology of Fishes 2: 261-284.

Mahon, R., E. K. Balon, and D. L. G. Noakes. 1979. Dis-
tribution, community structure and production of fishes in
the upper Speed River, Ontario: a preimpoundment study.
Environmental Biology of Fishes 4: 219-244.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada Bulletin
Number 184. 966 pp.

Williams, D. D., and B. W. Coad. 1979. The ecology of
temporary streams III. Temporary stream fishes in south-
ern Ontario, Canada. International Revue Gesamter
Hydrobiologie 64: 501-515.

Received 10 May 1980
Accepted 14 November 1980

Distribution and Harvest of Canada Geese (Branta canadensis)
in Southern Manitoba Prior to Development of Oak Hammock Marsh

DENNIS G. RAVELING! and CHARLES C. DIXON2

'Canadian Wildlife Service, Winnipeg, Manitoba R3T 2N6
Present address: Division of Wildlife and Fisheries Biology, University of California, Davis, California 95616
2Manitoba Department of Mines, Natural Resources and Environment, Winnipeg, Manitoba R3N 0H6

Raveling, Dennis G., and Charles C. Dixon. 1981. Distribution and harvest of Canada Geese (Branta canadensis) in
southern Manitoba prior to development of Oak Hammock Marsh. Canadian Field-Naturalist 95(3): 276-280.

The numbers, distribution, and harvest of different subspecies of Canada Geese (Branta canadensis) in autumn in southern
Manitoba were studied between 1967 and 1970 prior to the restoration of Oak Hammock Marsh near Winnipeg. Small
numbers of Canada Geese were found in many locations, but only about a dozen sites were used by 1000+ birds. Peak
populations in southern Manitoba varied from about 15 000 to 30 000 Canada Geese. The Interlake region between Lakes
Manitoba and Winnipeg contained the most geese and was the site of an estimated 45% of the kill by hunters. Giant Canada
Geese (B. c. maxima) predominated in the harvest in most of the Interlake, in southeast and west-central Manitoba. Brantac.
interior and a similar goose were the major forms killed in the southern Interlake near where the redeveloped Oak Hammock
Marsh now attracts autumn peak populations of up to 120 000 migrant Canada Geese. Small Canada Geese (B. c. hutchinsii

and similar forms) predominated in the kill on the Red River, at Delta, and in southwest Manitoba.

Key Words: Branta canadensis, Canada Goose; distribution, numbers, hunter kill, subspecies.

Canada Goose (Branta canadensis) occupies a
broad geographic range, from the west to the east
coast and from the arctic islands to the prairies (Dela-
cour 1954), and is highly variable in morphology.
Proper understanding and management of Canada
Geese depends on increasing knowledge of the interre-
lationships of different subspecies and populations.
The resident breeding Canada Geese in southern
Manitoba are the large and relatively pale-colored
B. c. maxima (Hanson 1965; Raveling 1976, 1978).
The dominant form in northern Manitoba is the
smaller and darker B. c. interior which nests primarily
on the Hudson Bay Lowlands (Hanson and Smith
1950; Vaught and Arthur 1965; Malecki et al. 1980),
but some also nest on the Canadian Shield in the
interior (Raveling 1977). They stop during migration
in southern Manitoba as do small Canada Geese (B. c.
Pparvipes—hutchinsii complex of MaclInnes 1966),
which nest on the tundra adjacent to, and on the
islands of, Hudson Bay and along the arctic coast.

Habitat loss or restoration may have pronounced
effects on the distribution of migratory birds and, for
game species, these changes may greatly alter the
impact of hunters on populations. Oak Hammock
marsh is a 3360-ha managed wetland 23 km north of
Winnipeg, Manitoba. The area was acquired between
1967 and 1972 and developed to desired water levels
by 1974 (Oetting and Dixon 1975). A major effect of
Oak Hammock has been a change in distribution and
numbers of geese in southern Manitoba during their
autumn migration. Annual concentrations of Canada

Geese have averaged 80 000 between 1974 and 1979 in
the Oak Hammock area.

In anticipation of an expanded program of goose
management and harvest in Manitoba, the Canadian
Wildlife Service (CWS) initiated a study in 1967 in
cooperation with the provincial Department of Natu-
ral Resources (DNR) to determine the status and
distribution of Canada Geese in southern Manitoba.
The purpose of this paper is to present data on the
occurrence of Canada Geese in autumn in southern
Manitoba and on the relative importance of different
subspecies to hunters in different geographic areas
during the late 1960s. These data also form a baseline
with which to compare subsequent results brought
about by changing population numbers, migration
patterns, and habitat.

Methods and Materials

Weekly aerial inventories of goose numbers were
conducted, weather permitting, in areas A, B, D, and
F (Figure 1) by the CWS and in areas E and G by the
DNR between mid-September and early November
1967-1970. Aerial searches in area C were irregular
and supplemented with counts made by agency per-
sonnel on the ground. Specific locations visited regu-
larly are indicated on Figure |. Although we recognize
that Canada geese occurred over a broader range,
these locations were generally acknowledged to be of
most importance during migration.

The distribution of harvest of Canada Geese was
indicated by the return of goose tail feathers by hun-

276

1981

(Survey boundaries) y}

(o,)

+

ONTARIO

— A
100 98°

FiGurE |. Locations of autumn aerial inventories and har-
vest of Canada Geese in southern Manitoba. Area A
—l. Marshy Point Goose Sanctuary, 2. Dog-
Moosehorn Lakes, 3. Reykjavik Point, 4. Peonan
Point — Portage Bay, 5. Pinemuta Lake - Lake St.
Martin, 6. Sturgeon Bay, 7. Saint Lakes, 8. Fisher
Bay, 9. Washow Bay, 10. Hecla Island - Riverton
Marshes, 11. Sleeve Lake. Area B — 1. Shoal Lakes,
2. Oak Hammock, 3. Netley-Libau Marshes. Area C
— 1. Alf Hole Goose Sanctuary, 2. Whitemouth
Lake, 3. Red River. Area D — Delta Marshes. Area E
— |. Whitewater Lake, 2. Oak-Plum Lakes, 3. Pem-
bina Valley Lakes. Area F — Big Grass Marsh. Area
G — The Pas vicinity: Kelsey, Little Kelsey, Niska,
Helldiver, Cul de Sac, Mawdesely, Cumberland
Lakes.

ters cooperating in the CWS parts collection survey
(Canadian Wildlife Service 1968). The raw data were
provided by G. Cooch (CWS) so we could proportion
the harvest to our areas of interest which differed from
the zones used by CWS.

Determinations of the subspecies of Canada Goose
killed by hunters were made by the authors. Some
hunters were contacted in the field, and the bag of
some others was examined at check stations periodi-
cally established along major road systems by the
DNR. Most specimens were, however, saved for us by
a group of cooperative sportsmen who were contacted
personally and who were provided with plastic bags,

RAVELING AND DIXON: CANADA GEESE, MANITOBA

100 fe
kilometres

BUT)

tags, and detailed diagrammatic instructions for sav-
ing parts from each goose that they or their friends
had killed. Parts saved from each goose were head and
neck including the feathering below the black neck, a
wing, a leg (below the feathers), and tail feathers,
preferably cut off above the pygostyle so that the
entire tail “fan” was preserved and measurements
could be compared to those available from live and
museum specimens.

Measurements of these parts were compared to
those available in the published literature and records
available from trapped geese from different popula-
tions and museum specimens. Paramount among
these sources were Hanson (1951, 1965), MacInnes
(1966), and Raveling (1976, 1977). Some hunters
knew how to identify sex of geese and provided us
with that information, but most did not. As adult
males of one race may overlap in size with females or
immatures of a larger size race, discrimination of sub-
species depended on personal experience and knowl-
edge of color and proportions as well as obvious mea-
surements such as culmen, tarsus, toe, wing, and tail
lengths.

Results and Discussion
Distribution and Numbers of Canada Geese

Small numbers of Canada geese occurred in many
areas in autumn, but few locations attracted over 1000
birds. Marshy Point (area Al, Figure 1) regularly held
2000-4500 geese during autumn. Other locations
where a peak of over 1000 (but less than 2500) Canada
Geese were seen on at least one occasion were (see
Figure | for locations indicated by letter-numeral)
Dog-—Moosehorn Lakes (A2), Peonan Point — Por-
tage Bay of Lake Manitoba (A4), Washow Bay in
Lake Winnipeg (A9), Hecla Island — Riverton
Marshes in Lake Winnipeg(A10), Sleeve Lake (A11),
Shoal Lakes (BI), Alf Hole Goose Sanctuary (C1),
Delta Marsh shoreline of Lake Manitoba (D),
Whitewater Lake (El), Swan Lake in the Pembina
Valley (E3), Big Grass Marsh (F), and Kelsey and
Little Kelsey lakes near The Pas (G).

In general, no area attracted a major concentration
of Canada Geese (Table 1) relative to many other
places in North America. Most Canada Geese in areas
A, C, and G were giant Canada Geese that nested in
the vicinity (see Raveling 1976, 1978, and below).
Based on knowledge of the size and migration pat-
terns of other populations of Canada Geese killed in
Manitoba (MaclInnes 1966; Vaught and Kirsch 1966),
it is obvious that several 100 000 migrants passed over
southern Manitoba and those that stopped did so for
short periods, in many areas, and in relatively small
numbers. Regular aerial inventories did not always
coincide with transitory peak populations; therefore,

278

TABLE !|—Peak numbers of Canada Geese observed in
southern Manitoba during autumn aerial inventories,
1967-1970

Peak numbers
of Canada Geese

Area
(see Figure 1)

A 6 400-14 100
B 1 400-4 900
C 1 300-3 100
D 700-1 700
E 800-3 200
Je 600-2 500
G 2 300-3 000
Totals (peaks in a given year)" 15 400-29 900

“Totals differ from sum of columns because peak numbers in
different areas did not occur in the same year.

estimates in Table | are minimal. For example, an
estimate (from the ground) of 9500 Canada Geese was
made at West Shoal Lake 3 d after an aerial inventory
had revealed 2300. Despite these problems, Table 1
reveals the general diffuse nature of Canada Goose
distribution in southern Manitoba in the late 1960s.
Geese stopping during days of major migration
movements did not stay for more than a few hours or
days.

Distribution of Harvest

The widespread distribution of Canada Geese
shown by aerial inventories was also reflected by the
variety of locations from which hunters reported
killing birds from which they sent in tail feathers
(Table 2). About 20% of the kill occurred in locations
where inventories had not been taken. In total, how-
ever, the Interlake region between Lakes Manitoba
and Winnipeg was the most important area of harvest
of Canada Geese with about 45% of the reported kill
(areas A and B, Figure |). This was divided approxi-
mately equally between the Shoal Lakes to Netley
Marsh area (area B) and the remainder of the southern
Interlake region (area A).

Subspecies Composition of Harvest
Small Canada Geese were of two types: those typi-
cal of McConnell River and Southampton Island,

THE CANADIAN FIELD-NATURALIST

Vol. 95

Northwest Territories, described by MacInnes (1966),
and an even more diminutive form. For purposes of
this report, they were combined. While most “middle-
sized” Canada Geese were clearly B. c. interior, there
was also a form slightly smaller (but distinctly larger
than the “small” Canadas) in all measurements. This
form was not obviously present in Wisconsin and
Illinois samples reported by Hanson (1951, 1965) or
Raveling (1977). It probably represents an undes-
cribed subspecies. For this paper, they were combined
with B. c. interior as middle-sized Canada Geese. The
giant Canadas were B. c. maxima as described by
Hanson (1965).

The type of Canada Goose killed by hunters varied
greatly by geographic area and even between closely
spaced locations within an area (Table 3). Small Can-
ada geese predominated at Delta, Big Grass Marsh, in
the southwest, and along the Red River (area C3).
Their migration tended to be more irregular than that
of the larger geese and their appearance in the harvest
fluctuated accordingly in these areas (Table 2).
Middle-sized Canada Geese were the major form
killed in area B[primarily in the vicinity of West Shoal
Lake, data came from 143 geese measured in 1965 and
1966 by L. Bidlake (1967 unpublished report of the
Manitoba Wildlife Branch) and from 77 geese exam-
ined by the senior author in 1967]. Giant Canada
Geese were most common in the harvest in areas A,
Cl, and G which also correspond to their main nesting
areas (see Raveling 1976, 1978). The most even distri-
bution of harvest of different forms of Canada geese
occurred at Big Grass Marsh (area F) and Red River
(C3).

Comparison of the distribution of geese (Table 1),
and harvest (Table 2), with subspecies composition
(Table 3) demonstrates the importance of local breed-
ing giant Canada Geese in the Interlake to harvest by
hunters in the late 1960s. This population was expand-
ing rapidly (Gulden and Johnson 1968; Raveling
1978), but the kill was predominantly in Canada. Just
a few kilometres away, middle-sized geese were the
major component of the harvest in area B.

Since the establishment of Oak Hammock Marsh,
large numbers of migrant geese of the Eastern Prairie

TABLE 2—Distribution of Canada Goose tail fans received from hunters by area in southern Manitoba’

Year N A B C

1968 120 30.8 24.2 1.7
1969 246 15.0 20.3 5.3
1970 383 pT 24.5 11.5
Totals 749 AMS) 23.1 7.9

% in area (see Figure 1)

D E F G Other
0.8 8.3 4.2 6.7 238}
8.9 16.7 5.7 1.6 26.4
7.0 10.7 1.6 6.5 15.4
6:7 12.3 3.3 4.9 20.3

“Excludes tail fans for which no latitude/longitude coordinates were available and those from north of 54°.

1981

TABLE 3—Types of Canada Geese in the hunter harvest by
area in southern Manitoba, 1965-1971

Nae Percent
Location® examined Giant” Mid-size’ Small”
Area A°
Marshy Point 100 67 30 3
Dog Lake 51 92 8 0
Fisher Branch —

Hodgson -—

Sleeve Lake 89 53 36 11
Total area A 240 67.1 DY) 5.4
Area B° 220 1.8 80.5 Vea
Area C

Alf Hole 34 77 18 6

Red River 20 35 25 40
Total area C 54 61 20 19
Area D® 23 17 4 78
Area E° 27 4 11 85
Area F 56 34 21 45
Area G 92 72 28 0
“See Figure |.
See text.

“1968, 1969, and 1970.

“Almost all from West Shoal Lake in 1965, 1966, and 1967.
“Seventeen from Whitewater Lake, 8 from Pilot Mound,
and 2 from Baldur.

Population (EPP) of B. c. interior have stopped in
southern Manitoba during autumn. They tradition-
ally stopped in spring when this area, as well as
another location (Grants Lake vicinity) 19 km
northwest of Winnipeg, were flooded by snow melt in
early-mid April. Upwards of 70 000 geese were esti-
mated to use these areas in spring 1969 and 1970, and
their identity as EPP geese was established by obser-
vation of several geese neck-banded at Swan Lake,
Missouri, the main wintering site for this population
which nests in northern Manitoba (Vaught and Kirsch
1966). The restoration of Oak Hammock as a year-
round marsh led immediately to its use in autumn by
this population.

The varying proportions of different subspecies,
and changes in numbers and distribution as habitat
and population levels change, present problems and
opportunities for management. The subspecies origi-
nate from totally different breeding habitats, and fac-
tors affecting their reproductive success and popula-
tion size will vary independently. Therefore, surveys
undertaken to assess age composition as reflected in
the kill of geese by hunters must take into account the
races involved. Regulations designed to meet goals
established for one population might have entirely
unintended effects on other populations. Knowledge
of the location of geese of different origins presents the
Opportunity to be flexible in recommendations for

RAVELING AND DIXON: CANADA GEESE, MANITOBA

279

harvest or habitat management specific to different
populations. Sampling programs need to be continu-
ally reassessed as changes occur. Combining results
just as Canada Geese, or just as large and small Can-
ada Geese, can obscure recognition of the dynamics of
these independent populations, of which small units
may be particularly vulnerable to heavy harvest (cf.
Raveling 1978).

Acknowledgments

We are grateful to numerous individuals who
assisted in surveys, facilitated the collection of geese,
or provided data from the national parts survey, espe-
cially L. Bidlake, E. Bossenmaier, D. Davies, R.
Robertson, R. Urban, R. Webb, and many other
regional staff of the DNR; G. Adams, G. Cooch, A.
Dzubin, R. Halladay, and R. Hutchinson of CWS; H.
A. Hochbaum and P. Ward of the Delta Waterfowl
Research Station; L. King and the P. Curry and A.
Vincent families of East Meadows Ranch at Clark-
leigh; B. Frenchand V. Jeffries of Rennie; R. Fortin of
Clarkleigh; R. Otto and family of Ashern; C. Obs-
zarski of Fisher Branch; S. Macks of Plumas; and
numerous other sportsmen.

Literature Cited

Canadian Wildlife Service. 1968. Species of waterfowl
killed in Canada during the 1967-68 hunting season. Can-
adian Wildlife Service Progress Notes Number 7. 22 pp.

Delacour, J. 1954. The waterfowl of the world. Vol. 1.
Country Life, Limited, London. 284 pp.

Gulden, N. A., and L. L. Johnson. 1968. History, behav-
ior, and management of a flock of giant Canada Geese in
southeastern Minnesota. /n Canada Goose management.
Edited by R. L. Hine and C. Shoenfeld. Dembar Educa-
tional Research Services Incorporated, Madison, Wiscon-
sin. pp. 59-71.

Hanson, H.C. 1951. A morphometrical study of the Can-
ada Goose (Branta canadensis interior Todd). Auk 68:
164-173.

Hanson, H.C. 1965. The giant Canada Goose. Southern
Illinois University Press, Carbondale. 226 pp.

Hanson, H. C., and R. H. Smith. 1950. Canada Geese of
the Mississippi Flyway: with special reference to an Illinois
flock. Illinois Natural History Survey Bulletin 25: 67-210.

MaclInnes, C. D. 1966. Population behavior of eastern arc-
tic Canada Geese. Journal of Wildlife Management 30:
536-553.

Malecki, R. A., D. Caswell, K. M. Babcock, R. A. Bishop,
and R. K. Brace. 1980. Mayor nesting range of the East-
ern Prairie Population of Canada Geese. Journal of Wild-
life Management 44: 229-232.

Oetting, R. B.,andC. C. Dixon. 1975. Waterfowl nest den-
sities and success at Oak Hammock Marsh. Wildlife
Society Bulletin 3: 166-171.

Raveling, D. G. 1976. Status of giant Canada Geese nesting
in southeast Manitoba. Journal of Wildlife Management
40: 214-226.

280

Raveling, D. G. 1977. Canada Geese of the Churchill River
basin in north-central Manitoba. Journal of Wildlife
Management 41: 35-47.

Raveling, D. G. 1978. Dynamics of distribution of Canada
Geese in winter. North American Wildlife and Natural
Resources Conference Transactions 43: 206-225.

Vaught, R. W., and G. C. Arthur. 1965. Migration routes
and mortality rates of Canada Geese banded in the Hud-

THE CANADIAN FIELD-NATURALIST

Vol. 95

son Bay lowlands. Journal of Wildlife Management 29:
244-252.

Vaught, R. W., and L. M. Kirsch. 1966. Canada Geese of
the eastern prairie population, with special reference to the
Swan Lake flock. Missouri Department of Conservation,
Technical Bulletin Number 3. 91 pp.

Received 9 May 1980
Accepted 21 February 1981

Hill’s Oak (Quercus ellipsoidalis) in Southern Ontario

P. W. BALL

Department of Botany, Erindale College, University of Toronto, Mississauga, Ontario LSL 1C6

Ball, P. W. 1981. Hill’s Oak (Quercus ellipsoidalis) in southern Ontario. Canadian Field-Naturalist 95(3): 281-286.

Hill’s Oak (Quercus ellipsoidalis) is reported from several localities in southern Ontario near Galt and Brantford. This
represents an eastward extension of the range of the species of over 200 km. In most localities in this area the trees occur in
fence rows and hedgerows, and with the increasing urbanization and road widening that is occurring, many are in jeopardy.
The distinction between Q. ellipsoidalis, Q. coccinea (Scarlet Oak), and Q. velutina (Black Oak) was made from leaf, bud, and

acorn characteristics.

Key Words: Quercus ellipsoidalis, southern Ontario, distribution.

The occurrence of Hill’s Oak (Quercus ellipsoidalis)
in Canada in the Rainy River district of Ontario at the
Northwestern edge of the range of the species has
recently been reported by Maycock et al. (1980) (see
Figure | for the known North American distribution).
Hill’s Oak was first recorded from Canada by Mitchell
(1912) from Point Edward, Sarnia, Lambton County,
Ontario. This record is supported by a specimen, for-
merly in the Dearness Herbarium, now in the herba-
rium of the Biosystematics Research Institute, Ottawa
(DAO); it consists of some acorns and acorn cups and
a few detached leaves. Whereas this material can be
assigned to Q. ellipsoidalis, it could equally well be
assigned to Q. velutina (Black Oak). A more complete
specimen collected by Dodge and Tripp in 1915,
which, according to a letter accompanying the DAO
specimen, was from the same tree as that seen by
Mitchell, is in the herbarium of the University of
Michigan, Ann Arbor(MICH). As noted by Maycock
et al. (1980) this specimen is undoubtedly Q. velutina,
but the foliage is so different from the DAO specimen
that it is doubtful if it did come from the same tree.
Maycock et al. (1980) also cited a specimen collected
by Macoun from the same locality that could also be
Q. ellipsoidalis, but this again 1s incomplete so cannot
be determined with certainty.

While examining the material of Q. ve/utina in the
University of Toronto herbarium (TRT), I noticed a
specimen, consisting of twigs with well-developed
terminal buds, mature acorns, and sun-foliage, col-
lected in 1957 by Tamsalu from just west of Galt, that
appeared to be markedly different from most speci-
mens. It had been determined by Tamsalu as Q. cocci-
nea (Scarlet Oak), but had subsequently been rede-
termined as Q. velutina. The records of Q. coccinea
from southern Ontario were thoroughly investigated
by Fox and Soper (1954), who concluded that these
should be referred to other species (primarily Q.
palustris, (Pin Oak) and Q. velutina). Investigation in
the field during the summer of 1978 led to the discov-

ery of similar trees scattered through North Dumfries
Township in the Regional Municipality of Waterloo,
and in the northern part of South Dumfries Township
in Brant County. Three more localities have been
found south of Brantford.

Identification of the Brant and Waterloo Oaks
The identity of these trees presented a number of
problems. They clearly fell within a group of species
centered on Q. velutina and Q. palustris. This group of
species was the subject of a recent study by Jensen
(1977a, b), who was able to demonstrate that Q. rubra
(Red Oak), Q. velutina (Black Oak), Q. shumardii
(Shumard’s Red Oak), Q. nuttallii (Nuttall’s Oak), Q.
coccinea (Scarlet Oak), Q. ellipsoidalis (Hills Oak),
and Q. palustris (Pin Oak) were all reasonably distinct
species. It quickly became apparent that four of the
species discussed by Jensen were quite distinct from
the Waterloo—Brant trees and could be eliminated as
possibilities. These were Q. palustris, which has small
very flat acorn cups and small acorns, and Q. rubra,
Q. shumardii, and Q. nuttallii which have very large
flat acorn cups and large acorns. The remaining three
species, Q. coccinea, Q. ellipsoidalis, and Q. velutina,
cannot be so readily distinguished. Typical Q. ve/utina
can be identified by the large usually deeply lobed
leaves; large, acute, distinctly angled, densely gray
pubescent terminal buds; and the distinct fringe of
scales on the acorn cup. However, Q. velutina is
extremely variable and individuals can be found with
the characteristic terminal bud, but with small leaves
and the acorn cup not or scarcely fringed. Quercus
coccinea can usually be identified by the presence of
several concentric rings at the tip of the acorn, but this
character is occasionally missing. Quercus ellipsoida-
lis has no unique characters, but it can be readily
identified by acombination of characters, particularly
the slender petiole; small, glabrous or sparsely pubes-
cent terminal buds; and the small, turbinate,
unfringed acorn cup. Table | lists some of the charac-

281

282 THE CANADIAN FIELD-NATURALIST Vol. 95
TABLE |—Characters distinguishing Quercus coccinea, Q. ellipsoidalis, and Q. velutina derived from Jensen (1977b)
compared with data from Ontario specimens of Q. velutina and Q. ellipsoidalis
coccinea ellipsoidalis velutina ellipsoidalis velutina
(Jensen) (Jensen) (Jensen) (Rainy River, Kenora) (Ontario)
Length of leaf
blade (mm) 111-160 60-110 (60-)(a)1 11-160 90-130 110-190
Width of leaf
blade (mm) 101-150 57-100 (57-)101-150 75-100(-115) 80-145
Petiole diameter
(mm) 0.97-1.44 0.68-0.96 0.97-1.44 0.8-1 (0.9-)1.0-1.4
Terminal bud
length (mm) (2.6-)5.7-9 2.6-5.6 5.7-9 3.0-4.3 (4.5-)5.5-7.5
Terminal bud
strongly angled — (rarely +) - + = +
Terminal bud Pubescent at Glabrous or pubes- Densely Pubescent Densely
indumentum apex cent at apex pubescent at apex pubescent
Acorn cup
width (mm) (9.8-)16-26.5 9.8-16 9.8-26.5 13-16 13-17
Acorn cup fringed - ~ + (rarely —) - + (rarely —)
Acorn cup: Glabrous or Sparsely
pubescence of pubescent Glabrous Pubescent pubescent Pubescent
inner surface at base all over all over all over
Acorn cup scale 2.26-3.50
length (mm) 3.51-5.15 (-5.15) 3.51-5.16 3.2-4.0 (3.0-)3.5-5
Acorn with
apical rings + (rarely —) - - = —

“Numbers in parenthesis are exceptional sizes, found in usually less than 10% of the sample.

ters which, according to the data published by Jensen
(1977b), distinguish these three species. A number of
other apparently diagnostic characters (e.g., twig
diameter) used by Jensen appeared on Ontario mate-
rial to be much too variable to be of any value in
determining individual specimens. Jensen’s data were
not presented in sucha way that they could be reliably
used to determine individual specimens nor was this
the purpose of his work. His data were obtained by
calculating means of a series of measurements taken
from selected trees. Therefore in some cases the
extremes will be exceeded by individual measure-
ments. Jensen also presented his data ina two-, three-,
or four-character state form, which may grossly over-
state the range of variation found in a particular spe-
cies. Despite these shortcomings, these data seem to
be useful for determining specimens. In addition to
Jensen’s data, Table | also includes, for comparison,
data derived from Ontario specimens of Q. ellipsoida-
lis (Rainy River and Kenora districts) and Q. velutina.
The data from Ontario Q. ellipsoidalis do not differ
significantly from that of Jensen, except that the
leaves are a little larger and the inner surface of the

acorn cup is sparsely pubescent all over, not glabrous.
The data from Ontario Q. velutina are also not
markedly different from Jensen’s data. The large dif-
ference in the figures obtained for acorn cup width 1s
almost certainly due to the way in which Jensen scaled
these data. If the Ontario data were scaled in the same
manner a similar range would be obtained.

Table 2 presents comparable data derived from
specimens collected from individual trees in Brant and
Waterloo counties. The Tamsalu specimens and Ball
numbers 78095, 78445, 79099, and 79445 agree well
with Q. ellipsoidalis from northwestern Ontario and
with Jensen’s data. The other specimens show combi-
nations of characters which make it impossible to
attribute them to a particular species. For example,
Ball 79436 most closely matches Q. coccinea, but the
acorns are without the apical rings and this individual
was growing adjacent to Ball 79445 (Q. ellipsoidalis)
and about 200 m from Ball 79451 (Q. velutina). In
view of the extensive evidence of hybridization
between Q. ellipsoidalis and Q. velutina presented by
Jensen (1977a) and others, it seems reasonable to
assume that these individuals are of hybrid origin.

1981

BALL: HILL’S OAK IN SOUTHERN ONTARIO

TABLE 2—Characters of Quercus specimens from Waterloo and Brant. (? character could not be observed)

Tamsalu
1S. VII1.1957 Ball Ball
(TRT) 78095 78444

Length of leaf

blade (mm) 100 105 120-135
Width of leaf

blade (mm) 80 100 115-120
Petiole diameter

(mm) 0.7 0.8 0.9
Terminal bud

length (mm) 3.0 355) 4.0
Terminal bud

strongly angled - - -
Terminal bud Pubescent Pubescent Pubescent

indumentum at apex at apex at apex
Acorn cup

width (mm) 15 15 12
Acorn cup fringed - - ~
Acorn cup: Sparsely Sparsely

pubescence of pubescent pubescent

inner surface all over ? all over
Acorn cup scale

length (mm) 4.0 4.0 3.0
Acorn with

apical rings = =

283
Ball Ball Ball Ball Ball
78445 79099 79436 79437 79445
95-100 105-125 100-150 90-135 90-115
80-85 110-130 95-125 70-110 90-110
0.8 0.8-1.0 0.9-1 0.7-0.8 0.6-0.8
4.5 4.0-5.2 3.8-4.8 3.5-4.0 3.5)
Pubescent
Pubescent Pubescent Pubescent almost Pubescent
at apex at apex at apex all over at apex
11 13-16 16-17 16-19 12
Glabrous Glabrous
or hairy or sparsely
a at base Glabrous pubescent ?
DES DES=3).5 3.5 3.5-4..5 3.0

Distribution and Ecology of Q. ellipsoidalis
in Southern Ontario

Investigations in the field and ina number of herba-
ria (DAO, TRT) have shown that Q. ellipsoidalis or
trees that are probably derived from @. e/lipsoidalis x
velutina hybrids are largely restricted to North
Dumfries township in the Regional Municipality of
Waterloo and to South Dumfries township in Brant
County. A few individuals have been found in two
other localities in Brant County adjacent to the Grand
River. Recently (1981) John Sewell has discovered a
roadside tree in northwestern Norfolk County about
12 km southwest of Brantford. The known distribu-
tion of these trees, except for the Norfolk County
locality, is shown in Figure 2a.

This population of Q. ellipsoidalis represents a
major eastward extension of the range of this species,
being over 200 km east of the nearest localities in
Michigan (Figure |). Whereas sucha large disjunction
is unusual, the area of Waterloo and Brant counties in
question is well known for the large number of
uncommon species which occur there, many with dis-
junct distributions. These include species of dry open
habitats such as Aureolaria virginica (False Fox-

gloves) and Swertia caroliniensis (Columbo), and spe-
cies of open wet habitats such as Carex lupuliformis (a
sedge), Eleocharis rostellata (Spike Rush), and many
others (Herriott 1905; Montgomery 1945; Soper
1962).

Little information can be given on the ecology of Q.
ellipsoidalis in Waterloo and Brant counties. Almost
all of the stations are in old fence rows or hedgerows.
In several localities the trees are growing very close to
extensive tracts of natural forest, but despite extensive
searches either none were found in these forests or
only a few trees were found at the edge. In a few
instances the trees were also found growing close to
ponds or small lakes which are frequent in this area,
and in one locality they appeared to have invaded an
abandoned field. The soil in almost all cases had a
high sand content and in several localities the trees
were associated with herbs which are generally
regarded as prairie elements, e.g. Andropogon gerar-
dii (Big Blue Stem), A. scoparius (Little Blue Stem),
Sorghastrum nutans (Indian Grass), and Lespedeza
capitata (Bush clover). One station is very close to the
only known locality of Bouteloua curtipendula in
Brant County (Montgomery 1956).

284

THE CANADIAN FIELD-NATURALIST

Vol. 95

Ficure |. North American distribution of Quercus ellipsoidalis (after Little 1971). Note disjunct localities in Michigan and

Ohio.

The historical geography of Dumfries township was
the subject of a detailed study by Wood (1958, 1961).
The township was first surveyed in 1816-1817 and
Wood was able to construct a map of the vegetation of
that period based on the surveyor’s notes. Extensive
areas in the southern and eastern halves of the town-
ship were described as “oak plains,” “plains,” or
“meadows.” An area (about 5X4km) west and
southwest of Galt was covered by “thickets” consist-
ing of scrubby individuals of pine, oak, poplar, and
hazel. Also mentioned were “briars,” “white thorn,”
“plum bushes,” and “cherry.” Such “thickets” suggest
an area under recolonization following a relatively
recent major disturbance. At the northern end of the
“thickets” an area of about 4 X | km was described as
recently burnt, thus suggesting that the “thickets”
were the result of an earlier more extensive fire. The
approximate area covered by these vegetation types is
shown in Figure 2b. Another brief description of the
vegetation south of Galt was provided by Fergusson
(1833). He described the area (pp. 130-131) as“... an
extensive range of open, grove-like woodland, princi-
pally oak, and the trees so dispersed as to not interfere
materially with the plough. It had much of the
appearance of some of the wildest part of English
park-scenery . . . It was a lovely landscape, with a
greater range open to the eye than usually occurs in
the interior of Canada.” These early 19th century

descriptions of Dumfries township suggest that much
of it consisted of extensive areas of dry grassland with
numerous open-grown oak trees which did not forma
continuous canopy and which was subject to distur-
bance by fire. Such a description conforms well with
modern accounts of the ecology of Q. ellipsoidalis in
Wisconsin and northwestern Ontario (Curtis 1959;
Maycock et al. 1980). A comparison of Figure 2a and
b shows that there is a good correlation between the
present-day distribution of Q. ellipsoidalis in Dum-
fries township and the areas of vegetation that were
suitable for it in 1816-1817. Many of the apparently
suitable areas from which the species is now missing
(e.g. east of Paris) have been subject to total destruc-
tion by quarrying in recent times or by urbanization.

The absence of this species from other areas of
southwestern Ontario is a little puzzling and may be
due to oversight. Nevertheless, the absence of speci-
mens from these areas suggests that, if it does occur
there, it must be extremely rare. It should be noted
that this disjunct population of Hill’s Oak is in serious
jeopardy. Most of the localities mapped consist of less
than five trees often in a poor state of health. None
occur along roads which have been improved in recent
years (e.g. Highways 24, 24A, and the new Highway
97) and the locality 2 km south of Galt is ona dirt road
which is in the process of being widened and improved
with virtually all hedgerow trees being destroyed. The

1981

nar
WATERLOO hs G / A

97

ae

St George
|

4 7

/
Brantford __
ITI JIT fy

/ / / }
Le fy, // ly
J / / /)
HSL ff. /. Hopf ihe\)
Lap / /

BALL: HILL’S OAK IN SOUTHERN ONTARIO

a
| / [77 |
| WATERLOO

WITS 1/47
| YL INS /
| / ‘/ Sue /
| Vi, OL LA
| t/ // A, ESTA
| ee : Y/// .
| 4
|
| ;
/

| Y Se

\ Brantford =

FiGuRE2.(a) Distribution of Quercus ellipsoidalis in Brant County and Waterloo County (now the Regional Municipality of

- Waterloo). One locality in Norfolk County not mapped. Herbarium specimens, ®. Sight records,°. County boundaries

indicated by broken line. Ontario highways indicated by unbroken line with number. Location of insert indicated by
aSterisk.

2(b) Occurrence of “plains,” “

oak plains,” “thickets,” and “burnt areas” in Dumfries township as noted in 1816-1817

(after Wood 1958, 1961). Plains and oak plains — boundary marked by straight lines. Thickets and burnt areas —
boundary marked by crosses.

largest group of trees (probably over 100) seen is just
south of Blue Lake, 5 km west of St. George. These
occur along the road, in hedgerows and in an old field
which has been partially subdivided into large house
lots. A few individuals occur in Pinehurst Conserva-
tion area, 8 km north of Paris, and on the edge of
forest tracts being managed by the Ontario Ministry
of Natural Resources. Unfortunately the normal
methods of forest management employed in these

areas are not likely to encourage reproduction of Q.

ellipsoidalis. The species undoubtedly requires open,
somewhat disturbed, habitats to reproduce success-
fully, and it may also be encouraged by fire (Curtis
1959).

The following is a list of specimens of Q. ellipsoida-
lis and Q. ellipsoidalis X velutina on which Table 2
and Figure 2 are based. Also listed are some addi-
tional localities for Q. ellipsoidalis in northwestern
Ontario which were utilized in compiling Table | and
which were not listed by Maycock et al. (1980).

286

Specimens of Q. ellipsoidalis and
Q. ellipsoidalis X velutina
Kenora District

S. shore of Kakagi Lake, 2 miles (3 km) south of Blacky
Bay. 49°10’N. 93°49’W. 8.VIII.1957. P.A. Bentley 57535
(DAO, TRT) (sub Q. rubra).

Rainy River District

Quetico Provincial Park. Prairie Portage at Sucker Lake.
24.V11.1957. Garton 4910 (DAO, TRT) (sub Q. rubra).
Wild Land Reserve. 5 miles (8 km) below town on Rainy
River. 7. VII.1961. Garton 8861 (DAO, TRT) (sub Q. rubra).

Brant County

On the Grand River between Caledonia and Brantford, near
Middleport. 1970. Gilmour (DAO). Onondaga twp. 1/2 km
west of Middleport. 43°06’N. 80°0S’W. Large tree at edge of
roadside and cornfield. 28.VI.1979. Ball 79099 (TRTE).
Brantford twp. 4km south of Brantford in the Oxbow.
43°07'N. 80°14’W. Hedgerow-roadside trees. 28.VI.1979.
Ball 79088 (TRTE); 20.1X.1979. Ball 79436, Ball 79437, Ball
79445 (TRTE). South Dumfries twp. Spottiswood Lake.
17.VIII.1957. Dore & Montgomery 16850 (DAO). South
Dumfries twp. East side of Spottiswood Lake. 43°15’N.
80°22’W. Edge of open area dominated by Sorghastrum
nutans, Andropogon scoparius, Lespedeza capitata, Aster
azureus, Solidago nemoralis. 30.V111.1978. Ball (and May-
cock) 78381 (TRTE). South Dumfries twp. c. | km west of
junction of Highways 5 and 24 along road to Blue Lake.
43°14’N. 80°18’W. Roadside tree. 30.VI.1978. Ball 78110
(TRTE). South Dumfries twp. 2 km west of junction of
Highways 5 and 24. 43°14’N. 80°19’W. Roadside-hedgerow
trees. 26.VIII.1980. Ball 80169, 80171, 80172, 80173,
(TRTE). South Dumfries twp. South of Blue Lake.
43°13’30’N. 80°20’W. Numerous roadside-hedgerow trees.
26. VIII.1980. Ball 80178, 80179 (TRTE). South Dumfries
twp.c. | km east of Blue Lake. 43°14’N. 80°20’ W. Roadside-
hedgerow tree. Ball 78111 (TRTE). South Dumfries twp.
County Line Road c. 200 m east of Hwy. 24A. 43°15’/N.
80°23’W. Roadside-fencerow tree. 26.V1.1978. Ball 78095
(TRTE). South Dumfries twp. County Line Road c. | km
west of Hwy. 24A. 43°17’N. 80°24’W. Roadside at edge of
small lake, associated with Andropogon scoparius, A. gerar-
dii, Lespedeza capitata. 26.VI111.1980. Ball 80191, 80192
(GRSHE):

Regional Municipality of Waterloo, North Dumfries twp.
Roadside 1-1/2 (2 km) west of Galt. 15.VIII.1957. A.
Tamsalu (TRT, HAM). c. 2 km west of Galt along regional
road 46 (formerly Hwy. 97). 43°22’N. 80°22’W. Roadside
tree. 21.VI.1978. Ball 78048 (TRTE). 5 km south of Galt on
road running west from the Grand River. 43°19’N. 80°19’W.
Roadside-fencerow tree. 23.VI.1979. Ball 78065 (TRTE).
6 km south west of Galt, 3 km west of Grand River close to
Waterloo forest, Sudden tract. Hedgerow-roadside trees.
43°18’N. 80°21’W. 23.VI.1978. Ball 78064A, B (TRTE).
1 km south of Galt, | km east of the Grand River. 43°20/N.
80°17’W. Hedgerow-roadside trees. 6.1X.1979. Ball 79400,
79401 (TRTE). 1-1/2 km WSW of Branchton, on County
Line Road. 43°18’N. 80°16’W. Roadside-hedgerow tree.
26.V1.1978. Ball 78096 (TRTE). Concession VII, lot 31, near
Ayr. 6.1X.1941. Montgomery 849 (DAO). 2 km E. of Ayr.

THE CANADIAN FIELD-NATURALIST

Vol. 95

43°18’N. 80°26’W. Hedgerow-roadside trees. 5.1X.1978. Ball
78444, 78445 (TRTE).

Regional Municipality of Haldimand-—Norfolk

Windham twp. 2-1/2km SW of Scotland. 43°00'N.
80°24’W. Roadside tree. 22.11.1981. J. Sewell (TRTE).

Acknowledgments

My thanks to Craig Campbell for his helpful com-
ments and suggestions, William Crins for checking
specimens ina number of herbaria, and to the curators
of the herbaria cited. This work was financed by Natu-
ral Sciences and Engineering Research Council grant
No. A6494.

Literature Cited

Curtis, J. T. 1959. The vegetation of Wisconsin. University
of Wisconsin Press, Madison. 657 pp.

Fergusson, A. 1833. Practical notes made during a tour in
Canada and a portion of the United States in 1831. Black-
wood, Edinburgh. XV and 379 pp.

Fox, W.G., and J. H. Soper. 1962. The distribution of
some trees and shrubs of the Carolinian zone of southern
Ontario, part III. Transactions of the Royal Canadian
Institute 30: 99-120.

Herriott, W. 1905. Some new or little known Canadian
plants. Ontario Natural Science Bulletin 1: 26-30.

Jensen, R. J. 1977a. A preliminary numerical analysis of
Red Oak complex in Michigan and Wisconsin. Taxon 26:
399-407.

Jensen, R. J. 1977b. Numerical analysis of the Scarlet Oak
complex (Quercus subgen. Erythrobalanus) in the eastern
United States: relationships above the species level. Sys-
tematic Botany 22: 122-133.

Little, E. L., Jr. 1971. Atlas of United States trees, vol. 1.
Conifers and important hardwoods. United States
Department of Agriculture, Forest Service. Miscellaneous
publication Number 1146. Washington, D.C. Approx.
200 pp.

Maycock, P.F., D.R. Gregory, and A.A. Reznicek.

1980. Hill’s Oak (Quercus ellipsoidalis) in Canada. Cana-

dian Field Naturalist 94: 277-285.

Mitchell, F. 1912. A note. Ontario Natural Science Bulletin

TOM

Montgomery, F. H. 1945. A botanical survey of Waterloo

County, Ontario. Transactions of the Royal Canadian

Institute 25: 217-265.

Montgomery, F. H. 1956. The introduced plants of Onta-
r1o growing outside of cultivation. Transactions of the
Royal Canadian Institute 31: 90-102.

Soper, J. H. 1962. Some genera of restricted range in the
Carolinian flora of Canada. Transactions of the Royal
Canadian Institute 34: 1-56.

Wood, J.D. 1958. The historical geography of Dumfries
township, Upper Canada. M.A. thesis, University of
Toronto, Toronto, Ontario. 142 pp.

Wood, J.D. 1961. The woodland-oak plains transition
zone in the settlement of western Upper Canada. Cana-
dian Geographer 5(1): 43-57.

Received 12 November 1980
Accepted 6 March 1981

Seasonal and Daily Use of Plant Communities by Sharp-tailed Grouse
(Pedioecetes phasianellus) in the parklands of Alberta

Dc. MOYLES
Brooks Wildlife Centre, Bag 1540, Brooks, Alberta, T0J 0JO

Moyles, D. L. J. 1981. Seasonal and daily use of plant communities by Sharp-tailed Grouse ( Pedioecetes phasianellus) in
the parklands of Alberta. Canadian Field-Naturalist 95(3): 287-291.

Sharp-tailed Grouse ( Pedioecetes phasianellus) showed specific preferences for different plant communities at different times
of the year. Grassland and grassland — low shrub transition zones were selected throughout the year, with trees being used
primarily in winter and spring. Grouse were recorded using marsh vegetation only during winter. Except 1n winter and spring
when Sharp-tailed Grouse were seen in trees, they generally used open cover during early morning, moving into taller
vegetation by midday. In the afternoon and evening they moved back into more open vegetation. In summer, females with
broods selected grassland and grassland — low shrub transition zones throughout the day while males and females without
broods selected taller vegetation. The extent of Trembling Aspen (Populus tremuloides) cover around an arena was inversely
correlated with the numbers of males recorded on it and to the total number of grouse recorded within 0.8 km of the arena.

Key Words: Sharp-tailed Grouse (Pedioecetes phasianellus), parkland, habitat use, Alberta.

Sharp-tailed Grouse (Pedioecetes phasianellus)
occupy a variety of habitats in North America, from
open grasslands and sage brush through parklands to
early successional stages and open portions of mixed
woods and boreal forests (Aldrich 1963). They have
also adapted to areas where agriculture has created a
mosaic of native vegetation and cultivated lands that
provides food and cover (Evans 1968; Pepper 1972).
Descriptions of vegetation used in spring and in
summer, particularily by females with broods
(Hamerstrom 1963; Artmann 1970; Pepper 1972), and
during winter (G. A. Ammann. 1957. The prairie
grouse of Michigan, Game Division, Department of
Conservation, Lansing, Michigan; Marshall and
Jensen 1937, cited in Johnsgard 1973) are available,
but Evans (1968) noted that a proper understanding of
the habitat requirements of sharptails throughout the
year is lacking. In this paper I report preferences
shown seasonally and daily by Sharp-tailed Grouse
for certain plant communities in aspen parkland of
east-central Alberta.

Study Area

The study area was located in Camp Wainwright
Military Reserve (54°48’N; 110°55’W), 12 km
southwest of Wainwright, Alberta. The area (28 km?)
included grassland, grassland - low shrub transition
zones, patches of low shrubs, tall shrubs, and low
shrub — tall shrub transition zones, copses of trees,
and wetlands.

The grasslands (19.2%) were dominated by Blue Grama
Grass (Bouteloua gracilis) and Speargrass (Stipa comata),
with Brome (Bromus inermis) and fescue (Festuca spp.) also
in appreciable amounts. Predominate forbs in the grasslands
and grassland-shrub transition areas were Pasture Sage

(Artemisia frigida) and Prairie Sage (A. gnaphalodes), asters
(Aster spp.), Yarrow (Achillea lanulosa), Golden Aster
(Chrysopis villosa), goldenrod (Solidago spp.), and Wild
Strawberry (Fragaria glauca). Horsetail (Equisetum spp.)
was also common in these areas. Patches of Bearberry (Arc-
tostaphylos uva-ursi) were found in south-facing slopes. Less
commonly, forbs such as Golden Bean (Thermopsis rhombi-
golia), milk-vetches (Astragalus spp.), Purple Prairie Clover
(Petalostemon purpureum), Gaillardia (Gaillardia aristata),
vetches (Vicia spp.), Silverweed ( Potentilla anserina), \hree-
flowered Avens (Geum triflorum), and locoweed (Oxytropis
spp.) were present.

In the ecotone between aspen copses and grassland, shrubs
such as Chokecherry ( Prunus virginiana), Saskatoon (Ame-
lanchier alnifolia), Wolf Willow (Elaeagnus commutata),
roses (Rosa acicularis and R. woodsii), and snowberry
(Symphoricarpus albus and S. occidentalis) were often grow-
ing in a scattered pattern and intermixed with grasses and
forbs, such as sages, goldenrods, Yarrow, and Golden Aster.
This grassland — low shrub transition zone covered 25.5% of
the study area; 8.6% was covered by thick, discrete clumps of
low shrubs less than 1 min height, particularly Chokecherry,
Snowberry, Wolf Willow, and Saskatoon; these clumps var-
ied from 0.5 m?to 10 m2. Grasses dominated the understory
in these patches.

Low shrub - tall shrub transition zones (21.8%) were usu-
ally found on the edges of aspen copses and dominated by
Trembling Aspen (Populus tremuloides) saplings or Choke-
cherry bushes to 3 m in height. The understory was usually
dense small shrubs, usually Chokecherry and Saskatoon
intermixed with aspen saplings. Patches of 1 to 2 m high
Wolf Willow were usually open with grasses composing
almost all of the understory.

Aspen dominated all stands of tall shrubs (10.2%), some of
which were found in discrete clumps but most were located
beside trees. The understory was usually dense witha variety
of small shrubs including Chokecherry, Saskatoon, and
aspen. Northern Bedstraw (Galium boreale), Cream-

287

288

coloured Vetchling (Lathyrus ochroleucus), Harebell (Cam-
panula rotundifolia), and anemones (Anemone spp.) were
commonly intermixed with sparse grasses.

The aspen corpses (13.3%) varied from about 3 m? in area
up to about |1 ha; average size was about 0.7 ha. The under-
story beneath the mature trees was usually open with some
aspen, Chokecherry, or Saskatoon saplings, grasses, and
Vetchling and asters. These copses were intermixed with
patches of grassland and grassland-shrub transition zones
throughout the study area. The aspen was expanding into the
grassland community, probably as a result of no large-scale
burning for at least 12 years, and only limited grazing in
portions of the area between 15 May and 15 November each
year.

Wetlands occupied about 2% of the study area. These were
mesic areas, with little open water and occupied with sedges
(Carex spp.). Willows (Salix spp.) grew along the borders in
bands from | to 5 m thick. Balsam Poplar (P. balsamifera)
also grew along the borders of these wetlands.

The study area was undisturbed apart from military
activity which affected the area in two ways: (1) small
wildfires occurred sporadically, never exceeding 3 ha
during the study, and (2) extensive use of all-terrain
and tracked vehicles created trails and openings
through wooded and shrubby areas.

Methods

The study area was searched systematically with
dogs from mid-May 1975 to early January 1977. In
addition, seven male Sharp-tailed Grouse were fitted
with radio transmitters in autumn 1975, 12 males in
spring 1976, and 13 males inautumn and winter 1976.
Locations of all grouse encountered, both during
searches and by radiolocation, were noted before
flushing if possible, otherwise at the point of flushing.
Weather conditions (wind speed, temperature, cloud
cover, and presence or absence of moisture on the
vegetation) were noted at the time of each sighting.

The vegetation within a I-m radius of each location
of sighting was classified into one of seven classes
(Table 1). The distribution of these sightings was
compared with the availability of the same classes
within the study area.

The impact of weather on the type of vegetation
used by grouse was tested with a multiple regression
analysis, using a hierarchial approach (Nie et al.
1975).

To examine the impact of aspen forest expansion
on the use of display arenas, the area covered by aspen
stands within a radius of 800 m of nine arenas was
measured with a planimeter from aerial photographs
(1 cm = 190 m) taken in 1962 and 1975.

Results and Discussion

Based on all observations over the entire year,
Sharp-tailed Grouse were not distributed randomly
relative to available plant communities except for

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 1—Description of seven plant communities in the
Aspen Parkland of Alberta used by Sharp-tailed Grouse

No. Community

Description

l Grassland Dominated by grasses and
forbs
2 Grassland — low A mixture of grasses, forbs,

shrub transition and woody plants less than
1 m in height

Dominated by wood plants
less than | m in height

A mixture of woody plants less
than 3 m in height

Dominated by woody plants
greater than | m but less
than 3 m in height

3 Low shrub

4 Low shrub - tall
shrub transition
5 Tall shrub

6 Trees Dominated by woody plants
greater than 3 m in height
7 Marsh Dominated by willow and

sedge

grassland — low shrub transition and for tall shrubs
(Figure 1). They preferred grassland (chi-square test,
P< 0.05), low shrub and trees (P< 0.05), and
avoided low shrub — tall shrub(P < 0.05). Marsh was
used only in winter (P < 0:05). Considering the sight-
ings by season provided a somewhat different picture
(Figure 2). In spring (1 April to 15 June) grouse
selected tree cOmmunities, seemingly for cover and

40

30 B,n- 501

SIGHTINGS

OF

20

10

PERCENT

VEGETATIVE

TYPE

Ficure |. Selection of the plant communities (Table 1) by
Sharp-tailed Grouse, based on 2322 sightings in 1975
and 1976 (A) as compared to the relative coverage of
the plant communities (B).

1981

40 |
SPRING SUMMER
n-509 n-672
30
ps
20
wn 4
@ 10
z
=
3e
G)
Oo A 2
“ao
AUTUMN WINTER
n=-583 n- 558

PERCENT
wo
{o)
——
]

nN

oO

——
J

1 Dea e 4 5 6 7 1 6 7

VEGETATIVE TYPES

FiGuRE 2. Selection of various classes of vegetation(Table 1)
by Sharp-tailed Grouse during four seasons based on
2322 sightings in 1975 and 1976. Spring, | Aprilto 15
June; Summer, 16 June to 31 August; Autumn, |
September to 15 November; Winter, 16 November to
31 March.

food. Insummer (16 June to 31 August) they selected
primarily grassland and tall shrub communities, par-
ticularly tall shrub with sparse understory during
midday. In autumn (1 September.to 15 November)
about 70% of grouse sightings were in grassland or
grassland — low shrub transition communities. These
communities offered both food (insects and fruits)
and roosting cover for grouse. Grouse in winter (16
November to 31 March) used grassland and trees.
They fed on aspen buds and roosted in grassland and
in the lee of the trees. Marsh vegetation was used in
winter when sedges and willows provided cover and
food.

Sharp-tailed Grouse selected different plant com-
munities throughout the day (Figure 3). In spring,
grouse were observed feeding mainly in grass-
land - low shrub transition zone and trees during
early morning, remaining in the taller vegetation
throughout midday. They moved back into grass-
land — low shrub transition and grassland communi-

MOYLES: SHARP-TAILED GROUSE HABITAT USE, ALBERTA

289

ties after midday apparently seeking food and roost-
ing cover. During the summer the grouse apparently
fed in the grasslands during early morning, sometimes
moving into heavier cover, perhaps for shade and
protection from predators, during midday, and then
back into more open vegetative types, again probably
seeking feeding and roosting cover, in the evening
(Figure 3). Inautumn grouse used the open vegatative
communities in early morning, apparently for feeding,
occasionally moving into taller, heavier cover by
midmorning. From midday to dark, grouse selected
grassland — low shrub transition, apparently for food
and roosting cover. However, during winter, grouse
were often seen in the morning and midday perched in
aspen trees, often eating buds, then moving into more
open cover including (frozen) marsh during midday,
and roosting predominantly in the low cover in the
evening (Figure 3).

During summer females with broods used grassland
and grassland —-low shrub transition significantly
more than other adult grouse (Figure 4), including
both males and females without broods (G test,
P< 0.05). These observations agree with Hamer-
strom (1963) and Pepper(1972), who recorded broods
in similar habitats on study areas in Wisconsin and
Saskatchewan. Adult grouse without broods selected
taller vegetative types, particularly tall shrubs. The
heavier vegetation probably provided better cover
from avian predators as well as cooler microclimate
during the hottest part of the day. However, the rela-
tionship between weather and plant community
selected by the grouse was not strong. Only 3.7% of
the variation in cover chosen by grouse could be
explained by weather. Other factors such as food
requirements, presence of conspecifics, familiarity
with an area, and avoidance of predators may have
been of more importance in influencing choice of
vegetation by these grouse.

The relationship between the amount of continuous
aspen cover withina radius of 0.8 km of a display site,
or arena, of a group of male Sharp-tailed Grouse and
the number of these displaying males present on each
of nine arenas in 1976 (Figure 5) suggests that
numbers may be inversely related to the total coverage
of aspen. Data on numbers of sharptails attending
three arenas were available from 1968 to 1978.
Changes in aspen cover within a radius of 0.8 km of
these three arenas between 1962 and 1975 were com-
pared with the numbers present in 1968 and
1976-1978 (Table 2). The apparent inverse relation-
ship also suggests that with increasing percentage of
aspen cover fewer Sharp-tailed Grouse used the area.
Caldwell(1976) determined that arenas in aspen park-
land in central Manitoba ceased to be used by grouse
when the percentage area dominated by grasses within

290
SPRING SUMMER
60
DAWN 45 n=33 n=259
0800 BO
15
id)
O
z
[= n=215
ae
08:01 -
O
— 11:00
WY)
LL
Oo 66
n=98 n=131
be 11:01- 45
= 1500
Ww 30
O
oc 15
WwW
a
64
45 n=331 n=63
15:01 -
DARK 30
15
THis era OM ona 7, (iy we eS

VEGETATIVE

THE CANADIAN FIELD-NATURALIST

Vol. 95

AUTUMN WINTER
n=158 n=38
n=98 n=206

n=187
n=124

7 Tera Sh On os, 1

TYPE

23 45 67

FiGureE 3. Use of various plant communities (Table 1) by Sharp-tailed Grouse throughout the day during the four seasons.
(During winter the time periods used are dawn to 08:00; 08:01 to 11:00; 11:01 to 14:00; 14:00 to darkness; on account of

shorter daylight period at this season).

0.8 km radius of the arena fell to less than 58%.
Pepper (1972) reported positive correlations between
the number of summer observations and the amount
of upgrazed native grass-shrub and tame haylands
within 1.6 km of arenas and between attendance of
grouse on arenas during spring display and the
number of sharptails seen within 1.6 km of the arena
in summer. Thus a decrease in the number of display-
ing males also implies a decrease in the total
population.

In summary, I believe that Sharp-tailed Grouse
inhabiting the parklands of central Alberta have spe-
cific preferences for different plant communities at
different times of the year and of the day. This sug-
gests that a mosaic of plant communities, particularly
grasslands and grassland-shrub mixtures with exten-

sive ecotone, provides optimum habitat. Further-
more, expansion of the aspen community at the
expense of the grassland community reduces the rela-

TABLE 2—Relationship between increase in aspen cover
within a 0.8-km radius of three arenas between 1962 and 1975
and decline in numbers of males displaying on these arenas
from 1968 to 1978

No. of displaying males

Increase in
Arena aspencover(%) 1968 1976 1977 1978
I 16 18 5 (28) 1(6) 0
2 14 30 13 (43) 11 (37) 10 (33)
3 31 31 7 (23) 0 0

“Percentage of males in 1968.

1981

25

Fa - [ Jan=2322
y [48

20

PERCENT OF SIGHTINGS

™
|

>.
os he coo

1 2 3 4 5

6
VEGETATIVE TYPES

FiGureE 4. Use of various plant communities (Table 1) by
female Sharp-tailed Grouse with broods (A) and by
broodless adults (males and females) (B) during the
summers of 1975 and 1976.

25 e
0 Y=26.9-0.46X
r=-0.69

NUMBER OF LEKKING MALES

10 20 30 40 50

ASPEN COVER (PERCENT OF MAXIMUM)

FiGURE 5. Relationship between area of aspen cover
(expressed as a percentage of the total) within a circle
of radius 0.8 km from an arena and the number of
displaying male Sharp-tailed Grouse on the arena.

MOYLES: SHARP-TAILED GROUSE HABITAT USE, ALBERTA

25K

tive availability of the various vegetative types, and as
a consequence seems to reduce the utility of the area to
grouse as suggested by lower numbers on arenas.

Acknowledgments

I thank D. A. Boag for his assistance during the
field work and for comments on this manuscript. Two
anonymous referees also made valuable comments on
earlier drafts. D. Alton, P. Harris, L. Ramsay, K.
Smyth, and D. Wade assisted in the field at various
times. Major R. Bridgeman allowed me to work in
Camp Wainwright and R. Fyfe of Canadian Wildlife
Service kindly allowed me the use of the Canadian
Wildlife Service facilities at Wainwright. I received
financial support from the Alberta Fish and Wildlife
Division, a National Research Council of Canada
Postgraduate Scholarship, and an operating grant
(2010) to D. A. Boag. K. Smyth drew the figures.

Literature Cited

Aldrich, J. W. 1963. Geographical orientation of American
Tetraonidae. Journal of Wildlife Management 27:
529-545.

Artmann, J. W. 1970. Spring and summer ecology of the
Sharptail Grouse. Ph.D. Dissertation, University of Min-
nesota. Xerox University Microfilms, Ann Arbor,
Michigan.

Caldwell, P. J. 1976. Energetic and population considera-
tion of Sharp-tailed Grouse in the aspen parkland of Can-
ada. Ph.D. Dissertation, Kansas State University, Man-
hattan, Kansas.

Evans, K.E. 1968. Characteristics and habitat require-
ments of the Greater Prairie Chicken and Sharp-tailed
Grouse — a review of the literature. U.S. Department of
Agriculture Forest Service Conservation Research Report
Number 12.

Hamerstrom, F.N., Jr. 1963. Sharptail brood habitat in
Wisconsin’s northern pine barrens. Journal of Wildlife
Management 27: 793-802.

Johnsgard, P. A. 1973. Grouse and quails in North Amer-
ica. University of Nebraska Press, Lincoln, Nebraska.
Nie, N. H., C. H. Hull, J. G. Jenkins, K. Steinbrenner, and
D. H.Bent. 1975. Statistical package for the social scien-

ces. McGraw-Hill Book Co., New York.

Pepper, G. W. 1972. The ecology of Sharp-tailed Grouse
during spring and summer in the aspen parklands of Sas-
katchewan. Saskatchewan Department of Natural
Resources, Wildlife Report Number 1. Regina,
Saskatchewan.

Received 30 November 1978
Accepted | March 1981

Movements and Haulout Behavior of Radio-tagged Harbor Seals,

Phoca vitulina

KENNETH W. PITCHER and DENNIS C. McALLISTER

Alaska Department of Fish and Game, 333 Raspberry Road, Anchorage, Alaska 99502

Pitcher, Kenneth W., and Dennis C. McAllister. 1981. Movements and haulout behavior of radio-tagged Harbor Seals,
Phoca vitulina. Canadian Field-Naturalist 95(3): 292-297.

Movements, haulout area fidelity, and haulout frequency of Harbor Seals (Phoca vitulina) were studied in the Kodiak Island
area, Alaska, by relocating radio-tagged animals captured on a large hauling area. Eight of 35 radio-tagged seals were found
on hauling areas other than the capture site. The longest movement was 194 km and one seal crossed 74 km of open ocean.
Movement rates up to 27 km/d were recorded. There appeared to be considerable fidelity to one or two specific haulout
locations by individual radio-tagged animals. Resident, radio-tagged seals of a large hauling area were hauled out during 50%
of the daily radio checks in June and 41% from | August to 5 September. On an individual basis, frequency of haulout ranged

from 16 to 80% of the days.

Key Words: Harbor Seals, Phoca vitulina; Gulf of Alaska, radio-tagged, movements, haulout behavior.

In common with other pinnipeds, Harbor Seals
(Phoca vitulina) spend much of their lives in the
ocean. At times they haul out of the water onto reefs,
beaches, ice, or other suitable substrata where they
rest, give birth, and suckle their young. At this time
they can be most easily and accurately counted. Sev-
eral workers have attempted to census Harbor Seal
populations by counting hauled-out animals (Ros-
enthal 1968; Pearson and Verts 1970; Calambokidis et
al. 1979). However, because of insufficient informa-
tion regarding haulout area fidelity and the propor-
tions of populations visible, such counts served only
as minimum estimates. A recent study suggests that on
the average Harbor Seals spend about 44% of their
daily activity budget hauled out (Sullivan 1979). Most
published information, based primarily on repetitive
observations of recognizable individuals, suggests
that at least some animals return repeatedly to the
same hauling area (Boulva and McLaren 1979;
Calambokidis et al. 1978; Reijnders 1976; Knudtson
1974).

Harbor Seals are considered to be relatively seden-
tary animals with local movements associated with
tides, food, reproduction, and season (Scheffer and
Slipp 1944; Fisher 1952; Bigg 1969). These impres-
sions were the results of general observations rather
than records of individual movements. The only scien-
tific information of which we are aware on movements
of individual seals is from recoveries of animals tagged
as pups (Divinyi 1973; Bonner and Whitthames 1974;
Boulva and McLaren 1979).

In this study we used radio-tracking techniques to
monitor movements and haulout behavior of individ-
ual Harbor Seals thereby collecting new and more

comprehensive information on the range of move-
ments, movement rates, haulout area fidelity, and
frequency of haulout of Harbor Seals.

Methods

This study was conducted in the Kodiak Island area
of the Gulf of Alaska (Figure 1). The primary study
site was the southwestern hauling area on Tugidak
Island (56° 27’N, 154° 47’W) where up to 9000 Harbor
Seals have been counted on a 3.2-km stretch of gravel
beach. Field work was conducted between 8 May and
9 September 1978. Thirty-five seals (including 24
mature females, 5 immature females, 5 mature males,
and | immature male) were captured on the south-
western hauling area, immobilized with Ketamine
hydrochloride (Ketelar; Parke, Davis and Company),
and fitted with radio transmitters attached with a
bracelet around the base of a hind flipper. The first 21
seals were captured between 8 May and 2 June. Cap-
ture operations were then suspended to avoid distur-
bance during pupping. Fourteen more seals were fit-
ted with transmitters from 3 to 9 July. The
transmitters, fabricated by Cedar Creek Bioelectron-
ics Laboratory of the University of Minnesota, oper-
ated on separate frequencies in the 164 to 165-MHz
range. Signals could be received only when the seals
were hauled out. Maximum range of the transmitters
was about 8 km. Radio-tagged seals were also marked
with individually recognizable color combinations of
cattle ear tags placed in hind flippers and vinyl flag-
ging attached to the transmitters to serve as backup
identifiers to detect transmitter failure or loss.

Both numbers of radio-tagged seals and total
numbers of seals hauled out on the southwestern haul-

292

1981

40 km
20 MILES

Sees Ss
ree POR
CAPTURE SITE

f SEMIDI
¢,!SLANDS

TR-5

CHIRIKOF f 3
Stang, fa

PITCHER AND MCALLISTER: HARBOR SEAL MOVEMENTS 293

“KODIAK Sage
ISLAND .g4rttray 5
a UGAK

ibe

TR-18

f, TR-13
SS re 4

SITKINAK
ISLAND

>

Figure |. Gulf of Alaska study area showing locations of radio-tagged (TR) Harbor Seals found at haulouts other than the

capture site on southwestern Tugidak Island.

ing area were monitored on a near daily basis during
June and from | August to 5 September. Additional
observations of radio-tagged seals were made from
9-31 May, but behavior of the animals may have been
affected by the disturbance of ongoing capture opera-
tions. Radio checks and counts of seals on the
southwestern hauling area were made from the top of
30-m bluffs abutting the beach. A portable radio-
tracking receiver with hand-held Yagi antenna was
used for the onsite radio checks. Seals were directly
counted from the bluffs or from polaroid prints taken
from the bluffs. Hauled-out seals were examined with
10X binoculars to locate radio-tagged individuals and
the results compared with radio checks to detect radio

failures or losses. Radio checks and counts were timed
to coincide with daytime low tides when maximum
numbers were usually hauled out.

Aerial, radio-tracking surveys, using a scanning
receiver, were flown in a Bellanca Scout fixed-wing
aircraft or a Bell 206 helicopter. Twenty-seven sur-
veys, totaling 53.6 h, were flown to locate radio-
tagged seals that moved from southwestern Tugidak
to other hauling areas. Coverage of the surveys
included most of the shoreline and all of the known,
major haulouts in the Kodiak Island group. Chirikof
Island and the Semidi Islands were surveyed twice.
The coast between Wide Bay and Amalik Bay on the
Alaska Peninsula was surveyed once. Surveys were

294

flown during lower stages of the tide when the most
seals were usually hauled out.

Results and Discussion
Movements

Eight radio-tagged seals (six females and two
males) were relocated 17 times at hauling areas other
than the capture site. The longest movement was a
minimum of 194km to Ugak Island (Figure 1).
Another animal crossed 74 km of open ocean to
Chirikof Island and then returned to southwestern
Tugidak. The other movements ranged between 26
and 74 km. The general pattern for these animals
seemed to be for them to remain at the new site for
some period of time rather than using many locations.
Twelve of 17 relocations of radio-tagged seals on haul-
outs other than southwestern Tugidak were clustered
in the northern Tugidak, Sitkinak, southern Kodiak
area (Figure |), which are the nearest hauling areas to
southwestern Tugidak. Several different “types” of
movements occurred. One seal (TR-18) made a long
move to another hauling area which it appeared to use
for the remainder of the study period. Another (TR-5)
made a long move to a hauling area, then returned to
southwestern Tugidak where it appeared to stay for
the remainder of the summer. Two radio-tagged seals
(TR-4, 13) alternated between two haulouts. Three
seals (TR-8, 15,22) were found at hauling areas other
than southwestern Tugidak only once. No correlation
between sex and age of the radio-tagged seals and
extent or type of movement was apparent; however,
sample sizes of all groups except adult females were
very small.

It was probable that additional movements of
radio-tagged seals occurred. Many of the other
transmitter-equipped seals were absent from south-
western Tudigak for extended periods (Figure 2).
Only occasional radio-tracking surveys were flown
and radio-tagged animals which were not hauled out
at the time or in the area of the survey would not have
been found.

Information on average movement rates was
derived by dividing minimum distances between con-
secutive sighting by elapsed time. Rates for four
animals were 24, 19, 27, and 26 km/d. These rates
were minimums because actual routes traveled were
unknown and actual travel times were no doubt less
than observed in most cases.

Information previously available on movements of
individual Harbor Seals was from the recoveries of
animals which had been tagged within several weeks
of their birth (Bonner and Witthames 1974; Boulva
and McLaren 1979; Alaska Department of Fish and
Game, unpublished data). They documented dispersal
of juveniles up to 250 km from large pupping areas.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Mansfield (1967) remarked on the wandering or dis-
persal of young Harbor Seals, referring to them as
“rangers.” Additional insight into the range of Harbor
Seal movements comes from offshore sightings.
Spalding (1964) reported observations of Harbor
Seals 50-65 km offshore in the Gulf of Alaska, and
Wahl (1977) saw a seal 80 km off the coast of
Washington State.

Haulout Area Fidelity

There appeared to be considerable fidelity to spe-
cific haulout areas by individual radio-tagged seals.
Twenty-three of 31 (74%) seals that were relocated
after capture were found only at the capture site. Of
the eight animals that were found on haulouts other
than southwestern Tugidak, three were found on the
same haulout more than once. Only one seal was
found on more than two sites and it was found on only
three. Additional use of haulouts other than south-
western Tugidak may have gone undetected, but the
consistency of these data leads to the conclusion that
few sites are generally used.

Other investigators have also presented evidence
suggesting considerable site fidelity. Calambokidis et
al. (1978) suggested that both year-round site loyalty
and long-distance movements occurred in Harbor
Seal populations in Puget Sound. They based this on
repetitive observations of uniquely marked individu-
als and fluctuations of counts of seals on hauling
areas. Knudtson (1974) and Reijnders (1976) reported
observations of the same animals returning repeatedly
to the same hauling area. Divinyi (1973) collected a
tagged Harbor Seal on the southwestern Tugidak
haulout where it had been tagged as a pup three years
earlier. Boulva and McLaren (1979) saw recognizable
individuals hauled out in the same area day after day
during summer.

Frequency of Haulout and Proportion Hauled Out

Insight into what proportion of the population was
represented by counts of hauled-out seals was gained
by examining haulout histories of radio-tagged seals
(Figure 2). Since more than one hauling area was
sometimes used and since only southwestern Tugidak
was regularly monitored, it was impossible to quantify
precisely the haulout frequency of all radio-tagged
seals. To alleviate this problem we subsequently clas-
sified certain radio-tagged seals as southwestern
Tugidak residents by excluding those found hauled
out at other locations and those which were absent
from southwestern Tugidak for extended periods. We
then examined the haulout data from these animals
during two periods with frequent monitoring and
minimal disturbance (1-30 June and | August-—5
September).

In June, resident radio-tagged seals were found
hauled out during daily radio checks on an average of

1981

iA G Ge D SE A {t NUMBERS

RADIO

PITCHER AND MCALLISTER: HARBOR SEAL MOVEMENTS

tL)
20 o— 0 [}-—HTHHTHIMH
: _— oe
5 (0 [Reon ae py Coo—Tinon TOD
8 Cllr Erie Saw MHHO TH
7 O70
8
O75 u
10 O
italien Gl
12 O14
ies 1 THI
1 WA THI HOA TH
oeae 0 1
16 -—THHI ier ea
ee teem Pk
18 0
19
20
MAY JUNE AUGUST

SEPT

USD)

FiGure 2. Haulout patterns of radio-tagged Harbor Seals on southwestern Tugidak Island showing the presence or absence

of each individual during daily radio checks. Open box, present; dark box, found on other haulout; capture operation
caused some disturbance between 8 May and 2 June; 25 valid radio checks between | and 30 June; no onsite radio
checks 2-31 July; 31 valid radio checks between | August and 5 September.

296

50 + 7% (95% confidence interval) of the days, and
between | August and 5 September they were found
hauled out on 41 + 5% (95% confidence interval) of
the days (Table 1). On an individual basis, frequency
of haulout ranged from 16% (TR-20 during June and
TR-24 during August-September) to 80% (TR-19
during June) of the days. If the resident radio-tagged
animals were representative of the population, the
average number of seals hauled out during the daily
radio checks probably represented between about 35
and 60% of the total population.

Our findings agree quite well with the results of
certain other studies. Sullivan (1979) found that Har-
bor Seals spent an average of 44% of their daily activ-
ity budgets hauled out. Summers and Mountford’s
(1975) estimate of Harbor Seals in the Wash, Great
Britain, based on mark-recapture studies, was con-
siderably higher than numbers seen hauled out. Finley
(1979) speculated that midday counts of Ringed Seals
(Phoca hispida) represented about 70% of the popula-
tion based on a recognizable individual that was
observed 71% of the time.

The effects of capture, handling, and attachment of
the transmitter package on the behavior of radio-
tagged seals was largely unknown but cannot be
entirely ignored. Haulout behavior of radio-tagged
seals observed on southwestern Tugidak appeared to
be normal. However, some seals developed abrasions
where the bracelet encircled the ankle, apparently
because of excess rigidity of the bracelet. In one
instance, both the health and choice of haulout loca-
tion of a radio-tagged seal appeared to be influenced
by the abrasion resulting from the transmitter
attachment. TR-22 was relocated 56 d after capture
ona beach not normally used as a haulout by Harbor

THE CANADIAN FIELD-NATURALIST

Vol. 95

Seals. She appeared thin and weak. Her hind flipper at
the point of transmitter attachment was badly chafed
and infected. We recaptured her and removed the
transmitter.

Four radio-tagged seals were never relocated after
release. They may have moved beyond the range of the
aerial surveys or not have been hauled out during any
of the surveys. If the transmitters failed or were lost
they would not have been relocated during aerial sur-
veys, but would have been seen on southwestern
Tugidak where visual searches were routinely made.
Mortality, either related or unrelated to capture and
transmitter attachment, could explain lack of con-
tacts. Hammond and Elsner (1977) reported delayed
deaths of seals several hours after they appeared to be
recovered from immobilization with Ketamine
hydrochloride.

Acknowledgments

This study was supported by the Bureau of Land
Management through interagency agreement with the
National Oceanic and Atmospheric Administration,
under which a multi-year program responding to
needs of petroleum development of the Alaskan con-
tinental shelf is managed by the Outer Continental
Shelf Environmental Assessment Program (OCSEAP)
office. Field assistance was provided by G. Browning,
P. Smith,and D. Stevens. Pilots during aerial radio-
tracking surveys were P. Buker, R. Wright, and Lts.
(NOAA) Christman and Layden. Drafts of the
manuscript were critically reviewed by J. Burns, D.
McKnight, K. Schneider, and D. Siniff. Thanks are
due to R. Reicle and D. Siniff of the University of
Minnesota for their assistance and advice on radio-
tracking procedures.

TABLE |—Haulout frequency of resident radio-tagged Harbor Seals on southwestern Tugidak Island

Resident 1-30 June | August — 5 September
seal Days Days Proportion Resident Days Days Proportion
number checked hauled out days hauled out seal no. checked hauled out days hauled out
2 25 1] 0.44 5 31 19 0.61
7 25 16 0.64 6 31 9 0.29
14 25 16 0.64 7 31 22 0.71
16 25 1] 0.44 13 31 10 0.32
17 25 9 0.36 14 31 19 0.61
19 25 20 0.80 16 31 15 0.48
20 25 4 0.16 17 31 9 0.29
19 31 15 0.48
24 31 5 0.16
29 3] 13 0.42
34 31 10 0.32
35 31 7 0.23
Overall 175 87 0.50 Overall 372 153 0.41

1981

Literature Cited

Bigg, M. A. 1969. The Harbour Seal in British Columbia.
Bulletin of the Fisheries Research Board of Canada 172:
33 pp.

Bonner, W.N., and S. R. Witthames. 1974. Dispersal of
Common Seals (Phoca vitulina), tagged in the Wash, East
Anglia. Journal of Zoology, London 174: 528-531.

Calambokidis, J. A., R. D. Everitt, J.C. Cubbage, and
S. D. Carter. 1979. Harbor Seal census for the inland
waters of Washington, 1977-1978. Murrelet 60: 110-112.

Divinyi, C, A. 1973. Growth and movements of a known-
age Harbor Seal. Journal of Mammalogy 53: 824.

Finley, K. J. 1979. Haul-out behavior and densities of
Ringed Seals (Phoca hispida) in the Barrow Strait area,
NWT. Canadian Journal of Zoology 57: 1985-1997.

Fisher, H. D. 1952. The status of the Harbour Seal in Brit-
ish Columbia with particular reference to the Skeena
River. Bulletin of the Fisheries Research Board of Canada
93: 58 pp.

Hammond, D., and R. Elsner. 1977. Anesthesia in phocid
seals. Journal Zoo Animal Medicine 8: 7-13.

Knudtson, P.M. 1974. Mother-pup behavior within a
pupping colony of Harbor Seals ( Phoca vitulina richardi)
in Humboldt Bay, California. M.Sc. thesis, California
State University, Humboldt, California. 42 pp.

Mansfield, A. W. 1967. Distribution of the Harbor Seal,
Phoca vitulina Linnaeus, in Canadian Arctic waters.
Journal of Mammalogy 48: 249-257.

PITCHER AND MCALLISTER: HARBOR SEAL MOVEMENTS

Doi,

Pearson, J. P., and B. J. Verts. 1970. Abundance and dis-
tribution and Harbor Seals and Northern Sea Lions in
Oregon. Murrelet 51: 304-305.

Reijnders, P. J. H. 1976. The Harbour Seal (Phoca vitul-
ina) population in the Dutch Wadden Sea: size and com-
position. Netherlands Journal of Sea Research 10:
223-235.

Rostental, R. J. 1968. Harbor Seal censuses in Humboldt
Bay during 1966 and 1967. California Fish and Game 54:
304-305.

Scheffer, V. B., and J. W. Slipp. 1944. The Harbor Seal in
Washington State. American Midland Naturalist 32:
373-416.

Spalding, D. J. 1964. Comparative feeding habits of the
Fur Seal, Sea Lion, and Harbour Seal on the British
Columbia coast. Bulletin of the Fisheries Research Board
of Canada 146: 52 pp.

Sullivan, R. M. 1979. Behavior and ecology of Harbor
Seals, Phoca vitulina, along the open coast of northern
California. M.Sc. thesis, California State University,
Humboldt, California. 115 pp.

Summers, C. F., and M. D. Mountford. 1975. Counting
the Common Seal. Nature 253: 670-671.

Wahl, T. R. 1977. Sight records of some marine mammals
offshore from Westport, Washington. Murrelet 58: 21-23.

Received 8 November 1980
Accepted 5 March 1981

Reproduction, Distribution, and Population Size of Largemouth Bass,
Micropterus salmoides, in an Oligotrophic Precambrian Shield Lake

DAVID C. MARALDO! and HUGH R. MACCRIMMON

Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1
'Present address: Algoma Fisheries Assessment Unit, Ontario Ministry of Natural Resources, P.O. Box 130, 69 Church
Street, Sault Ste. Marie, Ontario P6A 5L5.

Maraldo, David C., and Hugh R. MacCrimmon. 1981. Reproduction, distribution, and population size of Largemouth
Bass, Micropterus salmoides, in an oligotrophic Precambrian Shield lake. Canadian Field-Naturalist 95(3): 298-306.

Numbers of self-sustaining, resident Largemouth Bass, Micropterus salmoides, (> 145 mm FL) in Tadenac Lake (45°04’N;
79°57’W) (308 ha) were estimated at 350 in 1975 and 213 in 1976, based on trap net data and angling returns of tagged fish.
Trap net catches included fish 1+ to 7+ years of age, but I+ and 2+ fish were not fully sampled because of gear Selsey: :
Biomass was calculated at 0.39 (1975) and 0.21 (1976) kg -ha’! for the entire lake, and 4.94 (1975) and 2.63 (1976) kg -ha'! for
littoral habitat used by Largemouth Bass. Adults spawned in shallow(< 1.4 m) sheltered bays usually (75%) on basal leaves of
Eriocaulon septangulare in late May and early June. Only 0.8% of the lake bottom provided potential nesting sites. Nine
completed nests were found in 1975, 15 in 1976. Only two nests failed to produce viable progeny each year. Mean survival rate
to hatch in successful nests was 0.978. Both juveniles and adults inhabited weeded littoral areas (8% of the lake surface area),
which offered the warmest water available, although maximum epilimnetic temperatures remained below the final preferen-
dum for the species. The size of natural populations of Largemouth Bass in oligotrophic Precambrian Shield lakes 1s

apparently limited by the extent of adequate warm weedy littoral areas.

Key Words: distribution, Largemouth Bass, Micropterus salmoides; nesting sites, oligotrophic, population estimates,

Precambrian, reproduction.

Largemouth Bass (Micropterus salmoides) inhabit
numerous oligotrophic lakes on the Ontario Precam-
brian Shield near the northern fringe of their native
range (Robbins and MacCrimmon 1974). Littoral fish
communities in these lakes may include the Small-
mouth Bass (M. dolomieui), Walleye (Stizostedion
vitreum vitreum), Northern Pike (Esox lucius), or
Muskellunge (E. masquinongy). Most of these lakes
have deeper waters suited to salmonids and coldwater
fish communities.

Tadenac Lake, one of these oligotrophic Precam-
brian Shield lakes, offers a unique opportunity to
study an indigenous self-sustaining population of
Largemouth Bass. This lake has no road access, no
habitations, very limited controlled angling, and,
therefore, has beenimmune from appreciable cultural
stresses. The purpose of our study was to describe the
reproduction, density, and biomass of the species in a
typical oligotrophic Precambrian Shield lake and
compare the data with that from warm eutrophic
waters where abundant populations of Largemouth
Bass normally occur.

Study Area

Tadenac Lake (45°04’N; 79°57’W) is located within
the Precambrian Shield of Ontario in Georgian Bay
Township. The lake has a surface area of 308 ha anda
maximum depth of 30 m, and is typically oligotrophic

in physical and chemical characteristics (mean depth,
8.2 m; August thermocline, 7-9 m; pH, 6.95; conduc-
tivity, 36 ws cm’).

Substrate within the littoral zone consists of sand,
gravel, barren rock, and silt. In the shallow bays bul-
rushes, Scirpus sp.; sedges, Cyperaceae family; pick-
erel weed, Pontederia sp.; pond lilies (white and yel-
lows), Nymphae sp.; Potamogeton sp.; and Pipewort,
Eriocaulon septangulare, were abundant. The lake is
surrounded by a conifer and mixed hardwood forest
which has regenerated after the area was clear-cut
before 1895. The ichthyofauna of the lake consists of
21 cold-water and warmwater species representing I 1
families (Maraido 1978).

A dam has maintained the level of Tadenac Lake
above that of the adjacent waters of Georgian Bay
since 1925. Midsummer surface levels of Tadenac
Lake were about 0.5 m above Georgian Bay during
the study. Before 1925, there would seem to have been
a free access to Largemouth Bass between Georgian
Bay and Tadenac Lake waters.

Material and Methods
Population Density

In 1975, six 1.8-m and four 2.4-m trap nets (vertical
measurements of box, lead, and wings) were operated
in Tadenac Lake from early May to mid-October. In

298

1981

1976, nine 2.4-m, one |.8-m, and one |.2-m trap nets
were fished from late April through September (Fig-
ure 1). Nets were lifted at weekly intervals, except
during the early spring when they had to be tended
more frequently. Catch per unit effort is the total
number of bass captured divided by the total number
of net-days in that month. Additional specimens of
Largemouth Bass were captured for examination,
tagging, and subsequent release by use of a 60-cm
small mesh trap net, a 2.5-cm monofilament nylon gill
net, Plexiglas fry traps (Casselman and Harvey 1973),
a 9-m seine, and by angling. Fish retained in anglers’
creels were also examined.

Fork length was measured to the nearest millimetre,
and fish were weighed to the nearest 25 g (if 1.0 kg)
or 10 g (if <1.0 kg). Fish greater than 140 mm FL
were tagged with either strap or spaghetti tags, and
finclipped. Smaller fish (<140 mm FL) were only
finclipped. Scale samples were taken from the body at
the posterior tip of the depressed pectoral fin. The
determination of age and growth from scales has

MARALDO AND MACCRIMMON: LARGEMOUTH BASS

299

proven valid for fish to age 7+ (Maraldo and Mac-
Crimmon 1979). Each fish was released at the point of
capture after tag numbers and vital statistics had been
recorded.

Population estimates were calculated from 1975
and 1976 trap net catches using the Schnabel method
with confidence limits determined by treating the
number of recaptures as a Poisson variable and, for
comparative purposes, were computed also by the
Petersen formula (Ricker 1975). The angling season
(July and August) was divided into four equal periods
with the number of unmarked and recaptured fish
summed over each. Annual Petersen population esti-
mates are the mean of the four values.

Proportions of the total catch in each age-class were
multiplied by the population estimates and then by the
mean weights for that year-class at the time of annulus
formation (Maraldo and MacCrimmon 1979). Total
biomass is the sum of these total weights for each
year-class, and is expressed as per unit area for both
the entire lake and the littoral area.

TADENAC LAKE

DEPTHS IN METRES

Ale AP SNEM@ SilitiES

eA -H

SPAWNING AREAS

FiGuRE |. Bathymetric map of Tadenac Lake showing trap net locations and spawning areas.

300 THE CANADIAN FIELD-NATURALIST

Reproduction

Daily searches of shallow areas (<2 m) for evidence
of nest building or reproduction by Largemouth Bass
began on 19 May 1975 and 11 May 1976. The bright
yellow eggs and the nearby presence of the male par-
ent identified active nests. The depth, distance from
shore, availability of cover, and substrate type of nests
were described. Nest sites were checked at I- to 3-d
intervals in 1975 and the status of embryos or larvae,
presence or absence of adult, and water temperatures
were recorded. In 1976, daily samples of embryos were
collected from nine nests by sweeping the mouth of a
large pipette across different areas of the nest while
applying suction. Ratios of living embryos (yellow
and translucent) to the total number of eggs per sam-
ples were then determined.

Distribution

Visual observation of the distribution of postemer-
gent juveniles was continued, at least weekly, to the
end of July in both years. From 9 August to 4 Sep-
tember of 1976, two Plexiglas fry traps (Casselman
and Harvey 1973) were placed in areas where schools
of Largemouth Bass had been observed previously.
Daily observations and catches of juveniles were
related to water temperature and depth, and to the size
of the fish. Data from trap net catches provided
information on distribution of larger fish, and on the
movements of those previously tagged.

Results
Spawning and Embryo Survival in Nests

A total of eight spawning areas were located in
shallow weedy bays, with fallen trees often present. In
1975, nine active nests were found in five bays between
21 May and 15 June, while 15 nests were located in six
bays between 20 May and 5 June of 1976 (Figure 1).
These nests were constructed between | and 10m
from shore at water depths of 0.2 to 1.4 m. They were
typically circular, about | m in diameter, and most
frequently (75%) situated on the basal leaves of E.
septangulare. The site was usually cleared of settled
organics by the male parent, but no depression was
formed in the substrate. Spawning occurred less than
24 h after the initiation of nest preparation. The eggs
were usually deposited in an area of about 0.3 m in
diameter, located near the center of the nest. There is
some evidence of spawning at the same locations in
successive years, as one nest was found at the same site
in both 1975 and 1976. :

The presence of tags on adult fish did not appear to
interfere with spawning activity. Tags were evident on
two males guarding nests in 1975, and on six in 1976,
one of which had been tagged only 8 d earlier. Guard-
ing males taken by angling from nests for tagging or
tag checks returned immediately to their nests on

Vol. 95

release. Guarding males generally remained within
2 m of their nests and defended the nest vigorously
against intruders until schooling juveniles moved
from the site. The embryos hatched 3 to 4d after
fertilization at water temperatures of 18.4 to 19.6°C.
Free-swimming juveniles were first evident at the nest
sites 4 to 6d after hatching.

Ambient nest temperatures ranged from 17.5 to
23.0°C between 21 May and 22 June in 1975, and from
16.5 to 25.5°C between 27 May and 9 June in 1976.
Water temperatures declined in 1975 from 22.8°C on
27 May to 18.0°C at some nests on the following day
and did not increase again until mid-June. In 1976,
however, the nest temperatures were lowest on the day
of discovery, and increased daily thereafter. The one
exception was a nest where low temperatures of 10.5
to 12.0°C prevailed between 20 and 23 May.

In 1975, two nests were unsuccessful; one became
exposed to air by declining water levels and the other
became almost completely covered by silt. Of nine
nests under scrutiny in 1976, all but two successfully
produced living embryos (sac-fry) at ambient incuba-
tion temperatures from 16.0 to 20.3°C. Mean daily
survival estimates of embryos for all nine nests ranged
from 0.062 to 0.995 (Table |). Despite survival rates of
0.963 and 0.638 in nest | on 21 and 22 May, no
embryos survived to emergence following exposure to
ambient water temperatures of 10.4 to 10.8°C. In nest
7, the other unsuccessful nest, survival rates were
0.165 and 0.062 on 29 and 30 May at temperatures of
19.5 and 19.0°C. Mean daily surface water tempera-
tures in all spawning areas during the sampling period
of 27 to 31 May ranged from 18.4 to 20.3°C. There
appeared to be no major changes in daily survival
rates over the incubation period at the seven success-
ful nests (Table 1).

Estimated survival of Largemouth Bass embryos in
nest 9 was 0.993 on 30 May 1976 while in nest 7
survival was only 0.062 on the same day (Table !).
These two nests were located within 10 m of each
other and were visually identical.

Distribution

A total of 23 sightings of schooling juveniles were
made between 10 and 26 June 1975. All but one of
these were in spawning areas. Water temperatures at
the sites of these schools (18.5°C on 10 June; 26.5°C
on 26 June) were similar to surface waters elsewhere in
the lake. In 1976, eight sightings of schooling juvenile
Largemouth Bass were made from 18 June to 4 July,
all in spawning areas. Water temperatures at the sites
of the schools were 23.0 to 25.0°C, typical of surface
temperatures elsewhere in the lake. All schools sighted
in both years were in <1.5 m of water with the fish
generally concentrated in the upper 0.3 m of the water
column.

1981 MARALDO AND MACCRIMMON: LARGEMOUTH BASS 301

In 1976 a total of 65 young-of-the-year bass were

Sih $3 captured between 10 August and 19 September in the
QS Se = two Plexiglas fry traps. Water temperatures at the
= es Ss S traps ranged from 20.5 to 25.0°C during August and
roe > were within 1.5°C of the maximum recorded at any
SSS S time in the lake.
Trap net fishing effort totalled 1740 and 1595 net-
days in 1975 and 1976. Mean monthly catch per effort
RM m enor (Table 2) varied considerably over the fishing season
S eas Ss with Largemouth Bass being most easily caught in
ES se S 3 Bs BS 2 May of both 1975 and 1976. Typically, catch per effort
Pees Wes ea es declined toa lowin July 1975 and June 1976, followed
by slight increases in August and September. The
numbers of fish caught at each of the eight net sites
Ze used in both years (Table 3) did not differ significantly
=a (t - test; P< 0.05).
al eseee Recaptures of tagged fish indicated little movement
2a IS & S28 Se of Largemouth Bass between areas of the lake, except
ssoocso during late spring and early summer. Only 32% (8 / 25)
and 23% (9/40) of the fish tagged in 1975 and 1976,
© respectively, were recaptured at a different site during
HN SI the same year. Fish that had moved between net loca-
Bl os Ss tions traveled minimum distances of 0.5 to 4.6 km in
PS |S ‘S 1975 and 1976. In 1975, 7 of the 14 recaptures in May
2 2 and June were at new locations while all 10 recaptures
3 in July, August, and September were at the site of
A original tagging. This pattern was repeated in 1976
ie with 7 of 15 recaptures in June and July recovered at
mc new locations. Individual fish tended to remain local-
SiS ized from one year to the next with only 4 of 10 fish,
sat tagged in 1975 and recovered in 1976, recaptured at a
S location other than that of original capture.

Information on daily and seasonal movements of
Largemouth Bass was also obtained from nesting
males. Nests of two males were located within 100 m
of the net where they had been tagged 8 and 30d
before, while two others had nests 0.4 and 1.8 km
from the point of tagging 32 and 21 d previously.
Later in the year, three males were recaptured in trap
nets located within 100 m of their former nest, and
another was captured by angling at distances of
1.1 km in July and 0.5 km in September.

Largemouth Bass aged 1+ to 7+ were caught in the
trap nets in both 1975 and 1976 (Table 4). Fish aged 3+

22
0.6380.0295

21
0.9630.0473

TABLE |—Estimated mean survival (SE) to hatch of Largemouth Bass embryos in nests during May 1976

8

B

=

el

°

e TABLE 2—Catch per effort (N) by month for trap nets set in
oe = Tadenac Lake for Largemouth Bass
2 i)
Sa ae aes Sule 1975 1976
°o P=

5 May 0.294 (72) 0.144 (36)

sal June 0.086 (25) 0.087 (28)
BS o July 0.047 (14) 0.089 (33)
gs —amnonwacl||” August 0.057 (15) 0.089 (25)

v7 September 0.064 (22) 0.098 (30)

302

(1973 year-class) dominated the 1976 catch, but this
year-class had not been dominant in the 1975 catch,
indicating that age 2+ fish were not fully recruited into
the trap net fishery. Largemouth Bass in the angler
catch ranged from 235 to 413 mm FL and 0.2 to
1.31 kg. Age 4+ fish dominated the catch of 1975 and
age 3+ fish in 1976 (Table 4).

Tag loss in the year of tagging was 29.7% in 1975
and 20.0% in 1976. Tag loss between 1975 and 1976
among 34 fish recaptured was 70.6%.

Population Estimates

Because of gear selectivity, only fish greater than
145 mm FI are included in the trap net estimates of
350 fish (1.14 ha') in 1975 and 213 fish (0.69 ha‘') in
1976. The 95% confidence interval of the trap net
estimates of 159 to 292 fish over the whole netting
season in 1976 overlapped the 95% confidence interval

THE CANADIAN FIELD-NATURALIST

Vol. 95

of the 1975 estimate of 246 to 520 fish, indicating that
the population size remained similar over the two
years. Trap net estimates for May and June only fell
within the 95% C.I. of the estimates over the entire
netting season in both years (Table 5).

Biomass of Largemouth Bass in Tadenac Lake was
estimated at 120.6 kg in 1975 and 64.2 kg in 1976, or
0.39 and 0.21 kg:-ha"! (Table 4). These are minimum
estimates because of the incomplete vulnerability of
the first three age-classes to the trap net fishery.

Means of the four annual population estimates
based on angler returns of tagged fish were 266 and
242 individuals in 1975 and 1976. These estimates are
within the 95% confidence intervals of the trap net
estimates of 246 to 520 in 1975, and 159 to 292 in 1976;
thus indicating good agreement between the two
methods. Again, population estimates from angler

TABLE 3—Catch statistics for Largemouth Bass at various trap net sites in Tadenac Lake

Water depth catien
Trap net at trap mouth Substrate Number Percent

Location * Time period size (m) (m) type of fish of total
1975

I 6 May-12 October 1.8 2.4 Silt 30 19.5

2 2 May-12 October 2.4 2.4-3.0 : Silt 16 10.4

3 4 May-12 October 1.8 37 Silt 13 8.4

4 :

5 4 May-29 June 2.4 2.4-3.0 Silt 6 3.9

6 6 May-—12 October 1.8 3.7-4.5 Gravel-rock 2 1S

7 2 May-12 October 1.8 2.4-3.0 Silt 6 4.5

8 1 July-12 October 2.4 4.5-6.7 Gravel-rock 0 0.0

9

10 6 April-29 June 1.8 3.7-5.5 Rock 0 0.0

1] 7 May-29 June 2.4 2.4-3.0 Silt 25 16.2

12 7 May-29 June 1.8 4.5-6.1 Sand-gravel 34 22.1

13 | July-12 October 1.8 3.7-4.5 Rock | 0.6

14 2 July—12 October 1.8 4.5 Gravel 4 2.6

15 3 May-12 October 2.4 4.5 Silt 16 10.4
1976

l 21 April-27 September 1.8 Doll Silt 35 21.6

2 20 April-27 September 2.4 2.4 Silt 18 11.1

3 20 April-27 September 2.4 4.3 Silt 28 14.2

4 21 April-27 September Dn 4.3 Sand-gravel — 7 4.3

5 21 April-27 September 12 1.8 Silt 3 1.9

6

7 20 April-27 September 2.4 307 Silt 15 9.3

8

9 21 April-27 September 2.4 4.9 Rock-gravel 0 0.0

10

11 26 April-27 September 2.4 1.8-3.7 Silt 21 13.0

12 20 April-27 September 2.4 5.8 Sand-gravel 24 14.8

13

14

15 20 April-27 September 2.4 Jol/ Silt 16 9.9

“Number refers to location in Figure 1.

303

LARGEMOUTH BASS

MARALDO AND MACCRIMMON

1981

‘(6L6] UOWUNID.eYW pur Opyeiey)) syIZua] payepno[eo-yoerq wWoI paurutiajaq ,
TZ 91Qe] Wor sajyeutysa suNf—ABP BuIsy,

8600 68°LI ce 80¢ 0 90°79
v00°0 lel Ole! 67 I ¢r0'0 98 E1
400°0 80°C OO! Ls C L¢£0°0 Lvl
c10°0 IL'¢ 0£6 VI v 0c0'0 cc9
L00°0 (ais Ors PI 17 £700 SIL
9c0'0 C18 OLE 6°79 CC £400 0S CC
c00°0 cs 0 OLT Ls C 800°0 9?
= a ee 0 0 100°0 pe 0
— ——— et 0 0 —— —
8900 L607 ef C6E 0 OL 071
1100 Of € OOTT 16 € 170'0 p9'Tl
c10°0 OLE Or6 Vl v 9110 EL Se
8100 CSS 069 CHC 8 8e10 CS CP
0c0'0 80°9 Ors p9E Cl 090'0 €¢ 81
900°0 961 06E ISI ¢ Vc0'0 BEL
100°0 ct 0 Ost Or I C100 09'£
a = ae 0 0 100°0 OL 0
(,-eu- 34) (34) (3) “41M yoieo 1y43nvd (,-eu- 34) (34)
eaIe 19g [210 | uray 12101 Joquiny BoIe 19g [8101
JSOAIV HY eae (dois dutpur}s)

ssewo!q payeuns|

soyojeo 1a[suy

102
C6 SI
SOL cl
Sc9 Ol
BLE 61
OSC 06
68 6c
SI €C

L6C
COL 81
1€8 eV
L69 19
ler eV
S0¢ 9E
89 es
L ev

g(3) ys uorjejndod
urd, ~—s-UT. OU [B}0}
poeunsy

soyojieo jou deiy

OL
OL
8
¢°6

yoyeo
ye101
jo
qUIdI19g

SOl

8
8

jysnes
Joquiny

[210 |

eee

"soyojeo Jajdue pue jou des] Wosy sseuorg payepnoyeo puke ‘s}yZI9M URS ‘UOIINGLISIP ad3yW—p ATEV_L

304

returns apply only to the size range of fish represented
in the angler catch.

Discussion

The distribution of Largemouth Bass in Tadenac
Lake is restricted to shallow (<3 m), weedy bays
offering the highest summer water temperatures, with
limited movements between these areas. Even in these
bays, midsummer water temperatures do not
approach the final preferendum of 30°C for Large-
mouth Bass young-of-the-year (Maraldo 1978) and
older fish (Siler and Clayton 1975). With fish selecting
those temperatures closest to the preferred tempera-
ture for the species, it is likely that 3 m is the lower
limit of the normal summer distribution since the
midsummer thermocline begins at 3-4 m. Although
trap net data (Table 3) show catches of Largemouth
Bass down to 6 m, these fish undoubtedly encoun-
tered the lead in much shallower water before entering
the trap.

Because spawning was localized in only five differ-
ent littoral sites in 1975 and six in 1976, there are
obvious assemblages of adult fish at favored spawning
locations, a phenomenon also noted by Kramer and
Smith (1962). Using catch per effort data from the
stationary trap nets as an index of movement (Table
2), it appears that local movements of fish within
Tadenac Lake were greatest in May with a decline in
June and July, followed by a slight increase in August
and September similar to findings of Hasler and
Wisby (1958).

To validate population estimates, we can reduce the
problems of non-random distribution of marked fish
and recruitment of new individuals into the fishery by
considering data from only May and June. This inter-
val coincides with the period of greatest movements of
Largemouth Bass in Tadenac Lake, and covers a short
time span during which there could be only a very
limited recruitment of new individuals into the
fishery. Combined May and June estimates of 297 and
201 Largemouth Bass in 1975 and 1976 (Table 5) are
similar to those of 350 and 213 individuals made over
the entire season. May and June estimates were within
the 95% confidence intervals calculated for the whole
season estimates in their respective years.

Computed estimates from fish recaptured by a dif-
ferent method (angling) than how initially captured
for tagging (trap) reduces possible biases associated
with differential vulnerability to capture between
tagged and untagged fish, and from non-random dis-
tribution of fishing effort. Population estimates from
angler returns were calculated from averaged data
over four 2-wk periods during the angling season. By
using this short time span for each estimate, the effects
of recruitment on the population estimate were
greatly reduced. Numbers of catchable-sized Large-

THE CANADIAN FIELD-NATURALIST

Vol. 95

mouth Bass in the lake estimated from angler returns
were 266 in 1975, and 242 in 1976. Both population
estimates were within the 95% confidence interval
calculated for the trap net estimates in that year, 1.e.,
246 to 520 individuals in 1975, and 159 to 292 individ-
uals in 1976.

Calculated biomasses of Largemouth Bass in
Tadenac Lake of 0.39 and 0.21 kg-ha'! in 1975 and
1976 represent very low figures relative to other waters
(Carlander 1977; Mahon and Balon 1977). About 25%
of the total surface area of Tadenac Lake is less than
3m deep with this percentage remaining relatively
constant because of the dam at the outlet. If only that
littoral area frequented by Largemouth Bass in used in
the calculation, the total biomass per area becomes
1.56 and 0.84 kg-ha ' in 1975 and 1976. The littoral
area actually providing habitat suitable for Large-
mouth Bass (1.e. <3 m deep, excluding areas of rocky
shelves and shoals, windswept sand beaches lacking
aquatic macrophytes) is only 24.4 ha, representing
32% of the littoral zone, 8% of the total lake surface
area, and 0.64% of the total volume. The biomass of
Largemouth Bass for these areas of favored habitat is
estimated at 4.94 kg-ha! (1975) and 2.63 kg-ha!
(1976) representing a significant localized abundance.

As in other waters (Miller and Kramer 1971; Hei-
dinger 1975), the spawning of Largemouth Bass in
Tadenac Lake was confined to shallow bays sheltered
from prevailing winds. Most nests were on one type of
substrate (1.e., the basal leaves of FE. septangulare).
Similarly in Minnesota, Kramer and Smith (1962)
found most nests located on mats of one kind of
aquatic vegetation, Eleocharis acicularis. The poten-
tial spawing habitat in Tadenac Lake was estimated to
be 2.6 ha representing about 0.8% of the lake bottom.

The spawning period of late May to early June for
Largemouth Bass in Tadenac Lake was similar to that
recorded in other waters in the northern parts of its
range (Mraz et al. 1961; Kramer and Smith 1962;
MacKay 1963). Water temperatures in most Large-
mouth Bass nests were within the 18.9 to 20.0°C range
typical of the species (Robbins and MacCrimmon
1974).

Depths of water over nests, ranging from 0.2 to
1.4 m, agree also with that in other waters (Mraz 1964;
Chew 1974: Robbins and MacCrimmon 1974), but are
considerably less than the 1.1 to 7.5 m reported by
Allan and Romero (1975). It is improbable that
spawning occurred in the deeper water of Tadenac
Lake because of the narrow muddy littoral shelf and
the abrupt change in average water temperature from
18.9 to 7.5°C at the 3-to 5-m thermocline during the
spawning season.

Meanestimates of survival to hatch of Largemouth
Bass embryos in successful nests (0.978) are greater

1981

MARALDO AND MACCRIMMON: LARGEMOUTH BASS

305

TABLE 5—Estimated populations of Largemouth Bass in Tadenac Lake

Confidence Number Size range
N interval marked Rt FL (mm) Wt. (kg)
1975
Schnabel estimate from trap nets
Over whole season 350 246-520 150 30 145-467 0.05-1.98
May-June (only) 297 180-531 93 15 162-467 0.06-1.98
Petersen estimates from
angler returns
Early July (1-15) 184 51-1840 92 2 300-393 0.50-1.05
Late July (16-31) 242 103-756 93 5 264-413 0.35-1.25
Early August (1-15) 285 79-2850 95 2 280-379 0.35-1.07
Late August (16-31) B58 120-1767 106 3 235-367 0.18—-0.90
Mean from angler returns 266
1976
Schnabel estimate from trap nets
Over whole season 213 159-292 127 45 174-411 0.07-1.60
May-June (only) 201 112-409 66 11 222-411 0.15—-1.60
Petersen estimates from
angler returns
Early July (1-15) 476 128- 68 | 273-420 0.34-1.39
Late July (16-31) 168 72-535 70 5) 249-388 0.28-0.97
Early August (1-15) 237 81-1185 79 3 278-401 0.32-1.11
Late August (16-31) 85 41-213 85 7 _ 0.20-0.95
Mean from angler returns 242

than the values of 0.804 and 0.922 reported in Utah
(Miller and Kramer 1971), and 0.65 to 0.94 in Minne-
sota (Kramer and Smith 1962). Incubation tempera-
ture fluctuations in successful Tadenac Lake nests
were well within the 10 to 13°C range of thermal
tolerance reported for Largemouth Bass embryos
(Swallow 1968).

The importance of direct temperature effects on
Largemouth Bass mortality is not clear. Coutant
(1975) considered small temperature changes not to be
responsible for high mortalities determining year-
class strengths. Badenhuizen (1969) also concluded
that early mortality in Largemouth Bass embryos may
not be adequately explained by temperature effects
alone. At both unsuccessful Tadenac Lake nests in
1976, males remained in attendance even after total
egg mortality had occurred. This is in contrast to the
suggestion by Kelly (1968) and Heidinger (1975) that
lowered temperatures may induce nest desertion by
the male. That the shallow littoral areas of Tadenac
Lake are suitable for Largemouth Bass reproduction
is confirmed by the successful hatching of embryos
and the subsequent production of young-of-the-year
fish in both years of study, by the presence of all
year-classes from 1968 through 1975 in the trap net
fishery, and by the continuous, although modest,
sport fishery over the past century.

The ability of small oligotrophic lakes on the Pre-
cambrian Shield to sustain natural Largemouth Bass

populations probably depends on the extent of warm
littoral areas and the abundance of aquatic macro-
phytes. In typical northern oligotrophic lakes these
conditions occur only in shallow backbays sheltered
from wind action, where a rapid warm-up of water in
the spring and early summer provides 18°C or higher
temperatures satisfactory for reproduction while
more open epilimnetic waters remain comparatively
cold. Fry and Watt (1957) noted, for example, a uni-
que population of Largemouth Bass present in Tho-
mas Bay (45°30’N) on the south Georgian Bay shore
of Manitoulin Island, and explained its presence by
the more sheltered location and consistently warmer
water than elsewhere in the locality. While there are
numerous small eutrophic lakes within the Precam-
brian Shield thermally suitable for the Largemouth
Bass, these may be too shallow to prevent diel or
winter anoxia.

Although Largemouth Bass populations in oligo-
trophic Precambrian lakes are low, growth rates of
individual fish are not suppressed (Maraldo and
MacCrimmon 1979) and localized biomass may be
adequate to support limited sport fishing, as in
Tadenac Lake. In fact, the very limited Largemouth
Bass habitats of these lakes are generally similar to the
more generous environments prevailing in more
southerly eutrophic waters. Thus it would appear that
the extent of suitable habitat limits Largemouth Bass
production at more northerly latitudes.

306

Acknowledgments

We thank the Tadenac Club Limited for providing
access to their facilities and for financial assistance,
Barra L. Gots for excellent technical support, and
R. B. MacGregor for cooperation in the field. A
research grant awarded to H. R. MacCrimmon from
the National Research Council of Canada helped fund
the study.

Literature Cited

Allan, R. C., and J. Romero. 1975. Underwater observa-
tions of Largemouth Bass spawning and survival in Lake
Mead. /n Black bass biology and management. Edited by
H. Clepper, Sport Fishing Institute, Washington, D.C.,
USA. pp. 104-112.

Badenhuizen, T. R. 1969. Effects of incubation tempera-
ture onsurvival of Largemouth Bass embryos. M.S. thesis,
Cornell University, Ithaca, New York, USA. 88 p.

Carlander, K. D. 1977. Handbook of freshwater fishery
biology, Volume 2. The Iowa State University Press,
Ames, Iowa, USA. 431 p.

Casselman, J. M., and H. H. Harvey. 1973. Fish traps of
clear plastic. Progressive Fish-Culturist 35: 218-220.

Chew, R. L. 1974. Early life history of the Florida Large-
mouth Bass. Florida Game and Freshwater Fisheries
Commission, Bulletin 7. 76 p.

Coutant, C. C. 1975. Responses of bass to natural and arti-
ficial temperature regimes. /n Black bass biology and
management. Edited by H. Clepper. Sport Fishing Insti-
tute, Washington, D.C., USA. pp. 272-285.

Fry, F.E.J., and K.E.F. Watt. 1957. Yields of year
classes of the Smallmouth Bass hatched in the decade of
1940 in Manitoulin Island waters. Transactions of the
American Fisheries Society 85: 135-143.

Hasler, A. D., and W. J. Wisby. 1958. The return of dis-
placed Largemouth Bass and Green Sunfish to a “home
area.” Ecology 39: 289-293.

Heidinger, R. C. 1975. Life history and biology of the Lar-
gemouth Bass. /n Black bass biology and management.
Edited by H. Clepper. Sport Fishing Institute, Washing-
ton, D.C., USA. pp. 11-20.

Kelly, J. W. 1968. Effects of incubation temperature on
survival of Largemouth Bass eggs. Progressive Fish-
Culturist 30: 159-163.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Kramer, R.H., and L. L. Smith, Jr. 1962. Formation of
year classes in Largemouth Bass. Transactions of the
American Fisheries Society 91: 29-41.

MacKay, H.H. 1963. Fishes of Ontario. Ontario Depart-
ment of Lands and Forests, Toronto, Canada. 300 p.

Mahon, R., andE. K. Balon. 1977. Ecological fish produc-
tion in Long Pond, a lakeshore lagoon on Long Point,
Lake Erie. Environmental Biology of Fishes 2: 261-284.

Maraldo, D.C. 1978. Ecology of Largemouth Bass
Micropterus salmoides Lacépéde in a Precambrian Shield
Lake. M.Sc. thesis, University of Guelph, Guelph, Can-
ada. 120 p.

Maraldo, D. C., and H. R. MacCrimmon. 1979. Compari-
son of ageing methods and growth rates for Largemouth
Bass, Micropterus salmoides Lacépéde, from northern lat-
itudes. Environmental Biology of Fishes 4: 263-271.

Miller, K.D., and R.H. Kramer. 1971. Spawning and
early life history of Largemouth Bass, Micropterus sal-
moides, in Lake Powell. /n Reservoir fisheries and limnol-
ogy. Edited by G. E. Hall. American Fisheries Society
Special Publication 8. pp. 78-83.

Mraz, D. 1964. Observations on Large and Smallmouth
Bass nesting and early life history. Wisconsin Conserva-
tion Department, Fish Research Report 11. 13 p.

Mraz, D., S. Kmiotek, and L. Frankenberger. 1961. The
Largemouth Bass. Its life history, ecology and manage-
ment. Wisconsin Conservation Department, Publication
2322 i3ip:

Ricker, W. E. 1975. Computation and interpretation of
biological statistics of fish populations. Fisheries Research
Board of Canada, Bulletin 191. 382 p.

Robbins, W. H., and H. R. MacCrimmon. 1974. The Black
Bass in America and overseas. Biomanagement and
Research Enterprises, Sault Ste. Marie, Canada. 187 p.

Siler, J. R., and J.P. Clayton. 1975. Largemouth Bass
under conditions of extreme thermal stress. /n Black Bass
biology and management. Edited by R. H. Stroud and H.
Clepper. Sport Fishing Institute, Washington, D.C.,
USA. pp. 333-341.

Swallow, W. H. 1968. The relation of incubation tempera-
ture to the mortality of fish embryo. M.S. thesis, Cornell
University, Ithaca, New York, USA. 45 p.

Received 16 October 1980
Accepted 12 March 1981

Forestry Management Practices and Populations of Breeding Birds
in a Hardwood Forest in Nova Scotia

B. FREEDMAN, C. BEAUCHAMP, I. A. MCLAREN, and S. I. TINGLEY

Department of Biology and Institute for Resource and Environmental Studies, Dalhousie University, Halifax, Nova Scotia
B3H 4J1

Freedman, B., C. Beauchamp, I. A. McLaren, and S. I. Tingley. 1981. Forestry management practices and populations of
breeding birds in a hardwood forest in Nova Scotia. Canadian Field-Naturalist 95(3): 307-311.

Effects of forestry practices on the breeding birds of a hardwood forest in Nova Scotia were studied. T otal breeding density on
three uncut control plots was (x + SD) 663 + 145 pairs/km?. Total breeding densities on three plots clear-cut 3-5 yr earlier
were somewhat lower (588 + 155 pairs/km2), as were the densities on one thinned plot (550 pairs/ km?) and on two strip-cut
plots (475 and 575 pairs/km2), but differences between treatments were not statistically significant.

Marked differences, however, occurred in the species composition of the bird communities in the various plots. The most
important breeding species on the control plots were Least Flycatcher (Empidonax minimus) and Ovenbird (Seiurus
aurocapillus). In the clear-cut plots the bird community was dominated by Chestnut-sided Warbler (Dendroira pensylvanica),
Common Yellowthroat (Geoth/ypis trichas), and White-throated Sparrow (Zonotrichia albicollis). The species mixtures of

the thinned and strip-cut plots were intermediate between those of the control and clear-cut plots.

Key Words: forest birds, breeding density, hardwood forest, silviculture.

The forest-related industries are among North
America’s largest. Nevertheless, there are relatively
few studies in which the effects of forestry on wildlife
communities have been examined, especially for those
perceived as being of little economic significance, such
as small mammals and songbirds. Recent studies of
changes in populations of breeding birds indicate that,
for many types of coniferous forest communities,
intensive harvest techniques, such as clear-cutting,
generally cause short-term decreases in both abun-
dance and breeding diversity, followed by rapid re-
covery to abundance and diversity levels that fre-
quently exceed those of uncut control forests (recently
reviewed in Crawford and Titterington 1979; Scott et
al. 1980; Smith 1980). The rates and degrees of recov-
ery in various coniferous forest types have been
related to several factors, especially the rates of
revegetation and the habitat diversity within the
regenerating forest. Far fewer studies have examined
the effects of forestry on the breeding birds of hard-
wood forests, and we are aware of only one study that
has investigated such forests in the northeast (Webb et
al. 1977). In addition, we are aware of only one study
(Welsh and Fillman 1980) that was done in habitats
modified by logging or silviculture in Canada, in spite
of the large areas of land that are disturbed here each
year.

In this paper we examine the changes in popula-
tions of breeding birds that occurred subsequent to
various forest management practices in a hardwood
forest in Nova Scotia.

Methods

All study plots were located in Kings County, Nova
Scotia, near 44°50’N, 65°44’W. This falls within the
Red Spruce-Hemlock-Pine zone (Picea rubra —
Tsuga canadensis — Pinus sp.) of Louck’s (1962) clas-
sification of maritime forests. However, the particular
forest that we studied originated after a fire 70 years
ago, and is now dominated by a mixture of northern
hardwood species, especially Red and Sugar Maple
(Acer rubrum and A. saccharum) and White and Yel-
low Birch (Betula papyrifera and B. allegheniensis).

Our study area has been subjected to experimental
forestry manipulations, including strip clear-cuts of
two widths, and silvicultural thinning. These were
intended to examine effects on regeneration and
growth of desirable tree species. In addition, opera-
tional clear-cutting of patches of forest had been done
in the immediate vicinity. Thus, we were able to census
breeding birds of hardwood forests in patch clear-
cuts, strip-cuts, silvicultural thins, and control, uncut
forests. All treatments were located within 3 km of
each other, in a fairly homogenous hardwood forest.

All plots were rectangular and were laid out by
compass and chain. The perimeter of each plot was
marked at 30-m intervals with flagging tape, and all
but the relatively small thinned treatment plot had a
transect through the long axis of the plot, which was
also flagged at 30-m intervals. The sizes of the plots
were determined by the sizes of the various silvicultu-
ral treatments examined: the sizes and shapes of the
control, uncut plots were similar to those of the patch

307

308

clear-cut plots. The thinned plot had been selectively
cut to 45% residual basal area. The wider strip-cut plot
had two 30-m-wide clear-cut strips; 45% of the plot
was cut, leaving 55% in one central uncut strip. The
other strip-cut plot had three 20-m-wide cut strips:
38% of the plot was cut, leaving 62% in two central
residual strips. Relative amounts of cut and uncut
habitat on the strip-cut plots was somewhat artificial,
and we might have obtained different results to those
described below if we had, for example, included addi-
tional uncut strips on the edges of our plots.

Birds on each plot were surveyed 10 times between
29 May and 9 July 1980. Eight censuses were made
between dawn and 11:30, and two between 19:00 and
21:50. The spot-map method was used to census birds
(after Williams 1936; Robbins 1978). We calculated
the densities of breeding bird species having at least
0.5 territories on a plot. Species richness was esti-
mated as the number of breeding species having at
least 0.5 territories per plot. Some scarcer species may
have been eliminated by this conservative treatment of
richness, if they held large territories or were seldom
observed.

Quantitative measurements of the plant communi-
ties were made in five evenly spaced 20 X 20 m quad-
rats per plot. Species and diameter at breast height of
all trees on the plots were recorded. From these mea-
surements, we calculated tree density (stems/ ha), stem
basal area (m2/ha), and frequency of occurrence for
individual tree species, and summed for the plot as a
whole. Canopy height was measured on dominant
trees using a clinometer. Shrubs in 10 evenly spaced
5 X 5 m quadrats per plot were tallied as to species
and number of individual stems, from which we calcu-
lated species and plot shrub densities (stems/ ha). The
percent ground cover of the various ground flora spe-
cies was estimated in twenty | X | m quadrats per
plot. Owing to foliage overlap, some plots had ground

THE CANADIAN FIELD-NATURALIST

Vol. 95

cover values greater than 100%. In addition, at each of
the 20 points where ground cover was measured, esti-
mates were made of the percent canopy cover, by
sighting vertically through a paper cylinder (4.2 cm
diameter, 10.5 cm length), and estimating the surface
area covered by tree foliage. These methods were
modified from James and Shugart (1970).

Scientific names of birds are given in Appendix 1.

Observations and Discussion

The characteristics of the vegetation of the various
plots are summarized quantitatively in Tables | and 2.
Qualitatively, the vegetation of the control plots was
dominated by the overstory, with only moderate
shrub and ground flora contributions. Conversely, the
clear-cut plots had virtually no overstory (a very low
density of unmerchantable trees had been left stand-
ing), and the vegetation was dominated by shrubs
(mainly stump sprouts of Acer spp., Betula papyrif-
era, and Viburnum cassinoides (Raisinbush), seed-
lings of Prunus pensylvanica (Pin Cherry), and
thickets of Rubus spp. (raspberry), and by a relatively
rich ground flora, compared with the uncut plots. The
vegetation of the thinned plot was intermediate to that
of the control and clear-cut plots. The overstory was
composed of relatively large and well-spaced stems of
Acer saccharum and Fraxinus americana (White
Ash). These produced a canopy cover only about
one-third of the control average. On the other hand,
the density, species composition, and cover of the
shrub and ground flora strata were roughly compara-
ble to that of the clear-cut plots. The vegetation of the
strip-cut plots was also intermediate to that of the
control and clear-cut plots, although of course the
vegetation occurred in discrete strips of uncut forest,
and shrub-dominated cut strips.

The breeding bird census data are summarized in
Table 3. Note that data for the three control plots and

TaBLe 1—Characteristics of the uncut mixed-species hardwood forest where the census plots were located, based on five

20 X 20 m quadrats in each of the three control plots

Relative Relative Relative
Density Basalarea Frequency density dominance frequency Importance
Species (trees/ ha) (m?/ ha) (%) (%) (%) (%) value!
Betula papyrifera 470 8.08 100 PUY SiEZ 17.9 76.8
Acer rubrum 412 7.18 93 24.3 27.8 16.6 68.7
Acer saccharum 265 4.65 67 15.6 18.0 12.0 45.6
Betula allegheniensis 203 2.04 67 12.0 We) 12.0 31.9
Fagus grandifolia? 50 0.24 60 2.9 0.9 10.7 14.5
Fraxinus americana ay les 2 13 1.6 Doll D3 9.0
Other species 49 0.72 60 Bel 2.8 10.7 16.6
Dead trees ANY 1.63 100 12.8 6.3 17-9 37.0
Total 1693 25.86 — = = i a

'Sum of relative density + relative dominance + relative frequency.

2Beech.

1981

FREEDMAN ET AL.: BREEDING BIRDS IN HARDWOOD FOREST

TABLE 2—Characteristics of the vegetation of the plots censused for breeding bird populations

Plot Canopy
Plot Age area height
Treatment number (yr) (ha) (m)
Control 3 ~70 3.3 of A19
6 ~70 5.6 16
8 ~70 3.4 17
Clear-cut l 3 4.1 0
7 3 3.4 0
5 5 3.4 0
Thin 9 5 1.8 23
Narrow Strip-cut 2 4 6.2
(a) Cut strips 0
(b) Uncut strips 19
Wider Strip-cut 4 4 2.9
(a) Cut strips 0
(b) Uncut strips 16

for the three clear-cut plots were reported in combined
form elsewhere (Freedmanet al. 1981). Interms of the
total density of breeding birds, the three control plots
(mean + SD= 663 + 145 pairs/km?) fell near the
middle of the range of densities for eight recent breed-
ing bird censuses in maple or birch-dominated hard-
wood forests of the northeastern United States or
eastern Canada (mean + SD = 635 + 328 pairs/km2?,
range = 260-1200 from 1979 censuses reported in:
American Birds, Vol. 34, No. 1). The breeding bird
community in the uncut forest was dominated by such:
species as Least Flycatcher, Hermit Thrush, Solitary
Vireo, Red-eyed Vireo, and various wood warblers,
including Black-and-white Warbler, Northern Par-
ula, Black-throated Green Warbler, American Red-
start, and Ovenbird (Table 3).

The total breeding bird densities of the clear-cut
plots fell within the range of those of the control plots
(mean + SD = 588 + 155 pairs/km?). Species _rich-
ness, however, averaged less (12.3 = 3.5 species per
control plot vs. 8.3 = 1.5 species per clear-cut plot). In
addition, there were great differences in species com-
position. In the clear-cut (low shrub) plots the com-
munity was dominated by Chestnut-sided Warbler,
Common Yellowthroat, Dark-eyed Junco, White-
throated Sparrow, and Song Sparrow.

The total density of the thinned plot was
550 pairs/km2, which was within the range of densi-
ties of both the clear-cut and the control forest. The
low richness of only seven species occurring on this
plot may in part be an artifact of the small plot size.
The thinned plot contained species that also occurred
either in the control forests or the clear-cut plots,
notably Least Flycatcher and Chestnut-sided
Warbler.

The total breeding densities of the strip-cut plots
were 475 pairs/ km? for the narrower strip-cuts, and

309
Tree Stem basal Canopy Shrub Ground
density area cover density cover

(stems/ha) (m2/ha) (%) (stems/ ha) (%)
1800 28.3 87 3 200 50
1400 19:7 73 7300 75
1900 29.6 id 2 400 60
0 0 0 29 400 120
0 0 0 23 600 100
0 0 0 36 800 170
460 13.3 26 41 900 160
0 0 0 40 600 160
1400 26.1 73 6200 100
0 0 0 29 200 180
1600 20.7 81 9 100 60

575 pairs/ km? for the wider strip-cuts. Both of these
were lower than the mean total density observed for
the control plots, although only the density for the
narrow strip-cut fell below the range observed among
the three control plots. Species richness of 10 and 13
species per plot was only slightly lower than that
observed on the control plots. Similarly to the thinned
plot, the bird species of the strip-cut plots included
those dominant in either the control or clear-cut plots.
Forest birds, such as Least Flycatcher, Red-eyed
Vireo, Ovenbird, and American Redstart were present
on the strip-cut plots, as were species typical of the
clear-cuts, such as Chestnut-sided Warbler and
Common Yellowthroat. We stress that, because of the
small size of our plots, we only have reliable data for
breeding birds that have relatively small territories.
Other species that are characterized by larger territory
size, such as woodpeckers or corvids, might also be
affected by these forestry manipulations, but were not
satisfactorily censused in our study.

Analysis of the total densities of birds among plots
showed no significant heterogeneity (chi-square test,
P> 0.05).

Only in one other study have the impacts of forestry
operations on populations of breeding birds in north-
eastern hardwood forests been examined (Webbet al.
1977). These authors examined stands which had 0,
25, 50, 75, or 100% of the merchantable volume
removed, and used counts of the numbers of singing
males heard at permanent sampling points as an index
of breeding bird abundance. Webb et al. (1977)
observed no significant changes in breeding bird
abundance. There were, however, changes in species
composition on the various plots, the most notable of
which was a decline in Ovenbird abundance in the
75% and 100% removal treatments.

In summary, we found few differences in the total

310

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 3—Densities of breeding birds in plots variously affected by cutting practices in a mixed hardwood forest in Nova
Scotia. Only species having at least 0.5 territorial pairs on a plot are indicated

Control plots

Species 3 6 8
Common Snipe 0 0 0
Ruby-throated Hummingbird 0 0 0
Alder Flycatcher 0 0 0
Least Flycatcher 290 120 0
Eastern Wood Pewee 0 10 0
Tree Swallow 0 0 0
American Robin 0 20 0
Hermit Thrush 60 40 30
Veery 50 10 0
Solitary Vireo 60 30 0
Red-eyed Vireo 80 50 30
Black-and-White Warbler 15 50 40
Northern Parula Warbler 15 30 40
Black-throated Blue Warbler 0 0 30
Black-throated Green Warbler 50 30 30
Chestnut-sided Warbler 0 0 0
Ovenbird 150 120 200
Northern Waterthrush 0 20 0
Mourning Warbler 0 0 0
Common Yellowthroat 0 0 0
American Redstart 15 80 100
Rusty Blackbird 0 0 0
Rose-breasted Grosbeak 15 10 0
Dark-eyed Junco 15 20 15
White-throated Sparrow 0 20 0
Song Sparrow 0 0 0
Total 815 660 515
Richness 12 16 9

density or richness of the breeding birds of a hard-
wood forest that was affected by several forestry prac-
tices, including clear-cutting, strip-cutting, and thi-
ning. However, there were marked species changes,
with obligate forest birds being replaced by early suc-
cessional species on the clear-cuts. The strip-cuts and
thin, being intermediate in vegetation structure, also
had intermediate mixtures of breeding birds.

Acknowledgments

We gratefully acknowledge the field assistance of J.
Beckwith, J. Dale, A. Duinker, O. Maessen, S.
Manuel, J. McLaren, and R. Morash. A. J. Erskine
helped with the interpretation of some bird territories.
Funding for this work was provided by a research
grant to B.F. and I. A.M. from the Canadian
National Sportsmen’s Fund.

Literature Cited

Crawford, H.S., and R. W. Titterington. 1979. Effects of
silvicultural practices on bird communities in upland
spruce-fir stands. Jn Management of north central and

Density (pairs/ km?)

Clear-cut plots Thinned plot Strip-cut plots

l 7 5 9 2 4
10 15 0 0 0 0
25 30 15 0 0 0
0 0 30 0 0 0
0 0 0 140 60 100
0 0 0 0 0 0
10 0 0 0 0 0
10 0 0 0 0 0
0 0 0 0 15 0
25 0 0 0 15 0
0 0 0 0 15 0
0 0 0 80 30 70
0 0 0 0 70 15
0 0 0 0 10 40
0 0 0 0 0 20
0 0 0 0 15 0
100 40 190 110 50 70
0 0 0 0 60 100
0 0 0 0 0 0
0 0 90 80 20 0
25 300 130 80 0 50
0 0 0 30 90 90
0 30 0 0 0 0
0 0 0 0 0 0
50 70 30 0 5) 20
90 190 100 30 0 0
90 70 0 0 0 0
435 745 585 550 475 575
10 8 7 7 13 10

northeastern forests for nongame birds. USDA Forest
Service General Technical Report NC-51, North Central
Forest Experiment Station, St. Paul, Minnesota. pp.
110-119.

Freedman B., C. Beauchamp, I.A. McLaren, and S.I. Ting-
ley. 1981. Maple-birch forest; Maple—birch forest, 5-
year clear-cut. American Birds 35: 50, 71.

James, F.C., and H.H. Shugart. 1970. A quantitative
method of habitat description. American Birds 24:
727-736.

Loucks, O. L. 1962. A forest classification for the maritime
provinces. Proceedings of the Nova Scotia Institute of
Science 25: 85-167.

Robbins, C. S. 1978. Census techniques for forest birds. /n
Management of southern forests for nongame birds.
USDA Forest Service General Technical Report SE-14,
Southeastern Forest Experiment Station, Asheville,
North Carolina. pp. 142-163.

Scott, V.E., J. A. Whelan, and P.L. Svoboda.
1980. Cavity-nesting birds and forest management. /n
Management of western forests and grasslands for non-
game birds. USDA Forest Service General Technical
Report INT-86, Intermountain Forest and Range Exper-
iment Station, Ogden, Utah. pp. 311-324.

1981

Smith, K. G. 1980. Nongame birds of the Rocky Mountain
spruce-fir forests and their management. Jn Management
of western forests and grasslands for nongame birds.
USDA Forest Service General Technical Report INT-86,
Intermountain Forest and Range Experiment Station,
Ogden, Utah. pp. 258-279.

Webb, D. L., D. F. Behrend, and B. Saisorn. 1977. Effect
of logging on songbird populations in a northern hard-
wood forest. Wildlife Monographs, Number 55. 35 pp.

Welsh, D.A., and D.R. Fillman. 1980. The impact of
forest cutting on boreal bird populations. American Birds
34: 84-94.

Williams, A. B. 1936. The composition and dynamics of a
beech-maple climax community. Ecological Monographs
6: 317-408.

Received 21 November 1980
Accepted 6 March 1981

APPENDIX 1—Scientific names of all bird species mentioned
in the text

Common Snipe
Ruby-throated Hummingbird

Capella gallinago
Archilochus colubris

FREEDMAN ET AL.: BREEDING BIRDS IN HARDWOOD FOREST

Alder Flycatcher

Least Flycatcher

Eastern Wood Pewee
Tree Swallow

American Robin

Hermit Thrush

Veery

Solitary Vireo

Red-eyed Vireo
Black-and-white Warbler
Northern Parula Warbler
Black-throated Blue Warbler
Black-throated Green Warbler
Chestnut-sided Warbler
Ovenbird

Northern Waterthrush
Mourning Warbler
Common Yellowthroat
American Redstart
Rusty Blackbird
Rose-breasted Grosbeak
Dark-eyed Junco
White-throated Sparrow
Song Sparrow

StI

Empidonax alnorum
Empidonax minimus
Contopus virens
Tridoprocne bicolor
Turdus migratorius
Catharus guttatus
Catharus fuscescens
Vireo solitarius

Vireo olivaceous
Mniiotilta varia

Parula americana
Dendroica caerulescens
Dendroica virens ;
Dendroica pensylvancia
Seiurus aurocapillus
Seiurus noveboracensis
Oporornis philadelphia
Geothlypis trichas
Setophaga ruticilla
Euphagus carolinus
Pheucticus ludovicianus
Junco hyemalis
Zonotrichia albicollis
Melospiza melodia

Breeding, Feeding, and Chick Growth of the Black Guillemot
(Cepphus grylle) in Southern Quebec

DAVID CAIRNS

Département de biologie, Université Laval, Québec, Québec GIK 7P4
Present address: Department of Biology, Carleton University, Ottawa, Ontario KIS 5B6

Cairns, David. 1981. Breeding, feeding, and chick growth of the Black Guillemot (Cepphus grylle) in southern Quebec.

Canadian Field-Naturalist 95(3): 312-318.

Black Guillemots (Cepphus grylle) at a colony in the estuary of the St. Lawrence River bred about 3 wk earlier than those of
another in the northeastern Gulf of St. Lawrence; mean hatching dates were 26 June and 15 July, respectively. Chicks spent a
mean of 36.5 din the nest in both colonies, and fledged at about 88% of adult weight. Four chicks were observed to fledge with
no parental involvement. Chick diets consisted largely of blennies and other littoral fishes. The daily feeding rhythm was
bimodal, with mid-morning and mid-afternoon peaks. Hatching and fledging rates per egg laid were much lower in single-egg
nests than in two-egg nests. Chick mortality was highest during the first few days after hatching and affected first and second

chicks equally.

Key Words: Black Guillemot, Cepphus grylle; breeding chronology, feeding, fledging, Gulf of St. Lawrence.

The Black Guillemot (Cepphus grylle) is a pursuit-
diving sea bird of inshore arctic and northern Atlan-
tic waters. Breeding sites are extensively distributed
within this range, and most take the form of small
loose colonies, although some birds nest solitarily,
and some large arctic colonies are known (Brownet al.
1975).

Most currently available data on the life history of
the Black Guillemot come from near the southern
margin of its range, in the Bay of Fundy (Winn 1950;
Preston 1968) and northeastern Denmark (Asbirk
1979 a, b). The colonies examined in these areas are
limited in population by number and quality of nest-
ing cavities, and are thus probably atypical of most
guillemot breeding sites.

In the present paper I describe breeding chronolo-
gies, chick diets and growth rates, and patterns of
nesting success in two eastern Canadian colonies.
Breeding habitat was available in excess in both
locations — a fact significant to breeding phenology
and to the relation between habitat morphology and
nesting success (Cairns 1980).

Study Area and Methods

This study was conducted on Brandypot Island
(47°52’N, 69°41’W) in the St. Lawrence estuary, 21
April to 23 August 1976, and onthe St. Mary’s Islands
(50° 18’N, 59°39’W) in the northeastern Gulf of St.
Lawrence, 6 May to 25 August 1977.

Brandypot Island lies roughly midway between the
north and south banks of the St. Lawrence, at a point
where the estuary is 27 km wide. Tidal amplitude in
the area is about 5 m, and the brackish waters have a
surface salinity of about 20 °/oo (Lavoie and Beaulieu
1971). Brandypot consists of three wooded islets

underlain by quartzite and conglomerate, which are
exposed at the shoreline as rocky slopes, cliffs, or
boulder heaps. About 200 breeding pairs of Black
Guillemots share the island with nesting Double-
crested Cormorants (Phalacrocorax auritus), Great
Blue Herons (Ardea herodias), Black-crowned Night
Herons (Nycticorax nycticorax), Black Ducks (Anas
rubripes), Common Eiders (Somateria mollissima),
Great Black-backed Gulls (Larus marinus), Herring
Gulls (L. argentatus), and Razorbills (Alca torda).

The four islands of the St. Mary’s chain lie 11 km
off the Quebec North Shore of the Gulf of St. Law-
rence, inan area influenced by cold Labrador Current
waters entering the Gulf through the Strait of Belle
Isle. Tidal range in the area is about | m. Central parts
of the islands are covered by low heath vegetation, but
there are broad rocky slopes around their peripheries.
West and Middle Islands, where the study was con-
ducted, harbor about 145 breeding pairs of Black
Guillemots. Red-throated Loons (Gavia stellata),
Common Eiders, Great Black-backed Gulls, Herring
Gulls, Common Terns (Sterna hirundo), Arctic Terns,
(S. paradisaea), Razorbills, Common Murres (Uria
aalge), and Atlantic Puffins (Fratercula arctica) also
breed on these islands. Further physical and avifaunal
details are given by Reed (1973) for Brandypot and by
Bédard (1969) and Chapdelaine (1980) for the St.
Mary’s Islands.

Data on nesting success and chick growth rates
were gathered during nest visits spaced 4-5 d apart on
Brandypot, except for a small number of easily access-
ible nests which were checked daily. On St. Mary’s,
nests were visited daily in two high-density subcolo-
nies, and at intervals of 4 d elsewhere.

Chicks were weighed with Pesola spring balances.

S12

1981

Chick diets were examined on Brandypot by collect-
ing fish from the vicinity of the nests, and by constrict-
ing chicks’ throats with pipe cleaners for periods of
3-6 h to prevent them from swallowing food items
delivered by their parents. At St. Mary’s, food items
were identified and counted visually, and fledging
behavior was observed, from a blind that gave a view
of six active nests. Feeding and fledging times are
given in Atlantic Standard Time.

Results
Breeding Phenology and Behavior

Black Guillemots were present in the waters around
both study islands by the beginning of field work; I
counted 400 around Brandypot on 26 April 1976, and
185 around St. Mary’s on8 May 1977. On Brandypot,
birds were first observed on the rocks below the breed-
ing sites on 25 April, with the first copulations
recorded on 29 April. These events occurred over 2 wk
later on St. Mary’s where birds were first seen on land
on 12 May, and the first copulation noted on 17 May.

Social interactions on the water were frequent dur-
ing the prelaying period and most often took the form
of pursuits. On Brandypot, the birds took part in
massed communal flights, involving up to several
hundred individuals and lasting several minutes. Dur-
ing these flights the guillemots often rose in tight
formation up to about 150 m above the water.

After the first landings, the guillemots gradually
spent more time on shore, and by the beginning of

St. Mary's

[_] Brandypot

IS
Oo

= as Dy)
fe) On Oo

Number of clutches hatchin

[Ga ZOOM Zee Sire.
June

CAIRNS: BLACK GUILLEMOT BREEDING

313

egg-laying the diurnal attendance patterns displayed
through the remainder of the breeding season were
established. On Brandypot, attendance was higher in
the morning but peaked only at high tides, whereas on
St. Mary’s attendance peaked regularly in the early
morning and late evening (Cairns 1979).

The nesting cycle was about 3 wk earlier on Bran-
dypot than on St. Mary’s (Figure 1); mean hatching
dates were 26 June (n= 118) and 15 July (n= 111) in
the two areas, respectively. The duration of the nest-
ling period was similar at both sites and its combined
mean was 36.5 d (SD = 1.5, n= 16, range 34-39).

Fledging of chicks was observed on four occasions
at St. Mary’s. All departures took place well after
sunset, between 19:35 and 20:30. Chick exits were
preceded by considerable movement within the nest
cavity, which I could hear from the blind, and chicks
were observed briefly at the nest entrance up to three
times before their final departures. After leaving the
nest, the fledglings made their way to the water 40 to
50 m away in bouts of scrambling and flapping that
alternated with rest periods of up to 10 min. None of
the fledglings was capable of sustained flight, and
their flapping leaps usually carried them no more than
ore mae

On Brandypot most fledglings must have left the
vicinity of the colony immediately after leaving the
nest, because only two birds in juvenal plumage were
ever seen on the water there. These birds probably
dispersed towards the lower St. Lawrence estuary,
because bands from four birds I had banded as chicks

ii 22° 26 30 © fa
July August

lO ea:

FIGURE |. Black Guillemot hatching dates on Brandypot Island in the St. Lawrence estuary (1976) and on the St. Mary’s

Islands in the Gulf of St. Lawrence (1977).

314 THE CANADIAN FIELD-NATURALIST

TABLE 1—Food items delivered to Black Guillemot chicks

Prey

Crustacea
Osmeridae

Capelin (Mallotus villosus)
Gadidae

Atlantic Tomcod (Microgadus tomcod)
Ammodytidae

American Sand Lance (Ammodytes americanus)
Blennioidea

Rock Gunnel (Pholis gunnellus)

Rock Gunnel/ Wrymouth (Cryptacanthodes maculatus)*

Arctic Shanny (Stichaeus punctatus)
Polar Eelpout (Lycodes turner!)
Ocean Pout (Macrozoarces americanus)
Unidentified
Cottidae
Arctic Staghorn Sculpin (Gymnocanthus tricuspis)
Grubby (Myoxocephalus aeneus)
Mailed Sculpin (Trig/lops murrayi)
Unidentified

Vol. 95
Brandypot St. Mary’s
No. % No. %
— — 2 I
2 15 ——
] 8 119 33
I 8 2 8
2 15
65 18
_— 101 28
8 =
3 23 =
_ 20 6
8 ee
8 wey
1 8 —
— 28 8

“These species could not be reliably distinguished from the blind.

on Brandypot were returned from this area from late

August to October 1976. Adult birds left the Brandy-

pot area simultaneously with the young.
Postfledging dispersal of adults and young from St.

Meals delivered: chick '. h!

04:00

06:00 08:00 10.00

12:00 14:00 16:00 18.00

Mary’s had not begun by 26 August. Juvenile birds
were regularly seen on the water after the onset of
fledging, and no reduction in the number of adult
birds in the area was noted during the fledging period.

20.00

Time

FIGURE 2. Daily rhythm of food deliveries to Black Guillemot chicks on St. Mary’s, based on 20 to 36 chick-hours of
observation for each hour. No observations available between 12:00 and 13:00.

1981

Feeding of the Chick

Thirteen fish were collected from Black Guillemot
nests at Brandypot. At St. Mary’s 362 food deliveries
to six nests were recorded during 51 h of observation
between 2 and 17 August. Chicks in the observed nests
were aged 11 d to fledging. Prey items observed or
collected are listed in Table 1. All food loads that were
clearly seen contained a single fish, except two on St.
Mary’s in which the prey appeared to bea shrimp-like
crustacean. Eight percent of food deliveries on St.
Mary’s were not identified, but in most cases this was
due to my attention being drawn by the simultaneous
arrival of a food-carrying adult at another nest. It is
therefore likely that most unidentified prey belonged
to species observed at other times. About half the food
items from both study areas were Blennioidea, small
mainly benthic fishes. At St. Mary’s, Atlantic Tomcod
(Microgadus tomcod) and American Sand Lance
(Ammodytes americanus) comprised most of the
remainder of the sample.

Several of the guillemots in the St. Mary’s feeding
observation area were banded and could be distin-
guished from their mates, allowing the feeding patterns
of individual birds to be monitored. A total of 127
food deliveries by eight recognizable birds was
recorded. The frequency distributions of prey items
delivered by these birds differed markedly (P < 0.005,
G-test), which suggests that Black Guillemots have

e St Marys

400 a Brandypot

300

200

Weight (g/

100

CAIRNS: BLACK GUILLEMOT BREEDING

315

individual preferences either of prey species or of feed-
ing area. One guillemot completely specialized in
lance and was not observed to deliver any other prey
(n = 17).

The daily rhythm of food deliveries on St. Mary’s
was bimodal with a strong peak in the morning and a
lesser one in the afternoon (Figure 2). Rates of food
delivery were 0.70 items:chick '-h'! for two-chick
broods, and 0.95 items-chick'-h! for singletons
(P<0.025, G-test), with an overall rate of 0.79
items:chick '-h |.

Terns nested as close as a few metres from Black
Guillemot sites on St. Mary’s, as did gulls on both
study islands. However, kleptoparasitism was infre-
quent, and of the four attacks I witnessed (all on St.
Mary’s), only one, by a Great Black-backed Gull, was
successful.

Growth of the Chick

Weight curves for 23 chicks on Brandypot and 45
chicks on St. Mary’s show that growth in both areas
was similar (Figure 3). Because the chicks in most
cases were not weighed daily, the individuals repre-
sented in each age-class varied from day to day. This
variation in the sample content is likely responsible
for most of the irregularity in the curves, particularly
near fledging, when sample sizes were smallest.

Mean weights SD (n) of chicks measured within
24h of fledging were 356 = 10.6 g (7) at Brandypot

20 24 28 BZ 36 40

Age (days)

FIGURE 3. Growth in weight of Black Guillemot chicks. Number of chicks varied between 2 and 16 on Brandypot, and between

1 and 33 on St. Mary’s.

316

TABLE 2—Black Guillemot nesting success for one-egg (25
nests) and two-egg (187 nests) clutches (data from both study
areas). Differences tested by G-test (*P < 0.05; **P < 0.01).

Clutch size
One-egg Two-egg

Eggs hatched/ nest 0.32 Nelly
Eggs hatched/egg laid 0825 0.58*
Chicks fledged/ nest 0.12 0.78
Chicks fledged/ egg laid On 0.39**
Chicks fledged / chick hatched:

one-chick broods 0.38(8)° 0.63(32)°

two-chick broods — 0.67(93)

“Number of individuals in parentheses.

and 355 + 23.8 g(13) at St. Mary’s. Fledging weights
were about 88% of breeding adult weights
(401 + 14.9 g (7) for Brandypot, 407 + 31.8 (47) for
St. Mary’s).

Egg and Chick Survivorship

Mean clutch size was 1.88 in the 212 nests moni-
tored (Table 2). Production from single-egg nests was
reduced not only by the smaller number of eggs, but
also through lowered hatching and fledging rates.
Mean production from two-egg nests was 6.5 times
greater than for one-egg nests.

Most chick mortality occurred during the first few
days after hatching, and no deaths were recorded after
24d (Figure 4). Chick mortality was unrelated to
order of hatching: of 10 chicks found dead whose
hatching sequence was known, five hatched first and
five hatched second.

Discussion

The onset of breeding in sea birds is ultimately
determined by annual cycles of water temperature and
other oceanographic variables that control prey
abundance (Immelmann 1971; Salomonsen 1955).
These cycles only roughly correlate with atmospheric
trends; thus the I-mo lag in springtime air tempera-
ture curves between Brandypot and St. Mary’s cor-
responds to a similar delay in guillemot nesting,
although the guillemots of Kent Island in the Bay of
Fundy also nest later than those of Brandypot, despite
similar spring temperature regimes (Preston 1968;
weather data from Riviére-du-Loup, Harrington
Harbour, and Grand Manan show spring tempera-
tures close to long-term means during study years,
Atmostpheric Environment Service 1975, 1976-1977).

Despite their later breeding date, Kent Island guil-
lemots first come to land at roughly the same time as
Brandypot birds (Winn 1950; Preston 1968), presum-
ably because early arrivals have a better chance of
securing or retaining one of the colony’s small number
of good-quality nesting cavities.

THE CANADIAN FIELD-NATURALIST

Vol. 95

The occurrence of massed communal flights on
Brandypot Island is related to the fact that most of the
birds there bred in high density along a single strip of
straight coastline and were thus in visual contact with
each other while on the water below the breeding sites.
Such flights were not observed at St. Mary’s nor at
other colonies where guillemots breed in small sub-
colonies separated by headlands (Preston 1968;
Bergman 1971; Asbirk 1979a, b). Massed high-
altitude flights have rarely been reported for alcids,
which normally fly low over the water (Tuck’s (1961)
record of Common Murre flights is an exception).
Because such flights require a much greater energy
expenditure than normal flying, they probably have a
ritual function. Asbirk (1979b) speculated that com-
munal gatherings of guillemots on the water play a
role in establishing a social hierarchy that would
determine proprietorship of nest sites, but this is
unlikely to have been the case on Brandypot where
sites were available in excess.

According to Winn(1950) and Thoresen and Booth
(1958), the Black Guillemot and the closely related
Pigeon Guillemot (Cepphus columba) lure their

36

S2

28

24

20

Number of chicks found dead

O-4 5-9) 10-14 IS=I9

Age (days)

2ORee

FiGurReE 4. Age at death of Black Guillemot chicks (data from
both study areas).

198]

young from the nest by dangling a fish in front of the
nest entrance. I observed no adult behavior that could
be interpreted as an attempt to coax chicks from the
nest. Moreover, my observations of fledgings
occurred after the last feeding visits of the adults in the
evening, and adults on three occasions were observed
to bring fish to nests from which the young had
already fledged.

The Black Guillemot is basically a littoral species
and is seldom sighted far offshore (Brown etal. 1975).
Although the adults feed on both fishes and inverte-
brates (Belopol’skii 1957; Uspenski 1958; Hartley and
Fisher 1936), the young are fed almost exclusively on
fish. Small benthic Blennioidea were the chief prey on
Brandypot and St. Mary’s, as they are in the Bay of
Fundy (Winn 1950; Preston 1968), the Gulf of Finland
(Bergman 1971), and Denmark (Asbirk 1979a).

Individual feeding specialization as found on St.
Mary’s has been previously documented by Slater and
Slater (1972) for Black Guillemots and by Drent
(1965) for Pigeon Guillemots. Although colony
attendance patterns of Danish and St. Mary’s guille-
mots are very similar (Asbirk 1979a; Cairns 1979),
feeding rhythms differed markedly. The feeding
rhythm at St. Mary’s resembled the bimodal atten-
dance pattern, but feeding in Denmark was relatively
uniform throughout the day (Asbirk 1979a). Overall
feeding rates were lower in Denmark (Asbirk 1979a)
and higher in the Bay of Fundy (Preston 1968) than
those found on St. Mary’s but in the absence of relia-
ble measures of meal weights, valid comparisons of
food intake cannot be made.

Growth rates, fledging weights, and nestling peri-
ods of Brandypot and St. Mary’s chicks were similar
to those reported in the literature (Winn 1950; Bianki
1967; Belopol’skii 1957; Asbirk 1979a), although
Asbirk (1979a) found a substantially longer nestling
period (39.5 d vs. 36.5 d).

The normal clutch size of Black and Pigeon Guille-
mots is two eggs, although one-egg clutches are regu-
larly encountered. Production per egg laid is
markedly lower in one-egg nests than in two-egg nests
(Preston 1968; this study), and the difference is due to
depressed viability of both eggs and chicks. Because
breeding success on Brandypot and St. Mary’s was
not related to habitat structure or nesting density
(Cairns 1980), it can be inferred that production of
single-egg clutches, with subsequent lowered hatching
and fledging rates, is associated with couples of infe-
rior parental quality. Asbirk (1979a) found single eggs
to be laid more often in inexperienced breeders, and
breeding success has been negatively linked to paren-
tal weights in several sea-bird species (e.g., Coulson
1968). Furthermore, any weakness of parental drive
on the part of pairs that produced one-egg clutches
could be exacerbated by the depressing effect of

CAIRNS: BLACK GUILLEMOT BREEDING

37

observer disturbance on breeding success (Cairns
1980).

Acknowledgments

This study was funded by grants from the National
Research Council of Canada to J. Bédard, and by a
Canadian Wildlife Service grant-in-aid of university
research. G. Rochette provided invaluable field
assistance on Brandypot, and prepared the figures.
The Canadian Ministry of Transport generously per-
mitted the use of their facilities for personal lodging
on both Brandypot and St. Mary’s. G. Foreman and
L. Chislett, lightkeepers on St. Mary’s Islands, pro-
vided advice, assistance, and warm hospitality during
field work there. A. Cairns, W. Cairns, and D.N.
Nettleship furnished useful advice during preparation
of the manuscript.

Literature Cited

Asbirk, S. 1979a. The adaptive significance of the repro-
ductive pattern in the Black Guillemot, Cepphus grylle.
Videnskabelige Meddelelser dansk naturhistorisk Fore-
ning 141: 29-80.

Asbirk, S. 1979b. Some behaviour patterns of the Black
Guillemot Cepphus grylle. Dansk Ornithologisk Forening
Tidsskrift 73: 287-296.

Atmospheric Environment Service. 1975. Canadian nor-
mals, Vol. 1-S1 1941-1970. Environment Canada,
Downsview, Ontario.

Atmospheric Environment Service. 1976-1977. Monthly
records: meteorological observations in Canada. Envir-
onment Canada, Downsview, Ontario.

Bédard, J. 1969. Histoire naturelle du gode, Alca torda, L.,
dans le golfe Saint-Laurent, province de Québec, Canada.
Etude du Service canadien de la faune, numéro 7. 79 pp.

Belopol’skii, L. O. 1957. Ecology of sea colony birds of the
Barents Sea. English translation from Russian, Israel Pro-
gram for Scientific Translations, Jerusalem 1961. 346 pp.

Bergman, G. 1971. Gryllteisten Cepphus grylle in einem
Randgebiet: Nahrung, Brutresultat, Tagesrhythmus und
Ansiedlung. Commentationes Biologicae, Helsinki 42:
1-26.

Bianki, V. V. 1967. Gulls, shorebirds and alcids of Kanda-
laksha Bay. English translation from Russian, Israel Pro-
gram for Scientific Translations, Jerusalem 1977. 250 pp.

Brown, R. G. B., D. N. Nettleship, P. Germain, C. E. Tull,
and T. Davis. 1975. Atlas of eastern Canadian seabirds:
Canadian Wildlife Service, Ottawa. 220 pp.

Cairns, D. 1979. Censusing hole-nesting auks by visual
counts. Bird-Banding 50: 358-364.

Cairns, D. 1980. Nesting density, habitat structure, and
human disturbance as factors in Black Guillemot repro-
duction. Wilson Bulletin 92: 352-361.

Chapdelaine, G. 1980. Onzieme inventaire et analyse des
fluctuations des populations d’oiseaux marins dans les
refuges de la cote nord du golfe Saint-Laurent. Canadian
Field-Naturalist 94: 34-42.

Coulson, J. C. 1968. Differences in the quality of birds nest-
ing in the centre and on the edges of a colony. Nature
(London) 217(5127): 478-479.

318

Lavoie, R., and G. Beaulieu. 1971. Salinité des eaux de
surface dans l’estuaire du Saint-Laurent.
canadien 98: 191-193.

Preston, W. C. 1968. Breeding ecology and social behavior
of the Black Guillemot, Cepphus grylle. Ph.D. thesis, Uni-
versity of Michigan. 138 pp.

Reed, A. 1973. Aquatic bird colonies in the Saint Lawrence
estuary. Service de la faune du Québec, Bulletin 18.54 pp.

Salomonsen, F. 1955. The food production in the sea and
the annual cycle of Faeroese marine birds. Oikos 6:
92-100.

Slater, P. J. B., and E. P. Slater. 1972. Behaviour of the
tystie during feeding of the young. Bird Study 19: 105-113.

Thoresen, A. C.,and E. S. Booth. 1958. Breeding activities
of the Pigeon Guillemot, Cepphus columba columba (Pal-
las). Walla Walla College Publications, Number 23. 36 pp.

Tuck, L. M. 1961. The murres. Canadian Wildlife Service

THE CANADIAN FIELD-NATURALIST

Naturaliste -

Vol. 95

Drent, R. H. 1965. Breeding biology of the Pigeon Guille-
mot, Cepphus columba. Ardea 53: 99-160.

Hartley, C. H., and J. Fisher. 1936. The marine foods of
birds in an inland fjord region in West Spitsbergen. Jour-
nal of Animal Ecology 5: 370-389.

Immelmann, K. 1971. Ecological aspects of periodic repro-
duction. /n Avian biology. Vol. 1. Edited by D. S. Farner
and J. R. King. Academic Press, New York. pp. 341-389.
Monograph Series, Number |. 260 pp.

Uspenski, S. M. 1958. The bird bazaars of Novaya Zemlya.
Translations of Russian Game Reports, Canadian Wild-
life Service, Ottawa. 159 pp.

Winn, H. E. 1950. The Black Guillemots of Kent Island,
Bay of Fundy. Auk 67: 477-485.

Received 23 June 1980
Accepted 16 March 1981

Food Habits of Deer Mice (Peromyscus maniculatus)

in Northern Ontario

ARTHUR M. MARTELL! and ANN L. MACAULAY?

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 3V1
2Present address: Canadian Wildlife Service, 1725 Woodward Drive, Ottawa, Ontario KIA 3Z7

Martell, ArthurM.,and AnnL. Macaulay. 1981. Food habits of Deer Mice( Peromyscus maniculatus) in northern Ontario.

Canadian Field-Naturalist 95(3): 319-324.

Summer (May-September) food habits of 712 Deer Mice (Peromyscus maniculatus) were examined on uncut, selectively cut,
clear-cut, and recently burned sites near Manitouwadge, Ontario. Foods taken showed a distinct seasonal pattern. The
number of food ‘items taken was much lower in May and June than later in the summer. The major food items in May-June
(arthropods and seeds) and in July-September (arthropods, seeds, and berries) varied in amount taken among the sites and
was likely a reflection of their availability. The diet of Deer Mice in selectively cut stands was highly similar to that in both
clear-cuts and recent burns. Deer Mice, because of the diversity and the plasticity of their diet, are able to exploit successfully
the limited food resources available on disturbed and successional sites.

Key Words: Deer Mice, Peromyscus maniculatus, food habits, diet, clear-cut, burn.

Deer Mice (Peromyscus maniculatus) are rare in
uncut upland Black Spruce (Picea mariana) — Feather
Moss (Pleurozium schreberi) forests in northern Onta-
rio. After those forests are clear-cut, Deer Mice invade
from adjoining mixed wood stands and predominate
in the small mammal community by the end of the
second summer following harvest (Martell and Rad-
vanyi 1977; Martell, unpublished data). They main-
tain that status for at least 15-25 yr following harvest.
Deer Mice are also common on recent burns.

Deer Mice, through their seed-feeding activities,
have been considered a serious hindrance to regenera-
tion of forests in many parts of North America (see
Pank 1974). Because of the major ecological changes
caused by timber harvesting and subsequent scarifica-
tion, the availability of preferred foods on clear-cuts
may be altered drastically. However, few researchers
have examined the changes in food habits of Deer
Mice on clear-cuts or in the ensuing successional
stages (Schloyer 1976).

As part of an investigation of the role of small
mammals in the regeneration of upland Black Spruce
forests in northern Ontario, we examined the food
habits of Deer Mice on clear-cuts, on recent burns,
and in uncut and selectively cut stands. The primary
aim of the study was to determine if Deer Mice con-
sume significant quantities of Black Spruce seeds, but
we also hoped that an examination of food habits
would help explain why Deer Mice are so successful at
exploting recent clear-cuts and burns while other spe-
cies, such as the Southern Red-backed Vole (Cle-
thrionomys gapperi), show a drastic decline in
numbers following harvest.

Study Area and Methods

The study area was located on the Ontario Paper
Company lease near Manitouwadge, Ontario
(49°07’'N, 85°50’W). Before harvest, upland sites sup-
ported mature stands of Black Spruce — Feather Moss
forest. After clear-cutting and subsequent scarifica-
tion with small flanged barrels and spiked anchor
chains, the sites consisted of exposed peat, mineral
soil, and rock, with an extensive cover of slash, logs
and stumps. Only a few small patches of living plants
remained (< 5% cover). Vegetation cover increased
with time (after 5 yr 7% moss and lichen cover and
30% vascular plant cover; after 12 yr 22% moss and
lichen cover and 67% vascular plant cover). Recovery
of vegetation on burns was more rapid than on clear-
cuts. Dry knolls in the study area supported mixed
stands of Trembling Aspen (Populus tremuloides),
Paper Birch (Betula papyrifera), White Spruce (Picea
glauca), and Balsam Fir (Abies balsamea). The conif-
ers were selectively harvested from many of those
stands leaving the aspen, Paper Birch, young conifers,
and the thick shrub and herb layer. Patches of
unharvested mixed wood occurred throughout the
selectively harvested stands.

Deer mice were taken with snap traps set in pairs at
approximately 15-m spacing either in a single line
(375 m long) or ina grid (approximately 2 ha in area).
Traps were run for 72 h and were checked each 24 h.
The following sites were sampled during the summers
of 1976 and 1977: (A) uncut upland Black Spruce
(2694 trap nights), (B) selectively cut mixed wood
(4344 trap nights), (C) upland Black Spruce clear-cut
during the winter of 1975-1976 (8058 trap nights), (D)

SH)

320

upland Black Spruce clear-cut during the winter of
1972-1973 (6096 trap nights), (E) upland Black
Spruce clear-cut during the winter of 1965-1966 (3888
trap nights), (F) selectively cut mixed wood harvested
and burned during early summer 1977 (300 trap
nights), (G) upland Black Spruce clear-cut and burned
during the summer of 1975 (1164 trap nights).

Stomachs were removed and the contents were
treated by the method outlined by Williams (1959).
The contents were washed through a 0.25-mm sieve
with hot water to remove fat and gastric juice and were
then dried on filter paper and stored in envelopes for
further study. Before examination the contents were
soaked in water in a watch glass for a few minutes to
restore their original appearance. The contents were
then examined under a dissecting microscope at
20-30 X and the volume of each food item was esti-
mated to the nearest 5%. Identification of items was
made by comparison with a reference collection made
on the study area and by feeding known foods to
captive animals and later examining their stomach
contents. Seeds and fruits were identified with the aid
of Montgomery (1977) and spores of hypogeous fungi
were identified by J. M. Trappe, Pacific Northwest
Forest and Range Experiment Station, Forestry
Sciences Laboratory, Corvallis, Oregon 97330.

In most cases seeds and berries could be identified
to species. Although finely masticated seed endo-
sperm was included with seeds, some were from berry

40

Relative Volume

wW
°o
l

20-

| MAY ]

JUNE | JULY

THE CANADIAN FIELD-NATURALIST

Vol. 95

seeds that were cracked open. No attempt was madeto
identify arthropods, epigeous fungi, or chlorophyl-
lous plant matter to species. Stones, fragments of
peat, and hair were not considered food items and
were not included in the analysis. Food habits
between sites were compared by percentage similarity
(Pielou 1975).

Results and Discussion

Five mouse stomachs from uncut Black Spruce
stands, 187 stomachs from selectively cut stands, 335
stomachs from clear-cuts, and 185 stomachs from
burns were examined. Because only five stomachs
were obtained from uncut Black Spruce stands,
because the contents of those stomachs were not con-
spicuously different from the contents of stomachs
taken at the same time in selectively cut stands, and
because most of the selectively cut stands contained
patches of uncut mixed wood, the five stomachs from
uncut Black Spruce stands were combined with those
from selectively cut stands.

Food habits showed a distinct seasonal pattern
which was, in general, common to all sites (Figure 1).
Arthropods were extremely important from May
through early July and were less important later in the
summer. Seeds were most important in early May and
in September, while berries were important from early
July through early September. The pattern is similar
to that observed by Dyke (1971) for Deer Mice in

TH.
vi

Chlorophyllous
Plant Matter

Achlorophyllous
Plant Matter

AUGUST [SEPTEMBER]

FiGuRE |. Seasonal pattern insummer foods of Deer Mice in northern Ontario. Samples sizes are given above the bars.

1981

uncut boreal forest near Great Slave Lake, Northwest
Territories.

The number of food items taken was much lower in
May and June (x = 9.7 items) than later in the summer
(x = 19.6 items). The mean percentage similarity for
May-June was 67% and for July-September was
71%, but the two periods were only 57% similar to
each other. Food habits were, therefore, combined
into those two periods for comparison among areas.
In the May-June period, sample sizes in three of the
areas are so Small that some food items may have been
missed (Obrtel and Holisova 1977), but sample size
was not a problem in the July-September period.

Although the major dietary items were similar in all
habitats, there were marked differences among areas
in the proportions of items taken both in early
summer (Table 1) and in late summer (Table 2). In
early summer arthropods made up most © 70%) of
the diet of Deer Mice on the 1975-1976 and 1965-1966
clear-cuts and on the 1977 burn, while arthropods and
seeds made up most of the diet on other sites. The
average percentage similarity was 77% within the first
group, 64% within the second group, and 64%
between groups. Seeds of graminoids predominated
in the selectively cut stands and on the 1972-1973
clear-cut while seeds of Pink Corydalis (Corydalis
sempervirens) predominated on the 1975 burn. There
was no consistent pattern of change in any food item
with age of either clear-cuts or burns. Deer Mice likely
used whatever was available on the sites.

In late summer, arthropods and berries made up
most 65%) of the diet on the older (1972-1973 and
1965-1966) clear-cuts while arthropods and seeds
made up most of the diet on other sites. The average
percentage similarity was 62% within the first group,
69% within the second group, and 55% between
groups. Seeds of graminoids predominated in the
selectively cut stands and on the 1977 burn while seeds
of Pink Corydalis predominated on the 1975-1976
clear-cut and on the 1975 burn. Blueberries (Vacci-
nium angustifolium) were the most important berries
taken by Deer Mice. Consumption of blueberries
increased with age of both clear-cuts and burns and
was likely related to availability.

The high proportion of arthropods taken by Deer
Mice in the present study is consistent with other
studies. Arthropods have been reported to be an
important component of the summer diet of Deer
Mice in uncut forests in the Northwest Territories
(48%) (Dyke 1971), Quebec (35%) (Hamilton and
Hamilton 1954), and northeastern United States
(71%) (Hamilton 1941). Williams (1959), however,
examined the food habits of Deer Mice taken primar-
ily in August in uncut forest in Wyoming and Colo-
rado and found that arthropods made up only 13% of

MARTELL AND MACAULAY: DEER MICE, NORTHERN ONTARIO BAI

the diet, while seeds (primarily conifer seeds) made up
63%. Conifer seeds were not identified as food items in
the present study. It is unlikely that anything but a
rare occurrence was missed as we had no difficulty
identifying conifer seeds in the stomach contents of
Deer Mice kept in the laboratory and fed small quan-
tities of seeds.

Few studies have been conducted on the changes in
food habits of Deer Mice on clear-cuts. Tevis (1956)
reported that the frequency of insects in the diet of
Deer Mice in northwestern California was higher in
cutover stands (60%) than in uncut stands (44%).
Schloyer (1976) found that Deer Mice in West Virgi-
nia ate more invertebrates © 45% volume) in very
young (<Syr) clear-cuts and in mature forests
(© 25 yr old) than in clear-cuts in the intermediate
stages (30% volume). He suggests that the proportions
consumed are related to the abundance of insects on
those sites. In our study, the proportion of the late
summer diet made up of arthropods was lower on
young clear-cuts (36%) than in selectively cut stands
(53%), and was lowest on the | 1- to 1 2-yr-old clear-cut
(20%). Those results are consistent with Schloyer’s
findings and may also be related to the availability of
arthropods.

In general, there was a high degree of similarity
between the diet of Deer Mice in selectively cut stands
and that in both clear-cuts (65%) and burns (68%).
The major dietary items differ more in proportion
taken than in kind. Many of the differences are likely
due more to variability among sites than to site treat-
ment or age. Deer Mice, because of the diversity of
their diet, appear able to exploit successfully the food
resources available on disturbed and successional
sites. Southern Red-backed Voles, on the other hand,
have a more limited diet and appear dependent on
food items which are scarce or absent on those sites
(Martell 1981).

Our observed absence of conifer seeds in the diet of
Deer Mice supports other studies which suggest that
conifer seed predation is minimal on recent upland
Black Spruce clear-cuts (Martell and Merritt 1979)
and that Deer Mice do not actively search for Black
Spruce seeds or Jack Pine (Pinus banksiana) seeds
but, rather, encounter them by chance during a search
for other foods (Martell 1979). Black Spruce seeds
were available in uncut Black Spruce stands, White
Spruce seeds were available in selectively cut mixed
wood stands, and both Black Spruce seeds and Jack
Pine seeds were available on the clear-cuts and burns.
Also, Southern Red-backed Voles on the same sites
consumed small quantities of conifer seeds (Martell
1981). Therefore, based on summer food habits, Deer
Mice cannot be considered to bea serious predator of
conifer seeds on upland Black Spruce clear-cuts in
northern Ontario.

Vol. 95

THE CANADIAN FIELD-NATURALIST

B22

(€)1'0 =

(S)LI
(E)11
87C a
(€)E0 es
(€1)9°0 =r
(Op)r' 07 =

roa =e
(E)1 1 ar,
(L€)6'01 es
OCI Te
(06)7' 9 (001)0°001

(6Z)€'F1

a

(Lr il

(OD8'r

cs (€)€0
a (L1)8'1

ONZI —

cl
ee (9)0'r
($)z'0
(LS)8°6€
0'0P

(06)6°€S

Qe =u Qaiu
CL6l ‘LLO1

L=u
9961-S96I

uing

Ic =u
‘€L61-CL61

(E17

68

(001)0°L8

Ce au
“9L61-SL6I

ynd-1e3[D

r9) |

(LI)€9

101

(LI)L9

(L1)0°S1
EAC

(OO1)€ 79

Jo}}eul
juejd snoyAydoiojyoy
slayjue pure SUdUIeIS
Sasseid pure sqio4
SaSSOW
suoyory]

Jayjew jued snoyAydosopya
djnd paynuopiuy
SLUDINIIID DSOY

salllag
ulladsopua paas
Spses paljnuapiu yy
suddAdadwuas SijDPAIOD
SploulwieIyH

sp2as
IZUNJOINVU PaljIUs plu yy
‘dds snwojy

dun

spodoiywy

Cl =u
3nd ATIAIIOIIIS
pue jnouy

d

q pueyv

Wd} pooy

poajuasaid

1B SW9}I Poo] Jo (sasayjuaied ut) auaiin990 Jo AQuanbaly puv (WUddIOd) JWINIOA dATVLIIY “OlLIUGO UJIYIIOU UT IdIPY 199 Jo sjiqey pooj sunf—AVP—] ATAV_L

323

DEER MICE, NORTHERN ONTARIO

MARTELL AND MACAULAY

(€1)6'1 (ODE (6)€'1 (6)7'1 (SIE I (DL ssnew ued snoyAydoso[yoy
(O1L'0 = (91)S'0 (91)6'0 (ZI) (€DE] SIoyjUe PUB SUIUIEIS
(p)b'0 (S)€0 (Z)E0 (€)€°0 (O17 7 (6)0'I sassvid pure sqio4
(€)7'0 (S)1'0> (L)E'0 (p)7'0 (f)€0 (L)E0 SOssoW
(€)1°0 == an (¢1)9'0 == (L)r'0 sudyoly]
vl £0 i 07 QE O€ Joyew jurjd snojAydoiojy)
(L)S*I (S)0°1 (L)8°I (p1)7'7Z (€)s°0 (Z1)6'°7 djnd parynuapiuy)
se i (Z)r'0 (€)Z'°0 (Z)s°0 (1)10> SaplOjpMAAU WUNIUIIID A
(pr)8°0I = (18)1°7s¢ (99)L'€7 (97)0°L (S)L'I WNYOfISNBUD WNIUIIID A
= = = (1)s‘0 = — SLUDINIIID DSOY
Re = (WDE I (SDE E = (€1)8'1 snsosiajs snqny
(10> = (6)Z'1 (170 (€)1I (p)S°0 DUDIUIBAIA DIADSDA{
matt = a = = (Z)r'0 Ad1SNID] SAGIY
7 =i eRe = = (10 DPIAl] DAPUDULOD
Cl Ol 8°96 10¢ 16 Ie SoLI9g
(€Z)S 01 (01)6'7 (L1)9'P ()L°7 (91)8°6 (81)s°9 wiiadsopua p22
(911 (Zr)8°S (Z)1'0> (g)€ "I (ONLI (L)1'Z Spses paljua plus)
= == (TDL'E — — — suadas va3sidy
= re (Z)1'0 = — (Z)r'0 ‘dds pypapy
= a = (D)t0> = (DLO ({DI1BAAAOU) DIJL1UI1Od
(7)6'6 = = (Z)r'0 (6b) 1'SZ (9)S'I suaatasaduas sypptsoD
=F = = = = (CUI piafiaddnod viniag
(1)L°0 =r = = = = “ds xaiD)
(LI)p'L (L€)0'01 (91)0'S (LZ)9°6 (€1)p'9 (IZ)r'01 SPIOUuIUIeIN)
8'6C L8i pel Ov OEP TC? SP29S
(€)9'1 (OI) (L)EE (€)0'1 (8)6'1 (€)e1 ISUNJOINVW PotftjUa plu /}
(6) 1'F1 (L€)9'01 (1€)6'€ (6$)8°91 (S7)6'E (Sr)7'8 ‘dds snwoj5
LSI LCI CL SLi 8s ¢6 isuny
(68)P'8€ (¢6)0'99 (pL)L°61 (€6)0'SE (SQELE (6) PES spodosyiy
OZ@I =u 61 =u gg =u eS] =u 19 =u O8| =u Wid}I poo
CL6I ‘LLO1 9961-S96I “€L61-CL6I ‘9L61-SL6I ind ATIATIIIIIS
pure jnouy)
uing ynd-IeIID
D 4 4 d @) q pure Vv

1981

pajuasaid aie
SUID} POO} Jo (sasoyjuased ur) sUaIINI90 Jo AOuanbady pue (jUsdI19d) sUINJOA JANLIIY “OIUO W1OYIOU UI ITY 199 JO SHUqey Poos Jaquiajdag—A[ng¢—Z a1dV 1

324

Acknowledgments

We are grateful to Great Lakes Forest Research
Centre, Ontario Ministry of Natural Resources, and
Ontario Paper Company for their cooperation. We
thank D. R. Fillman for preparing Figure 1, and J. E.
Bryant and T.C. Dauphine for reviewing the
manuscript.

Literature Cited

Dyke, G. R. 1971. Food and cover of fluctuating popula-
tions of northern cricetids. Ph. D. thesis, The University of
Alberta, Edmonton. 245 pp.

Hamilton, W. J., Jr. 1941. The food of small forest mam-
mals in eastern United States. Journal of Mammalogy 22:
250-263.

Hamilton, W. J., Jr.. and W. J. Hamilton, III. 1954. The
food of some small mammals from the Gaspe Peninsula,
P.Q. Canadian Field-Naturalist 68: 108-109.

Martell, A. M. 1979. Selection of conifer seeds by Deer
Mice and red-backed voles. Canadian Journal of Forest
Research 9: 201-204.

Martell, A.M. 1981. Food habits of Southern Red-backed
Voles (Clethrionomys gapperi) in northern Ontario. Can-
adian Field-Naturalist 95(3): 325-328.

Martell, A. M., and W. F. Merritt. 1979. Preliminary trials
of radio-tagging Black Spruce seed with manganese-54 for
seed fate studies. Canadian Wildlife Service, Progress
Note 94. 4 pp.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Martell, A. M., and A. Radvanyi. 1977. Changes in small
mammal populations following clearcutting of northern
Ontario Black Spruce forest. Canadian Field-Naturalist
91: 41-46.

Montgomery, F.H. 1977. Seeds and fruits of plants of
eastern Canada and northeastern United States. Univer-
sity of Toronto Press, Toronto. 232 pp.

Obrtel, R., and V. Holisova. 1977. Sufficient sample size
for analysis of small rodent diets. Folia Zoologica 26:
193-205.

Pank, L. F. 1974. A bibliography on seed-eating mammals
and birds that affect forest regeneration. United States
Department of the Interior, Fish and Wildlife Service,
Special Scientific Report, Wildlife Number 174. 28 pp.

Pielou, E. C. 1975. Ecological diversity. John Wiley and
Sons, New York. 165 pp.

Schloyer, C. R. 1976. Changes in food habits of Peromys-
cus maniculatus nubiterrae Rhoads on clearcuts in West
Virginia. Proceedings of the Pennsylvania Academy of
Science 50: 78-80.

Tevis, L., Jr. 1956. Responses of small mammal popula-
tions to logging of Douglas-fir. Journal of Mammalogy
37: 189-196.

Williams, O. 1959. Food habits of the Deer Mouse. Journal
of Mammalogy 40: 415-419.

Received 5 January 1981
Accepted 6 April 1981

Food Habits of Southern Red-backed Voles (Clethrionomys gapperi)
in Northern Ontario

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 3V1

Martell, Arthur M. 1981. Food habits of Southern Red-backed Voles (Clethrionomys gapperi) in northern Ontario.
Canadian Field-Naturalist 95(3): 325-328.

Summer (May-September) food habits of 258 Southern Red-backed Voles (Clethrionomys gapperi) were examined on
uncut, selectively cut, and clear-cut sites near Manitouwadge, Ontario. Foods taken showed a distinct seasonal pattern with
new green vegetation, seeds, berries, and mushrooms appearing in the diet as they became available. Conifer seeds were an
insignificant part of the diet. The major dietary items, lichens and fungi, were common to all sites. Because environmental
conditions on clear-cuts are unsuitable for those foods, Southern Red-backed Voles on clear-cuts are likely dependent ona

declining food source.

Key Words: Southern Red-backed Vole, Clethrionomys gapperi, diet, food habits, clear-cut.

Southern Red-backed Voles (Clethrionomys gap-
peri) are common in uncut conifer and mixed wood
forests in northern Ontario. When upland Black
Spruce (Picea mariana) — Feather Moss ( Pleurozium
schreberi) forests are clear-cut, C. gapperi decline in
numbers until they are rare or absent by the end of the
second or third summer following harvest (Martell
and Radvanyi 1977; Martell, unpublished data). The
scarcity of red-backed voles on clear-cuts has been
observed in other parts of North America and has
been attributed to a decrease in either cover, or food
supply, or both (Gashwiler 1970; Lovejoy 1975).
However, comparative studies of food habits of red-
backed voles in uncut and clear-cut stands are few
(Schloyer 1977).

As part of an investigation of the role of small
mammals in the regeneration of upland Black Spruce
forests in northern Ontario, | examined the food hab-
its of C. gapperi on clear-cuts and in uncut and selec-
tively cut stands. The primary aim of the study was to
determine if C. gapperi consume significant quantities
of Black Spruce seeds, but I also hoped that an exami-
nation of food habits would help explain the decline in
numbers of C. gapperi on clear-cuts.

Study Area and Methods

The study area was located on the Ontario Paper
Company lease near Manitouwadge, Ontario
(49°07’N, 85°50’W). Before harvest, upland sites sup-
ported mature stands of Black Spruce — Feather Moss
forest. After clear-cutting and subsequent scarifica-
tion only a few small patches of living plants remained.
Dry knolls supported mixed wood stands from which
the conifers were selectively harvested.

Southern Red-backed Voles were taken with snap
traps during the summers of 1976 and 1977 in uncut

325

upland Black Spruce (2694 trap nights), selectively cut
mixed wood (4344 trap nights), upland Black Spruce
clear-cut during the winter of 1975-1976 (8058 trap
nights), and upland Black Spruce clear-cut during the
winter of 1976-1977 (1884 trap nights). Vole stomachs
were removed and the contents were treated by the
method outlined by Williams (1959). A total of 44
stomachs from uncut stands, 141 stomachs from selec-
tively cut stands, and 73 stomachs from clear-cuts
were examined; 66 of the stomachs from clear-cuts
were from the first summer after harvest and the
remaining 7 were from the second summer after har-
vest. Because the food items were so finely masticated
it was difficult to make specific identifications; how-
ever, food items could be placed in categories with
more confidence. Stones, fragments of peat, and hair
were not considered food items and were not included
in the analysis. All of those items together, however,
made up only 0.2% by volume of total stomach
contents.

A more detailed description of the study area, of
trapping methods, and of the analysis of the stomach
contents is given by Martell and Macaulay (1981).

Results and Discussion

Food habits showed a distinct seasonal pattern
which was similar in all stands (Figure 1). Lichens,
mainly Cladina spp. and Cladonia spp., dominated
the diet in early May and remained an important part
of the diet throughout the summer. Fresh green plant
matter was taken primarily between mid-May and
mid-July as it became available. Green plant matter
became available earlier on the uncut and selectively
cut sites than on the clear-cuts. Seeds were taken
primarily in June and early July, followed by berries
in late July. Mushrooms grew in importance from

326 THE CANADIAN FIELD-NATURALIST Vol. 95
1004 gener | il ‘e MY Gy
90 “Gg isl ZZ )} | rthropods
80 Wy EY ag Wy Y Lichens
UG EV) EV) WAY
e Wy EY VY) ahs
et Ag ng Y AY] IE uns)
2 IW BY ae a7 We
: 50 of ae IH k : Seeds
a Es ise tal me Palo ieennee
el) EW FR EL : ALN
AT) LY i eee
10 eI + tae ao! Achlorophyllous
: ico ine TN Plant Matter

| MAY | JUNE |

| SEPTEMBER|

FiGureE |. Seasonal pattern in summer foods of Southern Red-backed Voles in northern Ontario. Uncut Black Spruce stands
are combined with selectively cut mixed wood stands to achieve sufficient sample sizes, and the combined sample (U) is
compared with clear-cut upland Black Spruce stands (C). Sample sizes are given above the bars.

early summer until they were one of the primary food
items in August and September. The pattern is similar
to that reported for C. gapperi in conifer forest in the
Northwest Territories (Dyke 1971) and in Manitoba
(Perrin 1979).

Food habits were averaged over the summer to
examine differences among sites (Table 1). Voles in
uncut stands took much less arthropod material,
berry material, and chlorophyllous plant matter, and
much more achlorophyllous plant matter than voles
in other stands. That may be related in part to availa-
bility, but is also influenced by the distribution of
stomachs sampled; most of the stomachs from uncut
stands were taken in early May and late September,
whereas stomachs from the other stands were more
evenly distributed throughout the summer. Voles in
selectively cut stands took much less lichen and much
more fungi than voles in either uncut or clear-cut
stands. That difference is likely related to availability.
In general, foods taken on clear-cuts were little differ-
ent from those in uncut and selectively cut stands.
Lichens and fungi made up most (80-89%) of the diet
in all stands.

Schloyer(1977) examined the food habits of 139 C.
gapperi on clear-cuts in West Virginia ranging in age
from lessthan 1 yrto more than 25 yr after harvest. In

that mixed deciduous forest C. gapperi did not disap-
pear from clear-cuts after harvest as they did in north-
ern Ontario. Schloyer found that the primary summer
food items, by volume, were vascular plant matter
(76%) and fungi (22%), primarily hypogeous fungi
(19%), with no significant variation among ages of
clear-cut. Insects made up only a small part of the diet
(1%) but were significantly higher in the 16- to 20-yr
successional stage (7%) than in the other stages. The
food habits I observed were similar to those found by
Schloyer (1977), except that most of the fungi con-
sumed were epigeous (mushrooms) rather than
hypogeous (Glomus spp.).

Summer (May-September) food habits of C. gap-
peri in conifer forest have been examined in the
Northwest Territories (Dyke 1971), Colorado (Mer-
ritt and Merritt 1978), Manitoba (Perrin 1979), and
Ontario (present study). Fungi were the most impor-
tant food item in all studies except Manitoba; how-
ever, the absence of mushrooms in that study may
have been due to the technique used to determine food
habits. The proportions of other food items taken
varied considerably among areas. Other major foods
taken were berries in the Northwest Territories, green
dicotyledonous vegetation and berry seeds in Mani-
toba, and lichens in the present study. Winter foods

1981

MARTELL: SOUTHERN RED-BACKED VOLES, NORTHERN ONTARIO

Sy]

TABLE 1|—Summer (May-September) food habits of Southern Red-backed Voles in northern Ontario. Relative volume
(percent) and frequency of occurrence (in parentheses) of food items are presented

Food item

Arthropods
Lichens
Fungi
Unidentified mushrooms
Glomus spp. (spores)
Seeds
Conifer seed coats
Unidentified seed coats
Seed endosperm
Berries
Fragaria virginiana
Rubus pubescens
Vaccinium angustifolium
Rosa acicularis
Unidentified pulp
Chlorophyllous plant matter
Anchlorophyllous plant matter

also varied among areas with most of the winter diet
being lichens and berries in the Northwest Territories
(Dyke 1971), seeds and fungi in Colorado (Merritt
and Merritt 1978), and dead dicotyledonous vegeta-
tion in Manitoba (Perrin 1979). Because essentially all
of the major food items taken were present in the diet
in all areas, the great differences among areas suggest
that C. gapperi can adapt to the foods available. How-
ever, the similarity in food habits between uncut and
clear-cut stands (Schloyer 1977, present study) sug-
gests that this adaption does not take place rapidly.
On clear-cuts in northern Ontario, C. gapperi con-
tinued to use lichens and fungi as the major items of
their diet. Freshly scarified upland Black Spruce
clear-cuts are usually quite dry and undergo great
fluctuations in daily temperature. Those conditions
make clear-cuts unsuitable for mushrooms and
lichens. Therefore, C. gapperi are likely feeding ona
declining food source. Maser et al. (1978) note the
importance of fungi to several species of red-backed
voles and suggest that the reason California Red-
backed Voles (C. californicus) disappear from clear-
cuts within a year after logging and burning is that the
voles are left without their specialized food supply,
hypogeous fungi. The disappearance of C. gapperi
following harvest in northern Ontario may also be
related to a diminishing specialized food supply, as
well as to a marked decrease in vegetative cover.
Conifer, primarily Black Spruce, seeds were taken
more frequently onclear-cuts than in uncut and selec-
tively cut stands, but conifer seeds were likely more

Uncut Selectively Clear-cut
Black Spruce, cut mixed wood, Black Spruce,
n= 44 n= 141 n= 73
0.7 (30) 2.0 (34) 3.1 (44)
58.6 (91) 30.1 (86) 56.4 (92)
30.6 (55) 52.6 (82) 23M la(83))
30.1 (50) 47.9 (72) 22.2 (49)
0.5 (7) 4.7 (24) 0.8 (11)
2.4 (25) 2.2 (18) 3.2 (32)
<0.1 (2) <0.1 (3) 0.1 (11)

<0.1 (2) <0.1 (1) —
2.4 (23) 2.2 (15) 3.1 (29)
2.8 (14) 6.1 (12) 7.4 (26)
— 0.2 (1)
= 7a) —
0.3 (9) _ 4.0 (12)
ce wat 0.6 (1)
DENS) 4.3 (7) 2.9 (12)
BE21(6)) 8.6 (20) 6.8 (22)
2.4 (18) 0.5 (16) eS)

abundant on clear-cuts than in uncut stands because
cones in the slash open on the warm, dry clear-cuts
and release their seeds. Conifer seeds, however, made
up an insignificant part of the diet. This supports
other studies which suggest that conifer seed preda-
tion is minimal on those recent clear-cuts (Martell
and Merritt 1979) and that C. gapperi do not actively
search for Black Spruce seeds or Jack Pine (Pinus
banksiana) seeds but, rather, encounter them by
chance during a search for other foods (Martell 1979).
Based on their summer food habits and their rapid
decline in numbers following clear-cutting, C. gapperi
cannot be considered to be a serious predator of
conifer seeds on upland Black Spruce clear-cuts in
northern Ontario.

Acknowledgments

I am grateful to Great Lakes Forest Research Cen-
tre, Ontario Ministry of Natural Resources, and
Ontario Paper Company for their cooperation. |
thank A. L. Macaulay for assistance in the field,
S. M. Martell for assistance in the laboratory, D. R.
Fillman for preparing Figure |, and J. E. Bryant and
T. C. Dauphine for reviewing the manuscript.

Literature Cited

Dyke, G. R. 1971. Food and cover of fluctuating popula-
tions of northern cricetids. Ph.D. thesis, The University of
Alberta, Edmonton. 245 pp.

Gashwiler, J.S. 1970. Plant and mammal changes in a
clearcut in west-central Oregon. Ecology 51: 1018-1026.

328

Lovejoy, D. A. 1975. The effect of logging on small mam-
mal populations in New England northern hardwoods.
University of Connecticut. Occasional Papers (Biological
Science Series) 2: 269-291.

Martell, A. M. 1979. Selection of conifer seeds by Deer
Mice and Red-backed Voles. Canadian Journal of Forest
Research 9: 201-204.

Martell, A. M., and A. L. Macaulay. 198t. Food habits of
Deer Mice (Peromyscus maniculatus) in northern Onta-
rio. Canadian Field-Naturalist 95(3): 319-324.

Martell, A. M., and W. F. Merritt. 1979. Preliminary trials
of radio-tagging Black Spruce seed with manganese-54 for
seed fate studies. Canadian Wildlife Service, Progress
Note 94: 4 pp.

Martell, A. M., and A. Radvanyi. 1977. Changes in small
mammal populations following clearcutting of northern
Ontario Black Spruce forest. Canadian Field-Naturalist
91: 41-46.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Maser, C., J. M. Trappe,andR. A. Nussbaum. 1978. Fun-
gal — small mammal interrelationships with emphasis on
Oregon coniferous forests. Ecology 59: 799-809.

Merritt, J. F.and J. M. Merritt. 1978. Population ecology
and energy relationships of Clethrionomys gapperi in a
Colorado subalpine forest. Journal of Mammalogy 59:
576-598.

Perrin, M. R. 1979. Seasonal variation in growth, body
composition, and diet of Clethrionomys gapperi in spruce
forest. Acta Theriologica 24: 299-318.

Schloyer, C. R. 1977. Food habits of Clethrionomys gap-
peri on clearcuts in West Virginia. Journal of Mammalogy
58: 677-679.

Williams, O. 1959. Food habits of the Deer Mouse. Journal
of Mammalogy 40: 415-419.

Received 5 January 1981
Accepted 6 April 1981

The Biological Flora of Canada

2. Vaccinium myrtilloides Michx., Velvet-leaf Blueberry*

S. P. VANDER KLOET! and I. V. HALL?

‘Department of Biology, Acadia University, Wolfville, Nova Scotia BOP 1X0
2Research Station, Agriculture Canada, Kentville, Nova Scotia B4N 1J5

Vander Kloet, S. P., and I. V. Hall. 1981. The biological flora. 2. Vaccinium myrtilloides Michx., Velvet-leaf Blueberry.
Canadian Field-Naturalist 95(3): 329-345.

Vaccinium myrtilloides Michx. is a deciduous low shrub endemic to North America with a range extending from central
Labrador to Vancouver Island and from the Northwest Territories (61° N) to West Virginia (39°N). To produce fruit the
flowers require cross-pollination by bees or andrenids. The fruit of V. myrtilloides is edible and matures during July and
August. Only in New Brunswick and Maine does the species occur in commercially viable quantity on the blueberry barrens.

Key Words: Vaccinium myrtilloides, Velvet-leaf Blueberry; Sourtop Blueberry, biology, ecology, physiology, distribution,
economic importance.

1. Name

Vaccinium myrtilloides Michaux; section Cyanococcus; Ericaceae;

V. canadense Richardson in Franklin’s Journal (1823 p. 736);

V. angustifolium var. integrifolium Lepage 1951;

Sour-top Blueberry, Velvet-leaf Blueberry, Airelle du Canada (Marie-Victorin 1935), Airelle
Fausse-Myrtille (Marie-Victorin 1964).

2. Description of the Mature Plant

(a) Raunkiaer life-form. Chamaephyte, rarely a nanophanerophyte. Although both Scoggan (1950) and
LeBlanc (1963) list this species as a nanophanerophyte for Bic, Gaspé, and Mount Yamaska, respectively, the
average height for V. myrtilloides is less than 50 cm (Camp 1945; Vander Kloet 1977) and we think their
classification reflects the potential rather than the actual height of the species. Winter-deciduous, broad-leaved,
low, colonial shrub with ascending branches, edible blue fruits, and deep tap root; reproduces by seeds and
rhizomes.

(b) Shoot morphology. Stems woody, average height 25 cm with a maximum of 75 cm, usually densely
pubescent (Figure | C), occasionally pilose, verrucose, terete, bark dirty brown or green; buds of two types, the
larger flower buds are borne terminally, and the more lanceolate vegetative buds, whose bracts are sharply
acuminate (Figure 1D), are borne proximally; leaves alternate in a spiral, simple, pinnately netted, entire,
elliptic, blade green and pubescent, rarely glabrous (Figure | C); rhizomes woody, covered with a brown bark,
bearing numerous shoots and adventitious roots; new rhizome growth white or pinkish. Rhizomes have a
prominent central whitish pith when cut horizontally as opposed to the solid core of roots.

(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 like many closely related genera and species are devoid of any
root hairs and protected by a root sheath (Leiser 1968).

(d) Inflorescence. The members of Vaccinium section Cyanococcus are characterized in part by flowers borne
in racemes; members of section Myrtillus by contrast have flowers borne singly in the leaf axils. Flowers of V.
myrtilloides (Figure 1B) are pentacyclic with five green sepals, five petals fused into a short urceolate, white to
pink corolla, 4 + 1 mm long, 10 stamens in two whorls of five and fused to the corolla, pollen grains in tetrads
32 + 3 um in diameter, and a single pistil with inferior ovary. Fruit (Figure 1A) is a true berry composed of 10
pseudolocules each with several small seeds. We found that the average number of perfect seeds was 12 + 10 and
that of imperfect seeds sensu Bell (1957) up to 71 per berry.

*Acronyms follow Index Herbariorum (Holmgren and Keuken 1974).

329

330 THE CANADIAN FIELD-NATURALIST Vol. 95

FiGureE |. A and B. Parts of the mature shoot of Vaccinium myrtilloides: (A) fruiting shoot with characteristic pubescent
stems and entire leaves; (B) a single floret.

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 331

D

FiGureE 1. Cand D. Parts of the mature shoot of Vaccinium myrtilloides: (C) berry and pilose twig; (D) winter twig with
sharply acuminate bracts.

(e) Subspecies. None reported.

(f) Varieties and forms. Lepage (1951) described the taxon integrifolium which he placed in V. angustifolium.
These plants have mostly glabrous leaf blades — only the margins and midveins are pubescent or pilose; the leaf
margins are entire; the twigs of the current season are pubescent in lines; and the perennating bud scales are
sharply accuminate. Phenotypically at least these (var. integrifolium) plants fit better in V. myrtilloides than V.
angustifolium. Moreover, the plants we have grown from seed are diploid, a characteristic yet to be found in V.
angustifolium (see Hall and Aalders 1961; Bent and Vander Kloet 1976; and Hersey and Vander Kloet 1976;
among others). Furthermore, experimentally this group interbreeds freely with V. myrtilloides but not with V.
angustifolium. Consequently we consider this taxon of somewhat glabrous plants hybrids of V. myrtilloides.

Vaccinium myrtilloides forma chicoccum (Deane) Fernald is the white-fruited form. White fruit in V.
myrtilloides is inherited as a simple recessive mendelian factor (Aalders and Hall 1962).

Boivin (1966) treated V. myrtilloides as a variety of V. angustifolium but as Tables | and 2 of Vander Kloet
(1977) amply show, this taxon is both morphologically and biologically distinct from V. angustifolium.

(g) Ecotypes. None reported, but likely.

(h) Chromosome numbers. Longley (1927) first reported 12 bivalent chromosomes for V. myrtilloides. Subse-
quent counts by Darrow et al. (1944), Hall and Aalders (1961), Whitton (1964), Love and Love (1966), and
Hersey and Vander Kloet (1976) have verified 2n = 24 for both American and Canadian specimens. The only
chromosomal anomaly found so far was a diploid-tetraploid chimera from Walden, Maine (Whitton 1964).

3. Distribution and Abundance

(a) Geographic range. Vaccinium myrtilloides is a Nearctic endemic, extending from central Labrador to
Vancouver Island (Figure 2). It extends from almost 61°N in the Northwest Territories southward to several
isolated uplands in the Appalachian Mountains, 39°N. Although Fernald (1950), Ryan (1974), and Scoggan
(1978) report V. myrtilloides from the Island of Newfoundland, no herbarium specimens are extant at A*,
ACAD, BM, CAN, DAO, GA, GH, K, MICH, MIN, NCSC, NCU, NEBC, NFLD, NY, NYS, PH, QK, TEX,

332 THE CANADIAN FIELD-NATURALIST Vol. 95

800 Kilometres

Vaccinium

myrtilloides

Michx.

FiGuRE 2. Distribution of Vaccinium myrtilloides.

TRT, US, and WIS, nor have the authors, after repeated searches on the Island, found the plant there.
Szcezawinski(1962) thought that V. myrtilloides was introduced to the Fraser Delta (he listed no specimens from
Vancouver Island); however, Hebda (1979) and in /itt. regards the species as native both in Fraser Delta and on
Vancouver Island, but quite disjunct — nearest stations lie some 200 km ENE into the interior lodgepole pine
forest of south central British Columbia.

(b) Altitudinal range. Throughout its range V. myrtilloides occurs from about sea level to rarely above 1200 m
above sea level on mountain slopes.

4. Physical Habitat

(a) Climatic relations. Since V. myrtilloides occurs throughout several of Rowe’s (1972) Forest Regions, viz.,
Subalpine, Boreal, Great Lakes — St. Lawrence and Acadian, we infer that the species is adapted to a wide
range of climatic conditions. It tolerates a potential growing season of 200 d in the south to 100 d in the north
(Atlas of Canada 1974), and occurs in subarctic areas of widespread permafrost (Brown 1967) to temperate
regions. Moreover, following the Thornthwaite Classification system as applied to Canada by Sanderson
(1948), V. myrtilloides ranges from a perhumid climatic type on the East Coast to a dry subhumid type in the
west. Numerous small-scale climatic descriptions are also available, e.g., Gaudreau (1979: pp. 12-15).

Moore (1965) regards the species as winter hardy. Indeed Bell and Burchill (1955) observed that in the absence
of snow cover during winter both V. myrtilloides and V. angustifolium are killed back to ground level, and that
during spring cold snaps, up to 16% dead florets are found in exposed buds. Reader (1979) found that spring
frost damage in blossoms began at -1°C and was complete at —10°C.

Bell and Burchill (1955) conclude that their study of winter bud hardiness supports Wulff’s (1943) hypothesis
that the Ericaceae did not evolve in an area of marked climatic periodicity and that many of its genera, including
Vaccinium, are in disharmony with low winter temperatures.

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 333

Camp (1945) reported that V. myrtilloides is quite shade tolerant, and we have observed plants just surviving
in closed canopy White Spruce (Picea glauca) — Balsam Fir (Abies balsamea) stands in central Nova Scotia and
Cape Breton. In canopy openings, however, the plants both flowered and fruited. Likewise Smith (1962) found
that populations of V. myrtilloides in Alberta occurred under low light intensity, indeed had a wider adaptabil-
ity to shade than the associated conspecific V. vitis-idaea.

Seed germination occurs only after periods of wet weather in late summer and autumn.

(b) Physiographic relations. One of the most important factors limiting flowering and subsequent fruit
development is the occurrence of late spring frosts in low-lying areas (Reader 1979). Vaccinium myrtilloides
germinates, thrives, and reproduces well by rhizomes on both mineral and organic soils provided adequate
moisture and aeration are available. These soils are quite acidic: 32 soil samples collected from V. myrtilloides
sites in Nova Scotia, New Brunswick, Maine, Quebec, Eastern Ontario, Alberta, and British Columbia gave pH
values ranging from 3.3 to 5.6; organic matter content varied from 3 (a gravel bar at Blue River, British
Columbia) to 93% (hummocks near Louisbourg, Cape Breton, Nova Scotia). Similar pH values were recorded
by Smith (1962) for Rocky Mountain House, Alberta, and by Gaudreau (1979), who also gives extensive
information on the availability of exchangeable cations in podzolic soils of the Tanginan Hills near Chicobi
Lake, Abitibi- West, Quebec.

The rhizome system of this species, once established, shes an important role in preventing slope erosion.
Should soil particles be washed into the network of rhizomes and shoots, new roots and shoots develop in the
additional soil, favoring the plant and retaining the soil.

Jeglum (1971) found that the presence of V. myrtilloides in the peatlands of Saskatchewan indicated very
oligotrophic conditions (pH 3.0-3.9) as well as indicating that the water level was more than 80 cm below
ground surface.

(c) Nutrient and water relations. The nutrient requirements of a slow-growing, woody perennial such as V.
myrtilloides are much less than many herbaceous species especially those of agronomic value. The nutrient
levels found in mature leaves of V. myrtilloides (Table 1) are similar to those reported for V. angustifolium
(Townsend et al. 1968) and V. macrocarpon (Townsend and Hall 1971). The nutrient occurring in greatest
amounts is N which generally occurs in the concentration range of 1.5-2.0%. Phosphorus is lowest at about
0.1%. Townsend and Hall (1970), working with V. angustifolium, cited the range of concentration of K as
0.40-0.55%, Ca as 0.40-0.65, and Mgas 0.15-0.20, and a perusal of Table | indicates a fairly close concurrence.
The values for the minor elements Fe, Mn, and B are also similar to those reported in the papers cited above.
Manganese values are often exceedingly high in leaves of Vaccinium species first reported by Lockhart and
Langille (1962). Some of the observed tendencies in nutrient levels can be ascribed to the advancing growing

TABLE 1—Nutrient levels in mature leaves of Vaccinium myrtilloides from various locations, based on dry weight"

% ug/ 2
Clone” N P K Ca Mg Fe Mn B
NS 221673 2.00 0.14 0.52 0.38 0.10 20 452 16
NBI‘ 1.34 0.21 0.39 0.67 0.28 104 1343 25
NB2° 1.15 0.19 0.48 0.67 0.31 106 735 22
NB3° 1G 0.10 0.36 0.55 0.24 105 TT 25
NB 829877 1.70 0.09 0.44 0.66 0.20 34 382 40
Me 623877 1.60 0.09 0.34 0.62 0.22 70 304 54
PQ 322674 1.80 0.13 0.38 0.42 0.14 24 282 30
PQ 513774 1.80 0.10 0.30 0.48 0.18 96 400 24
E. Ont 75-35 1.80 0.10 0.26 0.56 0.22 44 458 58
Wisc. 1.45 0.15 0.44 0.52 0.27 192 2177 32
BC 128879 1.40 0.11 0.24 0.56 0.22 148 202 24

“We thank B. Dykeman for performing the leaf tissue analysis.

’Clones are numbered by letters indicating Canadian province or state of the United States, followed by sample number, day,
month, and year, e.g., NS 221673, Nova Scotia; sample 2, collected on 2! June 1973. Vouchers in ACAD.

“Provenance, sandy soil at Hoyt, New Brunswick (coll. B. Dykeman, NB Dept. of Agriculture).

“Gerloff et al. (1964).

334 THE CANADIAN FIELD-NATURALIST Vol. 95

season. Townsend and Hall (1970) found the nitrogen concentrations decrease during the growing season while
calcium and magnesium accumulate in the leaf tissue. Both NS221673 and PQ322674 (collected on the first day
of summer) have the lowest levels of Ca and Mg, and highest of N; the remainder were collected in July and
August.

Gorham and Gordon (1960) discovered that V. myrtilloides is very sensitive to sulfur dioxide pollution and
therefore we think the species has promise as an indicator species to monitor acid-rain.

The taproot (Hall 1957) may penetrate to >1 m into the soil, allowing use of subsoil moisture reserves. Smith
(1962), however, found no taproots on the V. myrtilloides colonies he excavated along slopes at about 1100 m,
Rocky Mountain House, Alberta. These may have been destroyed because of slope soil creep since seedlings
(Figure 3) from Alberta and British Columbia do develop taproots. Smith (1962) also found that the depth of
the rhizomes varied from 4.5 + 0.9 to 9.0 + 2.8 cm depending on the thickness of the organic horizon. The
thicker the organic horizon, the shallower the rhizome system.

In the Acadian Forest Region of New Brunswick, Flinnand Wein (1977) gave a rhizome depth of 8 + 1 cm for
V. myrtilloides.

5. Plant Communities. Within the greater part of its transcontinental range from Nova Scotia to British
Columbia (Figure 2), V. myrtilloides is a common understory shrub in a variety of drier and less fertile
coniferous forest and woodland communities, achieving highest abundance and fidelity on well-drained sandy
sites dominated by Pinus spp. (Table 3). It also occurs, however, on xeric rock outcrops and hygric peatlands,
and Maycock and Curtis (1960) have shown that it has a bimodal distribution along the moisture gradient in
forests of the upper Great Lakes region, 1.e., peaks of presence and abundance on dry sites and wet sites.

The species is probably most abundant, in terms of total biomass and fruit production, in young, postdistur-
bance communities, e.g., recently burned pine forests from western Quebec to British Columbia, and recently
clear-cut forests in New Brunswick and Maine now managed as blueberry fields by commercial growers (Hall
1959).

On heath barrens in Nova Scotia (Hall and Aalders 1968), old fields in New Brunswick (Hall 1959), and
granite—gneiss outcrops in eastern Ontario (Hall et al. 1979) V. myrtilloides is present but usually much less
abundant than V. angustifolium. In the upper Great Lakes region and northwestern Ontario these two species

24 38 56

FIGURE 3. Vaccinium myrtilloides seedlings. Berry collected from alder forest at Blue River, British Columbia, on 29 August
1979 and seedlings grown from seed in the greenhouse at Acadia University, Wolfville, Nova Scotia.

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 335

TABLE 2—Summary of the mean +SE values for exchangeable cations of Vaccinium myrtilloides soil from several granitic
outcroppings in the Thousand Island Region of eastern Ontario

Cation content (mmol/g dry soil)

Site N Cag Na Mg” K* Mn™

Washburn, Frontenac County 18 1.79+0.39 0.14+0.01 0.5340.11 0.3140.05 0.40+0.14
Mt. Fitzsimmons, Ivy Lea, Leeds County 10 1.72+0.60 0.1340.02 0.47+0.09 0.35+0.05 0.1640.11
Rock Dunder, Morton, Leeds County 19 2.75+0.54 0.15+0.01 0.81+0.13 0.40+0.06 0.56+0.20
Gananoque, Leeds County 4 3.92+1.04 0.1340.02 0.6240.14 0.29+0.02 0.66+0.27
Charleston Lake, Leeds County 2 425 e239 Onl OL02i e052 0250 ian Or3 O== OF O23 et ONIbl

frequently co-dominate the dwarf shrub strata of xero-mesophytic pine woodlands (Maycock and Curtis 1960)
(Table 3). West of Lake Winnipeg on the boreal interior plains, however, where V. angustifolium does not occur
(Hallet al. 1979), V. myrtilloides is the only blue-fruited Vaccinium in many Pinus banksiana woodlands (Table
3) except on colder sites where V. caespitosum sometimes occurs with it. On higher-elevation, moister sites in
Alberta and British Columbia V. membranaceum, V. myrtillus,and V. scoparium replace V. myrtilloides (Wali
and Krajina 1973; La Roi and Hnatiuk 1980). In northern boreal woodlands beyond the northern limits of V.
myrtilloides the dominant blueberry is V. u/iginosum (Viereck and Dyrness 1980).

Hall (1959) reported V. myrtilloides cover values >2Z5% in cleared woodlots of New Brunswick (Table 3).
Segadas-Vianna (1955) found V. myrtilloides in numerous bog communities of Ontario and Quebec (Table 3).
La Roi(1967) tallied V. myrtilloides in 11 of 19 Picea glauca — Abies balsamea stands from Cape Breton to NW
Manitoba, but its cover was never >1% in them; in contrast, the species occurred in 16 of 20 P. mariana -
feather moss stands from central Alberta to New Brunswick with cover values up to 5% east of Lake Superior
(Table 3). Curtis (1959) recorded V. myrtilloides in 73% of his northern wet forests in Wisconsin. Rowe (1956)
classified V. myrtilloides as a xero—mesophytic species characteristic of fresh forest types in Manitoba and
Saskatchewan. Kabzems et al. (1976) found the species to be important in their Pinus banksiana — Vaccinium
vitis-idaea — Pleurozium schreberi forest type on well-drained sites in Saskatchewan. Moss (1953) reported that
V. myrtilloides was abundant in P. banksiana and P. contorta woodlands as well as locally abundant on the
drier hummocks of peatlands in Alberta. Wali and Krajina (1973) recognized a P. contorta — V. myrtil-
loides — Arctostaphylos uva-ursi — Cladina gracilis association for very well-drained aeolin sands and sandy
outwash materials in the northern interior of British Columbia; in some of their stands V. myrti/loides attained
cover values >40% (Table 3). Within its range V. myrtilloides invariably has much higher presence and
abundance in needle-leaf evergreen than in broad-leaf deciduous forests.

Table 3 shows the estimated cover of terrestrial bryophytes, terrestrial lichens, and vascular plant species ina
representative set of seven North American plant communities in which V. myrtilloides is a quantitatively
important component. In the recently cleared New Brunswick woodlot managed for blueberry production, V.
myrtilloides has the highest cover of any vascular species; V. angustifolium, Cornus canadensis, Spiraea alba,
Dennstaedtia punctilobula, and Kalmia angustifolia were the main associated species. In the Black
Spruce — Leatherleaf bog of Quebec, Picea mariana, Chamaedaphne calyculata, and Rubus chamaemorus were
the dominant species and V. myrtilloides had only 1% cover. In the Black Spruce — Feather Moss forest of
western Quebec, P. mariana, Ledum groenlandicum, C. canadensis, Gaultheria hispidula, K. angustifolia, and
V. myrtilloides were the leading vascular species. In the White Spruce — Balsam Fir forest of northern Ontario
P. glauca, A. balsamea, Diervilla lonicera, Aralia nudicaulis, Clintonia borealis, and Cornus canadensis were
the cover-dominant species and V. myrtilloides had only 0.7% cover. In the mixed Jack, Red, and White Pine
forest of upper Michigan, Pinus banksiana, Pteridium aquilinum, Pinus strobus, Vaccinium angustifolium, and
V. myrtilloides were the dominant vascular species. In the Jack Pine forest of northern Alberta V. myrtilloides
was the third-ranked vascular species after P. banksiana and V. vitis-idaea, with C. canadensis and Linnaea
borealis next. In the Lodgepole Pine forest of northern British Columbia V. myrtilloides ranked second after
Pinus contorta with an estimated cover of 41%; the main associated species were Arctostaphylos uva-ursi, V.
caespitosum, and C. canadensis. The latter species was recorded in six of the seven stands in Table 3 and was a
leading member of the understory in five stands. The heath family (Ericaceae) has high quantitative importance
in plant communities containing V. myrtilloides; 9 of the 24 leading vascular species listed above are heaths.
Both bryophytes and lichens can attain very high cover values in V. myrtilloides communities (Table 3).

The material in this section was prepared and written by George H. La Roi (Department of Botany,
University of Alberta, Edmonton, Alberta T6G 2E9).

336 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 3—Species-stand table showing the percent cover of terrestrial bryophytes, terrestrial lichens, and vascular plant
species ina representative spectrum of North American plant communities in which Vaccinium myrtilloides is a quantitatively
important component. Communities include (1)* a recently logged forest in New Brunswick; (2) a Black Spruce — Leatherleaf
bog in Quebec; (3) a Black Spruce — Feather Moss forest in western Quebec; (4) a White Spruce — Balsam Fir forest in
northern Ontario; (5) a mixed Jack, Red, and White pine forest in upper Michigan; (6) a Jack Pine forest in northern Alberta;
and (7) a Lodgepole Pine forest in northern interior British Columbia. Very rare species present in a single stand are not
included

Stand number I 2 3 4 5 6 7

Terrestrial bryophytes — 63.0 86.6 21.3 4.5 90.0 3.0

Terrestrial lichens — 0.5 0.3 4.8 B25 2.0 70.0

Vascular species

Trees
Abies balsamea** — _ 0.1 10.0 0.1 0.1 —
Acer rubrum — — — — 0.4 _ —
Betula papyrifera — — 0.5 5.0 0.1 — —
B. populifolia 1.0 a _ _ ~~ — —
Picea glauca a — — 20.0 0.1 3.0 —
P. mariana — Biles 58.0 0.1 = 0.5 —
Pinus banksiana — — 0.1 0.1 18.6 30.0 —
P. contorta — _— _— — — _ 42.0
P. resinosa _ _ — _ 3.6 —
P. strobus — — — 3.0 4.1 = =
Populus tremuloides 0.4 _ 0.1 2.0 _— 0.5 —
Sorbus americana — -- 0.1 2.4 _ _ —
Shrubs

Acer spicatum - — — 4.9 — — =
Alnus crispa _ _ 0.1 3.7 — = =
Amelanchier alnifolia — — — — — a 0.5
A. stolonifera _ _. -- 0.1 0.4 = —
Andromeda glaucophylla - 0.5 — — — = =
Arctostaphylos uva-ursi — — = — — 2.0 8.0
Chamaedaphne calyculata — BES — — — = =
Corylus cornuta — — — 5.0 — = =
Diervilla lonicera — — 0.1 7.6 0.1 = =
Epigaea repens — — 0.1 0.1 0.8 — _
Kalmia angustifolia 4.4 1.0 3.8 — = = =
K. polifolia — 5.0 — — == = <=
Ledum groenlandicum - 5.0 10.7 _ a 0.1 —
Lonicera canadensis a — _— 0.1 — = =
L. involucrata — — — — = 0.1 —
Oxycoccus microcarpus — — 1.0 — — = =
Prunus pensylvanica Bp. _ — 0.3 — = =
P. virginiana — — — 0.3 0.2 = =
Pyrus arbutifolia 1.8 _ — — — — =
Rhododendron canadense 3.0 a _ — — = —
Ribes glandulosum — — — 0.2 — _ =
Rosa acicularis — — _ 0.1 — 0.1 =
R. gymnocarpa — — _ — _ — 0.1
Rubus idaeus — — 0.1 0.4 — = =
Salix scouleriana — — _ _— _ _— 0.1
Shepherdia canadensis — — — — ~ 0.1 —
Spiraea alba 5.8 — — — — = =
Vaccinium angustifolium 22.8 1.0 Tel 0.3 4.0 = =
V. caespitosum — — = — = — 7.0
Vaccinium myrtilloides 26.2 1.0 2.1 0.7 3.3 9.9 41.0

V. vitis-idaea
Viburnum edule — _ 0.1 — — 0.3 =

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES ROT)

TABLE 3—(continued)

Stand number l 2 3 4 5) 6 a

Forbs
Actaea rubra — — — 0.1 a — —
Apocynum androsaemifolium — — 0.1 0.1 — —
Aralia nudicaulis — — — 7.0 — — —
Aster conspicuus _— — — — — — 0.5
A. lateriflorus 0.6 -- — — = —

A. macrophyllus — _— — DD — — —
Campanula rotundifolia — — — —_ a 0.1 —
Chimaphila umbellata — — — — _ a 0.5
Clintonia borealis _— — 0.1 5.4 — — —
Coptis trifolia — 0.5 0.1 1.0 oa 0.1 —
Cornus canadensis 7.8 0.5 4.8 Sal _ 5.6 3.0
Epilobium angustifolium 0.6 — 0.1 — 0.1 0.4 0.1
Fragaria virginiana — — — a — 0.3 —
Galium boreale — — — _— — 0.2 —
G. triflorum — — 0.1 — = a
Gaultheria hispidula — 0.5 4.1 0.3 — — =
G. procumbens — _ — — 1.1 a —
Geocaulon lividum — 0.5 0.3 — — —- —
Hieracium pratense 0.8 — — — — — —
Hypopitys monotropa = = — — 0.1 — —
Lathyrus ochroleucus — — = — — 0.8 —
Linnaea borealis _— _— 0.3 2.0 — BE 0.5
Listera cordata _ — 0.1 — — — —
Maianthemum canadense — — 0.4 Do) — 2.0 —
Melampyrum lineare _ — — — 0.8 _— 0.5
Moneses uniflora — — = 0.1 0.1 = =
Pyrola secunda — — 0.1 0.3 — 0.1 =
P. virens — — — 0.1 — — 0.5
Rubus chamaemorus — 37.5 0.1 — — — —
R. pubescens — — —

Rumex acetosella 1.3 — — — — = —
Sarracenia purpurea — 0.5 — — = = =
Smilacina trifolia — 5.0 — — — — =
Solidago rugosa 0.6 — — — — = =
Streptopus roseus — — — 0.6 — = =
Trientalis borealis — _— — 0.6 — 0
Viola adunca — — — — 0.2 0.
V. renifolia _— — — 0.2 — 0

Graminoids
Agrostis tenuis 0.4 a Les = a Se = a
Calamagrostis canadensis — _ — — _ 0.4 oo
Carex aenea = — — — a 0.1 —
C. oligosperma aoa 5.0 — — = = ae
C. pauciflora ese 5.0 ast tes aS fas or
C. paupercula _ 1.0 a Ee. as ws: .
C. trisperma — 5.0 0.1 — — = =
Danthonia spicata 0.8 — — — 0.2 — —
Elymus innovatus — = = Be ae 1.4 es
Eriophorum vaginatum — 0.5 — _ _ _ —
Oryzopsis asperifolia — — — — 0.1 — 0.5
O. pungens = _ == = 0.4 0.1 =
Schizachne purpurascens ~ _ — 0.1 — = =

Pteridophytes
Dennstaedtia punctilobula 5.4 — — — — = ==
Dryopteris austriaca — — — 0.1
Gymnocarpium dryopteris — — _ 0.1 —

338 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 3—(concluded)

to
Wo
£
Nn
oO
)

Stand number l

Pteridophytes (continued)
Lycopodium annotinum -- — --
L. clavatum a _ a
L. complanatum _ — _ — 0.1 _ a
L. tristachyum — — — — — 0.1 —
Pteridium aquilinum 3.0 — _ 0.1 17.8 _— a

os

*(1) Data from Hall (1959); (2) data from relevé no. 6, Table 95 in Gaudreau (1979), with Braun—Blanquet scale converted to
cover; (3) data from Black Spruce stand no. 18 in La Roi (1967); (4) data from White Spruce — fir stand no. 22 in La Roi
(1967); (5) unpublished data from Marquette County supplied by G. H. La Roi; (6) unpublished data from House River
area supplied by G. H. La Roi; (7) data from relevé no. 44, Table 3 in Waliand Krajina (1973), with Domin-Krajina scale
converted to cover.

**Nomenclature follows Scoggan (1978).

6. Growth and Development

(a) Morphology. The seed, when sown following procedure of Vander Kloet (1978), sends out a radicle after
20 + 7 d, cotyledons emerge after 31 + 8 d, and the first leaves unfold after 48 + 6 d. These first leaves (Figure 3)
are much smaller and usually more ovate than mature leaves, moreover they are often coarsely serrate, making
identification of seedlings difficult until the first perennating buds are formed. Seedlings germinated ina misting
chamber during October, transferred to a cold frame the following June, and transplanted to the field in
October, do not flower or produce rhizomes until the third summer. Once the plant has attained a diameter of
about 20-30 cm and rhizome growth occurs in several directions, expansion of the clone is more rapid. Smith
(1962) found that in mature plants rhizome length ranged from 100 to 1400 cmand produced an average of one
to two shoots for each 100 cm of rhizome length, depending on the density of the overstory. Hall (1957)
observed a maximum of five growth rings in the eldest rhizome of V. myrtilloides while the taproot of the same
specimen had 15 growth rings and a diameter of | cm.

(b) Physiology. Little is known about the physiology of V. myrtilloides. Usually this species is lumped with V.
angustifolium under the general rubric of lowbush blueberry culture, but the physiological work is invariably
carried out on V. angustifolium (e.g., Hall et al. 1970). Wood and Wood (1963) reported that there was more
sucrose in the nectar of V. myrtilloides than in V. angustifolium. Hilton and Barker (1962) have unpublished
data which suggests equimolar quantities of glucose and fructose and only traces of sucrose in frozen berries of
V. myrtilloides.

The physiology of flowering is also poorly known. We found that apical abortion precedes the onset of flower
bud induction. However, the causality of apical abortion is poorly understood. Hilton and Barker (1962)
suggested an endogenous system linked in part to auxin production and apical activity. We are currently testing
responses of V. myrtilloides under different photoperiodic regimes.

(c) Phenology. Vaccinium myrtilloides overwinters ina leafless state, with greenish, yellowish, or brown twigs.
Flower primordia are formed in the previous late summer, shortly after the cessation of vegetative growth, but
may continue until late October if air temperatures remain >0°C with extended periods greater than 10°C (Bell
and Burchill 1955). These workers found that by early winter, the immature flowers have a completely turned
ovule, the archegonia are mature, the integument is not differentiated, but the anthers are fully formed and
mature pollen mother cells are present. Thus temperature is an important regulator of phenology.

At Ivy Lea, Leeds County, eastern Ontario, the vegetative and flower buds begin to swell in late April or early
May, if night air temperatures have exceeded 6°C for 4-5 d. Anthesis occurs there usually between 22 May and
28 May, a week earlier in central New York, but in Maine, Gaspé, New Brunswick, and Nova Scotia a week or
10d later (Wood 1965). In northern Quebec, Labrador, Ungava, northern Ontario, northern Manitoba,
Saskatchewan, northern Alberta, and the Northwest Territories, flowering usually occurs from late May to
early July while in British Columbia flowering occurs from late May in the Fraser Delta to late June inland
(personal observations; Taylor and MacBryde 1977).

Leaf and vegetative shoot development precedes, is concomitant with, or follows, flowering depending on the
particular clone. The formation ofa black tip at the apical meristem indicates cessation of shoot growth. At Ivy
Lea, leaves harden by mid-June, turn red in early October (or which is more likely will have crisped during one
of the periodic droughts that strike these outcroppings during July and August), and absciss in late October.

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 339

Berries ripen 49-68 d after flowering, depending on precipitation, temperature, and the effectiveness of the
pollinators. Fruit set may be delayed (Aalders and Hall 1961) if the pollinators carry a mixed load, which they
frequently do (Vander Kloet 1976).

7. Reproduction

(a) Floral biology. In eastern Ontario, the flowers are primarily pollinated by andrenids especially Andrena
vicina and A. carlini (Vander Kloet 1972). The 4- to 5-mm-long corolla is not readily manipulated by Bombus,
nevertheless B. ternarius and B. terricola are frequent visitors along with andrenids in Maine and in the
Maritimes (Boulanger et al. 1967). Likewise Reader (1977) observed Apis mellifera, Bombus spp., andrenids,
and dialictids on V. myrtilloides in western Ontario. The latter also noted that the visitation rate to V.
myrtilloides flowers was about 9 per minute for Apis mellifera, 13 for Bombus spp., and 4 for andrenids and
dialictids. Nectar volume is 0.3 + 0.11 yl per flower of which some 70% are dissolved solids (Reader 1977).

The 10 stamens are functional as soon as the flowers fully open and produce 2.4 X 103 tetrads (Reader 1977),
but the stigmata do not become receptive until 2-3 d later. After pollination, the greenish-cream colored corolla
turns pink and a few days later abscises.

Experimental pollination trials indicate that selfing is generally not successful, but both Reader (1977) and
Whitton (1964) reported successful selfing in the wild. The results of controlled intercrossing and selfing of six
populations of V. myrtilloides are given in Table 4.

The vascular anatomy of the ovary has been described by Palser (1961).

(b) Seed production and dispersal. It is rare to find a seedless berry of V. myrtilloides even when the number of
berries per unit area of plant is low. The number of seeds per berry in crosses is givenin Table 4. The American
Robin (Turdus migratorius) is a major seed disperser in southwestern New Brunswick as it often feeds and
migrates just before the berries are harvested by man (Eaton 1957). By growing seeds from scats of Black Bear
(Ursus americanus), we have found inter alia, seedlings of V. myrtilloides. The number of large, plump, viable
seeds per berry in the field is 16 + 10 which weigh 26 + 7 mg/ 100 seeds based on 250 samples. Number and
weight of seeds are not correlated (Vander Kloet, unpublished data) and depend on genetic factors and
environmental conditions. Should the weather be warm and dry during anthesis, pollinators tend to be more
active and berries have higher seed counts.

(c) Seed viability and germination. Viability is reduced by at least 10% after excretion by birds and mammals
(Krefting and Roe 1949). Nichols (1934) reported a germination success of 20%. We obtained 30% and could not
find any difference among samples that had been frozen, held in the refrigerator at 1°C, and freshly harvested
ones. Two of 16 batches showed a bimodal germination, i.e., several seeds germinated after 18 d and these had
their second set of leaves when a second lot of seeds began to germinate 76-82 d after sowing. Seeds of V.

TABLE 4—Effect of self-pollination and cross-pollination on fruit set and seed number under greenhouse conditions in
Vaccinium myrtilloides for the years 1976 to 1978

Number of
Parentage Number of Percent seeds per Percent
of cross flowers pollinated set berry germination
Self-pollinations
ONT 75-41* (1 plant ) 50 0 — =
PQ 324776 = (3 plants) 59 0 — =
NS 222773 (5 plants) 50 6 | 25
NS 517773 (3 plants) 50 4 7 44
NS 13873 (2 plants) 50 0 — —
NS 221873 (2 plants) 50 10 2 25
Cross-pollinations
ONT 41 X NS 222773 119 32 9+2 II
PQ 324776 X NS 517773 125 51 8+2 46
NS 221873 X NS 517773 150 WS 31+7 23
NS 13873 X NS 222773 100 65 16+6 51
NS 222773 X NS 517773 100 59 20+ 11 37

*Vouchers in ACAD. Plants are numbered by letters indicating province followed by sample number, day, month, and year,
e.g. NS 222773, Nova Scotia; sample 2 collected on 22 July 1973.

340 THE CANADIAN FIELD-NATURALIST Vol. 95

myrtilloides germinate best when sown ona |:! sand—peat mix (pH 4.5) ina misting chamber in a greenhouse.

(d) Vegetative reproduction. Asexual reproduction occurs when the older rhizomes senesce, are cut by slope
creep, partially killed by fire, by shading, burrowing, and/or frost heaving. Smith (1962) found that rhizome
depth and shoot number were influenced by location.

8. Population Structure and Dynamics.

(a) Dispersion patterns. Lateral growth of the rhizome system below ground is a means for continuous
advancement of ramets of a particular clone. Branching of rhizomes is strongly dichotomous following the
cessation of growth in the dormant period. Injury to the growing point will produce a similar effect as two lateral
buds are then stimulated to grow one oneither side of the original growing point. Rhizome growth continues as
long as a few shoots develop and translocate foodstuffs back to the storage tissues of the rhizome such as the
parenchyma cells of the pith. Thus V. myrtilloides frequently shows a contagious, clonal dispersion.

Long-distance dispersal is by seed. The genetic diversity found in the seedlings of V. myrtilloides offers a
multitude of new genotypes in each new habitat (Hall and Aalders 1961). Under heavy competition in old fields
seedling growth is slow, requiring at least five years for a plant to reach 15 cm in diameter (Hall 1957; Eaton and
Hall 1961).

Once a plant becomes established it takes a major change in vegetation such as the development of a
multilayered overstory to destroy the clone. In the 25 to 30 years that are often required for the forest canopy to
develop, the V. myrtilloides clone may grow up to 10 m in diameter and spread to adjoining areas. The most
destructive factor is cultivation and even this may be beneficial since it cuts the rhizome into segments that
continue to grow and produce shoots (Hall 1963).

(6) Growthand turnover rates. Once a plant becomes established it can persist almost indefinitely, though often
in a weakened condition. Roland (1946) considered V. myrtilloides to be a relict in certain poplar woods of
well-drained sandy areas of the Annapolis Valley in Nova Scotia.

(c) Successional role. Vaccinium myrtilloides is one of the lesser shrubs in the successional sequence from
abandoned hayland to forest inthe Maritime Provinces (Hall 1959). Following clearfelling of A. balsamea and
Picea rubens in the same region, weak plants of V. myrtilloides and other relatively heliophytic woodland
species produce vigorous shoots which quickly restore their vigor and establish a dense pioneer stage leading
back to the climax fir—spruce forest (Hall 1955).

Farther west where V. myrtilloides is an important understory shrub in pine forests and woodlands (see
Section 5) which are periodically burned over by wildfires (Rowe and Scotter 1973), some rhizomes usually
survive even the hottest fires if the soil is deep enough to afford refuge (Flinn and Wein 1977). On relatively
infertile and dry sandy sites where competition from herbs is not intense, V. myrtilloides populations expand
rapidly, with peak flower and fruit production usually occurring 10—20 years after fire and prior to tree canopy
closure, after which they decline in cover and fruit production.

The material in this section was prepared and written by George H. La Roi (Department of Botany,
University of Alberta, Edmonton, Alberta T6G 2E9).

9. Interaction with Other Species

(a) Competition. In the Acadian Forest Region V. myrtilloides is an important component of some forests
(Loucks 1962); in these it frequently occurs with D. punctilobula (Hay-scented Fern), Lycopodium spp. (a club
moss), C. canadensis (Bunch Berry), Maianthemum canadense (Wild Lily-of-the-Valley), and the conifers of the
upper story (Hall 1955). Vaccinium myrtilloides has a much higher survival potential than its congener V.
angustifolium in the mature Acadian spruce-fir forest (Hall 1959).

In lowbush blueberry fields of the Acadian Forest Region that have recently been developed from woodlots,
D. punctilobula, Pteridium aquilinum (Bracken), S. alba (Meadowsweet), Prunus pensylvanica (Pin Cherry),
C. canadensis, Rhododendron canadense (Rhodora), and K. angustifolia (Lambkill) are the chief competitors
(Hall 1959) (Table 3).

(b) Symbiosis. The pollinators of V. myrtilloides, mainly bumblebees or related genera, have been described in
Section 7(a).

At present there is considerable interest in mycorrhizal associations in Vaccinium species (Pearson and Read
1973a, b). In Canada, the Department of Forestry at Laval University has a contract with Agriculture Canada to
demonstrate the presence and importance of mycorrhizae in lowbush blueberries. Englander (1979), working
with Rhododendron (Ericaceae), found that radioactive phosphorus (P32) applied to the fungal fruit body

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 341

appeared within a short time in the suspected host plant; when the isotype was applied to twigs of the host plant
it soon appeared in the fungus.

(c) Predation and parasitism. The insects found on V. myrtilloides are generally the same as those found on V.
angustifolium. G. W. Wood (personal communication) states, “the pubescence of the foliage of V. myrtilloides
appears to hinder the feeding and movement of the thrips larvae and thus acts as a factor of resistance. Injury to
V. myrtilloides by thrips is not typical. The leaves do not curlas tightly as in the case of V. angustifolium, and the
occurrence of aphids and other insects in the leaf galls is much more common.” Feeding by larvae of Actebia
fennica (Black Army Cutworm) starts in the early spring as soon as the flower buds of V. myrtilloides begin to
swell; they make a small hole in the side of the flower buds and eat out the inner tissue; feeding is mainly
nocturnal (Wood and Neilson 1956). From mid-June to late July immediately following bloom in bearing fields
and in recently burn-pruned areas Neochlamisus cribripennis (Blueberry Case Beetle) larvae may feed on
foliage, stems, and fruit. The adults emerge in late August and possibly feed on the bark until winter sets in
(Wood 1970). Eggs of Frankiniella vaccinii and Taencothrips vaccinophilus (Blueberry Thrips) are laid in the
shoots in late May or early June and the larvae feed and live in the shoots from June to August; their presence is
recognized by the characteristic wrapping of the reddish leaves about the stems (Wood 1956 and 1960). Larvae
of Altica sylvia (Blueberry Flea Beetle) attack blueberry fields from early to mid-June (Maxwell et al. 1953).

Orgyia leucostigma (Whitemarked Tussock Moth) larvae attack a variety of broadleaf trees and evergreens
and may feed heavily on V. myrtilloides (Wood 1979). As the fruit ripens the adults of Rhagoletes mendax
(Blueberry Maggot) begin to lay their eggs in fruit (Wood 1962); this species has not been found in Labrador and
Quebec.

Fruits of V. myrtilloides are eaten by several birds, most notably, American Robin. Foliage and twigs of V.
myrtilloides are eaten by a number of mammals chief of which are Odocoileus virginianus (White-tailed Deer)
and Sy/vilagus floridanus (Eastern Cottontail). Sheep selectively avoid V. angustifolium and V. myrtilloides
and consume the grasses and sedges between or associated with these species (Hall 1954). Fruits are eaten by
Procyon lotor hirtus (Raccoon), Odocoileus virginianus, Vulpes vulpes (Red Fox), Ursus americanus (Black
Bear), and Erethizon dorsatum (Porcupine).

Important fungal diseases of V. myrtilloides are Botrytis cinerea (Grey Mold), Exobasidium vaccinii Wor.
(Red-leaf), and Pucciniastrum géoppertianum (Witches-broom) (Conners 1967). We think Monilinia Blight
caused by Monilinia vaccinii-corymbosi occurs on V. myrtilloides, but it has not been reported as such because
of field reports on “lowbush blueberry” or “blueberries.” Parmelee (1977) records the presence of Microsphaera
vaccini on V. myrtilloides.

(da) Toxicity and allelopathy. Devlin (1980) reports a growth inhibitor in V. macrocarpon (Cranberry) leaves
but none is known for V. myrtilloides.

10. Evolutionand Migration. Camp (1942) classified V. myrtilloides as a member of the basic diploid group
2n = 24) which arose from phyletic divergence as opposed to his “mere speciation” which could occur whenever
several of the diploid taxa introgress. Subsequent allopolyploidy or autopolyploidy would produce polyploid
taxa — taxa, which, Camp argued, are the end of the evolutionary process and not the beginning.

We postulate that section Cyanococcus originated in South America, migrated through the Caribbean
Islands (there are no members of the section along the isthmus between North and South America, including
Mexico, and southwestern USA) onto the sand dunes, relict sand dunes, and pine flatwoods of peninsular
Florida, an area where the diploid V. darrowii still thrives. The trend has been to lose the evergreen and
xeromorphic characters such as small, thick, glaucous, inrolled leaves [glaucescence has been shown to retard
transpiration (Andersen et al. 1979; Freeman et al. 1979)] of the present V. darrowii to a more temperate
deciduous plant with fewer xeromorphic characters as in V. tenellum (Table 5), or with larger leaf blades as in V.
pallidum. A concomitant reduction in seed weight (Crouch and Vander Kloet 1980) and pollen size has also
been observed as the group drifted northward with the North American continent and was subjected to an era of
continuous climatic cooling which transformed the Arcto-Tertiary deciduous forest and V. pa/lidum into more
boreal types where the shorter growing season selected against larger leaf blades and heavier seeds.

The subsequent glacial epochs of the Pleistocene forced many of the northern diploid populations to migrate
southwards, which resulted in the formation of several hybrid populations and species suchas V. myrsinites, V.
hirsutum, V. corymbosum, and V. angustifolium (Camp 1942, 1945; Vander Kloet 1977, 1980).

Hybrids between V. myrtilloides and V. boreale have been found ina commercial lowland blueberry field at
Frizzleton, Inverness County, Nova Scotia. Hall and Aalders (1962) reported that a comparison of the

342 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 5— Means and standard deviations for selected attributes of the diploid, lowbush species in Vaccinium & Cyanococcus

Taxon
characters V. darrowii V. tenellum V. pallidum . V. myrtilloides V. boreale
Plant height (cm) 55+39 30+11 37+14 35+14 4+2
Habit Evergreen Deciduous Deciduous Deciduous Deciduous
Leaf blade Glaucous Glandular Pale Green Green
Glabrous Pubescent Glabrous Pubescent Glabrous
Leaf width (mm) 4+] 10+2 17+4 12+4 4+1.5
Leaf margin Entire + Entire Entire or serrate Entire Serrate
Corolla length (mm) Sae7) Yar? Yacs} 4+] 4+]
Pollen tetrads (um) 44+2 3642 Siite B23 3343
Seed weight (mg/ 100 seeds) 47+8 39+6 34+6 26+7 se)
Habitat Pine flatwoods Pine flatwoods Open oak woods Taiga Headlands
Alpine/ arctic
tundra
Geographical range
Latitude 27°N-32°N 30°N-37°N 33°N-44°N 39° N-61°N 44°N-S8°N
Longitude 80° W-91°W 76° W-88° W 70° W-94° W 60° W-125°W 53° W-74°W

suspected natural hybrids with artificially produced ones showed both to be intermediate between the parents in
general plant form and leaf shape.

11. Response Behavior

(a) Fire. In natural communities or managed forests, V. myrtilloides survives wild fires or controlled burning
below ground (Flinn and Wein 1977). Recolonization occurs by rhizome sprouting and the density of new
shoots varies significantly depending upon whether the burn occurs in May or September. According to these
same authors density of new emerging shoots is much higher after a September burn and can be explained as a
result from accumulation of food reserves in the rhizomes during the growing season. Commercial stands which
are burn-pruned every second year showed a marked decrease in V. myrtilloides abundance (Chandler and
Hyland 1941). Currently we are investigating the optimum burn cycle for this species.

(b) Grazing and harvesting. Grazing by deer removes the flower buds or terminal vegetative buds which in turn
stimulates dormant tissue further down the stem to grow. With flowers removed more metabolites and nutrients
are directed to vegetative growth. The destruction of the terminal growing point of either the shoot or rhizome
will destroy apical dominance and give rise to one or more new growing sites. The rakes used in fruit harvesting

-occasionally pull up pieces of rhizomes and there is considerable damage to flower buds. This poses no problem
in commercial stands which are generally burn-pruned following the taking of a single crop.

(c) Flooding. Flooding during the growing season has never been reported as a problem. Even on bogs or
around the margins of lakes V. myrtilloides is normally found growing on the tops of hummocks above the
water table (Moss 1953). Thus it may not tolerate inundation.

(d) Drought. Inthe Maritime Provinces there is generally sufficient rainfall to meet requirements. Indry years a
substantial rain of about 2 cm three to four weeks before harvest will have a favorable effect on berry size and
harvest yields. Over the rest of its transcontinental range the species frequently grows on thin and/ or coarse-
textured soil, and drought symptoms such as leaf-browning and fruit abortion are more common.

(e) Herbicides. Vaccinium myrtilloides is resistant to Asulam and Terbacil at recommended rates for control-
ling, respectively, P. aquilinum, and grasses and sedges. Dicamba and 2,4-D applied after leaf abscission of V.
myrtilloides for control of K. angustifolia may reduce the number of shoots of V. myrtilloides the year following
application. Generally the rhizomes are not affected and growth is normal the second year.

12. Relationship to Man

Fernald (1950) called this species “Sour-top”, an uncomplimentary name; we prefer his alternate colloquial
name, “Velvet-leaf.” Differences in the size, shape, color, and taste of berries exist among clones, but those of V.
myrtilloides, when ripe, are equally as sweet and pleasant as those of V. angustifolium. The crop of blueberries
in New Brunswick is about 2 million kg annually and of this perhaps 30% is from V. myrtilloides (Hall et al.
1979). Most of the berries are sold to processors in Maine who sell them as pie filling or as a component of

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 343

muffin mixes. Both V. angustifolium and V. myrtilloides are much sought after in the pine forests of the Great
Lakes region, but not commercially as yet, except at Sudbury. The Cree nation makes use of western
populations.

Literature Cited

Aalders, L. E., and I. ¥V. Hall. 1961. Pollen incompatability and fruit set in lowbush blueberries. Canadian Journal of
Genetics and Cytology 3: 300-307. ;

Aalders, L. E., and I. V. Hall. 1962. The inheritance of white fruit in the velvet-leaf blueberry, Vaccinium myrtilloides
Michx. Canadian Journal of Genetics and Cytology 4: 90-91.

Andersen, P. C., D. W. Buchanan, and L. G. Albrigo. 1979. Water relations and yields of three Rabbiteye Blueberry
Cultivars with and without drip irrigation. Journal of the American Society for Horticultural Science 104: 731-736.

Atlas of Canada. 1974. Ministry of Energy, Mines and Resources. MacMillan Co. of Canada Ltd., Toronto, Ontario.
254 pp.

Bell, HP. 1957. The development of the blueberry seed. Canadian Journal of Botany 35: 139-149.

Bell, H. P., and J. Burchill. 1955. Winter resting stages of certain Ericaceae. Canadian Journal of Botany 33: 547-561.

Bent, F. C., and S. P. Vander Kloet. 1976. Reports on Ericaceae. Jn IOPB chromosome number reports L, II. Taxon 25:
344-345.

Boivin, B. 1966. Enumération des plantes du Canada. Naturaliste Canadien 93: 371-437.

Boulanger, L. W.,G. W. Wood, E. A. Osgood, and C. O. Dirks. 1967. Native bees associated with the lowbush blueberry in
Maine and Eastern Canada. Maine Agricultural Experiment Station and Canada Agricutlural Research Station Bulletin
126. 22 pp.

Brown, R. J. E. 1967. Permafrost in Canada. Ministry of Energy, Mines and Resources, Geological Survey and National
Research Council, Division of Building Research, Publication Number NRC 9769.

Camp, W.H. 1942. The structure of populations in the genus Vaccinium. Brittonia 4: 189-204.

Camp, W. H. 1945. The North American blueberries with notes on other groups of Vacciniaceae. Brittonia 5: 203-275.

Chandler, F. B., and F. Hyland. 1941. Botanical and economic distribution of Vaccinium L. in Maine. Proceedings of the
American Society for Horticultural Science 38: 430-433.

Conners, I. L. 1967. Anannotated index of plant diseases in Canada and fungi recorded on plants in Alaska, Canada and
Greenland. Canada Department of Agriculture, Research Branch, Publication 1251.

Crouch, P. A.,and S. P. Vander Kloet. 1980. Variation in seed characters in populations of Vaccinium § Cyanococcus (the
Blueberries) in relation to latitude. Canadian Journal of Botany 58: 84-90.

Curtis, J. T. 1959. The vegetation of Wisconsin. University of Wisconsin Press, Madison. 657 pp.

Darrow, G. M., W. H. Camp, H. E. Fischer, and H. Dermen. 1944. Chromosome numbers in Vaccinium and related
groups. Bulletin Torrey Botanical Club 71: 498-506.

Devlin, R. M. 1980. Growth inhibitor in cranberry leaves. Cranberries 44: 3 and 12.

Eaton, E. L. 1957. The spread of blueberry seed through manure and by migrating robins. Proceedings of the American
Society for Horticultural Science 69: 293-295.

Eaton, E. L.,and I. V. Hall. 1961. The blueberry in the Atlantic Provinces. Canada Department of Agriculture Publication
754. The Queen’s Printer, Ottawa, Ontario. 36 pp.

Englander, L. 1979. Association of Clavaria fruiting bodies and Ericaceae indicated by field observations and nutrient
exchange phenomena. /n Programmes and Abstracts; Fourth North American Conference on Mycorrhiza. June 24-28,
1979 at Colorado State University, Fort Collins, Colorado.

Fernald, M. L. 1950. Gray’s Manual of botany. 8th edition. American Book Company, New York, New York. 1632 pp.

Flinn, M. A., and R. W. Wein. 1977. Depth of underground plant organs and theoretical survival during fire. Canadian
Journal of Botany 55: 2550-2554.

Freeman, B., L. G. Albrigo,and R. H. Riggs. 1979. Cuticular waxes of developing leaves and fruit of Blueberry, Vaccinium
ashei Reade cv. Bluegem. Journal of the American Society for Horticultural Science 104: 398-403.

Gaudreau, L. 1979. La végétation et les sols des collines Tanginan Abitibi-Ouest, Québec. Laboratoire d’écologie forestiére,
Université Laval, Québec. 391 pp.

Gerloff, G. C.,D. D. Moore, and J. T. Curtis. 1964. Mineral content of native plants of Wisconsin. Wisconsin Agricultural
Experiment Station Research Report 14. 27 pp.

Gorham, E., and A. G. Gordon. 1960. Some effects of smelter pollution northeast of Falconbridge, Ontario. Canadian
Journal of Botany 38: 307-312.

Hall, I. V. 1954. Ecological Studies. In Dominion blueberry substation, Tower Hill, New Brunswick progress report
1949-1953. Canada Department of Agriculture. pp. 18-23.

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. 1963. Note on the effect of a single intensive cultivation on the composition of an old blueberry stand. Canadian
Journal of Plant Science 43: 417-418.

344 THE CANADIAN FIELD-NATURALIST Vol. 95

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. 1962. A natural hybrid between Vaccinium myrtilloides and Vaccinium boreale on Cape
Breton Island. Canadian Field-Naturalist 76: 203-205.

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, N. L. Nickerson, and S. P. Vander Kloet. 1979. The biological flora of Canada. 1. Vaccinium
angustifolium Aiton, Sweet Lowbush Blueberry. Canadian Field-Naturalist 93: 415-430.

Hall, I. V., F. R. Forsyth, L. E. Aalders, and L. P. Jackson. 1970. The physiology of the lowbush blueberry. Economic
Botany 26: 68-73.

Hebda, R. J. 1979. Size, productivity, and paleoecological implications of ericaceous pollen from Burns Bog, southern
Fraser River Delta, British Columbia. Canadian Journal of Botany 57: 1712-1717.

Hersey, R., and S. P. Vander Kloet. 1976. Reports on Ericaceae. In I|OPB chromosome number reports LII. Taxon 25:
342-343.

Hilton, S. A.,and W. G. Barker. 1962. The research status of the lowbush blueberry (Vaccinium spp.). Proceedings of the
16th International Horticultural Congress (Brussels, Belgium). pp. 426-432.

Holmgren, P. K., and W. Keuken. 1974. Index herbariorum. Part |. The herbaria of the world. 6th edition, Oosthoek,
Scheltema and Holkema, Utrecht, Netherlands.

Jeglum, J. K. 1971. Plantindicators of pH and water level in peatlands at Candle Island, Saskatchewan. Canadian Journal
of Botany 49: 1661-1676.

Kabzems, A., A. L. Kosowan, and W. C. Harris. 1976. Mixedwood section in an ecological perspective: Saskatchewan.
Forestry Branch, Department of Tourism and Renewable Resources, Technical Bulletin No. 8.

Krefting, L. W.,and E. I. Roe. 1949. The role of some birds and mammals in seed germination. Ecological Monographs 19:
270-286.

La Roi, G. H. 1967. Ecological studies in the boreal spruce-fir forests of the North American taiga. I. Analysis of the
vascular flora. Ecological Monographs 37: 229-253.

La Roi, G. H., and R. J. Hnatiuk. 1980. The Pinus contorta forests of Banff and Jasper National Parks: a study in
comparative synecology and syntaxonomy. Ecological Monographs 50: 1-29.

LeBlanc, F. 1963. The life-forms of the flora of Mount Yamaska, Rouville County, Quebec. Canadian Journal of Botany 41:
1425-1437.

Leiser, A. T. 1968. A mucilaginous root sheath in Ericaceae. American Journal of Botany 55: 391-398.

LePage, E. 1951. Entites nouvelles dans la flore a Québec. Naturaliste Canadien 78: 341-352.

Lockhart, C. L., and W. M. Langille. 1962. The mineral content of the Lowbush Blueberry. Jn Canadian Plant Disease
Survey. pp. 124-128.

Longley, A. E. 1927. Chromosomes in Vaccinium. Science 66: 566-568.

Loucks, O. L. 1962. A forest classification for the Maritime Provinces. Proceedings of the Nova Scotian Institute of Science
25: 85-167.

Love, A.,and D. Love. 1966. Cytotaxonomy of the alpine vascular plants of Mount Washington. University of Colorado
Studies, Series Biology 24: 1-74.

Marie-Victorin, Frére. 1935. Flore laurentienne. Imprimerie de la Salle, Montréal, Québec.

Marie-Victorin, Frére. 1964. Flore laurentienne. Deuxiéme édition. Les Presses de l’Université de Montréal, Montréal.
Québec. p. 442.

Maxwell, C. W. B., G. W. Wood, and W. Neilson. 1953. Insects in relation to blueberry culture. Jn Dominion blueberry
substation, Tower Hill, New Brunswick progress report 1949-1953. Canada Department of Agriculture. pp. 13-16.

Maycock, P. F., and J. T. Curtis. 1960. The phytosociology of the boreal conifer-hardwood forests of the Great Lakes
region. Ecological Monographs 30: 1-35.

Moore, J. N. 1965. Improving highbush blueberries by breeding and selection. Euphytica 14: 39-48.

Moss, E. H. 1953. Forest communities in northwestern Alberta. Canadian Journal of Botany 31: 212-252.

Nichols, G. E. 1934. The influence of exposure to winter temperatures upon seed germination in various native American
plants. Ecology 14: 364-373.

Palser, B. F. 1961. Studies of floral morphology in the Ericales. V. Organography and vascular anatomy in several United
States species of the Vacciniaceae. Botanical Gazette 123: 79-111.

Parmelee, J. A. 1977. The fungi of Ontario. 11. Erysiphaceae (mildews). Canadian Journal of Botany 55: 1940-1983.

Pearson, V., and D. J. Read. 1973a. The biology of mycorrhiza in the Ericaceae. I. The isolation of the endophyte and
synthesis of mycorrhizas in aseptic culture. New Phytologist 72: 371-379.

Pearson, V.,and D. J. Read. 1973b. The biology of mycorrhiza in the Ericaceae. II. The transport of carbon and phosphorus
by the endophyte and the mycorrhiza. New Phytologist 72: 1325-1331.

Reader, R. J. 1977. Bog ericad flowers: self-compatibility and relative attractiveness to bees. Canadian Journal of Botany
55: 2279-2287.

Reader, R. J. 1979. Flower cold hardiness: a potential determinant of the flowering sequence exhibited by bog ericads.
Canadian Journal of Botany 57: 997-999.

1981 VANDER KLOET AND HALL: VACCINIUM MYRTILLOIDES 345

Richardson, Sir John. 1823. Narrative of a journey to the shores of the Polar Sea in the years 1819, 20, 21 and 22. With an
appendix on various subjects relating to science and natural history. J. Murray, London. 768 pp.

Roland, A. E. 1946. The vegetation of the Annapolis Valley. I. Well-drained sand areas. Acadian Naturalist 2: 1-20.

Rowe, J. S. 1956. Uses of undergrowth plant species in forestry. Ecology 37: 461-473.

Rowe, J.S. 1972. Forest regions of Canada. Department of the Environment, Canadian Forestry Service, Publication
Number 1300. 172 pp.

Rowe, J. S., and G. W. Scotter. 1973. Fire in the boreal forest. Quaternary Research 3: 444-464.

Ryan, A. G. 1974. Shrubs of Newfoundland. Parks Division, Department of Tourism, Province of Newfoundland.

Sanderson, M. 1948. The climates of Canada according to the new Thornthwaite classification. Scientific Agriculture 28:
507-517.

Scoggan, H. J. 1950. The flora of Bic and the Gaspé peninsula, Quebec. National Museum of Canada Bulletin No. 115. The
King’s Printer, Ottawa, Ontario.

Scoggan, H. J. 1978. The flora of Canada. National Museum of Canada Publications in Botany No. 7. National Museums
of Canada, Ottawa, Ontario. 1711 pp.

Segadas-Vianna, F. 1955. Ecological study of the peat bogs of Eastern North America. I]. The Chamaedaphne calyculata
community in Quebec and Ontario. Canadian Journal of Botany 33: 647-684.

Smith, D. W. 1962. Ecological studies of Vaccinium species in Alberta. Canadian Journal of Plant Science 42: 82-90.

Szczawinski, A. F. 1962. The Heather Family (Ericaceae) of British Columbia. British Columbia Provincial Museum,
Department of Recreation and Conservation. The Queen’s Printer, British Columbia.

Taylor, R. L.,and B. MacBryde. 1977. Vascular plants of British Columbia: a descriptive resource inventory. University of
British Columbia Technical Bulletin No. 4. p. 221.

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., and I. V. Hall. 1971. Nutrient levels in leaf and soil samples from three cranberry bogs in the Annapolis
Valley of Nova Scotia. Cranberries 36: 11-12.

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. 1972. The North American blueberries revisited: a taxonomic study of Vaccinium § Cyanococcus Gray.
Unpublished Ph.D. thesis, Queen’s University at Kingston, Ontario.

Vander Kloet, S. P. 1976. Nomenclature, taxonomy, and biosystematics of Vaccinium section Cyanococcus (the blueber-
ries) 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. The taxonomic status of Vaccinium pallidum, the hillside blueberries including Vaccinium
vacillans. Canadian Journal of Botany 56: 1559-1574. ,

Vander Kloet, S. P. 1980. The taxonomy of the highbush blueberry, Vaccinium corymbosum. Canadian Journal of Botany
58: 1187-1201.

Viereck, L. A., and C. T. Dyrness. 1980. A preliminary classification system for vegetation of Alaska. United States
Department of Agriculture, Forest Service. General Technical Report PNW-106. 38 pp.

Wali, M. K., and V. J. Krajina. 1973. Vegetation-environment relationships of some sub-boreal spruce zone ecosystems in
British Columbia. Vegetatio 26: 237-381.

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, G. W. 1956. Note on injury to blueberry sprouts by the blueberry thrips, Frankliniella vaccinii Morgan (Thysanop-
tera: Thripidae). Canadian Journal of Agricultural Science 36: 510.

Wood, G. W. 1960. Note on the occurrence of two species of thrips (Thysanoptera: Thripidae) on low-bush blueberry in
New Brunswick and Nova Scotia. Canadian Entomologist 92: 757-758.

Wood, G. W. 1962. The blueberry maggot in the Maritime Provinces. Canada Department of Agriculture Publication 1161.

Wood, G. W. 1965. Evidence in support of reduced fruit set in lowbush blueberry by pollen incompatibility. Canadian
Journal of Plant Science 45: 601-602.

Wood, G. W. 1970. Survival of blueberry casebeetle adults in burned blueberry fields. Journal of Economic Entomology 63:
1364.

Wood, G. W. 1979. Insects. Jn Lowbush blueberry production. Agriculture Canada Publication 1477. pp. 24-25.

Wood, G. W., and W. T. A. Neilson. 1956. Notes on the black army cutworm, Actebra fennica (Tausch.) (Lepidoptera:
Phalaenidae), a pest of low-bush blueberry in New Brunswick. Canadian Entomologist 88: 93-96.

Wood, G. W., and F. A. Wood. 1963. Nectar production and its relation to fruit set in the lowbush blueberry. Canadian
Journal of Botany 41: 1675-1679.

Wulff, E. V. 1943. Historical plant geography. Chronica Botanica Company, Waltham, Massachusetts. 223 pp.

Received 2 July 1980
Accepted 22 February 1981

Notes

Attempted Avian Predation by a Canadian Toad,

Bufo americanus hemiophrys

FRANCIS R. COOK! and JOYCE C. COOK?

'Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario K1A 0M8

?R.R. #3, North Augusta, Ontario KOG 1RO

Cook, Francis R.,and Joyce C. Cook. 1981. Attempted avian predation by a Canadian Toad, Bufo americanus hemiophrys.

Canadian Field-Naturalist 95(3): 346-347.

Vertebrates are rarely taken by toads ( Bufo), which are generally characterized as avoiding relatively large prey. Anexception
was a 62-mm (snout-vent) Canadian Toad, Bufo americanus hemiophrys, found near Beausejour, Manitoba, attempting to
swallow a young Red-winged Blackbird, Agelaius phoeniceus, which had apparently recently fallen from a nearby nest.

Key Words: Bufo americanus hemiophrys, Canadian Toad; predation, Red-winged Blackbird, Agelaius phoeniceus.

The feeding pattern of toads (Bufo) has been char-
acterized by Clarke (1974) as largely that of a predator
on small, often armored or noxious, insects, particu-
larly beetles, ants, and (in the tropics) termites. Clarke
acknowledged, however, that some earlier authors
regarded toads in indiscriminate feeders. He also
stressed that toads generally eat many small items
rather than a few large ones, largely independent of
the size of the toad (juvenile or adult). However,
Heatwole and Heatwole (1968), and Bragg (1957)
observed some increase in prey size with increase in
toad size.

Vertebrates have rarely been reported as food items
for toads. Campbell and Davis (1968) reported a non-
gravid female Gulf Coast Toad, Bufo valliceps, which
contained a Fence Lizard, Sceloporus undulatus, and
another which contained a juvenile Fowler’s Toad, B.
woodhousei fowleri. In their review of the literature
they found only one previous report, Smith and Bragg
(1949), who observed a captive juvenile B. woodhou-
sel fowleri eat a small lizard in a terrarium. Over-
looked by the authors were reports that a Giant Toad,
B. marinus, had eaten a small frog (Weber 1938),
“small batrachians” (= frogs) (Barbour 1930: 76) and
ducklings and rats (Myers 1931: 25). Also, Alexander
(1964) recorded that Giant Toads ate a variety of items
from dog food to overripe avocados: an exception to
the common Bufo response of preying only on moving
objects. Subsequently, Hahn (1976) and Hughes
(1978) cited additional reports of vertebrates taken by
the Asian and African toads Bufo melanostictus, B.
regularis, and B. funereus.

Because records of toads preying on vertebrates,
especially relatively large ones, are infrequent, our

observation indicates that toads cannot be completely
stereotyped regarding prey size.

On 24 June 1968 between 16:15 and 19:00 during a
field study of toads in southeastern Manitoba, a
lethargic Canadian Toad, B. americanus hemiophrys
(for nomenclature see Cook 1978) was found at the
edge of a water-filled roadside ditch 1.6 km north of
Beausejour, Manitoba. On turning the animal over,
we found most of a young bird, later identified as a
Red-winged Blackbird, Agelaius phoeniceus, pro-
truding from the toad’s mouth. Only the bird’s head
and part of its neck had been swallowed. Although
apparently near death when found, the toad revived
after the bird was removed. The toad measured
62 mm snout to vent and was a female with relatively
small ovarian eggs, both pigmented and unpigmented.
Comparative sizes of toad and bird are shown in
Figure |. Both were preserved at capture (NMC
11218). Among cattail (7yphus sp.) stems, 15 malong
the ditch, was the nest of a Red-winged Blackbird,
containing three young, about the size of the one the
toad had seized. Godfrey (1966) gives the usual
number of eggs for this species as four (range three to
five). As the nest was about 0.3 mabove the water, it is
most likely that one nestling had fallen out of the nest
into the water where its struggles attracted the toad.
Bragg (1957) suggested that the nature of a potential
prey’s movements was important in whether it was
attacked or not. Possibly its struggles were like those
of a lepidopteran, fluttering at the surface of the
water.

The only existing study of B. a. hemiophrys feeding
(Moore and Strickland 1954) fits this taxon into the
general pattern summarized by Clarke (1974). Lepi-

346

1981

Pra ; 2 3 4 § pea! < Ay a ee Ble oe 9 oO
Hane TOT TETcaTTTA TT TTT TT TPT TT TET

FIGURE |. Comparison of the size of the female Canadian
Toad, Bufo americanus hemiophrys, 62 mm snout-
vent, with its prey, a nestling Red-winged Blackbird,
Agelaius phoeniceus.

doptera feeding is apparently rare as only two are
noted from the 16 adult toads examined by Moore
and Strickland (1954). However, much of the range of
this toad is in moderately arid environment of aspen
parkland and the northern periphery of the short-
grass prairie. It shows morphological trends to arid
adaptations (Cook 1978). At least occasional oppor-
tunistic feeding on relatively large prey (such as Lepi-
doptera) could be an advantage in such habitats where
the potential active period is limited by periodic arid-
ity and generally cool evenings during a relatively
short annual activity period (May through Sep-
tember). This might be particularly true of females
that are maturing eggs for the subsequent breeding
season. Later examination of the toad’s stomach
revealed that it was moderately full, containing beetle
remains (wings etc.) and soft-bodied insects. Heat-
wole and Heatwole (1968) indicated that size of prey
attacked tended to decrease with amount of food pre-
viously taken, but this does not seem to have held true
in this individual.

NOTES

347

Weare indebted to W. E. Godfrey and H. Ouellet of
the National Museum of Natural Sciences Ornithol-
ogy Section, who confirmed the identification of the
avian remains; and to James A. Johnston, Herpetol-
ogy Section, who photographed the preserved
specimens.

Literature Cited

Alexander, T. R. 1964. Observations on the feeding behav-
ior of Bufo marinus (Linné). Herpetologica 20(4):
255-259.

Barbour, T. 1930. Some faunistic changes in the Lesser
Antilles. Proceedings of the New England Zoological Club
11: 73-85.

Bragg, A. N. 1957. Some factors in the feeding of toads.
Herpetologica 13(3): 189-191.

Campbell, P. M., and W. K. Davis. 1968. Vertebrates in
stomachs of Bufo valliceps. Herpetologica 24(4): 327-328.

Clarke, Raymond D. 1974. Food habits of toads, genus
Bufo (Amphibia: Bufonidae). The American Midland
Naturalist 91(1): 140-147.

Cook, Francis R. 1978. An analysis of toads of the Bufo
americanus group 1n a contact zone in central northern
North America. Ph.D. thesis, University of Manitoba,
Winnipeg, Man. 232 pp.

Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203. 428 pp.

Hahn, Donald E. 1976. Worm snakes in the diet of a toad,
Bufo melanostictus. Herpetological Review 7(4): 167.
Heatwole, Harold, and Audry Heatwole. 1968. Motiva-
tional aspects of feeding behavior in toads. Copeia

1968(4): 692-698.

Hughes, B. 1978 (Letter to Editor). Herpetological Review
9(1): 23-24.

Moore, J. E., and E. H. Strickland. 1954. Notes on the
food of three species of Alberta amphibians. American
Midland Naturalist 52(1): 221-224.

Myers, J. G. 1931. A preliminary report onan investigation
into the biological control of West Indian insect pests.
Empire Marketing Board Publication (London) 42:
1-173.

Smith, C. C., and A. N. Bragg. 1949. Observations on the
ecology and natural history of Anura. VII. Food and
feeding habits of the common species of toads in Okla-
homa. Ecology 30: 333-349.

Weber, Neal A. 1938. The food of the Giant Toad, Bufo
marinus (L.), in Trinidad and British Guiana with special
reference to the ants. Annals of the Entomological Society
of America 31: 499-503.

Received 13 March 1980.
Accepted 20 January 1981.

348 THE CANADIAN FIELD-NATURALIST

Vol. 95

Daily Movements of Red Squirrels, Tamiasciurus hudsonicus

STEVE G. FANCY

LGL Alaska Research Associates, P.O. Box 80607, Fairbanks, Alaska 99708

Fancy, S. G. 1981. Daily movements of Red Squirrels, Tamiasciurus hudsonicus. Canadian Field-Naturalist 95(3): 348-350.

My study of the daily movements of Red Squirrels ( Tamiasciurus hudsonicus) near Atlin, British Columbia, in 1978 showed
that the shortest movements were in the prebreeding season (averaging 34.3 m, 9 Q ; 88.1 m, dd _). Males made the greatest
movements (mean to 169.9 m) during breeding and gestation whereas females made long movements (mean to 190.1 m)
during lactation and postweaning. Both the breeding cycle and the availability of food supplies influence the daily movements.

Key Words: Red Squirrel, Tamiasciurus hudsonicus; daily movements, British Columbia.

The movements of Red squirrels (Tamiasciurus
hudsonicus) are relatively easy to study because the
animals are fairly numerous, territorial, diurnal, and
vocal, and thus are easy to locate. Study of this species
may help us to understand the daily movements of
other, less common, and wider-ranging territorial
species. Although several studies on Red Squirrel
ecology have included information on movements,
there has been no detailed quantitative study on the
daily movements of Red Squirrels. By “territory” I
refer to a more or less clearly defined area where an
animal tends to segregate itself from other individuals
of its species (Fitzwater 1941).

Methods

I studied Red Squirrel movements during the spring
and summer 1978 near Atlin, British Columbia
(59°38’N, 133°19’W). The vegetation on the 1.03-km?2
study area consisted of patches of White Spruce
(Picea glauca) and Lodgepole Pine (Pinus contorta)
separated by muskegs and dense stands of willows
(Salix spp.) (Fancy 1979).

Livetrapped squirrels were marked with fur dye and
metal ear tags. The location of marked squirrels on the
gridded study area was recorded during daily patrols
on foot. Although it was not possible to cover each
part of the study area equally, all territories were
visited at least once each field day. Observations of
squirrel behavior indicated that my presence had no
effect on squirrel movements. These methods resulted
in 4150 observations, including 502 trap captures, of
94 marked squirrels.

The study period was divided into five seasons
based on the female reproductive cycle. The greatest
distance between any two locations at whicha particu-
lar squirrel was seen on any given day was used to
compare movements of Red Squirrels by sex and
season. For each squirrel during each season, a mean
distance was calculated from the daily distances.
These mean distances for each squirrel were used in
subsequent analyses.

Results and Discussion

A mean distance was not available for every squirrel
each season, as some squirrels died or left the study
area, while others were first observed after the study
began. An analysis of variance was precluded by these
missing data, by a significant interaction between sex
and season, and by unequal variances, even after a
square root transformation was used. A Kruskal-
Wallis test (Hull and Nie 1979) was used, therefore, to
test seasonal differences in the mean distance between
sexes, while pairwise /-tests assuming unequal varian-
ces (Sokal and Rohlf 1969: 374) tested the differences
in movements between sexes for each of the five sea-
sons (Table 1).

For both males and females, the five seasonal
means were significantly different (Table 1). Males
made the greatest movements during the gestation
and breeding seasons, while females made longer
movements during the lactation and postweaning sea-
sons. The differences between the daily movements of
males and females was significant in all seasons except
during the postweaning season. Males moved further
than females during the prebreeding, breeding, and
gestation seasons, while females moved further than
males during the lactation and postweaning seasons.

Daily movements of Red Squirrels except during
the breeding season are mainly associated with food
gathering and territorial defence (C. C. Smith 1968).
C. C. Smith (1968) found that in the daylight hours,
adult squirrels spend about 80% of the time in food-
related activities, and less than 1% in territorial
defense. Because squirrels in habitat with a poor food
supply would need to travel greater distances to meet
their energy demands than would squirrels in food-
rich habitats, we would expect to find a high positive
correlation between the food supply of an area and the
mean daily distance of territory owners on that area.
Studies by Kemp and Keith (1970) and Rusch and
Reeder (1978) support this because the greatest
movements they detected were in poor-quality Trem-
bling Aspen (Populus tremuloides) areas, while shor-

1981

NOTES

349

TABLE 1—Comparison of mean daily distance (m) moved by squirrels using Kruskal-Wallis tests for seasonal differences in
distance moved by each sex, and /-tests assuming unequal variances for differences between sexes

Males Females t-test
Season n MeantSD n MeantSD IP
Pre-breeding! 15 88.1+53.4 8 34.3+33.9 < 0.05
Breeding? 19 156.9+114.3 11 62.4+27.6 < 0.05
Gestation3 16 169.9+125.1 12 79.34+43.7 < 0.05
Lactation? 21 99 .6-44.6 16 190.1+151.3 < 0.02
Postweaning> 19 106.8+161.6 12 151.4+93.7 ns
Bartlett’s test P< 0.001 P< 0.001
Kruskal-Wallis statistic P=0.029 P< 0.001

122 March — 11 April.
212 April —9 May.

310 May — 31 May.

4] June — 1 August.

52 August — 28 August.

ter movements were detected in White Spruce forests.
M. C. Smith (1968) reported that home ranges of
squirrels following a cone crop failure were consider-
ably larger than in preceeding years, as squirrels tra-
veled greater distances in search of food. I found a
spring density of 0.5 adult squirrels per hectare; this is
considerably lower than that found in other studies
(Brink 1964; Wood 1967; C. C. Smith 1968; Davis
1969; Rusch 1970). Because 27.7% of my study area
was covered by muskegs and dense willows, the avail-
able food supply in my study area was probably lower
than that in the other areas studied.

During the breeding season, food is no longer the
critical resource for male squirrels. Estrous females,
receptive on only one day (C. C. Smith 1968; Rusch
and Reeder 1978), represent a temporally and spa-
tially limited resource. Because receptive females
remain close to their middens during this season
(Fancy 1979), males must move over large distances to
increase the probability of encountering an estrous
female.

The longer movements made by females during the
lactation and postweaning seasons were made primar-
ily by parturient females, presumably as a result of the
increased energy demands from the birth and caring
of their young. During the lactation season, lactating
females each averaged 7.3 daily movements exceeding
100 m, while each male and barren female averaged
4.2 movements greater than 100 m during this season.

The results of this and other studies (C. C. Smith
1968; M. C. Smith 1968; Kemp and Keith 1970; Rusch
and Reeder 1978) indicate that daily movements of
Red Squirrels are influenced by the breeding cycle and
the availability of food supplies within an area.
Further work with this and other territorial species
may enable us to predict the extensiveness of daily

movements based on the available food supply of an
area.

Acknowledgments

This paper is based ona portion of my M.Sc. thesis.
I am indebted to Archie Mossman for his guidance
during the fieldwork and for his comments on the

paper.

Literature Cited

Brink, C. H. 1964. Spruce seed as a food of the squirrels
Tamiasciurus hudsonicus and Glaucomys sabrinus in inte-
rior Alaska. M.Sc. thesis, University of Alaska, Fair-
banks. 73 pp.

Davis, D. W. 1969. The behaviorand population dynamics
of the Red Squirrel ( Tamiasciurus hudsonicus) in Saskat-
chewan. Ph.D. thesis, University of Arkansas, Fayette-
ville. 229 pp. Diss. Abstr. 30: 902-B.

Fancy, S. G. 1979. Dispersal and daily movements of Red
Squirrels (Tamiasciurus hudsonicus). M.Sc. thesis, Hum-
boldt State University, Arcata, California. 60 pp.

Fitzwater, W. D., Jr. 1941. The Red Squirrel: territorial-
ism, activity, census methods. M.Sc. thesis, New York
State Coll. Forestry, Syracuse. 117 pp.

Hull, C. H., and N. H. Nie. 1979. SPSS Update. McGraw-
Hill, New York. 238 pp.

Kemp, G. A.,and L. B. Keith. 1970. Dynamics and regula-
tion of Red Squirrel (Tamiasciurus hudsonicus) popula-
tions. Ecology 51: 763-779.

Rusch, D. A. 1970. Population ecology and behavior of
Red Squirrels in central Alberta. M.Sc. thesis, University
of Wisconsin, Madison. 94 pp.

Rusch, D. A.,and W. G. Reeder. 1978. Population ecology
of Alberta Red Squirrels. Ecology 59: 400-420.

Smith, C. C. 1968. The adaptive nature of social organiza-
tion in the genus of tree squirrels Tamiasciurus. Ecological
Monographs 38: 31-63.

350 THE CANADIAN FIELD-NATURALIST

Smith, M. C. 1968. Red Squirrel responses to spruce cone
failure in interior Alaska. Journal of Wildlife Manage-
ment 32: 305-317.

Vol. 95

Red Squirrel ( Tamiasciurus hudsonicus) in Wood Buttalo
National Park. M.Sc. thesis, University of Saskatchewan,
Saskatoon. 97 pp.

Sokal, R.R., and F.J. Rohlf. 1969. Biometry. W.H.
Freeman & Co., San Francisco. 776 pp.
Wood,T. J. 1967. Ecology and population dynamics of the

Received 30 October 1980
Accepted 27 January 1981

Fate of Overwintered Clutches of the Common Snapping Turtle
(Chelydra serpentina) in Algonquin Park, Ontario

MARTYN E. OBBARD and RONALD J. BROOKS
Department of Zoology, University of Guelph, Guelph, Ontario NIG 2W1

Obbard, Martyn E.,and Ronald J. Brooks. 1981 Fate of overwintered clutches of the Common Snapping Turtle (Chelydra
serpentina) in Algonquin Park, Ontario. Canadian Field-Naturalist 95(3): 350-352.

The frequency of successful overwintering of Common Snapping Turtle (Chel/ydra serpentina) clutches in Algonquin Park,
Ontario, is very low. In a 4-yr study, 42 of 257 clutches emerged in the fall of the year of oviposition. Of the 215 clutches that
did not emerge in fall, only | overwintered successfully. This low success of overwintering clutches may be animportant factor
limiting the northern distribution of the Common Snapping Turtle.

Key Words: Common Snapping Turtle, Chelydra serpentina; nesting, delayed emergence, Algonquin Park, limiting factors.

Delayed emergence or overwintering of young in
the nest occurs regularly in many species of turtles
(Gibbons and Nelson 1977). There are few reports
(Toner 1940; Bleakney 1963; Ernst 1966), however, of
clutches of the Common Snapping Turtle (Chelydra
serpentina) surviving over winter and no indication of
how often this phenomenon might occur in a popula-
tion. Therefore, in this study, we examined the fre-
quency of successful overwintering in clutches of the
Common Snapping Turtle in Algonquin Park,
Ontario.

Study Area and Methods

From 1976 to 1979, we monitored nesting areas
used by female snapping turtles along the North
Madawaska River in Algonquin Park (45°35’N,
78°30’W) (see Obbard and Brooks 1979, 1980 fora
description of the study area). A number of potential
nesting areas was checked daily, morning and even-
ing, throughout the nesting period each year, and the
location of clutches was recorded. Each year the fate
of 60-70 clutches was followed, and these clutches
were checked frequently until late fall. Whenever pos-
sible, clutches that did not emerge in the year of ovipo-
sition were excavated and examined in May of the
following year. The viability of hatchlings and of
young in pipped eggs was recorded. If a clutch con-
tained only unpipped eggs when excavated, several
eggs were opened and viability of embryos was noted.

Each year, a sample of unpipped eggs was retained in
the laboratory to determine whether any young would
hatch.

Results and Discussion

The fate of 257 clutches studied is summarized in
Table 1. Only 42 clutches emerged in September or
October in the year of oviposition. Of the remaining
215 clutches, 129 were excavated in spring and the rest
were either destroyed by predators or were not
located. Only | of the 129 overwintered clutches that
were examined contained live hatchlings. In all other
cases, hatchlings and young in pipped eggs were dead.
All unpipped eggs that were opened contained dead
embryos, and no young hatched from eggs retained in
the laboratory. However, the possibility remains that
some unpipped eggs contained living embryos since
not all eggs were opened and examined. There was
considerable yearly variation in the frequency of suc-
cessfully emerging clutches and in the stage of devel-
opment reached by embryos in unhatched eggs. Fac-
tors affecting development and emergence will be
considered elsewhere.

The single clutch that had live hatchlings when
excavated in spring was laid 18 June 1978 on the
banks of the North Madawaska River 2.3 m from the
concrete abutment of a highway bridge. No hatchlings
had emerged from the nest by 2 November 1978 when
it was last checked. On 5 May 1979, there was still no

1981

NOTES 351

TABLE |—Fate of 257 clutches of the Common Snapping Turtle in Algonquin Park, Ontario

Total located Evidence of
during the emergence Destroyed by

Year nesting period in fall predators
1976 64 4 9
1977 69 6 3)
1978 60 20 10
1979 64 12 26

DSI 42 48

evidence that any young had emerged, so the clutch
was uncovered. The nest contained 16 live hatchlings,
11 dead hatchlings, and 3 infertile eggs. All young had
hatched and were free of the egg shell, and all had lost
the caruncle, but had unhealed yolk sac scars. There
appeared to be no correlation between depth in the
soil where the hatchlings were found and rate of survi-
val. However, the sandy soil had been frost-free for
some time so live hatchlings may have moved verti-
cally in the nest chamber.

Frequency of successful overwintering of Common
Snapping Turtle clutches in this study was only | in
129 and would be even lower if one considered all
clutches laid. It is likely that this rare occurrence was
fortuitous. In the case reported here, the clutch was
probably protected from lethal low temperatures by
deep snow cover accumulated from snow-clearing
operations on the bridge. However, the actual cause of
overwinter mortality is not known, and other factors
suchas lack of oxygen, starvation, or accumulation of
waste products in unpipped eggs may be responsible.

It has been suggested that overwintering in the nest
by hatchling turtles is an adaptation allowing survival
in northern portions of a species’ range (Carr 1952).
For example, hatchling Painted Turtles (Chrysemys
picta) commonly overwinter in the nest near Ann
Arbor, Michigan (Sexton 1957). Overwintering, in the
nest is common in many species of turtles in the south-
ern and central United States (Gibbons and Nelson
1977), whereas overwintering in the northern regions
has been reported as unsuccessful in the Stinkpot
(Sternotherus odoratus) in Michigan (Risley 1933),
the Spiny Softshell ( Trionyx spiniferus) in Minnesota
(Breckenridge 1960), and the Common Snapping Tur-
tle in South Dakota (Hammer 1970). These and our
data make it unlikely that delayed emergence is an
adaptation to northern climates in all turtles. As Gib-
bons and Nelson (1977) emphasized, the costs of
immediate emergence should be weighed against the
costs of delayed emergence in each species. If costs of
the latter are high, as found in our study, then selec-
tion should favor early dates of oviposition and fall
emergence in the year of oviposition.

Overwintered clutches

Total Hatchlings
analyzed in Only unpipped and pipped Live
spring eggs eggs hatchlings
22 20 2 0
39 26 13 0
4] 6 35 |
27 15 12 0
129 67 62 1

Common Snapping Turtle hatchlings usually
emerge in the fall in Michigan (Sexton 1957), South
Dakota (Hammer 1970), and Ontario (this study), and
successful overwintering is probably very rare in areas
where frost extends below the depth of the nest. The
inability of hatchlings to overwinter successfully in the
nest is probably a major factor limiting the northern
distribution of the Common Snapping Turtle, and
possibly other turtle species.

Acknowledgments

This study was supported by grants from the Cana-
dian National Sportsmen’s Fund (grant 3-R 39) and
the Ontario Ministry of Natural Resources. We thank
the Ontario Ministry of Natural Resources for per-
mission to work in Algonquin Park and for use of the

- facilities of the Wildlife Research Station. We thank

G. Nancekivell, D. Schluter, R. Stewart, D. Oliver, A
Green, R. Moses, T. Friesen, I Ross, and D. Campbell
for assistance in the field.

Literature Cited

Bleakney, J. S. 1963. Notes on the distribution and life his-
tories of turtles in Nova Scotia. Canadian Field-Naturalist
77: 67-76.

Breckenridge, W. J. 1960. A Spiny Soft-shelled Turtle nest
study. Herpetologica 16: 284-285.

Carr, A. 1952. Handbook of turtles. Cornell University
Press, Ithaca, N.Y. 542 pp.

Ernst, C. H. 1966. Overwintering of hatchling Chelydra
serpentina in southeastern Pennsylvania. Philadelphia
Herpetological Society Bulletin 14(2): 8-9.

Gibbons, J. W.,and D. H. Nelson. 1977. The evolutionary
significance of delayed emergence from the nest by hatc-
hling turtles. Evolution 32: 297-303.

Hammer, D. A. 1970. Reproductive behavior of the Com-
mon Snapping Turtle. Canadian Herpetologists’ Society
Quarterly 1(4): 9-13

Obbard, M. E., and R. J. Brooks. 1979. Factors affecting
basking in a northern population of the Common Snap-
ping Turtle, Che/ydra serpentina. Canadian Journal of
Zoology 57: 435-440.

Qbbard, M. E.,and R. J. Brooks. 1980. Nesting migrations
of the Common Snapping Turtle (Che/ydra serpentina).
Herpetologica 36: 158-162.

352 THE CANADIAN FIELD-NATURALIST Vol. 95

Toner, G. C. 1940. Delayed hatching in the Snapping Tur-
tle. Copeia 1940: 265.

Risley, P. L. 1933. Observations on the natural history of
the Common Musk Turtle, Sternotherus odoratus
(Latreille). Papers Michigan Academy Science, Arts, and
Letters 17: 685-711.

Sexton, O. J. 1957. Notes concerning turtle hatchlings.
Copeia 1957: 229-230.

Received 16 September 1980
Accepted 27 January 1981

Predation and Caching of Seabirds by Red Foxes (Vulpes vulpes)
on Baccalieu Island, Newfoundland

A. D. MACCARONE and W. A. MONTEVECCHI

Department of Psychology, Memorial University of Newfoundland, St. John’s, Newfoundland A1B 3X9

Maccarone, A. D., and W. A. Montevecchi. 1981. Predation and caching of seabirds by Red Foxes (Vulpes vulpes) on
Baccalieu Island, Newfoundland. Canadian Field-Naturalist 95(3): 352-353.

Red Foxes (Vulpes vulpes) prey on and cache many burrow- and cliff-nesting seabirds on Baccalieu Island, Newfoundland,
possibly for use in winter when food 1s scarce or unavailable. The large seabird population do not appear to be threatened by
the small number of foxes on the island, although the situation is potentially dangerous for burrow-nesting species (Leach’s

Storm-Petrels (Oceanodroma leucorhoa), Atlantic Puffins (Fratercula arctica)).

Key Words: seabirds, predation, foxes, caches, Newfoundland, Leach’s Storn-Petrel, Atlantic Puffin, Red Foxes.

Nesting on cliffs and in burrows are considered to
be antipredator adaptations (e.g. Tuck 1961). Both
aerial and terrestrial predators, however, are often
successful against these birds (Nettleship 1972; Mon-
tevecchi 1978; Jones and Byrd 1979). In this paper
cases of predatory and caching activities by Red
Foxes (Vulpes vulpes) on cliff— and burrow-nesting
seabirds on Baccalieu Island, Newfoundland
(48°07’N, 54°12’W), are described.

Foxes were first seen on the island in winter
1959-1960, when pack ice jammed the 3.2 km channel
separating the island from the mainland (E. Blundon
and P. Rice, Baccalieu Island, personal communica-
tion). Currently they are the only mammals that
inhabit the island. Baccalieu Island (6 X I km) sup-
ports hundreds of thousands of breeding pairs of
Leach’s Storm-Petrels (Oceanodroma _ leucorhoa,
possibly the largest colony in the world), and many
thousands of Black-legged Kittiwakes (Rissa tridac-
tyla), Atlantic Puffins (Fratercula arctica), and
Common Murres (Uria aalge) (Brown et al. 1975).

In 1976-1978, we found expanses (hundreds of
square metres) of petrel nesting ground on grassy
slopes dug out by foxes (Figure 1). In 1978, we found
fragments in fox scats or chewed remains of 11 egg-
shells of murres, kittiwakes, puffins, and Northern
Gannets (Morus bassanus). Some eggshells may have
been discarded by Common Ravens (Corvus corax)
and subsequently chewed up and ingested by foxes.

On I1 June 1978, in part of an active fox denand in
an adjacent burrow-like excavation (about 1.5 X 1 X
1 m), we found a cache containing fresh carcasses of
30 petrels, 3 kittiwakes, 3 puffins, a Common Murre,
and a Fox Sparrow (Passerella iliaca), all adults. In
August 1980, two other fox caches were located about
1.5 km away in an area where another den was sus-
pected (a kit sighted with an adult fox on 21 June).
One cache contained the fresh carcasses of at least 27
petrels (two chicks), 29 puffins (four chicks); the other
contained 15 petrels, 9 puffins, a fledgling kittiwake,
and a murre chick (K. Brink and D. Roby, University
of Pennsylvania, personal communication).

We do not know the extent to which foxes cache
prey elsewhere. Arctic Foxes (Alopex lagopus) are
known to horde substantial food ina single location,
though Red Foxes are not (D. Macdonald, Oxford
University, personal communication). Many uneaten,
discarded, petrel carcasses were found on the island,
especially in the wooded areas (cf. Kruuk 1972).

We first witnessed predation on 23 June 1979 at
11:25, when a Red Fox, walking along a cliff top,
caused a large number of nesting kittiwakes and
murres to scatter in panic from the ledges below. The
fox climbed down the nearly vertical cliff and carried a
murre egg up the cliff and out of view. Sixteen murre
eggs were taken in this way in 95 min. On 28 June
1979, we again sawa fox climb down this cliff, scaring
off the kittiwakes and murres. This time the fox duga

1981

NOTES

353

FIGURE |. An area where burrows of Leach’s Storm-Petrels were dug out by Red Foxes (photo: W. A. Montevecchi)

puffin out of a burrow in a grassy area among the
ledges and carried it off.

The Red Fox population on Baccalieu apparently
includes two or more adult breeding pairs, as kits have
been seen in most years, and two active dens were
found in 1978. In winter, Willow Ptarmigan (Lagopus
lagopus), a few small land birds, and berries may
provide foxes with limited food. The vertical cliffs
around the island’s perimeter and absence of beaches
preclude scavenging on marine invertebrates and car-
rion. Winter food shortages apparently keep the pop-
ulation in check, and foxes may have to cache sizable
quantities of seabirds to survive. No decline in seabird
numbers in the 20 years foxes have lived on the island
is evident, though such changes would not be easily
detectable in view of the absence of standardized esti-
mates of the large populations of burrow-nesters.
Marine birds with low reproductive rates may be very
seriously affected by additional sources of adult mor-
tality. Petrels and puffins, whose nest-site selection
patterns have evolved on islands free of land preda-
tors, are particularly vulnerable to foxes; adults of
cliff-nesting species are less likely to be preyed on even
in areas accessible to foxes. The extirpation by foxes
of any seabird species on Baccalieu Island seems
unlikely. However, in view of depredations foxes have
inflicted on seabird islands (Jones and Byrd 1979) and
the apparent lack of alternative prey on Baccalieu, we
conclude that systematic efforts should be made to
assess the foxes’ impact.

The Canadian Coast Guard provided housing on
Baccalieu, where E. Blundon, R. Hyde, F. Noonan, P.
Rice, and L. Walsh offered hospitality. B. D. Harvey,
I. R. Kirkham, and R. Purchase helped in searching
the fox cache. K. Brink, I. R. Kirkham, D. Roby, and
G. A. Sanger gave helpful criticism and information.
Research was supported by NSERC grant A0687 to
W. A. M.

Literature Cited

Brown, R. G. B., D. N. Nettleship, P. Germain, C. E. Tull,
and T. Davis. 1975. Atlas of eastern Canadian seabirds.
Canadian Wildlife Service, Ottawa. 220 pp.

Jones, R.D., Jr., and G. V. Byrd. 1979. Interrelations
between seabirds and introduced animals. /n Conserva-
tion of marine birds of northern North America. Edited by
J.C. Bartonek and D.N. Nettleship. United States
Department of the Interior, Fish and Wildlife Service,
Wildlife Research Report 11: 221-226.

Kruuk, H. 1972. Surplus killing in carnivores. Journal of
Zoology (London) 166: 233-244.

Montevecchi, W. A. 1978. Predator-prey interactions
between ravens and kittiwakes. Zeitschrift fiir Tierpsy-
chologie 49: 136-141.

Nettleship, D. N. 1972. Breeding success of the Common
Puffin (Fratercula arctica L.) on different habitats at
Great Island, Newfoundland. Ecological Monographs 42:
239-268.

Tuck, L. M. 1961. The Murres. Canadian Wildlife Service
Monograh No. I, Queen’s Printer, Ottawa. 260 pp.

Received 11 July 1980
Accepted 29 January 1981

354 THE CANADIAN FIELD-NATURALIST

Vol. 95

Twinning and Postpartum Activity in Barren-Ground Caribou

(Rangifer tarandus)

E. JANET McDONALD and ARTHUR M. MARTELL

Canadian Wildlife Service, 204 Range Road, Whitehorse, Yukon Territory YIA 3V1

McDonald, E. Janet,and Arthur M. Martell. 1981. Twinning and postpartum activity in Barren-ground Caribou (Rangifer

tarandus). Canadian Field-Naturalist 95(3); 354-355.

On 26 May 1980 twin Caribou calves ( Rangifer tarandus) were observed on the calving grounds of the Porcupine Caribou
herd in the northern Yukon Territory. Postpartum activity of one of the twins was delayed relative to that of the other twin

and to that of a single calf born at the same time.

Key Words: Caribou, Rangifer tarandus, twinning, calving.

Twin calves have not been reported in wild Caribou
(Rangifer tarandus) although twin fetuses have been
found (Shoesmith 1976) and twins have been born to
captive Caribou (McEwan 1971) and to semi-
domesticated Reindeer (also R. tarandus) (Nowosad
1973). On 26 May 1980 twin calves were observed in
the northern Yukon Territory (69°26’N, 140°46’W)
during investigations on the calving grounds of the
Porcupine Caribou herd. Weather was sunny and
warm (15-19°C) with a light NE wind (21 km/h).
From 09:24 to 14:32 solar time, about 500 Caribou in
seven loose bands of about 40-150 animals were under
observation as they moved northwest across a gently
rolling cotton grass (Eriophorum vaginatum) mea-
dow. They were mainly cows (97% of the cows were
antlered) although about 50 yearlings and 3-5 calves
were present. Between 13:54 and 14:09 a band of 123
animals broke up; 56 remained in the area, 50 moved
northwest and out of sight, and 17(11 antlered cows, |
unantlered cow, 5 yearlings) moved north and down-
slope to a flat alluvial herb heath which was about
25% snow covered and 15% melt puddles. They began
to lie down and, at 14:24, all 17 were lying when
continuous observations began on an antlered cow
(A) of that group that appeared to be in labor. Eight
minutes later, after repeatedly lying down and stand-
ing up, she gave birth while standing within the closely
spaced band of Caribou (5-10 m between individu-
als). Although attention was focused on that cow and
her calf, a second antlered cow (B), about 10 maway,
was in the same field of the scope and, 12 min later,
gave birth while standing. Cow B then lay down and
did not stand up again for 2 h. Cow B’s calf was
smaller than cow A’s calf. Cow B’s body patly ob-
scured her calf and, at times, the heads of two calves
appeared to be visible behind her body. The presence
of a second calf with cow B was not confirmed, how-
ever, until 2 h after the observed birth when she and

one calf stood up exposing to view a second, much
smaller and wetter appearing calf lying beside the first.

Continuous observations were made by a two-
person team, an observer and a recorder. The nature
of continuous observations is such that essentially all
of the observer’s attention was given to the focal
animal, in this case cow A. Therefore, the activities of
other animals in the field of the scope are recorded
only generally, as they relate to the focal animal, and
some short-term, unrelated activities may be missed.
The recorder, however, was able to scan the other
animals periodically. In this case, observations were
normally made 15X at a distance of about 350 m,
giving a field of view of about 18 m. Cow B was the
only other animal in the field of the scope up to the
time that the twins were confirmed, and no other
Caribou in the group were observed either to be in
labor or to give birth.

At 120 min after the observed birth, the larger twin
stood up, walked 5 m, and lay down. Cow B followed
and lay down beside the larger twin while the smaller
twin remained lying and made only a few small
movements. The larger twin stood up again about
180 min after the observed birth, walked back, and lay
down beside the smaller twin. Again cow B followed
and then lay down beside both calves. Following that
move the smaller twin began to move its head and
body and got up on two legs. Cow B was attentive to
both twins and frequently licked their bodies and
anuses. During the first 326 min following calving,
cow B stood up only four times, twice to follow her
calf and twice to stand for less than | min each. Cow
A, on the other hand, lay down, stood up, walked, and
fed repeatedly during the first 338 min following calv-
ing. Neither of the twin calves attempted to nurse,
whereas cow A’s calf nursed 31 times for a total of
12 min 26 s (x = 24+ 4.2 s) during the 171 min from
the beginning of the first successful nursing. Com-

1981

‘pared to the single calf, it appears unusual that neither
of the twins nursed during the first 326 min following
birth, although the first successful nursing in Reindeer
may not take place until 5-6 h after the birth (Baskin
1970).

Cow A passed the placenta while standing 32 min
after giving birth and began eating the placenta
134 min later. Cow B passed the placenta while lying
35 min after giving birth and began eating the pla-
centa 158 min later. Times from birth for the first
observations of several postpartum activities for cow
A’s calf are as follows: got up on two legs and dropped
down again, 9 min; got up on four legs and dropped
down again repeatedly, 22 min; stood on four legs for
a short while (8s), 40 min; attempted to nurse,
50 min; stood for 2 min, 84 min; walked and ran
around cow, 98 min; walked for 2 min, 117 min;
nursed successfully, 167 min. Those times are similar
to times reported previously (de Vos 1960; Kelsall
1957, 1960; Lent 1966; J. Curatolo and D. Roseneau,
unpublished observations). They are consistent with
those observations that the larger twin stood and
walked at 120 min after birth but it is unusual that the
smaller twin did not even successfully stand up on four
legs during the 206 min after its presence was
confirmed.

Observations were terminated at 20:11 because of
fatigue, cold, and hunger. When the observers
returned 24 h later, six cows, five calves, and an aban-
doned calf were present on the site. The abandoned
calf appeared to be too large to be either of the twin
calves. The observers returned again 12 h after the
previous check and observed five cows, five calves,
and an abandoned calf on the site. Those animals were
presumed to be the same group seen at the previous
check. The cows and calves moved off to the north-
west at that time when another band of Caribou
moved through. The site was then thoroughly
searched for dead calves but none were found. Possi-

NOTES

355

bly both twins left with cow B during the first day after
birth or the weaker twin may have been removed by a
scavenger. A Wolverine (Gu/o gulo) was seen near the
site on 28 May. Grizzly Bears ( Ursus arctos) are com-
mon in the area but none were seen near the site.

Acknowledgments

We are grateful to Polar Continental Shelf Project,
Western Arctic Scientific Resources Centre, and
Yukon Wildlife Branch for their support. We thank
K. Asquith, C. McEwan, and W. Nixon for their
assistance with the field work, and D. R: Flook and
D. R. Klein for their comments on the manuscript.

Literature Cited

Baskin, L.M. 1970. Reindeer ecology and _ behavior.
“Nauka” Publishing House, Moscow. 149 pp. (Translated
from Russian, Canadian Wildlife Service, Ottawa).

de Vos, A. 1960. Behavior of Barren-ground Caribou on
their calving grounds. Journal of Wildlife Management
24: 250-258.

Kelsall, J. P. 1957. Continued Barren-ground Caribou
studies. Canada Department of Northern Affairs and
Natural Resources, Canadian Wildlife Service, Wildlife
Management Bulletin, Series 1, Number 12. 148 pp.

Kelsall, J. P. 1960. Co-operative studies of Barren-ground
Caribou. Canada Department of Northern Affairs and
Natural Resources, Canadian Wildlife Service, Wildlife
Management Bulletin, Series 1, Number 15, 145 pp.

Lent, P. C. 1966. Calving and related social behavior in the
Barren-ground Caribou. Zeitschrift fur Tierpsychologie 6:
701-756.

McEwan, E. H. 1971. Twinning in Caribou. Journal of
Mammalogy 52: 479.

Nowosad, R. F. 1973. Twinning in Reindeer. Journal of
Mammalogy 54: 781.

Shoesmith, M. W. 1976. Twin fetuses in Woodland Cari-
bou. Canadian Field-Naturalist 90: 498-499.

Received 13 November 1980
Accepted 20 February 1981

356 THE CANADIAN FIELD-NATURALIST

Vol. 95

Seasonal Pelage Change of Marten (Martes a. americana) in Maine

EDWARD C. SOUTIERE! and J. DOUGLAS STEVENTON

School of Forest Resources, University of Maine, Orono, Maine 04469
'Present address: Remington Farms, Chestertown, Maryland 21620

Soutiere, Edward C., and J.Douglas Steventon. 1981. Seasonal pelage change of Marten (Martes a. americana) in Maine.

Canadian Field-Naturalist 95(3): 356

The twice yearly molt and seasonal coat color of Marten (Martes a. americana) in Maine is documented. The summer pelage is
dark whereas the winter pelage is usually paler with reddish-yellow hues predominating.

Key Words: Marten, Martes a. americana; pelage color, molt.

The pelage color of Marten (Martes a. americana) .

has been described as being highly variable. Seton
(1929, p. 482) stated that the general color ranges from
golden yellow to blackish umber. These color variants
are described as not being related to the seasonal molt;
the summer coat not differing greatly in color from
that of the winter coat (Seton 1929; Jackson 1961, p.
328). While live-trapping Marten in the Moosehead
Lake and northern Aroostook County regions of
northern Maine from 1974 to 1979, we noticed that,
much like the Ermine (Mustela erminea), the Marten’s
pelage color is distinctly different in summer and
winter.

We use the Munsell system (United States Depart-
ment of Agriculture 1951) to specify and identify pel-
age colors. For example, R4/ 14 means that the hue is
red, the value is 4, and its chroma is 14.

We handled 211 individual Marten of which 63
were observed through one or more molts. The pat-
tern and timing of fur replacement followed that des-
cribed for the European Pine Marten(Martes martes)
by Schmidt (1943). The molt into the summer pelage
begins in late April and is complete by mid-June. The
body color of Marten captured during summer
(N=71) is very dark grayish brown (10YR3/2) or
black (1OYR2/ 1) while the tail is lighter (1IOYR4/3),
tipped with black. The throat patch is reddish yellow
GES ARTS):

The molt to winter coat is complete by mid-October
in northern Maine. In contrast to the dark summer
pelage, the winter coat includes a grayish head and
neck and a body color suggestive of a Red Fox
(Vulpes vulpes). The body color is variable both on
and between individuals. The predominant colors of
winter Marten (N = 176) are yellowish red (SYRS5/8),
strong brown(7.5YR5/8), reddish yellow (7.5YR6/8),
yellowish brown (10YR5/6), and brownish yellow
(10YR6/6). The head, neck, and occasionally the fore
shoulders are light gray (10YR7/1) or very pale brown
(10YR8/4). The color of the legs and tail gradates
from the body color to a very dark brown (10YR2/2)
on the feet and tip, respectively. The throat patch is
reddish yellow (7.5YR7/8) or yellow (10YR8/6).

Two specimens exemplifying the extremes of winter

University of Maine. One skin (Catalogue No. 73-2) is
color are filed in the Wildlife Resources Museum,
dark brown (10YR3/3) with a very dark brown
(10YR2/ 2) stripe along the back from the neck to root
of tail. This specimen has a white throat patch
(10YR8/2). The second skin (Catalogue No. 276) is
nearly white (1OYR8/1) except for the legs and tail
which are dusky red (10YR3/2). In Ontario, de Vos
(1952, pp. 21-25) described Marten in winter pelage as
ranging from very pale yellow, a partly albinistic type,
to very dark blackish-brown. He made no reference to
color variation associated with the seasonal molt.

Of 1055 pelts we examined at fur buyers in Maine,
only 2.4% were the dark brown color variant, and
0.7% the nearly white variant. It is clear that in Maine
the reddish-yellow winter pelage described is most
common and the dark brown or white Marten, such as
the museum specimens described or those described
by de Vos, are rare.

We thank Art Soukkala, Kate Wynne, and John
Major for providing some of the live-trapping data
and assisting in examination of pelts. We also thank
the fur buyers who allowed access to pelts. M. W.
Coulter reviewed the manuscript. Most of this study
was conducted during research supported by the
School of Forest Resources, University of Maine; the
Penobscot County Conservation Association; and
MclIntire-Stennis Research Project ME-09606.

Literature Cited

de Vos, A. 1952. The ecology and management of Fisher
and Marten in Ontario. Ontario Department of Lands and
Forests Technical Bulletin. 90 pp.

Jackson, H. H. T. 1961. Mammals of Wisconsin. The Uni-
versity of Wisconsin Press, Madison. 436 pp.

Schmidt, F. 1943. Naturgeschichte des Baum und des
Steinmarders. Monographie des Wildsaugetierre 10:
100-102.

Seton, E. T. 1929. Lives of game animals. Vol 2. Country
Life Press, Garden City, New Jersey. 746 pp.

United States Department of Agriculture. 1951. Soilsurvey
manual. Handbook 18. 483 pp.

Received 5 January 1981
Accepted 4 March 1981.

1981

NOTES 357

Marsh Nesting of Common Loons (Gavia immer)

ROBERT ALVO

Department of Biology, Queen’s University, Kingston, Ontario K7L 3N6

Alvo, Robert. 1981. Marsh nesting of Common Loons (Gavia immer). Canadian Field-Naturalist 95(3): 357

The results of a survey conducted on four lakes on the Rideau Canal System, southern Ontario, showed that 58% of the nests
of Common Loons (Gavia immer) were in marshes. This nest site choice may be attributable to the high levels of human
disturbance on these lakes. This finding has implications both for loon and marsh conservation.

Key Words: Common Loon, Gavia immer; nest site, island, marsh, human activity.

The Common Loon (Gavia immer) is a summer
resident of lakes in northern United States and most
of Canada. Olson and Marshall (1952),-Vermeer
(1973), and McIntyre (1975) reported that 93, 96, and
88% of the loon nests that they found were on islands.
According to Vermeer (1973), island nesting probably
reduces predation; mainland sites are avoided because
of their vulnerability to mammalian predators. How-
ever, when islands are not present, loons usually nest
in marshes (Munro 1945).

The loon’s habit of nesting close to the water’s edge
makes the nest especially susceptible to disturbance by
humans. The presence of cottagers and boats often
keeps loons off their nests during the incubation
period, and may ultimately affect nest site selection.
All of the published studies on Common Loon nesting
preferences have been conducted in relatively undis-
turbed areas. The present study was undertaken in a
disturbed area to determine whether nest site selection
may be affected by human activity.

Study Area

Four lakes (Clear, Indian, Opinicon, and Sand) in
the Canadian Shield (44°33’N, 76°20’W), 40-60 km
north of Kingston, Ontario, varying in area from 1.72
to 7.86 km2, served as the study area. The lakes are
mesotrophic to eutrophic. Cottage density varies from
16.5 to 43.2 cottages/km2 of lake area. All four lakes
are on the Rideau Canal System and therefore have a
large amount of boat traffic during the summer. There
is also much fishing, sailing, and waterskiing on these
lakes.

Methods

A survey of nesting loons was carried out by canoe
2 May - 31 August, 1980. Nests found were checked
every | or 2 wk to determine the outcome of each.
Hatching success was based on the observation of at
least one chick.

Results and Discussion

Of 17 nests found, 7 were situated on islands and 10
were in marshes. Islands with flat sections of shore-
line, which would have provided suitable nesting sites,
were present near all marsh nests. However, in each of

these cases the potential nesting islands were closer to
cottages and to the main routes of boat travel than was
the marsh site;-marsh nests were generally at least
100 m from cottages and from boat traffic, while the
potential nesting islands usually were located beside
routes of boat traffic and within 50 m of the nearest
cottage. The areas with marsh nests were generally
very shallow, the nest usually being situated in water
about 0.5m deep; boats were thus very seldom
observed in these areas.

Four of 10 marsh nests were successful, compared
to 2 out of 7 island nests. The sample sizes are too
small to compare the success rates in different situa-
tions statistically.

Olson and Marshall (1952) suggested that human
activity may influence the selection of nest sites by
loons. I suggest that loons in developed areas are
giving up their favored island nesting sites to escape
such disturbance, and asa result marsh nesting may be
increasing in frequency. The small number of shel-
tered marshy bays may impose a limit to the number
of breeding pairs of loons ona lake. All of this adds to
the importance of marsh conservation.

Acknowledgments

This work was funded by an Experience’80 grant to
Raleigh Robertson. I thank Barbara Black for her
kind help. Frank Phelan, Raleigh Robertson, and
Steve Dukoff offered invaluable editorial assistance.

Literature Cited

McIntyre, J. M. 1975. Biology and behavior of the Com-
mon Loon(Gavia immer) with reference to its adaptability
ina man-altered environment. Ph.D. dissertation, Univer-
sity of Minnesota, Minnesota.

Munro, J. A. 1945. Observations of the loon in the Cariboo
Parklands, British Columbia. Auk 62: 38-49.

Olson, S. T., and W.M. Marshall. 1952. The Common
Loon in Minnesota. Occasional Papers Minnesota
Museum of Natural History 5: 1-77.

Vermeer, K. 1973. Some aspects of the nesting require-
ments of Common Loon in Alberta. Wilson Bulletin 85:
429-435.

Received 29 December 1980
Accepted | March 1981

358 THE CANADIAN FIELD-NATURALIST

Vol. 95

Evidence of Arctic Grayling (Thymallus arcticus)

Spawning in a Highway Culvert

L. F. KRATT

Environmental Management Associates, 1530-10 Avenue S:W., Calgary, Alberta T3C 0J5

Kratt, L. F. 1981. Evidence of Arctic Grayling (Thymallus arcticus) spawning in a highway culvert. Canadian Field-

Naturalist 95(3): 358.

Arctic Grayling (Thymallus arcticus) eggs were collected in gravel deposits within a circular highway culvert. Because gravel
at stream crossings may providea suitable spawning substrate where none existed before, this alteration is a beneficial effect of

road construction.

Key Words: Arctic Grayling, Thymallus arcticus,; spawning, culverts, road construction, Yukon Territory.

During a fisheries study in the spring of 1980 in
southern Yukon Territory, several potential Arctic
Grayling (Thymallus arcticus) spawning streams were
examined. In addition to direct observations of
reproductive behavior, evidence of spawning was
gathered by checking areas of suitable substrate for
the presence of eggs. Samples were collected by hold-
ing the edge of a fine-mesh dip net to the stream
bottom at a right angle to the current and stirring the
area immediately upstream with a stick. In this
manner eggs were dislodged and were swept into the
net. I employed this technique successfully during

previous Arctic Grayling early life history studies

(Kratt and Smith 1977).

The construction and maintenance of the Alaska
Highway have resulted in gravel deposits at many
stream crossing locations. These deposits provide
suitable spawning areas in creeks which would nor-
mally not be utilized by Arctic Grayling for reproduc-
tive purposes. One such stream drains a small muskeg
area before flowing into the Koidern River (61°58’N,
140°25’W). Surveys of this stream in 1980 were
designed to determine if the creek had again been
utilized by Arctic Grayling; hence, efforts were con-
centrated on locating eggs within the substrate.

On 28 May, Arctic Grayling eggs were found in
pockets of gravel located upstream and downstream
of the road crossing. Eggs were also collected in a
small sand and gravel deposit within the culvert.
Because the stream culvert is of the circular variety,
substrate materials undoubtedly originated in up-
stream areas and were deposited by the current.

I assumed that eggs were laid in the culvert because
no eggs were observed on the surface of the gravel
substrate in any area or were evident in the stream
drift. Despite the absence of nest-building behavior,
Arctic Grayling eggs are deposited at a substrate
depth of 2 to 3cm (Kratt and Smith 1977). Eggs
dislodged from upstream spawning sites would likely

pass through the culvert as they remain adhesive for
only a short period of time after spawning (Bishop
1971; personal observations).

Guidelines for the protection of aquatic habitats
frequently suggest the use of open-bottom culverts for
road developments as stream substrates remain rela-
tively undisturbed and disruption of fish passage is
minimized (e.g., Dane 1978; Dryden and Stein 1975;
Metsker 1970). Although I support these recommen-
dations, the data presented here indicate that mate-
rials deposited within closed culverts may also be of
importance where available spawning habitat is
limited. Thus in some circumstances road construc-
tion materials may enhance aquatic habitats.

I thank Foothills Pipe Lines (Yukon) Ltd. for per-
mission to publish this manuscript.

Literature Cited

Bishop, F. G. 1971. Observations on the spawning habits
and fecundity of the Arctic Grayling. Progressive Fish-
Culturist 33(1): 12-19.

Dane, B. G. 1978. A review and resolution of fish passage
problems at culvert sites in British Columbia. Fisheries
and Environment Canada, Fisheries and Marine Service,
Technical Report Number 810. 126 pp.

Dryden, R.L., and J. N. Stein. 1975. Guidelines for the
protection of the fish resources of the Northwest Territo-
ries during highway construction and operation. Envir-
onment Canada, Fisheries and Marine Service, Technical
Report Number CEN/T-75-1. 32 pp.

Kratt, L. F., and R. J. F. Smith. 1977. A post-hatching
sub-gravel stage in the life history of the Arctic Grayling,
(Thymallus arcticus). Transactions of the American
Fisheries Society 106(3): 241-243.

Metsker, H. E. 1970. Fish versus culverts, some considera-
tions for resource managers. United States Department of
Agriculture, Technical Report Number ETR-7700-5. 19

Pp.

Received 14 January 1981
Accepted 5 March 1981

1981 NOTES 359

Freshwater Fishes from Northern Newfoundland

PETER J. ROMBOUGH,'” STEPHEN E. BARBOUR,! and JOSEPH J. KEREKES3

1Biology Department, Dalhousie University, Halifax, Nova Scotia B3H 4J1

2Present address: Pacific Biological Station, Nanaimo, British Columbia V9R 5K6

3Environment Canada, Canadian Wildlife Service, c/o Biology Department, Dalhousie University, Halifax, Nova Scotia
B3H 4J1

Rombough, Peter J., Stephen E. Barbour, and Joseph J. Kerekes. 1981. Freshwater fishes from northern Newfoundland.
Canadian Field-Naturalist 95(3): 359-361.

Atlantic Salmon, Salmo salar; Brook Trout, Salvelinus fontinalis; Arctic Char, Salvelinus alpinus; Rainbow Smelt, Osmerus-
mordax; American Eel, Anguilla rostrata, Threespine Stickleback, Gasterosteus aculeatus; Fourspine Sticleback, Apeltes
quadracus; and Ninespine Stickleback, Pungitius pungitius, were the only freshwater fishes collected from three ponds in
L’Anse aux Meadows National Historic Park at the northern tip of insular Newfoundland. The record of the Fourspine
Stickleback extends its known range northward by 45 km.

Key Words: Salmo salar, Salvelinus fontinalis, Salvelinus alpinus, Osmerus mordax, Anguilla rostrata, Gasterosteus
aculeatus, Apeltes quadracus, Pungitius pungitius, L_ Anse aux Meadows National Historic Park, Newfoundland, records,
range extension.

The Great Northern Peninsula of Newfoundland 1s
the closest part of the island to mainland North Amer-
ica. Garside (1970) and Dadswell (1972) raised the
possibility that it may have served as an avenue for the
invasion of freshwater fish from the mainland follow-
ing the Wisconsin glaciation. The freshwater fish
fauna of the peninsula is therefore potentially impor-
tant in the zoogeography of the island but has been
only cursorily examined. Van Vliet (1970) surveyed
several streams throughout the peninsula, but always

within estuarine influences. Dadswell (1972) sampled
freshwater ponds in Gros Morne National Park near
the base of the peninsula. The present study, to our
knowledge, is the first intensive sampling of fresh-
waters near its northern tip.

L’Anse aux Meadows National Historic Park is
located on the Strait of Belle Isle near the northern
extremity of the island of Newfoundland (51°35’N,
55°30’W). Three ponds in the park, each ina separate
watershed, were surveyed from 6 to 26 August 1977

Pa

] (0) Tkm

FIGURE |. Location of the three ponds surveyed in L’Anse aux Meadows National Historic Park on the Great Northern
Peninsula of Newfoundland. 1, Duck Pond; 2, Skin Pond; 3, Black Duck Pond; H, Historic Site.

360

TABLE |—Physical and chemical characteristics of the ponds
sampled in L’Anse aux Meadows National Historic Park,
Newfoundland

Black

Duck Skin Duck

Pond Pond Pond
Surface area (ha) 29.4 7.4 S92
Maximum Depth (m) 16 161 2.4
Mean Depth (m) 4.5 — 1.2
pH 6.5 6.7 6.7
Conductance (uS/cm) 54 42 Sy
Temperature (°C) 11-17 12 11
Color (Hazen Units) 65 60 120
Predominant bottom type Rubble, Mud, Sand

gravel sand mud

and 8 to 17 August 1978. The ponds were all situated
on the coastal barrens within | km of the sea (Figure
1). They were relatively shallow, unstratified in
August, and highly colored. The physical and chemi-
cal characteristics of the ponds are summarized in
Table |.

Samples were collected using variable-mesh gill
nets (2.5-7.5 cm stretched mesh) and fine-mesh trap
nets (0.5-1.0 cm stretched mesh). Representatives of
each species were preserved and stored in the Depart-
ment of Biology, Dalhousie University.

The species diversity of the freshwater fish fauna of
L’Anse aux Meadows National Historic Park (Table
2) is impoverished compared with more southern
areas of Newfoundland. Only 8 of the 23 species
recorded in Newfoundland freshwaters (Scott and
Crossman 1964) were collected in the Park. Although
two adult Arctic Char, Salvelinus alpinus, were cap-
tured in Black Duck Pond in 1977, a resident popula-
tion is unlikely because no adults were captured in
1978 or juveniles in either year. The fish captured in
1977 were probably strays from rivers that enter
nearby Pistolet Bay and are known to contain Arctic
char.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Rainbow Smelt, Osmerus mordax, and American
Eel, Anguilla rostrata, were common in Black Duck
Pond and Duck Pond. However, Jeffers (1932)
reported that according to the local inhabitants both
species were rare in the Pistolet Bay region in the late
1920s. These apparently low population levels coin-
cided with low surface water temperatures in the
northwest Atlantic. Because this region is near the
northern limit of the range of both species, low ocean
temperatures may have restricted their numbers. The
subsequent warming trend (Lauzier and Hull 1969)
may account for their present relative abundance.

The capture of a Fourspine Stickleback, Apeltes
quadracus, in Duck Pond extends the known range of
the species to the northern tip of insular Newfound-
land. The previous most northerly record was from
brackish water near the mouth of Western Arm
Brook, about 120 km southwest (45 km south) of the
present location (Van Vliet 1970). The Fourspine
Stickleback is typically a marine species but is occa-
sionally found in freshwater (Scott and Crossman
1973; Dadswell 1972). The specimen collected in Duck
Pond probably had strayed from the salt marsh at the
outlet of Duck Pond Brook.

The present freshwater ichthyofauna of Newfound-
land appears to be derived solely from invasions of
euryhaline species from the sea. Garside (1970) raised
the possibility that unreported populations of obligate
freshwater fishes may have become established on the
Great Northern Peninsula through migration across
the Strait of Belle Isle shortly following the last glacial
recession. No obligate freshwater fishes were recorded
during our survey of an area along the proposed inva-
sion route, and the possibility that they have become
established naturally in insular Newfoundland
appears remote.

Acknowledgments

Our thanks are extended to P. G. Daye, A. J. Ers-
kine, E. T. Garside, and S. W. Speller for their helpful
comments on the manuscript. Financial support was

TABLE 2—Species and numbers of freshwater fishes captured in ponds in L’Anse aux Meadows National Historic Park in
1977/1978. Plus signs (+) indicate that the species was observed but not captured

Salmo salar, Atlantic salmon

Salvelinus fontinalis, Brook Trout

Salvelinus aplinus, Arctic Char

Osmerus mordax, Rainbow Smelt

Anguilla rostrata, American Eel”

Gasterosteus aculeatus, Threespine Stickleback
Apeltes quadracus, Fourspine Stickleback

Black Duck Pond Skin Pond Duck Pond
65/43 —/- -/- ,
18/50 36/67 28/13
2/- -|- -|-
12/16 —/- 28 / >2000°
8/12 ~/- +/+
63/>1072 -/- +/314

-/- -/- =
—/2 +/199 —/20

Pungitius pungitius, Ninespine Stickleback

“Larvae except for two adults.

>The presence of a fairly abundant eel population was indicated by damage to gill-netted fish and by the presence of eel marks

in gill nets.

1981

provided by Parks Canada and the Canadian Wildlife
Service.

Literature Cited

Dadswell, M. J. 1972. New records of freshwater fishes
from the northwest coast of insular Newfoundland. Cana-
dian Field-Naturalist 86: 289-290.

Garside, E. T. 1970. Newsamples of the piscifauna of insu-
lar Newfoundland. Canadian Field-Naturalist 84:
385-386.

Jeffers, G. W. 1932. Fishes observed in the strait of Belle
Isle. Contribution to Canadian Biology and Fisheries 7:
203-211.

Lauzier, L. M., and J. H. Hull. 1969. Coastal station data
temperatures along the Canadian Atlantic coast 1921-

NOTES 361

1969. Fisheries Research Board of Canada Technical
Report 150. 27 pp.

Scott, W. B., and C. J. Crossman. 1964. Fishes occurring
in the fresh waters of insular Newfoundland. Department
of Fisheries, Canada, Queen’s Printer. 124 pp.

Scott, W. B., and C. J. Crossman. 1973. The freshwater
fishes of Canada. Fisheries Research Board of Canada
Bulletin 184. 966 pp.

Van Vliet, W. H. 1970. Shore and freshwater fish collec-
tions from Newfoundland. National Museum of Canada,
Publication in Zoology 3: 1-30.

Received 16 January 1978
Revision received 17 December 1980
Accepted 10 March 1981

Sightings of the Hop Merchant (Polygonia comma) Butterfly

on Insular Newfoundland

BERNARD S. JACKSON

Oxen Pond Botanic Park, Memorial University, St. John’s, Newfoundland AIC 5S7

Bernard S. Jackson. 1981. Sightings of the Hop Merchant ( Po/ygonia comma) butterfly on insular Newfoundland. Cana-

dian Field-Naturalist 95(3): 361-362.

The status of the Hop Merchant (Po/ygonia comma) on Insular Newfoundland is undetermined. The controversy as to
whether or not this species should be included on the island’s list of Macrolepidoptera has been recently revived by the

photograph and records of new sightings presented here.

Key Words: Hop Merchant, Polygonia comma; insular Newfoundland, Lepidoptera, Nymphalidae, range extension.

The Hop Merchant, Polygonia comma (Lepidop-
tera: Nymphalidae), ranges from the maritime pro-
vinces, Anticosti Island, Quebec, and Ontario, south-
ward to North Carolina (in mountains) and westward
to Kansas, Nebraska, and Iowa (Klots 1951). Its status
in Newfoundland is undetermined. Although Dos
Passos (1935) states that he recognized P. comma as
occurring in southern Newfoundland, he almost
immediately corrected this error (Dos Passos 1935).

Krogerus (1954), in his study of Newfoundland
Lepidoptera, made no mention of P. comma. W. W.
Gregory (1975, checklist of the butterflies and
skippers of Canada, Lyman Entomological Museum
and Research Laboratory, Ste. Anne de Bellevue,
Quebec. 44 pp.) included Newfoundland in the range
of this species, but recently advises (personal com-
munication, 1980) that he now considers this earlier
record doubtful and has dropped this taxon from the
revised checklist, shortly to be published. Morris
(1980) did not include P. comma as occurring in the
province.

Because previous records of this species in New-

foundland are unsubstantiated and considered con-
troversial, | have reported new sightings here.

On 23 May 1979 an unidentified butterfly was
observed feeding from the flowers of a native Prim-
rose (Primula laurentiana) and Leatherleaf (Chamae-
daphne calyculata) in the Oxen Pond Botanic Park,
St. John’s, Newfoundland (47°34’N, 52°43’W), and
another smaller specimen was briefly observed flying
around the same area, simultaneously. Some hours
later, one was observed in the same general area,
feeding from the flower of a Dandelion (Taraxacum
officinale) and assumed to be one of the original two
sighted.

Because Oxen Pond Botanic Park is a nature
reserve, especially for butterflies, neither specimen
was collected. However, a close-up Kodachrome
transparency (see Figure 1) showing the ventral sur-
face of the wings was sent independently to J. Donald
Lafontaine and Eugene G. Monroe, both of the Bio-
systematics Research Institute, Ottawa. Both identi-
fied it as a photograph of P. comma.

Howe (1975) has stated that the host plants of this

362

FiGuRE |. Print from the 35-mm color transparency taken on
23 May 1979 of the butterfly subsequently identified
as a Hop Merchant (Polygonia comma).

species are hops (Humulus), nettle ( Urtica), false net-

tle (Boehemia), and elm (U/mus); the Stinging Nettle
(Urtica dioica), Hop (Humulus lupulus), and English
Elm (U/mus procera) are to be found within the gen-
eral area of the sightings. False nettle (Boehemia) does
not occur in Newfoundland.

Of the five species or subspecies of the genus Poly-
gonia currently found in Newfoundland, only the
Green Comma (P. faunus) is reasonably common. All

THE CANADIAN FIELD-NATURALIST

Vol. 95

others are uncommon and not readily identifiable in
the field.

Both hops and Stinging Nettle are to be found in
many Newfoundland outports, especially on the
Avalon Peninsula, having originated with the early
settlers. English Elm trees also are scattered and well
established throughout the city of St. John’s. There-
fore, in areas where one or more of its host plants are
present, the Hop Merchant could occur at a low popu-
lation level similar to that of other recorded Polygo-
nia. Its apparent anonymity could largely be due to
the scarcity and difficulty in identifying members of
this genus and to the lack of experienced field
observers in the area.

Literature Cited

Dos Passos, C. F. 1936. Further notes on butterflies of
southern Newfoundland. Canadian Entomologist: 68-98.

Dos Passos, D. F. 1943. A correction. Canadian Entomol-
ogist 75: 178.

Howe, W.H. 1975. The butterfiles of North America.
Doubleday and Company, Inc., New York.

Klots, A. B. 1951. A field guide to the butterflies of North
America, east of the Great Plains. Peterson Field Guide
Series, Houghton Mifflin Company, Boston, Riverside
Press, Cambridge, Massachusetts. 349 pp.

Krogerus, H. 1954. Investigations on the Lepidoptera of
Newfoundland, I. Macrolepidoptera. Acta Zoologica
Fennica 82, edidit, Societas Pro Fauna et Flora Fennica.
Helsingflorsiae. 80 pp.

Morris, R. F. 1980. Butterflies and moths of Newfound-
land and Labrador, the Macrolepidoptera. Publication
1691. Canadian Government Publishing Centre, Hull,
Quebec, Canada K1A 0S9. 407 pp.

Received 28 November 1980
Accepted 23 March 1981

1981

NOTES

363

Application of the Varrié Snow Index to Overwintering
North American Barren-Ground Caribou (Rangifer tarandus arcticus)

WILLIAM O. PRUITT, JR.

Department of Zoology, University of Manitoba, Winnipeg, Manitoba R3T 2N2

Pruitt, William O., Fr. 1981.

Application of the Varrio Snow Index to overwintering North American Barren-ground

Caribou (Rangifer tarandus arcticus). Canadian Field-Naturalist 95(3): 363-365.

The Varrio Snow Index (VSI) that I derived to model the wintering activity and movements of feral and domesticated
Reindeer in northeastern Finland was used successfully to model the wintering activity and movements of wild North
American Barren-ground Caribou ( Rangifer tarandus arcticus). This index is a potentially useful tool for determining nivally

suitable areas for Caribou overwintering.

Key Words: Caribou, Reindeer, snow ecology, Mammalia: Cervidae, winter activity.

During the winter 1957-1958, as part of the Coop-
erative Caribou Investigation, I studied the api (snow
cover of the forest floor; see Pruitt 1979 for terminol-
ogy related to snow characteristics) of northern Sas-
katchewan in relation to overwintering Barren-
ground Caribou (Rangifer tarandus arcticus).
Throughout the snowseason I established, and some-
times repeated, standard api profiles (Klein et al.
1950) at both forest and lake sites. Most of these
profiles were established in February 1958. On the
basis of obvious relations in the data, I concluded
(Pruitt 1959) that Caribou could detect differences in
thickness, hardness, and density of the api, and that
there were thresholds to these morphological charac-
teristics that the Caribou did not transgress.
Moreover, most of the animals’ winter movements
could be explained by analysis of api morphology.

Later, after I derived the concept of a Snow Index in
relation to the small mammals of the low arctic tundra
of the Ogotoruk region in northwestern Alaska
(Pruitt 1966), I made several unsuccessful attempts to
derive a similar index for my Saskatchewan Caribou
observations. It was evident that my 114 api profiles
were insufficient to permit such derivation.

The problem remained unsolved until winter
1976-1977 when I was able to establish 460 api pro-
files in relation to feral Reindeer ( Rangifer tarandus)
at Varrio Subarctic Research Station in northeastern
Finland. The number of observations, as well as
access to more recent studies such as those by Hen-
shaw (1968), Stardom (1975), and Skogland (1978)
enabled me to derive the Varrio Snow Index (VSI)
(Pruitt 1979). This resulted from my observation that
overwintering Rangifer react not only to the gross
factors of total thickness, hardness, and density of the
api, but to other, more subtle, characteristics as well.
A hard layer in the upper half of the api seems to be
more important in deterring feeding than a layer of

equal hardness in the lower half. Surface hardness is
probably more important than any other single char-
acteristic because Caribou test this directly (Pruitt
1979). I combined the characteristics of the api ina
variety of trial indices; the combination that best
modeled the wintering activity and movements of the
feral Reindeer in Salla North Reindeer Association |
called the Varrio Snow Index (Pruitt 1979).

Yi (ee (ah J) UE) = Cen OOO

where H~” = hardness of hardest layer more than
halfway between the substrate and the
top of the api.

H,T, = hardness times thickness of basal layer.

VT. = vertical hardness of surface times
- thickness of surface layer.

H,T, = hardness times thickness of hardest
layer (if not H,7,). If basal layer is the
hardest then term H,7, drops out.

T , = total thickness of api.

The Saskatchewan data had been stratified into “no
Caribou,” “occasional Caribou,” and “Caribou con-
centration” on the basis of extensive survey flights (see
Pruitt 1959 for definitions of these strata). The data
were Stratified further into forest sites and, usually, a
nearby lake site. The forest profiles in the areas of
“Caribou concentration” had hardnesses between
6.5 g-cm” and 60 g-cm”, densities of 0.13 and 0.20,
and thicknesses less than 59 cm. The profiles in the
areas of “occasional Caribou” could have hardnesses,
densities, and thicknesses in the same range as in the
“Caribou concentration” areas, but they also could
exceed these limits. The profiles in the “no Caribou”
areas could have hardnesses, densities, and thick-
nesses in the same range as in the “Caribou concentra-
tion” and “occasional Caribou” areas; they also could

364

exceed these limits and, in addition, they could exhibit
ice layers or “sigulik” in the profiles.

The Finland data were stratified into “Reindeer —

feeding” and “no Reindeer feeding.” These data are
also a time-related series so only portions can be com-
pared directly to the Saskatchewan data.

The purpose of this paper was to document the
successful application of the VSI to North American
Barren-ground Caribou.

Results

Using the Saskatchewan data I calculated VSI
values (Figure 1). All possible pairs proved to be
significantly different (one-tailed /-test, P< 0.05).
The VSIs of api used by Saskatchewan Rangifer in
February were numerically equivalent to VSIs of api
used by Finnish feral Reindeer in mid-November. I
had previously shown that the “Reindeer feeding”
VSIs of Varri6 region api increase until spring (Pruitt
1979). What Saskatchewan Rangifer would do in
VarriO-type api is an unanswerable question, but it is
clear that Finnish feral Reindeer have a significantly

CARIBOU
CONCENTRATION

OCCASIONAL
CARIBOU

NO
CARIBOU
FIGURE |. Api profiles (log VSI) in northern Saskatchewan,

winter 1957-1958, in relation to Caribou distribution:
Mean (horizontal lines) + SD (solid rectangle),
range (vertical lines); n, number of Caribou. Note that
“Caribou concentration” data indicate that most
animals were markedly restricted in their distribution
with regard to nival factors.

THE CANADIAN FIELD-NATURALIST

Vol. 95

greater threshold of sensitivity to the factors encom-

_passed by VSI than do Saskatchewan Rangifer.

Discussion

VSI models the api relations of Canadian wild
Barren-ground Caribou equally as well as it does
those of feral Finnish Reindeer. The differences
between Canadian and Finnish api relations are prob-
ably the result of artificial selection or culling in Fin-
nish Reindeer. One fundamental precept of domesti-
cation is that any Reindeer that tend to “wander” (..e.
instinctively to obey signals of api hardness, density,
and thickness) are slaughtered. Even after some thou-
sands of years of artificial selection, contemporary
Reindeer still retain measurable reactions to the same
api characteristics as their wild relatives.

Up to now it was not known whether discrepancies
between thresholds reported for North American wild
Caribou and Eurasian Reindeer (Nasimovich 1955;
Skogland 1978) were artifacts of different instrumen-
tation and methods or indications of an actual biolog-
ical difference. According to my comparison of VSIs
the difference between Saskatchewan and Finland
Rangifer-api relations appears to be real. Indeed, this
difference can be viewed as a measure of amount of
domestication or departure from the wild type.
Because api conditions in Finland are not typical of all
of the range of Eurasian Rangifer, a logical next step
would be to extend the comparison to include Old
World wild tundra Reindeer. One can conclude, how-
ever, that VSI is potentially a useful tool in determin-
ing nivally suitable areas for Caribou overwintering.

Literature Cited

Henshaw, J. 1968. The activities of the wintering Caribou
in northwestern Alaska in relation to weather and snow
conditions. International Journal of Biometeorology
12(1): 21-27.

Klein, G. J., D. C. Pearce, and L. W. Gold. 1950. Method
of measuring the significant characteristics of a snow-
cover. Ottawa National Research Council Technical
Memo. Number 18, Publication 2269. 22 pp.

Nasimovich, A. A. 1955. The role of snow cover conditions
in the life of ungulates in the USSR. Institute of Geo-
graphy, Academy of Science USSR, Academy of Science
Press, Moscow. 402 pp.

Pruitt, W. O. 1959. Snow as a factor in the winter ecology
of Barren Ground Caribou. (Rangifer arcticus) Arctic
12(3): 158-179.

Pruitt, W.O. 1966. Ecology of terrestrial mammals. Jn
Environment of the Cape Thompson region, northwestern
Alaska. Chapter 20. United States Government Printing
Office, Washington. pp. 519-564.

Pruitt, W. O. 1979. A numerical “Snow Index” for Rein-
deer (Rangifer tarandus) winter ecology. (Mammalia:
Cervidae). Annales Zoologici Fennici 16(4): 271-280.

Skogland, T. 1978. Characteristics of the snow cover and

1981

its relationship to wild mountain Reindeer (Rangifer
tarandus tarandus) feeding strategies. Arctic and Alpine

NOTES

365

International Reindeer—Caribou Symposium: pp. 324-
334.

Research 10(3): 569-580.
Stardom, R.R.P. 1975. Woodland Caribou and snow
conditions in southeast Manitoba. Proceedings First

Received 19 November 1980
Accepted 4 March 1981

First Canadian Record of Bering Cisco (Coregonus laurettae) from the
Yukon River at Dawson, Yukon Territory

DIRK A. DEGRAAF

Beak Consultants Limited, 3530 Ila Street N.E., Calgary, Alberta T2E 6M7
Present Address: LGL Limited, Environmental Research Associates, P.O. Box 13248, Station A, St. John’s, Newfoundland
AI1B 4A5

deGraaf, Dirk A. 1981. First Canadian record of Bering Cisco (Coregonus laurettae) from the Yukon River at Dawson
Yukon Territory. Canadian Field-Naturalist 95(3): 365

A Bering Cisco (Coregonus laurettae) was captured in the Yukon River at Dawson. This constitutes the first record of this

species in Canadian waters and a range extension of 500 km upstream.

Key Words: Coregonus laurettae, Bering Cisco; Yukon River, range extension.

On 21 September 1977 a specimen of Bering Cisco
(Coregonus laurettae) was captured by gill net during
fish inventory studies in the Yukon River at Dawson.
This constitutes the first record of the species in Cana-
dian waters and a range extension of 500 km upstream
in the Yukon River from previously recorded catches
at Fort Yukon (Alt 1973). The specimen was cata-
logued (NMC 77-1771) at the National Museum of
Natural Sciences in Ottawa; D. E. McAllister con-
firmed the identification.

The specimen was a male with moderately large
testes, but it did not appear to be in spawning condi-
tion. Proportions and meristics agree with published
descriptions (McPhail and Lindsey 1970; Scott and
Crossman 1973) except that there are 10 principal anal
rays instead of the usual 12 to 14. There were 37 gill
rakers with 23 on the lower arch; an X ray showed 62
vertebrae including the urostyle. Its total length was
350 mm and the standard length 306 mm, which is
near the maximum size (36 cm) reported by McPhail
and Lindsey (1970), but well within the range of up to
475 mm reported at Alt (1973).

The biology of this species has not been described in
any detail in the published literature; the only account
since McPhail (1966) established that they were suffi-
ciently distinct from Arctic Cisco (Coregonus autum-
nalis) to warrant species status is by Alt (1973). Bering
Cisco approaching spawning condition were captured
from the Yukon River, Alaska, near the mouths of
both Hess Creek and the Notwitna River during early
and mid-June; the run peaked at Rampart in early
September (Alt 1973). Relatively large catches were

also reported from a fish wheel at Fort Yukon in late
August. The data of Alt (1973) and this record suggest
that Bering Cisco probably spawn in the main stem
Yukon River between Fort Yukon, Alaska, and the
Canadian border.

If substantial numbers of Bering Cisco penetrated
the Yukon River system as far as Dawson ona regular
basis, they would probably occur regularly in fish
wheels operated between Dawson and the Alaskan
border, though, because of their size, not in salmon or
whitefish nets. A small catch by fish wheel operators
would probably not be noted because some salmon
fishermen questioned referred to all coregonids as
“inconnu” and routinely discarded them.

This range extension resulted from impact-related
studies funded by Foothills Pipe Lines (Yukon) Ltd.

Literature Cited

Alt, K. T. 1973. Contributions to the biology of the Bering ©
Cisco (Coregonus laurettae) in Alaska. Journal of the
Fisheries Research Board of Canada 30: 1885-1888.

McPhail, J. D. 1966. The Coregonus autumnalis complex
in Alaska and northwestern Canada. Journal of the
Fisheries Research Board of Canada 23: 141-148.

McPhail, J. D.,and C. C. Lindsey. 1970. Freshwater fishes
of northwestern Canada and Alaska. Fisheries Research
Board of Canada, Bulletin Number 173. 381 pp.

Scott, W. B., and E. J. Crossman. 1973. Freshwater fishes
of Canada. Fisheries Research Board of Canada, Bulletin
Number 184. 966 pp.

Received 26 September 1980
Accepted 26 January 1981

Reports of Significant Range Extensions

Sharp-shinned Hawk, Accipiter striatus (Accipitriformes:
Accipitridae) on Victoria Island, Northwest Territories

Holman, Northwest Territories (70°43’N,
117°30’W) early November 1979, in the village.

The bird was captured inside a garage by a local
Inuk hunter. The live bird appeared to be blind in one
eye at the time of capture. It was killed, photographed,
kept frozen, and later given to me. Unfortunately, the
specimen was lost in shipment to our laboratory; thus
the sex was not determined.

Two photographic slides of the specimen are on file
in the Ornithology Section, National Museum of
Natural Sciences, Ottawa. Confirmation of the identi-
fication was made by W. Earl Godfrey, Curator Eme-
ritus of Ornithology.

Sharp-shinned Hawks are known to breed in the

northern Yukon and southern District of Mackenzie
but have not been recorded as accidentals in the Can-
adian arctic islands. This constitutes an extension of
947 km in a direction of 56°97’ from the nearest
record at Old Crow, Yukon Territory.

THOMAS G. SMITH

Arctic Biological Station, Department of Fisheries and
Oceans, 555 St. Pierre Blvd., Ste. Anne de Bellevue, Quebec
H9X 3R4

Received 11 July 1980
Accepted | March 1981

Two-lined Salamander, Eurycea bislineata
(Amphibia: Caudata: Plethodontidae), in Labrador

Cache River tributary, Labrador, Province of New-
foundland (53° 12’N, 62°16’W), 16 August 1979.

Several larval Two-lined Salamanders were cap-
tured by electrofishing during a baseline population
study of Brook Trout (Sa/velinus fontinalis). The
stream followed a deeply incised valley in open Black
Spruce (Picea mariana) lichen forest, which is typical
of the Labrador Plateau. At the collection site, situ-
ated 100 m downstream of a small lake, the stream
was 3.9 m wide, the mean depth was 0.14 m, and the
mean cross-section water velocity was 0.3 m/s. The
stream substrate consisted of boulders and rock rub-
ble, most of which were covered with aquatic mosses.
The site was shaded by overhanging alders (A/nus
spp.), willows (Salix spp.), and Black Spruce. At the
time of collection water temperature was 14°C, dis-
solved oxygen 8 mg/L, pH 4.76, and specific conduc-
tance 14 wS/cm; these conditions are similar to those
of other streams in the area (Jamieson 1979). Such low
specific conductance is generally associated with low
biological productivity (Ryder et al. 1974) which was
further evidenced by a sparse benthos (122.0 ani-
mals/m7?). Despite the indications of low productivity,
the population of Brook Trout at this site was rela-
tively high (39.4 fish/ 100 m2) when compared with

other sites in Newfoundland and Labrador. Most of
the trout were smaller than 100 mm (fork length). The
only other fish species captured in this stream was
Lake Chub (Couesius plumbeus).

The only previous record of a Two-lined Sala-
mander from Labrador was a juvenile found near
Labrador City (Cook and Preston 1979), over 250 km
W of the Cache River site. Based on verbal reports of a
small salamander in the Lake Melville region (Bleak-
ney 1958), Cook and Preston (1979) hypothesized the
existence of the Two-lined Salamander in central and
eastern Labrador. Another species, the Blue-spotted
Salamander (Ambystoma laterale), is known from the
Goose Bay area (Cook and Folinsbee 1975). Although
the Lake Melville salamanders could be either species,
our-record supports the suggestion by Cook and Pres-
ton (1979) that the Two-lined Salamander has a wider
Labrador range than their single specimen indicated.

One specimen was retained for identification and
has been deposited in the National Museum of Natu-
ral Sciences, Ottawa, Canada; catalogue number
NMC 20562. This larva was 61 mm in total length and
30 mm from snout to the anterior margin of the vent.

We acknowledge the Lower Churchill Develop-
ment Corporation Ltd. which funded the aquatic

366

1981

investigations resulting in the collection of the
specimen.

Literature Cited

Bleakney, J.S. 1958. A zoogeographical study of the
amphibians and reptiles of eastern Canada. National
Museum of Canada Bulletin 155: 1-119.

Cook, F. R., and J. Folinsbee. 1975. Second record of the
Blue-spotted Salamander from Labrador. Canadian
Field-Naturalist 89: 314-315.

Cook, F. R., and J. Preston. 1979. Two-lined Salamander,
Eurycea_ bislineata, in Labrador. Canadian Field-
Naturalist 93: 78-179.

Jamieson, A. 1979. A water quality atlas for streams and
lakes of Labrador. Canadian Fisheries and Marine Service
Data Report Number 148. 53 pp.

Ryder, R.A., S.R. Kerr, K.H. Loftus, and H.A.

RANGE EXTENSIONS

367

Regier. 1974. The Morphoedaphic index, a fish yield
estimator—review and evaluation. Journal of the Fisher-
ies Research Board of Canada 31: 663-688.

DIRK A. DEGRAFF!
BRUCE K. BOLES?
JAMES A. LOVISEK3

'Atlantic Biological Services Ltd., P.O. Box 13248, Station
A, St. John’s, Newfoundland AIB 4A5

*Northland Associates Ltd., P.O. Box 1734, St. John’s, New-
foundland AIC 5P5

3Royal Ontario Museum, Toronto, Ontario M5S 2C6

Received 22 September 1980
Accepted 28 January 1981

News and Comment

John M. Powell — Honoured

The sixth annual Loran L. Goulden Memorial
Award for outstanding contributions to natural his-
tory in Alberta was awarded to John M. Powell of the
Northern Forestry Research Centre, Edmonton, at
the Annual General Meeting of the Edmonton Natu-
ral History Club on 13 January 1981. Dr. Powell isa
well-rounded naturalist, who has published on such
diverse areas of nature study as climatology, botany,
entomology, ornithology, mammalogy, and history of
naturalists. He was particularly recognized in this
award for his outstanding dedication to the Federa-
tion of Alberta, Naturalists (FAN) and its corporate
member clubs, having served as FAN’s first Secretary,
third President, and a continuous board member in

Report on New Titles of Books

The year 1980 (volume 94) was rather successful for
the book review section of The Canadian Field-
Naturalist. The number of reviews published, 48, was
down from 1979, but greater than 1978. However, 70
reviews were completed and submitted for publica-
tion, which will mean a significant increase in 1981.
This trend should continue because 67 books were
sent to reviewers. The 93 complimentary New Titles
received from publishers represents a substantial
increase Over previous years indicating either an
increase in natural history literature or in the reputa-
tion of our journal.

Request for Information — Marked Snowy Owls

Since 1977 I have banded over 60 Snowy Owls
wintering in the Duluth, Minnesota harbor area. Owls
were individually marked with black dye on wrists,
wing, and/or tail feathers, and fitted with a green
patagial wing tag with white letter-number combina-
tion on the right wing. Fifteen of 32 owls banded in
1978-1979 and 1979-1980 returned to the harbor

' study area in subsequent winters. Observers are asked
to look for and report any marked owls seen, espe-
cially breeding owls. Where feasible, banding of
marked owl’s young would be greatly appreciated.

FAN’s 10-yr history, and also as a former President of
the Calgary Field Naturalists and Director of the
Edmonton Natural History Club. He has also repres-
ented FAN on various government committees. A
biography of John M. Powell by Charles D. Bird will
appear in the forthcoming history of the Federation of
Alberta Naturalists and its corporate member clubs,
and his publications will be listed in the March 1981
issue of the Alberta Naturalist.

MARTIN K. MCNICHOLL

Chairman, Loran L. Goulden Memorial Award Committee

Authoritative book reviewers are always in
demand. Persons are especially needed to review New
Titles listed as available, but the Book Review Editor
will make every effort to obtain review copies of other
appropriate titles that are requested by prospective
reviewers. Further information and guidelines for
preparation of reviews can be obtained by contacting
Dr. Wilson Eedy, Book Review Editor, R.R. 1, Mof-
fat, Ontario LOP 1JO.

Reports should include place, date, color marks (may
be quite faded), and wing tag number, general habitat,
evidence of breeding, and abundance of small mam-
mal prey. Adult females occupying adjacent winter
territories have returned to occupy adjacent territories
in subsequent winters; they may be breeding in the
same locality. All reports will be acknowledged and
should be sent to: David L. Evans, 2928 Greysolon
Road, Duluth, Minnesota 55812 (218-724-0261) and
to the Bird Banding Laboratory, Office of Migratory
Bird Management, Laurel, Maryland 20811.

368

1981

Restoration and Management Notes

Free copies of the first issue of Restoration and
Management Notes, a new publication from the Uni-
versity of Wisconsin-Madison Arboretum, will be
available to interested persons on request. The publi-
cation, to be made up mostly of short notices dealing
with the techniques and principles of restoring and
managing communities of native plants and animals,
is intended to encourage communication between

NEWS AND COMMENT

369

researchers, managers, naturalists, landowners, and
others involved in the active conservation of natural
and semi-natural areas. The first issue is scheduled for
publication in April 1981. Those interested in receiv-
ing a complimentary copy are invited to contact Dr.
W.R. Jordan, III, The University of Wisconsin-
Madison Arboretum, 1207 Seminole Highway, Madi-
son, Wisconsin 53711. (608) 263-7888.

Notice of Motions to Amend the Constitution of The Ottawa Field-Naturalists’ Club

Notice of four motions 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 8 now reading:
“OFFICERS. The officers of the Club shall be a
President, a Vice-President, a Recording Secre-
tary, a Corresponding Secretary, and a
Treasurer.”
be amended to read as follows:
OFFICERS. The officers of the Club shall be a
President, a First Vice-President, a Second Vice-
President, a Recording Secretary, a Correspond-
ing Secretary, and a Treasurer.

Article 10, now reading:
“STANDING COMMITTEES. Four standing
committees, each consisting of at least five
members, shall be appointed by the Council,
namely: a Publications Committee, an Excur-
sions and Lectures Committee, a Finance Com-
mittee, and a Membership Committee. The
Chairman of each standing committee shall be
chosen from among the members of the Council.
The Editor of The Canadian Field- Naturalist and
the Business Manager of The Canadian Field-
Naturalist shall be members of the Publications
Committee. The Vice-President, the Treasurer
and the Business Manager of The Canadian
Field- Naturalist shall be members of the Finance
Committee. The Chairmen of the Excursions and
Lectures Committee and the Membership Com-
mittee shall have power to add to their
Committees.”
be amended to read as follows:

STANDING COMMITTEES. Six standing com-
mittees, each consisting of at least six members,
shall be appointed by the Council, namely: an
Executive Committee, a Publications Commit-

tee, an Excursions and Lectures Committee, a
Finance Committee, a Conservation Committee,
and a Membership Committee. The Chairman of
the Executive Committee shall be the President.
The Chairman of each other standing committee
shall be chosen from among the members of the
Council.

Both Vice-Presidents and the Recording Secre-
tary shall be members of the Executive Commit-
tee. The membership of each standing committee
shall contain at least one of the Vice-Presidents.
The Treasurer and the Business Manager of The
Canadian Field- Naturalist shall be members of
the Finance Committee. The Editor of The Can-
adian Field- Naturalist and the Business Manager
of The Canadian Field-Naturalist shall be
members of the Publications Committee. The
Chairmen of the Excursions and Lectures Com-
mittee, the Conservation Committee, and the
Membership Committee shall have power to add
to their Committees.

Article 17, now reading:
““DUTIES OF THE VICE-PRESIDENT. In the
absence of the President, or at his request, the
Vice-President shall act in his stead. The Vice-
President shall be a member of the Finance
Committee.”

be amended to read as follows:
DUTIES OF THE VICE-PRESIDENTS. In the
absence of the President, or at his request, either
the First or Second Vice-President shall act in his
stead. Both Vice-Presidents shall be members of
the Executive Committee. The membership of
each standing committee shall contain at least
one of the Vice-Presidents.

Article 18, now reading:
“DUTIES OF THE RECORDING SECRETARY. The
Recording Secretary shall keep minutes of the
proceedings of the Council, the Annual Business

370

Meeting and Special Business Meetings. He shall
give previous notice to each member of the
Council of its meeting and to the general mem-
bership of the Annual Business and Special Busi-
ness Meetings. He shall be the custodian of the
Constitution and the By-laws and of the records
of the Club. He shall be the compiler of the
Annual Report of the Council and shall make it
available to the General Membership at the
Annual Business Meeting. He shall receive and
deal with proposed motions to amend the consti-
tution pursuant to Article 23.”
be amended to read as follows:

DUTIES OF THE RECORDING SECRETARY. The
Recording Secretary shall keep minutes of the
proceedings of the Council, the Annual Business
Meeting, and Special Business Meetings. He
shall give previous notice to each member of the
Council of its meeting and to the general mem-
bership of the Annual Business and Special Busi-
ness Meetings. He shall be the custodian of the
Constitution and the By-laws and of the records
of the Club. He shall be the compiler of the
Annual Report of the Council and shall make it
available to the General Membership at the

THE CANADIAN FIELD-NATURALIST

Vol. 95

Annual Business Meeting. He shall receive and
deal with proposed motions to amend the consti-
tution pursuant to Article 23. The Recording
Secretary shall be a member of the Executive
Committee.

Motions proposed by Daniel F. Brunton, seconded by
Paul Catling.

Frank Pope
Recording Secretary

Explanatory Note

There are two basic reasons for proposing these motions.
First, the expanding programs and activities of The Ottawa
Field-Naturalists’ Club are placing a heavy burden on the
President and Vice-President and in order to share this work-
load more equitably, a Second Vice-President position is
proposed. The two Vice-Presidents should be considered
equal in all respects. Second, it is felt that the important and
fundamental roles of the Executive and Conservative Com-
mittees in Club activities should be recognized by designat-
ing them as standing committees of Council.

Notice of The Ottawa Field-Naturalists’ Club Annual Business Meeting

The 103rd Annual Business Meeting of The Ottawa Field-Naturalists’ Club will be held in the auditorium of
the Victoria Memorial Museum Building, Metcalfe and McLeod Streets, Ottawa, on Tuesday, 12 January 1982,

at 20:00.

Frank Pope
Recording Secretary

Call for nomination 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 1982, as
required by the Constitution.

Club members may also nominate candidates as
officers and other members of Council. Such nomina-
tions 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 K1 Y 4JS, to arrive
no later than 13 November 1981.

The Committee will also consider any suggestions
for nominees which members wish to submit to it by |
November 1981. It would be helpful if some relevant
background on the proposed nominees were provided
along with the suggested names.

DANIEL F. BRUNTON
Chairman, Nominating Committee

1981

NEWS AND COMMENT

The Ottawa Field-Naturalists Club

Special Publications

1. Autobiography of John Macoun
A reprint of the 1922 edition of the fascinating life story of one of Canada’s outstanding early
naturalists, with a new introduction by Richard Glover and bibliographical essay, footnotes,
and index by William A. Waiser, plus three maps of John Macoun’s western travels.
Individuals $12.50 plus $2 postage and handling
Libraries $15.00 plus $2 postage and handling

2. Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index.
Compiled by John M. Gillett
A complete author, title, and subject index to the predecessors of The Canadian Fiela-
Naturalist, the first thirty-nine volumes of the publications of The Ottawa Field-Naturalists’
Club.
$25 plus $2 postage and handling

Centennial Bird Record

Songs of the Seasons
More than fifty eastern North American birds and amphibians are presented in full stereo-
phonic sound as recorded in the wild by wildlife recording expert F. Montgomery Brigham.
$9.11 (postage and handling included but
Ontario residents must add 7% sales tax

Please send orders to:

The Ottawa Field-Naturalists’ Club
Box 3264 Postal Station C
Ottawa, Ontario, Canada

KIY 4J5

371

Book Reviews

ZOOLOGY

Birds of Pacific Rim National Park

By David F. Hatler, R. Wayne Campbell, and Adrian Dorst.
1978. Occasional Paper Series Number 20. British
Columbia Provincial Museum, Victoria. 194 pp., illus.
$3.00.

Although Pacific Rim National Park, British
Columbia, forms the focal point for this study, the
authors chose to work “from an ecological rather than
a real estate perspective.” The study area thus consists
of the west coast of Vancouver Island from Qlayoquot
Sound south to Port San Juan, including several
pelagic records well beyond sight of the shore. The
book begins with a series of short introductory sec-
tions, including the history of work on birds in the
area, methods of research, and a checklist of 247 bird
species “known or strongly suspected” to occur in the
park, to which Clark’s Nutcracker is added in the
Preface. The bulk of the text consists of detailed spe-
cies accounts, followed by a series of transects at
specific places within the park. Several appendices
and an index complete this volume.

The species accounts range in length from one
paragraph to about four pages. Each documents the
status of the species in the park area, often in compari-
son with its status on other parts of Vancouver Island
and/or the lower British Columbia mainland. A chart
of the number of records per month is given for many
species and serves as a rough guide to seasonal occur-
rence. Records are based ona combination of histori-
cal information, incidental and exploratory observa-
tions by 104 observers, and systematic surveys
conducted for Parks Canada. General ecology and life
history data are included for many species, and anec-
dotes of behavioral observations are scattered
throughout. Species accounts on cormorants, Great
Blue Heron, wintering swans, Bald Eagle food and
feeding behavior, Osprey, Black Oystercatcher,
Glaucous-winged Gull, several alcids, and Northwest-
ern Crow are in sufficient detail that the book should
be included in the bibliographic files of any researcher
working on these species. Other tidbits of information
which may be missed by people not inclined to read
local avifaunal works include a comparison between
courting behavior of the two goldeneye species, a
comparison of the feeding areas of Surf and White-
winged Scoters, a record of an apparent hybrid
Glaucous-winged X Western Gull, speculation on
feeding habits of Chestnut-backed Chickadees, and
observations of Starlings apparently copying turn-
stone feeding methods. Naturalists concerned about

a7

their effects on birds should read the account of the
Common Merganser for an example of why attempts
to “rescue” ducklings separated from the brood are
unwise.

When distinct races of a species are found to be
conspecific, there is an unfortunate tendency for many
observers to stop noting which race they see in particu-
lar areas. Therefore, I was pleased to read the authors’
differentiation between the breeding and common
migrant Audubon’s race and the rare migrant Myrtle
race of the Yellow-rumped Warbler. The authors also
state clearly that Oregon Junco is the race of the
Dark-eyed Junco in the park, thus indicating that the
Slate-colored has not yet been identified there. An
appendix comparing recent names (as used in the
book) with those still included in many field guides

will be helpful to casual observers.

The transects near the end of the book should prove
useful as local bird-finding guides to visiting natural-
ists. Each contains a table of numbers of each species
seen on various dates, plus a map on which 1s indi-
cated species seen at particular times of the year (or all
year) and “special” species sometimes found in the
vicinity.

Birds of Pacific Rim National Park is an attractive
volume, well written and amply illustrated with both
photographs and drawings. Data on many species
(especially waterbirds) are more extensive than usu-
ally included in local avifaunal works. Inclusion of
such data in this book will irritate some researchers
who are more likely to overlook some information
pertinent to their interests than if the authors had
summarized the data in a separate journal publica-
tion. On the other hand, inclusion of these facts adds
interest to the book and might stimulate lay people to
make (or report) further observations. Moreover, the
publishing priorities of the three active authors may
preclude at least some of these data from ever being
published if they were not included in the book.

The book is remarkably error free, but statements
in two species’ accounts are slightly misleading. In
discussing three territorial male Blue Grouse, the
authors mention that the females “belonging to them”
were not seen, implying at least a temporary pair
bond. The best evidence to date, collected also on
Vancouver Island, suggests that this species is promis-
cuous. Observations of Townsend’s and Wilson’s
warblers feeding young Brown-headed Cowbirds are
apparently taken as evidence these warblers were the
hosts. While this may well be true, non-host adult

1981

birds will sometimes feed demanding young cowbirds,
and such observations are not sufficient in themselves
to establish the feeding bird as the host. Incidently,
Friedmann and his co-workers have compiled two
host lists since the one cited (Auk 88: 239-255. 1971;
and Smithsonian Contributions to Zoology 235.
1977). To these minor points, I add one additional
criticism. The authors use “Hypothetical” status in
two ways: both for species identified but without sub-
stantiating evidence, and for species expected to occur
on the basis of occurrence nearby. The former use is
more common, and the latter species would probably
have better been relegated to an appendix.

BOOK REVIEWS

S13)

This book is a “must” item on the bookshelves of all
west coast ornithologists. Compilers of local avifau-
nal works elsewhere would do well to use it as a
standard. At the low price, it should easily be in the
budget for bird-watching visitors to Vancouver Island
and for researchers on species for which extensive
accounts are included. I recommend it highly.

MARTIN K. MCNICHOLL

128 Silvergrove Hill N.W., Calgary, Alberta T3B 4Z5

The Alaskan Bird Sketches of Olaus Murie with Excerpts from his Field Notes

Compiled and edited by Margaret Murie. 1979. Alaska
Northwest Publishing, Anchorage. 64 pp. illus. (51 paint-
ings). $14.50.

It is difficult to consider just one Murie brother.
Both Olaus and Adolph Murie were pioneers in the
early days of game management, bridging the time in
wild Alaska from the dog sleigh to the snowmobile,
and pushing back the frontiers of biological knowl-
edge in such widely separated and wildly different
places as the turbulent Aleutians, cold-impoverished
Labrador, and islands of wilderness in the south such
as Wyoming’s Grgnd Teton Mountains. It was
Adolph who, perhaps, published most memorably
about the north (The Wolves of Mount McKinley in
1944: A Naturalist in Alaska in 1961), but his southern
contributions were formidable too (The Moose of Isle
Royale in 1934; Ecology of the Coyote in Yellowstone
in 1941). Olaus’ large and definitive work The Elk of
North America, published in 1951, is of course
“southern,” but his first major pioneering work,A/as-
ka-Yukon Caribou in 1935, was remarkable for its
ecological approach to data gathered along lonely
miles behind a dog team. His Food Habits of the
Coyote in Jackson Hole, Wyoming, published in the
same year, is an early example of the geographic
dichotomy of interest shared by both brothers. The
last major work published by Olaus documents
studies done in earlier years. Fauna of the Aleutian
Islands and Alaska Peninsula was published in 1959.

Olaus Murie was a leader through the early exciting
years in wildlife biology, when Aldo Leopold’s Game
Management \ed the way to “wise use” of wildlife
resources, and whena maturing knowledge of ecology
combined productively with wildlife biology, the
principles of land use, and geographical explorations
into mountains still unmapped and along shores still
unknown. It was when wildlife management was com-
ing of age; when its leaders were broadly based natu-

ralists and were proud of it. From this base of know-
ing and loving the land and its life evolved today’s
more specialized approaches to wildlife research and
management.

Olaus Murie died in 1963. His wife, Margaret, sur-
vives him, and in the book reviewed here brings
together a collection of his bird paintings made in
Alaska, many in the Aleutians, through the 1920s and
1930s. Appropriate passages from his field notes of
the time accompany the sketches.

The introduction by Margaret Murie puts the
times, the places, and the work into context. She is
well qualified for the role. At a time when society was
not sure that it approved and when life in the wilder-
ness had few of today’s comforts, she was usually with
Olaus in the field. So too were their children even
before they could walk.

This is a book of pictures, all in full color, and the
word “sketches” in the title is correctly used. Most are
field sketches, the “soft parts” paintings of the thor-
ough ornithologist collecting little known birds, and
so usually consisting of head and neck portraits in
color accompanied often by a foot and leg in similar
detail. The main purpose of such sketches is to catch
the fleeting colors of birds that soon fade after death,
such as the hues of the bill, eye, bare areas on the head
and neck when they occur, and the foot and leg. This
need of accurate record is the main incentive to pro-
duce such works, but despite the rough to nearly
impossible conditions under which they must be exe-
cuted, they occasionally qualify as superb ornitholog-
ical portraiture, and more rarely as good art aside
from their dedication to visual truth.

The sketches in this book have unusual value, for
they feature little known details of Alaskan species of
birds still intimately known to few scientists, and to
even fewer with active paint brushes. Those of Whis-
kered Auklet, Crested Auklet, Parakeet Auklet,

374

Sabine’s Gull, downy young of Wandering Tattler, the
same of Spectacled Eider, Bristle-thighed Curlew, and
Red-faced Cormorant are among the special ones. As
a bonus, some to me are high ornithological art,
among them the curlew and downy eider mentioned
above, and a superb portrait of an immature male
Common Eider.

Olaus Murie was a productive and careful scientist
with wide interests and many talents. In his later years
he was a leading force in the Wilderness Society, and
he had much to do with the United States Govern-
ment’s Wilderness Act, brief facts speaking volumes

A Bird-Finding Guide to the Toronto Region

By Clive E. Goodwin, 1979. The Toronto Field Naturalists
Club, Toronto. V + 97 pp. $2 plus 50¢ postage

This guide is intended for the newcomer to bird
watching in the Toronto region, either those coming
from other areas or for long-time residents just getting
interested in the sport of finding birds.

After an introductory chapter on the plan of the
book and abbreviations used init, there isa chapter on
the habitats in the area and their associated bird spe-
cies, then a chapter on these species indicating when to
look for them on a month-by-month basis.

This is followed by several chapters giving helpful
directions for finding specific bird sites, from the lake-
shore back to the moraine country to the north, witha
summary of special associated birds to look for at
different seasons. I am personally most familiar with
sites in the eastern part of the Toronto region where I
found the author’s directions to key areas straightfor-
ward and the highlights appropriate. The booklet
concludes with chapters “for the newcomers” and with
a systematic list with the best month and best areas
suggested for each species.

This is a sturdy paperback, 6 x 9% in. (1 in. =

The Black Bear in North America

Edited by D. Burk. 1979. Boone and Crockett Club, Alexan-
dria, Virginia. 299 pp. U.S. $10.

This book contains the proceedings of a workshop
on the management of the North American Black
Bear. The goal of the workshop was threefold: First,
to review the state of existing knowledge and the
present status of the Black Bear in each state and
province in which the species is found; second, to
develop policy on all aspects of management; and
third, to produce a document that will serve to assist

THE CANADIAN FIELD-NATURALIST

Vol. 95

about what he thought important. What he saw and
thought and did through his life will be built upon and
enjoyed for a very long time. This book of sketches
should be followed by other publications based on his
journals and papers, for those who know about him,
and for many more who will discover him.

Y ORKE EDWARDS

British Columbia Provincial Museum, Victoria, British
Columbia V8V 1X4

25.4 mm) with an attractive cover photograph of a
Saw-whet Owl. It contains three maps, one of the
Halton Region Forest, one of Metropolitan Toronto,
and one of the Toronto Region as a whole, each with
arrows or numbers showing the general locality of
points of particular interest to bird watchers.

The book is meant to be used in conjunction with a
good road map of the region, as the maps in the book
are not adequate for finding specific areas, even
though the written directions are very good with a
map in hand. I would have preferred bar-type graphs
to show the seasonal occurrence of each bird rather
than the method used, but this might have added to
the expense of what is now an exceptionally inexpen-
sive guide. A title on the spine of the booklet would
have been helpful in locating it on a crowded
bookshelf.

This is a very helpful guide for anyone interested in
the bird life of the Toronto region.

J. MURRAY SPEIRS

1815 Altona Rd., Pickering, Ontario LIV 1M6

managers and biologists in the planning and decision-
making process.

The book is divided into four parts. Part one is a
summary of existing knowledge relevant to Black
Bear management. There are reports from 37 states, 9
provinces, the Yukon and Northwest Territories, and
Mexico. In general, representatives from each juris-
diction endeavored to review habitat, laws and regula-
tions, protection, management, population trends,
hunting, public attitudes, and current problems. The

198]

representatives from Michigan presented three papers
that dealt with hunting with hounds, age determina-
tion from premolar sections, and age determination
from cementum annuli. A review of the Black Bear’s
status was not outlined for Michigan.

It was genérally felt that compared to other major
wildlife species, knowledge of the Black Bear is pres-
ently insufficient to enable biologists to develop suita-
ble management strategies. Many jurisdictions have
initiated research and management programs only in
the last decade. There is much still left to learn about
the Black Bear in Central and North America.

Part two consists of a paper presented by Stephen
Herrero of the University of Calgary, entitled “Black
Bears — the Grizzlies Replacement.” Herrero pre-
sents the theory that Black Bears, despite their appar-
ent adaptibility to man and altered habitat, will not
succeed the Grizzly Bear in its habitat for evolved
morphological and behavioral reasons.

Part three is comprised of the reports from the
Pacific, Rocky Mountain, southeast, and central and
northeast Canada and the United States working
groups. Consideration is given to: habitat, land use,

The Freshwater Fishes of Alaska

By James E. Morrow. 1980. Alaska Northwest Publishing
Company, Box 4-EEE, Anchorage, Alaska. 248 pp., illus.
U.S. $29.95 (+ $.75 U.S. Postage and handling).

This in an informative, pleasingly designed, well-
illustrated book on the 56 species of fishes occurring in
the freshwaters of Alaska. A considerable amount of
information is packed into the two and a half (814 X
1] in.) pages devoted on the average to each species.
The text, which is written in readable prose rather
than telegraphic statements, should appeal to the
angler as well as the scientist, especially as the text’s
layout and typography is pleasing to the eye.

The book is divided into an introductory section, a
key to the families, family and species accounts, a
25-page bibliography, a glossary, and an index. The
color photos and illustrated plates are grouped
together in the middle of the book. Some color photos
are excellent (the elegant spawning livery of the Dolly
Varden) while others are based on moribund speci-
mens. The carbon dust illustrations by Marion J.
Dalen combine detail, precision, and the spark of life.
I care less for her water colors which do not seem to
capture the luminosity of live pigments.

A chapter is devoted to each family or subfamily.
The chapter usually begins with a key to species and
an account of the distinctive morphological and bio-
logical features of the family. Important diagnostic
characters are often illustrated to help the reader use

BOOK REVIEWS

S75

management, hunting, population trends and density,
illegal activities, bear-man conflicts, public education,
and research needs. Corresponding recommendations
are set forth in the text. For example, timber man-
agement guidelines designed to protect and enhance
Black Bear habitat were submitted by the Pacific
working group. They include recommendations that
deal with the size, spacing, and frequency of clear cuts,
as well as herbicide use, road construction, and access.

Part four, entitled “Final Session Comment,” is a
dialogue between the participants during which the
chairmen summarize the recommendations developed
by their respective working groups.

The text is well organized; however, a few of the
figures referred to in the text are missing. The book is
a useful reference document and is recommended to
those involved in Black Bear management.

PAUL A. GRAY

Wildlife Service, Government of the Northwest Territories,
Yellowknife, Northwest Territories X1A 2L9

the keys. The species accounts have the following
sections after the common and scientific names: dis-
tinctive characters, description, range and abundance,
habits, and importance to man. Habits, the longest
section, presents an account of the life history of the
species. Outline drawings and maps with ranges indi-
cated by cross-hatching are included for most species.
The reviewer prefers spot distribution maps for the
greater amount of information they convey. Little
emphasis is given to systematics.

The book is well edited and there are few typogra-
phic and scientific errors. The range of Arctic Charr,
however, extends to Alert, northern Ellesmere Island;
and Ninespine Stickleback, contrary to several refer-
ences, are not found in Greenland. The lateral line
“pores” of Blackfish and head “pores” of the Arctic
Lamprey are instead surface neuromasts. Plate 30D
shows eight left and seven right branchiostegals on the
coastrange sculpin; normally six are found on each
side in this species.

This beautifully designed, informative, and reada-
ble book should appeal to the fisherman, naturalist,
and biologist. The author, artist, and editor are to be
congratulated.

Don E. MCALLISTER

Ichthyology Section, National Museum of Natural Sciences,
Ottawa, Ontario KIA 0M8

376

North American Birds of Prey

By William Marshall. Paintings by Gary Low. 1980. Gage,
Toronto. 176 pp., illus. $29.95

North America has had several excellent wildlife
artists. Among bird illustrators, there are many
superb painters from Audubon to Landsdowne. With
the publication of North American Birds of Prey,
another Canadian seems destined to become a select
member of this group.

In this beautiful book, Gary Low has presented 27
full-color paintings of the most common hawks and
owls of northeastern North America. Also provided
for each species is a baseball-card-like facial portrait.
With their penetrating eyes, different beaks, and var-
ied feathering on the face, birds of prey make excellent
subjects for these portraits. Low excels in his color
pictures. An important decision for a wildlife artist is
how purely representative he should be. Roger Tory
Peterson’s paintings for his field guides are to help us
identify birds: they are like pictures. Other artists,
Parnell, for example, take considerable liberties with
the subject, often producing a symbol rather than a
bird. Low demonstrates his ability in both areas. His
paintings of the Turkey Vulture, Broad-winged
Hawk, and Snowy Owl are superb pictures of these
birds. In some of his other paintings, Low dazzles us
with something unexpected which jumps out from the
picture; the Goshawk about to pounce ona rabbit, the
Saw-whet Owl with a kinglet, a Long-eared Owl
‘nustled’ against a tree. These pictures present more
than just bird paintings. My favorite is the picture
chosen for the cover: our most common hawk, the

Wolf and Man: evolution in parallel

Edited by R. L. Halland H. S. Sharp. 1978. Academic Press,
New York. 210 pp., illus. U.S. $22.95.

Wolfand Man, as the title suggests, is a collection of
papers on the topic of human-—wolf co-evolution. The
editors’ purpose is clearly to draw a parallel between
the adaptive behaviors of humans and wolves. This
aim is based on the premise that both humans and
wolves evolved as group hunters. Although this
hypothesis has appeared reasonable to natural scien-
tists for sometime, to my knowledge this is the first
published account of a multidisciplinary analysis of
the topic.

The text is organized in a manner that describes the
cultural attributes, social features, and history of
Homo and Canis. Part one presents four chapters
concerning group behavior of wolf and man as it
relates to culture. Specific reference is made to group

THE CANADIAN FIELD-NATURALIST

Vol. 95

Red-tailed Hawk, sitting on a post which is part of an
elegant fence.

The only disappointment with the book is with the
falcons — to some the essence of avian predators.
Unfortunately, all four of the falcons are depicted
settled, three on trees and one ona rock. I hope Gary
Low will attempt something more daring with these
magnificent birds next time.

Each illustration is accompanied by a short text
written by William Marshall. Like Low, Marshall
faces a choice as well: be very biological and scientific
or be informal and ‘chatty.’ He combines approahces
and offers readers insight into the bird’s behavior and
role. Of special interest are the two back sections, one
on falconry and one on pesticides. These are both
extremely interesting.

Finally, for each bird depicted there is a food sum-
mary at the back of the book, which provides interest-
ing information. It certainly reminds us that the birds
we have been looking at are predators.

North American Birds of Prey is a beautiful book
both for birders as well as those who appreciate fine
paintings. It will be very interesting to watch Gary
Low develop as a painter of wildlife. He clearly shows
he intends to do more than “hold a mirror up to
nature.” This is a bold step. His first book launches
him well on his journey.

DAVID LOVE

World Wildlife Fund (Canada), 60 St. Clair Avenue East,
Suite 201, Toronto, Ontario M4T 1N5

hunting characteristics. A chapter on Coyote natural
history is also included. The following three chapters
represent a treatise of wolf communication features
and functions as they compare to communicative evo-
lution features in man. The final two chapters exam-
ine the paleobiology of dogs and man.

The editors have by and large successfully accom-
plished their endeavor. There are, however, some
areas of assumption made in the text. For example, in
the sections on communication, two papers are pres-
ented (both previously published) that clearly demon-
strate the fact that olfactory and auditory factors
(scent, marking, howling) have an indirect commu-
nicative function in wolves. This premise seems quite
reasonable. The editors next analyze early human
forms of visual communication (1.e., interpretation of
visual signs) on a comparative basis. It seems more

1981

plausible to assume that these communication fea-
tures are in fact divergent evolutionary trends.

The comparative analysis of wolf-human group
hunting behavior on a cultural basis is somewhat
overstated in the text. It is important to realize that
internal wolf pack regulatory behaviors are most criti-
cal to group hunting characteristics. Therefore,
although wolf-human hunting characteristics may be
similar (i.e., Comparative Ethnology of the Wolf and
the Chipewyan), the sociological factors critical to
group hunting characteristics in the two generic
groups are somewhat variable.

In summary, the editors have done an admirable
job of combining the study of behavior from both an

The Squirrels of Canada

By S. E. Woods, Jr. 1980. National Museums of Canada,
Ottawa. 200 pp., illus. $29.95.

This book has a pleasing appearance and is well
designed and well illustrated, as it should be for the
price. The color photographs of each of the 22 species
of marmots, ground squirrels, chipmunks, tree squir-
rels, and flying squirrels discussed are superb, the line
drawings evocative (if one ignores the odd perspective
of eagle and marmot on page 39). The text runs
smoothly and is divided into sections entitled: Names,
giving derivations of the common and specific names;
Description; Range, accompanied by maps of the
Canadian and North American distribution; Behav-
iour; Personality, incorporating a suitable quote from
a naturalist or zoologist; Habitat; Feeding Habits;
Life Cycle; Natural Enemies; Relations with humans;
and Where to observe. This format is handy for refer-
ence and for browsing, but involves much repetition
of information between species (we are told six times
that Spermophilus means “lover of seeds”), and some
within a species (such as the timing of hibernation,
and information on burrows and populations.)

The text is curiously old-fashioned, referring to all
individual animals as “he”; describing the skunk as
“more valuable” than the Woodchuck; calling the
Yellow-bellied Marmot society a patriarchal one; ref-
erring frequently to individuals standing like “picket
pins”; and having animals “perform bodily functions”
rather than urinate and defecate. There are a number
of interesting anecdotes which help to bring the sub-

BOOK REVIEWS

S77

organism and social standpoint. This alone makes the
text well worth reading. Some areas of the text are
superficially treated, such as the chapter on Coyote
natural history which lacks current references.
Persons interested in wolf biology from an anthro-
pological perspective will enjoy the complete text.
Individuals looking for a biological slant to wolf—hu-
man behavioral evolution should seek elsewhere.

ALAN KENNEDY

Environmental Studies Group, Esso Minerals Canada,
500-6th Avenue, S.W., Calgary, Alberta T2P 0S1

jects to life.

Unfortunately there are several errors of fact as wel.
as some typographical errors. It is not correct to say
that primates other than man walk upright, to general-
ize that zoologists use muscle arrangements of a
rodent’s jaw for classification, or to define incisors as
only located on the premaxillary bone. I also find it
difficult to believe that one could confuse a Red
Squirrel and a Fox Squirrel even at a distance, given
their different habits and their great difference in size.
In connection with size, I should mention here discre-
pancies in various measurements. A. W. F. Banfield,
in The Mammals of Canada which 1s listed in the
bibliography, gives the average Red Squirrel weight as
about 190g, while Woods reports 230g; for the
Northern Flying Squirrel, Banfield quotes weights
ranging from 75 to 139 g, while Woods gives an aver-
age weight of 160 g.

This book is aimed at the layman rather than the
professional zoologist. Although a short list of refer-
ences is included, over one-third are pre-1950 and
there are only four original papers cited for the 1970s.
None of the data in the text itself is referenced. There
is a short glossary, a short index, and a seven-line table
of metric conversion figures.

ANNE INNIS DAGG

Integrated Studies, University of Waterloo, Waterloo, Onta-
rio N2L 3G1

378

BOTANY

The Flora of Canada

By H. G. Scoggan. 1978-1979. (Four volumes). Publications
in Botany Number 7 (1-4), National Museum of Canada,
Ottawa. xiii+ 1711 pp. $131.00 (sold only as set).

This massive work sets out to list all species growing
without cultivation in Canada. It recognizes 3218
native species and a further 884 aliens, for a total of
4102 species. To provide treatments for each of these
and their subspecific taxa is a mammoth task and is
the result of many years of study and research.

The resulting product is presented in four parts.
Part 1: General Survey provides an introduction to
the study as well as an analysis of Canadian plant
distribution, the factors affecting it and the resultant
patterns that emerge, a life form classification of vas-
cular plants, and a description of floral regions in
Canada. A brief list of nomenclatural changes result-
ing from the study is presented, followed by a list of
abbreviations and the herbarium acronyms utilized in
the text, a tabular conspectus of the flora, a glossary,
and finally, the literature cited. Parts 2, 3, and 4 pro-
vide the systematic treatments. These are divided into
Pteridophyta to Monocotyledoneae, Dicotyledoneae
(Saururaceae to Violaceae) and Dicotyledoneae (Loas-
aceae to Compositae), respectively. Part 4 concludes
with an index to the complete work.

The four hardbound books constitute a handsome
set of volumes. The printing quality is excellent, with
clear, well-arranged copy being presented on good-
quality, firm, paper stock. Bindings are sturdy yet
flexible, facilitating easy use of the books and ensur-
ing durability. Although type size is necessarily small
for economy’s sake, readability has not been lost.
Different type styles have been skillfully used with a
minimum of negative space to provide a simple,
clearly delivered text to enhance readability further.
The index in Part 4 is printed on differently colored
paper. This isa small thing, but helps the user of these
volumes considerably. The absence of indices for each
volume, necessitating constant reference to Part 4, is
the only negative criticism I would raise in this regard.
Overall, the physical preparation and presentation of
the work are excellent.

Part 1: General Survey includes an interesting and
informative, if somewhat dated, discussion of plant
distributions in Canada and the factors affecting
them. It seems to me, however, that the sections deal-
ing with floral distribution formulae, and families and
genera of restricted distribution, tend to be overly
complex and academic in their application. In the
same vein, I question the utility of defining distribu-
tion patterns so finely . .. to the point of delimiting a

THE CANADIAN FIELD-NATURALIST

Vol. 95

distinct geographical pattern on the basis of two spe-
cies of Jsopyrum, one each in southern Ontario and
western British Columbia.

The discussion of floral regions of Canada is very
complete, perhaps overly so. I wonder if so many
species must be listed to describe characteristic floral
elements for each region? The addition of common
names, however, is appreciated. I found the anecdotal
tangents (such as a discussion of fall colors in the
Great Lakes — St. Lawrence Forest Region) to be
interesting but out of context here. This is particularly
true for the ‘eco-tours’ described in discussion of the
western forest region. These somewhat rambling
divergences become rather confusing and irritating. A
map illustrating these floral regions would have added
to this section.

The glossary is excellent. The author defines a much
wider range of terms than one normally expects to
find in floral manuals and does so in clear, concise
language. I would add similar applause for the Litera-
ture Cited section, were it more complete. For some
reason, many literature references are cited only in the
species’ accounts (e.g. Alyssum desertorum, Arabi-
dopsis saluginea).

Parts 2, 3, and 4 are structured in a clear and
obviously well-thought-out manner. A technical des-
cription introduces each family, followed by a key to
the species therein. Genera are then listed alphabeti-
cally (a very useful feature), with references to pe-
tinent literature accompanying each. Individual spe-
cies’ accounts follow the generic keys and are also
placed in alphabetical order. Specific treatments start
with the distribution formulae described in Part I:
General Survey, followed by a symbol representing
their life form classification. I suspect that these will
be largely overlooked by the reader, because of their
complexity. These are followed by a very general but
adequate habitat description. The account is con-
cluded with a verbal description of the Canadian and
North American range of each species, a listing of
published distribution maps and of synonymy, largely
as that synonymy relates to previously published
Canadian literature. The distribution map feature is
an excellent one that is too often overlooked in such
manuals. The synonymy cited seems to be quite
thorough. Range data, however, are quite inadequate.
The method devised for describing these is cumber-
some and misleading, especially for northern taxa.
Describing the northern limit for such species as
Streptopus amplexifolius and Ranunculus macount
is far less useful than defining their southern limits

1981

(which is not done) and results in a serious loss in
understanding. Some range statements (e.g. Smilax
herbacea “... north to Thunder Bay [Ontario]. . .”)
are far too sweeping and lead to further confusion.
The author seems to have relied very heavily on pub-
lished range descriptions, without conducting a very
critical evaluation of these (e.g. Dryopteris simulata
(= Thelypteris simulata) is cited in Ontario on the
basis of a literature citation; the voucher at DAO was
revised to 7. palustris years ago).

Subspecific taxa, including microspecies, subspe-
cies, varieties, and forms, are keyed out within species’
accounts and are discussed in the same manner as full

species.
The author describes his taxonomic treatment of
the flora of Canada as representing “.. . a relatively

conservative species concept ...”. I found this difficult
to evaluate, as the treatment varies from group to
group. Ferns, for example are very conservatively
treated (e.g. Dryopteris cristata var. clintoniana
rather than D. clintoniana) whereas sedges are both
‘split’ (Carex cumulata, C. merritt-fernaldii, C. brev-
ior) and ‘lumped’ (C. /axiflora s.1.)

A number of inconsistencies run through the text.
Species which seem to be accepted as validly repres-
ented in the flora (e.g. Ulmus pumila, Potamogeton
capillaceus (= P. bicupulatus)) are discussed within
the accounts of other species and are not included in
the keys. The reason for this confusion is not clear.
Lists of hybrids for Carex and Salix are placed after
the general account, but not so for Quercus or Dryop-
teris. (Indeed, the many hybrids of Dryopteris are not
even discussed.) Some taxa which are discussed
separately and are considered to be synonymous with
other accepted species are not listed under the synon-
ymy for that species (e.g. Polygonum persicaria,
including P. puritanorum). Relatively small but
annoying errors are duplicated repeatedly through the
text (e.g. Algonquin Park, Ontario, is in Nipissing
District, Ontario, not Renfrew County), suggesting
that proofing was not as exacting as it could have
been.

The most serious criticism I have of the work is the
degree to which its content is dated. Although the

Multivariate Analysis in Vegetation Research

By Laszlo Orloci. 1978. Junk, The Hague (North American
distributor: Kluwer, Boston.), 2nd edition, ix + 451 pp.,
illus. U.S. $49.00.

This book should be useful to advanced naturalists
interested in mathematical derivations and calcula-
tions of complex statistical analyses involving large

BOOK REVIEWS

1379

author states that research was ended in mid-1972,
one is hard pressed to find many post-1969 references.
As a result, nomenclature and taxonomy are not sig-
nificantly more up-to-date than that of Gray’s Manual
of Botany or Britton and Brown’s Illustrated Flora.
Important literature for such groups as Draba and
Woodsia, available before and/or during 1972, was
not used. This is particularly critical for such a diffi-
cult genus as Draba and leaves the present treatment
virtually unworkable. Recent studies of this latter
group recognized 40 species, in contrast to the 27
recognized in this work. Only 5 of the 7 species of
Spiranthes and 13 of the 15 species of Cirsium, for
example, are discussed here. Important literature
dealing with the grasses of Ontario, the rare plants of
various provinces, the thistles of Canada...andsoon
... also could not be incorporated.

The age of these data probably is also responsible
for many range discrepancies (e.g. the northern limits
of many southern species in Ontario seem to be 100
km or more south of their known extremes).

One could reasonably expect that this long-awaited
manual would be a ‘state-of-the-art’ study .. . a confi-
dent, standard reference which the Canadian botani-
cal community could refer to as their ‘bible.’ This
work has some excellent features — such as a gener-
ally superior layout and physical construction, the
reference to published range maps, a fine glossary —
but these successes are severely compromized by out-
dated data, vague range descriptions, and a number of
serious inconsistencies.

I must conclude that those considering its acquisi-
tion would be better advised to invest the few addi-
tional dollars required to obtain Britton and Brown's
Illustrated Flora, or perhaps save some money by
purchasing Boivin’s Enumeration des Plantes du
Canada and Gray's Manual of Botany.

This Flora of Canada is, unfortunately, not the
‘bible’ that has been eagerly awaited... more for that
which is not included than for that which 1s.

DANIEL F. BRUNTON

Southwick Drive, R. R. 3, Manotick, Ontario KOA 2NO

data sets. The book is unique because it bridges the
gap between advanced plant ecology and complex
statistical analyses. It explains the many different
techniques used to classify the importance of a plant
species in its community using easily collected data
from a few small sampling sites. Most of the tech-

380

niques could easily be used by the animal ecologist.
The book is written in the style of a long discussion or
lecture, and it treats almost all known multivariate
analysis methods.

Orloci describes multivariate methods that are
extremely time-consuming to compute by examples
with tiny data sets making the calculations simpler
and easier to follow. He also provides listings of 27
BASIC computer programs with sample runs which
handle many of the calculations described. These pro-
grams should be easily used by those with some com-
puter training and their results easily interpreted by
those with some statistical background.

In many areas the text becomes highly mathemati-
cal with little obvious connection to vegetative phe-
nomena. For those naturalists without advanced
mathematical training, these sections will be difficult
to read.

Typographical errors are numerous. Eventhougha
good author index appears, the subject index and
glossary have poor coverage limiting the usefulness of

THE CANADIAN FIELD-NATURALIST

Vol. 95

the book as a quick reference. No list of tables and
figures with pagination is given but tables and figures
are cited in the text without page numbers. This
becomes a problem because tables and figures are
often many pages removed from the discussion con-
cerning them and are thus difficult to find.

This 2nd edition is not much different from the first;
a few (7) programs, new references, and some statisti-
cal methods are added. The “List of Symbols” has
been dropped which, in my opinion, should not have
been done.

Perhaps the best way to use this book 1s to read it in
its entirety while taking notes for future reference. In
this spirit, I strongly recommend this book for use by
serious naturalists.

RICHARD M. ZAMMUTO

Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7

Vascular Plants of Continental Northwest Territories, Canada

By A. E. Porsildand W. J. Cody. 1980. National Museum of
Natural Sciences, Ottawa. vill + 667 pp., illus. $80.00.

This monumental work fills a significant gap in
bringing together keys and descriptions of all the spe-
cies of ferns and flowering plants of the mainland
portion of the Northwest Territories. It represents
many years of work and it is exciting to at last have it
available. Gone are the days of tramping the southern
Northwest Territories with backpacks full of provin-
cial and Alaskan floras. It is sad that Dr. Porsild
passed away before the book was published: his con-
tributions will not be forgotten by northern botanists
and students of natural history, for his books are the
basic references on vascular plants.

In addition to their roles as Chief Botanist in Cana-
da’s National Herbarium (A. E. Porsild) and Curator
of Canada’s Department of Agriculture Vascular
Plant Herbarium (W. J. Cody), both authors have
had extensive field experience in the Northwest Terri-
tories. Dr. Porsild especially was no “summer bota-
nist”: he knew the north in all seasons and the value of
plants to native peoples and to wildlife. The descrip-
tions include numerous extra details from mosquito
pollination of the northern bog-orchid to the parts of
plants which are important to northern herbivores
such as Caribou, Muskoxen, and ptarmigan.

The flora covers the 1112 known species plus addi-
tional species which are expected to occur here. There
are line drawings for almost all species and distribu-

tion maps for all the native species. The drawings and
distribution maps are conveniently located together
and immediately following the descriptions of the spe-
cies of each family. The distribution maps also indi-
cate the extent of a plant’s range in the rest of Canada,
in Iceland, Greenland, Alaska, and the western tip of

the USSR.
There is a nine-page section that documents botani-

cal collectors who contributed specimens through
1976, the last date for inclusion of most distribution
information. The collections are by no means exhaus-
tive. It remains to be seen if the ranges of over one
third of the species are as restricted as Cody suggests
in Vascular Plants of Restricted Range in the Conti-
nental Northwest Territories, Canada, another
National Museum publication. It is hoped that this
excellent new flora will stimulate dozens of new dis-
tribution records! A more thorough knowledge of the
Northwest Territories flora would assist in assessing
which of the species with apparently restricted distri-
butions are potentially rare or endangered.

The book is clearly a scientific, academically
oriented publication and an essential acquisition for
botanical and science libraries, and it will be of inter-
est to many northerners and visitors to the north.
Unfortunately, its high price will discourage many
potential purchasers, especially among nonbotanists
who want to take a book of this type with them for
field identifications. (No. There are no color photo-

1981

graphs!) It is unlikely that it will foster northern scien-
ces as muchas the authors might have intended: at this
price it isn’t likely to be readily available to nor-
therners in local settlement schools and libraries.

BOOK REVIEWS 381

KAYE L. MACINNES

Land Resources Division, Department of Indian and North-

ern Affairs, Yellowknife, Northwest Territories X1A 2R3_

The Arctic and Antarctic: their division into geobotanical areas

By V. D. Alexsandrova. 1977. Nauka, Leningrad. English
Translation by Doris Love. 1980. Cambridge University
Press, New York. 247 pp. illus. U.S. $34.50

Plant taxonomists and ecologists who are inter-
ested in circumpolar ecosystems and who have a
limited facility with the Russian language will be
pleased when they learn that V. D. Aleksandrova’s
1977 book has been translated into English. The trans-
lation has been made by Doris Love who is well
known for her work oncircumpolar plant taxonomy-
geography subjects and who is therefore admirably
suited to provide this translation.

The book brings Aleksandrova’s personal expe-
rience, from a career that spans at least 40 years (her
earliest cited publication is 1937), together with the
circumpolar literature. Well over 500 references are
used and, of these over 300 are in the Russian lan-
guage. About 30% of the references predate 1949,
while 10, 30, and 30% of the references were produced
in the 1950s, 1960s, and 1970s, respectively. The book
consists of two short chapters on geobotanical regions
of the Arctic (Chapter 1, 18 pages) and Antarctic
(Chapter 4, 18 pages) plus two chapters that describe
the arctic tundra (Chapter 2, 125 pages) and arctic
polar deserts (Chapter 3, 25 pages) in greater detail.
Alexsandrova divides the Arctic and Antarctic into
hierarchal classification of regions, subregions, pro-
vinces, subprovinces, and districts. Although the sub-
ject is described as geobotanical, the work is primarily
a description of plant species distrubution. There is
very little quantitative information on plant commun-
ity composition, on plant succession or community

response to disturbance, on soils and geology, or on
climatic regimes.

This book is a very subjective approach to the div-
ision of vegetation into units and it is therefore impor-
tant to understand the philosophy of the writer. I
would suggest that the reader begin with Love’s Fore-
word and then proceed through the Preface and Con-
clusions (Chapter 5), before beginning to abosrb the
detailed species information in Chapters | through 4.
Readers who are not familar with the European and
Russian plant community terminology may find some
initial difficulty; however, these terms present few
problems in understanding the work.As very small-
scale maps are provided in the text, I found it useful to
have a somewhat larger-scale map of the circumpolar
region in front of meas I read. Many small geobotani-
cal divisions could have been effectively presented in
photographs or topographical soil-vegetation
profiles.

This work will be very satisfactory to plant taxon-
omists who are interested in species distribution in the
circumpolar area. It will be of limited value, however,
to plant ecologists who are interested in plant eco-
physiology or quantitative plant synecology. The
book serves as a useful introduction to the Arctic of
the Old World and the Antarctic for students who are
initiating studies in polar plant ecology or who may be
preparing for visits to these areas.

Ross W. WEIN

Department of Biology, and Fire Science Centre, University
of New Brunswick, Fredericton, New Brunswick E3B 5A3

Flore des champignons au Québec et régions limitrophes

By René Pomerleau. 1980. Les Editions la Presse, Montréal.
652 pp., illus. $65.

Congratulations to Dr. Pomerleau, who, with the
publication of this tome, has given us the most com-
prehensive text on the mushrooms (a term used here
to include all fungi over about 0.5 cm in size) not only
of Canada but of North America. As the author
admits, the flora is not complete, descriptions of all
the mushrooms of Quebec are not included, but nearly

1400 of the larger species are treated with many more
mentioned in the comparative notes. No one in any
other book has treated such a diverse and large
number of the North American fungi.

The book is well illustrated with 305 species pic-
tured in color. Unfortunately some plates are tinted an
unnatural pink. Line drawings show the shape, the
principal gross features, and some microscopic fea-
tures of 809 species.

382

The book is divided into two principal sections. The
general part of 165 pages lucidly introduces the reader
to the mushrooms. In the chapter on the mushrooms
in the history of man, we are told that the first written
report of intoxication caused by mushrooms
appeared more than 400 years before Christ. Further
that “mushroom” is derived from “mousseron,” the
common name in Normandy for the Tricholoma of
St. George. The role of mushrooms in the origins of
several religions is discussed. Other sections outline
their biology, illustrate and define their characters,
and formulate the systematics section of the book.
Their ecology and uses by man, especially their edibil-
ity or inedibility, are the subjects of other chapters.
The general section is directed at the beginning stu-
dents, who may be confused when they see, as in Plate
1, both microscopic and macroscopic features with no
scale to indicate that haustoria are considerably
smaller than even small mushrooms.

The descriptive part of the book is divided into
chapters each treating similar or allied groups of
fungi. The final sections of the book are a glossary,
seven pages of bibliographic citations and indices of
the French, English, and Latin names. The book is
large, 28.5 X 22 cm, and heavy. 2.8 kg, but the bind-

ENVIRONMENT

THE CANADIAN FIELD-NATURALIST

Vol. 95

ing does not appear to be strong enough to survive
even moderate use.

The order of arrangement of the genera, 1.e., the
systematics, follows the most modern schemes in most
cases. In the family Polyporaceae (the wood rot
fungi), however, a number of generic names last used
in 1928 have been unexplainably resurrected.

This book will be the authoritative work on the
mushrooms for many years, not only of Quebec but
northeastern North America. That this impressive:
flora, which was centered on the southermost region
of Quebec, is now the most detailed flora for Canada
emphasizes our ignorance of the mushroom flora of
Canada. Without knowledge of the species we can
hardly understand their varied role in nature, neither
can we fully use their potential contribution to the
fields of forestry, horticulture, and mycophagy.

This is a magnificent contribution. It will stimulate
interest and research on all aspects of the ecology and
biology of the mushrooms.

J. GINNS

Biosystematics Research Institute, Canada Agriculture,
Ottawa, Ontario KIA 0C6

Bibliography of the Natural History of Newfoundland and Labrador

By M. Laird. 1980. Academic Press, London. 1xxxi + 376
pp. $57.50.

Professor Laird, in this most useful reference book,
has brought together about 4000 published and a few
unpublished references to the natural history of New-
foundland and Labrador. These, in many instances,
are accompanied by a sentence or two outlining the
significance of the publication.

This work, which was begun by the author in 1967
on his arrival at Memorial University, is the result of
much searching of both European and North Ameri-
can literature. It is without question a labor-of-love.
No such work is ever complete, but from the number
of obscure references noted it is obvious that the
author delved deeply and no doubt had much cooper-
ation from other interested individuals. I was sur-
prised, however, to find that there was no reference
made to Bibliography of Canadian Plant Geography
by Adams and Senn which appeared in Transactions
of the Royal Canadian Institute over the years 1930 to
1947 and was continued in Canada Department of
Agriculture Publication 863 (1951). I have noted a
number of references in this series to occurrences of

plant species in Newfoundland and Labrador not
found in the present work.

That Professor Laird is well versed in the natural
history, both past and present, of his area is well
brought out in over 50 introductory pages in which he
touches on many early references to visits by well-
known naturalists such as Banks and Audubonas well
as less tangible hearsay mentions such as “A monster
of the sea.” Here too, he expresses concern regarding
the enormous increase in exploitation of the Grand
Banks since World War II as well as the implications
of importations of non-native species to the island and
unforeseeable consequences related thereto.

The work is made more useful by the inclusion of an
index. It will be a welcome tool for anyone beginning a
study on some phase of the natural history of New-
foundland and Labrador.

WILLIAM J. CODY

Biosystematics Research Institute, Agriculture Canada,
Ottawa, Ontario KIA 0C6

1981

Terrestrial Environments

By J.L. Cloudsley-Thompson. 1979. Halsted Press (Wiley),
New York. 253 pp. U.S. $14.95

Cloudsley-Thompson attempts to write about the
“more important terrestrial habitats of the earth, with
special respect to their influence on the fauna.” He has
“tried to illustrate the selective influence of the envir-
onment on its fauna and, conversely, the influence of
animals on their habitat.” Essentially the book is an
introduction to global terrestrial ecology.

Terrestrial Environments has 15 chapters. The first
chapter is an introduction of zoogeography, a pream-
ble on the theories of continent formation and basic
evolution. Environmental factors, specifically climate
and soil, are reviewed in the second chapter. In chap-
ters 3-10 the important terrestrial habitats: tropical
forest, savannah, desert, steppe, temperate forest,
taiga, tundra and snowlands, and mountains are
discussed. |

Cloudsley-Thompson briefly reviewed the climate
and vegetation of each habitat, but devoted most of
the text to the faunal component. For some reason he
did not review the soil for each habitat, one of the
important environmental factors outlined in Chapter 2.

The book is full of rambling, ambiguous terms that
serve only to confuse the reader. The outstanding
problem lies with the author’s use of nomenclature.
Common names used to describe plant and animal
species differ throughout the world. For example, the
common name for the Moose in Europe is Elk. The
European Elk and the North American Moose are
given the same Latin binomial (Alces alces) because
they are the same species, which is different from the
North American Elk (Cervus elaphus). Cloudsley—
Thompson did not discuss this difference in his book.
To confuse the reader further, he incorrectly listed
some subspecies of North American Moose, “. . .
(Alces americana and A. gigas)” (p. 132), without
explanation. In North America there are four recog-
nized subspecies of Moose (A. alces): A. a. americana,
A. aandersoni, A. a. gigas, and A. a. shirasi (Peterson
1955).

The author banters about terms such as “forest
reindeer,” “tundra deer,” “caribou,” “domesticated
reindeer,” “wild reindeer,” and “deer” without at least
briefly discussing the common name differences (e.g.,
the North American Caribou is the European and
Asian Reindeer equivalent) and the Latin binomials.
In the context of the author’s discussion, all of the
above named animals belong to the species Rangifer
tarandus; however, there are many subspecies.
Cloudsley-Thompson states that “forest reindeer
(Rangifer tarindus) (sic) are larger than tundra deer.”
This is most confusing to the novice reader. In Canada

99 66

99 66

BOOK REVIEWS

383

the “forest reindeer” or Woodland Caribou is R. t.
caribou and the “tundra deer” or Barren-ground
Caribou is R. t. arcticus.

The problem could have been alleviated if
Cloudsley-Thompson had identified the global varia-
tion among common names, followed the correct
procedure for the use of Latin binomials, and used the
most accepted common name(s) in conjunction with
the Latin binomial.

The book is plagued with many incorrect state-
ments. For example, on page 135 it is stated that
“Tundra, a word of Finnish origin, meaning an open
forestless stretch of country, is applied to a high tract
of land north of the Arctic Circle.” The Arctic Circle
lies well north of 65°N, but the tundra in the eastern
Canadian Arctic extends south of 60°N.

On page 138 the author wrote that “mammals of the
tundra have no special protection against the winter.”
This statement is incorrect. Shvarts (1963, cited by
Pruitt 1978) concluded the storage of energy reserves
to be a most important adaptation of northern mam-
mals. He also noted that species best adapted to tun-
dra conditions evolved the ability to maintain a nor-
mal biological function with a lowered metabolism.

In the Chapter entitled “Tundra and Snowlands,” it
is stated that “during the winter, the gound is covered
with a deep layer of snow.” It is true that snow reaches
considerable depth over certain portions of the Rus-
sian tundra (Formozov 1964), but there are areas
where only small amounts of snow fall (e.g., on the
coast, near the Yana River, USSR). In Canada, pre-
cipitation decreases northward; the continental tun-
dra and the High Arctic Islands are virtual deserts.

On page 148 the author stated that “carnivorous
animals nevertheless fare miserably in the Arctic win-
ter and drag out a bare existence. Bears, wolves, and
foxes suffer extremeties of famine; they become very
emaciated and their stomachs empty for long peri-
ods.” I don’t doubt that there are instances of starva-
tion; however, it is not as commonas the author leads
the reader to believe.

Cloudsley-Thompson made some statements that
leave the reader hanging. On page 130 he stated, in
reference to montane and subalpine forest, that “this
type of northern coniferous forest is due in part to
local physiographic factors and, again, differs from
the true taiga.” The author does not tell the reader
how it differs from the taiga.

The remaining chapters deal with: microenviron-
ments, fresh waters, the selective influence of the
organism on its habitat, and ecological regulation. |
fail to see the purpose behind devoting a chapter to
limnology when the book is supposed to dwell on
terrestrial habitats.

384

There are two appendices; they are lists of world
climates and vegetation, and the deserts of the world.
In addition, there is a bibliography, an index to the
authors cited in the text, and a general subject index.
There are ink drawings, tables, and maps throughout.

I recommend this book only to those willing to sort
out the annoying errors throughout the text.

Literature Cited

Formozoy, A. N. 1964. Snow cover as an integral factor of
the environment and its importance in the ecology of
mammals and birds. Boreal Institute, University of
Alberta, Occasional Paper Number |. 176 pp.

Biogeography

By E. C. Pielou. 1979. Wiley, Toronto. 1x + 351 pp., illus.
U.S. $22.50.

This book should be useful to those wanting an
overview of earth’s evolutionary history based upon
past and present distributions of plants and animals.
Most aspects of biogeography are discussed, but
plants are dealt with more often than fishes, amphibi-
ans, or reptiles. The author, taxonomic, and subject
indexes make the text a useful reference.

The book reads more easily than expected for the
subject matter concerned. Continental drift, ice ages,
dispersal, migration, disjunctions, and biogeographic
realms, phylogenies, and genetics, and marine, island,
and ecological biogeography are heavily dealt with in
their own chapters or sections. The subject is properly
introduced in early chapters, and later chapters con-
cern advanced study. The chapter on phylogeny and
geographic spread (Chapter 3) is perhaps the best in
the book and should be read by all persons even
vaguely interested in biogeography. More time is
spent on Canadian biogeography than in other bio-
geography texts.

Some sections require somewhat complex mathe-
matical ability, but the author has intentionally

The Mystery of Migration

Edited by Robin Baker. 1980. Wiley, Toronto. 256 pp., illus.
$29.95.

Migration has puzzled and fascinated man for cen-
turies. Anyone aware of the natural world cannot help
but be struck by the migratory phenomenon. An
introductory student in migration will find some
stunning individual feats: the circumpolar, annual
flight of an Arctic Tern; the transcontinental journey
of a Monarch Butterfly; the reverse-salmon cycle of

THE CANADIAN FIELD-NATURALIST

Vol. 95

Peterson, R. L. 1955. North American Moose. University
of Toronto Press. 280 pp.

Pruitt, W. O. Jr. 1978. Boreal ecology. Edward Arnold
(Publishers) Limited, London. 73 pp.

Shvarts,S.S. 1963. Adaptation of terrestrial vertebrates to
sub-arctic conditions. Academy of Science USSR, Ural
Branch, Institute of Biology. Trudy 33. 132 pp. (cited by
Pruitt 1978.)

PAUL A. GRAY

Wildlife Service, Government of the Northwest Territories,
Yellowknife, Northwest Territories X1A 2L9

arranged these sections so they are easily skipped
without loss of continuity. Even though some of the
mathematical functions are difficult to follow, several
of those dealt with should be used in the near future by
biogeographical researchers.

Typographical errors are numerous. Many referen-
ces are cited in the text but are absent from the biblio-
graphy. Most figures are difficult to interpret because
few labels appear to aid in orientation. Although the
Geological Time Scale is located in a useful place (the
end papers), it is hard to understand as diagramed.

Other items of interest throughout the text include
the following: the percentage of plants with non-lobed
leaves is associated with mean annual temperature (p.
45). Plants evolve slower and have larger geographic
ranges than mammals (p. 49-50). Plants (50 species in
21 families) with branches emerging at right angles to
the main stem are found only in New Zealand (p. 214).
Evidence suggests that congeneric species are most
often found with sympatric ranges (Chapter 7).

RICHARD M. ZAMMUTO

Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7

the European and North American eel; and much
more.

The other particularly evident fact about migration
is that it is practised, in one form or another, by nearly
every living organism. The Mystery of Migration
makes a point of defining migration in its broadest
terms: “migration — movement from one place to
another.” Such a definition opens a new world of
migrants: plants seeds, snails on a beach, plankton in

1981

the ocean, Moose moving up or downa mountainside,
and of course, Man. Homo sapiens is seenasa remark-
able migrant having colonized the world from his
beginnings about 3 000 000 years ago in the grass-
lands of Africa.

The book applies this broad definition to all life
forms from plants through invertebrates, insects, fish,
amphibians, and reptiles, birds, bats, aquatic animals,
land mammals, to man. A comprehensive theory of
migration and some interesting comparisons are devel-
oped. The studies of “lifetime tracks” (the path traced
out in space by an individual between birth and death)
is especially interesting.

Dr. Baker has pulled together a very professional
group of contributors. The writing throughout is clear
and concise and will accommodate either a casual
glance at the text under illustrations, ora more careful
study of the complete text. The book is lavishly illus-
trated with color and black and white photographs.
Excellent additions to the illustrations are the maps,
charts, and graphs which have been produced for the
book. Some of the graphs convey a great deal of

Biology of Communication

By D. Brian Lewis and D. Michael Gower. 1980. Tertiary
Level Biology Series, Halsted Press, John Wiley and Sons,
Toronto and New York. 239 pp., illus. U.S. $27.95.

The title Biology of Communication evokes scenes
of fledglings eliciting food from their parents, and of
insects sending messages to other insects via phero-
mones. While these events are considered in this book,
the subject matter probes far more deeply, into neuro-
physiology and the nature of the media through which
signals travel and which necessarily affect the signals
and their reception. Indeed the final chapter, dealing
with methods of analyzing sequences of behavior by
matrices, information analysis, and multivariate
analysis, will be of interest only to researchers. The
authors say that the previous knowledge assumed
before one reads this work isa basic understanding of
physiology and of the principal concepts in behavior.
Even so, one wonders if the presentation need be as
erudite as it is. An early section begins “In the visual
modality, animals have the capacity for coding inten-
sity, temporal patterning, frequency (colour) and spa-
tial patterning (by line coding) to a sophisticated
level.” For this introduction would it not be simpler to
write that animals can see well, and are able to judge
an object’s relative intensity, motion, color, and
shape?

A general impression one receives from this book is
how much is known about communication, especially

BOOK REVIEWS

385

information in a very concise way.

Not only is there a good index, essential to such a
book, but there are also subjects printed on the top-
right corner of every page. This superb indexing will
make the book very accessible to the student-in-a-
hurry. The contributors are all English, and thus there
is an emphasis on ‘old-world’ creatures. This emphasis
is well placed, however, because most of the pioneer-
ing work in migration was done in Europe and espe-
cially the United Kingdom.

The Mystery of Migration is an excellent book both
for the inquisitive browser and for the more serious
student of migration. While acomprehensive look ata
spectacular natural phenomenon is provided, it seems
fitting that part of the mystery still remains. We have
more to learn from the salmon, the marine turtles, the
terns, the whales, and the worms.

DAVID LOVE

World Wildlife Fund (Canada), Suite 201, 60 St. Clair
Avenue East, Toronto, Ontario M4T IN5

in insects which lend themselves to physiological
experiments, and in birds which have rather stereo-
typed and discrete behavior patterns. On further
reflection, however, when one remembers that there
are perhaps one million species of animals in all, one is
awed at the infinite amounts of knowledge which must
yet be undiscovered. In any case the intricacies of
communication reported here are amazing. In discuss-
ing mimicry, the authors explain that an edible insect
mimicking in color an evil-tasting one cannot become
too abundant relative to the model, because the first
sampling of a predator is likely to shape its future
responses. One insect mimic therefore emerges later in
the season than its model to help solve this problem.
As another example, the female cabbage looper moth
has learned to disperse her male-attracting scent in
relation to the wind velocity, so little is wasted. At low
wind speeds she exposes her pheromone-producing
glandular surfaces for about 20 min, with the expo-
sure time decreasing to 5 min at wind speeds of 3 m
per second.

Much of the information discussed is illustrated in
figures which vary greatly in elaborateness. Some
express simple ideas in straight-forward fashion while
others, taken from primary sources, are almost
incomprehensible to the average reader. The subject
matter ranges equally widely, from how flowers com-
municate to insects, to the functioning of a nerve cell,

386

to the properties of water, to examples of courtship in
butterflies and vertebrates.

A casual naturalist will probably think this book
too academic, but professional ethologists will find
much in it to interest them. They will be able to ponder
statements querying some of the work of Tinbergen
and Lorenz, and assess claims such as that “Hybrids
almost always show reduced ‘fitness’ in some respect.”

THE CANADIAN FIELD-NATURALIST

Vol. 95

They will also enjoy browsing through the nearly 300
references, almost all from the past three decades.

ANNE INNIS DAGG

Integrated Studies, University of Waterloo, Waterloo,
Ontario N2L 3G]

Environmental Effects of Dams and Impoundments in Canada: experience and prospects

By R. M. Baxter and P. Glaude. 1980. Canadian Bulletin of
Fisheries and Aquatic Sciences 205. Supply and Services
Canada, Hull. 34 pp. $2.50 in Canada; $3.00 elsewhere.

This Canadian government bulletin is an extensive
review of the literature in a rapidly developing area.
Using a previous review (Baxter, R. M. 1977, Environ-
mental Effects of Dams and Impoundments. Annual
Review of Ecology and Systematics 8: 255-283) as a
basis, the authors have included more recent studies
and have presented the subject ina Canadian context.
It is written in a comfortable style and is very well
edited; this reviewer found only one error in the text.

For a general readership interested in learning
something about the environmental issues involved in
dam projects, this publication will provide a valuable
introduction. It covers a wide range of potential prob-
lems and benefits, citing Canadian examples where-
ever possible. The caveat urging caution in applying
foreign experience to northern Canadian environ-
ments is well-founded. The table of contents, unusual
in sucha short work, provides a quick guide to specific
topics.

This communication will also be a useful guide to
the literature for the student, scientist, or engineer
who intends to study dam-related problems in depth.

MISCELLANEOUS

It contains 243 references and, fortunately, uses the
author and date-alphabetized citation system rather
than the sequentially numbered reference format used
in many reviews.

While this bulletin succeeds admirably in consider-
ing most conceivable impacts, I was mildly disap-
pointed by the section on climatic effects. The authors
recognize the effect of water bodies on evaporation
and heat budgets, but do not mention the potential
additivity of this effect. Because the total number of
reservoirs can only be expected to increase, the total
surface area covered by water must also increase. It is
therefore possible that impacts which seem only local
when viewed individually may ultimately have
regional or even global climatic consequences at some
point in the future.

As with most literature reviews, this one will
become outdated as more studies appear. As the
authors note, the post impoundment studies on the
James Bay project, when they are completed, will bea
major influence on our thinking.

P. E. Ross

Departement de Sciences biologiques, Universite de Mont-
real, C.P. 6128, Succursale A, Montreal, Quebec H3C 3J7

Morphometrics: the multivariate analysis of biological data

By Richard A. Pimentel. 1979. Kendall/ Hunt, Dubuque,
Iowa. x + 276 pp., illus. U.S. $19.95.

This book is a complete introduction to meaningful
analysis of large data sets from studies in field biology.
It is a useful guide for field biologists interested in
using multivariate biometry without having to con-
sider the in-depth, mathematical terminology used by
most multivariate texts. Mathematical proof of
explained methodology appears as an afterthought

instead of the primary focus it receives in most texts
that propose biological orientation. This fact makes
this text the only one of its kind written by and fora
field biologist. It could serve as a text in a biometry
course subsequent to that by R. R. Sokal and F. J.
Rohlf (1969, Biometry: the principles and practice of
statistics in biological research).

Elementary matrix algebra is explained in such a
way that anyone with algebraic training can under-

1981

stand. The shortcomings of multiple correlation and
regression are discussed. Multivariate analysis of vari-
ance and covariance, principal component, factor,
canonical correlation, ordination, cluster, and dis-
criminant analyses are explained in their respective
chapters. In short, these subjects provide the field
researcher with the most up-to-date statistical tools
known to us for explaining biological observations.

Perhaps most important from the viewpoint of a
researcher wanting to learn about multivariate biome-
try is the 23-page glossary. There are more than 450
terms listed with definitions written in plain, compre-
hensible English.

Shadow of the Hunter — stories of Eskimo life

By Richard K. Nelson. Illustrated by Simon Koonook.
1980. The University of Chicago Press, Chicago, Illinois.
xiii + 282 pp. U.S. $12.50.

“The Eskimos who live in widely scattered settle-
ments along the Arctic Coast and north Slope of
Alaska call themselves Inupiat, which means ‘the real
people.’ In this way they. set themselves apart from,
and above, the rest of humanity. Perhaps, considering
the harshness of the environment in which they live,
they are entitled to this distinction. I hope the chapters
that follow will give you some basis to judge for
yourself.”

With these words Richard Nelson begins to weave a
tale of a year in the lives of the Tareogmuit (sea-
oriented Eskimos) of the fictional village of Ulurunuk.
Weare taken from Siginyasaq Tatqiq — moon of the
returning sun (January), through /rniivik Tatqiq —
the moon when animals give birth (June), to Siginri-
lyaq Tatgiq — the moon with no sun (December).
While on our fascinating journey we experience the
cold loneliness of winter seal hunting; the excitement
of spring whaling; and the danger and adventure of
hunting Walrus and Polar Bear during the times
before snow machines, outboard motors, down
clothing, soda pop, and television. Nelson provides us
with considerable insight into the monumental every-
day logistical effort required of the women, children,
and men to support the more “exciting” activities. He
exposes us to the social interrelationships, competi-
tion, fear, and pride among the people of Ulurunuk.
Shadow of the Hunter is basically a collection of 10
short stories, each one corresponding to a “moon” in
the Tareogmuit year. The stories are factual and excit-
ing. Nelson’s style is relaxed and readable. He is an
astute observer and is able to convey detail in an
interesting manner. Individually, each story can stand
on its own merits, but the stories are linked by the

BOOK REVIEWS

387

Typographical errors are moderate and some pages
seem to have whole sentences missing (eg. pp. 25, 197).
Perhaps the major problem with this book is that a
single data base on Painted Turtles is analyzed and
reanalyzed by various methods throughout the text
which limits the variety of real world examples shown.
Overall, I strongly recommend this text for use by
field biologists.

RICHARD M. ZAMMUTO

Department of Zoology, University of Western Ontario, _
London, Ontario N6A 5B7

changeable environment near Ulurunuk. The book is
relatively free of typographical errors and the type is
attractive. The drawings by Simon Koonook, a Point
Hope Eskimo and student at the University of Alaska,
greatly enhance the book. Both Richard Nelson and
The University of Chicago Press should be com-
mended for their inclusion. I hope that we will see
more of Simon Koonook’s art in the future. One
shortcoming which I felt the book had, although
briefly touched upon in the Appendix, was the lack of
a comparison between Eskimo life as depicted in this
book and present life in the coastal villages of Alaska.
I am afraid that some people will assume that this
book portrays life as it is today. It does not.

As with other books about Eskimos (Brower, Freu-
chen, Stefansson, and others), and based upon my
own experiences in the coastal villages of northern
Alaska, it is always amazing how attuned to their
environment are the Tareogmuit. They are able to
predict changes in weather or sea-ice conditions and
are reacting to these changes before most Janik (non
natives) realize that a change is underway. The feel-
ings and insights portrayed in this book could only be
presented by a person who has experienced what is
written.

Nelson spent 14 months in the village of Wain-
wright on the northwest coast of Alaska studying
“Eskimo methods of hunting, traveling and surviving
on the sea ice.” This Air Force funded project was
designed to gather information for survival manuals
and it produced Nelson’s first book Hunters of the
Northern Ice (The University of Chicago Press, 1969).
Although ethnographic by nature, Shadow of the
Hunter continues where Hunters of the Northern Ice
leaves off by reminding us that we are dealing with
living people and a living culture, people who have
hopes, desires, and frustrations like the rest of us.

Go
oo
oo

Researchers who have written for the general public
have generally been suspected by the scientific com-
munity of being charlatans or at least not sufficiently
serious about science. In addition, scientific publica-
tions are, as arule, not interesting reading toa layper-
son. Consequently most of us fail to find out about
some very interesting research and why we are sup-
porting it with our tax dollars. Fortunately in recent
years several top-level researchers have come forward
to explain or present their research findings in a read-
able form. To this list of scientist-writers we must add
Richard K. Nelson, who is a naturalist of the highest
order as well as an interesting and informative writer.

NEW TITLES

Zoology

Advances in the study of behavior, Volume II. 1980. Edited
by J. S. Rosenblatt, R. A. Hinde, C. Beer, and M.-C. Busnel.
Academic Press, New York. 392 pp. U.S. $35.

Arctic animal ecology. 1980. By Hermann Remmert. Trans-
lated from German by J. Wieser. Springer-Verlag, New
York. 280 pp., illus. U.S. $24.80.

Atlas of benthic shelf Foraminifera of the southwest Atlan-
tic. 1980. Edited by E. Boltovskoy, G. Giussani, S. Wata-
nabe, and R. Wright. Junk, The Hague. vi+ 148 pp., illus.
U.S. $57.90.

The birdwatcher’s A-Z. 1980. By Alan J. Richards. David
and Charles (Canadian distributor McGraw-Hill Ryerson,
Toronto). 336 pp., illus. £12.50.

British coastal shrimps and prawns: keys and notes for the
identification of the species. 1979. By G. Smaldon. Academic
Press, New York. vi+ 126 pp., illus. U.S. $11.50.

+The Canada Goose (Branta canadensis): an annotated bibli-
ography. 1981. Compiled by Scott R. Craven. Special Scien-
tific Report — Wildlife No. 231. United States Department
of the Interior, Washington. 66 pp. Free.

The desert bighorn: its life history, ecology, and manage-
ment. 1980. Edited by Gale Monson and Lowell Sumner.
University of Arizona Press,Tucson. 370 pp. Cloth U.S.
$27.50; paper U.S. $14.95.

Dossier Caribou: écologie et exploitation du Caribou du
Québec—Labrador. 1979. Edité par Francois Turdel et Jean
Huot. Recherches Amérindiennes au Québec, Montréal. 166
pp., illus. $7.50.

Easy identification guide to North American snakes. 1979.
By Hilda Simon. Dodd, Mead, New York. 128 pp., illus.
U.S. $8.95.

THE CANADIAN FIELD-NATURALIST

Vol. 95

In addition to Shadow of the Hunter and Hunters of
the Northern Ice, Nelson has written Hunters of the
Northern Forest. This latter was his second book and
doctoral dissertation. It isa human ecological study of
the Athapaskan speaking Kutchin Indians of east-
central Alaska. I encourage you to read Shadow of the
Hunter and Nelson’s other works, as they will provide
you with many hours of enjoyment and entertainment.

THOMAS J. ELEY

S.R. Box 60977, Fairbanks, Alaska 97701

Entomology. 1980. By Cedric Gillott. Plenum, New York.
675 pp., illus. U.S. $49.50 ($59.50 in Canada); text edition
U.S. $22.50.

Evolution of the vertebrates. 1980. By Edwin H. Colbert. 3rd
edition. Wiley, New York. 510 pp. U.S. $25.

* A field guide to the birds east of the Rockies. 1980. By Roger
Tory Peterson. 4th edition. Houghton, Mifflin (Canadian
distributor Thomas Allen, Markham, Ontario). 384 pp.,
illus. Cloth $19.50; paper $12.95.

*A guide to the birds of Alaska. 1981. By Robert H. Arm-
strong and the editors of Alaska magazine. Alaska North-
west, Anchorage. 320 pp., illus. U.S. $18.95 plus U.S. $1
postage.

Insect photoperiodism. 1980. By Stanley D. Beck. 2nd edi-
tion. Academic Press, New York. 392 pp. U.S. $32.50.

Invertebrate animals: collection and preservation. 1980.
Compiled by Roger J. Lincoln and J. Gordon Sheals. Cam-
bridge University Press, New York. vilit+ 150 pp., illus.
Cloth U.S. $22.50; paper U.S. $6.50.

The island waterfowl. 1980. By Milton W. Weller. lowa State
University Press, Ames. 144 pp., illus. U.S. $10.95.

*Manual of Nearctic Diptera. Volume 1. 1980. Sponsored by
Biosystematics Research Institute. Agriculture Canada,
Ottawa. Monograph 27. 674 pp., illus. $40 in Canada; $48
elsewhere.

Merlins of the Welsh marches. 1980. by D. A. Orton. David
and Charles (Canadian distributor McGraw-Hill Ryerson,
Toronto). 160 pp., illus. £6.95.

*Migrant shorebird ecology with special reference to shore-
bird migration along the north-eastern shore of Lake Onta-
rio. 1980. By Peter William Strahlendorf. Napanee District,
Ontario Ministry of Natural Resources, Napanee. 96 pp.,
illus. $5.

1981]

*The mystery of migration. 1980. Edited by Robin Baker.
Wiley, Toronto. 256 pp. $29.95.

{ Naked species of Gondolella (Conodontophorida): their dis-
tribution, taxonomy, and evolutionary significance. 1980.
By Peter H. von Bitter and Glen K. Merrill. Life Sciences
Contributions 125. Royal Ontario Museum, Toronto. 49
pp., illus. $3.

*North American birds of prey. 1980. by William Marshall.
Gage Publishing, Toronto. 1976 pp., illus. $29.95.

*Les Oiseaux familiers du Québec. 1980. Par Julien Boisclair.
Stanké, Montréal. 184 pp., illus. $24.95.

*The Peregrine Falcon. 1980. by Derek Ratcliffe. Buteo
Books, Vermillion, South Dakota. 416 pp., illus. U.S.
$42.50.

{Principles of fishery science. 1981. By W. Harry Everhart
and William D. Youngs. 2nd edition. Cornell University
Press, Ithaca. 349 pp., illus U.S. $16.50.

The role of animals in biological cycling of forest-steppe
ecosystems. 1980. By R.I. Zlotin and K.S. Kodashova.
Translated by W. Lewus and W. E. Grant. Edited by N. R.
French. Dowden, Hutchinson, and Ross (distributed by
Academic Press, New York). 240 pp. U.S. $19.50.

Rotatoria. 1980. Edited by H. J. Dumont and J. Green.
Proceedings of a symposium. Hydrobiologia volume 73.
Junk, The Hague. xii + 268 pp. U.S. $79.

*Les salmonidés des eaux de la plaine de Montréal. 1. Histo-
rique, 1534-1977. 1980. Par Vianney Legendre, Jean-René
Mongeau, Jean Leclerc, et Jocelyne Brisebois. Rapport
Technique 06-27. Service de aménagement et de l’exploita-
tion de la faune, Québec. 280 pp.

*Les salmondés des eaux de la plaine de Montréal. 2. Biom-
étrie, biogéographie, 1970-1975, et registre de péches,
1941-1976. 1980. Par Vianney Legendre, Jean Leclerc, et
Jocelyne Brisebois. Rapport Technique 06-28. Service de
Paménagement et de l’exploitation de la faune, Québec. 139
Pp.

*Small mammals. 1981. By Robert E. Wrigley. Canadian
Album Series: a coloring adventure in Canadian themes.
Hyperion Press, Winnipeg. 40 pp., illus. $4.95.

Strictly for the chickens. 1980. By Frances Hamerstrom.
lowa State University Press, Ames. 136 pp., illus. U.S.
$11.95.

Time in animal behavior. 1980. By M. Richelle and H.
Lejeune. Pergamon Press, Elmsford, New York. 282 pp.,
illus. Cloth U.S. $50.50; paper U.S. $24.20.

Venomous snakes of the world: a checklist. 1980. By K. A.
Harding and K. R. G. Welch. Pergamon Press, Elmsford,
New York. 200 pp.

BOOK REVIEWS

389

Wildlife management techniques manual. 1980. Edited by
S.D. Schemnitz. 4th edition. The Wildlife Society,
Washington. 722 pp., illus. U.S. $20.

Wolves, bears, and bighorns: wilderness observations and
experiences of a professional outdoorsman. 1980. By JohnS.
Crawford. Alaska Northwest, Edmonds, Washington. 192
pp., illus. Cloth U.S. $23.95; paper U.S. $15.50.

Zoogeography and diversity of plankton. 1979. Edited by S.
van der Spoeland A. C. Pierrot-Bults. Halsted (Wiley), New
York. viii + 410 pp., illus. U.S. $69.95.

Botany

Atlas of the flora of Pennsylvania. 1979. By Edgar T.
Wherry, John M. Fogg, Jr., and Herbert A. Wahl. Univer-
sity of Pennsylvania Morris Arboretum, Philadelphia.
xxx + 390 pp. U.S. $9.95.

*The biology of the bromeliads. 1980. By David H. Benzing.
Mad River Press, Eureka, California. xvit+ 305 pp., illus.
+ plates. U.S. $14.40.

Flora of the Bahamian archipelago. 1981. By D. S. Correll,
J. Cramer, Braunschweig, Germany. c. 1300 pp., illus. c. U.S.
$96.

Flora of Baja California. 1980. By Ira L. Wiggins. Stanford
University Press, Stanford, California. 1025 pp. U.S. $65.

*Living with plants: a guide to practical botany. 1980. By
Donna N. Schumann. Mad River Press, Eureka, California.
xx + 325 pp., illus. U.S. $14.20.

Plant classification. 1979. By Lyman Benson. 2nd edition.
Heath, Lexington, Massachusetts. 901 pp., illus. $26.95.

+Plants of Quetico and the Ontario Shield. 1980. by Shan
Walshe. University of Toronto Press, Toronto. xvit+ 152
pp., illus. Cloth $25; paper $7.95.

And some brought flowers: plants in a new world. 1980. By
Mary Alice Downie and Mary Hamilton. University of
Toronto Press, Toronto. 160 pp., illus. $24.95.

*The vegetation and phytogeography of coastal southwestern
James Bay. 1980. By J. L. Riley and S. M. McKay. Life
Sciences Contributions 124. Royal Ontario Museum,
Toronto. 81 pp., illus. $5.50.

Environment

*The Arctic and Antarctic: their division into geobotanical
areas.1980. By V. D. Alek Sandrova. Translated by Doris
Love. Cambridge University Press, New York. xiii + 274 pp.,
illus. U.S. $34.50.

*The boreal ecosystem. 1980. By James A. Larsen. Academic
Press, New York. 500 pp., illus. U.S. $45.

390

*The Canadian Environment: data book on energy and envi-
ronmental problems. 1980. By Madelyn Webb. Saunders,
Toronto. 166 pp., illus. $9.95.

Cape Cod environmental atlas. 1979. Edited by Arthur H.
Brownlow. Boston University Department of Geology, Bos-
ton. xvit+ 62 pp.+ plates. U.S. $5.

Conservative biology: an evolutionary-ecological perspecti-
ve. 1980. Edited by Michael E. Soule and Bruce A. Wilcox.
Sinauer, Sunderland, Maine. xv + 395 pp., illus. U.S. $14.95.

*Ecology and field biology. 1980. By Robert Leo Smith. 3rd
edition. Harper and Row, New York. xii + 835 pp., illus.
U.S. $18.95.

Ecology: a textbook. 1980. By Hermann Remmert. Trans-
lated from German by M. Biederman-Thorson. Springer-
Verlag, New York. 189 pp., illus. U.S. $21.30.

Ecology of highlands. 1980. By M.S. Mani. Monographiae
Biologica, 40. Junk, The Hague. xiv + 236 pp., illus. U.S.
$58.

Erosion and environment. 1980. By Milos Holy. Pergamon
Press, Elmsford, New York. 266 pp., illus. U.S. $55.

Euphrates and Tigris: Mesopotamian ecology and destiny.
1980. Edited by J. Rzoska. Monographiae Biologicae, 38.
Junk, The Hague. x + 122 pp., illus. U.S. $31.50.

Examination of water for pollution control: a handbook for
managers and analysts. 1981. Edited by M.J. Suess. Per-
gamon Press, Elmsford, New York. 1700 pp., illus. U.S.
$374.

The global predicament: ecological perspectives on world
order. 1979. Edited by David W. Orrand MarvinS. Soroos.
University of North Carolina Press, Chapel Hill. xvi + 398
pp. U.S. $19.

How to save the world: strategy for world conservation.
1980. By Robert Allen. Barnes and Nobles, Totowa, New
Jersey. 150 pp., illus. U.S. $12.95.

Hypertrophic ecosystems. 1980. Edited by L. Mur and J.
Barica. Developments in Hydrobiology, 2. Junk, The
Hague. 330 pp. U.S. $87.

Island ecology. 1979. By M. L. Gorman. Chapman and Hall,
London, 80 pp., illus. U.S. $4.95.

Landscape ecology. 1979. Edited by P. Muller and C.
Rathjens. Junk, The Hague. vit 224 pp., illus. U.S. $52.65.

Limits to action: the allocation of individual behavior. 1980.
Edited by J.E.R. Staddon. Academic Press, New York. 320
pp. U.S. $24.

*The mitigation symposium: a national workshop on mitigat-
ing losses of fish and wildlife habitats. 1980. Co-ordinated by

THE CANADIAN FIELD-NATURALIST

Vol. 95

Gustav A. Swanson. General Technical Report RM-65. U.S.
Department of Agriculture, Fort Collins, Colorado. 685 pp.
Bree:

Neusiedlersee: the limnology of a shallow lake in central
Europe. 1979. Edited by H. Loffler. Monographiae Biologi-
cae, 37. Junk, The Hague. x + 559 pp., illus. U.S. $102.65.

*Overshoot: the ecological basis of revolutionary change.
1980. By William R. Catton, Jr. University of Illinois Press,
Champaign. xvi + 298 pp. U.S. $16.50.

Perspectives on lake ecosystem modeling. 1980. Edited by
Donald Scavia and Andrew Robertson. Ann Arbor Science,
Ann Arbor. 326 pp., illus. U.S. $33.95.

The River Volga and its life. 1979. Edited by P. D.
Mordukhai-Boltovskoi. Monographiae Biologicae 33. Junk,
The Hague. xvi+ 473 pp., illus. U.S. $73.70.

Shallow lakes: contributions to their limnology. 1980. Edited
by M. Dokulil, H. Metz, and D. Jewson. Developments in
Hydrobiology, 3. Junk, The Hague. 218 pp. U.S. $59.50.

Sharing nature with children. 1980. By Joseph Bharat Cor-
nell, Ananda, Nevada City, California. 144 pp., illus. U.S.
$4.95.

+Transuranic elements in the environment. 1980. Edited by
Wayne C. Hanson. DOE/ TIC-22800. United States Depart-
ment of Energy, Oak Ridge, Tennessee. xvii + 728 pp., illus.
U.S. $18.50.

A year in the life of a field. 1981. By Michael Allaby. David
and Charles (Canadian distributor McGraw-Hill Ryerson,
Toronto). 192 pp., illus. £7.95.

Miscellaneous

The art of Canadian nature photography. 1980. Edited by
J. A. Kraulis. Hurtig, Edmonton. 128 pp., illus. $27.50.

+ Autobiography of John Macoun, Canadian explorer and
naturalist, 1831-1920. 1980. 2nd edition. Special Publication
No. |. The Ottawa Field-Naturalists’ Club, Ottawa. 361 pp.
$12.50 plus $2 postage.

Careers in conservation. 1980. By Ada Graham and Frank
Graham. Sierra Club, San Francisco. ix + 166 pp., illus. U.S.
$9.95.

+Dictionary of theoretical concepts in biology. 1981. By
Keith E. Roe and Richard G. Frederick. Scarecrow Press,
Metuchen, New Jersey. 380 pp. U.S. $17.50.

Fossils in the making: vertebrate taxonomy and paleoecol-
ogy. 1980. Edited by Anna K. Behrensmeyer and Andrew P.
Hill. Papers from a conference, Austria, 1976. University of
Chicago Press, Chicago. xii + 338 pp., illus. Cloth U.S. $18;
paper U.S. $7.

1981

Biological sciences at the National Research Council of Can-
ada: the early years to 1952. 1979. By N. T. Gridgeman. Sir
Wilfrid Laurier University Press, Waterloo. xxii + 154 pp.,
illus. $7.50.

Interactions of energy and climate. 1980. Edited by Wilfrid
Bach, Jurgen Pankrath, and Jill Williams. Reidel, Dor-
drecht, Holland. Cloth Dfl. 110; paper Dfl.50.

The kindly fruits of the earth: recollections of an embryo
ecologist. 1979. By G. Evelyn Hutchinson. Yale University
Press, New Haven. xiv + 264 pp., illus. U.S. $18.50.

McGraw-Hill encyclopedia of ocean and atmospheric scien-
ces. 1980. Edited by Sybil P. Parker. McGraw-Hill, New
York. 580 pp., illus. U.S. $34.50.

Practical physics: the production and conservation of
energy. 1980. By Joseph F. Mulligan. McGraw-Hill, New
York. xvii. 526 pp., illus. U.S. $16.95.

Sensory assessment of water quality. 1980. by B.C. J.

BOOK REVIEWS

Sol

Zoeteman. Pergamon Press, Elmstord, New York. 160 pp.,
illus. U.S. $38.

Sourcebook of hydrologic and ecological features: water
resource regions of the conterminous United States. 1980. By
R.M. Cushman, S. B. Gough, M.S. Moran, and R. B.
Craig. Ann Arbor Science, Ann Arbor. 126 pp. U.S. $24.

The stargazer’s bible. 1980. By W.S. Kals. Doubleday,
Garden City, New York. vii + 168 pp., illus. U.S. $3.50.

Stars and planets. 1980. By Robin Kerrod. Arco, New York.
125 pp., illus. U.S. $6.95.

+ Water, wells, and ground water supplies in Ontario. 1980. By
Gunther Funk, William A. McClenaghan, and Cyril Hol-
land. Ontario Ministry of the Environment, Toronto. 95 pp.,
illus. Free.

* Assigned for review
t Available for review

The Ottawa Field-Naturalists’ Club

Minutes of the One Hundred and First Annual Business Meeting of The Ottawa Field-Naturalists’ Club

The 101st Annual Business Meeting of The Ottawa
Field-Naturalists’ Club (OFNC) was held in the audi-
torium of the National Museum of Natural Sciences
on 15 January 1980. President R. Taylor called the
meeting to order at 20:04, a quorum of 54 persons
being present. (The final attendance total of 65 was
assumed to be a reflection of the good weather that
evening!) The Recording Secretary read the minutes
of the 100th Annual Business Meeting, which were
approved on motion by E. Beaubien (2nd, E.
Dickson).

R. Taylor referred to business arising from the min-
utes. On the question of whether the Club needs two
auditors, a motion to amend the Constitution (as
previously published in The Canadian Field-
Naturalist (CFN) in accordance with Article 23 of the
Constitution) was read out by C. Gilliatt and put to
the vote. The motion was carried.

A letter was sent to the Munros after the last
Annual Business Meeting, thanking them for looking
after the Moodie Drive bird feeder for so long. An
advertisement was placed in Trail & Landscape to find
more feeder helpers.

Under ‘Correspondence’ the President read a memo
sent to him by M. Singleton of the Federation of
Ontario Naturalists publicizing the Ontario Natural-
ist wetlands issue and asking for OFNC help in build-
ing an awareness of this problem throughout the
province.

The financial statement was presented by the
Treasurer, B. Henson, who pointed out that the major
difference between 1979 and 1978 was the expenses for
the Centennial projects this year. G. McGee noted
that our assets are extremely large and asked what we
intended to do with them. C. Gilliatt pointed out that
there were still some Centennial projects outstanding
to which we have committed $31,000. The Finance
Committee would be suggesting worthwhile projects
on which to spend our excess funds. A motion to
adopt the 1978-1979 financial statement was moved
by R. Foxall (2nd, H. L. Dickson) and approved.

The President read the Annual Report of Council,
which will be published in The Canadian Field-
Naturalist. Acceptance of the report was moved by
H. L. Dickson (2nd, E. Dickson) and the motion was
carried.

H.L. Dickson, Chairman of the Nominations
Committee, was called upon by The President to pres-
ent the slate of officers and Council for 1980. The
nominations were as follows:

392

President: R. Taylor; Vice-President: H. L. Dick-

son; Treasurer: B. Henson; Recording Secretary:

D. F. Brunton; Corresponding Secretary: F. Pope;

additional members of Council: R. Bedford, F.

Bell, W. J. Cody, E. Dickson, R. A. Foxall, C.

Gilliatt, F. Goodspeed, W. Gummer, P. Hall, J. D.

Lafontaine, D. Laubitz, H. MacKenzie, J. K.

Strang, and K. Taylor.

The proposed slate was approved on a motion by
H. L. Dickson (2nd, J. Reddoch). The President
expressed his thanks to the outgoing Council
members for their work on behalf of the OFNC: Eli-
sabeth Beaubien, ex-chairperson of the Education
and Publicity Committee; Charlie Beddoe, ex-
chairperson of the Excursions and Lectures Commit-
tee during the hectic Centennial year, Jane Proulx
(formerly Diceman), Marc Forget, and Valerie Hume,
ex-Corresponding Secretary. He expressed particular
thanks to Ewen Todd, former OFNC President, who
after many years on Council took on the Chairman-
ship of the Conservation Committee and guided it
during a time when an enormous amount was
achieved on behalf of the club. C. Gilliatt had
expressed the desire to step down as OFNC Vice-
President and the President thanked him very much
for his assistance both as Vice-President and as
chairman of the Finance Committee. D. Laubitz had
felt that three years of being recording secretary was
enough and the President thanked her for her tre-
mendous help in keeping Council meetings to their
agenda.

Gilliatt said that F. M. Brigham had agreed to serve
as Club auditor and therefore moved (2nd, W. J.
Cody) that Brigham be appointed OFNC Auditor for
the 1979-1980 fiscal year (Motion Carried).

It was announced that copies of the Autobiography
of John Macoun will probably be available in about a
month at a cost of about $10. Those interested in being
informed of its publication were asked to put their
names on a list that would be circulated during the
social hour.

The OFNC has been approached by the Nakkertok
Ski Club to see if the Club would be interested in
sharing the use of a large area of land to which Nak-
kertok has access. An excursion to the area has been
arranged for 9 March 1980 and Council would like to
have members’ reactions both to the area and to the
proposal.

The President stated that he would be away from 19
January to 30 March 1980. He also wished to under-

1981

line Hue MacKenzies’s remarks on the hard work of
all who contributed towards our tremendously suc-
cessful Centennial year and the success of the many
projects.

E. Beaubien expressed thanks to R. Taylor, on
behalf of the OFNC membership, for his work as
President during the year.

THE OTTAWA FIELD-NATURALISTS’ CLUB

393

The meeting was adjourned at 21:00 on motion by
H. L. Dickson (2nd, C. Beddoe).

During the social hour slides of various activities
that had taken place during Centennial Year were
shown.

D. LAUBITZ
Recording Secretary

Report of Council to The Ottawa Field-Naturalists’ Club for the Year 1980

Nineteen hundred and eighty was a year of new
initiatives for the OFNC, seeing the establishment of a
new committee, the reorganization of another, and
the reevaluation of our publication policies. The club
appears to be entering a new phase in its history, one
in which the need to streamline operations and
increase organizational efficiency is very important.
The club is at an awkward stage in its development...
almost too large to be managed totally by volunteer
labor and perhaps not large enough to support full-
time staff. Serious discussions on the need for a per-
manent OFNC office were held and initial explora-
tions were conducted and will be continued. The club
presently enjoys an excellent financial condition, due
in large part to careful planning and investments in the
past. Discussion continues on ways to utilize these
accessible funds for the benefit of OFNC objectives
and membership without jeopardizing future finan-
cial security.

Roger Taylor presided over club affairs for a second
term as OFNC President. During his absence from
Canada in February and March, H.L. Dickson,
OFNC Vice-President, and R. A. Foxall, past presi-
dent, adeptly handled presidential duties.

Natural history study groups have been established
(under the coordination of H. L. Dickson) to encour-
age skill development within the membership and to
gather additional life sciences data for the Ottawa
Region. This program, an important part of early
OFNC history but sadly neglected in later years, is
being enthusiastically received by club members.

A thorough review of existing publications policies
and procedures was conducted by the Ad Hoc Com-
mittee on OFNC Publications. The committee tabled
its report to Council late in 1980. That report now
forms the basis for further discussion and evaluation
of publication policies, objectives, and procedures
(and their implementation) within the OFNC.

With the closing of the Ottawa Journal the superla-
tive bird column of the late John Bird (and subse-
quently F. M. Brigham and B. Morin) ceased publica-
tion. The OFNC approached the Ottawa Citizen with
suggestions for an additional, broadly based natural

history column in that newspaper. Initial responses
have been positive.

The OFNC continues to deposit historical club
materials in the National Archives. A large quantity of
bird data as well as materials from several long-time
OFNC members (including the late Herbert Groh)
were deposited this year. Members are encouraged to
send any OFNC material in their possession which
may have historical value to any member of Council;
they will see that it gets to the Archives.

The following committee reports are abstracted
from presentations made to Council by committee
chairpersons (as indicated):

Awards Committee

This committee was established in 1980 to initiate
the nomination of deserving OFNC members for
awards offered by other organizations and to recog-
nize the achievements of members within the club.
This is due to the realization that a tremendous
amount of effort by individual members goes into the
success of OFNC projects and that such contributions
should be recognized. OFNC awards have been estab-
lished in the following categories:

1) Member of the Year Award

2) OFNC Service Award

3) Naturalist Award

4) Conservation Award
The nomination of OFNC Honorary Members will
also fall within the mandate of this committee in the
future.

The committee is still in the development stage but
we are hopeful that it will become fully functional in
1981.

(H. L. Dickson)

Birds Committee

The Birds Committee underwent a major reorgani-
zation in 1980. Formerly the Bird Records Committee
which was sponsored by (but not formerly a commit-
tee of) the OFNC, the Birds Committee is now respon-
sible for all aspects of birding activity within the club.
It includes the Bird Feeder Subcommitee (responsible

394 THE CANADIAN FIELD-NATURALIST

for the maintenance of OFNC bird feeders) and the
Bird Records Subcommittee (responsible for rare bird
reports reviewing and other technical matters). The
overall committee has been expanded to meet the
needs of this broader mandate.

The committee organized the Spring, Fall, and
Christmas Bird Counts in the Region again this year,
largely through the efforts of B. Ladouceur. Over 100
observers participated in the Christmas Count, mak-
ing it one of the largest in Canada. An ad hoc group
has been established to evaluate the whole question of
bird counts within the OFNC and will be reporting
back early in 1981.

Four OFNC bird feeders were maintained in the
Region again this year and were enjoyed by thousands
of hikers, skiers, and birders from the Ottawa area.
Over 635 kg (1400 lb) of seed was consumed by an
appreciative clientele! The National Capital Commis-
sion (Interpretive Section) has kindly agreed to cover
the seed requirement for one of the feeders, thus
reducing the increasingly large financial committment
for feeder supplies.

The Ottawa Rare Bird Alert and Bird Feeder Inven-
tory were both revised this year. The committee is
seriously investigating the possibility of establishing a
phone-in ‘Bird Hot-line’ for the Ottawa Region.

Ten rare bird reports were processed in 1980,
including the (accepted) Scissor-tailed Flycatcher
report which was the first record for the Ottawa-Hull
area. The photo duplicate file of rare bird records was
increased substantially in 1980.

The tremendous effort put into the reorganization
and programs of the Birds Committee by the various
subcommittees and members have really paid off and
1981 should be an extremely successful and produc-
tive year for OFNC birding activities.

(D. F. Brunton)

Centennial Committee

The activities of the committee are winding down
with the completion of the majority of centennial
projects. The Autobiography of John Macoun and
Transactions of The Ottawa Field- Naturalists’ Club
and The Ottawa Naturalist: Index were both pub-
lished in 1980 and have been well received. Two pro-
jects initiated in the Centennial Year (the Naturalists’
Guide to the Ottawa Region and Orchids of Ottawa)
are on-going. The former at least may be ready for
publication late in 1981.

My thanks once more to all of those who have
contributed so much time and effort towards the suc-
cess of the Centennial projects.

(H. MacKenzie)

Conservation Committee
In 1980 the committee found itself devoting a great
deal of time to various Regional and Municipal con-

Vol. 95

cerns. Briefs and/ or plan critiques were presented ona
variety of local topics and to a variety of local authori-
ties, including:

Ottawa—Rideau River Corridors Plan (Regional

Municipality of Ottawa—Carleton)

Gloucester Official Plan (Gloucester Township)

Gatineau Park Master Plan (National Capital

Commission)

Greenbelt Management Plan (National Capital

Commission)

Management of conservation lands (Ontario Min-

istry of Natural Resources)

Bridlewood Buffer Zone (Regional Municipality of

Ottawa-—Carleton)

Carp Hills development (Regional Municipality of

Ottawa-—Carleton)

Mer Bleu preservation (Nature Conservancy of

Canada)

Communications with the appropriate authorities
were also made regarding a number of provincial
and/or national issues, including:

Long-range land use in Southern Ontario (Ontario

Ministry of Natural Resources)

National Grasslands Park (Environment Canada)

Madawaska River Park Reserve (Ontario Ministry

of Natural Resources)

Preservation of Oshawa Second Marsh (Environ-

ment Canada, Ontario Environment)
Polar Bear Pass IBP Site protection (Environment
Canada)

Niagara Escarpment management & planning
(Niagara Escarpment Commission)

Backus Woods preservation (Ontario Ministry of
Natural Resources)

Wetlands conservation (Federation of Ontario
Naturalists)

BIOP Program: arctic oil spill research (Environ-

ment Canada)

Environmental impacts of power line development

(Ontario Hydro)

The implications of conservation areas manage-
ment by the Ontario Ministry of Natural Resources
under the authority of the Forestry Act was a major
concern of the committee in 1980. Although a great
deal of discussion and correspondence with local
OMNR personnel took place, little real progress was
apparent. There is a serious and fundamental differ-
ence in the perception of conservation management
ideals and approaches between OMNR management
staff and that of OFNC personnel, who are less con-
cerned for commercial forest production. This
undoubtedly will remain a major concern of the com-
mittee in 1981.

The OFNC was unable to stop the importation of
Raccoon-dogs from Europe into southern Ontario
but was able to rouse widespread attention to this

1981

potentially devastating situation. The animal is a
serious problem for agriculture and wildlife in the Old
World. Through contact with Tom MacMillan, P.C.
Environment Critic in Parliament, a question on this
matter was brought before the House of Commons. It
is hoped that our efforts will at least see a strengthen-
ing of regulations controlling such potentially dan-
gerous imports in the future.

J. Reddoch of the committee was a main speaker at
the York University conference on Natural Areas in
1980. She discussed the highly successful experience
of the OFNC regarding the establishment of desig-
nated natural areas within the Regional Municipality
of Ottawa—Carleton.

The Committee had little time for field work this
year (we are hopeful of better in 1981) but did manage
to visit the Carp Hills where important life sciences
data were gathered.

We are receiving more and more requests from
various groups and agencies for support of particular
conservation issues. To cope with this, the member-
ship of the committee is increasingly emphasizing
provincial and national expertise, in addition to local
knowledge. It has been a demanding but largely
satisfying year.

I want to thank everyone on the committee for their
support and hard work. Their input has made 1980 a
very busy but successful year for conservation in the
Ottawa Region.

(H. L. Dickson)

Education and Publicity Committee

This was a year for reorganization for the commit-
tee. Only two members from the previous year stayed
on into 1980. With additional volunteers, however,
the committee is now back up to strength.

Early in the year Council was approached for a
reevaluation and clarification of the committee’s func-
tion. After lengthy discussions, it was decided that the
committee should concentrate its efforts on helping
the people of the Ottawa Region to become more
aware of the OFNC and its programs.

Two major displays (at the Billings Estate in June
and at the Ottawa Duck Club annual show in
November) were planned and managed by the com-
mittee and other interested OFNC members. Atten-
dance at both was very good.

Club members selected by the committee selected
winners and presented $100 in prize money for the
best biological and life sciences exhibits at the Ottawa
Regional Science Fair. A subscription to Trail &
Landscape was also awarded to each winner.

The committee arranged for leaders to conduct out-
ings for several groups, including the Ottawa Girl
Guides, Beavers, several school groups, and the

THE OTTAWA FIELD-NATURALISTS’ CLUB

395

Nepean Parks & Recreation Association.

Regular publicity for OFNC activities was estab-
lished in 1980. This included notifying local news-
papers, radio and television stations, and the monthly
magazine What’s On in Ottawa. Because attendance
was fairly good over the past year, increased publicity
was felt to be unnecessary.

Advertizing for the Macoun Autobiography was a
major concern for the committee. With the assistance
of Sally Gray, the committee wrote and mailed 800
fliers to individuals and organizations. Review copies
were also sent out to selected journals.

The committee assisted George McGee in the revi-
sion of the ‘Birds, Botany & Geology’ brochure that is
designed to help visitors to the National Capital. The
National Capital Commission is reprinting the
publication.

The committee also coordinated efforts to market
OFNC sales items. They are now available at all
OFNC meetings and special displays. A special
Christmas pamphlet of OFNC sales items was
included in a mailing of Trail & Landscape.

(P. Hall)

Excursions and Lectures Committee

The committee organized 44 excursions, evening
meetings, and workshops in addition to the nine
monthly meetings of the OFNC, the annual business
meeting, and the annual dinner. Bird outings continue
to be the most frequent form of excursion... 17 inall.
Two of those were long-distance outings (to Presqu’il
Park, Ontario, and Derby Hill, New York). Five other
(general natural history) excursions were also long-
distance efforts requiring bus transportation and
proved to be very popular. The general-type excur-
sions appear to be increasing in popularity and as
such, the relative number of bird outings was reduced
in 1980.

Two new types of outings were held this year...an
overnight excursion to the NCC cabins at Lac
Lapeche in Gatineau Park and a midweek bird hike.
Both were very successful.

Weare pleased to note an increase in monthly meet-
ing attendance (with over 100 in attendance at one
meeting).

My thanks to all those who led outings and/or
provided excellent monthly meeting presentations.
The National Museum of Natural Sciences continues
to be of important assistance through their willingness
to provide meeting space and other facilities. Their
‘Dinobus’ provided valuable transportation on sev-
eral OFNC excursions in 1980 — for which we are
most grateful.

(F. Bell)

396

Finance Committee

Three meetings of the committee were held during
the year.

Proposals were made to Council regarding the sel-
ling price of OFNC Special Publications (Autobi-
ography of John Macoun and Transactions of The
Ottawa Field- Naturalists’ Club and The Ottawa Nat-
uralist: Index). In addition, we determined minimal
financial reserves that would be required by both the
CFN and the OFNC in general. These were estab-
lished as $20,000 and $12,000, respectively. The gen-
eral OFNC reserve figure may be somewhat high and
will be reevaluated.

The committee recommended the establishment of
ad hoc committees to evaluate two possible expendi-
tures of accessible OFNC funds: the establishment of
an OFNC scholarship, and the microfilming of all
back issues of CFN. The latter group will also exam-
ine the worthiness of selling all back numbers of CFN
to a dealer, thus reducing inventory.

A budget was proposed to Council for the 1981
fiscal year in October 1980. It calls for revenues of
$15,200 and expenditures of $14,470, for a budget
surplus of $730. Centennial expenditures amounting
to about $9400 are still outstanding. ;

A new typewriter for the Treasurer’s Assistant
(costing up to $850) was authorized by Council and
will be acquired.

(C. Gilliatt)

Macoun Club Committee

The Macoun Club continues to be important for
enthusiastic young naturalists in the Ottawa Region.
Most of the credit for the club’s success this year must
be given to the youngsters themselves. Susan Hulley,
Alex Hubbard, and Ian Bell served as Presidents for
the Senior, Intermediate, and Junior groups, respec-
tively. Arnet Shepard continued as Club Chairman in
1980 and is due thanks for his important efforts.
Through the year he was assisted by Elisabeth Beau-
bien, Fazal Mohammed, Andrew MacFarlane, and
Michael Rankin.

Canadian

Membership Canadian (other

(local) areas)
Individual 460 (455) 337 (339)
Family 233 (219) 26 (29)
Sustaining 15 (15) 3 (1)
Life 13 (12) 14 (12)
Honorary 9 (6) 5 (4)
Total 730 (707) 385 (385)
Change +23 2S

Figures in parenthesis represent 1979

THE CANADIAN FIELD-NATURALIST

Vol. 95

The second annual Macoun Symposium was held
last March, concerning such topics as carcinogens,
pesticides, nuclear wastes, garage and pulp and paper
industry pollution.

Weekend outings were characterized by rainy
weather, but this did not dampen the enthusiasm of
the participants. Many of the outings were to the
Club’s study area near Moodie Drive or to the Conroy
Road Nature Trail. Other trips were held too, includ-
ing a tour of the Carleton University greenhouses and
a trip to the Adirondack Mountains of northern New
York State.

Projects this year included the overhauling of the
library card catalogue, the establishment of a club
herbarium, and continued investigations in the study
area.

Rod Story was awarded the 1980 Baldwin Scholar-
ship and the $150 prize that goes with it.

A meeting between OFNC and National Museum
of Canada personnel was held in November to discuss
the level of Museum participation in Macoun Club
activities. It was agreed that the Club was a worth-
while venture and that Museum staff should become
more directly involved; this involvement should be
formalized.

With the continued support of the National
Museum of Natural Sciences and the direction
received from the OFNC, the Club should continue its
successful program through the 1980s.

(K. Taylor)

Membership Committee

On | December 1980, OFNC records listed 1220
members, compared with 1198 in December 1979.
This modest increase continues the trend of recent
years.

This year the OFNC was proud to elect four new
Honorary Members:

Clarence Frankton, Ottawa, Ontario

Yorke Edwards, Victoria, British Columbia

Mary E. Stuart, Ottawa, Ontario

Douglas B. O. Savile, Ottawa, Ontario.

USA Foreign Total
93 (96) 4 (3) 894 (893)
1 (1) 4 (1) 261 (250)

— — 18 (16)

3 (3) 2 (2) 32 (29)
1(-) = 15 (10)
98 (100) 7 (6) 1220 (1198)

-2 +] 22

1981

We thank the more than 60 members who offered
their services by filling in the questionnaire on the
back of the 1979 membership renewal form. We pro-
pose to have this (and other volunteer information)
coded into our computer file in 1981 so that commit-
tees seeking volunteers can have up-to-date inform-
ation.

I thank all committee members for their willing
efforts, with a special thank you to Ellaine Dickson,
who voluntarily devotes so much time and energy to
maintaining OFNC membership records and ensuring
that our computer file is up to date... . along with her
many other Club interests and responsibilities.

(F. Goodspeed)

Publications Committee

Three publications fall within the responsibilities of
the committee; The Canadian Field- Naturalist, Trail
& Landscape, and The Shrike.

The Canadian Field-Naturalist was published in
four issues comprizing Volume 94, Issues | to 4, witha
total of 43 articles, 40 book reviews, and 44 notes. A
listing of 285 new titles related to the interests of the
readership was also provided. We were able to publish
this year without financial assistance from a scientific
publications grant, primarily due to a higher rate of
payment of page charges by contributors.

Trail & Landscape was published as Volume 14,
Issues | to 4, containing Club news and articles related
to natural history and conservation in the Ottawa
Region. A total of 184 pages of material was
published.

THE OTTAWA FIELD-NATURALISTS’ CLUB

397

The Shrike contains seasonal summaries of bird
sightings in the Ottawa Region and news of interest to
local birders. It has gone through a difficult period in
1980 but is now back on track and will be on a solid
footing in 1981.

Both the Autobiography of John Macoun and
Transactions of The Ottawa Field- Naturalists’ Club
and The Ottawa Naturalist: Index were made availa-
ble in 1980. They mark the inauguration of the “Spe-
cial Publications” of the OFNC. This designation will
be applied to future publications sponsored by the
Club when they are of significant value.

(J. K. Strang)

By the end of 1981 we should be reporting that
many of the ‘loose ends’ identified above have been
wrapped up. The OFNC is moving steadily towards a
more stable and secure status, both financially and
organizationally, while at the same timé enhancing the
field-naturalist orientation that has been our strength
and tradition for over a century. It has been a produc-
tive, exciting, and (at times) controversial year in
OFNC history. The momentum established in 1980 is
truly an appropriate and satisfying start to the new
decade.

DANIEL F. BRUNTON

January 1981 Recording Secretary

Auditor’s Report

To: Members of The Ottawa Field-Naturalists’ Club

I have examined the balance sheet of The Ottawa Field-Naturalists’ Club as at September 30, 1980, and the
related Income Statements for the year then ended. My examination included a general review of the accounting
procedures and such tests of the records and supporting vouchers as considered necessary under the

circumstances.

In my opinion, these financial statements present fairly the financial position of the organization as at
September 30, 1980, and the results of its operations for the year then ended in accordance with generally

accepted accounting principles.

December 3, 1980

(Signed) F. Montgomery Brigham

398 THE CANADIAN FIELD-NATURALIST

The Ottawa Field-Naturalists’ Club Balance Sheet

as of September 30, 1980

Assets

Current
Gashvand:tenmudepOsitsias vc cms tices ise
/NECOWMS RACANAIDIS cooobobooodcnboccca00b008
INCCHUWE UmINTE GES teare a ek noe eey coeiahe erento meas eicensee
PrepardeexPensest vse ccramin alte ony caer ss fera's 2/5

Fixed at cost
FSGQUIEPINEN egies er pee aN teh okseae Veneer aoe
Bessraccumulatedidepreciationuer. aces ioe

Total Assets

Current liabilities
FANCCOUNTSEPAV AID Caw ie ore cues oesteretepogoutrovetel shesesinipsicdedons
Deternedwincomes oss wk ke eee

Memorial funds
Bal iwi etches he te ee ree eee eee

Surplus
Balance: Octoben limi eae cre aeere
Add: income over expenditure
The Ottawa Field-Naturalists’ Club ......... $3,913.17
The Canadian Field-Naturalist ............. 5,281.53
9,194.70

Less: net expenditure — Centennial projects .... 15,413.66
Balance;septemberes 01980) emrnieew teers

Total liabilities and surplus ....... joooeodououeus

$49,250.62
10,693.69
1,448.51
650.00

529.50
469.00

SIS Se7S
7,413.00

197.50
50.00

$58,902.99

6,218.96

Vol. 95

$62,042.82
60.50
$62,103.32
SOM wale

247.50
$9,419.29

52,684.03
$62,103.32

1981 THE OTTAWA FIELD-NATURALISTS’ CLUB

The Ottawa Field-Naturalists’ Club
Statement of Income and Expenditure

for the year ended September 30, 1980

Income

Apportionment of membership fees
eNTRITUEN = S dos 8 Sie Grate ce Caticec rerteetanah eS eeraDk neces eo
[LINO Sebel nee ae acer oe eee ee ee rer oR

Trial & Landscape
SUIDSCHIPCIOMS wrasse s cite e ncie eee chats
BackenWinbers: vc.ceks eerste ices ace comune tage wiaeete

Shrike subscriptions
DONE OMS eg ee: Ae ccoweyn etre Veteierer cake uaa Momence ene

NWETRERE oS ce ee eR ct pnelrte Neeeen a ater eet

Expenditure

Trail & Landscape
| PUL DIT oan, Ai ae eal et WA seamen sath ie
inc WIALION Sal wee ee heh Ue etae «meee anes
Baditimesandi Once: fm thee to ae ue eee he ee
LOMO TAT TAP ev. oy seen es Seve tas Slane eM ees

SHrike publishing and expenses 2.) ...:.54--.--

Committee activities — net
SXCuRSTONS#andulectunesy sence ote eee
INMEmIDETS pp te Fak cust erscite cies ceeds ues
Niacoung@lulbiese at oct ase ctas Bk ore taaeeaeron tes
GOnSeivatlOMe = ts tes ey holes nes eae aes
BiGdQReC ond Sie -asaiey scene atelier Ce
Bindeeeedersmerecas crea eae sister
AUC i toe veces eres eseeetauclete seh mates Mena
ORCHIGESUERVEW tri ory comers wes atv acne evabeneeeeoee

Baldwins Scholarship) sees enxcimayere ect tees
SPECIAIPACUIVIDIES | Sere erty .c ate teteuaa., Sete sys cas seee oe
WOTINCIEEXPEMSES ook eines ie ele erase tees
OihiceryASSiStante ce ciirosesaweue oat Rune conver pamrs moes
Ofmcer Sup mlieSenoeae crest toe mens eae he Serena
WIS CINE OUS Ms cw youats sees eines shemsLe Aoaenerels

$7,524.00

234.00

370.00
102.00

3,402.55
221.74
WA) AAI
415.00

(89.79)
823.65
149.74
28.85
5.54
628.69
17.00
55.76

150.00
258.22
368.41
300.00
607.37

98.94

399

$7,758.00

472.00

490.00
550.55

2,549.51 $11,820.06

$4,168.50
336.01

1,619.44

1,782.94 7,906.89
SSO aly

400 THE CANADIAN FIELD-NATURALIST

The Ottawa Field-Naturalists’ Club
Statement of Centennial Project Expenditures

for the year ended September 30, 1980

Recording — Songs of the seasons 1979-1980

REASON OUST Reasons Gh Stele mi ciateliay ak ciethcocr a Amer rates Ne $7,887.45

FSC MIG aera acre oN tree aul rare toe SUE ot ae 6,966.76 $ (920.59)
Club pin

FREVS TUL Barerens coe re rice creri spent rage a te canteen pee eae 207.20

JS <foXSUVOlRA WU eN Ae ie eaa rman athd wea ene iala LNs porn aren ere 1,188.36 981.16
Bird record cards

NEN ETI Cero keyg: ieee e tec eral etc daret mist tins apenas 169.75

| SbeoyevralGAASeitiass orto, oly ONG ORT eure ee OO 405.95 236.20
Hasti-notes

| ACAUSIMDIS Ay ales geal ie enero a eater a om sige oe aia tes 61.60

Ee XMEMGITUTES a terse mus errs ie et ire ap aurtmranara 89.07 27.47
Macoun autobiography reprint

REV CMC Reerasiicus se tne sis cites as RNOMR eai Mo cerete ahs 2,700.00

| ES OXSTaKaU IRS i ah tee sea Ree aN a mca h See RS 10,079.50 ° S250
CFN Index

| RENUS TOT a sees fess hc RSPR eB tn Rie rs 225.00

Fc PEM AIG e gerne tera aoe ees Jenny ents UA OMtA aiainatal ads 10,484.50 10,259.50
Club T-shirts

I(EN TID calesen eas Nga eicet aa, Cate ee 10.00

Exe emGitune es -zie mu eusiae fae. cea we sederaee eta ope hates 185.90 175.90
Orehidsyorthe Ottawa District °<. ene es eee 1,072.89
EROVects complete dain O 79) ares aes ee eee eons 2,589.49

IEESS*COMALIO NS teers st citiate a ene eemporea meer saat S2ier2 L7Ol 77
INEtE XE MACUL! eres can cen enced oe Pan. oieas wane aires $20,973.80

1980

$3,861.91
1,047.64

47.60

91.00

2.80

2,700.00
9,561.04

225.00
10,484.50

10.00
185.50

Vol. 95

$(2,814.27)

(47.60)

(91.00)

(2.80)

6,861.04

10,259.50

175.90
1,072.89

$15,413.66

1981

THE OTTAWA FIELD-NATURALISTS’ CLUB

The Ottawa Field-Naturalists’ Club

Statement of Income and Expenditure — The Canadian Field-Naturalist

for the year ended September 30, 1980

Income

Apportionment of membership fees

Annual
Life .

Subscripti

Ce

CC

OMS cocecocssgdpocco00bdd0ooDBDC008

Publication

Reprint

BS cdd0odod0b000 FO KC Od 00 G00 CODD O00

PAIES AnGh alo) SCUNNE Sagoo o bea e oon see eos

Extra P

BUSS.” "eileen taka esees ores eae ene emule es

BackeNuimbensic otc tac cis ace sleh eae

Other — i

MNIVEMES ean tars este ta Nerang a che te acva.e Gos emceees

XC AMC Meera eepee olor ere oc Nanas) eae a kes

Expenditure

FUSS nS ierewnes ie rok ye tect eect a ey ge Ee SAN 0S

Reprints
Circulatio

ee eee ec eee ee ee we ee we ee ee ee we wwe eee eee

IM oodoqddcvocoD oC bN UH HOOOUDU DOO DOO MOOS

Gitinnerandwe xpPenSesiy nec Sere ete We: Becky vee
CRIES ASSISUZINTIEN era ratote oS Belo erence eadialigee Seman ase

Postage

Ce

CHEMES SW/Ofo) See neoted chores a oe eee eer ec eis
UO IN@ Hela Agee estes ees es ate koa oie Te ceed real

$5,014.00
156.00
5,170.00

15,421.40

6,120.60
2,618.00
2 SEZ)

616.49

5,892.56
722.09

$28,792.67
4,292.96

$20,591.40

AON AeAdS)

6,614.65

$33,085.63
4,770.71
1,187.30
1,890.00
688.15
401.02
1,473.00
41.00

401

$48,818.34

$43,536.81
N) pASIIo3)

TABLE OF CONTENTS (concluded)

Book Reviews

Zoology: Birds of Pacific Rim National Park — The Alaskan bird sketches of Olaus Murie with 372
excerpts from his field notes — A bird-finding guide to the Toronto region — The Black Bear in
North America — The freshwater fishes of Alaska — North American birds of prey — Wolf
and man: evolution in parallel — The squirrels of Canada

Botany: The flora of Canada — Multivariate analysis in vegetation research — Vascular plants of 378
continental Northwest Territories, Canada — The Arctic and Antarctic: their division into
geobotanical areas — Flore des champignons au Québec et régions limitrophes

Environment: Bibliography of the natural history of Newfoundland and Labrador — Terrestrial 382
environments — Biogeography — The mystery of migration — Biology of communication —
Environmental effects of dams and impoundments in Canada: experience and prospects

Miscellaneous: Morphometrics: the multivariate analysis of biological data — Shadow of the Hunter- 386
stories of Eskimo life

New Titles 388

The Ottawa Field-Naturalists’ Club 392

Mailing date of previous issue 23 April 1981

The Ottawa Field- Naturalists’ Club

Honorary Members

William J. Cody
William G. Dore
R. Yorke Edwards
Clarence Frankton
W. Earl Godfrey

President: R. Taylor
Vice-President: D.F. Brunton

Recording Secretary: E.F. Pope
Corresponding Secretary: W.K. Gummer

Treasurer: B.C. Henson

Louise de K. Lawrence
George H. McGee
Hugh M. Raup

Loris S. Russell
Douglas B.O. Savile

1981 Council

R.E.Bedford
F.H. Bell
D.R. Bewley
P.M. Catling
W.J. Cody

S. Darbyshire
E. Dickson
S. Gawn

Pauline Snure

J. Dewey Soper
Charles M. Sternberg
Mary E. Stuart
Sheila Thomson
Robie W. Tufts

C.S. Gilliatt

F.E. Goodspeed
J.A. Jackson

D. Laubitz

A. Mason

R.C. Montgomery
J.K. Strang

K. Taylor

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.

TABLE OF CONTENTS (continued)

Articles (continued)
Breeding, feeding, and chick growth of the Black Guillemot (Cepphus grylle)
in southern Quebec DAVID CAIRNS

Food habits of Deer Mice (Peromyscus maniculatus) in northern Ontario
ARTHUR M. MARTELL and ANN L. MACAULAY

Food habits of Southern Red-backed Voles (Clethrionomys gapperi) in
northern Ontario ARTHUR M. MARTELL

The Biological Flora of Canada

2. Vaccinium myrtilloides Michx., Velvet-leaf Blueberry
S. P. VANDER KLOET and I. V. HALL

Notes

Attempted avian predation by a Canadian Toad, Bufo americanus hemiophrys
FRANCIS R. COOK and JOYCE C. COOK

Daily movements of Red Squirrels, Tamiasciurus hudsonicus STEVE G. FANCY

Fate of overwintered clutches of the Common Snapping Turtle (Che/ydra serpentina)
in Algonquin Park, Ontario MARTIN E. OBBARD and RONALD J. BROOKS

Predation and caching of seabirds by Red Foxes (Vulpes vulpes) on Baccalieu
Island, Newfoundland A. D. MACCARONE and W. A. MONTEVECCHI

Twinning and postpartum activity in Barren-ground Caribou (Rangifer tarandus)
E. JANET MCDONALD and ARTHUR M. MARTELL

Seasonal pelage change of Marten (Martes a. americana) in Maine
EDWARD C. SOUTIERE and J. DOUGLAS STEVENTON

Marsh nesting of Common Loons (Gavia immer) ROBERT ALVO
Evidence of Arctic Grayling (Thymallus arcticus) spawning in a highway culvert Pa P SK RAI

Freshwater fishes from northern Newfoundland
PETER J. ROMBOUGH, STEPHEN H. BARBOUR, and JOSEPH J. KEREKES

Sightings of the Hop Merchant (Po/ygonia comma) butterfly on insular Newfoundland
BERNARD S. JACKSON

Application of the Varrio Snow Index to overwintering North American Barren-
ground Caribou (Rangifer tarandus arcticus) WILLIAM O. PRUITT, JR.

First Canadian record of Bering Cisco (Coregonus laurettae) from the Yukon River
at Dawson, Yukon Territory DIRK A. DEGRAAF

Reports of Significant Range Extensions

Sharp-shinned Hawk, Accipiter striatus (Accipitriformes: Accipitridae) on

Victoria Island, Northwest Territories THOMAS G. SMITH
Two-lined Salamander, Eurycea bislineata (Amphibia: Caudata: Plethodontidae,
in Labrador DIRK A. DEGRAAF, BRUCE K. BOLES, and JAMES A. LOVISEK

News and Comment

312

319

325

B29

346
348

350

352

354

356
39)
358

859)

361

363

365

366

367

368

concluded on facing page

THE CANADIAN FIELD-NATURALIST Volume 95, Number 3 1981

Editorship of The Canadian Field- Naturalist
Editor’s thanks

Editorial
Editor’s farewell musings LORRAINE C. SMITH

Viewpoint

Is the Impact Factor a meaningful index for the ranking of scientific research journals?
ROGER TAYLOR

Articles

Vegetation with Atlantic coastal plain affinities in Axe Lake, near
Georgian Bay, Ontario P. A. KEDDY

Low DDT residues in plasma of Bald Eagles (Haliaeetus leucocephalus)
wintering in Colorado and Missouri
CHARLES J. HENNY, CURTICE R. GRIFFIN, DALE W. STAHLECKER,
ALAN R. HARMATA, and EUGENE CROMARTIE

Changes in small mammal populations following clear-cutting
in upper Michigan conifer swamps Louis J. VERME and JOHN J. OZOGA

Nest-tree sharing by herons and cormorants in Montana LARRY S. THOMPSON

Occurrence and growth patterns of the American Brook Lamprey,
Lethenteron lamottenii, in the Ottawa River
J. LANTEIGNE, J. M. HANSON, and S. U. QADRI

A time-activity budget for breeding Mallards (Anas platyrhynchos)
in Manitoba RODGER D. TITMAN

Invasion of a new reservoir by fishes: species composition, growth,
and condition ROBIN MAHON and MARK FERGUSON

Distribution and harvest of Canada Geese (Branta canadensis) in
southern Manitoba prior to development of Oak Hammock Marsh
DENNIS G. RAVELING and CHARLES C. DIXON

Hill’s Oak (Quercus ellipsoidalis) in southern Ontario P. W. BALL
Seasonal and daily use of plant communities by Sharp-tailed Grouse

(Pedioecetes phasianellus) in the parklands of Alberta D. L. J. MOYLES
Movements and haulout behavior of radio-tagged Harbor Seals,

Phoca vitulina KENNETH W. PITCHER and DENNIS C. MCALLISTER

Reproduction, distribution, and population size of Largemouth Bass,
Micropterus salmoides, in an oligotrophic Precambrian Shield lake
DAVID C. MARALDO and HUGH R. MACCRIMMON

Forestry management practices and populations of breeding birds
in a hardwood forest in Nova Scotia
B. FREEDMAN, C. BEAUCHAMP, I. A. MCLAREN, and S. I. TINGLEY

231
232

233

236

241

249

258

1S) i)

261

266

272

276

281

287

292

298

307

continued on inside back cover

ISSN 0008-3550

Ge. AN us, COMP. ZOCL:

“3 ARY

* The CANADIAN Ws &

‘\RVA RR
4

FIELD-NATURALIST*™

Published by THE OTTAWA FIELD-NATURALISTS’ CLUB, Ottawa, Canada

Volume 95, Number 4 October-December 1981

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.

Honorary Members
William J. Cody Clarence Frankton George H. McGee Douglas B.O. Savile Mary E. Stuart
William G. Dore W. Earl Godfrey Hugh M. Raup Pauline Snure Sheila Thomson
R. Yorke Edwards Louise de K. Lawrence Loris S. Russell J. Dewey Soper Robie W. Tufts

1981 Council

President: R. Taylor R.E. Bedford S. Darbyshire D. Laubitz
Vice-President: D.F. Brunton F.H. Bell E. Dickson A. Mason
Recording Secretary: E.F. Pope D.R. Bewley S. Gawn R.C. Montgomery
Corresponding Secretary: W.K. Gummer P.M. Catling C.S. Gilliatt J.K. Strang
Treasurer: B.C. Henson W.J. Cody F.E. Goodspeed K. Taylor

F.R. Cook J.A. Jackson

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) 733-3050.

The Canadian Field-Naturalist

The Canadian Field- Naturalist is published quarterly by The Ottawa Field-Naturalists’ Club. Opinions and ideas
expressed in this journal do not necessarily reflect those of The Ottawa Field-Naturalists’ Club or any other agency.

Editor: Francis R. Cook, Herpetology Section, National Museum of Natural Sciences, Ottawa, Ontario
KIA 0M8

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

Associate Editors:

C. D. Bird A. J. Erskine Charles J. Krebs Stephen M. Smith
E. L. Bousfield Charles Jonkel W. O. Pruitt, Jr.

Chairman, Publications Committee: R. E. Bedford

All manuscripts intended for publication should be addressed to the Editor.

Urgent telephone calls may be made to the Editor’s office (613-996-1755), or his home on evenings and weekends (61 3-269-
3211), or to the Business Manager’s office (613-996-1665).

Subscriptions and Membership
Subscription rates for individuals are $13 per calendar year. Libraries and other institutions may subscribe at the
rate of $25 per year (volume). The Ottawa Field-Naturalists’ Club annual membership fee of $13 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.

Cover: Smith’s Longspur, Calcarius pictus, female photographed at nest in the Chilkat Pass area, British Columbia, by Earvio
Siam (British Columbia Provincial Museum Photo No. 661). See Note by Martin, Hannon and Moses p. 469.

The Canadian Field-Naturalist

Volume 95, Number 4 October-December 1981

Announcement

The Ottawa Field-Naturalists’ Club is pleased to announce the appointment of Dr.
Francis R. Cook as Editor of The Canadian Field-Naturalist starting with this issue.
Dr. Cook, a former editor (1962-1966) and associate editor (1972-1981) of The
Canadian Field-Naturalist, is Curator, in the Herpetology Section, at the National
Museum of Natural Sciences (1960-). The appointment was made after a lengthy
search involving consultation with many natural history experts across Canada. We
are confident that-under Dr. Cook’s editorship The Canadian Field-Naturalist will
continue to maintain its high standards.

In 1979 anad hoc committee was appointed to examine and to make recommenda-
tions concerning all aspects of existing Ottawa Field-Naturalists’ Club periodicals; in
particular The Canadian Field- Naturalist, Trailand Landscape, and The Shrike. The
Executive Committee of the Council is preparing a detailed publication policy of the
Ottawa Field-Naturalists’ Club based on the ad hoc committee report and its subse-
quent reviews. When ready, it will be published in The Canadian Field- Naturalist.
Readers can be assured that the Club will continue to publish The Canadian Field-
Naturalist and to provide the support necessary to maintain the present standard of
excellence.

ROGER TAYLOR
President, The Ottawa Field-Naturalists’ Club

R. E. BEDFORD
Chairman, Publications Committee

403

Grazing Intensity Effects on the Breeding Avifauna of

North Dakota Native Grasslands

H. A. KANTRUD

Northern Prairie Wildlife Research Center, P.O. Box 1747, Jamestown, North Dakota 58401

Kantrud, H. A. 1981. Grazing intensity effects on the breeding avifauna of North Dakota native grasslands. Canadian

Field-Naturalist 95(4): 404-417.

A breeding bird census and plant survey was conducted on 180 samples of lightly, moderately, and heavily grazed and hayed
native grasslands in North Dakota in 1974. The ten most important cover plants on each of eight major physiographic
landforms in three of the four regions (the Agassiz Lake Plain excluded) overlapped so extensively that only 19 species were
involved: 13 grasses or sedges, four forbs, one shrub, and one clubmoss. Bird densities were generally highest in (i) regions and
landforms containing numerous natural basin wetlands, (ii) flatter, glaciated landforms containing more fertile soils, and (iii)
landforms of greater relief and high habitat heterogeneity. Avian species richness tended to decrease with increased grazing
intensity, but total bird density increased due to higher populations of a few species, and hayland that had been mowed and
raked during the previous growing season was highly attractive to some species.

Key Words: Grassland birds, North Dakota, landforms, soils, vegetation, grazing, mowing.

Grasslands in North Dakota have evolved under
varying climatic, geomorphologic, and edaphic condi-
tions. There are differences in the type and genesis of
the various physiographic landforms created by Pleis-
tocene glaciation. Studies of the avian communities
on different glacial landforms on the Northern Great
Plains and Interior Lowlands have not been under-
taken. This study was initiated because of the rapid
destruction of native grasslands in North Dakota, the
extent of which varies greatly by landform. For con-
servationists, it is important to know the general floral
and faunal composition of the remaining grassland
areas before protective measures are undertaken. In
this report, I describe differences in the density and
species richness of breeding birds on variously grazed
native grasslands throughout the state of North
Dakota.

The surface of North Dakota generally slopes
downward, from 1069 m above sea level in the
southwestern corner of the state to 230 m above sea
level in the northeast corner. Deposits of sediment and
till related to Wisconsinan glaciation cover about 75%
of the state lying north and east of the Missouri River;
earlier glaciations covered an additional 12% of the
state but their deposits have nearly been eliminated by
subsequent erosion. The Missouri Coteau divides the
state, with the Central Lowland lying east of it and the
Great Plains lying to the west of it. Additional details
of state geology may be found in Bluemle (1977).

North Dakota has been classified into various
regions and subregions based on physiography and
the biota (Stewart and Kantrud 1972, 1973; Stewart
1975). I recognize four regions in this report (Figure
1). The 13 landforms on which samples were located

404

are based on Bluemle (1977, and personal communi-
cation), and are described in Appendix A.

Methods
Selection of Sample Units

One hundred and eighty samples of uncultivated
native grassland were selected (Figure 1), in numbers
roughly proportional to the areas of the several physi-
ographic landforms (Table 1) within the four major
physiographic regions of the state. Some landforms
occur in more than one region. There were 21 combi-
nations of region and landform.

Sample site locations were initially selected at ran-
dom from numbered legal townships, sections, and
quarter-sections within each of the 21 strata. How-
ever, if a selected area was devoid of native grassland,
the nearest example on the desired landform was
chosen. Many sites were rejected because field exami-
nation revealed old furrows, rockpiles, domestic grass
plantings, or other evidence of past disturbance by
cultivation. Although most of the 180 sites were legal
quarter-sections (65 ha, 160 acre), several as large asa
section (259 ha) oras smallas 16 ha were also utilized.
Most sites were square but occasionally it was neces-
sary to utilize odd-shaped areas.

Bird Censuses

Field work was conducted by two teams, each con-
sisting of a botanist and an ornithologist. The breed-
ing bird census began 22 May and ended 19 July 1974.
This period includes the peak nesting season for
nearly all species studied (Stewart 1975). Bird cen-
suses were conducted from sunrise to sunset when
temperatures did not exceed 32°C, precipitation did

1981

e - SOUTHWESTERN
en SHORE

KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA

405

FiGureE |. Major physiographic regions of North Dakota showing location of sample plots. Physiographic landforms cannot
be shown at this scale because of their extremely complex pattern and small size.

not occur, and sustained wind velocities did not
exceed 24 km/h.

Jarvinen etal. (1976) found no temporal changes in
bird observability on transects in open fields and
meadows. Comparison of five 3-h time periods (0600-
2100) in our study revealed that (i) peak densities for
at least three species occurred during each of the peri-
ods, and (ii) the largest decrease in total bird registra-
tions compared to the peak counts at 0900-1200 h was
only 37% during the period 1800-2100 h. Temporal
variation in bird counts undoubtedly influenced the
density indices for many species but was not consi-
dered a serious problem because of the random way
we conducted our censuses.

Analyses based on single censuses are not recom-
mended during intensive studies where population
estimates are required but have been used successfully
to derive indices to populations or multispecies diver-
sity measurements during extensive surveys (Robbins
and Van Velzen 1967; Stewart and Kantrud 1972;
United States Department of Interior and Canadian
Department of Environment 1977; Rotenberry 1978).
Speirs and Orenstein (1967) have shown that, in open
Ontario habitats, average efficiency of single censuses

is 66-76% that of 6-10 temporally spaced censuses in
estimating breeding populations. I felt justified in bas-
ing indices to bird density on single counts in the
grassland habitats because most of the species possess
behavioral adaptations, including lengthy, loud, and
often elaborate flight songs, perch songs, and syn-
chronous displays, that tend to greatly increase their
detectability compared to birds inhabiting wooded
areas. Birds are also rather easily flushed in grass-
lands. The large areas of the sample plots also resulted
in relatively high absolute numbers of bird observa-
tions per plot. However, I caution against the use of
single censuses in studies where precise population
estimates are desired.

All species were identified in the field by sight or
sound. Counts were based primarily on number of
pairs as indicated by territorial males or segregated
pairs. Each male Red-winged Blackbird (polygynous)
or Brown-headed Cowbird (promiscuous) was consi-
dered to represent only one pair. Single raptors,
grouse, shorebirds, and nighthawks ona sample plot
were considered to represent a pair, but if more than
one was observed the total was halved and rounded to
the nearest lower integer. Additional details on

406 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 1—General features of the major physiographic regions of North Dakota. The number of plots (total 180) on which
bird censuses and vegetative surveys were conducted are shown for the predominant physiographic landforms within each
region.

Region
Southwestern Slope Missouri Coteau Drift Plain Agassiz Lake Plain
A. Topography broadly rolling hilly gently flat
undulating
Drainage integration well poor moderate moderate
Basin wetlands few many many few
Characteristic soils® typic borolls typic and udic borolls udic borolls

udic borolls
Elevation (x m above

sea level)’ 730 580 460 270
Thickness of glacial

sediments thin or absent very thick thick thick
Precipitation

(x cm/yr) 37 41 44 50
Height of dominant

upland grasses &

sedges® short, mid mid, short mid, short tall, mid

B. Thirteen physiographic

landforms/ No. of plots Total
Ground moraine 19 6 25
Terminal moraine 12 12 24
Lake plain 3 14 6 23
Glacial outwash deposits 11 9 20
Eolian sand deposits 8 4 5 17
Unglaciated bedrock 17 17
Dead-ice moraine 16 16
Kames 3 8 11
Sheet moraine 10 10
Glaciated bedrock 5 5
Lake shore deposits 5 5
Deltaic deposits 4 4
Alluvial river deposits 3 3
Total no. of sampled plots 57 28 66 29 180

*Borolls are frigid soils (mean annual temperature <8°C) with nearly black surface horizons high in organic matter and
bases. Udic borolls have greater than average soil moisture than typical (typic) borolls (Aandahl 1972).

From Bluemle (1977) and personal communication.
“From Jensen (n.d.).
“From Stewart (1975).

determination of breeding status have been described
previously (Stewart and Kantrud 1972).

The verges of many plots bordered other habitat
types. To reduce the influence of birds occupying
fencelines or adjacent habitats, census transects were
restricted to the interior of each plot except in the case
of the few that were less than 65 ha in area. The
“hollow square” technique of Speirs and Orenstein
(1967) was used on the standard 65 ha plots; the
1609 m (1 mile) transect route used lay 201 m inside
the boundary (Figure 2). Birds were counted within
101 m of the route, yielding an areal coverage of

31.5 ha. Transect routes were variable on plots of
other sizes and shapes.

Transects were walked at a standard speed of
40 m/min. Indices to bird density were expressed in
pairs/min X 103, allowing standardization of data
from plots of unequal size and shape. The technique of
expressing bird densities in units of time has been used
in similar extensive studies (e.g. Grinnell and Storer
1924; Skinner 1975).

The phenology during late spring and early summer
is about two weeks earlier in southwestern North
Dakota than in the northeastern portion of the state.

1981

Census Transect (1609m)

SW

65ha Square (quarter section)

FIGURE 2. Quarter section sample plot showing transect
route. Shaded areas censused = 31.5 ha.

The effects of these differences on within-season vari-
ations in the bird censuses were partially compensated
for by beginning the survey in the southwestern part of
the state; plots were then visited sequentially on a
broad front moving toward the northeast.

The scientific names of the bird species are provided
only in Table 2.

Vegetation Surveys

A rapid field reconnaissance of the vegetation was
conducted immediately after the bird census on each
plot. An areal cover index was assigned to each species
on the plot. The cover classes and range of cover (%) in
each class were as follows: (1) 0-1, (2) 1-10, (3) 10-25,
(4) 25-50, (5) 50-75, (6) 75-90, (7) 90-100. Class mid-
point values were used in the analysis. Scientific
names of plants follow the Atlas of the Flora of the
Great Plains (Great Plains Flora Association 1977).

Land- Use

Grazing intensity was estimated by comparing the
percentage bare soil, amount of standing and fallen
dead vegetation, and average vegetative height on the
study plots with that of nearby protected or ungrazed
areas. Based on the above information, plots were
placed into three subjective categories: heavily, mod-
erately, and lightly grazed.

Seven of the 180 plots consisted of native grassland

KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA

407

used for hay. These plots had been mowed and raked
in 1973, but not yet mowed in 1974 at the time of
census.

All categories of grazing were represented in each of
the four regions. Hayland was represented in all but
the Southwestern Slope region.

Results

All of the bird species considered in this report nest
below, at, or slightly above the ground in moist to dry
grasses or low shrubs. Many species also breed in
vegetation other than grassland or select other types
of nest sites.

Statewide, the five most abundant species were the
Chestnut-collared Longspur (34% of the total indi-
cated pairs), Western Meadowlark (12%), Clay-
colored Sparrow (10%), Savannah Sparrow (8%), and
Horned Lark (6%), totalling 70% of the avifauna.

In the following sections the occurrence of birds is
considered (i) for the four physiographic regions, (ii)
the nine most important landforms, and (111) the four
land-use categories.

Physiographic Regions

Avian abundance and species composition among
the four physiographic regions are shown in Table 2.
Total bird density and species richness followed sim-
ilar patterns, with highest values occurring in the two
central regions, and lowest values in the Agassiz Lake
Plain. The number of species recorded in each region
showed rather wide variation, ranging from 28 in the
Drift Plain to 19 in the Agassiz Lake Plain.

Nine of the 30 species have a limited distribution
among the regions of the state as indicated by the
maps of Stewart (1975): (1) Long-billed Curlew and
McCown’s Longspur presently occur only in the
extreme western part of the state but formerly were
common farther east; (2) Ferruginous Hawk occurs
regularly in the Southwestern Slope and Missouri
Coteau and appears to be reestablishing breeding
populations in the Drift Plain region; (3) Sprague’s
Pipit, Lark Bunting, and Baird’s Sparrow are rare or
local in the Agassiz Lake Plain region; (4) Marbled
Godwit, Short-billed Marsh Wren, and LeConte’s
Sparrow occur regularly in the three easternmost
regions but are rare or absent in the Southwestern
Slope. The results of our study fully support Stewart’s
(1975) conclusions regarding the nine aforementioned
species.

Several species which occurred in all regions
showed increases or decreases across the regional gra-
dient. From west to east, densities of Killdeer, West-
ern Meadowlark, Red-winged Blackbird, and Savan-
nah Sparrow increased, whereas Horned Lark and
Chestnut-collared Longspur showed reduced density.

408 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 2. Density indices (mean pairs/min X 103) and species richness of 30 grassland birds among major physiographic
regions and land-use types in North Dakota, 1974.

Region Land-use
South- Agassiz Moder- Hayland
western Missouri Drift Lake Heavily ately Lightly Mowed
Slope Coteau Plain Plain Grazed Grazed Grazed 1973

Species No. of plots — (57) (28) (66) (29) (53) (63) (57) (7)
Ferruginous Hawk

(Buteo regalis) 0 3 As bd 1 ] 0 0
Marsh Hawk

(Circus cyaneus) 3 3 ] ; 0 yD 2 1 0
Sharp-tailed Grouse

( Pedioecetes

phasianellus) 1 0 T Uv 0 T 1 0
Killdeer (Charadrius

vociferus) 5 6 14 22 16 9 8 15
Long-billed Curlew

(Numenius americanus) 1 A a - 0 0 1 0
Upland Sandpiper

(Bartramia longicauda) 3 10 18 18 20 7 11 13
Willet (Catoptrophorus

semipalmatus) 2 4 3 0 4 1 1 0
Marbled Godwit

(Limosa fedoa) * 1 a3 14 3 3 5 4
Wilson’s Phalarope

(Steganopus tricolor) 3 1 10 11 6 4 7 30
Mourning Dove

(Zenaida macroura) 8 5 10 20 13 7 12 0
Burrowing Owl

(Athene cunicularia) ] 0 ] 0 l ] 0 0
Common Nighthawk

(Chordeiles minor) ] 0 2 0 ] 0 2 0
Horned Lark (Eremophila

alpestris) 92 67 61 3 154 41 9 0
Short-billed Marsh Wren

(Cistothorus platensis) 2 0 1 13 0 0 3 0
Sprague’s Pipit (Anthus

spragueil) 10 16 13 * 11 15 6 0
Common Yellowthroat

(Geothlypis trichas) 5 32 4 10 0 8 18 33
Bobolink (Dolichonyx

oryzivorus) 4 3 19 73 0 7 39 145
Western Meadowlark

(Sturnella neglecta) 118 140 146 167 130 146 136 189
Red-winged Blackbird

(Agelaius phoeniceus) 15 30 38 101 17 34 61 80
Brown-headed Cowbird

(Molothrus ater) 31 74 68 45 62 49 51 56
Dickcissel (Spiza

americana) 1 0 1 0 0 | ] 0
Lark Bunting (Cala-

mospiza melanocorys) 72 47 15 e 25 57 Dil 0
Savannah Sparrow

(Passerculus

sandwichensis) 12 34 99 295 33 59 147 412
Grasshopper Sparrow

(Ammodramus

savannarum) 63 65 76 32 40 74 75 43

1981 KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA 409
TABLE 2. (concluded)
Region Land-use
South- Agassiz Moder- Hayland
western Missouri Drift Lake Heavily ately Lightly Mowed
Slope Coteau Plain Plain Grazed Grazed Grazed 1973

Species No. of plots > (57) (28) (66) (29) (53) (63) (57) (7)
Baird’s Sparrow

(Ammodramus bairdit) 37 73 62 a 29 57 49 59
LeConte’s Sparrow

(Ammospiza leconteii) ee 0 li 4 0 0 2 7
Vesper Sparrow

(Pooecetes gramineus) 16 6 19 8 14 19 12 0
Clay-colored Sparrow

(Spizella pallida) 56 166 155 61 65 135 139 0
McCown’s Longspur

(Calcarius mccownii) | a we * 0 1 0 0
Chestnut-collared Long-

spur (Calcarius ornatus) 529 505 370 12 606 422 170 87
Index of total bird density
(Mean pairs/ min
X 103/plot) 1094 1290 1210 911 1254 1160 1001 1173
Species richness
(Mean no. species/ plot) 6.18 7.14 U2 6.90 6.28 7.03 7.11 7.29

“Asterisks indicate region(s) in which species currently is rare or absent (Stewart 1975).

°T = density <0.5.

The Southwestern Slope supported distinctly
higher concentrations of two species: Horned Lark
and Lark Bunting. In the Missouri Coteau, Sprague’s
Pipit and Common Yellowthroat were noticeably
commoner than elsewhere. The Agassiz Lake Plain
had the highest number of species (nine) reaching their
greatest density: Killdeer, Marbled Godwit, Mourn-
ing Dove, Short-billed Marsh Wren, Bobolink, West-
ern Meadowlark, Red-winged Blackbird, Savannah
Sparrow, and LeConte’s Sparrow. Species showing
strongest attraction to the grasslands in the two cen-
tral regions, the Missouri Coteau and Drift Plain,

were Brown-headed Cowbird, Baird’s Sparrow, and
Clay-colored Sparrow. The Drift Plain and Agassiz
Lake Plain shared equally high values for densities of
Upland Sandpiper and Wilson’s Phalarope.

Physiographic Landforms

Landforms in which 10 or more plots were cen-
sused, regardless of region, are compared for mean
density of total birds and species richness in Table 3.
Total bird density was more variable than species
richness among landforms. Glacial outwash deposits,
dead-ice moraine, and ground moraine supported

TABLE 3. Total bird density index and species richness on nine major physiographic landforms in four physiographic regions

in North Dakota, 1974’.

No. of
Land Form sample plots
Glacial outwash deposits 20
Dead-ice moraine 16
Ground moraine 25
Lake plain 23
Terminal moraine 24
Sheet moraine 10
Kames 11
Eolian sand 17
Unglaciated bedrock 17

*See Table | for distribution of landforms among regions.

(mean pairs/min X 103/ plot)

Total bird
density index
Mean no. species/ plot

1436 8.35
1434 8.31
1350 7.60
1238 7.74
1182 6.04
1123 5.30
1088 24
816 6.41
810 5.29

410

highest densities of breeding grassland birds. Inter-
mediate densities occurred on lake plain, terminal and
sheet moraine, and kames. Total bird density was
much lower on eolian sand and unglaciated bedrock.
Species richness reached highest values on glacial
outwash deposits and dead-ice moraine, was interme-
diate on lake plain and ground moraine followed by
eolian sand and terminal moraine, and was lowest on
kames, unglaciated bedrock, and sheet moraine.
Comparisons of use of landforms at the species level
are shown in Table 4 by region, but only landforms in

THE CANADIAN FIELD-NATURALIST

Vol. 95

which 10 or more plots were censused in any region
are included. Species comparisons are valid within
and among regions but the landforms listed may not
represent the optimum landform fora species because
only major landforms are listed.

In the Southwestern Slope region, grasslands deve-
loped over glacial outwash deposits supported a
denser and richer avifauna than those developed over
sheet moraine or unglaciated bedrock. Most of the
increased use of outwash deposits was attributable to
high concentrations of Red-winged Blackbird,

TABLE 4. Density indices (mean pairs/min X 103) and species richness of 29 grassland birds among eight major physiographic
landforms within three physiographic regions of North Dakota, 1974’.

Southwestern
Slope Region

Glacial Ungla-

outwash Sheet ciated

deposits moraine bedrock
Species No. of plots > (11) (10) (17)
Ferruginous Hawk
Marsh Hawk 5 4 1
Sharp-taiicd Grouse 4
Killdeer 11 5
Long-billed Curlew 5
Upland Sandpiper 9 1
Willet 2 1
Marbled Godwit
Wilson’s Phalarope 9
Mourning Dove 8 11 =)
Burrowing Owl 3 4
Common Nighthawk 2 1
Horned Lark 101 33 141
Short-billed Marsh Wren
Sprague’s Pipit 12 23 4
Common Yellowthroat 6 8
Bobolink 17 3
Western Meadowlark 128 76 134
Red-winged Blackbird 40 3
Brown-headed Cowbird 70 35 3
Lark Bunting 81 44
Savannah Sparrow 44 7 7
Grasshopper Sparrow 53 52 61
Baird’s Sparrow 111 23 26
LeConte’s Sparrow
Vesper Sparrow 9 30
Clay-colored Sparrow 81 106 2
McCown’s Longspur 5
Chestnut-collared

Longspur 782 723 329

Index of total bird density
(mean pairs/ min
X 103/ plot) 1575 1123 810
Species richness
(mean no. species/ plot) 8.91 5.30 5.29

Missouri

Coteau Region Drift Plain Region

Dead-ice Terminal Ground Lake Terminal
moraine moraine moraine plain moraine
(16) (12) (19) (14) (12)

2 4

3 2 4

1
9 2 9 23 25)
14 5 2 333} 50

6 3 9
1 10 2

2 11 25
6 3 7 15 9

3

79 50 23 76 101

1 1

27 15 28

17 Sy 8 8
6 12 15 12
143 137 154 129 183
22 41 21 71 40
87 56 66 VT 104
52 40 8 7
29 41 156 114 34
84 40 136 59 50
119 12 122 75 6

4
1 13 5 8 23
138 205 118 106 177
586 397 462 316 443
1434 1099 1342 1207 1266
8.31 5.58 7.32 9.07 6.50

*“Dickcissel is omitted from this table because it did not occur on the landforms shown.

1981

Brown-headed Cowbird, Lark Bunting, Savannah
Sparrow, Baird’s Sparrow, and Chestnut-collared
Longspur. Sheet moraine was especially attractive to
Sprague’s Pipit and Clay-colored Sparrow. Grass-
lands developed over unglaciated bedrock supported
noticeably high numbers of Horned Lark and Vesper
Sparrow and they were the only grasslands in the state
harboring breeding populations of Long-billed
Curlew and McCown’s Longspur.

In the Missouri Coteau region, values of both mul-
tispecies measures were higher on dead-ice moraine
than on terminal moraine. Most of the increased use
of the former landform was due to higher population
densities of Sprague’s Pipit, Grasshopper Sparrow,
Baird’s Sparrow, and Chestnut-collared Longspur.
However, terminal moraine was important for a few
species including Common Yellowthroat, Vesper
Sparrow, and Clay-colored Sparrow.

Major landforms in the Drift Plain were less varia-
ble in total bird density and species richness than in
the other two regions. Grasslands developed over
ground moraine supported slightly greater total bird
density but lake plain was highest in species richness.
Noticeably higher numbers of Savannah, Grass-
hopper, and Baird’s Sparrow occurred on ground
moraine, while lake plain was especially attractive to
Marbled Godwit, Wilson’s Phalarope, Sprague’s
Pipit, and Red-winged Blackbird. Terminal moraine
supported highest densities of Upland Sandpiper,
Horned Lark, Western Meadowlark, Brown-headed
Cowbird, and Clay-colored Sparrow.

Land- Use

Density indices for breeding birds are related to
four categories of land-use in Table 2. Although the
total number of tracts of hayland censused was small
(7), this habitat has been included because of its
apparently strong attractiveness or aversiveness to
some species.

Mean density of birds tended to increase as grazing
intensity increased but species richness tended to
decrease. Of the three grazing types, heavily grazed
areas supported only 22 species compared to 26and 27
for moderately and lightly grazed areas, respectively.
Grassland mowed at the end of the previous growing
season but currently unmown at time of the census
supported a density of birds comparable with the
moderately grazed areas, but was slightly higher in
species richness than any of the grazed areas.

Heavily grazed grasslands were especially attractive
to Killdeer, Upland Sandpiper, Horned Lark, and
Chestnut-collared Longspur. The two latter species
comprised almost two-thirds (61%) of the total bird
density on these areas. In addition, Sprague’s Pipit
apparently preferred both heavily and moderately
grazed plots over lightly grazed plots.

KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA

411

Only one species, the Lark Bunting, showed a
strong tendency to attain peak density under moder-
ate grazing. Three common sparrows, including Grass-
hopper, Baird’s, and Clay-colored, were about equally
highly abundant under both moderate and light graz-
ing. Four species, including Common Yellowthroat,
Bobolink, Red-winged Blackbird, and Savannah
Sparrow showed much higher densities under light
grazing.

Species attracted to hayland that had been mowed
the previous year included Wilson’s Phalarope,
Common Yellowthroat, Bobolink, Western Meadow-
lark, Red-winged Blackbird, and Savannah Sparrow.
Grasslands under this type of land-use were avoided
by Mourning Dove, Horned Lark, Sprague’s Pipit,
Lark Bunting, Vesper Sparrow, and Clay-colored
Sparrow.

Vegetation

Dominant plants of the major physiographic land-
forms are shown in Table 5. Among the grasses, Poa
pratensis, an introduced species, greatly dominated
the flora in central and eastern North Dakota. Among
the native grasses, Bouteloua gracilis, Koeleria
Pyramidata, and Stipa comata were important in all
major landforms in the three westernmost regions.

Agropyron smithii was important in all regions but
was replaced by A. caninum on eastern lake plain.
Native grasses of importance primarily in eastern
regions included Andropogon scoparius and Stipa
viridula. Two short grasses, Buchloe dactyloides and
Festuca octoflora, were important only in the west.
Three short carices (Carex eleocharis, C. filifolia, C.
pensylvanica) were important in two or more regions.

The short clubmoss, Selaginella densa, was an
important floral component on outwash deposits in
the Southwestern Slope and on dead-ice moraine in
the Missouri Coteau. Symphoricarpos occidentalis
was the only shrub of importance on the major physi-
ographic landforms. It was found in especially large
amounts on landforms in the Drift Plainand Missouri
Coteau. The only forbs of importance included
Antennaria rosea on lake plain in the Drift Plain
regionand Plantago patagonica, Psoralea argophylla,
and Artemisia frigida on unglaciated bedrock in the
Southwestern Slope region.

Discussion
Physiographic Regions

Grasslands in the four physiographic regions of
North Dakota range in type from subhumid tallgrass
prairie in the Agassiz Lake Plain to semiarid plains in
the Southwestern Slope dominated by short and mid-
grasses. The total bird density in particular areas of
the North Dakota grasslands can be attributed largely

412

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 5. Mean % cover of the ten most abundant plant species in eight major physiographic landforms within three

physiographic regions of North Dakota, 1974.

Southwestern
Slope Region

Glacial Ungla-

outwash Sheet ciated

deposits moraine bedrock
Species No. of plots — (11) (10) (17)
Selaginella densa 7
Psoralea argophylla 6 5)
Plantago patagonica 12
Symphoricarpos

occidentalis 4

Antennaria rosea
Artemisia frigida 6 7
Carex eleocharis 16 12
Carex filifolia 10 12 17
Carex pensylvanica 8 9
Agropyron caninum
Agropyron smithii 9 1] 22
Andropogon scoparius
Bouteloua gracilis 17 23 21
Buchloe dactyloides
Festuca octoflora 9
Koeleria pyramidata 25 21 22
Poa pratensis 21
Stipa comata 17 21 16
Stipa viridula 12

to a few species (Table 2). The Missouri Coteau and
Drift Plain not only supported 10-29% greater total
bird density than the other regions but also a 3-15%
greater species richness. The increased bird density in
these two regions is due to their moderate attractive-
ness to species that reach greater densities, either in
more arid areas to the west or in moister grasslands to
the east, combined with markedly higher populations
of a few characteristic species. Increased species rich-
ness in the two regions may be related to the abun-
dance there of natural basin wetlands (the “Prairie
Pothole Region” of waterfowl ecologists; Stewart and
Kantrud 1971, 1973). The undulating topography is
ecologically heterogeneous and soils bordering the
wetlands generally support growths of taller grasses.
Patches of mesic mixed-grass prairie occur adjacent to
the tallgrass prairie and shorter, xeric mixed-grass
prairie occurs nearby but at higher elevations. Each of
these grassland types may be large enough to attract
their characteristic bird species within a few hectares,
thus increasing species richness. Although Wiens
(1974) found relatively uniform densities of breeding
birds among five types of North American grasslands,
representing a much larger range of environmental
conditions than occur among North Dakota grass-
lands, plot-to-plot and regional variations in grazing

Missouri

Coteau Region Drift Plain Region

Dead-ice Terminal Ground Lake Terminal
moraine moraine moraine plain moraine
(16) (12) (19) (14) (12)

7 7
16 16 14 7 27
5
5 6
6 19 6 8
10 6
8 7 6 20
7
10 7 10 8
5 5
15 13 22 25 34
10 12 12 7 6
32 40 49 54 54
12 10 4 6 5
5 5 6 5

intensity may have obscured regional differences that
may have existed.

There are indications that the number of bird spe-
cies may also be increased in the Missouri Coteau and
Drift Plain because the grasslands are predominately
of intermediate height compared to the other two
regions. Cody (1968, 1974) concluded that, while the
shortgrass prairie was divided on the basis of foraging
behavior and the tallgrass prairie according to vertical
foraging range, in grasslands of intermediate height
(about 0.75 m) the avifauna coexists by subdividing
the habitat itself into patches of different height that
are species-specific, further enhancing species
richness.

Of 11 passerines found throughout North Dakota,
three of the four largest in body size (Bobolink, West-
ern Meadowlark, Red-winged Blackbird) occurred in
highest density in the Agassiz Lake Plain, the region
of greatest annual precipitation (Table 1). This tends
to corroborate Wiens’ (1974) suggestion that the
standing crop biomass of passerine species may
decrease across a gradient from subhumid tallgrass
prairie to arid shrubsteppe.

On the Agassiz Lake Plain only 19 species were
recorded compared to 22-28 for the other regions.
The lower number is probably due in part to habitat

1981

destruction by agricultural cultivation: Ferruginous
Hawk, Sharp-tailed Grouse, Burrowing Owl, Long-
billed Curlew, and Sprague’s Pipit have been extir-
pated, or now survive only in very low numbers, in the
Agassiz Lake Plain.

Physiographic Landforms

Grasslands developed over landforms lacking
extensive soil development (sheet moraine, ungla-
ciated bedrock) and those on coarse, nutrient-poor
glacial deposits (kames, eolian sand) supported rela-
tively low concentrations of birds and few species.
Increased density and species richness was found on
flatter landforms (outwash deposits, lake plain), land-
forms with the most fertile glacial soils (ground
moraine, lake plain), and landforms with a greater
variety of habitats (dead-ice moraine, ground
moraine).

Differential use of landforms by individual species
may be caused by a variety of factors. The higher
densities of Baird’s Sparrow and Chestnut-collared
Longspur, in the Southwestern Slope region, on grass-
lands developed on outwash deposits, may be due to
the flat topography characteristic of that landform.
Flat topography has been noted as favorable for those
species by Thompson (1890), Bird (1930), and Rand
(1948). Terminal moraine, which is hillier than dead-
ice moraine and contains fewer wetlands, supported a
more dispersed, species-poor avifauna in the Missouri
Coteau. Grasshopper Sparrow (Taverner 1927),
Baird’s Sparrow (Thompson 1890), and Chestnut-
collared Longspur may have been more attracted to
dead-ice moraine because of its more gentle relief. On
the other hand, brush is more common on terminal
moraines which explained their importance to such
species as Common Yellowthroat, Vesper Sparrow,
and Clay-colored Sparrow.

In the Drift Plain, the flat, moist lake plain was
attractive to Marbled Godwit, Wilson’s Phalarope,
Sprague’s Pipit, and Red-winged Blackbird. Ground
moraine (which occupies intermediate elevations)
supported higher populations of mixed-grass prairie
species, including Grasshopper and Baird’s Sparrows.
The high, arid terminal moraines had higher densities
of Horned Lark. Terminal moraines also often sup-
port well-developed stands of Symphoricarpos occi-
dentalis which are highly attractive to Clay-colored
Sparrow.

Land-Use
Grazing

During this study, mean total bird density was
highest on heavily grazed plots, intermediate on mod-
erately grazed, and lowest on lightly grazed. Heavy
grazing resulted in reduced species richness, and

KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA 413

increased dominance bya few species. The response of
breeding birds to grazing (and other land-use practi-
ces) in mixed-grass prairies has been previously stu-
died in two areas of North Dakota, Billings County
(Hopkins 1980) and Stutsman County (Johnson 1972-
1974), and also at Cottonwood, South Dakota (Wiens
1973); Nebraska Sandhills (Wiens and Dyer 1975);
Matador, Saskatchewan (Maher 1973); and two sites
in Alberta (Owens and Myres 1973; Karasiuk et al.
1977). Total bird density was always higher on idle or
lightly grazed sites, except on the Saskatchewan area,
which is in contrast to what is reported in this paper
and so deserves discussion.

(i) Horned Lark and Chestnut-collared Longspur

During this study, over 60% of the total bird density
on heavily grazed areas was attributable to the
Horned Lark and Chestnut-collared Longspur. Dur-
ing the Saskatchewan study, the Horned Lark and the
longspurs comprised over 85% of the bird population
on grazed plots. A similar situation apparently exists
in Manitoba where Harris (1944) stated that all but
one of the Chestnut-collared Longspur nests were
poorly concealed in cover thinned by cattle and grass-
hoppers. In Saskatchewan, Belcher (1961) and Roy
(1964) also noticed the affinity of this species for over-
grazed pastures. However, in Alberta, Owens and
Myres (1973) and Smith (1976) stated that the semico-
lonial nature of the Chestnut-collared Longspur there
results in great local variation in numbers.

The effects of grazing on rangeland avifaunas may
apparently vary markedly in different western grass-
land types (Wiens and Dyer 1975). For example, on
the shortgrass prairie of Colorado, Giezentanner
(1970) found only small differences in density of
Horned Lark among lightly, moderately, and heavily
summer-grazed plots and the Chestnut-collared
Longspur was absent under heavy grazing in that
area. Bailey and Niedrach (1938) noted that Chestnut-
collared Longspur were restricted to valleys in Colo-
rado. In Montana, DuBois (1935) observed that this
species nested in lower, moister areas than the Horned
Lark.

(ii) Clay-colored Sparrow, Grasshopper Sparrow,
and Baird’s Sparrow

For these species, the negative response to heavy
grazing, noted during this study, agrees with the
results of the previous seven Canadian and American
studies of the effects of grazing on the avifauna of
mixed-grass prairie. Our findings also agree with
those of Walkinshaw (1940) for the Grasshopper
Sparrow and of Todd (1947) and Lane (1968) for the
Baird’s Sparrow.
(iii) Sprague’s Pipit

Our findings on the effects of grazing on Sprague’s

414 THE CANADIAN FIELD-NATURALIST

Pipit are in disagreement with the results of the three
Canadian studies: Maher (1973) and Owens and
Myres (1973) found much higher use of idle than of
grazed plots by this species, and Karasiuk et al. (1977)
found much greater numbers on lightly than on heav-
ily grazed grasslands. In contrast, moderately and
heavily grazed areas received greater use by Sprague’s
Pipit in North Dakota. This finding may again reflect
(as in the cases of the Horned Lark and Chestnut-
collared Longspur) the variable effects of grazing ona
bird species in various grassland types. In Montana,
Saunders (1914) also noted that the Sprague’s Pipit
commonly occurred in areas of sparse grass.

(iv) Western Meadowlark

In North Dakota, the Western Meadowlark was
about equally common under the three levels of graz-
ing intensity. Johnson (1972-1974) and Hopkins
(1980) also found that grazing had negligible effects
on this species in North Dakota. In both Alberta
studies, it was about equally common under idle or
grazed conditions. Inthe Nebraska Sandhills, it was a
dominant species on grazed but not ungrazed prairie.
In the South Dakota and Saskatchewan studies the
Western Meadlowlark apparently preferred less heav-
ily grazed areas. In Oklahoma, Smith (1940) found
that only severe overgrazing made conditions unsuit-
able for it. In Montana, Thompson and Dahmer
(1978) noted that severe grazing caused the species to
abandon territories early in the breeding season.

(v) Upland Sandpiper

Upland Sandpiper showed little response to grazing
pressure during this study, and that of Johnson (1972-
1974), in North Dakota. It has been shown that, in
Wisconsin and North Dakota, Upland Sandpiper
prefer dense vegetative cover for nest sites (Ailes 1976;
Higgins et al. 1969) and that grazing can be detrimen-
tal (Buss and Hawkins 1939; Kirsch and Higgins
1976). In one South Dakota study it occurred only in
ungrazed plots (Wiens 1973), but in another (Loke-
moen and Duebbert 1974) a very high nesting popula-
tion occurred under moderate grazing. Upland Sand-
piper may feed and rear their broods in very different
habitats from that which they prefer for nest sites
(Ailes 1976).

(vi) Lark Bunting

Lark Bunting showed a fairly strong affinity for
moderately grazed areas during this study. Hopkins
(1980) found small numbers only under moderate
grazing in the North Dakota mixed-grass prairie while
Johnson (1972-1974) recorded the species only under
heavy grazing. On shortgrass prairie in the west, Lark
Bunting is rare or absent under heavy summer grazing
(Rand 1948; Finzel 1964; Giezentanner 1970).

Vol. 95

(vii) Bobolink and Savannah Sparrow

In this study, these two species were greatly reduced
or extirpated by heavy grazing. Johnson (1972-1974)
found highest densities of Bobolink under lightly
grazed conditions in North Dakota grasslands. Other
studies of Bobolink (Kendeigh 1941; Tester and Mar-
shall 1961; Hibbard 1965, 1972; Cody 1968; Wiens
1969; Martin 1971; Potter 1972; Harrison 1974) indi-
cate a strong preference for tall, dense vegetation.

Johnson (1972-1974) found Savannah Sparrow
about equally common under lightly and heavily
grazed conditions in North Dakota grasslands but
only small numbers of birds were observed. A prefer-
ence for idle or lightly grazed areas was noted by
Lincoln (1925), Rand (1948), Owens and Myres
(1973), Karasiuk et al. (1977), and Maher (1979). This
species shows an affinity for areas possessing dense
ground cover (Tester and Marshall 1961; Lein 1968;
Wiens 1969; Potter 1972).

(vill) Other Species

Short-billed Marsh Wren, Common Yellowthroat,
Red-winged Blackbird, and LeConte’s Sparrow were
greatly reduced or extirpated by heavy grazing. John-
son (1972-1974) found highest densities of Common
Yellowthroat under ungrazed conditions in North
Dakota.

Mowing

Comparisons of data for plots of hayland with
those from past studies are complicated by a number
of factors. During this study, haylands censused had
been mowed and raked the previous year, but not
during the year of census. Thus by late spring they had
grown appreciably, and attained many of the aspects
of idle grassland without a well developed litter layer.
This condition was characterized by dense popula-
tions of species which prefer taller grass for nesting
cover: these included Bobolink, Red-winged Black-
bird, Savannah Sparrow, and LeConte’s Sparrow.

Two plots studied by Owens and Myres (1973) in
Alberta also had been mown and raked the previous
year, but by mid-June of the year of census they
contained vegetation only 20-30 cm tall; in this case
the avian community was dominated by Sprague’s
Pipit and Chestnut-collared Longspur, two species
which in North Dakota grasslands showed favorable
response to short vegetation created by moderate or
heavy grazing.

Though nowhere common, LeConte’s Sparrow
reached peak numbers on hayland during this study,
whereas Murray (1969), also in North Dakota, noted
the importance of litter for breeding of this species and
believed annual mowing would extirpate it. This dis-
parity may be related not only to the time of harvest
but also to differences in forage-harvesting tech-

1981

niques. Thus, removal of grassland vegetation has
been shown to have different effects on important
characteristics of the vegetation such as height, den-
sity, and amount of litter (Owens and Myres 1973;
Weaver and Fitzpatrick 1934; Wiens 1973), and sev-
eral studies (Tester and Marshall 1961; Wiens 1969)
have shown that changes in litter appear to cause
changes in distribution and abundance of Bobolink,
Savannah Sparrow, and LeConte’s Sparrow.

Bobolink peak densities occurred on hayland in this
investigation and those of Bailey (1915), Kendeigh
(1941), and Springer (1965). Hibbard (1972) attrib-
uted lower numbers of this species on burned tallgrass
prairie to lack ofa litter layer. Martin (1971) reported
that Bobolink males which attracted the most females
inhabited fields with increased vegetation density and
horizontal heterogeneity, but male pairing levels were
not correlated with litter depth. Without detailed
experimental studies it is difficult to determine which
vegetational factors these populations were respond-
ing to, because Penfound (1964) has shown that denu-
dation of the litter layer in tallgrass prairie through
mowing ultimately results in increased vegetative
biomass, productivity, and number of species.

In conclusion, the results of this study indicate that
the distribution and abundance of most grassland bird
species in North Dakota have been negatively affected
by agricultural and pastoral activities. Grassland plots
on the flatter, more fertile landforms were difficult to
find, especially in the intensively-cultivated Agassiz
Lake Plain region. This region supported highest
measured densities of most species found there, but
was also the region in which the fewest species were
recorded, indicating that fragmentation of grasslands
by agriculture has not only resulted in extirpation of
species originally present but that, acre-for-acre, cul-
tivation of eastern tallgrass prairie has destroyed bet-
ter habitat for more bird species than in the shorter
grasslands to the west. Abuse of native grasslands by
overgrazing has also had a negative impact as evi-
denced by the small number of species that reached
highest densities and the reduced species richness
under heavy grazing.

I recommend that protection be given to larger
remaining areas of unplowed grassland representative
of the various types found throughout the state. Graz-
ing and burning management should then be applied
in order to promote the continued survival of the
native grassland birds that are now much less com-
mon than formerly.

Acknowledgments

R. E. Stewart supervised this investigation and par-
ticipated in the field work. Assistance in the field was
received from L. M. Cowardin, D. S. Gilmer, A. T.

KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA 415

Klett, and J. T. Lokemoen. I thank D. Alan Daven-
port and D. H. Johnson for their help with computer
programming and C. A. Faanes and J. R. Serie for
reviewing earlier drafts of the manuscript. The study
was supported by the U.S. Fish and Wildlife Service
through the Migratory Birds Program and the Coal
Project, Western Energy Land Use Team, Office of
Biological Services.

Literature Cited

Aandahl, A. R. 1972. Soils of the Great Plains. P.O. Box
81242, Lincoln, Nebraska 68508.

Ailes, I. W. 1976. Ecology of the Upland Sandpiper in cen-
tral Wisconsin. M.S. thesis, University of Wisconsin,
Stevens Point. 63 pp.

Bailey, A. M. and R. J. Niedrach. 1938. The Chestnut-
collared Longspur in Colorado. Wilson Bulletin 50:
243-246.

Bailey, F. M. 1915. Characteristic birds of the Dakota
prairies I. In the open grassland. Condor 17: 173-179.

Belcher, M. 1961. Birds of Regina. Saskatchewan Natural
History Society. Special Publication 3. 76 pp.

Bird, R. D. 1930. Biotic communities of the aspen parkland
of central Canada. Ecology 11: 356-442.

Bluemle, J. P. 1977. The face of North Dakota — the geo-
logic story. North Dakota Geological Survey Educational
Series 11. 73 pp.

Buss, I. O.and A. S. Hawkins. 1939. The Upland Plover at
Faville Grove, Wisconsin. Wilson Bulletin 51: 202-220.
Cody, M. L. 1968. On the methods of resource division in
grassland bird communities. American Naturalist 102:

107-147.

Cody, M. L. 1974. Competition and the structure of bird

communities. Princeton University Press, Princeton. 318

Pp.

DuBois, A. D. 1935. Nests of Horned Larks and longspurs
ona Montana prairie. Condor 37: 56-72.

Finzel, J. E. 1964. Avian populations of four herbaceous
communities in southeastern Wyoming. Condor 66:
496-510.

Giezentanner, J. B. 1970. Avian distribution and popula-
tion fluctuations on the shortgrass prairie of north-central
Colorado. U.S. International Biological Program, Grass-
land Biome Technical Report 62. 112 pp.

Great Plains Flora Association. 1977. Atlas of the flora of
the Great Plains. Iowa State University Press, Ames. 600

pp.

Grinnell, J. and T.I. Storer. 1924. Animal life in the
Yosemite. University of California Press, Berkeley. 752
pp.

Harris, R. D. 1944. The Chestnut-collared Longspur in
Manitoba. Wilson Bulletin 56: 105-115.

Harrison, K. G. 1974. Aspects of habitat selection in grass-
land birds. M.A. thesis, Western Michigan University,
Kalamazoo. 82 pp.

Hibbard, E. A. 1965. Comments on the distribution and
abundance of the Lark Bunting and other prairie fringil-
lids in Minnesota and North Dakota. Loon 37: 70-72.

Hibbard, E. A. 1972. Burned and unburned prairie. Amer-
ican Birds 26: 1004-1005.

416

Higgins, K.F., H.F. Duebbert and R.B. Oetting.
1969. Nesting of the Upland Plover on the Missouri
Coteau. Prairie Naturalist 1: 45-48.

Hopkins, R. B. 1980. Mixed prairie I. American Birds 34:
67-68.

Jarvinen, O., R. A. Vaisanen, and Y. Haila. 1976. Estimat-
ing relative densities of breeding birds by the line transect
method. III. Temporal constancy of the proportion of
main belt observations. Ornis Fennica 53: 40-45.

Jensen, R. E. (no date). Climate of North Dakota. North
Dakota State University, Fargo. 48 pp.

Johnson, D. H. 1972-1974. Breeding bird populations of
selected grasslands in east-central North Dakota. Ameri-
can Birds 26: 971-975; 27: 989-990; 28: 1030-1031.

Karasiuk, D., H. Vriend, J. G. Stelfox and J. R. McGillis.
1977. Study results from Suffield, 1976. Jn Effects of lives-
tock grazing on mixed prairie range and wildlife within
PFRA pastures, Suffield Military Reserve. Compiled by
J. G. Stelfox. Range-Wildlife Study Committee, Cana-
dian Wildlife Service, Edmonton, Alberta. pp. E33-E44.

Kendeigh, S. C. 1941. Birds of a prairie community. Con-
dor 43: 165-174.

Kirsch, L. M. and K. F. Higgins. 1976. Upland Sandpiper
nesting and management in North Dakota. Wildlife
Society Bulletin 4: 16—22.

Lane, J. 1968. Baird’s Sparrow. Jn Life histories of North
American cardinals, grosbeaks, buntings, towhees,
finches, sparrows, and allies. By A. C. Bent. Dover, New
York. pp. 745-765.

Lein, M. R. 1968. The breeding biology of the Savannah
Sparrow, Passerculus sandwichensis (Gmelin) at Saska-
toon, Saskatchewan. M.A. thesis, University of Saskat-
chewan, Saskatoon. 171 pp.

Lincoln, F. C. 1925. Notes on the bird life of North Dakota
with particular reference to the summer waterfowl. Auk
42: 50-64.

Lokemoen, J. T.and H. F. Duebbert. 1974. Summer birds
for a South Dakota prairie. South Dakota Conservation
Digest 41(2): 18-21.

Maher, W. J. 1973. Birds: I. Population dynamics. Cana-
dian Committee for the International Biological Pro-
gramme. Matador Project, Technical Report 34. 56 pp.

Maher, W. J. 1979. Nestling diets of prairie passerine birds
at Matador, Saskatchewan, Canada. Ibis 121: 437-452.

Martin, S. G. 1971. Polygyny in the Bobolink: habitat
quality and the adaptive complex. Ph.D. dissertation,
Oregon State University, Corvallis. 181 pp.

Murray, B.G., Jr. 1969. A comparative study of the
LeConte’s and Sharp-tailed Sparrows. Auk 86: 199-231.

Owens, R. A.and M. T. Myres. 1973. Effects of agriculture
upon populations of native passerine birds of an Alberta
fescue grassland. Canadian Journal of Zoology 51:
697-713.

Patterson, D. D., G. A. Johnsgard, M. D. Sweeney, and
H. W. Omodt. 1968. Soil Survey Report; county general
soil maps. North Dakota Agricultural Experiment Station
Bulletin 473. 150 pp.

Penfound, W. T. 1964. Effects of denudation on the pro-
ductivity of grassland. Ecology 45: 838-845.

Potter, P. E. 1972. Territorial behavior in Savannah Spar-
rows in southeastern Michigan. Wilson Bulletin 84: 48-59.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Rand, A. L. 1948. Birds of southern Alberta. National
Museum of Canada Bulletin 111. 105 pp.

Robbins, C. S.,and W. T. Van Velzen. 1967. The breeding
bird survey, 1966. U.S. Fish and Wildlife Service Special
Scientific Report-Wildlife 102. 43 pp.

Rotenberry, J.T. 1978. Components of avian diversity
along a multifactorial climatic gradient. Ecology 59:
693-699.

Roy, J. F. 1964. Birds of the Elbow. Saskatchewan Natural
History Society, Regina. 33 pp.

Saunders, A. A. 1914. The birds of Teton and northern
Lewis and Clark counties, Montana. Condor 16: 124-144.

Skinner, R. M. 1975. Grassland use patterns and prairie
bird populations in Missouri. /n Prairie: A multiple view.
Edited by M. K. Wali. University of North Dakota Press,
Grand Forks. pp. 171-180.

Smith, A. R. 1976. Comparable results from similar areas
at Matador, Saskatchewan and Hand Hills, Alberta. In
Effects of livestock grazing on mixed prairie range and
wildlife within PFRA pastures, Suffield Military Reserve.
Compiled by J. G. Stelfox. Range-Wildlife Study Com-
mittee, Canadian Wildlife Service. Edmonton, Alberta.
pp. E45-E54.

Smith, C. C. 1940. The effect of overgrazing and erosion
upon the biota of the mixed-grass prairie of Oklahoma.
Ecology 21: 381-397.

Speirs, J. M. and R. Orenstein. 1967. Bird populations in
fields of Ontario County, 1965. Canadian Field-Naturalist
81: 175-183.

Springer, P.F. 1965. 51. Tall and midgrass prairie.
Audubon Field Notes 19: 618-619.

Stewart, R. E. 1975. Breeding birds of North Dakota. Tri-
College Center for Environmental Studies, Fargo. 295 pp.

Stewart, R. E. and H. A. Kantrud. 1971. Classification of
natural ponds and lakes in the glaciated prairie region.
U.S. Fish and Wildlife Service, Resource Publication 92.
57 pp.

Stewart, R. E. and H. A. Kantrud. 1972. Population esti-
mates of breeding birds in North Dakota. Auk 89:
766-788.

Stewart, R. E.and H. A. Kantrud. 1973. Ecological distri-
bution of breeding waterfowl populations in North
Dakota. Journal of Wildlife Management 37: 39-50.

Taverner, P. A. 1927. Some recent Canadian records. Auk
44: 217-228.

Tester, J. R.and W. H. Marshall. 1961. A study of certain
plant and animal interrelations on a native prairie in
northwestern Minnesota. University of Minnesota
Museum of Natural History, Occasional Papers 8. 51 pp.

Thompson, E. E. 1890. The birds of Manitoba. Proceed-
ings of the U.S. National Museum 13: 457-643.

Thompson, L. S. and T. D. Dahmer. 1978. Breeding birds
of prairie grassland and shrubland habitats in northeast-
ern Montana. American Birds 32: 107-109.

Todd, W. E. C. 1947. Notes on the birds of southern Sas-
katchewan. Annals, Carnegie Museum 30 (Art. 22):
383-421.

U.S. Department of the Interior and Canadian Department
of the Environment. 1977. Standard operating proce-
dures for aerial waterfowl breeding ground population
and habitat surveys. Section IV. 33 pp.

1981 KANTRUD: GRAZING EFFECTS ON AVIFAUNA OF NORTH DAKOTA 417

Walkinshaw, L. H. 1940. Some Michigan notes on the
Grasshopper Sparrow. Jack-Pine Warbler 18: 50-59.

Weaver, J. E.and T. J. Fitzpatrick. 1934. The prairie. Eco-
logical Monographs 4: 109-295.

Wiens, J. A. 1969. An approach to the study of ecological
relationships among grassland birds. American Ornithol-
ogists’ Union. Ornithological Monographs 8. 93 pp.

Wiens, J. A. 1973. Pattern and process in grassland bird
communities. Ecological Monographs 43: 237-270.

Wiens, J. A. 1974. Climatic instability and the “ecological
saturation” of bird communities in North American grass-
lands. Condor 76: 385-400.

Wiens, J. A. and M.I. Dyer. 1975. Rangeland avifaunas:
their composition, energetics, and role in the ecosystem. In
Proceedings of the symposium on management of forest
and range habitats for nongame birds. Coordinated by
D. R. Smith. U.S. Department of Agriculture. Forest Ser-
vice, General Technical Report WO-1. pp. 146-182.

Received 5 May 1980
Accepted 28 March 1981

APPENDIX A. Descriptions of 13 physiographic landforms on which sample sites were located in North Dakota.

Landform
Alluvial River Deposits

Dead-ice Moraine

Deltaic Deposits
Eolian Sand

Glacial Outwash Deposits

Glaciated Bedrock

Ground Moraine

Kames

Lake Plain

Lake Shore Deposits
Sheet Moraine

Terminal Moraine

Unglaciated Bedrock

Description

Generally flat topography underlain by glacial sediments deposited by modern rivers on
valley floors.

Hilly areas where large amounts of rock and sediment, incorporated into and on top of
advancing glaciers, slumped and slid as the glaciers stagnated and melted. Dead-ice
moraine contains large numbers of both deep and shallow wetlands.

Flat or gently undulating sandy or silty areas formed where large rivers flowed into
glacial lakes.

The wind-worked surfaces of sandy areas such as glacial river deltas or outwash plains.
Topography is generally of choppy dunes with a few shallow wetlands.

Broad, flat plains of sand and gravel deposited by water flowing from melting glaciers.
Saline wetlands are common in many outwash areas. In some areas, outwash was
deposited over stagnant ice which resulted in more rugged topography and numerous,
less saline wetlands.

Various types of sedimentary bedrock (shales, sandstones, siltstones, etc.). Glaciated but
glacial remnants usually restricted to a few scattered boulders. These sediments are well
drained due to post-glacial erosion.

Gently rolling topography covered with moderate amounts of glacial till. This is the most
common glacial landform in North Dakota and is characterized by fertile soils and
numerous shallow basin wetlands.

Mounds and conical hills of gravel, sand, silt, and clay which were formed when
depressions in glaciers filled with these materials slumped during the glacial melt period.

Flat deposits of sediment which formed the bottoms of lakes created by glacial blockage
of major drainageways. The most fertile agricultural soils in North Dakota occur in lake
plains (Patterson et al. 1968).

Gently undulating sand and gravel beaches or wave-cut scarps of other material formed
at the edges of glacial lakes.

Areas occupying the outer limits of glaciation which contain a continuous or patchy thin
layer of glacial till.

Hilly, thick deposits of glacial till formed at the edges of glaciers. Terminal moraine
commonly occur as areas a few km wide and several tens of km long and contain
moderate numbers of small, shallow wetlands.

Bedrock of various types lying south and west of the furthest advance of Wisconsin age
glaciation. Gently undulating grasslands on this type usually have developed ona single
parent material but several parent materials may be represented over large areas. Well
drained due to erosion with few natural basin wetlands.

Relative Abundances of Birds in Boreal and Subarctic Habitats of
Northwestern Ontario and Northeastern Manitoba

MARGARET A. MCLAREN and PETER L. MCLAREN

LGL Ltd., 414-44 Eglinton Ave. W., Toronto, Ontario M4R IAI

McLaren, Margaret A. and Peter L. McLaren. 1981. Relative abundances of birds in boreal and subarctic habitats of
northwestern Ontario and northeastern Manitoba. Canadian Field-Naturalist 95(4): 418-427.

Indices of abundance of breeding birds based on transect surveys are presented for an area stretching | 000 km NW from the
Albany River, Ontario to the Caribou River, Manitoba. Data are summarized for six geographic areas and nine habitats.
Overall abundance changed little with latitude. Species composition changed considerably and these changes are discussed in
relation to habitat. The highest abundance indices of birds were found in deciduous scrub and the lowest in coniferous forest

with no understory.

Key Words: Breeding birds, relative abundance, boreal forest, subarctic, Ontario, Manitoba.

The avifauna of the interior of northwestern Onta-
rio and northeastern Manitoba is poorly known.
James et al. (1976) summarized the known distribu-
tions of birds in Ontario and made qualitative assess-
ments of abundance. Their data base was very small
for interior areas north of 51° N and consisted primar-
ily of records from visits to Favourable Lake in 1938
(Royal Ontario Museum records) and Big Trout Lake
in 1975 (Lee 1978). The interior of northeastern
Manitoba is similarly poorly known although one
quantitative study has been done there (Gillespie
1960). ;

In this paper we present indices of abundances
(expressed as birds/km2) of breeding birds in terres-
trial habitats along a proposed pipeline corridor inan
area reaching from the Albany River, Ontario,
northwest 1000 km to the Caribou River, Manitoba.
Supplementary information about limits of distribu-
tion and nesting has been presented in McLaren and
Renaud (1980) and McLaren and McLaren (1981).

Methods

We conducted line transect surveys at 55 sites from
3-27 June 1977 (Figure 1). Sites within 75 km of Pickle
Lake, Ontario and at Gillam, Manitoba, were reached
by car; all others were reached by helicopter. Sites
were distributed along the pipeline corridor; logistic
considerations with the helicopter (possible landing
sites, cruising range) precluded regularly spaced sam-
pling. About 2.6 km (range: | 280-4 570 m) of tran-
sect were surveyed at each site and all but 2 sites were
surveyed between 04:40 and 10:00. Transects nor-
mally followed the form of a square or three sides of a
rectangle. ‘Edge effects’ were avoided by starting and
finishing transects at least 50 m from a road, lake or
river.

We conducted line transect surveys, rather than
plot surveys, because of the necessity of sampling a

large area (ca. | 000 km in length) in a single season.
The method used was a variant of those described by
Graber and Graber (1963) and by Emlen (1971,1977).
At each site two observers walked parallel courses
20 m apart; one recorded the birds detected while the
other recorded habitat information. Bird data
included the species, number detected, estimated dis-
tance of the bird from the centre-line of the transect,
vocalizations (if any) and when possible, behaviour,
age, and sex of the bird(s) detected. All birds detected,
regardless of their lateral distances, were recorded.
The second observer used a hand-held counter to tally
the number of paces covered and recorded habitat
type and variables every 200 paces. This observer also
noted the pace number where the habitat type
changed. A factor converting paces to metres was
determined for each ‘habitat recorder’ and applied to
the pace data. The fact that we could regularly cir-
cumnavigate a square by compass, returning to within
about 50 m of the starting point, suggests that pacing
was consistent.

Only birds estimated to be within 50m of the
centre-line of the transect strip were used in the calcu-
lation of indices. Records of flocks of five or more
birds and records of birds flying over the transect strip
from a location off-transect but not landing on the
transect were not included in calculations.

The index of abundance was calculated as the
number of birds per square kilometre. This represents
only minimum densities of birds present in an area or
habitat. Absolute densities cannot be determined
from single replicate transect surveys since not all
birds will be detected. Methods to estimate absolute
densities using correction factors determined from
replicate transects have been developed (Emlen 1977;
Jarvinen and Vaisanen 1976; Jarvinen et al. 1976,
1977). However, the scope of our study did not permit
us to replicate transects and thus determine correction

418

1981

FOREST-
TUNDRA |
TRANSITION &’

CHURCHILL 2

RIVER ' | TYRRELL 5
a! SEA 4
1@, BEACH

‘e i

NELSON

UDSON BAY
OWLANDS

SEVERN RIVER

@,
‘@ NORTH CARIBOU L.
@! TO

;o LITTLE SACHIGO L.

ie

Gy ‘
POSTELNIC

25 50 75 IOOKM
& LAKE

ICKLE LAKE
EA

PICKLE | AR
LAKE ‘4

FIGURE 1. Sites surveyed in northwestern Ontario and
northeastern Manitoba.

MCLAREN AND MCLAREN: ABUNDANCE OF BIRDS 419

factors. Because we recorded all birds detected, we
express our results as ‘birds/km” rather than as
‘males/km2’. Nevertheless, the large majority of pas-
serines were identified as males (80%) and the indices
for most passerines represent pairs detected per km2.

Factors affecting detection of birds include weather
conditions, time of day, rate of progress, observer
ability and, especially, conspicuousness of the bird
species. Most of our surveys were conducted in the
early morning and under similar weather conditions.
We attempted to keep our rate of progress constant
although transects with large numbers of birds were
slower (because recording took longer) than transects
with fewer birds. In addition, progress was probably
slower in dense habitats (e.g. deciduous scrub) than in
open habitats, even though we deliberately walked
slowly in the latter. Conspicuous birds, in this study,
are those that vocalize frequently and loudly. Since
the majority of birds recorded were heard but not
seen, even among the quiet species, the ‘openness’ of
the habitat should not have affected detection rate
greatly. Because detection rate is unknown but
undoubtedly differs among species we do not compare
the relative indices of abundance among or between
species but only among geographic areas or among
habitats for individual species or all birds combined.
Similarly, because the indices presented are not exact
and represent only minimum densities we consider
only relatively large differences to be real.

The 55 sites where surveys were conducted were
grouped into six geographic regions (Figure 1). These
regions were chosen partly on the basis of geographic
proximity and partly on the basis of similarity of
habitats at the sites within each group (Figure 2). All
sites were within the boreal forest region (Rowe 1972);
the three southern groupings were in predominantly
continuous forest, whereas the northern three were in
a zone of forest interspersed with barrens, although
most sites sampled on the Tyrrell Sea Beach were in
forest.

Habitat Descriptions

The work presented here was part of a study of bird
populations in a much larger area stretching from the
Canadian high arctic to central Ontario. Because of
the wide latitudinal range in the larger study area,
habitat classification was based on the very general
scheme of Fosberg (1967). The primary characteristic
in that scheme is the degree of openness of the canopy
plants. Secondary and tertiary characteristics are
height of woody vegetation (forest, > 5m; scrub,
0.5 m-5 m; heath, < 0.5 m; and tundra) and physiog-
nomy (needle-leaved vs. broad-leaved plants).
Because intrusions of physiognomically different
plants can affect bird communities (Erskine 1977), we
also recognized a ‘mixed’ category defined as having

420

100

aN

75

Wes

in
-

S70 SIE Dor |
ONIFE RO oN

VAWS

ay

50

~~.
2

\

VoD
ZA Aen
\

LZ
N

7

SNK

25

PERCENT OF AREA SURVEYED

‘e) Bene
PICKLE NORTH CARIBOU L. WETIKO
LAKE TO HILLS

AREA LITTLE SACHIGO L.

THE CANADIAN FIELD-NATURALIST

‘

Vol. 95

Ke KE
KO xy

*
\\
x

0

SS

x
XY

y;
RY
OX)

RY
Y

a4,
RAXKX®
AX RX RY
wy
Wy
iy

EK
,

‘

*

QY

.
%)
XxXY

es

0%
0
X)
X

/
£5
9
XX)

\

9,

O)
oe

/ CX
XY
X

Z a
()
(x)
xX
1,

0)
O

KKK)
rani
WK
wy
WX
dy

aK KX)
AXRY
YK
ry

9
0

eK)
Wo
a
)
‘
‘
XX
xX
Wed

S,
)
\
MY
4
¥
M6

x
*
9,
VN
it
y

V/

X
xX
0
wy)

KX)
tone
aC
y
relat

‘'
~
N

A

Q

= 7,4

INDY ATL

hs~ ase N
~

or

KH
WI”
DENG,
<n
x

ss
Bali

NIN
(

\\

EeSU Tacs
AN INE
> N77> J4n, 7s

HUDSON BAY

TYRRELL FOREST-
LOWLANDS SEA TUNDRA
BEACH TRANSITION

FIGURE 2. Distribution of habitats in northwestern Ontario and northeastern Manitoba based on habitat sampling during

transects.

ratios of 30:70 to 70:30 deciduous to coniferous vege-
tation in the canopy. In order to increase the sample
size in individual habitats we combined closed and
open canopy categories for each height and physiog-
nomy grouping. Within any region, a given habitat
was usually either closed or open, so this combination
is not seen as contrary to the classification scheme.

We identified 9 general habitats that comprised
99% of the area surveyed. Deciduous forest occurred
in the southern three regions (Figure 2) and was char-
acterized by Trembling Aspen (Populus tremuloides)
with occasional White Birch (Betula papyrifera).
Coniferous forests occurred in all six regions and were
primarily Black Spruce (Picea mariana). Coniferous
forests with understory averaged 45% understory
cover and without understory averaged 15% cover.
Understory, when present, was about equally divided
between coniferous (young coniferous trees) and
deciduous [Alder (A/nus spp.) and willow (Salix
spp.)] vegetation. Coniferous forest with understory
was generally an open canopy forest; all other forest
habitats were closed canopy except for coniferous
(lichen) forest on the Tyrrell Sea Beach.

Coniferous scrub was composed primarily of Black
Spruce and occasionally (north to the Wetiko Hills)
Jack Pine (Pinus banksiana). Deciduous scrub was
mainly alder and willow but occasionally Dwarf Birch
(B. glandulosa). Both mixed forest and mixed scrub
were composed of the same species as the respective
pure deciduous or coniferous habitats.

Typical plants in heath were Labrador Tea (Ledum
groenlandicum), Ledum decumbens, Leatherleaf
(Chamaedaphne calyculata), blueberry (Vaccinium
spp.) and Dwarf Birch. In the northern part of the
study area heath tended to be open with a ground
cover of grasses (Gramineae) or sedges (Cyperaceae)
whereas farther south there was little gramineous
vegetation. Tundra occurred only in the extreme
north of the study area and was dominated by dwarf
shrubs such as Crowberry (Empetrum nigrum) and
Alpine Azalea (Loiseleuria procumbens), by sedges,
or by a combination of dwarf shrubs and sedges.

A dominant feature of the boreal forest is the pres-
ence of muskeg. Muskeg supports a variety of vegeta-
tion types ranging from sedge to spruce forest but all
are characterized by the presence of organic soils

1981

(Radforth 1969). Although we did not distinguish
muskeg habitat from similar habitats on mineral soils,
many of our survey sites, especially in the North Cari-
bou to Little Sachigo Lake area, the Hudson Bay
Lowlands and the Forest-Tundra Transition, were in
muskeg.

Mixed and deciduous forest occurred only from the
Albany River to the Wetiko Hills. Mixed and decidu-
ous scrub habitats were encountered more frequently
in the three southern areas than in the north. Scrub
habitats (including coniferous scrub) in these south-
erly areas tended to consist of young regeneration
(after fire) rather than stunted forest which dominates
the northwestern Hudson Bay Lowlands (Rowe
1972). Coniferous forest with understory was a rela-
tively common habitat throughout the study area. On
the Wetiko Hills, the Tyrrell Sea Beach and, toa lesser
extent, in the Pickle Lake area, coniferous forest with
understory tended to occur on well drained uplands
whereas in the other three areas it tended to be on
muskeg. Coniferous forest without understory was
encountered only in the two southern areas, where the
ground cover was usually moss, and on the Tyrrell Sea
Beach, where ground cover for all habitats was lichen.

Heath occured primarily in the Hudson Bay Low-
lands, where it was encountered in about equal pro-
portions of open muskeg and regeneration after fire
on mineral soil, and in the Forest-Tundra Transition.
Heath and tundra together comprised almost half
(Table 1) of the habitat surveyed in this area.

Very small amounts of three other habitats
occurred on the transect strips — grassy marsh, lake
and recently burned forest with little or no living
vegetation.

Spruce Budworm (Choristoneura fumiferana),
which may markedly affect densities of birds in the
boreal forest (Kendeigh 1947; Erskine 1977), was not
conspicuously abundant at any of the sites surveyed.

Results

Lowest abundance indices were recorded in conif-
erous forest without understory and heath (Table 1).
All of the high densities, in deciduous forest, decidu-
ous scrub and mixed scrub, were based on very small
samples. There was considerable variation in these
indices within one habitat type among different areas.
For example, the abundance index in coniferous
forest with understory in the Wetiko Hills was much
higher than in any other region. Similarly, the abun-
dance index for mixed scrub in the Hudson Bay Low-
lands, based ona very small sample, was much higher
than the other values for this habitat (Table 1).

Table 2 presents abundance indices of all species
that were recorded during the surveys. Many species,
particularly some of the non-passerines, were detected
very rarely, and their occurrence in a given habitat

MCLAREN AND MCLAREN: ABUNDANCE OF BIRDS

421

should not be interpreted as a preference for that
habitat. It is readily apparent from the table that none
of the more common species was restricted to only one
of even the very general habitat types considered, and
that many species occur in four or more different
habitat types.

Nevertheless, each habitat did support a generally
distinct combination of bird species. The species most
representative of deciduous forest (i.e. highest abun-
dance indices) were the Least Flycatcher, Red-eyed
Vireo and Ovenbird, although Ruffed Grouse and
Yellow-bellied Sapsuckers were also typical. The spe-
cies most representative of spruce forest with or with-
out understory, were the Gray Jay, Ruby-crowned
Kinglet and Yellow-rumped Warbler. The Golden-
crowned Kinglet was more abundant in mixed forest
than in spruce forests but was, nevertheless, found
only in habitats containing spruce trees. The Black-
poll Warbler was the most abundant species in spruce
scrub but the Palm Warbler, Rusty Blackbird, Dark-
eyed Junco and, toa lesser extent, Lincoln’s Sparrow,
were also common. In deciduous scrub, Tennessee
and Wilson’s warblers and White-throated and
White-crowned sparrows were the most abundant
species. Swainson’s Thrush was found in all forest and
scrub habitats but was considerably more abundant in
deciduous scrub than in the others.

The Lapland Longspur was the dominant species in
both heath and tundra. Other common tundra species
included the Horned Lark, Common Redpoll and
Savannah Sparrow. Common heath species included
the Tennessee Warbler, Blackpoll Warbler, Rusty
Blackbird, Savannah Sparrow, Dark-eyed Junco and
Tree Sparrow. No species was more abundant in
heath than in any other habitat.

Concomitant with changes in vegetation, the com-
position of the avian community changed with
increasing latitude. Figure 3 summarizes the changes
in species composition and shows graphically the
gradual disappearance of species associated with
deciduous and mixed forests (e.g., Red-eyed Vireo,
Nashville Warbler, Ovenbird) and the appearance,
first of species of low coniferous forests (e.g., White-
crowned Sparrow, Tree Sparrow) and then of species
of treeline and tundra (e.g., Harris’ Sparrow, Lapland
Longspur). Interestingly, although the abundance of
most species varied considerably over the study area,
the overall density of birds remained quite similar
over a north-south distance of approximately
1 000 km (Table 2).

Discussion

The indices reported here are not directly compara-
ble with most previous quantitative studies of bird
populations in the boreal forest (e.g., Ward 1975,
Erskine 1977) because of differences in survey method

THE CANADIAN FIELD-NATURALIST Vol. 95

422

"(1X9] 998) syeIIQeY J9YI0 JO sjUNOWe [[BUIS Jo 3oUasaId 9Yy} Jo asnedaq [e10} 243 0) dn ppe skemye jou Op saoueIsSIC,

(9°€Z)
O€l

(S11)
p71

(p°97)
I8I

(L°91)
L8I

(60£)
zal

(9're)
9TI

1 TIVYdAO

ny

(p°€1)
rol
(p°L)
86

(69)
07

(9°0)

+

SW

(0°97)
Lrl

(08)
OPT
(90)

+

(Lv)
9ST

(S°0)

+

(ol)

+

(€1)
$6

SO

(pp)
€Le
(10)

+

(v0)

+

(8°0)

+
(8°0)
+
(L'1)
SIZ
(9°0)

+

sd

(6'S1) (9°02) (S 9p) (6p) TIV
orl 98 Shl 061 -UAAO
(€°0) (1'€)
— + 607 - Lid
(10>) (8°€) (L°9)
f SIT LEI ad asl
(9p) (10>)
= — 8sI + 14H
(9°9) (10) (p's) (€"€)
671 + IbZ Il HM
(LZ) (L's) (€°91) (10>)
Iv €8 871 + SION
(S°9) (9°01) (S01) (¢1)
£91 LL 901 VIZ Id

AW dO NAO Ad

‘PpoAdAINs UY IIe sosayuored ul sloquinyy ‘yUaseid a1aM SPIIq Jey

SO}BIIPUI ,+4, {UIY | > JO SAIAINS IO} Paje[No[vd JOU 919M SIdIPUT "VIPUN] = NY] ‘WIwIH = aH “QnJ9g paxlp = SW ‘Qnos snolajuoD = So ‘qniosg snonpidag = Sq
4Sa10J Paxil] = AW ‘AlojsJapuy] NOYIM 4S910,J SNOIIJIUOD = FD ‘AIOISIOPUL] YIM JS910.4 SNOIAJIUOD = QAO ‘S204 snonpioeq = 4C ‘| 2NB17 99s ‘uoIpIsUueIL
eIPUN | -IS2IOF = LL ‘Yoeag eas [Josh], = {SL ‘spuejmoy Aeg uospny = TAH ‘SIH OXIA = HAA ‘OAC OF1YOeS 9]9317] 01 BY] NOQIUIeD YON = STON ‘O48
BAH = Td) ‘eqowuepy usa}svayiOU puv OLIeJUG U19}saMYIIOU UI $}ezIQeY OUIU PU svaIe XIS Ul (QdeJPlog) (;WIy/Spsiq) 9DUePUN|E Jo SIdIPU [[eI9AO “| ATAVL

1981 MCLAREN AND MCLAREN: ABUNDANCE OF BIRDS 423

TABLE 2. Indices of abundance of birds (birds/km2) in six areas and nine habitats in northwestern Ontario and northeastern
Manitoba. Area and habitat designations are as in Table 1. ‘+’ indicates species present but not recorded on the transect strip.
Numbers in parentheses are either the number of sites or the number of sites at which the habitat type was recorded.

Area Habitat
PL NCLS WH HBL TSB FIT DF CFU CF MF DS CS MS He Tu

Species (14) (12) (7) (I (4) (7) C1) (45) (21) (23) (12) (32) (15) (19) (3)
American Bittern

Botaurus lentiginosus — ] + <1 1 —-— —
Canada Goose Branta canadensis — + — + — ] Se ee ae eee
Mallard Anas platyrhynchos + + + <1 —- — —- —- ~—+ —- —- ~~ ~—- S| =
Green-winged Teal Anas crecca —- = 1 Se ey ee ran cas] —
Blue-winged Teal Anas discors — <I — ] —- — — 1+ ~~ ~ ~ ~ ~ —
American Kestrel Falco sparverius — + + — 1 —- — + <1 +
Spruce Grouse Canachites canadensis | ! ] — ] SS SIE 3 — — | ~—> —
Ruffed Grouse Bonasa umbellus ! 1 1 <li — — 6 <I — 1! ~—~ | ~ ~— —
Common Snipe Capella gallinago <1 1 + 2 + Sl = <b <h == + — —
Whimbrel Numenius phaeopus S— Fe 2 Se heey Caen os) Mee aS)
Solitary Sandpiper Tringa solitaria = + P Zs it — <I — — — 1 —- —
Greater Yellowlegs

Tringa melanoleuca ea Se) aie 4 Sle Sle, Se SS SSF Sa ae
Lesser Yellowlegs Tringa flavipes — + — 5 2 4 — <I 1 — — 5 jn ere)
Least Sandpiper Calidris minutilla ites eer 2 — 3 Sl Ss SS SS ae 2

Short-billed Dowitcher

Limnodromus griseus —- -— — <1 — ] eS SS SD
Stilt Sandpiper

Micropalma himantopus — —- -—- =—- —+ =
Northern Phalarope Lobipes lobatus —
Bonaparte’s Gull Larus philadelphia + —- — +
Short-eared Owl Asio flammeus ee a ee ae
Belted Kingfisher Megaceryle alcyon
Common Flicker Colaptes auratus + — !
Yellow-bellied Sapsucker

Sphyrapicus varius 2 _ 2 —- —- — 6) i eS 2 6 oS Se
Black-backed Three-toed

Woodpecker Picoides arcticus — ] 1 <!) — ~— ~ ~ —~ <1 —~ ~—~ — 1 =
Yellow-bellied Flycatcher

Empidonax flaviventris 8 7 10 3 5 3 GO 1eOerg oS 7 ale a) 2 —
Alder Flycatcher

tr Empidonax alnorum 2 ] y 5) — <I 2 See Pee al 9 —- —
Least Flycatcher

Empidonax minimus 8 2 8. 31" =
Olive-sided Flycatcher

Nuttallornis borealis <1 ! + + —_- — F
Horned Lark Eremophila alpestris — —_ 4 |
Tree Swallow /ridoprocne bicolor | — = F
Gray Jay Perisoreus canadensis 3 11 6 Ss 13 2 — 12 5 ne)

+ + _— + + a=
| 5 2

+
|
+
+
---|
|
|
\
—_
|
|
|

|
oe
N
Ww
N
lon)
S
|

|

Common Raven Corvus corax
Boreal Chickadee Parus hudsonicus
Red-breasted Nuthatch

Sitta canadensis l
Brown Creeper Certhia familiaris 3
Winter Wren Troglodytes trogloydtes 1
Hermit Thrush Catharus guttatus ]
Swainson’s Thrush

Catharus ustualtus 8 8 J Sil ho Si NG SAO 4 <p ka ea
Gray-cheeked Thrush

Catharus minimus —- — — <i l 5 — t|—_—— + 2 3 2 —
Golden-crowned Kinglet

Regulus satrapa 5 2 3 —- —- — — 1 GT

424 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 2 (Continued)

Area Habitat

PL NCLS WH HBL TSB FIT DF CFU CF MF DS CS MS He Tu
Species (14) G2) (7) (1) (4) (@~ (QI) (45) (21) (23) (12) (32) (15) (19) (3)
Ruby-crowned Kinglet
Regulus calendula
Waxwings Bombycilla spp.

9 8 2 5 TS —
|
Northern Shrike Lanius ludovicianus — = =
1
5

1 + 6
— — 1 + — 6 + 1 —
1 ee (a —- — — <1l~—~ ~ —
— — — 4 — 1 3 — <1 1 —- —
10 — — — 32 Poh 9 ar 3 — —

Solitary Vireo Vireo solitarius
Red-eyed Vireo Vireo olivaceous
Philadelphia Vireo

Vireo philadelphicus <1 — 2 - —- ~~ ~~ ~ ~ ~ —
Black-and-white Warbler

Mniotilta varia PPS) eI ce Sel Win Rees 4 <1 — 1 HW — 122 — —
Tennessee Warbler

Vermivora peregrina 1 10 40 12 #15 3 10 16 1 14 5! 6 10 6 —
Orange-crowned Warbler

Vermivora celata <1 1 ! 2 1 —- — 1 <1 — § 2 1 —-— —
Nashville Warbler

Vermivora ruficapilla 5 1 4 —- —- — Bay Aa I bees) ] 7 — 4 — —
Yellow Warbler Dendroica petechia + 2 ] 2 —- — 8 <I — <1 11 — 1 — —
Magnolia Warbler

Dendroica magnolia 5 3 7 1 —_— = 4 See 5) 5 1 10 — —
Yellow-rumped Warbler

Dendroica coronata 5 10 6 2 10 3 — 10 10 4 7 PS
Black-throated Green Warbler

Dendroica virens 1 + <1 — 1 -— ~ ~ ~~
Chestnut-sided Warbler

Dendroica pensylvanica <b SS aS ee SS SS 2 SS aS ee
Bay-breasted Warbler

Dendroica castanea 3 2
Blackpoll Warbler Dendroica striata + 26 nt) — 12 2

!

nah
ox
|
|
|
BSS

Palm Warbler Dendroica pailmarum + 1 12 + 1 — 4
Ovenbird Seiurus aurocapillus Taal O88 —- — — 24 1
Northern Waterthrush

Seiurus noveboracensis <1
Connecticut Warbler Oporornis agilis | | — 1! <I ~ ~—~ | ~~ ~—~ —
Mourning Warbler

Oporornis philadelphia 1 2. IS es Se Se
Common Yellowthroat

Geothlypis trichas <1 —
Wilson’s Warbler Wilsonia pusilla 2 4 | 4 l — 2
Canada Warbler Wilsonia canadensis < | —
Rusty Blackbird Euphagus carolinus | 2 + 9 ]
Pine Grosbeak Pinicola enucleator — — — | 4
Common Redpoll Carduelis flammea =
Pine Siskin Carduelis pinus 2 ! + ar —
White-winged Crossbill

Loxia leucoptera — -— —
Unidentified Crossbill Loxia sp. 1 ] it l ie Se SSB I << ee
Savannah Sparrow

Passerculus sandwichensis + =
Dark-eyed Junco Junco hyemalis 3 5 fl

2 !

i)
BN
|
/\
BN
+

|
~
N
Ww
|

|

Tree Sparrow Spizella arborea
Chipping Sparrow Spizella passerina
Harris’ Sparrow Zonotrichia querula
White-crowned Sparrow

Zonotrichia leucophrys —- — — 18 4 wy) Sy E28 OI HO) anaes
White-throated Sparrow

Zonotrichia albicollis 2 5 4 6 3 + 2 ay) ot lies 2 owls Bin) at
Fox Sparrow Passerella iliaca Laer] 2 3 + 2 2 1

h— NAMA

1981

TABLE 2 (Concluded)

MCLAREN AND MCLAREN: ABUNDANCE OF BIRDS

425

Area Habitat
PL NCLS WH HBL TSB FIT DF CFU CF MF DS CS MS He Tu
Species (14) (2) () (dQ) 4@ @~ d)z 45) (1) (23) (12) (32) (15) (19) ©)
Lincoln’s Sparrow Melospiza lincolnii + ] ] 12 3 ] Se De ee 5 1 — —
Song Sparrow Melospiza melodia + + — ] ee ee ee gs S| ey ie
Lapland Longspur .

Calcarius lapponicus 0) ee en eee) AeA ()
Sritnismleonesput Calcarius picts. == = a= ares)
All passerines! 117 119 179 162 115 114 175 140 79 137 365 132 204 84 98
All birds! 126 124 187 181 124 130 190 145 86 146 373 147 220 104 130
% passerines 93 96 96 90 93. 88 92 97 92 94 98 90 93 81 72

'The totals exceed the sums of the individual densities because records of unidentified birds are not included in the species list.

and in habitat classification. Densities derived from
plot surveys (Erskine 1977) or narrow transects (Ward
1975) tend to be similar and higher than densities
derived from wide transects (LGL Ltd. unpubl. data).
The most nearly comparable data are those of James
(1980) in the Pickle Lake area. Using one observer and
a 50 m wide transect, he recorded an average index of
155 pairs/km2, a value not greatly different from our
results in the same area.

Although quantitative comparisons are not possi-
ble, a number of qualitative similarities between our
results and those of other workers are evident. Gilles-
pie (1960) and Carbyn (1971) both surveyed plots that
correspond in habitat type to our coniferous forest
without understory and both reported their lowest
densities on these plots. We also found the fewest
birds in this habitat. Considerable variation in the
calculated densities in habitats corresponding to our
coniferous forest with understory were also noted by
other workers (Gillespie 1960; Carbyn 1971; Ward
1975). Reasons for such variation are not apparent,
but may reflect, at least partly, both the amount and
the type of undergrowth in the forest.

Most of our samples of deciduous scrub were very
small and open to sampling error. Nevertheless, the
very high relative density we found is similar to the
pattern seen in other studies. Salter and Davis (1974),
Tull et al. (1974), Ward (1975) and Wisely and Tull
(1977) all surveyed several habitats in the Mackenzie
Valley and found that relative densities were highest in
scrub habitats with a deciduous component. Ward
(1975) found, as we did, that in scrub habitats relative
densities were highest in deciduous scrub, lowest in
coniferous scrub and intermediate in mixed scrub.
Gillespie (1960) also found very high densities in
deciduous scrub along a stream near Churchill,
Manitoba. He reported that densities in coniferous
forest and scrub were only 28% to 39% of the density
in deciduous scrub. Our results are remarkably sim-
ilar; minimum densities in coniferous forest and scrub

varied from 23% to 39% of those in deciduous scrub.

Few species of birds were confined to only one of
the general habitats distinguished in this study. Sev-
eral of the habitats included a common vegetational
feature (e.g. coniferous trees) and it is possible that
birds were responding to the presence of this single
feature rather than to the broad habitat mosaic. On
the other hand, some species are not limited to a
particular type of vegetation. McLaren (1975) found
that several wood warblers in central Ontario demon-
strated considerable latitude in the vegetation used for
foraging and nesting. The blurring of avian habitat
preferences as a consequence of broadly-defined habi-
tat types may be an unavoidable aspect of transect
surveys where the likelihood of encountering substan-
tial amounts of pure and finely-defined habitats is
small.

Nevertheless, the species that we found to be typical
of the habitats we encountered are in fairly close
agreement with Erskine’s (1977) conclusions about
boreal bird communities and our data show that these
representative species are likely to be found even in
relatively isolated outposts of particular habitats. For
example, deciduous and mixed forest along the
Wetiko Hills are rare habitats in this part of the boreal
forest (Rowe 1972), yet several species previously
reported only from areas 100 to 150 km farther south
occur there in minimum densities of 3-8 birds/km2
(e.g., Golden-crowned Kinglet, Black-and-white and
Nashville warblers, Ovenbird). The Tyrrell Sea Beach,
an ancient beach ridge, is dominated by well-drained
spruce forest generally taller than 7 m, and is sur-
rounded by extensive areas of black spruce muskeg
scrub generally less than 5 m tall. On the beach ridge
Ruby-crowned Kinglets occur in detected densities up
to 8 birds/km2 whereas detected densities in the sur-
rounding scrub are only 1-2 birds/km?2. Yellow-
rumped Warblers show a similar pattern of low densi-
ties in the scrub forests of the Hudson Bay Lowlands
and Forest-Tundra Transition and high densities

426 THE CANADIAN FIELD-NATURALIST Vol. 95

POSTELNIC LAKE
SEVERN RIVER
LITTLE SACHIGO LAKE

ALBANY RIVER

WETIKO HILLS
NELSON RIVER
CHURCHILL RIVER
SEAL RIVER
CARIBOU RIVER

PURPLE FINCH
COMMON GRACKLE
CHESTNUT-SIDED WARBLER
MOURNING WARBLER
BLACK-THROATED GREEN WARBLER
BROWN THRASHER

COMMON YELLOWTHROAT
CONNECTICUT WARBLER
PINE SISKIN

BROWN CREEPER
YELLOW-BELLIED SAPSUCKER
GOLDEN-CROWNED KINGLET
SOLITARY VIREO
RED-EYED VIREO

RUFFED GROUSE
NASHVILLE WARBLER
BAY-BREASTED WARBLER
OVENBIRD
BLACK-AND-WHITE WARBLER
MAGNOLIA WARBLER

LEAST FLYCATCHER
WINTER WREN

YELLOW WARBLER

TREE SWALLOW
OLIVE-SIDED FLYCATCHER
BELTED KINGFISHER
COMMON FLICKER

SPRUCE GROUSE

HERMIT THRUSH

ALDER FLYCATCHER
WHITE-THROATED SPARROW
CHIPPING SPARROW
RUBY-CROWNED KINGLET
RED-BREASTED NUTHATCH
SWAINSON’S THRUSH
YELLOW-BELLIED FLYCATCHER
BOREAL CHICKADEE
AMERICAN ROBIN

GRAY JAY

DARK-EYED JUNCO

RUSTY BLACKBIRD
YELLOW-RUMPED WARBLER
SAVANNAH SPARROW
COMMON RAVEN

NORTHERN WATERTHRUSH
COMMON SNIPE

GREATER YELLOWLEGS
BLACKPOLL WARBLER
TENNESSEE WARBLER
WILSON’S WARBLER
ORANGE-CROWNED WARBLER
FOX SPARROW

PALM WARBLER

LINCOLN’S SPARROW
LESSER YELLOWLEGS
WHITE-CROWNED SPARROW
LEAST SANDPIPER
GRAY-CHEEKED THRUSH
TREE SPARROW

HARRIS’ SPARROW
LAPLAND LONGSPUR
COMMON REDPOLL

HORNED LARK

WHIMBREL

FIGURE 3. Observed ranges of some commonly detected species in northwestern Ontario and northeastern Manitoba, based
on both on- and off-transect data.

1981

along the Tyrrell Sea Beach. This is apparently a
response to the height of the trees since these species
are as abundant in the muskeg forests between Pickle
Lake and Little Sachigo Lake as they are on the
Tyrrell Beach ridge.

Despite the general agreement between our results
and Erskine’s (1977), there are a few anomalies. Some
of the species Erskine reports as typical were not
found in large numbers (or at all) during this study,
probably because we were near the limits of (or
beyond) their breeding ranges. For example, the
American Redstart (Setophaga ruticilla) has been
reported as common north to about the Severn River
in our study area (James et al. 1976). However, we
recorded none. Erskine (1977) reported the Dark-eyed
Junco to be a typical species in deciduous scrub;
although we recorded it commonly, we found none in
this habitat. In fact deciduous scrub was the only
habitat other than tundra where juncos were not seen.
On the other hand, we found Wilson’s Warblers and
White-crowned Sparrows to be among the most
abundant species in deciduous scrub. Gillespie (1960)
also found White-crowned Sparrows abundant in this
habitat. That Erskine (1977) does not list these species
as typical of deciduous scrubis likely again a function
of range. Most of Erskine’s study plots were well to the
south of the range of these two species.

Acknowledgments

The work reported here was part of a larger study of
bird populations funded by the Polar Gas Project. We
thank L. Doran and B. Ross of the Polar Gas Project
for their logistic support. W. McKeever of Viking
Helicopters Ltd. piloted the helicopter.

The surveys were conducted by the authors and by
C. Holdsworth, W. G. Johnston, E. Nol, and W.
Renaud of LGL Ltd. We thank W. J. Richardson of
LGL Ltd. and L. Doran and J. Riddick of the Polar
Gas Project for their comments on the manuscript.

Literature Cited

Carbyn, L. N. 1971. Densities and biomass relationships of
birds nesting in boreal forest habitats. Arctic 24: 51-61.

Emlen, J. T. 1971. Population densities of birds derived
from transect counts. Auk 88: 323-342.

Emlen, J. T. 1977. Estimating breeding season bird densi-
ties from transect counts. Auk 94: 455-468.

Erskine, A. J. 1977. Birds in boreal Canada. Canadian
Wildlife Service Report Series 41. 71 pp.

Fosberg, F. R. 1967. A classification of vegetation for gen-
eral purposes. /n Guide to checklist for IBP areas. Edited
by G.F. Peterken. Blackwell Scientific Publishers,
Oxford and Edinburgh. pp. 73-120.

Gillespie, W. 1960. Breeding bird and small mammal popu-
lations in relation to the forest vegetation of the subarctic
region of northern Manitoba. Ph.D dissertation, Univer-
sity of Illinois, Urbana. 120 pp.

MCLAREN AND MCLAREN: ABUNDANCE OF BIRDS

427

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.

James, R. D. 1980. Notes on the summer birds of Pickle
Lake, Ontario, Ontario Field Biologist 34(2): 80-92.

James. R. D., P. L. McLaren and J. C. Barlow. 1976. An-
notated checklist of the birds of Ontario. Life Sciences
Miscellaneous Publications, Royal Ontario Museum. 75
Pp.

Jarvinen, O. and R. Viisanen. 1976. Estimating relative
densities of breeding birds by the line transect method. IV.
Geographical constancy of the proportion of main belt
observations. Ornis Fennica 53: 87-89.

Jarvinen, O., R. Vaisanen and Y. Haila. 1976. Estimating
relative densities of breeding birds by the line transect
method. III. Temporal constancy of the proportion of
mainbelt observations. Ornis Fennica 53: 40-44.

Jarvinen, O., R. A. Vaisanen and Y. Haila. 1977. Bird cen-
sus results in different years, stages of the breeding season
and times of the day. Ornis Fennica 54: 108-116. ;

Kendeigh, S. C. 1947. Bird population studies in the conif-
erous forest biome during a spruce budworm outbreak.
Ontario Department of Lands and Forests Biological Bul-
letin 1: 1-100.

Lee, D. L. 1978. An annotated list of the birds of the Big
Trout Lake area, Kenora District. Ontario Field Biologist
32: 17-36. :

McLaren, P. L. 1975. Habitat selection and resource utili-
zation in four species of wood warblers (Aves: Parulidae).
Ph.D, dissertation, University of Toronto, Toronto. 168
Pp.

McLaren, P. L. and M. A. McLaren. 1981. Bird Observa-
tions in northwestern Ontario, 1976-1977. Ontario Field
Biologist 35: 1-6.

McLaren, M. A. and W. E. Renaud. 1980. Some observa-
tions of birds in northeastern Manitoba — 1975-1977.
Blue Jay 38: 31-35.

Radforth, N. W. 1969. Muskeg as an engineering problem.
In Muskeg engineering handbook. Edited by I. C. Mac-
Farlane. University of Toronto Press, Toronto. pp. 3-30.

Rowe, J. S. 1972. Forest regions of Canada. Department of
the Environment, Canadian Forestry Service Publication
Number 1300. 172 pp.

Salter, R. E. and R. A. Davis. 1974. Surveys of terrestrial
bird populations in Alaska, Yukon Territory, Northwest
Territories and northern Alberta, May, June, July 1972.
Arctic Gas Biological Report Series 12. Chapter 2. 384 pp.

Tull, C. E., 1. D. Thompson and P. E. Taylor. 1974. Con-
tinuing surveys of terrestrial bird populations in North-
west Territories, Yukon Territory, and Alaska: June and
July, 1973. Arctic Gas Biological Report Series 29. Chap-
ter 3. 217 pp.

Ward, J.G. 1975. Continuing surveys of terrestrial bird
populations in the Mackenzie Valley, June, 1974. Arctic
Gas Biological Report Series 30. Chapter 4. 93 pp.

Wisely, A. N. and C. E. Tull. 1977. Ground surveys of ter-
restrial breeding bird populations along the Fort Simpson
realignment of the proposed Arctic Gas pipeline route,
Alberta and Northwest Territories, June, 1975. Arctic Gas
Biological Report Series 35. Chapter 6. 59 pp.

Received 28 April 1980
Accepted 25 March 1981

Status and Breeding Success of New Brunswick Bald Eagles

R. F. STOCEK! and P. A. PEARCE?

'Maritime Forest Ranger School, Fredericton, New Brunswick E3B 4X6
2Canadian Wildlife Service, Fredericton, New Brunswick E3B 4Z9

Stocek, R.F.,and P.A. Pearce. 1981. Status and breeding success of New Brunswick Bald Eagles. Canadian Field-Naturalist

95(4): 428-433.

Aerial surveys of New Brunswick for Bald Eagle (Haliaeetus leucocephalus) nests in 1974-80 showed 19 occupied sites, all but
one in the southwest of the province. The maximum noted in any year was 12. Fifty-three percent of breeding pairs produced
young, an average of 0.73 young per occupied site annually, fitting the pattern of marginal production in Atlantic Canada and
adjacent Maine. Past brood size of the eagle, never a common breeding bird in New Brunswick, appears to have been
comparable to the present. Many former breeding sites are still occupied. There was no relation between breeding success and
remoteness or accessibility of the nest, but coastal nesters seemingly were more successful than those in lake and river habitats.

Key Words: Bald Eagle Haliaeetus leucocephalus, breeding success, New Brunswick.

The Bald Eagle (Haliaeetus leucocephalus) occurs
in New Brunswick as both a resident and a transient
bird (Squires 1976). Declining populations have been
documented in several regions of the United States in
the last three decades (Broley 1958, Sprunt et al.
1973). In Canada, Godfrey (1970) regarded the species
as endangered, but the Committee On the Status of
Endangered Wildlife in Canada (COSEWIC) has not
recognized it as such. More recently, scattered
attempts have been made to assess regional Bald Eagle
populations in Canada, and large, healthy popula-
tions have been reported in Saskatchewan and
Manitoba (Whitfield et al. 1974, Leighton etal. 1979).
In New Brunswick, the bird is now protected under
provisions of both the provincial Endangered Species
Act and the Fish and Wildlife Act. The present inves-
tigation was undertaken to define the past and present
reproductive status of the Bald Eagle in that province.

Methods

Published and unpublished records of the breeding
occurrence and productivity of the Bald Eagle in New
Brunswick provided the data base on which the plan
field surveys. Sources of information included New
Brunswick Museum, the former Northeastern Wild-
life Station at University of New Brunswick, New
Brunswick Departments of Natural Resources and of
Tourism, federal Departments of Fisheries and
Oceans, Environment, and Indian and Northern
Affairs, and the Canadian Nature Federation.

Field work during 1974-80 consisted mainly of
aerial surveys, for identification of important nesting
areas and assessment of production. Most flying was
done in Cessna 172, 180, 336, and Piper Cherokee
aircraft, with helicopter support (Jet Ranger II, and
Huey and Kiowa military units), particularly in 1979
and 1980. The pilot and one other observer usually

428

constituted the crew, although as many as five
observers were used at one time. Flying altitude and
air speed varied between 35 and 65 m and 110 and
160 km/h, respectively.

During 1974a strip approximately 0.4 km wide was
surveyed along the entire coast, following the forest
edge, a distance of about | 600 km. Flying was under-
taken inland to search lake shores and rivers, includ-
ing those with known or suspected breeding sites.
Thirty-four hours of surveys were flown in that year.
During 1975, 1976, and 1977 only known and sus-
pected mainland nests were checked from the air.
Only ground checks were made in 1978. A more inten-
sive aerial search of lakes and rivers and the southern
coast was conducted in 1979, while all known and
suspected nests were again inspected in 1980.

Nests were checked from the air at least twice in
each breeding season, during incubation and again
when advanced nestlings were expected to be present.
Supplementary ground checks were infrequent and
often consisted of only one visit. Assessment of repro-
duction is based on terminology proposed by Postu-
palsky (1974), as follows: an area containing at least
one nest was termed a breeding site; a site was judged
to be occupied if two adults were present on or near
the nest, or if one bird was near a new or repaired nest;
and an occupied nest in which at least one young was
reared to an advanced stage of development was
deemed productive or successful.

A wilderness value was developed for each occupied
site in 1979, using a modification of the criteria and
method of Mathisen (1968). Each factor — human
activity, presence of roads, timber cutting, surround-
ing habitat, and nest visibility — was evaluated from
the air, ona scale of 0 to 4. All factors were averaged
for each breeding site. A score of 4.0 represented a site
that was remote and considered relatively free of

1981

potential human disturbance, while a value of 0.0
indicated a nest site that was very accessible and dis-
turbance prone. The wilderness value is somewhat
subjective but it allows quantification and compari-
son of breeding sites.

Results and Discussion
Breeding Occurrence and Density

The Bald Eagle seemingly was never a common
breeding bird in New Brunswick (Moore 1928). There
are records of only 30 nestings in 1900-49, and 75 in
1950-73, no more than five being noted in any one
year. Eagles regularly nested on Grand Manan Island,
in the Bay of Fundy (Herrick 1873, Pearsall 1879,
Boardman 1903, Pettingill 1939). Three or four pairs
continue to nest each year on the Canadian Islands
south of the Charlotte County mainland including the
Grand Manan archipelago. Nesting also occurred on
the mainland coast of Charlotte County (Chamber-
lain 1882) and in the St. Croix Valley (Boardman
1903). Today, two or three pairs still nest near the
coast, and the St. Croix River watershed has some
sites that have been occupied for over 25 years. No
breeding is known from the Bay of Fundy east of Saint
John.

In northern New Brunswick, the only early record
of nesting eagles is from Northumberland County
(Philipp and Bowdish 1977). Very few nests have been
seen in the northern counties of Victoria, Madawaska,
Restigouche, and Gloucester. The only recent occu-
pied site in that region was on the Tabusintac River,
Northumberland County, where eagles have nested
for 20 years. The large Miramichi watershed in that
county supported only a few nesting pairs, the last
being reported in the 1950s.

The estuary of the St. John River up to Fredericton
contains preferred nesting habitat for Bald Eagles.
They are known to have nested there since at least
1900. A cliff nest was reported at Upper Greenwich,
Kings County, in 1911 (Townsend 1912), and a nest at
the mouth of Noonan Stream, Sunbury County,
about 1920 was said to have been occupied for many
years (N.E. Wildlife Station files). There were four
occupied sites in the estuary in the 1970s. Between
1950 and 1980, 36 occupied eagle nests involving at
least 13 breeding sites, were recorded for that part of
the river, although up to 100 birds summered there in
1949-50 (Wright 1953). That summering population
was composed largely of the southern subspecies (H. /.
leucocephalus) (Squires 1976), and has since declined
(Stocek 1979).

The distribution of nest sites in New Brunswick in
1974-80 is shown in Figure 1. All but one were in
southwestern New Brunswick between the estuary of
the St. John River and the border with the State of

STOCEK AND PEARCE: NEW BRUNSWICK BALD EAGLES

429

Maine. During our investigation, 19 occupied sites
were found in that part of New Brunswick. A density
of 0.11 occupied sites per 100 km? was estimated for
that region in 1980. Other published estimates of eagle
nest density in eastern Canada included 0.08 per
100 km? for western Labrador in 1970-73 (Wetmore
and Gillespie 1976), and 0.77 occupied sites per
100 km? in Cape Breton Island, Nova Scotia (Stocek
1980). Three-quarters of the occupied sites in New
Brunswick were near lakes or along rivers and
streams, including the Tabusintac, Kennebecasis, St.
Croix, St.John, Mactaquac, Canoose, and Porto-
bello. Most of the remainder were on coastal islands.
Province-wide, 22 occupied sites have been identified
since 1970. During any one year, the maximum
number was 12 (0.02 per 100km2). Since 1950, Bald
Eagles have nested in all counties except Kent, West-
morland, Albert, and Saint John. Only two breeding
sites are known in neighboring Maine above 46°N
latitude (C.S. Todd and R.B. Owen, unpublished
data). We found only one breeding site above that
latitude in New Brunswick.

To determine the amount of available nesting habi-
tat used by Bald Eagles we measured the shoreline of
major lakes and rivers. Primary and secondary habi-
tat were identified, for purposes of comparison of nest
density, according to criteria of Leighton et al. (1979),
i.e. primary shoreline comprised lakes 11 km or
greater in circumference and rivers which were drawn
as two distinct lines on 1:250 000 National Topogra-
phic Series maps. We searched | 100 and 2 100 km of
total lake and river shoreline, respectively. That
included 98% of primary lake habitat and 60% of
primary river shoreline in the province. The five
southwestern counties contain 80% of the province’s
total lake area, all the larger lakes being found there,
and 30% of all the rivers, by length. The density of
breeding sites is given in Table 1. The greater use of the
southwestern part of the province and the preference
for lake habitats are clearly indicated. The apparent
lack of discrimination between primary and secon-
dary shoreline suggests that the criteria used in their
identification may not be applicable at the low densi-

TABLE |. Density of Bald Eagle breeding sites in New
Brunswick, by primary and secondary shore habitat

Number of breeding sites/ 100 km shoreline

Southwestern
Whole province counties
Habitat primary secondary primary secondary
Lake 0.76 0.71 0.93 0.88
River 0.41 0.26 0.80 0.74
Total 0.52 0.45 0.86 0:82

430

THE CANADIAN FIELD-NATURALIST

Vol. 95

NEW BRUNSWICK

Ficure 1. New Brunswick, showing locations of Bald Eagle nest sites, 1974-80.

ties of eagles prevailing in New Brunswick. Marine
breeding areas were found only on the southwest
coast, where breeding site density was 1.39/100 km
shoreline, compared with 0.38/ 100 km for the total
provincial coastline. On the outer islands, the breed-
ing site density was 2,.23/100 km shoreline. In con-
trast, breeding site densities of eagles were as high as
5.79/100 km primary shoreline in Saskatchewan in
1974.

Breeding Site Characteristics
Bald Eagle nests were usually constructed in the

upper crown of a dominant tree in the forest or ina
large, solitary tree on intervale land. Nest habitats
ranged from flooded hardwood stands in lowland
river valleys to mixed stands on steep hillsides. Seven-
teen of 25 nests were built in hardwood trees. All nests
in softwoods were in white pine. Twenty-one of 25
nest trees were living. We know of no recent cliff nests
in New Brunswick. Gittens (1968) reported that 24 of
the 50 nests he investigated in Nova Scotia were in
hardwoods, especially aspen, and 16in white pine. He
also noted that four of 50 nests of Cape Breton Island
were on cliffs.

1981

Of 26 nest sites we studied, 13 were within 100 m of
water and only two were more than 500 m from water.
Four of the inland nests were located on small lakes or
streams near major bodies of water rather than on the
larger waterways themselves. Seven of 18 breeding
sites had alternate or supernumerary nests, ranging up
to 5 kmaway. Based on years of occupancy and breed-
ing history at the site, we considered those distant
nests to be parts of the same breeding sites. One nest
tree contained two nests, only one occupied at a time.

The wilderness value scores of 11 sites occupied in
1979 ranged from 1.2 to 4.0. There appeared to be no
relationship between the success of a nest and the
wilderness rating of the site. The unsuccessful and
successful sites had mean scores of 3.5 and 3.1, respec-
tively. The two most remote sites failed to produce
young, and the site most susceptible to actual human
activity was successful. However, because of typically
adverse weather in New Brunswick during early
spring when Bald Eagles start breeding, human activ-
ity on lakes and rivers is often not underway until
June; by then eggs have hatched and the chances of
nest desertion are reduced, although snowmobiling
and winter cutting may create disturbance in some
areas earlier in the breeding cycle.

Breeding and Productivity

Bald Eagles start moving to nesting areas in New
Brunswick in February and March. Incubating birds
have been noted as early as 25 March and eggs have
been seen from 6 April to 16 May. On Grand Manan
Island, Herrick (1873) reported finding eggs with well-
developed but frozen embryos on 20 February. Peak
egg laying occurs in early April. Information in the

STOCEK AND PEARCE: NEW BRUNSWICK BALD EAGLES

431

Maritimes Nest Records Scheme indicates that the
mean clutch size of the Bald Eagle in the Maritime
Provinces, based on 13 full clutches, was 2.15 + 0.55 -
1 x le, 9 x 2e, 3 x 3e. Our investigation showed that
nests contained young from 23 May to 10 August. We
found 19, nine, and one, nests with one, two, and three
young, respectively. Peak fledging takes place in late
July. In Nova Scotia, eggs hatched between 10 and 25
May and young fledged between 15 and 30 July (Git-
tens 1968).

Figures for eagle productivity are shown in Table 2.
During our study, 53% of the breeding pairs were
successful. The average number of young produced
annually was 0.73 and 1.38 in occupied and successful
nests, respectively. Data on 17 successful nests in
1960-73 gave an average fledgling brood size of 1.43.
Sprunt et al. (1973) estimated that at least 50% of
breeding pairs should rear young, with a mean of 0.7
young yearly per occupied nest, to maintain a stable
population. Only 60% of the breeding pairs in New
Brunswick were successful at least half of the time.
Generally poor nesting success has resulted in low
annual production of young. Information on the
breeding success of eagle populations elswhere in
Atlantic Canada and in Maine (Table 3) indicates that
New Brunswick fits the regional pattern of marginal
production.

Broods of two young were seen in 9 of 29 successful
nests. Two young also were noted in about one-third
of successful eagle nests in Nova Scotia (Gittens 1968)
and in Maine (C.S. Todd and R.B. Owen, unpub-
lished data). In the larger, more productive popula-
tions in Saskatchewan and Manitoba (Whitfield et al.

TABLE 2. Productivity of Bald Eagles in New Brunswick, 1974-80!, by year and habitat

Number of breeding sites

ellenes) Young raised /nest
Year e Examined through)? Successful Occupied Successful
Year
1974 11 10 (8) 4 0.63 1245)
1975 12 10 (9) 3 0.56 1.67
1976 12 9 (5) ] 0.20 1.00
1977 12 8 (7) 5 1.00 1.40
1978 6 6 (3) 2 1.00 1.50
1979 17 12 (11) 6 0.64 1. 7/
1980 18 12 (12) 8 1.00 1.50
Habitat
Coastal 15 11(6) 6 1.66 1.66
Lake 40 33(27) I] 0.48 1.18
River 33 23(22) 12 0.77 42

'No aerial surveys were conducted in 1978.
2Not all occupied nests could be followed through to success or failure.

432 THE CANADIAN FIELD-NATURALIST Vol. 95
TABLE 3. Bald Eagle breeding success in Atlantic Canada and adjacent Maine, U.S.A.
Number of :
occupied breeding % Young raised/nest
Area (Source) Period sites Successful Occupied Successful
Labrador 1970-73 24 45 0.60 1.20
(Wetmore and Gillespie 1976)
Maritime Provinces 1975 28! 54 0.80 1.53
(Stocek and Pearce, unpublished
data)
Nova Scotia 1968 23 54 0.72 1.36
(Gittens 1968)
New Brunswick 1974-80 67 53 0.73 1.38
(this study)
Maine 1977-78 112 39 0.60 1.52

(C.S. Todd and R.B. Owen,
unpublished data)

‘Includes 9 sites in New Brunswick also used in this study.

1974), northwestern Ontario (Grier 1974), and Alaska
(Sprunt et al. 1973), two young were also produced in
about one-third of successful attempts. Almost 80%
(67/88; Table 2) of the breeding sites examined were
occupied during our investigation. Five of 13 sites
occupied in 1974-75 were no longer being used in
1980.

Birds nesting in marine environs appeared to be
more successful than those occupying lake and river
breeding sites (P< 0.01, t-test) (Table 2). Breeding
success in freshwater habitats showed no significant
difference. A habitat-related disparity in breeding
success was also mentioned by C.S. Todd and R.B.
Owen (unpublished data) for eagles in Maine in 1977-
78: they credited birds on the coast and inland lakes
with greater success than those in river habitats.

Acknowledgments

Thanks are expressed to J. C. Baird and the late
B. S. Wright for their assistance, and to biologist-pilot
A. Boer who gave freely of his time, often on short
notice. The authors are particularly indebted to the
4224 Tactical Helicopter Squadron, Canadian Forces
Base Gagetown, for permission to travel with the unit
on training missions. Financial support was provided
by Canadian Wildlife Service, New Brunswick
Departments of the Environment and of Natural
Resources, World Wildlife Fund, Canadian Wildlife
Federation, Samuel and Saidye Bronfman Family
Foundation, Quebec-Labrador Foundation, Inc.,
Tractors and Equipment 1962 Ltd., Fredericton Fish
and Game Association, and Moncton Fish and Game
Association. New Brunswick Wildlife Federation
assisted in obtaining funding. The help of the New
Brunswick Fish and Wildlife Branch, Forest Service,
Electric Power Commission, and Museum is also
acknowledged with gratitude. H.W. Blenis, Maritime

Forest Ranger School, provided encouragement and
administrative support.

Literature Cited

Boardman, S. L. 1903. The naturalist of the St. Croix. C.H.
Glass and Co., Bangor. 351 pp.

Broley, C. L. 1958. The plight of the American Bald Eagle.
Audubon Magazine 60: 162-163, 171.

Chamberlain, M. 1882. A catalogue of the birds of New
Brunswick. Bulletin of the Natural History Society of New
Brunswick 1: 23-68.

Gittens, E. F. 1968. A study of the status of the Bald Eagle in
Nova Scotia. Unpublished M.Sc. thesis, Acadia Univer-
sity, Wolfville, N.S.

Godfrey, W. E. 1970. Canada’s endangered birds. Canadian
Field-Naturalist 84: 24-26.

Grier, J. W. 1974. Reproduction, organochlorines, and mer-
cury in northwestern Ontario Bald Eagles. Canadian
Field-Naturalist 88: 469-475.

Herrick, H. 1873. A partial catalogue of the birds of Grand
Manan, New Brunswick. Bulletin of the Essex Institute 5:
3-16, 28-41.

Leighton, F. A., J. M. Gerrard, P. Gerrard, D. W. A. Whit-
field, and W. J. Maher. 1979. An aerial census of Bald
Eagles in Saskatchewan. Journal of Wildlife Management
43: 61-69.

Mathisen, J. E. 1968. Effects of human disturbance on nest-
ing of Bald Eagles. Journal of Wildlife Management 32:
1-6.

Moore, W. H. 1928. A list of the birds of New Brunswick,
Canada. 67th Annual Report. 1927. New Brunswick
Department of Lands and Mines. pp. 91-112.

Pearsall, R. R. 1879. Grand Manan notes, summers of 1877
and 1878. Forest and Stream 13: 524-525.

Pettingill, O. S., Jr. 1939. The bird life of the Grand Manan
archipelago. Proceedings of Nova Scotian Institute of
Science 19: 293-372.

Philipp, P. B.,and B. S. Bowdish. 1917. Some summer birds
of northern New Brunswick. Auk 34: 265-275.

Postupalsky, S. 1974. Raptor reproductive success: some

1981

problems with methods, criteria and terminology. Raptor
Research Report 2: 21-31.

Sprunt, A. IV, W. B. Robertson Jr., S. Postupalsky, R. J.
Hensel, C. E. Knoder, and F. J. Ligas. 1973. Comparative
productivity of six Bald Eagle populations. Transactions
of 38th North American Wildlife and Natural Resources
Conference. pp. 96-106.

Squires. W. A. 1976. The birds of New Brunswick. Second
Edition. Monographic Series No. 7. The New Brunswick
Museum, Saint John.

Stocek, R. F. 1979. Decline of summering Bald Eagles in
central New Brunswick. Canadian Field-Naturalist 93:
443-445.

Stocek, R. F. 1980. The current status of the Bald Eagle in
the Maritime Provinces of eastern Canada. Jn Proceedings
of the 1980 Bald Eagle Days. Edited by T.N. Ingram. Eagle
Valley Environmentalists, Apple River, Illinois. pp.
201-203.

STOCEK AND PEARCE: NEW BRUNSWICK BALD EAGLES

433

Townsend, C. W. 1912. Notes on the summer birds of the
Saint John River Valley, New Brunswick. Auk 29: 16-23.

Wetmore, S. P., and D. I. Gillespie. 1976. Osprey and Bald
Eagle populations in Labrador and northwestern Quebec,
1969-1973. Canadian Field-Naturalist 90: 330-337.

Whitfield, D. W. A., J. M. Gerrard, W. J. Maher, and
D. W. David. 1974. Bald Eagle nesting habitat, density,
and reproduction in central Saskatchewan and Manitoba.
Canadian Field-Naturalist 88: 399-407.

Wright, B. S. 1953. The relation of Bald Eagles to breeding
ducks in New Brunswick. Journal of Wildlife Manage-
ment 17: 55-62.

Received 17 September 1980
Accepted 21 March 1981

The Biological Flora of Canada.
3. Vaccinium vitis-idaea L. var. minus Lodd. Supplementary Account..*

IVAN V. HALL! AND JENNIFER M. SHAY2

! Research Station, Agriculture Canada, Kentville, Nova Scotia, B4N 1J5
2 Deptartment of Botany, University of Manitoba, Winnipeg, Manitoba, R3T 2N2

Hall, Ivan V., and Jennifer M. Shay. 1981. The Biological Flora of Canada. 3. Vaccinium vitis-idaea L. var. minus Lodd.
Supplementary account. Canadian Field-Naturalist 95 (4): 434-464.

Vaccinium vitis-idaea var. minus is a widely distributed evergreen dwarf shrub of northern temperate, boreal and arctic areas
in North America. Rapid dispersal of seeds by birds and mammals through their droppings, followed by good seed
germination and seedling establishment, permits this species to colonise burned forests and open habitats. Its evergreen
characteristics and low growth form permit effective photosynthesis and growth in northern climates because its niche is the
warmer microclimate at the soil surface. Plants flower in Canada from mid-June to early August depending on location, and
the berries generally mature in late August or early September. Berries are of low pH, moderate caloric content and are high in
tannin and anthocyanin.

Key Words: Vaccinium vitis-idaea var. minus, Partridgeberry, Fox Berry, Lingonberry, biology, ecology, physiology, distribu-
tion, economic importance.

1. Name

Vaccinium vitis-idaea L. var. minus Lodd., section Vitis-Idaea; Ericaceae

Vaccinium vitis-idaea spp. minor (Lodd.) Hultén (Hultén 1937, 1949),

Vaccinium vitis-idaea L. (Scoggan 1979); Mountain-cranberry, Rock Cranberry, Cowberry, Lingen,
Lingberry, pommes de terre (Fernald 1950), Partridgeberry (Rouleau 1956), airelle-d’Ida (Marie-
Victorin 1964), Lingonberry (Hultén 1968), Fox Berry (Roland and Smith 1969), Redberries
(Bourchard et al. 1978); Dry-ground Cranberry (Looman and Best 1979); berris, graines rouges
(Scoggan 1979).

2. Description of the Mature Plant
a) Raunkiaer life-form. Chamaephyte. Evergreen dwarf creeping Sits, reproducing by seeds and rhizomes.

(b) Shoot morphology. Stems semi-woody, up to 15 cm high, 1-2 mm diameter, puberulous, the bark with a
reddish tinge in ae autumn; terminal flower buds larger than the vegetative buds; evergreen leaves alternate ina
spiral (Figure 1), simple, petiolate, 5-18 mm long, 3-9 mm wide, somewhat leathery, pinnately net veined,
margin slightly revolute, obovate, apex mucronate to emarginate, base cuneate, the upper surface dark green,
purplish in autumn, the lower surface waxy, pale green with black glandular dots; slender creeping stems
semi-woody, bearing numerous shoots, new growth white or pinkish. Occasionally produces rhizomes not
morphologically distinct from stems (Shaver and Cutler 1979).

(c) Root morphology. Tap root with finely divided rootlets at the extremities (Hall and Beil 1970) (Figure 2),
adventitious roots occurring at nodes along the creeping stems and rhizomes (Viereck and Little 1972). Leiser
(1968) has described a mucilaginous root sheath surrounding the root tip.

(d) Inflorescence. Flowers occurring singly or in racemose clusters (Figure 1), “the most common numbers
being five and six” in Newfoundland (Torrey 1914), at Kentville, Nova Scotia, 5.2 + 1.6 (x + standard error
used throughout paper) (n = 167). Stems bearing inflorescences at anthesis are 9 cm tall (Pojar 1974). Flowers
pedicellate; sepals 4, green (pinkish in dried specimens), deltoid, 1.5 + 0.0 mm diameter (n = 42); petals 4, the
corolla pinkish-white, bell shaped, corolla lobes free for half their length, 4.7 + 0.5 mm long (n = 42), 4.8 + 0.6
mm wide (n = 42); stamens 8, in 2 whorls, filaments hairy. Pollen is borne in tetrads, and shed from the anther
through a terminal pore.

* See J.C. Ritchie (1955).

434

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 435

< / & : A
2 . 2 &® ~ . rf Dara
0.5 : : —>
cm \ i ee
FIGURE 1. Flowers and leaves of Vaccinium vitis-idaea L. Ficure 2. Seedlings of Vaccinium vitis-idaea L. var. minus
var. minus Lodd. Lodd. Approximate age of largest plant | yr.

The stigma and part of the style protrude beyond the corolla (Figure 1). The ovary is inferior with many
ovules and together with the calyx produce a true berry. Berries are carmine in colour when ripe, globose, nearly
1 cm diameter (Gleason 1958), acid, and have several small seeds per berry (see 7b); seeds are dark brown, elliptic
to semicircular, 1.0 x 0.6x 0.5 mm, with a reticulate surface (Montgomery 1977).

(e) Subspecies. Hultén (1949) treats V. vitis-idaea L. var minus as a subspecies.

(f) Varieties. We have followed Fernald (1950) who considered the smaller North American form as a variety
(Vaccinium Vitis-Idaea L. var. minus Lodd.) and the larger European plant as variety vitis-idaea.

(g) Ecotypes. None recognized in North America.

(h) Chromosome numbers. 2n = 24 has been reported by Newcomer (1941) from plants collected in N. Carolina
and Tennessee, by Taylor and Mulligan (1968) from a collection southeast of Port Clements in the Queen
Charlotte Islands, B.C., and from the IBP sites at Barrow Alaska by Packer and McPherson (1974). The
European V.vitis-idaea var. vitis-idaea also has 2n = 24 (Ritchie 1955, Sorsa 1962, Hedberg and Hedberg 1964,
Rousi 1966, 1967, Laane 1969).

3. Distribution and Abundance.

(a) Geographic range. Vaccinium vitis-idaea var. minus, a native species listed by Macoun (1884), occurs
widely in north temperate, boreal and arctic regions. In North America it extends from eastern Newfoundland
to Alaska and British Columbia (Figure 3, Table 1). It has been collected along the north coast of Alaska, in the

436 THE CANADIAN FIELD-NATURALIST Vol. 95

Vaccinium
vitis-jidaea L.
var. minus Lodd.

Ficure 3. Distribution of Vaccinium vitis-idaea L. var minus Lodd. from specimens in the herbaria of Canada Agriculture,
Ottawa, Ontario (DAO), National Herbarium, National Museum of Canada, Ottawa, Ontario (CAN), Acadia
University, Wolfville, Nova Scotia (ACAD), University of Manitoba, Winnipeg, Manitoba (WIN), and University of
Alberta, Edmonton, Alberta (ALTA).

Mackenzie District, Victoria Island, Keewatin (Scoggan 1979) and southern Baffin Island (Polunin 1948). It
ranges southward to northern Minnesota and New England (Scoggan 1979). Porsild and Cody (1980) map the
N. American distribution, Ritchie (1962) gives Manitoba sites and a map of Alaska sites is contained in Viereck
and Little (1975). Beyond North America it has been recorded in Greenland up to 78° N and in eastern Asia
(Hultén 1948). Eurasian distribution of var. vitis-idaea is given in Ritchie (1955). Distribution maps are listed in
Hultén (1971) and include those of Meusel (1943), Raup (1947) Szczawinski (1962), Porsild (1964), Hultén
(1968, 1971).

(b) Altitudinal range. In Eastern Canada this species extends from about sea level to the summits of the
Shickshock (elevation 1290 m), Gaspé Peninsula, and Torngat Mts. (elevation 1200 m), Newfoundland. On
Mount Washington (elevation 1700 m), New Hampshire it is found on north and west slopes (Bliss 1963). It
occurs from valley bottoms to well above treeline in the Alberta Rockies (elevation 2250 m) (Hrapkoand LaRoi
1978). Throughout the northwest it occupies habitats of various altitudes including the lower elevations of the
Mackenzie Mts., Northwest Territories, and the Ogilvie (1350m) (See and Bliss 1980), and St. Elias Mts. (up to
1500 m) (Birks 1977), and the Alsek River region (2100-2400 m) in the Yukon (Douglas 1974). Populations of
this species at the highest elevations are less vigorous and have small leaves with short internodes. At sea level in

southern Alaska it grows vigorously (Cushwa and Coady 1976). It occurs around Hudson Bay on uplands,
islands and in estuarine habitats.

4. Physical Habitat
(a) Climatic relations. The major part of the plant’s distribution lies within Thornthwaite’s (1931) Dfc climatic

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 437

region, characterised as semi-arid with short, cool summers and long, cold winters. It occurs within the arctic,
subarctic, cryoboreal, boreal and mesic climatic soil classes (Clayton et al. 1977a).

Larsen (197 1a) suggests that in central northern Canada boreal forest and tundra, correlations with air mass
frequencies are sufficiently high to warrant the conclusion that the distribution of V. vitis-idaea, one of the
dominant species in Black Spruce, White Spruce and Jack Pine forests, is related to arctic air mass distribution
and frequency. But in rock field (fell-field) tundra and tussock muskeg tundra, V. vitis-idaea distribution and
frequency are related to warm, moist Pacific air mass frequency.

Due to their northerly distribution, plants must withstand extreme cold and prolonged cover by snow. The
latter may be an advantage because ample snowcover is necessary to protect the leaves from snow abrasion
(Savile 1972). On Mt. Washington it dominates dwarf shrub communities where winter snow is deep but melts
early in the spring (Bliss 1966). In contrast, Hrapko and LaRoi (1978) report that V. vitis-idaea var. minus has
high cover in the Dryas octopetala* (Dryas) — Empetrum nigrum (Crowberry) community type where
snowmelt is late. In Greenland it can survive snow cover for more than a year (Warming 1908). Resumption of
growth is governed by the timing of snow melt (Wielgolaski 1974). Both light (Oldemeyer and Seemel 1976) and
moisture are important influences on development and growth. In Nova Scotia, maximum flower bud
formation occurs in open exposed sites, although in Alberta bud formation was greatest in the shade of Populus
tremuloides (Trembling Aspen) (Smith 1962) where moisture was greater. Lehmushovi (1977) states that in
Finland the more open and better illumined the habitat, the higher the fruiting percentage. The effect of
temperature is shown by later flower anthesis in coastal and more northerly sites. The low growth habit and
shallow root system of V. vitis-idaea enable it to take advantage of the warmer microenvironment found at
ground level. Ritchie (1955) suggests that temperature is the critical factor that limits its southern distribution in
England. He reported significant temperature differences between the southern heath areas where V. vitis-idaea
was present and absent. In field plantings at Kentville following the normal winters of 1978-79 and 1979-80,
when maximum January temperature was 15°C and the minimum -19.5°C, all native Vaccinium species,
including V. boreale (Sweet Hurts), V. myrtilloides (Sour-top Blueberry), V. angustifolium (Lowbush Blueb-
erry) and V. vitis-idaea var. minus survived without winter injury, whereas the European V. vitis-idaea
(Cowberry) plants had mostly brown, withered leaves and V. myrtillus (Dwarf Bilberry) had the stems killed to
ground level (personal observation I.V.H.).

(b) Physiographic relations. Vaccinium vitis-idaea occupies a wide range of habitats growing in extreme
exposure on headlands, sea cliffs, rocky hilltops, eskers and mountain summits, and in more sheltered sites in
Black and White spruce forests. It is found on low flat areas, gentle to steep slopes (Larsen 1972b) and
mountainous terrain (Hrapko and LaRoi 1978).

At Caribou Lake, northern Manitoba, it was a principal component of the south- and east-facing slopes of
eskers and moraines but was absent on the snowless northwest side (Ritchie 1960a). In contrast, near Inuvik
(Ritchie 1977) V. vitis-idaea was absent in the open forest on south-facing slopes and had a frequency of 40%
and cover of 3% on north-facing slopes. North-facing slopes in the Pelly Lake area had the greatest moisture and
supported more or less continuous vegetation which included V. vitis-idaea with a mean frequency of 80%
(Larsen 1972b).

Parent material may be sandstone (Hall er a/. 1964), limestone (Polunin 1948, Rickard ef al. 1965) schist,
gneisic or granite rock, glacial outwash plains of coarse sand and gravel (Larsen 197 1a) or peat (Ritchie 1960a,
Haag 1974). Vaccinium vitis-idaea is found on gray luvisols, humoferric podsols, dystric brunisols, orthic and
cryic regosols, gleysols and cryic gleysols, fibrisols, mesisols, and rockland (soil classification after Clayton et al.
1977a) and histic cryaquepts (peaty phase of orthic gleysol) with 15 cm peat (Haag and Bliss 1974). Ritchie
(1960a) reports it on peat up to 1.5 m deep. Soil pH is generally acid, it may be 3.0-3.9 (Dugle 1972), 4.5-4.7
(Barclay-Estrup and Nuttall 1974), 4.9 (Swan and Dix 1966), or 5.4 (Haag 1974). Vaccinium vitis-idaea var.
minus was common in a Larch fen (Larix laricina) with a pH 7.0-8.2 (Ritchie 1960b).

The Orthic Gleysols described by Hrapko and LaRoi (1978) had the following characteristics: pH 5.2, N = 0
ppm, p = 5 ppm, and K = 127 ppm. Their field capacity was 46%, permanent wilting point 27% and available
water 19%. Eutric Brunisols and Orthic Gleysols at Tuktoyaktuk had a 10-15 cm organic layer with a total
exchange capacity of 77.4 meq/ 100 g overlying a thin, discontinuous Bg horizon (total exchange capacity 118.1
meq/ 100 g). The permafrost maximum active layer was 35-50 cm at the end of the summer (Haag 1974), and
ranged from 29-112 cm.

In the southern part of its range V. vitis-idaea var. minus occurs most frequently in bogs and relatively moist

*Nomenclature follows Scoggan 1979

438 THE CANADIAN FIELD-NATURALIST Vol. 95

sites, although further north it frequents dry to moist sites (Polunin 1948, Larsen 1971b, 1972a) or those that are
well to moderately drained (Ritchie 1960b; Lavkulich 1973). It grows in moist sites as well as those with a slight
moisture deficit. Larsen (1972b) pointed out that differences in moisture rather than soil type governed V.
vitis-idaea distribution at Pelly Lake, for it flourishes best where there is some moisture retention. In the peat

_ soils around Little Catalina on the Bonavista Peninsula, Newfoundland, the linear growth was three to four
times that on the drier mineral soils nearby (personal observation I.V.H.).

(c) Nutrient and water relations. Vaccinium vitis-idaea nutrient requirements are low being satisfied by soils
that are shallow or of low fertility. It is generally shallowly rooting (Shaver and Chapin III 1980) but may root
from 0-30 cm (Hrapko and LaRoi 1978). Rhizomes and roots grow well in peat, but often penetrate and growin
mineral layers (Weber 1974, Shaver and Cutler 1979), the latter also found roots concentrated in the structured
dead zone, below the living moss. In Britain the rhizomes are invariably confined to the humus layer (Ritchie
1955).

Haag (1974) fertilized a dwarf shrub-heath community dominated by Betula nana ssp. exilis (Dwarf Birch)
and Salix glauca (Willow) witha prostrate ground cover of V. vitis-idaea var. minus, Ledum palustre (Northern
Labrador-tea) and Empetrum nigrum var. nigrum using 100 and 200 kg/ha ammonium nitrate (34% N) and 100
and 200 kg/ha superphosphate (20% P). The applications of N increased production by 15% and 30% by 12
July, and to 45% and 68% by 6 August at low and high rates respectively. Phosphorus at low rates decreased
total production 20% but increased it 4% at high rates. When N and P were combined, total production
increased 15% and 30% on7 July and 35% and 75% by 6 August. This indicates that total primary production is
limited more by available soil N than P.

5. Plant Communities

Vaccinium vitis-idaea var. minus isa northern plant and like its European counterpart (Ritchie 1955) occurs
primarily in the Boreal Forest and Tundra regions (Rowe 1972). It is found ina range of communities including
forests (Moss 1953a, 1953b; Ritchie 1956, 1959, 1960a, 1960b; LaRoi 1967., Blouin and Grandtner 1971,
Bouchard et al. 1978), various types of muskeg (Larsen 1965, Ritchie 1960b), bog, fen (Ritchie 1960b), peatlands
with feather mosses and/ or Sphagna (Calder and Taylor 1968, Railtonand Sparling 1973); arctic-alpine tundra
(Porsild 1937; Bliss 1966; Porsild and Cody 1980), and other communities characterized by their physiography
such as rock outcrops, eskers and rockfields (Larsen 1965), dunes (Grandtner 1977), and heath barrens along
coasts (Torrey 1974; Marie-Victorin 1964).

Data showing the abundance of Vaccinium vitis-idaea var. minus and the location of a selection of
communities containing it are given in Tables | and 2. LaRoi (1967) found it present in 35% of the White Spruce
and 42% of the Black Spruce stands in an east-west arc across the Boreal Forest. In the east its presence in the
Acadian (Roland and Smith 1969) and Great Lakes-St.Lawrence Forest Regions (Blouin and Grandtner 1971)
is more limited. In the west it forms part of Rowe’s (1972) Mixedwood Forest Region (Swan and Dix, 1966,
LaRoi 1967, Larsen 1972a, Shay and Shay 1979) and it occurs westwards to the Northern Pacific Coast (Pojar
1974). Many studies have reported it in arctic and alpine locations (Polunin 1948, Cody 1954b, Bliss 1963, 1966,
Birks 1977, Hrapko and LaRoi 1978).

6. Growth and Development

(a) Morphology. The oblong-elliptical cotyledons are ca. 2 mm in length. The first true leaves are nearly ovate
and less than one-third the size of fully mature leaves. Rhizomes begin to grow on seedlings 5 cm in height
(Figure 2). Lateral growth of plants is slow, being ca. 2 to 4cm per year at Kentville. Bliss (1966) studied growth
at 1600 -1800 m on Mt. Washington, New Hampshire and found that plants elongated 0.05-0.1 cm in 3-day
periods, and that over four years, mean shoot elongation in V. vitis-idaea ranged from 0.8-1.8 cm per year. Bliss
(1962b), in a review article on the adaptations of arctic and alpine plants to environmental conditions, states that
work in Russia by Zhuikova showed that most plants of V. uliginosum (Alpine Bilberry), V. myrtillus, and V.
vitis-idaea did not flower until 14-20 years of age. Ritchie (1955) indicated that, in cultivation, few flowers are
formed in V. vitis-idaea before plants are 5-10 years old.

In a pine forest in Sweden, Tear (1972) found that in four year old plants, stems of V. vitis-idaea were
monopodial with 2 to 3 shoots per stem and 3 to 4stems per plant. At least one stem (approximately 4cm long) is
produced each year, the number depending on plant size. Fertile shoots have more leaves than sterile ones and
produce one flower bud containing 5-6 flowers. On the other hand, plants on farmland were more branched and
stems were sympodial with 6 to 12 stems per plant (2cm long). The number of leaves per running cm was greater
than in the pine forest and two flower buds per stem were produced instead of one. Vaccinium vitis-idaea from

1981

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS

439

TABLE |. Vaccinium vitis-idaea L. var. minus Lodd. in various North American plant communities. Relative abundance
based on author’s estimates: a = abundant, f = frequent, o = occasional, r= rare, x = present.

Vegetation region*
section

BOREAL FOREST
B.2 Gaspé

B.8 Central Plateau

B.9 Superior

B.11 Upper English
River

B.14 Lower English
River

B.15 Manitoba
Lowlands

B.18a Mixed wood

B.18a Mixed wood

B.18ab Mixed wood

— Hay River

B.18b Hay River

B.19a Lower Foot-
hills

Location and
elevation (m)

Bic and Gaspé
peninsula
Klotz Lake, Ontario

Bowman Island,
Ontario

Raven Lake, Ontario

West Hawk Lake,
Manitoba
Elma, Manitoba

Clear Lake,
Manitoba

Deep Lake,
Manitoba

Prince Albert Nat.

Relative
abundance of

Park, Sask. (600 m) 106°37’ W

Waskesiu Lake,
Saskatchewan
Candle Lake, Sask.

(600 m) Sask.
(480 m) Alta.

Fort Providence,
N.W.T.
(300 m) Alta.

Horn Plateau,
N.W.T.
Northwestern Alta.

Northwestern Alta.

Crimson Lake, Alta.
(950-1010 m)
Swan Hills, Alta.
(1300 m)
(870 m) Alta.

Latitude and V. vitis-idaea
longitude Plant community var. minus Reference
48°-49° N Bog, alpine cliff x Scoggan (1950)
64°-68° W
60°50’ N Black Spruce forest Larsen (1972a)
73° 40’ W
48°44’ N Black Spruce, Balsam Barclay-
87° 59’ W Fir (0) Estrup & Nuttall (1974)
Krummholtz a
Lichen heath (0)
49°16’ N Black Spruce forest r Larsen (1972a)
91°12’ W
49° 53’ N Black Spruce forest f Larsen (1972a)
95° 15’ W
49°53’ N open bog (0) Reader and Stewart
95°54’ W (1971)
50° 43’ N Black Spruce forest (0) Larsen (1972a)
100°01’ W
50°53’ N Black Spruce bog f Shay & Shay (1979)
100° 51’ W
Sa SS) INI Black Spruce forest r LaRoi (1967)
SSeoogIN| Black Spruce forest ff Larsen (1972a)
106° 15’ W
53°50’ N Trembling Aspen forest T Swan & Dix (1966)
105° 20’ W White Birch forest r
Jack Pine forest a
White Spruce forest Co)
Black Spruce forest (0)
S39 S77 IN| White Spruce forest r LaRoi (1967)
106° 29’ N
57° 36’ N Black Spruce forest T LaRoi (1967)
117°31’ W
61°13’ N White Spruce forest a Larsen (1972a)
117°31’ W
59° 28’ N White Spruce forest T LaRoi (1967)
117° 11’ W
62°08’ N Black Spruce forest (0) Thieret (1961)
118°07’ W Lichen tundra f
55-60° N White Spruce forest o-f Moss (1953a)
114-120° W Black Spruce forest f-a
Sphagnum bog f Moss (1953b)
55-60° N White Spruce forest f Moss (1953a)
114-120° W si Pine forests r-f
Poplar forests r-f
Black Spruce bog forest f-a
Spagnum bog f Moss (1953b)
52°26’ N String fen w/ Tamarack I Slack et al. (1980)
115°01’ W
55° N Patterned fen w/ Black r Vitt et al. (1975)
116° W Spruce
54°24’ N Black Spruce T LaRoi (1967)

116°47’ W

440

TABLE |. (continued)

Location and
elevation (m)

BOREAL FOREST (continued)

Vegetation region*
section

B.19b Northern (750 m) B.C.
Foothills
(840 m) N.W.T.
B.21 Nelson River,
Manitoba
B.22a Northern Cranberry Portage,
Coniferous Manitoba
Gods Lake,
Manitoba

Ilford, Manitoba
(360 m) Sask.
(375 m) Sask.

McBride and Tod
Lakes, Manitoba

B.22b Athabasca

South
Cluff Lake, Sask.
B.23a Upper (240 m) N.W.T.
Mackenzie
(240 m) N.W.T.
Norman Wells,
N.W.T.
B.23a Upper Lower Liard River,
Mackenzie N.W.T.
(540-690 m)
(480-780 m)

(780-1140 m)
(1050-1110 m)

Wapata Lake, Sask.

Latitude and

longitude

58°42’ N
123° 40’ W
58°59’ N
125° 47’ W
54° 36’ N
100° 12’ W
54° 35’ N
101°23’ W
54°23’ N
94°26’ W
56°07’ N
95° 20’ W
55° 20’ N
105°05’ W
55°20’ N
104° 58’ W
56° 52’
99° 57’
56° 45’
101° 48’
58°55’
105° 37’
58° 20’
104° 35’ W

ZezeZ2<e2

60° 35’ N
116°07’ W
61°03’ N
117°22’ W
65°17’ N
126°51’ W
60-61° N
123° 30’ W to
124° W

THE CANADIAN FIELD-NATURALIST

Plant community

Black Spruce
White Spruce
White Spruce forest
Black Spruce
Black Spruce forest
Black Spruce forest
Black Spruce forest
White Spruce forest

Black Spruce forests
Jack Pine woodland
White Spruce forest

Black Spruce forest

Trembling Aspen
forest

Mixed conifer-
deciduous forest

Black Spruce-
Tamarack bog forest

Shrub, shoreline

Black Spruce forest

White Spruce forest
Black Spruce forest

Black Spruce-
feathermoss forest
on flood plains

Black Spruce-
Tamarack
forest on floodplains

Deciduous forest on
flood plains

Black Spruce-
Trembling Aspen
forest

Black Spruce forest

Lodgepole Pine-lichen
woodland

Mixed coniferous
forest

Subalpine fir

Mixed coniferous
forest to timberline

Relative
abundance of
V. vitis-idaea

var. minus Reference
t LaRoi (1967)
f LaRoi (1967)
r LaRoi (1967)
0 Zoltai and Tarnocai
(1971)

(0) Larsen (1972a)
a Larsen (1972a)
T LaRoi (1967)
r LaRoi (1967)
f-a Ritchie (1956)
o-f

a

a Larsen (1972a)
f-a Harms (1978)
f-a

f
o-f

f Larsen (1972a)
f Larsen (1972a)
f Cody (1960)

a Jeffrey (1961)

Vol. 95

1981

TABLE |. (continued)

Location and
elevation (m)

BOREAL FOREST (continued)

Vegetation region*
section

B.23a Upper Lower Liard River,
Mackenzie N.W.T.
(1140 m)

B.24 Upper Liard Nahanni National

Park & vicinity,
N.W.T.

(510 m) N.W.T.
(420 m) N.W.T.

B.26b Central
Yukon

Atlin Provincial
Park, B.C. -
Birch Mountain
(2061 m)

B.26c Eastern (900 m) B.C.

Yukon
(1290 m) N.W.T.

B.26d Kluane St. Elias Mountains,

N.W.T.

Alsek River, N.W.T.

B.27 Northwestern
Transition

Caribou Lake,
River, Manitoba

Long Bay, Southern

Indian Lake, Man.

Seal River, Man.

Lynn Lake, Man.

Yellowknife, N.W.T.

Abitau-Dunvegan
Lakes, N.W.T.

Latitude and
longitude

61°03’ to
61°58’ N
123° 30’ to
128° 14" W
60° N
127° 43’ W
59° 38’ N
126°50’ W
59° N
133° W

59°55’ N
131°41’ W
60°07’ N
130° 44’ W
61°32’ N
140° 30’ W

60° 30’ N
137° 30’ W

59° 20’ N
95° 10’ W

56°40’ N
100° 10’ W
59° N

96° 45’ W

56°50’ N
101°30’ W
62°54’ N
114°56’ W
62° N
110° W

Plant community

Alpine dwarf scrub
tundra
Spruce forest

Black Spruce forest
White Spruce forest

Alpine fell fields
Alpine meadow and
scrub

White Spruce forest
White Spruce forest

Open Black Spruce
forest

Dwarf Birch scrub
tundra

White Spruce forest

White Spruce treed
fen

White Spruce-
Trembling Aspen
forest

Willow shrub

White Spruce forest

Open Black Spruce
forest with lichen
scrub

Open Black Spruce
forest with moss
scrub

Tamarack forest

Black Spruce forest

Open Black Spruce bog

Open Black Spruce
forest with lichen
scrub

Dwarf scrub heath

White Spruce forest
on flood plains

Black Spruce forest

Black Spruce forest
White Spruce forest
Post-fire succession
Moss-lichen to closed
spruce forest

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS

Relative
abundance of
V. vitis-idaea
var. minus

=> =

o-f

44]

Reference

Scotter and Cody (1974)

LaRoi (1967)

Buttrick (1977)

Birks (1977)

Douglas (1974)

Ritchie (1960a)

Shay et al. (nd)

Ritchie (1959)

Larsen (1972a)

Maikawa and Kershaw
(1976)

442 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 1. (continued)

Relative
abundance of

Vegetation region* Location and Latitude and V. vitis-idaea

section elevation (m) longitude Plant community var. minus Reference
BOREAL FOREST (continued)
B.27 Northwestern __ Fort Reliance, 62° 40’ N Black Spruce forest a Larsen (1972a)
Transition N.W.T. 109° 07’ W White Spruce forest f
Tussock muskeg a
tundra
Aubrey & Colville 67° 20’ N Open White Spruce x Riewe (1979)
Lakes, N.W.T. 126° 15’ W with lichen scrub
Cottongrass-sedge on x
shores and river
margins
Dwarf scrub-lichen x
heath
Fens and bogs x
B.28a Grand Falls (270 m) Nfld. 48° 46’ N Black Spruce forest I LaRoi (1967)
56°14’ W
B.29 Northern St. Barbe South 49° 30’ to Raised bogs of coastal f Bouchard et al. (1978)
Peninsula District, Nfld. 50° 50’ N plain
56° 40’ to
58° 20’ W
Gros Morne National 49°40’ to Black Spruce dwarf a Bouchard and Hay
Park, Nfld. 50° N laurel dwarf scrub (1976)
57° 40’ to Ericaceous dwarf scrub a
58° W Black Spruce scrub f
Bog (0)
Herbaceous communi- (0)
ties along sea shore
barrens
Balsam Fir forest r
B.30 Avalon Western Bay, Nfld. 47°50’ N Coastal barren f Hall (unpublished)
53°10’ W
Little Catalina, Nfld. 48°40’ N Coastal barren it
53°05’ W
B.31 Newfound- St. Barbe South 49° 30’ to Alpine scrub “tucka- x Bouchard et al. (1978)
land-Labrador District, Nfld. 50°50’ N moor”
Barrens 56° 40’ to Juniper scrub on f
58° 20’ W serpentine table-
land
Buchans Plateau, 48°45’ N Black Spruce forest r Bergerud (1971)
Nfld. 57° 10’ W
B.32 Forest tundra Great Whale River, 55°17’ N Open forest with lichen f Forest and Legault (1977)
Quebec 77° 46’ W or moss
E. Pen Island 57° N Bare rock on coast
Coast of Hudson 88° W Moss heath hummocks T Kershaw (1974)
Bay, Ontario Lichen heath r-f | Kershaw and Rouse
(1973)
Lower Hayes River,
Manitoba 57° N White Spruce forest f Ritchie (1960b)
92°20’ W on flood plains
Open White Spruce f
with lichen or moss
scrub
Tamarack forest (0)
Churchill, Manitoba 58°47’N Black Spruce forest a Larsen (1972a)

94° 10’ W

1981

TABLE |. (continued)

Vegetation region*
section

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS

Location and
elevation (m)

BOREAL FOREST (continued—

B.32 Forest tundra

B.33 Alpine Forest
Tundra

MONTANE

M.5 Douglas-fir
and Lodge-
pole pine

Alpine tundra

COASTAL FOREST
C.3 Northern
Pacific Coast

GREAT LAKES-ST.

Ennadai Lake,
N.W.T.

Artillery Lake,
N.W.T.

Campbell-Dolomite
Uplands, N.W.T.

Inuvik, N.W.T.

Nahoni (1200 m) and
Ogilvie (1350 m)
Uplands, N.W.T.

Banff and Jasper
National Parks,
Alberta (1280-
1830 m)

Jasper National
Park, Signal Mt.
(2180-2260 m)

Pacific Rim National
Park, Vancouver
Island, B.C.

LAWRENCE FOREST

L.6 Temiscouata-
Restigouche

L.12 Rainey River

Bic and Gaspé
Peninsula, Quebec

Southeastern
Manitoba

Latitude and

443

Relative
abundance of
V. vitis-idaea

longitude Plant community var, minus Reference
60° 42’ to Black Spruce forest a
61°10’ N+ White Spruce forest a
100° 55’ to Tussock muskeg a
101°45’ W tundra
Rock field a
63°05’ N Black Spruce forest a
108°05’ W White Spruce forest a
Tussock muskeg a
tundra
Rock field a
68° 20’ N Open Black Spruce (0) Ritchie (1977)
133° 40’ W woodland
Black Spruce forest (a)
68° 23’ N Picea mariana-Vac- T-a Black and Bliss (1978)
133°42’ W cinium uliginosum
woodland
65° 36’ N Alpine lichen T-O See and Bliss (1980)
136°41’ W communities
64° 18’ N
137°21’ W
50°55’ to Lodgepole Pine- T
53°10’ N lichen woodland forest
115° to 119° WLodgepole Pine forest T
52°51’ N Dryas rock tundra r Hrapko and LaRoi
117°59’ W Heath tundra T-O (1978)
Sedge-wood rush snow r
bed tundra
49°10’ N Sphagnum bog r Pojar (1974)
125°55’ W
48° to Slaty sea ledges and x Scoggan (1950)
49° 15’ N sea cliffs
64° 15’ to
68° W
49°00’ to Jack Pine forest (0) Mueller-Bombois (1964)
49°40’ N Black Spruce-feather f
95° 10’ to moss forest
96°20’ W_— Deciduous forest ie
(Trembling Aspen,
Balsam, Poplar)
Jack Pine-deciduous r
forest
Tamarack-White Birch T

forest

444

TABLE 1. (continued)

Vegetation region*
section

THE CANADIAN FIELD-NATURALIST

Location and
elevation (m)

USA HEMLOCK-WHITE PINE
NORTHERN HARDWOODS

N. Appalachian
Highlands

Great Lakes-
St. Lawrence

Presidential Range,
New Hampshire

Lake Agassiz peat-
lands, Minnesota

LOW ARCTIC TUNDRA

New Quebec

South Coast of
Baffin Island

Western Coast of
Hudson Bay

Port Burnell, N.E.
coast of Ungava
Bay, Quebec

Wakeham Bay, S.
Shore of Hudson
Strait, Quebec

Puvirnitug, Quebec

Lake Harbor

Cape Dorset

Coral Harbor,
Southampton
Island

Chesterfield Inlet

Latitude and
longitude

44° 15’ N
71°20’ W

48°05’ N
92°45’ W

60° 25’ N
64°52’ W

61°36’ N
71°57’ W

60°02’ N
77° 15’ W

62°52’ N
69°53’ W

64° 10’ N
76° 30’ W

64°08’ N
83° 10’ W

63° 20’ N
90° 42’ W

Plant community

Dwarf shrub heath
with sedge & rush

Dwarf shrub heath

Diapensia

Bog

Mixed swamp forest

Cedar string bog and
fen

Tamarack string bog
and fen

Black Spruce-feather
moss forest

Sphagnum-Black

Relative
abundance of
V. vitis-idaea

var. minus Reference
o-f Bliss (1963)
o
r
r
r Heinselman (1970)

Spruce-Leather Leaf

bog forest

Sphagnum-Leather
Leaf-laurel-spruce
heath

Dwarf scrub heath on o-d Polunin (1948)
lowlands
Lichen heath on low- a
lands
Herbaceous tussocks f
in marsh areas
Dwarf scrub heath on f-a
lowlands
Dwarf scrub heath, f
snow patches
Communities on x Bournérias (1971)
shores and dry
gravel
Dwarf scrub heath on f Polunin (1948)
lowlands
Dwarf scrub heath on (0)
summits and slopes
Birch-willow dwarf f
scrub
Herbaceous com- f
munities on south
facing slopes
Dwarf scrub heath on f-a
lowlands
Dwarf scrub heath f
Dwarf scrub heath a
Dwarf scrub heath, f

snow patches

Vol. 95

1981

TABLE 1. (continued)

Location and
elevation (m)

Vegetation region*
section
LOW ARCTIC TUNDRA (continued)
Keewatin District Northern Keewatin,
N.W.T.

Snow Bunting Lake,
N.W.T.

Pelly Lake, N.W.T.

Dubawnt Lake
N.W.T.

Mackenzie District Clinton Colden Lake,

Eastern Area N.W.T.

Matthews Lake,
N.W.T.
Mackenzie Delta Various sites

and Tuktoyaktuk

Peninsula
Various sites
Tuktoyaktuk
Tuktoyaktuk
ALASKA, USA

INTERIOR FOREST
Eastern Alaska
Yukon-Tanana
Upland (390 m)

Latitude and
longitude

64° to
69° N

91° to
98° W

66° 10’ N
94°25’ W

66°03’ N

101°03’ W

64°01’ N
99° 25’ W

64° 08’ N
107° 27’ W

64°05’ N
111° 15’ W
68° 45’ to
69°55’ N
131° 20’ to
134° 40’ W

68° 20’ to
69° 55’ N
131° 20’ to
134° 45’ W
69° N
131°52’ W

69° 10’ N

133° 10’ W
69°27’ N
133° W

63°42’ N
144° 23’ W

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS

Relative

abundance of
V. vitis-idaea

Plant community

Sedge meadow

Willow-sedge
meadow

Orthophyll scrub

Lichen-heath plateau

Lichen uplands

Barrens

Low meadow

Rock field

Scattered lichens on
esker summits

Wet Cotton Grass
meadow

Dry lichen heath

Dwarf scrub-herb
tundra

Rock field

Black Spruce forest

Tussock muskeg
tundra

Rock field

Dwarf scrub

Tussock muskeg
tundra

Rock field

Marshy hummocks

Dwarf scrub heath
Medium scrub heath
Dwarf Birch on raised
center polygons
Sedge heath and sedge-
Cottongrass heath
Sedge meadow
Dwarf scrub heath
Cottongrass dwarf
birch heath

Black Spruce forest-
tundra

White Spruce-birch
forest

Cottongrass-lichen

Dwarf shrub heath

Dwarf shrub heath

Black Spruce forest

var. minus

[—7) (=n) (©) tay 5) ta} fa)

445

Reference

Thompson (1980)

Larsen (1972b)

Larsen (1971a)

Cody and Chilcott
(1955)
Corns (1974)

Hernandez (1973)

Ritchie (1974)

Haag (1974)

Wein and Bliss (1973)

LaRoi (1967)

446

TABLE 1. (continued)

THE CANADIAN FIELD-NATURALIST

Vol. 95

Location and
elevation (m)

Vegetation region*
section
ALASKA, USA
INTERIOR FOREST (continued)
Eastern Alaska

Yukon-Tanana
Upland (600 m)

(345 m)
(600 m)
(705 m)

Near Fairbanks

Sites near
Fairbanks

National Moose
Range, Kenai
Peninsula

COASTAL FOREST
Glacier Bay, S.W.
Alaska (5-45 m)

LOW ARCTIC TUNDRA
Upper Firth River
Valley, Alaska-
Canada

Umiat

Latitude and
longitude

62°06’ N
145°52’ W
65° 15’ N
146°47’ W
63° 33’ N
148° 48’ W
61°59’ N
146°53’ W
64°80’ N
148°06’ W

58°22’ N
135°37’ W

69° N
141° W

69° 22’ N
152°08’ W

abundance of

Plant community

Black Spruce forest

White Spruce-fir
forest

White Spruce-fir
forest

White Spruce-fir
forest

Open Black Spruce
forest with feather-
moss and lichen

Closed Black Spruce
forest with feather-
moss

Open Black Spruce
woodland with
moss and lichen

White Spruce forest
with alder

Black Spruce forest

Mature spruce-
deciduous forest

Mature deciduous
forest (Trembling
Aspen, White Birch)

Spruce regrowth after
1947 fire

Birch-spruce regrowth
after 1947 fire

Hemlock-spruce forest
Lodgepole Pine
muskeg

Dwarf scrub-herba-
ceous communities
on centers of frost
polygons

String fen and bog

Sedge meadow

Willow scrub

Alder scrub

Cottongrass-dwarf
scrub heath

Sedge-dwarf shrub
heath

Relative

V. vitis-idaea
var. minus

f-d

f-d

>m310 K

Reference

LaRoi (1967)

Dyrness & Grigal (1979)

Oldemeyer and Seemel
(1976)

Reiners et al, (1971)

Drew and Shanks (1965)

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 447

TABLE |. (concluded)

Relative
abundance of
Vegetation region* Location and Latitude and V. vitis-idaea
section elevation (m) longitude Plant community var. minus Reference
LOW ARCTIC TUNDRA (continued)
Ogotoruk Valley 68°06’ N Cottongrass tussock (0)
165° 46’ W tundra
Cape Thompson 65° 45’ to Wet herbaceous T Johnson et al. (1966)
region 69°00’ N meadows
162° 00’ to Cottongrass tussocks (0)
166°15’ W on soilifluctia
slopes and with ice-
wedge polygons
Sedge communities of (0)
high center polygons
Meadows on dry T

rocky uplands

*Vegetation Regions in Canada after Rowe (1972), Alaska after Viereck and Little (1975), and the conterminous United
States after Braun (1950). Low arctic climate designation after Bliss (1979).

Jay Peak, Vermont, a sub-alpine site had a leaf surface-to-biomass ratio of 51 + 9 cm?/g (Svoboda and Taylor
1979). In Norwegian birch forest, this ratio for V. vitis-idaea was 96 cm?/g dry wt. on 18 June, attained its
maximum of 211 cm2/g dry wt. on 20 July and fell to 121 cm?/g dry wt. by the end of August (Berg et a/. 1975).
Most of the leaves were held at angles of less than 30° from the horizontal.

(b) Physiology. Chlorophyll content of Vaccinium vitis-idaea var. minus in tundra associations at Tuktoyak-
tuk (Hutchinson et al. 1976) was 11.0 wg/leaf on 15 July 1973, rose to 18.4 wg/leaf by 20 August but dropped to
9.5 wg/leaf by 31 August. Plants contained 2.4 mg/g dry wt .chlorophyll a, 1.3 mg/g dry wt. chlorophyll B and
0.8 mg/g dry wt. carotenes. Total chlorophyll content in plants from a mature forest at Norman Wells was only
1.92 mg/g dry wt. in August.

Field transpiration when measured between 14.10 and 21.10 hr on 14 July 1973 had a mean value of 1.6 = 0.6
mg H,O/ min/g dry wt./leaf. It had risen to 2.4 + 1.2 mg H,O/ min/g dry wt./leaf when measured between 11.50
and 15.50 hr on 24 August (Hutchinsoneta/. 1976). Pigment summaries for plants affected by oil spills indicate
an enhancement of pigments in regrowth after summer spills but a reduction in pigments in regrowth after
winter oil spills (Hutchinson et a/. 1976). At Hardangervidda, Norway, in a birch forest, the previous years’ V.
vitis-idaea var. vitis-idaea leaves were harvested on 15 and 20 July and 15 August. They yielded an average of
3.37 mg chlorophyll a and B per g dry wet.; 176 mg chlorophyll per g nitrogen; 2.06 mg chlorophyll per 100 cm?
leaf area; the carotenoid ratio was 1.82and nitrogen content 2.31% (Berg 1975). Data for V. vitis-idaea in lichen
heath and dry meadow, and for older leaves are also given by Berg (1975).

The distribution of '4C photosynthates (Berg et a/. 1975) indicates that at budbreak, green parts of V.
vitis-idaea accounted for 22% of the recovered radiocarbon in lichen heath and 39% in birch forest, while
below-ground parts contained 55% in the lichen heath, and 35% in the birch forest. Two weeks after budbreak
green parts accounted for 37% (lichen heath) and 49% (birch forest) and below ground parts 39% (lichen heath)
and 21% (birch forest). At both sampling periods the remainder of !4C was found in the non-green above-ground
parts.

Hadley and Bliss (1964) studied the energy reserves of several alpine species including V. vitis-idaea var.
minus on Mt. Washington. For this species they found that new shoots produced 5270 + 24.4 cal/g and a lipid
content of 3.12 + 0.08%, quite similar to old shoots with 5288 + 25.9 cal/ ganda lipid content of 3.86 + 0.12%.
The protein content of new shoots (10.3%) was significantly higher than that of older shoots (8.7%). The calories
per gram of rhizomes and roots (4996 + 17.5) were nearly equal to those of the shoots, but the lipid
(1.43 + 0.08%) and protein (5.3%) contents were significantly below values for the shoots. In a preliminary
study Bliss (1962a) had shown that caloric values for the entire plant increased during early July but declined
somewhat as the berries matured. Wielgolaski and Kjelvik (1975) give caloric values for V. vitis-idaea in

Vol. 95

THE CANADIAN FIELD-NATURALIST

448

€€ asuapouds wunwayluvIDp
+ 6r SisuappuDdds snusO)D
6b Saployijadiu winiuisdv4
S v1 I ppnu vjjaiiyy
LI I DUId]D vavIAID
67 I DAOL{iun sasauopy
€€ iG wnsop fia] windy
+ Ol SE Z SNAADIOAIIW SNIIOIAXE)
99 €P 8 DAOLfISUO] “IRA S1VaAOG DaDUUIT
0S 6 pajvidaj xadv)
P os 9 suaosaqnd snqny
ep ZI DIOfill DULIDIIUS
ZTE I LS L avouIWeIH
LS ZI pulsadsip xaIvy
CZ 9 DINUdAOI snjd10D
9¢ pinpidsiy visay1jnvy
Ol snjpjadiuponb snd2094xEQ
I I 6l DInjnIAjp2 auydppapuvy)
9 iG DIDJUap1A] DI11Ua10g
A I 62 C €C v6 ce €l ¢ uNJIpUunjuUao1s WNpaT
9 Ol pyofisnsup piujoy
v7 I LS ¢ 69 cl wuinisiu windjaduy
0Z €Z wunyofisnsup ‘4
00T CC 18 (ZG v v6 6S 69 v 8 02 SNUIU “IBA DAODPI-SI1A WNIUIIIDA
%A %D MA VASO uemer aa) %A %A %A %D %AU %D VY SsqioH pue sqniyg
— — — 0 — — — I % punols seg
— _— as S — polea mw = = yoadsy
= polreA 6-€ I €-0 Pollva 0 0 polrea (0) adoys
2 O€-SI O0S-SLZ S9EI OSI 00L-0SS 0S9 0s (w) spninty
OE of ET LOTO0l 90.8h1 OF.0r1 SS00I 0CSOl AS 0001 VS 6 C0 0o€S “Bu0T
61 089 £0099 M8009 CE oI9 OT 19 10S ofS £6 00S £5 060 ££ o8P ‘VeT
xI IIIA IIA IA A AI Il II I SUS
52 Be pe ee Fee 0 Oe ye mo Fg
Es = 6 pene Sy & me BS Oo © 3 0 3 fp
= 8 < a cs iS) es) Ss 53 6 as oR Ps 2 eS
a: ee a Tee eS ra eo 3 ©
zZ&8 a ~ 8 8 a Ts 8 > 09 po
28 Ze oo S58 3, 7 o& Zz 5 - B
= < >s a8 re £ s Eyes 5 43
i=l : a > Sao canal aa 5.8 o eq
ns a me <o us ape oS © =
ES pe re ee ac :
3 R ) : ae
Aas x 5 =| =
— 3 = : >
No 3 a

‘sartunu0d juRld sAQeUasoidal ouIU UI Sotdads pajEIoOsse pu ‘PpOT SNUuIU “IVA “T Vavpi-siiiA UNIUIIIDA “7 ATAV

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 449

1981

v9

OV

91
(G3
00!

LI

LC

€l

68

-m

MN

I hs a)
re)

81

(44

17
66
6£
8s
18
06

loa)

NDNarronwrsasrtonm

pyofiusnsup vaanssnvg
DIAOPDAGH S1ADINIIpag
pyofinop ‘dss vanvjs xyvs
DZ1YAAOPlOYI XALDD
wuinoipajds unjasinbZ
SIISnpod vII11Ualog
pjiydojoav “S§

pyofiunjd ‘dss vyofiaydyd xyvg
psojnpun]s ojniag

DY Ofiin] S11SO13DII4p
DINIID XS

parvqday X10

S1jnvID aUuallg

11M0]1281g XadDD
puospvijal adoissv)
‘dds/‘ds xaun9

DUId]D aojYyIOsalH]
winssids ‘dss winjpuispa uinsoydouq
DIDpunNIOd XaIDD
pyofyod ppawoipuy
winsou1syn wniuizID4A
Sniowapiupys snqny
aajsnjod wnpaT

puidjp sojdydpisojoap
SISUBPDUD) S1jSOABDULDIDD
ponv]s vag

suadiaA DjOLAT

adpauy wniddunjay
DIOfiavsp DIOLAg
SYNDIIpNU DIDI
wuindipodyon4) wniodsig
vdsido snujpy

ajnpa wnuingi
sisuapouDd vIO1A
unyofiusnsup wnigojidq
ajpa10q winypy
snonajo1yoo snsdyivy
suasund sisdozkio
$1]Da10q SDIuUald
ppunsas njosdg
snipujod sajispiag
isan-pan sojdydpisojo4p
DUDIUIBAIA DILDSDL]
Sapiodaigs uinjasinby
SLUDINIIID DSOY
Ssnipaouul snuajq

Vol. 95

THE CANADIAN FIELD-NATURALIST

450

(4 Mpaquajyjanu <7
Cl (6 uinjpsuoja unudi2ig
vc I winjnjoaad wnuday
~Z puldafisuns DUIpD]D 7 ‘ds wnusvyds
97 + unijanbiujopnasd wndag
cE I € wnsosna wnipidyy
uinjjayojnd wniyoudinq
91 + ‘dds wnisayidyonag
OV LI 8Y ‘dds wnusvyds
v8 I I suagsaosnf ‘q
cl (6 unjpjnpun ‘q
ce € wunisnsup wnudsziq
09 S winpi3in} “y
VC C WN DUILUNID UNIUuWOoIDjINy
vl I HMO PuUniq Unipolar
rl Z SISUAAISDI-DISIAD UNI
67 C ‘ds wniupy
+ Z P 6c 6 suapuajds wniwos0j Ay
67 LS 67 1Maqgadyos wUNnIzOANajd
rl Z ¢ adisnjod wniuwosnjnp
8 I vl €l tl uingsnf{ wnusvydsy
Tl + v I GI wnuiadiunt wnys141d] 0d
Sassoj
(6 DAO fipupdis DjOAAg
%A %D %A %) *O %A %A %A %)D %AU %)D VV Sqiox{ puke sqniys
= = = 0 — = = = I % punois sieg
= = as S — poles A — — yoadsy
= PouwA 6-€ I £-0 Bonus 0 0 Pollea (c) ado[s
= Of-SI O0S-SL7 = S9ET OSI 00L-0SS 0S9 0S (Ww) spranty
OE of ET O10] 908h1 OFObl SS.00I 07SO0I AS 001 VS S6 Oks “Bu0T
61 089 £0099 O8o0b9 EID Olol9 10S of iS 00S LS 6b iff 08P Vey]
XI IIIA IIA IA A Al II] II I aus
Py ee Eye
s.8 <2 of ms aie a§ 38 BS bo
* 3 ieee Seren Bog BS aS: is zg =
Be PAG a eee ee ass eS ze Se aes
=e : > §& is S Seite N &. > S = Za
ee ee SE he:
ns = Spee eeu es eS oe 2
o> 0 tot) > n . (tie) <
53 S aes = is 5
o> og 5 . a
—§ = SS 5
to o 3 =e

(paenuuod) Z ATaVL

451

DIJDWOAAIAY DJJAUDAIIG.
supinu pjYyog
Dippixdd piuopyjD
DUUIIUOD XadDD

po fiisajul svdiq

‘dds snpojs0undaiq

suajiu wnuddyiuawo] ppojz0ydiu
aADY1D UNIPY id Ie pdivsdyovig xX1v¢
DIJOfi1D] SISOABDIIAY ‘dds vog

asuaaiD unjasinby
SNUIA]D Sn|DSDAISp

(8261) SSI pue yorTg

suaquingodd p14najasioT
DUDIIDUL DAI
wnyofisnsup wnioydougq
suvjs ‘)

SsuDjjidvs XaivyD

DSO]JIA DINIINBUIg
DILIOpDAGH] S1ADINIIpag

xI

sadisuo] v1ADIa1S
psnfuod pjnznT
(€L61) ussite] IIIA

DUDIIAAUID D1I1A
pyofisadsp sisdozdiQ.
SNIDIOYII 4a1Sp

wngij24D DUOAYydan
(6L61) [e81H pure ssouikgq

suajiu wnudAyjuawmoy
sInpop snqny

pyofial sido

DIDP4OD DAAISIT
DIDSNIGO Dildauaqvy
DIOsIpUunjo4 DAasSOA
DauUDS]Dg Saiqy
DUIJIAD] X1ADT

IIA

suarspindind auys0z1yos
unpIAl] uojnDI0ayH)
wunjounjduos wnipodostT

DUDIADUL DIAIJID

pyjdydoonnjs ppawoipup

wuinsojnpuvs saqry

(6L61) Aeys pue Aeys III
‘3[Qe UT JOU Saroads [eUOnIppy

(LL61) SXtG IA (S961) UssIeT A (9961) XIS pueuemg A]

ee
——————————SSSSSSSS SS

HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS

1981

99 py INajO1yIo “"y

Sc DINPIIU DILOJIa]pv

v uinsojJUuaUlo} UOjNDIOaLal§

v8 € G6 pID]JNINI “Dd
Cl I DIIpUuvjs! “Dd

Cl I C7 G SIJDAIU DIADAIAD
OV Cl pulafisund *D
v8 CI Siu DUIPDID
v DUIUDI “q

Cl + + G psoylydp “gq
€ pjAjoppdjod psasiljag

S1jsad]p “Dd

€ DaDAJOANDUID “D

08 cl «OO! € pinosnqav “Dd
VA 6 91 ‘dds niuopyjD
08 cI «OO! OL saioads uayory
sugyory

v8 9 WIOMIQATT
96 €v 98 06 satoads ssop|
cl ‘ds winijiuosmyy

a4isnjod wuniuuoopjny

Vol. 95

THE CANADIAN FIELD-NATURALIST

452

“SIOYINE [VUIZIIO SMOT[OJ JINJeOUSWION A
“poljIpow ANYSIs s1oyINe [eUIZIIO ‘(6/6]) UeBBOIS SMO][OJ JIN} eSUSWION V7
“OHS Joyjoue ul PoISst] Apealye satoads 10j MTZ <<
ISAOD + “%Z = “7y0) “HE = Ore | oAbY popnyout satoads 9] BOS OI-I ulwod I3A09 — x ‘% Aduanbaly dAT} POI = WA ‘Aouanbady yuaol1od = WA SI9A00 qugo1od = yO)
‘WI 0-SZ “XI
‘2 [-O7 “IITA
:Syo[d 7 ¢Z7-8p “ITA
‘10]d ;W OT ‘IA
‘Wl [-O81 ‘A
‘Wl $Z'0-0€ “AI
‘2 670-1 ‘TIT
‘sjuiod wopuel 0s ‘TI
Syoosued} Ww 00! RB suoje sjuiod OOI ‘T :OHS 9ZIS ajduies
‘(8L61) SSI PUL YOuTIG — XI *(6L61) [es1IH pue
ssaukiq — ITA *(LL61) S41 — IA ‘(S961) UasseT — A ‘(9961) XI puke ueMs — AI ‘(6L61) Aes pue Keys — IIT ‘(1 L61) WeMaIS pue Jopeoy — IT ‘(6L61) 2H —I

aque 10j sajouj004

(panuljuod) 7 a1aV L

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 453

Norwegian tundra dry meadow sites as 4895 + 25 calories per g dry wt. for green material, 4734 + 32 for
non-green material above the roots, 4731 + 52 for living roots and 4634 + 83 for dead material.

The light compensation point for this species on Mt. Washington was 200 ftc and the light saturation point
2000 ftc. Following the cessation of terminal shoot growth, respiration decreased with a resulting increase in net
photosynthesis (Hadley and Bliss 1964). Vaccinium vitis-idaea was one of the species selected for detailed
physiological study in Norway. Nygaard (1975) reports the effects of low temperature pretreatment (8-12 hr at
+2-3° Cand 12-16 hrat+10° C) and high temperature pretreatment (8-12 hrat+15° Cand 12-16 hrat +23-25° C)
at a relative humidity of 80% using an infrared gas analyzer. The apparent photosynthesis in V. vitis-idaea
pretreated at high temperatures was higher than when pretreated at low temperatures. For example with high
temperature pretreatment apparent photosynthesis at 15°C was 4.2 mg CO,/g/hr and with low temperature
pretreatment photosynthesis at 15° C was 2.1 mg CO,/g/hr (see Nygaard 1975, p. 165). Such acclimation effects
were smaller in experiments carried out in summer than in fall. The CO, compensaticn point as a function of
acclimation temperature was 63 ppm at 25° Cin low temperature material, 51 ppm at 25° C for high temperature
material, and 86 ppm for untreated material.

Skre (1975) reports CO, exchange data for V. vitis-idaea under different experimental conditions and
compares his Norwegian data with other studies. The values for apparent photosynthesis of 5-6.7 mg CO,/g/hr
are higher than those of Hadley and Bliss (1964) and Scott and Billings (1964) but agree with those of Nygaard
(1975).

The composition of stems and leaves of V. vitis-idaea var. minus was monitored at various seasons on the
Kenai Peninsula, Alaska (LeResche and Davis 1973, Oldemeyer and Seemel 1976, Oldemeyer et a/. 1977). Their
data indicate a slight protein peak in summer (Table 3) also shown by Scotter (1965) in northern Saskatchewan.
However there was a dramatic decrease from summer to winter in values for Ca, Mg, K, Na, Cu, Fe, Mg, Znand
Mn.

In the test garden at Kentville, Nova Scotia, the levels of various elements found in V. vitis-idaea var. minus
are similar to those given by Gerloff et a/. (1964) for several other ericaceous species. At Kentville, levels of N
and P in current leaves (Table 4) were greatest in June and decreased to September. Calcium in the current
growth increased as the season progressed undoubtedly due to the buildup of calcium pectate in the cell walls of
the leaf tissue. Similar trends were previously reported for Vaccinium angustifolium (Townsend et al. 1968). The

TABLE 3. Composition of stems and leaves of V. vitis-idaea L. var. minus Lodd. from northern Saskatchewan and the Kenai
Peninsula, Alaska.

Scotter* LeResche Oldemeyer and Seemel Oldemeyer
and Davis et al.
1965 1973 1976 1977
3 analyses 3 sites 6 sites 6 sites
July 1960 Sept. 1960 Mar. 1961 May 1971 Aug. 1973 Feb. 1974 July 1974
Carbohydrate %
Cell walls 67 28.iae Ay Byo7/ ae We) 50.5
Acid detergent fibre 17.6 meas MAN Si) ae AW) 44.6
Lignin 50.0 1226) se O4 el Seta KG 23.8
Protein % 6.33 7.87 6.34 5.4 S./25 Oye 5.4 + 0.30 7.6
Minerals ppm
Ca 4700 5500 5500 6300 49905452 8ee2 Ou S223 699
Mg - = - 1700 1328 +27360 4620.41 1426
K - - - 3400 438 +288.68 29.8 + 5.94 3691
Na = - - 80 55) 2s S23 2228221074) 72
Cu - - - - oy) ae Sts? 0.2 + 0.03 13.8
Fe - - - - IESE =O IS6 3.2 + 0.88 44
Zn - - - - Gsjae She (0,3) ac (037 8.6
Mn - - - - WOae so? ESE OR29 111.8
iP 1000 1600 1100 900 - - - - -
Ether extract % = 2 = 2.1 E . = s z
Ash % De Del DDS 2.1 - - - - -
Crude fat % 2.81 2.81 3.16 2 : = : : -
fibre % 15.63 20.37 17.70 - - - - - -

*Northern Saskatchewan, leaves only were analysed.

454 THE CANADIAN FIELD-NATURALIST Vol. 95

TABLE 4. Nutrient levels in Vaccinium vitis-idaea L. var. minus Lodd leaves from the test garden at Kentville, Nova Scotia

% composition on dry wt. basis

Leaf cohort Date N P K Mg Ca
pyg! 2 June 79 0.872 0.060 0.27 0.12 0.61
c 2 June 79 1.43 0.145 0.58 0.12 0.29
pyg 5 July 79 0.81 0.067 0.42 0.13 0.55
c 5 July 79 0.92 0.103 0.59 0.14 0.39
pyg 2 Aug 79 0.70 0.060 0.32 0.16 0.52
c 2 Aug 79 0.85 0.095 0.38 0.13 0.41
Pyg 5 Sept 79 0.76 0.072 0.43 0.15 0.61
c 5 Sept 79 0.87 0.093 0.48 0.15 0.50

1. pyg = preceding year’s growth, c = current growth.
2. samples dried at 70°C and analysed by standard procedures (A.O.A.C. 1970).

decline of N content in the previous year’s leaves during the early summer suggests that senescence has
commenced. Levels of N, Pand K were higher in the current leaves than the preceding year’s at all dates, whereas
with Ca, the preceding year’s leaves had the higher values. For Mg, values were more or less equal for the two
years at the various sampling dates. The unfertilized soil producing the leaves sampled (Table 4) was a well
drained Berwick sandy loam with a pH of 4.6 (Cann et al. 1965).

Analysis of mineral content of V. vitis-idaea in IBP tundra sites in Norway and Finland is given in Wielgolaski
et al. (1975). In general the data are similar to the Kentville analyses.

Stark et al. (1979) have shown that ripe fruit has a high tannin content (721 mg/ 100 g fresh weight), high
anthocyanin content (127 mg/ 100 g), low pH (2.5) and a total sugar content of 6.6%. Hoffman et al. (1967)
found the ascorbic acid of V. vitis-idaea var. minus on 15 September was 22.5 mg/ 100 g and the B-carotene on 23
September 1964 was 0.79 ug/g of berries.

(c) Phenology. Bell and Burchill (1955) found that V. vitis-idaea var. minus overwintered with ovule develop-
ment barely evident and sporogenous tissue present in some flowers. According to these authors it was the 12th
ericaceous species to bloom in Nova Scotia, from mid-June to early July. A major difference between V.
vitis-idaea var. minus and its European counterpart var. vitis-idaea is the fact that the latter had two flowering
periods per season (Ritchie 1955). On 15 May 1979 vegetative growth measured | cm on plants in the transplant
garden at Kentville (lat. 45°04’N) and full bloom occurred on 25 May. This was an unusually early season,
approximately two weeks early. With plants further north the date is later, blooming in the Yukon about
mid-July. The first flowering of this species at Sheep Mountain, southwest Yukon, (lat. 61°21’N) occurred
about | July and continued for nearly a week; it was the 48th of 60 species to flower (Hoefs 1979). On Mt.
Washington (lat. 44° 17’N), Hadley and Bliss (1964) reported that vegetative growth started 18 June, plants were
in flower on 2 July, and bore young fruits by 25 July. Ritchie (1955) states that root growth occurs in both early
spring and fallin V. vitis-idaea var. vitis-idaea. Old leaves of var. minus were starting to drop in August of the
second year in Nova Scotia. Svoboda and Taylor (1979) indicated that leaves persist for 3 years. Vaccinium
vitis-idaea shoots began to grow from 23-30 June in a Norwegian alpine heath 200-300 m above tree line
(Wielgolaski and Karenlampi 1975), the time of snowmelt being a very important feature that varies from year
to year. Growth in Finland begins about 20 days after snowmelt (Kjelvik and Karenlampi 1975) and lasts about
40 days. There isa shorter period between snowmelt and budbreak in Norway and length growth ends about 25
July in Norwegian birch forest and lichen heath. The delay in budbreak may be a form of protection for plants
growing onexposed sites with thin, early melting snow cover (Wielgolaskiand Karenlampi 1975). In Norwegian
alpine lichen heath, shoot growth of V. vitis-idaea was 7 mm in 1971 and almost the same in the warmer summer
of 1972. It grew about 10 mm ina dry alpine meadow in both years, while in subalpine birch forest it grew 39 mm
in 1971 and 42 mm in 1973, appearing to be strongly dependent on moisture (Wielgolaski and Karenlampi
1975). Maximum shoot length was attained in early July in the subalpine birch forest, and toward the end of
July in the alpine lichen heath. Vaccinium vitis-idaea in the birch forest had a higher fertility (3.2%) than in
lichen heath sites (0.6%) and dry meadow (0%).

7. Reproduction
(a) Floral biology. Vaccinium vitis-idaea is slightly protandrous, that is, the anthers ripen and release their
pollen before the stigma of the same flower is receptive but is partially self-compatible. It produces both odors

1981 HALL AND SHAY: VINIUM VITIS-IDAEA VAR. MINUS 455

and nectar (Pojar 1974). In Newfoundland cross pollination is by bees and butterflies (Torrey 1914), while in

southern B.C. it is pollinated by bumble bees and bee flies (Pojar 1974). Warming (1908) lists four bumble bees:
Bombus balieatus, B. consobrinus, B. lapponicus and B. jonellus as the chief pollinators in Greenland. Torrey
(1914) states that under field conditions in southeastern Newfoundland flowers remain open for about a week
and that under favorable conditions, 30% of the pollinated flowers set fruit. Flowers that were cross-pollinated
between clones set fruit more readily than selfed flowers (Halland Beil 1970). A significant positive correlation
(r = 0.466) was found between seed number and berry weight (Hall and Butler 1971). Two nectar samples from
50 blossoms collected on 26 May 1979 yielded 1.0 and 1.6% soluble solids.

(b) Seed production and dispersal. Seventy-five berries collected on Tancook Island, Nova Scotia in October
1978 were stored at 1° C until 21 March 1979, when their mean weight was 0.403 + 0.098 g and the mean number
of seeds was 15.2 + 7.4. Incontrast, an average of 6.7 seeds per berry was reported for var. vitis-idaea in Britain
where seed set and berry production followed pollination by 2-4 weeks (Ritchie 1955).

Data froma stand of V. vitis-idaea var. minus on Tancook Island, Nova Scotia, show that the weight of sterile
shoots was not significantly greater than that of fruiting shoots (without berries) and the weight of berries
exceeded 1000 kg/ha. The weight of 25 immature berries (0.15 + 0.01 g) taken on 25 July 1979 from the same
locality was considerably less (berries will increase in weight by at least one-third in final month of growth) than
that of mature berries reported above. Veijalainen (1976) reports the average yield of V. vitis-idaea ina good
uncultivated site in Finland to be 500 kg/ha and the maximum yield in cultivation to be 8150 kg/ha. The yield
from a fertilized forest was 3000 kg/ha.

Seeds are dispersed in the droppings of many birds and mammals, such as Dendragapus obscurus (Grouse),
Calachites canadensis (Spruce Grouse), Piranga erythromelas (Scarlet Tanager), Sialia sialis (Eastern Blu-
ebird), Hylocichla spp. (Thrushes), and other songbirds, as well as Ursus americanus (Black Bear), Tamias
striatus (Eastern Chipmunk), Peromyscus leucopus (White-footed Mouse) and other mammals (Martin et al.
1961).

(c) Seed viability and germination. Seeds were collected from 13 sites in eastern Newfoundland on 13 and 14
September 1977 and held ina refrigerator at 1° C until 20 December when germination tests began. Seeds placed
on the surface of a peat-vermiculite medium under 16 h light at 26° Cand 8 h dark at 24° Chad 87% germination
after 59 days. The highest rate of germination was in seeds extracted from fresh, mature berries and a steady
decline occurred with less than 10% germinating by 12 to 16 months. Johnson (1975) studied buried seeds ina
subarctic lichen woodland east of Great Slave Lake, and reported that 65% of the sample cores contained seed of
Vaccinium vitis-idaea and V. uliginosum. Vaccinium vitis-idaea counts showed 3.3 seeds/ 1000 cc of which 0.7
seeds/ 1000 cc were fertile at Porter Lake (180 years post-fire), and 4.8 seeds/ 1000 cc with zero fertility as
Selwyn Lake (98 years post-fire). In Britain, Ritchie (1955) reports a lower percentage seed germination for
freshly harvested seeds but an increased rate for cold-treated seeds.

(d) Vegetative reproduction. Examination of ericaceous communities in the Atlantic Provinces and Quebec
suggests that lateral growth is by rhizome proliferation of ramets from the original seed-established clones (Hall
and Beil 1970). Harper and White (1974) report a similar situation in V. myrtillus. Large old clonal plants may
eventually break up by frost action, fire, burrowing, etc. into separate clones which, in turn, produce their own
vegetative shoots and rhizomes. Pojar (1974) reports vegetative reproduction by creeping stems, and Johnson
and Rowe (1977) indicate that this is an efficient means of vegetative reproduction in subarctic habitats.

Eighty-two percent of the stem cuttings from 14 Newfoundland clones produced roots, when held under
intermittent mist, from 15 September 1977 to 7 December 1977. Studies in Finland on propagation have shown
that rooting of stem cuttings is best in the spring (89%). Milled peat (85%) is superior to humus (55%) as a
rooting medium; indole butyric acid at 6000 ppm gave a slight increase in rooting (90%), and pieces of rhizome
behaved similarly to stem cuttings (Lehmushovi 1975). Rooting success was 83% for the controls.

8. Population Structure and Dynamics

(a) Dispersion patterns. Vaccinium vitis-idaea is generally dispersed contagiously, the creeping stems form-
ing patches, but it may form mats (Hrapko and LaRoi 1978) described as loose, in moist mossy situations, or
dense in dry, rocky sites (Viereck and Little 1972).

(b) Age distribution. In Sweden, Tear (1972) found that the age of plants ina woodland varied from one to nine
years with the average being four. His method of determining age was based on counting the number of annual
rings at the base of the aboveground stem as previously shown for V. myrtillus by Flower-Ellis (1971) and for V.
angustifolium — V. myrtilloides by Hall (1957). Harper (1977) points out that shrubs are difficult for

456 THE CANADIAN FIELD-NATURALIST Vol. 95

population biology studies as the aboveground parts may be of a different age than the underground system.
After any major disturbance such as a forest fire, habitat becomes available for colonisation but seedlings are
subject to severe inter- and intraspecific competition from annuals in the first year, perennials (mostly sedges,
grasses, legumes and composites ) from years two to five and later by shrubs and trees (Torrey 1914) (see also
Table 2). Once established V. vitis-idaea var. minus will persist indefinitely unless shaded out by conifers.
Within 25 years after fire its frequency becomes relatively stable (Johnson and Rowe 1977).

(c) Size distribution. Ten plants which had been previously grown in 10 cm clay pots were set out in the
transplant garden at Kentville in 1970. In 1979 they had an average maximum crown diameter of 70cm. In this
experiment plants grew free from competition, but under normal field conditions crown diameter would
probably have been less.

(d) Growth and turnover rates. Observations in the field in Nova Scotia and Newfoundland revealed that
seedlings showed high rates of survival and new crops of seedlings are produced in favorable years. Once
established, plants spread by slender creeping stems rooting at nodes (Viereck and Little 1972). Fresh weight of
V. vitis-idaea var. minus from Tancook Island, Nova Scotia, was 253.5 + 69.5 g/m? for sterile shoots,
207.5 + 43.7 g/m? for fruiting shoots and 112.5 + 19.8 g/m? for fruit (unpublished data). Stewart and Reader
(1972) reported 845 + 8% terminal growing points per m2, and 740 + 7 rooted shoots per m? with a mean weight
of 15 mg per rooted stem. Leaves contributed 27.8 g/m? (53%), flowers and fruits 13.8 g/m? (26%), new stems 8.5
g/2, (16%), old stems 3.6 g/m? (5%) to the 52.7 g/m? net primary production in aerial components ina S.E.
Manitoba bog. Ericoides, Ledum groenlandicum (Labrador-tea), Chamaedophne calyculata (Leather-leaf),
Kalmia polifolia (Bog-laurel), Oxycoccus quadripetalus (Small cranberry) and Vaccinium vitis-idaea var.
minus) had net aerial primary production of 298.2 g/ m2, their total annual subsurface production was 1461.1
g/m? with 1234.6 g/m? remaining after | year’s decomposition (Stewart and Reader 1972). Thirty nine percent of
the V. vitis-idaea leaves decomposed after | year in the bog. Incontrast inan Alaskan site Shaver and Chapin III
(1980) reported total aboveground biomass in a tussock tundra community to be 59 g/m? with above ground
productivity being 16 g/m?/yr. Leaves accounted for 35.4 + 8.8 g/m72., stems 18.3 + 3.3 g/m?, and 2.4+0.1
g/m? occurred in the unstructured organic layer (Shaver and Cuttler 1979). Vaccinium vitis-idaea var. minus
accounted for 22% of the average 2 year production. The mean annual aerial production at four sites near
Inuvik, Alaska (Wein and Bliss 1973a) was 6.1 g/m? with new leaf growth appearing in mid-July. Productivity in
the Kenai peninsula, Alaska, ranged from 1000-3000 kg/ha (Oldemeyer and Seemel 1976). Data for productiv-
ity in various Fennoscandian communities is given in Kjelvik and Karenlampi (1975). Leaves persist for at least
3 years.

(e) Successional role. Seeds can germinate on bare ground if conditions are favorable, but V. vitis-idaea var.
minus is not generally a pioneer species, although it grows on the summits of the highest rockfields at Pelly Lake,
N.W.T. with Rhododendron lapponicum (Lapland Rosebay), Hierochloé alpina (Holy Grass), Alectoria
ochroleuca (Lichen) and A. nitidula (Larsen 1972b). It invades the nearby tundra bog gravel communities
dominated by Alectoria ochroleuca, A. nitidula, Hierochloé alpina and Luzula confusa (Woodrush). It also
invades senescent Eriophorum vaginatum (Cotton-grass) tussocks in communities dominated by the Eriopho-
rum and areas of frost activity after the pioneers (Larsen 1972b). It combines the Empetrum nigrum and a lichen
mat to forma seral community leading towards White Spruce (Ritchie 1959). Mueller-Dombois (1964) reports
it in successionally advanced Jack Pine stands with 2-5 cm humus and rotten wood on the ground.

In the far north V. vitis-idaea var. minus assumes a climax role (Cody 1954a) whereas in the barrens of
Newfoundland it acts as a seral species before being displaced by Picea mariana (Black Spruce) and Abies
balsamea (Balsam Fir).

9. Interaction with Other Species

(a) Competition. Since light intensities below 2152 lux are limiting (Hadley and Bliss 1964), Picea glauca
(White Spruce), P. mariana, and Abies balsamea must be considered serious competitors for light. Similarly
Kalmia angustifolia (Sheep-laurel), Ledum groenlandicum and Potentilla tridentata (Three-toothed Cinqu-
efoil) and other species occupying the same habitat, compete for water, nutrients, etc. (Table 2).

Of some 15 species responsible for net primary productivity of bog vegetation in southeastern Manitoba V.
vitis-idaea var. minus with 7.8% relative cover is sixth in importance. Furthermore, it ranked third in total
annual above-ground net production being exceeded only by Chamaedaphne calyculata and Ledum groenlan-
dicum: respective values were 52.7, 106.1 and 68.1 g/m? (Reader and Stewart 1971). In the Mackenzie Delta
between Dempster (64° N 138° W elevation 1200 m) and Umiat (69° N 151° W), sites north of the tree line and

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 457

dominated by Eriophorum vaginatum ssp. spissum had 12-31% of their cover provided by Ledum decumbens,
V. vitis-idaea, V. uliginosum and Betula nana spp. exilis. Eriophorum vaginatum, L. palustre and V. vitis-idaea
contributed 78-81% of the annual production (Wein and Bliss 1974).

(b) Symbiosis. Warming (1908) stated that V. vitis-idaea has both endotrophic and ectotrophic mycorrhiza
associations. Ritchie (1955) cites additional European work which was carried out on V. vitis-idaea var.
vitis-idaea.

(c) Predation and parasitism. Macoun (1884) stated that the fruit of V. vitis-idaea var. minus was invaluable as
a source of food to birds migrating north in spring. Foliage of the species is eaten by a number of mammals such
as Alces alces (Moose), Lepus americanus (Snowshoe Hare), Rangifer tarandus groenlandicus (Barren-ground
Caribou), and Ursus americanus (Black Bear). In Alaska, Cushwa and Coady (1976) found that Alces alces ate
large quantities of this evergreen during the winter on the Kenai Peninsula but inland, deep snow prevented
consumption. In digestion trials Oldemeyer and Seemel (1976) report in vitro dry matter disappearance in
August of this species as 50.7%. The composition of stems and leaves showed carbohydrates of cell walls to be
33.1%, acid-detergent fiber 29.3%, lignin 12.6% and protein 5.7%. In February the dry matter was lower (41.8%)
carbohydrates were greater and protein about the same at 5.4%.

There is a suggestion by Kelsall (1968) that the green leaves of V. vitis-idaea as well as those of Ledum
groenlandicum, L. decumbens and Arctostaphylos uva-ursi (Common Bearberry) may supply some special
ingredient in the winter diet of Rangifer tarandus. In his quantitative analysis of stomach contents of caribou in
northwestern Saskatchewan and District of Mackenzie he showed that Vaccinium uliginosum and V. vitis-idaea
account for 3.8% of summer 21.5% of winter foods. Leaves of V. vitis-idaea consisted of 5.76% protein, 3.13%
fat and 20.20% crude fibre. Scotter (1965) pointed out that the nutritive value of lichens was lower than that of a
shrub group consisting of Arctostaphylos uva-ursi, Ledum groenlandicum and Vaccinium vitis-idaea. The
group average for protein was 7.38% crude fat 3.98% and crude fibre 18.25% dry matter basis. Porsild (1962)
states that species of Vulpes (foxes), Lagopus (ptarmigan), Corvus (ravens), Larus (sea gulls) and Anatidae
(geese) feed on Vaccinium spp. fruit.

In Newfoundland, Lepus americanus feeds heavily on the shoots in winter and Ursus americanus eats this
species in spring when snow has melted from the barrens but persists in the adjoining forests; U. americanus and
Vulpes vulpes (Red Fox) consume quantities of the fruit in late fall and this constitutes a large proportion of
their diet.

The fruit, in Newfoundland, is occasionally infested with Grapholitha sp. (Olethreutidae) whose larval stage
generally leaves the fruit by the second week of September before fruit harvest (Hall 1978).

The major fungal diseases listed by Conners (1967) are: Red leaf — caused by Exobasidium vaccinii,
Witches’-broom — Pucciniastrum goeppertianum, and leaf rust — Pucciniastrum vaccinii. Conners states that
E. vaccinii occurs on V. vitis-idaea var. minus in Alaska, and Savile (1959) cites several infested specimens from
British Columbia and Quebec.

(d) Toxicity and allelophathy. None Known.

10. Evolution and Migration

Camp (1945) placed this species in a section by itself and offered no explanation for its origin. The North
American plants are consistently smaller than those in Europe (Fernald 1950). Camp (1945), however, reported
that both dwarf and robust types were present in the east as well as west of this continent. Hultén (1949) states
that only spp. minus is found in North America.

Frost and Ising (1968) used thin layer chromatography of phenolic compounds in studying V. myrtillus, V.
vitis-idaea and the hybrid between them (V. intermedium). Differences in the chromatograms of the bilberries
from Breanas in southern Sweden and Varmdon near Stockholm were considerable. Between the hybrids from
the localities, differences were also noted in the phenolic compounds, identified only as to position.

11. Response Behavior

(a) Fire. This is an important factor within the botanical range of V. vitis-idaea var. minus. In fire-prone

ecosystems survival is related to the amount of moisture in the soil, the season of the year and depth of rhizome

layer. Inarctic Eriophorum tussock communities Wein and Bliss (1973a) report V. vitis-idaea ssp. minus and E.

vaginatum ssp. spissum (Hare’s Tail) had not recovered fully from fires after two growing seasons. The nutrient

levels in leaves of V. uliginosum ssp. alpinum were higher from burned areas than unburned sites.
Vaccinium vitis-idaea var. minus appears within the first six years after fire rapidly attaining its maximum

458 THE CANADIAN FIELD-NATURALIST Vol. 95

and maintaining a frequency of between 70 and 100% ina range of Black Spruce-lichen, Jack Pine-lichen, and
White Spruce-birch communities of all ages up to 280 years (Johnson and Rowe 1977). Black and Bliss (1978)
report similar findings in Picea mariana-Vaccinium uliginosum forests where fire does not change plant
succession as dominant vascular plants soon resprouted and achieved preburn prominence. After eight years V.
vitis-idaea var. minus had a frequency of 56% and cover of 8% rising toa maximum frequency (100%) and cover
(55%) at 144 years. It was the only species to increase in cover and frequency in stands aged 120-200 years, and to
maintain relatively high cover values in communities 200-300 years old. It attained a cover of 15% 130 years after
fire on drumlins of sand and gravel near Great Slave Lake with Stereocaulon paschale (22%) and Cladonia
stellaris (17%) (Maikawa and Kershaw 1976). It dominates both burned and unburned sites in the Kenai
Peninsula, Alaska (Oldemeyer et a/. 1977).

In Eastern Canada the rhizomes of V. vitis-idaea var. minus are much finer and generally closer to the surface
of the soil than those of V. angustifolium which can withstand regular burning (Hall er a/. 1979). Fire, therefore,
weakens the plants and reduces the proportion of V. vitis-idaea in mixed stands. Re-colonisation is possible by
regrowth from dormant buds on stems and rhizome pieces which escape lethal temperatures.

(b) Grazing and harvesting. Small mammals such as Snowshoe Hares clip the ends of stems, destroy apical
buds and cause increased branching, while Moose remove entire clusters of stems in a single mouthful. Sheep
will only graze V. vitis-idaea when more palatable species are wanting (Ritchie 1955).

(c) Flooding. The normal habitats of this species are not subject to flooding. In peatlands, it occurs on the tops
of hummocks which are drier than depressions (Larsen 1972b).

(d) Drought. Prolonged dry spells seriously reduce vegetative growth and decrease fruit size. In the drier part of
its range the vigor and frequency of shoots is less.

(e) Herbicides. 2,4-D causes browning of stems and leaves, leaf drop, and at high concentrations (1.3 to 1.8 L
per 450 L of H,O) death of the plant. Relaled Vaccinium spp. are classified as susceptible to 2,4-D; 2,4,5-T; and
intermediate to ammonium sulfamate (Ontario Weed Committee 1980).

(f) Chemical changes: oil spills. Vaccinium vitis-idaea var. minus with a frequency of 64% in a cottongrass
community, and 42% in a dwarf-shrub community was sprayed with Norman Wells crude oil in July 1973
(Freedman and Hutchinson 1976). Its leaves lost their chlorophyll slowly, the dead leaves remained on plants
for up to two seasons and regrowth leaves showed gigantism. A summer spill had greater effect than one in
February when plants were covered by 40 cm snow because devolatisation and partial detoxification of the oil
occurred before it contacted plant tissue. A year after the summer spill V. vitis-idaea had a frequency of 21% in
the cottongrass community and 17% in the dwarf shrub community. The authors suggest that a high degree of
recovery might be produced after 10 to 15 years, but species diversity would be reduced with bryophytes and
lichens eliminated (Freedman and Hutchinson 1976). In mature Picea mariana forest, V. vitis-idaea had a cover
of 48% and frequency of 98% prior to crude oil spills. Summer spills reduced cover to zero, while low and high
intensity winter spills reduced cover to 12% and 6%, respectively. Regrowth shoots lost water more rapidly than
controls, apparently due to their thinner cuticle (Hutchinson et al. 1976). At Tuktoyaktuk, in the cottongrass
winter spill site (Hutchinson and Hellebust 1978), recovery provided up to 78% cover in the second year; 58% of
this was Eriophorum, with V. vitis-idaea, Calamagrostis canadensis (Blue-joint), Ledum groenlandicum and
Rubus chamaemorus (Bake-apple) providing the remainder. Hutchinson and Hellebust (1978) give flowering
success and other details for V. vitis-idaea in cotton grass, mature forest and dwarf shrub communities after
crude oil spills.

(g) Frost. Opened flowers had a 50% mortality at -1.5° Cand the corresponding temperature for unripe berries
and flower buds was -3.1 to —3.5°C (Tear 1972). In Japan hardening at 0°C for 10 days enhances freezing
resistance in V. vitis-idaea, it was maximal after hardening at -3° C for 10 days when resistance to -70° C was
induced (Sakai and Otsuka 1970).

12. Relationship to Man

Fruit of Vaccinium vitis-idaea var. vitis-idaea has been an important ingredient in the diet of most Scandina-
vians for centuries and harvested with small comb-sieves by hand in open woodlands. The flavor of raw berries
of var. minus is very tart, but Porsild (1974) and Ryan (1978) claim that the flavor is improved by slight frost.
Porsild (1937) states that this species is a most valuable antiscorbutic due to its high vitamin C content. Hedrick
(1919) states, “This is the wi-sa-gu-mina of the Crees and the cranberry most plentiful and most used throughout
Rupert’s Land”. In Alaska, the Inuit names are ‘keepmingyuk’ (Shismaref north to Noatak), ‘keepmik’ (Seward

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 459

Peninsula) and ‘toomalgleet’ (Lower Kuskokwim) and the Indian name is ‘nutlut’ (Heller 1953). Heller states
that the berries can be mixed with rose hips to make an excellent jam. Biermann (1975) has written a brief
description of the plant and suggested the economic importance of the fruit among people of Scandinavian
origin in Minnesota and Wisconsin.

The berries make excellent jams, jellies, and preserves. In favorable years up to 100 tonnes are exported from
Newfoundland to Scandinavian countries for human food products.

Acknowledgments

The authors acknowledge the technical assistance of A.T. Lightfoot, A.C. Brydon, K.G. Cairns and B.
Brookes.

We also thank W.J. Cody, D.G. Dodds, H.J. Lloyd, J.E.H. Martin, C.T. Shay, S.P. Vander Kloet and G.W.
Wood for helpful suggestions in the preparation of this manuscript and C.T. Shay for compiling Table 1.

Literature cited

Association of Official Analytical Chemists. 1970. Official methods of analysis of the Association of Official Analytic
Chemists, Washington, D.C. 11th ed. pp. 8-58.

Barclay-Estrup, P.and D.V. Nuttall. 1974. Some aspects of the distribution and ecology of Crowberry (Empetrum nigrum L.)
on the north shore of Lake Superior. Canadian Field—Naturalist 88: 171-181.

Bell, H.P. and J. Burchill. 1955. Winter resting stages of certain Ericaceae. Canadian Journal of Botany 33: 547-561.

Berg, A. 1975. Pigment structure of Vascular Plants, Mosses and Lichens at Hardangervidda, Norway, /n Fennoscandian
Tundra Ecosystems. Edited by F.E. Wielglaski, Springer-Verlag, N.Y. 216-244.

Berg, A., S. Kjelvik and F.E. Wielgolaski. 1975. Measurement of leaf areas and leaf angles of plants at Hardangervidda,
Norway, Jn Fennoscandian Tundra Ecosystems. Edited by F.E. Wielgolaski, Springer-Verlag, N.Y., 103-110.

Bergerud, A.T. 1971. Abundance of forage on the winter range of Newfoundland caribou. Canadian Field—Naturalist 85:
39-52.

Biermann, J.E. 1975. A description of Vaccinium vitis-idaea. Fruit Varieties Jounral 29: 5-7.

Birks, H.J.B. 1977. Modern pollen rain and vegetation of the St. Elias Mountains, Yukon Territory. Canadian Journal of
Botany 55: 2367-2382.

Black, R.A. and L.C. Bliss. 1978. Recovery sequence of Picea mariana-Vaccinium uliginosum forests after burning near
Inuvik, Northwest Territories, Canada. Canadian Journal of Botany 56: 2020-2030.

Bliss, L.C. 1962a. Caloric and lipid content in alpine tundra plants. Ecology 43: 753-757.

Bliss, L.C. 1962b. Adaptations of arctic and alpine plants to environmental conditions. Arctic 15: 117-144.

Bliss, L.C. 1963. Alpine plant communities of the Presidential Range, New Hampshire, Ecology 44: 678-697.

Bliss, L.C. 1966. Plant productivity in alpine micro-environments on Mount Washington, New Hampshire. Ecological
Monographs 36: 125-155.

Bliss, L.C. 1979. Vascular plant vegetation of the Southern Circumpolar Region in relation to antarctic, alpine and arctic
vegetation. Canadian Journal of Botany 57: 2167-2178.

Blouin, J. and M.M. Grandtner. 1971. Etude écologique et cartographie de la vegetation du comté de Riviere-du-Loup.
Ministére des Terres et Foréts. Gouvernement du Québec. Mémoire No. 6. 240-246.

Bouchard, A. and S. Hay. 1976. The vascular flora of the Gros Morne National Park coastal plain, in Newfoundland.
Rhodora 78: 207-260.

Bouchard, A., S. Hay and E. Rouleau. 1978. The vascular flora of St. Barbe South District, Newfoundland: An interpretation
based on biophysiographic areas. Rhodora 80: 228-308.

Bournérias, M. 1971. Observations sur la flore et la végétation des environs de Puvirnitug (Nouveau-Queébec). Le Naturaliste
Canadien 98: 261-318.

Braun, E.L. 1950. Deciduous forests of eastern North America. Blakiston Co., Philadelphia. 596. pp.

Buttrick, S.C. 1977. The alpine flora of Teresa Island, Atlin Lake, B.C., with notes on its distribution. Canadian Journal of
Botany 55: 1399-1409.

Calder, J.A.and R.L. Taylor. 1968. Flora of the Queen Charlotte Islands. Part 1. Systematics of the vascular plants. Research
Branch, Canada Department of Agriculture, Monograph No. 4, Part 1. p. 473.

Camp, W.H. 1945. The North American Blueberries with notes on other groups of Vacciniaceae. Brittonia 5: 203-275.

Cann, D.B., J.I. McDougall and J.D. Hilchey. 1965. Soil survey of Kings County, Nova Scotia. Report No. 15, Nova Scotia
Soil Survey, Truro. 41-42.

Clayton, J.S., W.A. Ehrlich, D.B. Cann, J.H. Day and I.B. Marshall. 1977a. Soils of Canada. Vol | Soil Report. Research
Branch Canada Dept. Agriculture Minister of Supply and Services, Ottawa. 241 pp.

Cody, W.J. 1954a. New plant records from Bathurst Inlet, N.W.T. Canadian Field—Naturalist 68: 40.

Cody, W.J. 1954b. Plant records from Coppermine, Mackenzie District, N.W.T. Canadian Field—Naturalist 68: 110-117.

Cody, W.J. 1960. Plants in the vicinity of Norman Wells, Mackenzie District, N.W.T. Canadian Field—Naturalist 77: 226-228.

Cody, W.J. and J.G. Chilcott. 1955. Plant collections from Matthews and Muskox Lakes, Mackenzie District, N.W.T.
Canadian Field—Naturalist 69: 153-162.

460 THE CANADIAN FIELD-NATURALIST Vol. 95

Conners, I.L. 1967. An annotated index of plant diseases in Canada and fungi recorded on plants in Alaska, Canada and
Greenland. Canada Department of Agriculture, Research Branch. Publication 1251. 310-312.

Corns, I.G.W. 1974. Arctic plant communities east of the Mackenzie Delta. Canadian Journal of Botany 52: 1731-1745.

Cushwa, C.T. and J. Coady. 1976. Food habits of moose, Alces alces, in Alaska: a preliminary study using rumen content
analysis. Canadian Field—Naturalist 90: 11-16.

Douglas, G.W. 1974. Montane zone vegetation of the Alsek River region, southwestern Yukon. Canadian Journal of Botany
52: 2505-2532.

Drew, J.V.and R.E. Shanks. 1965. Landscape relationships of soils and vegetation in the forest-tundra ecotone, Upper Firth
River Valley, Alaska, Canada. Ecological Monographs 35: 285-306.

Dugle, J.R. 1972. Plant associations in the Whiteshell Nuclear Research Establishment Controlled Area. Whiteshell Nuclear
Research Establishment Pinawa, Manitoba. 72 pp.

Dyrness, C.T. and D.F. Grigal. 1979. Vegetation-soil relationships along a spruce forest transect in interior Alaska. Canadian
Journal of Botany 57: 2644-2656.

Fernald, M.L. 1950. Gray’s manual of botany. 8th edition. American Book Company, New York, New York. 1134 pp.

Flower-Ellis, J.G.K. 1971. Age structure and dynamics in stands of bilberry (Vaccinium myrtillus L.). Avdelningen for
Skogsekologi, Rapporter och Uppsatser. Nr 9. 108 pp.

Forest, P. and A. Legault. 1977. Analyse de la flore vasculaire de Poste-de-la-Baleine, Nouveau-Québec Le Naturaliste
Canadien 104: 543-566.

Freedman, W. and T.C. Hutchinson. 1976. Physical and biological effects of experimental crude oil spills on low Arctic
tundra in the vicinity of Tuktoyaktuk, N.W.T., Canada. Canadian Journal Botany 54: 2219-2230.

Frost, S. and G. Ising. 1968. An investigation into the phenolic compounds in Vaccinium myrtillus L. (bilberries), Vaccinium
vitis-idaea L. (cowberries), and the hybrid between them V. intermedium Ruthe employing thin layer chromatography.
Hereditas 60: 72-76.

Gerloff, G.C., D.D. Moore and J.T. Curtis. 1964. Mineral content of native plants of Wisconsin. Wisconsin Agricultural
Experiment Station Report 14. 27 pp.

Gleason, H.A. 1958. The new Britton and Brown Illustrated flora of the northeastern United States and adjacent Canada.
Vol. 3. The Sympetalous Dicotyledoneae. New York Botanical Press. 589 pp.

Grandtner, M.M. 1977. Contribution a l’étude ecologique des dunes a Empetrum nigrum du Parc National Forillon, Québec.
Documents phyto-sociologiques 1: 135-142.

Haag, R.W. 1974. Nutrient limitations to plant production in two tundra communities. Canadian Journal Botany 52:
103-116.

Haag, R.W. and L.C. Bliss. 1974. Energy budget changes following surface disturbance to upland tundra. Journal Applied
Ecology 11: 355-374.

Hadley, E.B. and L.C. Bliss. 1964. Energy relationships of alpine plants on Mt. Washington, New Hampshire. Ecological
Monographs 34: 331-357.

Hall, I.V. 1957. The tap root in lowbush blueberry. Canadian Journal Botany 35: 933-934.

Hall, 1.V. 1978. Vaccinium species of horticultural importance in Canada. Horticultural Abstracts 48: 441-445.

Hall, 1.V., L.E. Aalders, and L.R. Townsend. 1964. The effect of soil pH on the mineral composition and growth of the
lowbush blueberry. Canadian Journal of Plant Science 44: 433-438.

Hall, 1.V., L.E. Aalders, N.L. Nickerson and S.P. Vander Kloet. 1979. The Biological Flora of Canada. I. Vaccinium
angustifolium Ait. Canadian Field—Naturalist 93: 415-430.

Hall, I.V.and C.E. Beil. 1970. Seed germination, pollination and growth of Vaccinium vitis-idaea var. minus Lodd. Canadian
Journal Plant Science 50: 731-732.

Hall, I.V. and K.P. Butler. 1971. The relation between seed number and berry weight in the foxberry. Cranberries 36: 17.

Harms, V.L. 1978. A botanical inventory of the Cluff Lake area, northwestern Saskatchewan, Musk-ox 22: 31-62.

Harper, J.L. 1977. Population biology of plants. Academic Press, London. 892 pp.

Harper, J.L., and J. White. 1974. The demography of plants. Annual Review of Ecology and Systematics 5: 419-463.

Hedberg, I. and O. Hedberg. 1964. Documented chromosome numbers of Swedish plants. Svensk Botanisk Tidskrift 58:
125-129.

Hedrick, U.P. 1919. Sturtevant’s notes on edible plants. J.B. Lyon Company, State Printers, Albany, New York. 588 pp.

Heinselman, M.L. 1970. Landscape evolution, peatland types, and the environment in the Lake Agassiz peatlands natural
area, Minnesota. Ecological Monographs 40: 235-261.

Heller, C.A. 1953. Edible and poisonous plants of Alaska. Cooperative Extension Work in Agriculture and Home Econom-
ics, Extension Service University of Alaska and United States Department of Agriculture. pp. 108-9.

Hernandez, H. 1973. Natural plant recolonization of surficial disturbances, Tuktoyaktuk Peninsula Region, Northwest
Territories. Canadian Journal Botany 51: 2177-2196.

Hoefs, M. 1979. Flowering plant phenology at Sheep Mountain, Yukon Territory. Canadian Field— Naturalist 93: 183-187.

Hoffman, I., F. S. Nosowad and W. J. Cody. 1967. Ascorbic acid and carotene values of native eastern arctic plants.
Canadian Journal of Botany 45: 1859-1862.

Hrapko, J. O. and G. H. LaRoi. 1978. The alpine tundra vegetation of Signal Mountain, Jasper National Park. Canadian
Journal of Botany 56: 309-332.

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 461

Hultén. E. 1937. Outline of the history of arctic and boreal biota during the Quaternary period. Stockholm: Bokforlags
Aktiebolaget Thule. 168 pp.

Hultén, E. 1948. Flora of Alaska and Yukon. Part VIII Ericales-Hydrophyllaceae. Lunds Universitets Arsskrift N.F. Avd. 2
Bd. 44: 1262-1265.

Hultén, E. 1949. On the races in the Scandinavian flora. Svensk Botanisk Tidskrift Bd. 43: 383-406.

Hultén, E. 1968. Flora of Alaska and neighboring territories. Stanford University Press. Stanford, California. 731 pp.

Hultén, E. 1971. The circumpolar plants. II. Dicotyledons. Almquist and Wiskell, Stockholm. 463 pp.

Hutchinson, T. C. and J. A. Hellebust. 1978. Vegetation recovery in the Canadian arctic after crude and diesel oil spills.
Minister of Supply‘and Services Canada. 62 pp.

Hutchinson, T. C., J. Hellebust and M. Telford. 1976. Oil spill effects on vegetation and soil microfauna at Norman Wells

| and Tuktoyaktuk, N.W.T. Minister of Indian and Northern Affairs, Ottawa, ALUR 74-75-83 Information Canada. 65 pp.

Jeffrey, W. W. 1961. Notes on plant occurrence along Lower Liard River, Northwest Territories. National Museum of
Canada Bulletin 171: 32-115.

Johnson, E. A. 1975. Buried seed populations in the subarctic forest east of Great Slave Lake, Northwest Territories.
Canadian Journal of Botany 53: 2933-2941.

Johnson, E. A.and J. S. Rowe, 1977. Fire and Vegetation Change in the Western Sub Arctic. Ministry Supply and Services
Canada. 58 pp.

Johnson, A. W., L. A. Viereck, R. E. Johnson and H. Melchoir. 1966. Vegetation and flora, Jn Environment of the Cape
Thompson Region, Alaska. Edited by N.J. Willimovsky and J.N. Wolfe. U.S. Atomic Energy Commission Washington,
D.C. 277-354.

Kelsall, J. P. 1968. The migratory barren-ground caribou of Canada. Department of Indian Affairs and Northern Devel-
opment, Canadian Wildlife Service. pp. 67-105.

Kershaw, K. A. 1974. Studies on lichen-dominated systems. X. The sedge meadows of the coastal raised beaches. Canadian
Journal of Botany 52: 1947-1972.

Kershaw, K. A.and W. R. Rouse. 1973. Studies on lichen-dominated systems. V. A primary survey of a raised-beach system
in northwestern Ontario. Canadian Journal of Botany 51: 1285-1307.

Kjelvik,S.and L. Karenlampi. 1975. Plant biomass & primary production of Fennoscandian subarctic and subalpine forests
and of alpine willow and heath ecosystems. /n Fennoscandian Tundra Ecosystems. Edited by F. E. Wielgolaski. Springer-
Verlag, N.Y., 111-120.

Laane, M. M. 1969. Further chromosome studies in Norwegian vascular plants. Blyttia 27: 5-17.

LaRoi, G. H. 1967. Ecological studies in the boreal spruce-fir forests of the North American taiga. I. Analysis of the vascular
flora. Ecological Monographs 37: 229-253.

Larsen, J. A. 1965. The vegetation of the Ennadai Lake area, N.W.T.: Studies in subarctic and arctic bioclimatology.
Ecological Monographs 35: 37-59.

Larsen, J. A. 197la. Vegetational relationships with air mass frequencies: boreal forest and tundra. Arctic 24: 177-194.

Larsen, J. A. 1971b. Vegetation of Fort Reliance, Northwest Territories. Canadian Field—Naturalist 85: 147-178.

Larsen, J.A. 1972a. Vegetation and terrain (environment): Canadian Boreal Forest and Tundra. U.S. Army Research Office
(Durham). 231 pp.

Larsen, J. A. 1972b. The vegetation of Northern Keewatin. Canadian Field—Naturalist 86: 45-72.

Lavkulich, L.M. 1973. Soils — vegetation — landforms of the Wrigley area N.W.T. Task Force on Northern Oil Develo
ment Report No . 73-18. Information Canada, 257 pp.

Lehmushovi, A. 1975. Methods of propagating the cowberry. Annales Agriculturae Fenniae 14: 325-333.

Lehmushovi, A. 1977. Some aspects of the cowberry trials in Finland. Annales Agriculturae Fenniae 16: 57-63.

Leiser, A. T. 1968. A mucilaginous root sheath in Ericaceae. American Journal of Botany 55: 391-398.

LeResche, R. E.and J. L. Davis. 1973. Importance of nonbrowse foods to moose on the Kenai Peninsula, Alaska. Journal
Wildlife Management 37: 279-287.

Looman, J. and K. F. Best. 1979. Budd’s flora of the Canadian Prairie Provinces, revised and enlarged edition. Hull,
Canadian Government Eublication Centre, Supply and Services, Agriculture Canada, Research Branch Publication 1662.
863 pp.

Macoun, J. 1884. Catalogue of Canadian Plants. Part II. Gamopetalae. Dawson Brothers, Montreal, Que. p. 293.

Maikawa, E. and K. A. Kershaw. 1976. Studies on lichen-dominated systems. XIX. The postfire recovery sequence of black
spruce-lichen woodland in the Abitau Lake Region, N.W.T. Canadian Journal of Botany 54: 2679-2687.

Marie-Victorin, Frére. 1964. Flore laurentienne. Deuxieme ed. Les Presses de l'Université de Montréal. Montréal, Qué., pp.
439-440.

Martin, A. C.,H. S. Zimand A. L. Nelson. 1961. American wildlife and plants. Dover Publications, Inc. New York. 358 pp.

Meusel, H. 1943. Vergleichende Arealkunde 1-2. Berlin-Zehlendorf. pp. 466 and 492.

Montgomery, F. H. 1977. Seeds and fruits of plants of Eastern Canada and Northeastern United States. University of
Toronto Press, Toronto. 233 pp.

Moss, E. H. 1953a. Forest communities in northwestern Alberta. Canadian Journal of Botany 31: 212-252.

Moss, E. H. 1953b. Marsh and bog vegetation in northwestern Alberta. Canadian Journal of Botany 31: 448-470.

Mueller-Dombois, D. 1964. The forest habitat types of southeastern Manitoba and their application to forest management.
Canadian Journal of Botany 42: 1417-1444.

462 THE CANADIAN FIELD-NATURALIST Vol. 95

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.

Nygaard, R. T. 1975. Acclimatization effect in photosynthesis and respiration, /n Fennoscanadian Tundra Ecosystems. E-
dited by F. E. Wielgolaski. Springer-Verlag, N.Y., 163-167.

Oldemeyer, J. L., A. W. Franzmann, A. L. Brundage, P. D. Arneson and A. Flynn. 1977. Browse quality and the Kenai
moose population. Journal of Wildlife Management 41: 533-542.

Oldemeyer, J. L.and R. K. Seemel. 1976. Occurrence and nutritive quality of lowbush cranberry on the Kenai Peninsula,
Alaska. Canadian Journal of Botany 54: 966-970.

Ontario Weed Committee. 1980. Guide to chemical weed control. Ontario Ministry of Agriculture Publication 75. 115 pp.

Packer, J. G.and G. D. McPherson. 1974. Chromosome numbers in some vascular plants from northern Alaska. Canadian
Journal of Botany 52: 1095-1099.

Pojar, J. 1974. Reproductive dynamics of four plant communities of sourthwestern British Columbia. Canadian Journal of
Botany 52: 1819-1834.

Polunin, N. 1948. Botany of the Canadian eastern Arctic, Part III. Vegetation and ecology. National Museum of Canada
Bulletin 104, Biological Series 32. 304 pp.

Porsild, A. E. 1937. Edible roots and berries of Northern Canada. Canada Department of Mines and Resources, National
Museum of Canada. 18 pp.

Porsild, A. E. 1962. Plant life in the Arctic. Canadian Geographical Journal 15: 121-145.

Porsild, A. E. 1964. Illustrated flora of the Canadian Arctic Archipelago. National Museum of Canada Bulletin 146. 218 pp.

Porsild, A. E. 1974. Rocky mountain wild flowers. National Museums of Canada (National Museum of Natural Sciences),
Natural History Series 2. 454 pp.

Porsild, A. E.and W. J. Cody. 1980. Vascular plants of Continental Northwest Territories, Canada. National Museums of
Canada, Ottawa, Ontario. p. 491.

Railton, J. B.and J. H. Sparling. 1973. Preliminary studies on the ecology of palsa mounds in northern Ontario. Canadian
Journal of Botany 51: 1037-1044.

Raup, H. M. 1947. The botany of Southwestern Mackenzie. Sargentia 6: Plate 33.

Reader, R. J. and J. M. Stewart. 1971. Net primary productivity of bog vegetation in southeastern Manitoba. Canadian
Journal of Botany 49: 1471-1477.

Reiners, W. A., I. A. Worley and D. B. Lawrence. 1971. Plant diversity ina chronosequence at glacier Bay, Alaska. Ecology
52: 55-69.

Rickard, W. H., J. J. Davis, W. C. Hanson and D. B. Watson. 1965. Gamma-emitting radionuclides in Alaskan tundra
vegetation, 1959, 1960, 1961. Ecology 46: 352-356.

Riewe, R. R. 1979. Interactions between wildlife, trapper-hunters and seismic lines in the Mackenzie Valley Region, NWT,
Canada. Part | Aubry-Colville Lakes. Indian and Northern Affairs Canada. Environmental! Studies, Ottawa. 114 pp.

Ritchie, J. C. 1955. Biological flora of the British Isles. Vaccinium vitis-idaea L. Journal of Ecology 43: 701-708.

Ritchie, J. C. 1956. The vegetation of Northern Manitoba I. Studies on the southern spruce forest zone. Canadian Journal
of Botany 34: 523-561.

Ritchie, J. C. 1959. The vegetation of northern Manitoba. III. Studies in the subarctic. Arctic Institute of North America.
Technical Paper No. 3. 56 pp.

Ritchie, J. C. 1960a. The vegetation of Northern Manitoba. IV. The Caribou Lake Region. Canadian Journal of Botany 38:
185-199.

Ritchie, J. C. 1960b. The vegetation of Northern Manitoba. VI. The Lower Hayes River Region. Canadian Journal of
Botany 38: 769-788.

Ritchie, J. C. 1962. A geobotanical survey of Northern Manitoba. /n. Arctic Institute of North America. Montreal.
Technical Paper 9. pp. 1-47.

Ritchie, J. C. 1974. Modern pollen assemblages near the Arctic tree line, Mackenzie Delta Region, Northwest Territories.
Canadian Journal of Botany 52: 381-396.

Ritchie, J. C. 1977. The modern and late-Quaternary of the Campbell-Dolomite Uplands, near Inuvik, N.W.T. Canada.
Ecological Monographs 47: 401-423.

Roland, A. E.and E. C. Smith. 1969. The flora of Nova Scotia. Part II. The dicotyledons. Proceedings of the Nova Scotia
Institute of Science 26: 277-743.

Rouleau, E. 1956. Studies on the vascular flora of the province of Newfoundland (Canada). I]. Some Newfoundland
vernacular plant names. Contributions de I’Institut Botanique de |’Université de Montréal, No. 69. 25-40.

Rousi, A. 1966. The use of North-European Vaccinium species in blueberry breeding. Acta agriculturae Scandinavica,
Supplementurn 16: 50-54.

Rousi, A. 1967. Cytological observations on some species & hybrids of Vaccinium. Zuchter 36: 352-359.

Rowe, J. S. 1972. Forest regions of Canada. Department of the Environment, Canadian Forestry Service, Publication 1300.
172 pp.

Ryan, A. G. 1978. Native trees and shrubs of Newfoundland and Labrador. Parks Division, Department of Tourism,
Government of Newfoundland and Labrador. St. John’s. 39 pp.

Sakai, A. and K. Otsuka. 1970. Freezing resistance of alpine plants. Ecology 51: 665-671.

1981 HALL AND SHAY: VACCINIUM VITIS-IDAEA VAR. MINUS 463

Savile, D. B. O. 1959. Notes on Exobasidium. Canadian Journal of Botany 37: 641-656.

Savile, D. B. O. 1972. Arctic adaptations in plants. Canada Department of Agriculture. Monograph 6. 81 pp.

Scoggan, H. J. 1950. The flora of Bic and Gaspé Peninsula. Québec. National Museum of Canada Bulletin 115. 292 pp.

Scoggan, H. J. 1979. The flora of Canada. National Museum of Natural Sciences. 1213 pp.

Scott, D.and W. D. Billings. 1964. Effects of environmental factors on standing crop and productivity of an alpine tundra.
Ecological Monographs 34: 243-270.

Scotter, G. W. 1965. Chemical composition of forage lichens from Northern Saskatchewan as related to use by barren-
ground caribou. Canadian Journal of Plant Science 45: 246-250.

Scotter, G. W. and W. J. Cody. 1974. Vascular Plants of Mahanni National Park & Vicinity, Northwest Territories. Le
Naturaliste Canadien 101: 861-891.

See, M. G. and L. C. Bliss. 1980. Alpine lichen-dominated communities in Alberta and the Yukon. Canadian Journal of
Botany 58: 2148-2170.

Shaver, G. R. and F. S. Chapin III. 1980. Response to fertilization by various plant growth forms in an Alaskan tundra:
nutrient accumulation and growth. Ecology 61: 662-675.

Shaver, G. R. and J. C. Cutler. 1979. The vertical distribution of live vascular phytomass in cottongrass tussock tundra.
Arctic & Alpine Research 11: 335-342.

Shay, J. M., G. McCullough and C. T. Shay. (n.d.) Biomass and nutrient content of flooded boreal forest in Long Bay,
Southern Indian Lake. Unpubl. MS. 26 pp.

Shay, J. M. and C. T. Shay. 1979. Vegetation and impact study of Deep Lake, Riding Mountain National Park. Prairie
Region Parks Canada; Department of Environment, Winnipeg. 135 pp.

Skre, O. 1975. CO, exchange in Norwegian tundra plants studied by infrared gas analyzer technique, Jn Fennoscandian
Tundra Ecosystems. Edited by F. E. Wielgolaski. Springer-Verlag, N.Y., 168-183.

Slack, N. G., D. H. Vitt and D. G. Horton. 1980. Vegetation gradients of mineratrophically rich fens in western Alberta.
Canadian Journal of Botany 58: 330-350.

Smith, D. W. 1962. Ecological studies of Vaccinium species in Alberta. Canadian Journal of Plant Science 42: 82-90.

Sorsa, V. 1962. The chromosome number of Chamaedaphne calyculata (L.) Moench, Ericaceae. Hereditas 56: 395-396.

Stark, R., I. V. Hall and P. A. Hendrickson. 1979. The partridgeberry of Newfoundland. Canadex.

Stewart, J. M. and R. Reader. 1972. Some considerations of production: Accumulation dynamics in organic terrain. Jn
Virgin peatlands. Proceedings of the 4th International Peat Congress Otaniemi, Finland, June 25-30. 247-258.

Svoboda, J. and H. W. Taylor. 1979. Persistence of cesium-137 in arctic lichens, Dryas integrifolia, and lake sediments.
Arctic and Alpine Research 11: 95-108.

Swan, J. M. A. and R. L. Dix. 1966. The phytosociological structure of upland forest at Candle Lake, Saskatchewan.
Journal of Ecology 54: 13-40. ;

Szczawinski, A. F. 1962. The heather family of British Columbia. British Columbia Provincial Museum, Victoria. pp.
188-190.

Taylor, R. L.and G. A. Mulligan. 1968. Flora of the Queen Charlotte Islands. Part 2. Cytological aspects of the vascular
plants. Canada Department of Agriculture Monograph No. 4, Part 2. p. 97.

Tear, J. 1972. Vegetativ och fruktifikatio utveckling hos vieldvaxande och odlade lingon. [ Vegetative and fruit development
of wild and cultivated cowberry]. Alfa-lavals Offsettryckeri, Tumba. 127 pp.

Thieret, J. W. 1961. A collection of plants from the Horn Plateau, District of Mackenzie, N.W.T. Canadian Field—
Naturalist 75: 77-83.

Thompson, D.C. 1980. A classification of the vegetation of Boothia Peninsula and the Northern District of Keewatin,
N.W.T. Arctic 33: 73-99.

Thornthwaite, C. W. 1931. The climates of North America according to a new classification. Geographical Review 21:
633-655.

Torrey, G. S. 1914. The partridgeberry. Department of Agriculture and Mines, St. John’s, Newfoundland. 12 pp.

Townsend, L. R.,I. V. Halland L. E. Aalders. 1968. Chemical composition of rhizomes and associated leaves of the lowbush
blueberry. Proceedings of the American Society for Horticultural Sciences 93: 248-253.

Veijalainen, H. 1976. Effect of forestry on the yields of wild berries and edible fungi, Jn Manand the Boreal Forest. Edited by
C. O. Tann. Ecological Bulletins No. 21. Swedish Natural Sciences Research Council.

Viereck, L. A. and E. L. Little. 1972. Alaska trees and shrubs. Agricultural Handbook No. 410, Forest Service U.S.D.A.
Washington, D.C. 265 pp.

Viereck, L. A. and E. L. Little. 1975. Atlas of the United States Trees Volume 2. Alaska Trees and Common Shrubs.
Miscellaneous Publications No. 1293, U.S.D.A., U.S. Government Printing Office, Washington, D.C., 19 pp. + 105 maps.

Vitt, D. H., P. Achuff and R. E. Andrus. 1975. The vegetation and chemical properties of patterned fens in the Swan Hills,
north central Alberta. Canadian Journal of Botany 53: 2776-2795.

Warming, E. 1908. The structure and biology of Arctic flowering plants. I. Ericineae (Ericaceae, Pirolaceae) I. Morphology
and biology. Meddelelser om Gronland. XXXVI. 49-53 and 121-123.

Weber, M. G. 1974. Nutrient budget changes following fire in arctic plant communities, /n Recovery of vegetation in arctic
regions after burning. Edited by R. W. Wein. Environmental-social Committee, Northern pipelines Taskforce on Northern
Development No. 74-6. Information Canada. 43-63.

464 THE CANADIAN FIELD-NATURALIST Vol. 95

Wein, R. W. and L.C. Bliss. 1973a. Changes in arctic Eriophorum tussock communities following fire. Ecology 54:
845-852.

Wein, R. W.and L. C. Bliss. 1973b. Experimental crude oil spills on arctic plant communities. Journal of Applied Ecology
10: 671-682.

Wein, R. W. and L. C. Bliss. 1974. Primary production in arctic cottongrass tussock tundra communities. Arctic and Alpine
Research 6: 61-274.

Wielgolaski, F. E. 1974. Phenological studies in tundra, Jn Phenology and Seasonality Modeling. Edited by H. Lieth.
Springer-Verlag, New York. 209-214.

Wielgolaski, F. E.and L. Karenlampi. 1975. Plant phenology of Fennoscandian Tundra Areas, Jn Fennoscandian Tundra
Ecosystems. Edited by F. E. Wielgolaski. Springer-Verlag, New York. 94-102.

Wielgolaski, F. E. and S. Kjelvik. 1975. Energy content and use of solar radiation of Fennoscandian tundra plants. Jn
Fennoscandian Tundra Ecosystems. Edited by F. E. Wielgolaski. Springer-Verlag, New York. 201-207.

Wielgolaski, F. E., S. Kjelvik and P. Kallio. 1975. Mineral content of tundra and forest tundra plants in Fennoscandia. Jn
Fennoscandian Tundra Ecosystems. Edited by F. E. Wielgolaski. Springer-Verlag, New York. 316—332.

Zoltai, S. C. and C. Tarnocai. 1971. Properties of a wooded palsa in Northern Manitoba. Arctic and Alpine Research 3:
115-129.

Revision received 5 August 1981
Accepted 23 October 1981

Notes

Observations on Black-crowned Night Heron Breeding

Success in a North Dakota Marsh

RAYMOND J. GREENWOOD

Northern Prairie Wildlife Research Center, Jamestown, North Dakota 58401, U.S.A.

Greenwood, Raymond J. 1981 Observations on Black-crowned Night Heron breeding success in a North Dakota marsh.

Canadian Field-Naturalist 95(4): 465-467.

Storms and mammalian predation severely reduced production in a Black-crowned Night Heron (Nycticorax nycticorax)
colony in two years and drought prevented nesting in two other years from 1976 to 1980. Maximum estimated production was
2.2 young per pair in 1978, the only year production was sufficient to maintain the population.

Key Words: Black-crowned Night Heron Nycticorax nycticorax, productivity, marsh, weather, predation.

In the northern great plains Black-crowned Night
Herons (Nycticorax nycticorax) commonly nest in
prairie marshes (Stewart 1975; Wolford and Boag
1971) where success is influenced by factors such as
predation (Eberhardt and Sargeant 1977; Greenwood
1981; Wolford and Boag 1971), unstable wetland con-
ditions (Swanson and Meyer 1977), and violent
storms (Johnson 1979; Piehl 1979). Henny (1972)
assumed that the average annual recruitment rate
among Black-crowned Night Herons was 2.0-2.1
young per pair, based on limited information from
large east coast colonies where nests are located in
trees. I found very low nesting success in a Black-
crowned Night Heron colony in east-central North
Dakota during 1976. The observation prompted me to
return to the colony in subsequent years for additional
information. The present report describes my obser-
vations for 1976-1980, and discusses some factors that
influenced breeding success.

Study Area and Methods

The colony was in a 306 ha semi-permanent marsh
in Griggs County, 4 km southwest of Walum, North
Dakota. The area has gently rolling topography and is
intensively farmed for cereal grain (Greenwood 1981).
Several other large marshes are nearby. Black-
crowned Night Heron nests were concentrated in a
7.5 ha clump of Phragmites (Phragmites communis)
that was usually in standing water; water depth at the
colony site ranged from | m in 1976 to nil in 1977.
Cattail (Typha sp.) and Bulrush (Scirpus sp.) were
other dominant emergent plant species in the marsh.
Weather observations are from the U.S. Weather Ser-
vice station at Cooperstown, North Dakota, 42 km
northeast of the marsh.

465

I visited the colony on 30 June 1976, 1 July 1977, 29
June 1978, 12 July 1979, and 5 July 1980. On each visit
nests, eggs, and young were counted; age of young was
estimated by comparison with illustrations of
Mc Vaugh (1972). Iattempted to determine causes and
timing of mortality among dead nestlings.

Numbers of breeding pairs and production of
fledged young had to be estimated because I visited
the colony only once each year. Numbers of breeding
pairs was based on the total number of occupied nests
observed. Occupied nests contained at least one egg or
nestling. All nestlings alive at time of check were
assumed to fledge. Annual production per pair was
estimated by dividing number of young assumed to
fledge by the estimated number of breeding pairs.

Results and Discussion

Annual observations of nests and nestlings are
summarized in Table 1. Nests were loosely con-
structed of Phragmites stems and leaves, and were
elevated about 25-35 cm above water. No nests were
found in 1977 or 1980. In 1977 the marsh was dry
during the entire nesting period. In 1980 water depth
was 0.8 m at the colony site, reflecting adequate pre-
cipitation received in 1979 (Table 2). However, very
little precipitation was received in April and May
1980, and temperatures those months were above
normal. I believe the dry conditions and hot weather
prevented nesting.

Black-crowned Night Herons nested in 1976, 1978,
and 1979, but nest success was low in 1976 and 1979.
Size of largest nestlings found each year (28-36 days
old) indicated that egg laying began annually in late
April or early May, consistent with observations
reported by Stewart (1975) and Wolford and Boag

466

TABLE 1. Productivity of a Black-crowned Night Heroncol-
ony in a prairie marsh in east-central North Dakota,
1976-1980.

1976 1977 1978 1979 1980
Total nests 57 0 ay) Sy 0
Occupied nests* 30 — 46= = 53 _
with eggs only 95 — 7 11 —
with eggs and
nestlings 0 — 2 7 —
with nestlings only 20 — SY 35 —
with dead nestlings 19 — 3) 30 —
with predation or
scavenging 120 — 0 11 —
Total eggs a ey) —
Total nestlings 390 105 80 —
Live nestlings 7 — 99 30 a
Estimated fledglings/
pair. 032335" — 7 2:205, 0:57.

“Contain at least | egg or nestling.

Not recorded.

“Total live nestlings/estimated breeding pairs (occupied
nests).

TABLE 2. Mean April-June temperature and precipitation at
Cooperstown, North Dakota during 1976-1980.

Long-
term
mean 1976 1977 1978 1979 1980

April 5.3 RO NOB S54 OO. Dat
ilempi()) Maye) 125) 4s 18955 15.6) 79:5) 1620
dine = NWA M57 | MW@.3 MBS eID Used)

April 3.6 3.6 1.0 3.5 8.1 0
Ppt (cm) May 6.5 2 8.1 6.0 8.6 1S)
June 9.7 8.7 eal OG Wks} 7

(1971) for other colonies at northern latitudes. How-
ever, each year many broods with smaller nestlings
were also present, a few as young as six days old,
possibly indicating asynchrony in laying, but more
likely the result of renesting after loss of initial
clutches.

Adverse weather and mammalian predation were
important causes of egg and nestling losses. Some
nests in 1976, 1978, and 1979 were unoccupied when
found, were not soiled with feces, and probably never
had contained nestlings. A few of those nests may
have been built by subadults. Meyerriecks, as cited by
Henny (1972), stated that “many yearlings come to the
nesting colonies, a few try to build nests, but I have not
seen breeding by this age group.” However, many
unoccupied nests may have resulted from undetected
predation on eggs.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Major storms on 28 June 1976, and 28 June and 3
July 1979 appeared to have influenced nesting. Prairie
storms usually are accompanied by violent winds and
frequently hail, and are known to kill birds (Johnson
1979; Piehl 1979). During the 1976 and 1979 visits, 82
and 63% of the nestlings observed were dead, and
most appeared to have died simultaneously. In 1976
many were in the water and were just beginning to
decompose, indicating recent death. In 1979 carcasses
were also in the water, but were in a more advanced
stage of decomposition than in 1976 (nine days had
elapsed since the most recent storm). Either of the
1979 storms or both could have been responsible for
the mortality. Two adult Black-crowned Night Her-
ons also were found dead near nests in 1979, appar-
ently victims of hail.

Of the occupied nests observed, 40% in 1976 and
21% in 1979 showed direct evidence of mammalian
predation or scavenging in addition to storm damage;
no such predation was observed in 1978. Evidence
consisted of eggs opened by mammalian predators
and partly consumed nestlings. In 1976 the partly
consumed nestlings were sufficiently intact to show
they died at nearly the same time as others found
nearby, but I was unable to determine if they had been
killed by predators or the storm. Predators responsi-
ble were probably Raccoons (Procyon lotor) that fre-
quently consume carrion (Greenwood 1981), but
Mink (Mustela vison) also inhabited the marsh. In
1976 foraging by radio-equipped Raccoons was doc-
umented in the colony on three occasions during the
nesting season. An adult female foraged in the colony
on the night of 16-17 June before I collected her in
early morning. Stomach contents showed she had
consumed 12 Black-crowned Night Heron embryos.
A yearling and an adult male both foraged in the
colony on the night of 29-30 June. Several partly
consumed nestlings were found when I investigated
the next morning. The yearling foraged in the colony
again on the night of 30 June—1 July. When collected
the next morning its stomach contained shell and
contents of putrefied Black-crowned Night Heron
eggs.

Estimated production during the five years ranged
from 0 to 2.2 fledged young per breeding pair. The
weighted average for the three years when nesting
occurred was only 1.1 young per pair. Tremblay and
Ellison (1979) found that human disturbance at
Black-crowned Night Heron colonies early in the nest-
ing season inhibited laying, but my visits were all late
in the season.

Productivity estimates based on single observations
are admittedly less sound than estimates derived from
more intensive studies; however, only successful re-
nesting after mid-July (highly unlikely) could have

1981

raised my annual estimates of production substan-
tially. Black-crowned Night Herons in North Dakota
usually terminate nesting at the end of June (Stewart
1975). More precise estimates of the numbers of
breeding pairs (probably higher than my minimum
counts based on occupied nests) would have resulted
in lower estimates of annual production than I calcu-
lated. Additional mortality among nestlings after my
visits would also have lowered production estimates.
The productivity I observed in all years but 1978 was
far below the maintenance level of 2.0-2.1 young per
pair suggested by Henny (1972). Wolford and Boag
(1971) believed that the pioneering population they
studied could not have maintained itself at the level of
recruitment they observed. My observations suggest
similar status for a small population of Black-
crowned Night Herons in a North Dakota marsh.

Acknowledgments

I thank the following persons who assisted in nest
counts; B. Bicknell, J. Choromanski, B. Garlinger, M.
Goos, S. Hetherington, S. Evanoff, C. Lackney, L.
Pellack, J. Piehl, and D. Vanderhoef. C. Faanes, D.
Johnson, and A. Sargeant kindly reviewed early
drafts of the manuscript, and offered many construc-
tive comments.

Literature Cited
Eberhardt, L. E.and A. B. Sargeant. 1977. Mink predation

NOTES

467

on prairie marshes during the waterfowl breeding season.
In Proceedings of the 1975 predator symposium. Edited by
R. L. Phillips and C. Jonkel. University of Montana, Mis-
soula. pp. 33-43.

Greenwood, R. J. 1981. Foods of prairie Raccoons during
the waterfowl nesting season. Journal of Wildlife Man-
agement 45: 754-760.

Henny, C. J. 1972. Ananalysis of the population dynamics
of selected avian species. U.S. Fish and Wildlife Service
Wildlife Research Report Number |. 99 pp.

Johnson, D. H. 1979. Effects of a summer storm on bird
populations. Prairie Naturalist 11: 78-82.

McVaugh, W., Jr. 1972. The development of four North
American herons. Living Bird 11: 155-173.

Piehl, J. L. 1979. Avian mortality froma severe hail storm.
Prairie Naturalist 11: 75-77.

Stewart, R. E. 1975. Breeding birds of North Dakota. Tri-
College Center for Environmental Studies, Fargo, North
Dakota. 295 pp.

Swanson, G. A.and M. I. Meyer. 1977. Impact of fluctuat-
ing water levels on feeding ecology of breeding Blue-
winged Teal. Journal of Wildlife Management 41:
426-433.

Tremblay, J.and L. N. Ellison. 1979. Effects of human dis-
turbance on breeding of Black-crowned Night Herons.
Auk 96: 364-369.

Wolford, J. W. and D. A. Boag. 1971. Distribution and
biology of Black-crowned Night Herons in Alberta. Ca-
nadian Field-Naturalist 85: 13-19.

Received 19 January 1981
Accepted 19 October 1981

Historical Nest Records for the Ferruginous Hawk

in Manitoba
MARC J. BECHARD

Department of Biological Sciences, Marshall University, Huntington, West Virginia 25701

Bechard, Marc J. 1981. Historical nest records for the Ferruginous Hawk in Manitoba. Canadian Field-Naturalist 95(4):

467-469.

Seven previously unreported Ferruginous Hawk clutches from Manitoba have been located in museum egg collections. These
together with published records clarify the hawk’s past breeding history and provide stronger evidence for its disappearance

from the province.

Key Words: Ferruginous Hawk Buteo regalis, nest records, Manitoba, population decline.

The Ferruginous Hawk (Buteo regalis) was once
widely distributed in grassland and steppe areas of the
western United States and southwest Canada (Bent
1937; Olendorff 1973; Snow 1974). Its present status is
defined as “suffering from population declines and
habitat diminutions in all parts of its range” (Ano-
nymous 1979). Populations in Canada appear to have
declined in areas subject to intensive agriculture and

human activity (Fyfe 1976; Houston 1978). In Mani-
toba, cultivation has removed much of the native
prairie habitat, and the Ferruginous Hawk seems to
have experienced a decline (Criddle 1929; Knapton
1979). This note provides evidence for the historic
breeding population and confirms the disappearance
of Ferruginous Hawk in Manitoba.

Museums containing bird egg collections with egg

468 THE CANADIAN FIELD-NATURALIST

sets from western Canada (Kiff 1979) were contacted
to obtain data on past Ferruginous Hawk nests in
Manitoba. Nest records accompanying egg sets in the
Western Foundation of Vertebrate Zoology and
American Museum of Natural History include seven
previously unreported Ferruginous Hawk nests in
southern Manitoba prior to 1924. C. P. Forge, who
farmed near Carman, Manitoba (Forge 1901; 1902),
collected three eggs on 10 June 1907 froma nest 8.1 m
above ground in an elm (U/mus sp.) by the Boyne
River near Hyde Park in Dufferin municipality. Hyde
Park was a temporary school district (#916) three to
nine miles north of present Roseisle, Manitoba. J. D.
Currie found four nests with a mean of 4.0 eggs per
nest on 10 May 1908 in prairie habitats of southwest-
ern Manitoba. Perhaps because there were then few
villages in that part of the province, specific locations
for these nests were not given; one was reported to be
in the extreme southwest corner of the province.
Three were ground nests and one was built 2.0 m
above ground ina willow (Salix sp.). On 20 May 1923,
D. Ogg located a nest containing four eggs 3.8 m
above ground in a balm of Gilead (Populus tacama-
hacca) and on 2 may 1924 he found a nest with four
eggs 4.0 m above ground in a poplar (Populus sp.).
Both nests were near the town of Pierson, Manitoba.

Descriptions of the construction and dung contents
of all these nests indicate them to be typical of Ferrug-
inous Hawk. As an additional check of collector iden-
tifications, I compared volumes of these eggs with
those of Buteo species nesting in Manitoba and adja-
cent areas of Saskatchewan and North Dakota.
Volumes were calculated using the equation of
Romanoff and Romanoff (1949). Mean volumes of
77.30 (n = 20 clutches), 59.55 (n = 20), 69.35 (n = 20),
and 61.26(n = 11) ml were obtained for Ferruginous,
Swainson’s (Buteo swainsoni), Red-tailed (Buteo
jJamaicensis), and Rough-legged (Buteo lagopus)
Hawks, respectively. Mean egg volumes of 78.05,
78.94, 73.03, 78.66, 78.12, and 76.65 ml for six of the
seven clutches I am reporting were too large for any
species except Ferruginous Hawk and substantiated
the accuracy of collector identifications. Eggs from
one of the four nests reported by J. D. Currie have
been lost. Since the other egg sets he collected appear
to be accurately identified, I assumed that nest also to
be valid.

Before settlement, southwestern Manitoba sup-
ported mixed-grass prairie habitats interspersed with
temporary and semi-permanent ponds and sloughs
(Knapton 1979; Weir 1960). Trees were limited to
river bottoms and “bluffs” in marshy areas. Indica-
tions are that until the 1920’s the Ferruginous Hawk
was not an infrequent breeding resident, nesting
primarily along rivers and at the edges between prairie

Vol. 95

and forest habitats. Coues (1878) and Thompson
(1891) listed the Ferruginous Hawk as a probable
Manitoban, breeding in open prairie at the edge of the
Pembina Mountains. Hales (1927) found it in wooded
areas along the Assiniboine and Souris Rivers. W. E.
Godfrey (pers. comm.) has seen, courtesy of Vere
Hunt Scott of Winnipeg, two photographs of nestling
hawks taken by Talbot Criddle near the family’s farm
at Aweme on 4 July 1917. Although labelled as
Rough-legged Hawks, they are clearly identifiable as
Ferruginous Hawks. Norman Criddle (1920), Talbot’s
brother, also reported Ferruginous Hawks to be
common near Aweme in and around the Spruce
Woods Timber Reserve. Those reports, with the
records reported herein, indicate that the hawk nested
throughout southwestern Manitoba from the Red
River to the Saskatchewan border. With settlement
and the spread of agriculture, the Ferruginous Hawk
started a decline and by 1928 was “less common than
in 1890” (Criddle 1929). Details subsequent to 1930
are not available, but failure to locate any active nests
in southwestern Manitoba in the 1970’s indicate that
the Ferruginous Hawk has now ceased to be a breed-
ing resident of the province (Knapton 1979). Similar
Ferruginous Hawk declines have been reported in
Washington, Oregon, Idaho, Utah, and North
Dakota where agriculture has altered nesting habitats
(Snow 1974; Stewart 1975; Powers and Craig 1976;
Anonymous 1979). The decline in Manitoba also
appears to be related to the loss of nesting habitat.
Loss of native prairie due to invasion by forests (Bird
1961) and increased areas of cultivation may have
destroyed or made unsuitable habitats, nest sites, and
food supplies formerly used for Ferruginous Hawk
nesting (Schmutz and Schmutz 1980).

I am grateful to C. S. Houston for suggesting this
study, and for providing ideas and material concern-
ing the historical geography and biography, with the
help of Peter Bower and Barry Hyman of the Public
Archives of Manitoba. Lloyd Kiff assisted in locating
pertinent egg sets and data in the Western Foundation
of Vertebrate Zoology. W. E. Godfrey contributed
the Talbot Criddle nest record and R. W. Nero pro-
vided constructive criticism. John Bull of the Ameri-
can Museum of Natural History is acknowledged for
his help with nest records.

Literature Cited

Anonymous. 1979. The blue list of 1980. American Birds 33:
832.

Bent, A. C. 1937. Life histories of North American Birds of
prey. Order Falconiformes (Part 1). United States
National Museum Bulletin 167.

Bird, R. D. 1961. Ecology of the aspen parkland of western
Canada. Publication 1066. Canada Department of Agri-
culture, Queen’s Printer, Ottawa.

1981

Coues, E. 1878. Field notes on birds observed in Dakota and
Montana along the forty-ninth parallel during seasons of
1873 and 1874. Bulletin U.S. Geologic and Geographical
Survey of Territories. Volume 4, Article 25. U.S. Govern-
ment Printing Office, Washington, D.C.

Criddle, N. 1920. Birds that are little known in Manitoba.
Canadian Field-Naturalist 35: 133-134.

Criddle, N. 1929. Memoirs of the eighties. Canadian Field-
Naturalist 43: 176-181.

Forge, C. P. 1901. Field notes from Manitoba. Oologist 18:
26-28, 41-42, 155-156.

Forge, C. P. 1902. Field notes from Manitoba. Oologist 19:
5-8, 49-52, 152.

Fyfe, R. W. 1976. Status of Canadian raptor populations.
Canadian Field-Naturalist 90: 370-375.

Hales, B. J. 1927. Prairie birds. MacMillan, Toronto.

Houston, C. S. 1978. Changing patterns of raptor popula-
tions with settlement. Jn Nature and change on the Cana-
dian plains. Proceedings of the 1977 annual meeting of the
Canadian Nature Federation. Edited by W. A. Davies.
Canadian Plains Proceedings 6, Canadian Plains
Research Center, Regina.

Kiff, L. F. 1979. Bird egg collections in North America. Auk
96: 746-755.

Knapton, R. W. 1979. Birds of the Gainsborough-Lyleton
region. Saskatchewan Natural History Society, Special
Publication 10.

Olendorff, R. R. 1973. The ecology of nesting birds of prey in

NOTES

469

northeastern Colorado. U.S. International Biological
Program. Technical Report 211. Natural Resources Ecol-
ogy Laboratory, Colorado State University, Fort Collins,
Colorado.

Powers, L. R. and T. H. Craig. 1976. Status of nesting Fer-
ruginous Hawks in the Little Lost River valley and vicin-
ity, southeastern Idaho. Murreiet 57: 26-31.

Romanoff, A. L. and A. J. Romanoff. 1949. The avian egg.
Wiley and Sons, New York, New York.

Schmutz, J. K. and S. M. Schmutz. 1980. Status of the Fer-
ruginous Hawk. Committee on Status of Endangered
Wildlife in Canada.

Snow, C. 1974. Habitat management series for unique and
endangered species. Report 13, Ferruginous Hawk. U.S.
Department of the Interior, Bureau of Land Management,
Denver, Colorado.

Stewart, R. E. 1975. Breeding birds of North Dakota. Tri-
College Center for Environmental Studies, Fargo, North
Dakota.

Thompson, E. E. 1891. The birds of Manitoba. Proceedings
United States National Museum. 13: 457-643.

Weir, T. R. 1960. Economic atlas of Manitoba. Manitoba
Department of Industry and Commerce, Winnipeg,
Manitoba.

Received 3 April 1981
Accepted 4 September 1981

Nesting of Smith’s Longspurs in British Columbia

KATHY MARTIN!, SUSAN HANNON’, and RICHARD MOSES?

'Biology Department, Queen’s University, Kingston, Ontario K7L 3N6
2Department of Zoology, University of British Columbia, Vancouver, British Columbia V6T 1W5

Martin, Kathy, Susan Hannon, and Richard Moses. 1981. Nesting of Smith’s Longspurs in British Columbia. Canadian

Field-Naturalist 95(4): 469-470.

In 1979 and 1980, 7 nests of Smith’s Longspurs were found in the Chilkat Pass area, the first confirmed breeding records in

British Columbia.

Key Words: Smith’s Longspur, Calcarius pictus, breeding range, British Columbia.

Smith’s Longspurs (Calcarius pictus) nest in grassy
and hummocky tundra from Alaska (Kessel and Gib-
son 1978), east across the Yukonand Northwest Terri-
tories, to the northwest border of Hudson Bay (God-
frey 1966). There is limited information on the south-
ern extent of their breeding range in Alaska (Sage
1976) and indirect evidence for the southern Yukon
(Hoefs 1973). In British Columbia, J. B. Foster pho-
tographed an adult male on 11 July 1975 at Fire Flats,
Spatzizi (B.C.P.M. Photo #394), and a group of 10
adults and flying young was seen in this area by B. S.
Ford on 12 August 1976 (BCNRS). An adult male
collected near Atlin by I. McT. Cowan on 9 June 1958

is in the University of British Columbia Museum
(CVM #8580). Evidence of breeding in the extreme
northwest of British Columbia, although of long
standing, has been indirect. C.H.D. Clarke on 17 July
1944 collected a female Smith’s Longspur witha large
brood patch at Kelsall Lake, km 120, Haines Road
(Rand 1948). Since then, pairs and singing males have
been observed irregularly in this area during the
breeding season (Weeden 1960, S. G. Cannings in
Grunberg 1979).

During June and July, 1979 and 1980, we found six
Smith’s Longspur nests near Chilkat Pass in north-
western British Columbia. Five nests were discovered

470

THE CANADIAN FIELD-NATURALIST

Vol. 95

TABLE 1. Date found, stage of development, clutch size, and outcome of Smith’s Longspur nests found in the Chilkat Pass

area of northwestern British Columbia.

Nest-Year Stage when Clutch

no. Date found found size Outcome

1-79 18 June eggs 4 unknown

1-80 25 June hatching 6 3 young fledged 9-14 July

2-80 25 June eggs 5 hatched 26 June, 4 young 3 July; outcome unkown
3-80 28 June eggs 5 hatched 29 June, 5 young fledged 8-11 July

4-80 2 July nestlings — 2 young fledged 2-5 July

5-80 1 July eggs 5) unknown

6-80! 27 June nestlings — 4 very small young; outcome unknown

'Nest found by R. W. Campbell, British Columbia Provincial Museum.

in an area of approximately 60 ha close to Kelsall
Lake, km 128, Haines Road. The sixth was located at
Chilkat Pass, 25 km to the south. Nests were found
accidentally by flushing incubating or brooding
female longspurs while we were studying Willow
Ptarmigan (Lagopus lagopus). Species identification
was confirmed by R. W. Campbell of the British
Columbia Provincial Museum, and photographs of
adults at and near a nest were taken by E. Siam
(BCPM Photo No 661). A seventh nest was found by
R. W. Campbell near km 103, Haines Road, on 27
June 1980.

Nests were in subalpine tundra characterized by
shrub cover of willow (Salix spp.), Dwarf Birch (Bet-
ula glandulosa) and Shrubby Cinquefoil (Potentilla
fruticosa), and a ground cover of sedges, lichens,
mosses and perennial forbs. The nests, built on flat
exposed sites or in the sides of hummocks, were lined
with dried sedges, grasses, and three contained several
Willow Ptarmigan feathers.

We found nests on 18 June 1979 and between 25
June and 2 July 1980, and revisited four of them
several times (Table 1). Clutch size varied from 4 to 6
with a mean of 5.0 eggs, which is larger than the mean
clutch of 4.0 eggs (N = 26)! for Smith’s Longspurs
nesting near Churchill, Manitoba (Jehl 1968). Period
of hatch (25-29 June) for the Chilkat nests in 1980 was
about 5 days earlier than that described by Jehl.

Female longspurs at Churchill engaged in distrac-
tion displays when flushed from the nest during incu-
bation (Jehl 1968). In the Chilkat Pass, females flew
low and inconspicuously for a long distance, some-
times out of sight, when flushed from the nest. No
defense of eggs or young was observed, except at one
nest with two large nestlings, and here parental scold-
ing was minimal.

Smith’s Longspurs probably breed regularly in the
Chilkat Pass. We observed other females and singing
males for which nests were not located. The southern
limit of the breeding range is poorly defined, but
Smith’s Longspurs probably nest throughout the
southern regions of Alaska and the Yukon Territory,

and in northwestern British Columbia, wherever there
is suitable habitat.

We thank R. Wayne Campbell, Sydney G. Can-
nings, and Robert Montgomerie for critiques of the
manuscript. Brina Kessel and C. Eric Tull provided
helpful references. Grants from the Arctic Institute of
North America; the Canadian Wildlife Service and
the Frank M. Chapman Fund of the American
Museum of Natural History to S. Hannon supported
the ptarmigan project.

Literature Cited

Godfrey, W.E. 1966. The birds of Canada. National
Museum of Canada Bulletin 203. 428 pp.

Grunberg, H. 1979. Northwestern Canada Region. Jn: The
nesting season: June I—July 31, 1979. American Birds 33:
881.

Hoefs, M. 1973. Birds of the Kluane Game Sanctuary,
Yukon Territory, and adjacent areas. Canadian Field-
Naturalist 87: 345-355.

Jehl, J. R., Jr. 1968. Breeding biology of Smith’s Longspur.
Wilson Bulletin 80: 123-149.

Kessel, B.,and D. D. Gibson. 1978. Status and distribution
of Alaska birds. Studies in Avian Biology, No. 1, Cooper
Ornithological Society.

Rand, A. L. 1948. Distributional notes on Canadian birds.
Canadian Field-Naturalist 62: 175-180.

Sage, B. L. 1976. The breeding distribution of Smith’s
Longspur in Alaska. Condor 78: 116-117.

Weeden, R. B. 1960. The birds of Chilkat Pass, British
Columbia. Canadian Field-Naturalist 74: 119-129.

Received 28 January 1981
Accepted 22 March 1981

Addenda: While this manuscript was in press D. Mossop
communicated to the authors the following unpublished
records: Chilkat Pass: 22 June 1971, nest with 4 eggs; 11
August 1972: adults feeding two fledged young; Aishihik
Lake, 80 km W of Whitehorse; 2 August 1979, adult feeding a
fledged young.

1 We omitted one 2-egg clutch from the calculation since Jehl
(1968) reported that probably it had been reduced in size by
predation.

1981 NOTES 471

Myiasis by Wohlfahrtia opaca (Coq.):
A Cause of Mortality of Newly Hatched Wild Ducklings

GARY WOBESER!, ALVIN GAJADHAR?, GERRY W. BEYERSBERGEN?, and LAWSON G. SUGDEN?

\Department of Veterinary Pathology, Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon,
Sask. S7N 0WO
2Department of Veterinary Microbiology, Western College of Veterinary Medicine, University of Saskatchewan, Saskatoon,
Sask. S7N 0WO
3Prairie Migratory Bird Research Centre, Canadian Wildlife Service, Environment Canada, 115 Perimeter Road, Saskatoon,
Sask. S7N 0X4

Wobeser, Gary, Alvin Gajadhar, Gerry W. Beyersbergen and Lawson G. Sugden. 1981. Myiasis by Wohlfahrtia opaca
(Cogq.): a cause of mortality of newly hatched wild ducklings. Canadian Field-Naturalist 95(4): 471-473.

Three instances of fatal myiasis involving recently hatched Northern Shoveler (Anas clypeata) and Blue-Winged Teal (Anas
discors) ducklings were found in central Saskatchewan during June 1980. Adult flies reared from the larvae were identified as
the flesh fly Wohlfahrtia opaca (Coq.) (Diptera:Sarcophagidae); a species that has been reported as a cause of myiasis in
mammals, but not in birds. The condition occurred in approximately 3% of the successful duck nests and 0.7% of ducklings

known to be present on the area.

Key Words: ducklings, mortality, myiasis, Wohlfahrtia opaca (Coq.), Anas clypeata, Anas discors.

The causes of mortality of ducklings after hatching
are seldom known in detail, although some brood
attrition usually occurs between hatching and fledg-
ing. This lack of detailed information relates to diffi-
culties in finding sick or dead ducklings before their
removal by scavengers or predators. Myiasis, “the
infestation of live human or vertebrate animals with
dipterous larvae, which, at least for a certain period,
feed on the host’s dead or living tissue, liquid body
substances, or ingested food” (Zumpt 1965), has been
documented as a sporadic cause of mortality of young
mammals (Capelle 1971), and reported in domestic
waterfowl (Zumpt 1965). We are unaware of any des-
criptions of the condition in wild waterfowl. We
report here three instances of myiasis, involving the
death of seven ducklings, that occurred in central
Saskatchewan in 1980.

Study Area and Methods

The ducklings were found during intensive searches
for duck nests on a 388.5 ha Canadian Wildlife Ser-
vice study area near St. Denis, Saskatchewan. The
area is characterized by agricultural land interspersed
with native vegetation and numerous small wetlands.
Ducklings found dead were necropsied and histologic
examination was done in some cases. Some fly larvae
removed from the carcasses were identified to genus
on the basis of external morphology, posterior spira-
cle pattern and cephaloskeletal structure (James and
Gassner 1947, Faust and Russell 1964, Zumpt 1965).
The remaining larvae were reared to adult by two
methods. Some second- and third- instar larvae were
placed on sterile peat moss in fly-proof containers and

held in the dark at room temperature. The remaining
second- and third- instar larvae were placed on the
surface of 5% sheep blood agar in petri plates (stand-
ard microbiological media) in fly-proof containers
and held in the dark at room temperature. Flies that
emerged were identified with the key provided by
James and Gassner (1947).

Results

On 11 June 1980, four Northern Shoveler (Anas
clypeata) ducklings were found dead on the shore of a
small wetland. The birds were in a group, as though
the female had been brooding them at the location; at
least one additional live duckling was present nearby.
On the following day, 12 June, two Blue-winged Teal
(Anas discors) ducklings were found dead on the
shore of another wetland approximately 100 m from
where the first birds were found. On 24 June a par-
tially pipped Blue-winged Teal egg containing a dead
embryo was found in a nest in the same general area.
The remaining eggs from this nest had apparently
hatched normally.

The gross findings were similar in all hatched duck-
lings, differing only in number and size of fly larvae
present. There was wetting of the down with serous
exudate on the ventral abdominal wall at the site of
the umbilicus. A small circular opening, varying from
0.3—1.0 cm in diameter, was present in the skin at this
site. Active fly larvae were evident within this opening
and small 1-3 mm white larvae were present on the
skin around the opening in a few of the birds. On
dissection, there was an extensive subcutaneous
“pocket” containing maggots on the ventral abdomi-

472

nal wall and extending into the space between the
abdominal wall and legs. This cavity contained a small
amount of serous yellow-brown fluid. In one Blue-
winged Teal, larvae had entered the body cavity. The
fly larvae within this pocket varied from 0.3-1.1 cmin
length; 16, 23, 18 and 4 were recovered from the four
Northern Shovelers and 11 and 23 were found in the
two Blue-winged Teal.

The partially pipped egg was cracked in the normal
hatching pattern for approximately one-half of its
circumference; the tip of the duckling’s bill protruded
through the opening. A 0.6 cm perforation was pres-
ent in the duckling’s skin immediately caudo-ventral
to the left wing. A total of 12 fly larvae ranging in
length from 0.3 to 1.0 cm was present in the subcu-
taneous space underlying this perforation and within
the body cavity. There was extensive intra-hepatic
hemorrhage.

There was no gross or histologic evidence of other
disease in these ducklings.

The larvae examined from all ducklings were identi-
fied as those of Wohlfahrtia spp. The second-instar
larvae placed on peat moss failed to develop, but most
of the third-instar larvae placed on this substrate
pupated and developed to adult. Larvae placed on
blood agar burrowed in, and apparently digested, the
agar. The third-instar larvae pupated within 5 days,
most of the second-instar larvae molted and then
pupated within | to 2 weeks. Adult flies emerged from
both media after 10 to 28 days. Approximately 90% of
the adult flies that emerged were identified as the flesh
fly Wohlfahrtia opaca. A few flies (Family Callipho-
ridae) also emerged, but they were not identified
further.

Discussion

Wohlfahrtia opaca is widespread in western North
America and has been reported from cases of cutane-
ous myiasis in several mammals including man
(Gassner and James 1948, Haufe and Nelson 1957).
No reports were found of infection with this species in
birds. The fly is larviparous and the larvae are consi-
dered to be obligatory parasites of warm-blooded ver-
tebrates, although development may continue in the
carcass after death of the host (Gassner and James
1948). First-instar larvae can penetrate the unbroken
skin of young animals (Gassner and James 1948); in
the ducklings the larvae appeared to have gained
entrance through the umbilicus. Ford (1933) found
that females of a closely related species, W. vigil
(Walker), were attracted to the eyes, perhaps due to
moist exudate in this area; females of W. opaca may
deposit their larvae near the umbilicus of recently-
hatched ducklings, or through an opening in an egg,
for the same reason. The cause of death in myiasis is

THE CANADIAN FIELD-NATURALIST

Vol. 95

unclear; however, young cottontail rabbits (Sy/vila-
gus floridanus) died within 3 days after infection by
W. vigil (Yuill and Eschle 1963) and ranch fox pups
infected with as few as four larvae of W. opaca may die
(James and Kraft 1964).

The recognition of three instances of myiasis on one
small area within a 2 week period suggests that the
condition may not be rare in ducklings. Altogether,
110 nests in which ducklings were known to have
hatched were found on the area, and we estimate that
approximately 1000 ducklings hatched from these
nests. The three instances of myiasis found thus
represented occurrence of the disease in almost 3% of
known nests and in 0.7% of ducklings known to have
hatched.

The lack of prior reports of myiasis in young water-
fowl is understandable, because dead ducklings are
difficult to find and the presence of maggots ona dead
duckling would not seem unusual. A careful examina-
tion must be performed and the larvae identified to
differentiate between the presence of species known to
cause myiasis and the development of saprophytic fly
larvae on a carcass dead from other causes. The few
calliphorid flies reared from the ducklings probably
represented the latter situation.

Myiasis caused by larvae of Wohlfahrtia spp. can
produce severe losses among the young of domestic
species, particularly mink and foxes, but the signifi-
cance of these parasites for wild populations is
unknown. Yuill and Eschle (1963) suggested that
infection with W. vigil may affect populations of cot-
tontails adversely, but this remains unproven. Boon-
stra (1977) believed that W. vigil had a relatively
minor effect on a population of moles (Microtus
townsendii), although up to 5.4% of the population
was parasitized and only 5% of infected individuals
were known to have lived for 2 weeks.

Literature Cited

Boonstra, R. 1977. Effect of the parasite Wohlfahrtia vigil
on Microtus townsendii populations. Canadian Journal of
Zoology 55: 1057-1060.

Capelle, K. J. 1971. Myiasis. In Parasitic diseases of wild
mammals. Edited by J. W. Davis and R. C. Anderson.
Iowa State University Press, Ames. pp. 279-305.

Faust, E. C. and P. F. Russell. 1964. Craig and Faust’s
Clinical Parasitology. 7th Edition. Lea and Febiger, Phi-
ladelphia. 1099 pp.

Ford, N. 1933. Observations on the behaviour of the sarco-
phagid fly, Wohlfahrtia vigil (Walk.). Journal of Parasi-
tology 19: 106-111.

Gassner, F. X. and M. T. James. 1948. The biology and
control of the fox maggot, Wohlfahrtia opaca (Coq.).
Journal of Parasitology 34: 44-50.

Haufe, W. O. and W. A. Nelson. 1957. Human furuncular
myiasis caused by the flesh fly Wohlfahrtia opaca (Coq.)

1981

(Sarcophagidae: Diptera). Canadian Entomologist 89:
325-327.

James, M.T. 1953. Wohlfahrtia (screwfly) parasitism of
foxes and mink and its control. National Fur News 26(4):
14, 15, 26, 27, 30.

James, M.T. and F. X. Gassner. 1947. The immature
stages of the fox maggot, Wohlfahrtia opaca (Coq.). Jour-
nal of Parasitology 33: 241-244.

James, M. T. and G. F. Kraft. 1964. The identity of the
American producers of Wohlfahrtia myiasis. In Proceed-
ings of the First International congress of parasitology.

NOTES

473

Edited by A. Corradetti. Pergamon Press, Oxford. pp.
949-951.

Yuill, T. M.and J. L. Eschle. 1963. Myiasis of penned nest-
ling cottontails. Journal of Wildlife Management. 27:
477-480.

Zumpt, F. 1965. Myiasis in man and animals in the Old
World. Butterworth, London. 267 pp.

Received 23 February 1981
Accepted 15 April 1981

Attempted Predation of Juvenile Starlings by

Northwestern Crows
PAUL C. JAMES

Biological Sciences Department, Simon Fraser University, Burnaby, B.C. V5A 1S6
Present address: Edward Grey Institute for Field Ornithology, Zoology Department, Oxford, England OX1 3PS

James, Paul C. 1981. Attempted predation of juvenile Starlings by Northwestern Crows. Canadian Field-Naturalist 95(4):
473-474.

Northwestern Crows (Corvus caurinus) on Mitlenatch Island, British Columbia, were seen stooping on juvenile Starlings
(Sturnus vulgaris) eight times. On two occasions, a Starling was captured but escaped. The crows have also been seen chasing

juvenile Song Sparrows (Melospiza melodia).

Key words: Northwestern Crow Corvus caurinus, Starling Sturnus vulgaris, British Columbia, predation.

The Northwestern Crow (Corvus caurinus) is a
well-known coastal scavenger and predator on bird
eggs and nestlings (Butler 1974, 1979; Drent and Gui-
guet 1961; Vermeer 1963). However, its possible role
as an active predator on full-fledged juveniles has not
been recorded. This note reports attempted predation
on juvenile Starlings (Sturnus vulgaris) by North-
western Crows on Mitlenatch Island, British Colum-
bia (49° 57’N, 125° 00’W).

A large flock of recently fledged Starlings usually
feed and roost on Mitlenatch during the summer
months. On 4 June 1979, I saw an adult Northwestern
Crow attack a Starling in flight. The Starling had been
feeding with about 20 others, which suddenly took
flight. The crow flew down from above, where it had
been sitting on the hillside. It grasped a Starling’s back
with both feet about 2 m above the ground, and held it
for a second or two, while the Starling screamed
loudly. The crow released it, and it dropped into some
tall grass. The crow then turned sharply, and landed
onarock some 3 m away. It was soon joined by 10-12
other crows, which stood around calling. The original
attacker and another bird walked about, apparently
searching for the Starling, but gradually the crows
dispersed. When all had left, H. Richardson and I
searched and found the bird. It was not visibly dam-
aged, but screamed loudly when picked up. When

released, the bird flew strongly to a nearby tree, and
then left the area.

The following day, I saw five stoops by crows on
juvenile Starling flocks in flight in the same area. One
Starling was caught as before, but escaped and flew to
some nearby trees with the crow in pursuit. Six other
crows landed at the top of the trees calling. A few
minutes later, I heard loud screaming from a Starling,
but could not determine what happened in the thick
foliage. Further stoops on juvenile Starlings in flight
were seen on 17 June and 8 July. On two occasions, I
found the remains of juvenile Starlings, which may
have been predated, on the island. N. Verbeek and
J.N.M. Smith (pers. comm.) have also found
remains of juvenile Starlings and Song Sparrows
(Melospiza melodia) on Mandarte Island, B.C.
(48°37’N, 123°18’W); although both suspect North-
western Crow predation, they have never witnessed it.
On 19 June and 25 July, I observed a colour-banded
two-year-old male Northwestern Crow in pursuit of a
juvenile Song Sparrow. On both occasions, the spar-
row escaped into piles of rocks or logs on the beach.

Aerial predation or attempted predation has been
reported for several species of corvids. The Common
Crow (Corvus brachyrhynchos) has been seen pursu-
ing smaller birds, including a Starling (Erskine 1980).
The European Carrion Crow (Corvus corone) has

474

aerially attacked a number of species, again notably
the Starling (Burgess and Smout 1976; Warren 1969),
as has the Common Raven (Corvus corax) (Elkins
1964). Other corvid species recorded attacking birds
in the air are the Magpie (Pica pica) (Pulman 1978),
and Steller’s Jay (Cyanocitta stelleri) (Carothers et al.
1972).

Literature Cited

Burgess, J. and T. C. Smout. 1976. Crows killing Starlings.
Bird Study 23: 146.

Butler, R. W. 1974. The feeding ecology of the Northwestern
Crow on Mitlenatch Island, British Columbia. Canadian
Field-Naturalist 88: 313-316.

Butler, R. W. 1979. The breeding ecology and social organi-
zation of the Northwestern Crow (Corvus caurinus) on
Mitlenatch Island, British Columbia. Unpublished M.Sc.
thesis. Simon Fraser University, Burnaby, B.C. 100 pp.

Carothers, S. W., N. J. Sharber, and R. P. Balda. 1972.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Steller’s Jays prey on Gray-headed Juncos and a Pygmy
Nuthatch during periods of heavy snow. Wilson Bulletin
84: 204-205.

Drent, R. H. and C. J. Guiguet. 1961. A catalogue of British
Columbia sea-bird colonies. British Columbia Provincial
Museum Occasional Paper No. 12, Victoria. 173 pp.

Elkins, N. 1964. Raven catching Rock Dove in the air. Brit-
ish Birds 57: 302.

Erskine, A. J. 1980. A House Sparrow die-off. Nova Scotia
Bird Society Newsletter 22.

Pulman, C. B. 1978. Magpie killing Swift. British Birds 71:
363.

Vermeer, K. 1963. The breeding ecology of the Glaucous-
winged Gull (Larus glaucescens) on Mandarte Island, B.C.
British Columbia Provincial Museum Occasional Paper
No. 13, Victoria. 104 pp.

Warren, R. B. 1969. Carrion Crow taking Starling in the air.
British Birds 62: 237-238.

Received 12 February 1981
Accepted 22 April 1981

A Probable Franklin’s X Ring-billed Gull Pair

Nesting in Alberta
D. V. WESELOH

Canadian Wildlife Service, Canada Centre for Inland Waters, P.O. Box 5050, Burlington, Ontario L7R 4A6

Weseloh, D. V. 1982. A probable Franklin’s X Ring-billed Gull pair nesting in Alberta. Canadian Field-Naturalist 95(4):

474-476.

A Franklin’s and a Ring-billed Gull were observed defending a common, active nest in a California/ Ring-billed Gull colony
near Edmonton, Alberta. The Franklin’s Gull did all the observed incubation and was in attendance, witha nest, at the colony
in two successive years. This is presumably the first known case of a Franklin’s X Ring-billed Gull pair.

Key Words: Ring-billed Gull Larus delawarensis, Franklin’s gull Larus pipixcan, nesting, Alberta.

On 1 June 1976 and 18 May 1977, I observed a
single Franklin’s Gull (Larus pipixcan) nesting on the
ground in the midst of a large mixed California/ Ring-
billed Gull (L. californicus/L. delawarensis) colony
(Figure 1) at Miquelon Lake, SE of Edmonton,
Alberta. In both years the Franklin’s Gull vigorously
defended the nest against all adjacently nesting Ring-
billed Gulls (there were no California Gulls nesting
nearby).

On my visit in 1976 the nest contained two freshly
hatched young and a pipping egg. In 1977 the nest
contained three eggs. One of these was collected and is
now preserved in the Provincial Museum of Alberta.
The 1976 egg measured 60.2 mm X 45.0 mm, and the
1977 eggs averaged 60.8 mm X 45.2 mm. This is larger
than the maximum measurements for eggs of Frank-
lin’s Gulls (56.5 mm X 38.5 mm, N = 48) but is very
close to mean measurements for eggs of Ring-billed

Gulls (5.9.3 mm X 42.3 mm, N = 40) (Bent 1921). In
both years the eggs were similar in size and colour to
eggs in adjacent nests of homogeneously paired Ring-
billed Gulls. The two young, observed in 1976, also
appeared identical to young of similar size from adja-
cent Ring-billed Gull nests (though possibly lighter in
down coloration and with smaller bills).

The 1977 nest was located in the same area of the
colony as was the 1976 nest. In 1977, an adult Ring-
billed Gull was also attending the nest. This was pre-
sumably the Franklin’s Gull’s mate as it was the only
gull tolerated and not attacked by the former near the
nest. The Franklin’s Gull did all the incubation during
my observation (15-20 minutes) although both it and
the Ring-bill frequently defended the nest and
attacked other nearby gulls. In 1976, I was unable to
locate a second gull at the nest, but the 1977 sighting
and the large dimensions of the eggs in both years

Ficure |. A Franklin’s Gull at its nest in a colony of California and Ring-billed gulls, Miquelon Lake, Edmonton, Alberta.

strongly suggest that both nests involved both Ring-
billed and Franklin’s Gulls.
_ The origin of the nest contents and of the interspe-
cific pair is open to speculation. The eggs were 17%
wider (only 7% longer) than the largest Franklin’s
Gull egg and, therefore, probably laid by a Ring-billed
Gull. This could have been the result of a male Frank-
lin’s Gull paired and mated to a female Ring-billed
Gull forming a true hybrid pair. Alternately, it may
have resulted from the loss of one member of a mated
Ring-billed Gull pair and the survivor’s subsequent
behavioral pairing with the Franklin’s Gull. The orig-
inal Ring-bill pair may have been either heterosexual
or homosexual as Ryder and Somppi (1979) have
shown that the latter are able to produce fertile eggs,
presumably as a result of prior insemination.
Depending on how the interspecific pair arose, the
Ring-billed Gull may or may not have been the same
individual in each year. The Franklin’s Gull, however,
almost certainly was the same individual in each year.
The near identical location of its nest in both years
suggests a strongly developed site tenacity in the
Franklin’s Gull. I would more readily expect such site

tenacity from a true interspecific pair rather than from
birds only behaviorally paired.

The occurrence of a Franklin’s X Ring-billed Gull
pair at the colony of either species is an unusual occur-
rence; I am aware of no previous records, and Gray
(1958) records none. However, if this was a true
hybrid pair, there is a relatively obvious explanation.
Although the nesting habitat of the two species is quite
different, it is fairly easy for these two species to nest in
close proximity to one another. Ring-billed Gulls
usually nest on islands in lakes, whereas Franklin’s
Gulls nest in emergent vegetation in lakes. In Alberta
there are or have been colonies of both species, simul-
taneously, on at least Beaverhill Lake (colonies
6.5 km apart) and Frank Lake (0.6 km apart) (per-
sonal observation). To the best of my knowledge,
Franklin’s Gulls do not and have not nested on
Miquelon Lake. On lakes where both species nest, it is
possible that a young Franklin’s Gull could wander or
be washed onto a colony of Ring-billed Gulls and
become imprinted to them. This could eventually lead
to a Franklin’s X Ring-billed Gull pair as witnessed
here.

476

I wish to thank Terry Thormin, Al Wisely and Ludo
Bogaert who accompanied me to the island in one or
other of the years. The observations were made while I
was employed at the Provincial Museum of Alberta. I
also thank Pierre Mineau and two anonymous
reviewers who commented on an earlier version of the
manuscript.

Literature Cited

Bent, A. C. 1921. Life histories of North American Gulls and

THE CANADIAN FIELD-NATURALIST

Vol. 95

Terns. United States National Museum Bulletin No. 113.
1963 Dover Reprint edition.

Gray, A. P. 1958. Bird hydrids: a checklist with biblio-
graphy. Technical Communication No. 13, Common-
wealth Agricultural Bureaux, Farnham Royal, Buckshire,
England.

Ryders, J. P. and P. L. Somppi. 1979. Female-female pair-
ing in Ring-billed Gulls. Auk 96:1-5S.

Received 2 February 1981
Accepted 25 March 1981

Nesting of Brewer’s Blackbirds on Man-Made Structures and

Natural Sites in British Columbia

ROBERT W. BUTLER

Canadian Wildlife Service, Box 340, Delta, British Columbia V4K 3Y3

Butler, Robert W. 1981 Nesting of Brewer’s Blackbirds on man-made structures and natural sites in British Columbia.

Canadian Field-Naturalist 95(4): 476-477.

In British Columbia, Brewer’s Blackbirds (Euphagus cyanocephalus) that nested on man-made structures began egg laying at
approximately the same date and had similar clutch sizes to nests in natural sites. Nests in natural sites were built closer to the
ground, had a longer egg laying season, and fledged more young per egg laid, on average, than nests on man-made structures.
Man-made sites were apparently used chiefly early in the season when some natural sites lacked concealment.

Key Words: Brewer’s Blackbird, Euphagus cyanocephalus, British Columbia, breeding, man-made sites.

The Brewer’s Blackbird (Euphagus cyanocephalus)
commonly nests on the ground, in shrubs, and in trees
(eg. Bent 1958, Stepney 1975, Walkinshaw and Zim-
merman 1961). I can find only one reference to this
species nesting on man-made structures. Edwards &
Ritcey (1967) mentioned that Brewer’s Blackbirds
nested on buildings in Wells Gray Park, British
Columbia, but they did not document the nesting
success. The purpose of this paper is to document and
compare nesting data of the Brewer’s Blackbird in
nests built on man-made structures and in natural
situations in British Columbia.

Data were collected in the field and supplemented
from the British Columbia Nest Record Scheme with
a sample of natural site nests with completed clutches
and all man-made site nesting attempts. Nests built on
man-made structures in British Columbia were found
on support beams (11), wooden ledges (7), fence posts
(5), telephone poles (3), haystacks (2), and a road-
grader (1). Nests built on man-made structures
(N = 25) were significantly higher off the ground
(x = 3.3m) than nests built in natural situations,
including nests on the ground (N = 54, x = 1.1m;
p < 0.01).

The earliest records for first eggs were 17 April and
23 April, in nests in natural and man-made sites,
respectively. However, egg laying ceased by 28 May in
nests found on man-made structures but continued
until 28 June in nests in natural sites. Most late
clutches in natural sites were presumably second
attempts. There was no significant difference between
egg initiation dates in nests in natural and man-made
sites within the province. However, when the data
were pooled without regard for nest site, those nests
(x = 3 May) west of the Coast Range held first eggs
significantly earlier than nests (x = 16 May; p < 0.01)
east of the Coast Range. Mean clutch sizes were sim-
ilar but the number of young that fledged per egg laid
was significantly different between nest sites (Table 1).

Howard (1974) suggested that American Robins
(Turdus migratorius) nested in coniferous vegetation
early in the season because of a shortage of concealed
sites in deciduous vegetation. Horvath (1964) also
suggested that seasonal changes in micro-climate
around the nest caused the Rufous Hummingbird
(Selasphorus rufus) to shift its nest site. In this study,
some Brewer’s Blackbirds nested on man-made struc-
tures in the early part of the nesting season whereas

1981

TABLE 1. Reproductive data of Brewer’s Blackbirds nesting
on man-made structures and natural sites in British Colum-
bia (sample sizes).

Natural site Man-made site

Clutch initiation dates,

range 17 April-28 June 23 April-28 May
median 13 May (41) 18 May (22)
Clutch size,

mean = s.d. 5.0 + 0.8 (40) 5.4+ 0.9 (21)
Number of young

fledged per egg

laid.* 0.48 (144) 0.36 (72)

*significant difference exists, p< 0.01.

natural sites were used throughout the season. This
suggests that nests on man-made sites offered better
concealment than many natural sites early in the sea-
son. The data from the Nest Record Scheme seldom
contained information on whether the same individu-
als shifted their nest site. A population could consist
of early and late nesters that preferred different sites.
However, I am reasonably certain from my observa-
tions of nests at the Creston Valley Wildlife Interpre-
tation Centre in 1980 that the same Brewer’s Black-
birds shifted their nest sites as the season progressed
because all five second nests (in natural sites) were
within a few meters of first nests (mostly on struc-
tures). In addition, one female, recognizable by her
amber-coloured irises, shifted her nest site from under

NOTES

477

a walkway to the ground a few meters below.

J.P. Goossen assisted in the field and D. R. Flook
and J. Grant made useful comments on the manus-
cript. G. E. J. Smith provided statistical advice and
R. W. Campbell allowed access to the British Colum-
bia Nest Record Scheme. I thank all of you.

Literature Cited

Bent, A. C. 1958. Life histories of North American Black-
birds, Orioles, Tanagers, and Allies. United States
National Museum Bulletin 211. (Dover Reprint 1965).

Edwards, R. Y.and R. W. Ritcey. 1967. The birds of Wells
Gray Park. Parks Branch, Department of Recreation and
Conservation, Victoria, B.C.

Horvath, O. 1964. Seasonal differences in Rufous Hum-
mingbird nest height and their relation to climate. Ecology
45: 235-241.

Howard D. V. 1974. Urban robins: a population study. In
J. H. Noyes and D. R. Progulske (eds.). Wildlife in an
urbanizing environment. Proceedings of symposium held
27-29 November 1973, Springfield, Mass. Cooperative
Extension Service, University of Massachusetts, Amherst.

Stepney, P. H. R. 1975. Tree nesting sites and a breeding
range extension of Brewer’s Blackbird in the Great Lakes
region. Canadian Field-Naturalist 89: 76-77.

Walkinshaw, L. H. and D. A. Zimmerman. 1961. Range
extension of the Brewer’s Blackbird in eastern North
America. Condor 63: 162-177.

Received 22 September 1980
Accepted 14 October 1981

News and Comment

Amendment to By-laws of The Ottawa Field-Naturalists’ Club

1. By-law 3. Strike out the sentence “It shall recom-
mend to the Council candidates for Honorary
Membership”.

2. Addition of two new by-laws:

(19) Duties of the Awards Committee:
The Awards Committee shall be responsible
for nominating deserving members of the
Club for Honorary Memberships, Special
Club Awards and for awards offered by other
clubs and organizations.

(20) Duties of the Executive Committee:
The Executive Committee shall propose pol-
icy direction, deal with areas which are sensi-
tive or difficult, act as a continuum of
thought, deal with emergency matters and call
upon other persons for advice when
necessary.

E F-Pope
Recording Secretary

Changes in By-laws, Ottawa Field-Naturalists’ Club

A motion to change By-law 15 of The Ottawa Field-
Naturalists’ Club was passed by 13 votes to 2 by the
Council at the meeting of 23 November 1981. This
by-law now reads as follows:

15. Membership Dues
The schedule of dues shall be as follows:

Individual $13.00
Family 15.00
Sustaining 30.00

Life membership shall be granted upon

payment of a single sum of $300.00
A motion to change By-law 16 of The Ottawa Field-
Naturalists’ Club was passed unanimously by the

Council at the meeting of 14 December 1981. This
by-law now reads as follows:
16. Subscription Fees
The schedule of subscription fees shall be as

follows:

The Canadian Field-Naturalist
Individual $13.00
Libraries and Institutions 25.00

Trail & Landscape
Libraries and Institutions 13.00

E.F. Pope
Recording Secretary

New Honorary Members of the Ottawa Field-Naturalists’ Club

In 1980 four outstanding members of the OFNC
were awarded with Honorary Memberships in recog-
nition of their immense contributions to the Club
and/or to Canadian natural sciences. They join the
select group of important Canadian naturalists who
have previously been so honoured.

Dr. Clarence (Clarie) Frankton _

Clarie served on the OFNC Council for 15 years
between 1947 and 1961 and was instrumental in reor-
ganizing our Reserve Funds to provide a greatly
increased rate of return. He has also served in a wide
range of projects including the conservation areas
program, Christmas Bird Count and Club excursions.

478

He served as OFNC Auditor for 10 years and was
Treasurer of the Club between 1947 and 1949. His
detailed knowledge of the natural values in the west
end of the Ottawa area was particularly vital in the
efforts to establish Conservation Areas (such as the
Carp Hills, South March Highlands and Bridlewood)
in the Official Regional Plan of the Regional Munici-
pality of Ottawa-Carleton.

Perhaps his greatest contribution to the OFNC has
been the easy grace with which he shared his expert
botanical knowledge and has encouraged the devel-
opment of amateur naturalists within the Club. A
number of OFNC members who are now working
professionally in the natural sciences owe a great deal
of credit for their development to the inspiration and

1981

guidance of Clarie Frankton. His enthusiasm for and
love of the natural world transcends his considerable
scientific knowledge and is truly an inspiration for
those fortunate enough to share a day in the field with
him.

Dr. Frankton retired in 1970 as Head of Taxonomy
for the Plant Research Institute (Agriculture
Canada). His important studies of weedy plants have
had considerable economic and scientific significance.
Although he has published many scientific papers in
those areas for which he established an international
reputation, he is perhaps best known to Canadian
naturalists for his book The Weeds of Canada.
Although first published in 1955, it is still much in
demand, particularly after it’s revision in 1970 (with
G.A. Mulligan).

Dr. Frankton was presented the Lawson Award by
the Canadian Botanical Association shortly after his
retirement. This award is presented only when the
CBA deems that a particularly significant contribu-
tion merits such high honour.

Clarie Frankton exhibits a rare combination of
long and outstanding service to the OFNC and to
natural science. It is only fitting that the Club
acknowledge this extraordinary achievement by elect-
ing him an Honorary Member.

Mary E. Stuart

Since becoming a member of the OFNC in 1945
Mary Stuart has been a dedicated and valuable Club
enthusiast who has contributed to a great many ser-
vices and programs. She served on Council from 1948
until 1972 and on a number of committees, including
Membership, Special Lectures, Publicity, Christmas
Census and Excursions & Lectures. She was particu-
larly active in the organization, meetings and outings
of the very successful Bird, Fern, Orchid and Trail
study groups in the late 1950’s and early 1960’s. She
also was highly involved in the early days of the News-
letter. To fill in her idle (!) hours she organized field
trips, Annual Dinners, evening nature walks, and so
on.

If the Club ever had a social hostess, Mary was it.
Friendly and gracious, she welcomed new members at
Annual Meetings, club outings, picnics at the Beatty
Point Lodge, and wherever the OFNC gathered. She
made people feel right at home and always had a
supply of Club literature to encourage new members.
In later years she has been very generous in offering
her farm near Packenham for OFNC excursions. The
Macoun Club, in which she maintains an interest, has
made a tradition of enjoying regular camping trips to
her property to study natural history.

For her unassuming manner, enthusiastic support,

NEWS AND COMMENT

479

and dedicated service to the OFNC, we are delighted
to number Mary Stuart amongst our distinguished
Honorary Members.

D. B. O. Savile

By profession a mycologist with more than one
hundred research publications to his credit, Dr. Savile
has received world-wide recognition and acclaim for
his work on the plant rusts (Uredinales). His many
honours include membership in the Royal Society and
a recently confered honorary doctorate from McGill
University.

Doug has been an active member of the OFNC
since 1944. He served on Council for many years and
was particularly involved in financial affairs. He has
produced some of the most original and significant
observations on natural history of any Club member.
He has long advocated the necessity for a broad
knowledge in many fields of natural history in order to
fully understand and interpret the natural world. His
own talent for astute observation and his genius for
grasping the significance of what he sees can be seen
by mentioning only a few of the score of topics on
which he has published : bird navigation and flight,
evolution of the avian wing, gliding and flight
mechanics in vertebrates, splashcup and springboard
methods. of seed dispersal, arctic adaptations in
plants, meteorology, plant geography — and so on.
Doug Savile’s numerous and scholarly contributions
to our knowledge of natural history rank him with the
top naturalists of the century. The OFNC is proud to
consider him one of our Honorary Members.

R. Yorke Edwards

Yorke has been a member of the OFNC since 1942
although he was active in Toronto naturalists’ circles
somewhat before that. In this time he has become one
of Canada’s best known naturalists. He has written
over 200 articles on various natural history subjects
(although birds have always been a particular favorite
of his), as well as two books on natural history and
interpretation.

He is a pioneer of Canadian interpretation. His
persistence and persuasive arguments saw the first
British Columbia park interpretation program
initiated in 1949. As interpretation specialist for the
Canadian Wildlife Service, he developed the Wildlife
Service’s system of interpretive centers. Untold
numbers of Canadians have benefited either directly
or indirectly from Yorke’s interpretive efforts, and
have developed a new awareness of some of Canada’s
important natural values.

Yorke is presently Director of the British Columbia
Provincial Museum in Victoria. He continues to serve

480

various conservation organizations and to write
thought-provoking articles on interpretation and con-
servation. For over 30 years he has made significant
personal contributions in each of the areas of concern
outlined in the objectives of the OFNC. It is indeed

Support for Field Research

The Center for Field Research offers funding and
volunteer assistance to postdoctoral scholars needing
support for field research. Working cooperatively
with EARTHWATCH, a national volunteer organiza-
tion, The Center sponsors seventy field research pro-
jects each year in the sciences and humanities, and in
1981 will send $700,000 and 1200 volunteers into the
field. Proposals for 1982-3 research are now being
reviewed.

Eligibility: Qualified projects are reviewed for scho-
larly merit and their ability to constructively utilize
teams of volunteers in the field. EARTHWATCH volun-
teers, a work force and funding source in one, pay a
share of project costs and assist in the field work. For
example, a coastal geologist studying barrier island
migration needs three two-week teams, of minimum 4
— maximum 10 volunteers each, to assist in sampling
and beach profile surveys. If each volunteer contrib-
utes $500 toward project costs, a total grant of $6,000
— $15,000 is provided.

The Center invites proposals from postdoctoral

Grants Available for Bird Projects

The James L. Baillie Memorial Fund for Bird
Research and Preservation invites applications for
grants to support projects on Ontario birds in 1982.

The fund’s aim is to encourage field studies by
amateur naturalists and to support projects which
increase or disseminate knowledge of birds in their
natural environments or contribute to their preserva-
tion. Priority will be given to projects which draw on
the resources of volunteer naturalists in conducting
field research of fieldwork and to applicants who do
not have access to other sources of support.

Examples of eligible projects might include (but are
not limited to) individual or group research by ama-
teur naturalists, distributional or other surveys using
information contributed by volunteers (e.g. Christmas

THE CANADIAN FIELD-NATURALIST

Vol. 95

appropriate that we award Yorke Edwards with an
Honorary Membership.

D. F. BRUNTON
The assistance of notes from S. Thomson, D. Gray and
V. Humphreys is gratefully acknowledged.

scholars of all nationalities and is committed to spon-
soring the research efforts of qualified women and
minority investigators.

Areas of Interest: Projects in any recognized
academic discipline are welcome, 1.e., anthropology,
archaeology, art history, biology, folklore, geology,
marine science, musicology, zoology. Interdiscipli-
nary proposals are also invited.

How to apply: Submit a two-page Preliminary
Proposal outlining your research objectives, project
dates, planned use of volunteers, and need for funds.
Following favorable review, a Formal Proposal will
be invited, which must precede field work by nine
months.

For application guidelines, and a listing of projects
currently receiving support, write:

Nancy Bell Scott
Center for Field Research
Box 127-U, 10 Juniper Road
Belmont, MA 02178

Bird Counts) and publication of local guides or
checklists.

Grants will not normally exceed $750. Applications
are due by December 31, 1981, and should be submit-
ted on forms obtainable from the Secretary, James L.
Baillie Memorial Fund, c/o Long Point Bird Obser-
vatory, P.O. Box 160, Port Rowan, Ontario
NOE 1 MO.

The James L. Baillie Memorial Fund approved 10
applications totalling about $4,500 in 1981. The Fund
is financed in part from the proceeds of the Jim Baillie
Memorial Bird Count (Baillie Birdathon). Donations
to the fund are tax deductible and may be sent to the
address given above.

1981

NEWS AND COMMENT

481

Common Tern Colour-Marking by the Canadian Wildlife Service: Request for Information

During 1981, Dr. Hans Blokpoel of the Canadian
Wildlife Service, colour-marked Common Terns at
two large colonies in the Great Lakes area, with the
objective of determining the year-round distribution
of the birds, especially their migration routes and
wintering areas in Latin America. Adult Common
Terns were trapped on their nests at the Eastern Head-
land of the Toronto Outer Harbour (Lake Ontario)
and at Tower Island (Niagara River). Orange plastic
tags were attached to both wings of the trapped adult
birds. In addition, young Common Terns were
marked with pink plastic wing tags at those colonies.
One standard metal leg band and one coloured plastic
leg band (yellow with a black horizontal stripe) were
put on each of the tagged birds.

If you see a Common Tern with a pink or orange
wing tag please record the following details: place,
date and colour of the tag. If possible, also record the
combination of numbers and/or letters on the tag (the
two tags on any bird have the same color and the same
combination of letters and numbers) and note which
legs the plastic and metal leg bands are on. Thank you
very much for your assistance. All reports will be
acknowledged and should be sent to:

Bird Banding Office
Canadian Wildlife Service
Ottawa, Ontario, Canada
KIA 0E7

Wood Buffalo National Park Management Planning Program

Parks Canada is presently engaged in a Manage-
ment Planning Program, which, when completed, will
determine the long range direction for the future use
and management of Wood Buffalo National Park.

The park, which straddles the Alberta/N. W. T.
border, is 44,807 sq. km in size and protects a natural
area of Canadian significance representative of the
Northern and Southern Boreal Plains of Canada.

Significant resource features within the park
include portions of the Birch and Caribou Mountains,
one of the largest fresh water deltas in the world,
extensive karst fields, vast salt plains, the largest free
roaming herd of bison in the world, important archeo-
logical sites and nesting sites of the endangered
Whooping Crane and Peregrine Falcon.

The park is largely undeveloped but does contain a
limited number of facilities including a 36 site camp-
ground anda group camp located at Pine Lake, 66 km
S. W. of Fort Smith, N.W.T. Associated with this
development is a day use area, an interpretive theatre
and display building, a boat launch and seventeen
privately owned cottages situated on lots leased from
Parks Canada.

Access to visitor facilities in the park is provided
primarily by a gravel loop road that extends S. W.
from Fort Smith, N. W. T.

Management Planning issues centre largely around
resource management concerns such as fire manage-
ment policy, management of the Bison herds, hunting,
trapping and subsistance fishing privileges, commer-
cial logging practises, possible changes to the park
boundaries, preservation of special areas and features

in the park. At question also is the future of native
settlements within the park. What level of recreational
use and the location of any additional recreational
facilities must also be determined. At issue also is the
appropriate level of access into the park via road, air,
and water.

Several regional issues such as the Slave River
hydroelectric proposal and its impact on the park,
native land claims, regional development and tourism
must also be addressed in the planning process.

Public meetings are now underway to permit the
public to exchange information with the planning
team and identify and discuss planning issues. In the
fall of 1981, the planning team will meet with the
public again to present a number of alternate plans
and receive comments.

Following this alternative plan stage a draft man-
agement plan will be prepared and will be made avail-
able for public review and comment. After this public
review, the final plan will then be prepared and pres-
ented to the Minister responsible for Parks Canada
for final approval.

The public is invited to participate in all stages of
the planning process and the planning team, based in
Fort Smith, welcomes input. If you would like to
receive copies of newsletters dealing with the planning
program, make your views known, require informa-
tion, or wish to have your name placed on the mailing
list, please contact the Public Participation Co-
ordinator, Wood Buffalo National Park Management
Planning Program, P.O. Box 750, Fort Smith,
N. W. T. XOE OPO, phone: (403) 872-2649.

482

Environmental Monitoring: a Symposium

The Aiberta Society of Professional Biologists is
pleased to announce a symposium entitled “Environ-
mental Monitoring”, to be held on 20 and 21 April
1982, at the Westin Hotel (formerly Edmonton Plaza)
in Edmonton, Alberta.

The symposium will examine the question of why
environmental monitoring is required, examine the
design basis of environmental monitoring, the practi-
cal aspects of conducting environmental monitoring,
the results of existing environmental monitoring pro-
grams, and the value of biomonitoring programs.
Eminent speakers from universities, government and
private industry will be featured.

XVth Pacific Science Congress, 1983

The XVth Pacific Science Congress will be held in
Dunedin, New Zealand, 1-11 February 1983. Its
theme is to be “Conservation, development and utili-
zation of the resources of the Pacific”.

A session is planned on the diversity, distribution,
abundance and management of vertebrate popula-
tions in the Pacific region. Joint sessions will be
arranged with related disciplines. Speakers are now
invited to offer papers (with title and short summary)
on such topics as: —

Biogeography
Species diversity

Errata

Please note the following corrections to The Canadian
Field-Naturalist 95 (3) :
p. 241 KEDDY: Relict Lakeshore Vegetation : Table 1:
the annotations * and { are reversed at the bottom of
the table; correct to

* Species with costal plain affinities

+ Rare in Ontario (Argus and White 1979)
p. 356 SOUTHIERE and STEVENTON: Seasonal Pelege
Change of Martin: line 2 of the right hand column of
text should preceed line | in this column so that the
paragraph beginning at the bottom of the left hand
column reads:

THE CANADIAN FIELD-NATURALIST

Vol. 95

© Date: 20 and 21 April 1982
© Location: Westin Hotel (formerly Edmonton
Plaza)
© Registration and Further Information:
Don Thompson
Secretary
Alberta Society of Professional Biologists
P.O. Box 566
Edmonton, Alberta
T5J 2K8

Telephone: (403) 429-9110

Habitat requirements

Migration and movements

Population ecology

Ecosystem studies

Man-induced changes

Endangered species

Conservation and management

For further information, please write to Dr C W

Burns, Section Convener, (Ecology and Environmen-
tal Protection), c/o Department of Zoology, Univer-
sity of Otago, P O Box 56, Dunedin, New Zealand.

“Two specimens exemplifying the extremes of winter

color are filed in the Wildlife Resources Museum
University of Maine ...” etc.

p. 363 PRUITT: Application of Varrio Snow Index :
Figure |: substitute in the legend “n, number of api
stations” for “n, number of Caribou”.

p. 376 LOVE: [Review of] North American Birds of
Prey: “By William Marshall” should be changed to
read “By William Mansell”

The editor is grateful to Paul A. Keddy, J. Douglas
Steventon, W. O. Pruitt, Jr.,and William Mansell for
bringing these to his attention.

1981

NEWS AND COMMENT

The Ottawa Field-Naturalists’ Club

Special Publications

1. Autobiography of John Macoun
A reprint of the 1922 edition of the fascinating life story of one of Canada’s outstanding early
naturalists, with a new introduction by Richard Glover and bibliographical essay, footnotes,
and index by William A. Waiser, plus three maps of John Macoun’s western travels.
Individuals $12.50 plus $2 postage and handling
Libraries $15.00 plus $2 postage and handling

2. Transactions of The Ottawa Field-Naturalists’ Club and The Ottawa Naturalist — Index.
Compiled by John M. Gillett
A complete author, title, and subject index to the predecessors of The Canadian Fiela-
Naturalist, the first thirty-nine volumes of the publications of The Ottawa Field-Naturalists’
Club.
$25 plus $2 postage and handling

Centennial Bird Record

Songs of the Seasons
More than fifty eastern North American birds and amphibians are presented in full stereo-
phonic sound as recorded in the wild by wildlife recording expert F. Montgomery Brigham.
$9.11 (postage and handling included but
Ontario residents must add 7% sales tax

Please send orders to:

The Ottawa Field-Naturalists’ Club
Box 3264 Postal Station C
Ottawa, Ontario, Canada

K1Y 4J5

483

Book Reviews

ZOOLOGY

A Manual of Mammalogy; with keys to families of the world

By Anthony F. DeBlase and Robert E. Martin. 1981. 2nd
edition. Wm. C. Brown, Dubuque, Iowa. xii + 436 pp.,
illus. U S $9.95.

This is the most complete laboratory manual avail-
able in mammalogy. This edition is much improved
over the first. Six new chapters appear and others are
entirely rewritten. Illustrations are now abundant.
Discussions on mammalian abundance, appendicular
skeletons, antlers, claws, extoparasites, hoofs, horns,
locomotion, nails, population dynamics, reproduc-
tion, sign, skin, skulls, systematics and teeth are dealt
with in their own chapters. About 100 pages are
devoted to explaining methodologies concerning age
determination, data recording, diet analysis, identifi-
cation, literature searches, marking, specimen prepa-
ration and preservation, statistical analyses, and trap-
ping. Each mammalian order is described in detail.
Distinguishing characteristics, distribution, fossil his-

tory, identification keys, line drawings of representa-
tive species, skull diagrams, supplementary reading
lists and taxonomic discussions are given for most all
mammalian families of the world. There now is espe-
cially good treatment of bats and rodents. A good
glossary and index make the text a useful quick
reference.

This manual would work well as a guide in a mam-
malogy course, but it could easily be used for wildlife
management, vertebrate zoology, applied ecology, or
mammalian taxonomy courses. Serious naturalists
interested in the scientific study of mammals would
find a copy useful.

RICHARD M. ZAMMUTO

Department of Zoology, University of Western Ontario,
London, Ontario N6A 5B7

The Great Gray Owl: phantom of the northern forest

By Robert W. Nero. 1980. Smithsonian Institution Press,
Washington, 167 pp., illus. $21.

At long last, the account of Bob Nero’s extensive
studies of the Great Gray Owl in and about southern
Manitoba has been published in one volume — and
it’s a gem. It is well-bound, beautifully illustrated and
nicely laid-out. The copy is printed in two well-spaced
columns in an easy-to-read typeface. Thirteen chap-
ters, each dealing with some major topic of Great
Gray Owl biology, are accompanied by a thorough
bibliography and a well-detailed index. The copy is
remarkably free of errors.

Each chapter is introduced by a vignette of the topic
at hand, written in beautiful prose. One can easily see
that Nero comes from Aldo Leopold country! The
subject comes alive — and this is a great strength of
the study throughout. Nero quotes freely and comfor-
tably from his field notes (and those of his associates)
in the present tense. The reader is so effectively carried
along by the text in this way that he can “see ” the
scenes being described. Within each chapter, the tre-
mendous personal contribution, by Nero, to our
knowledge of the Great Gray Owl is evident — as is his
love for these creatures and their environment. No
clinically sterile data base, these birds!

The careful placement of the spectacular photo-

484

graphs of Robert R. Taylor compliment the text
enormously. If there was any doubt that Taylor is a
giant in his field, this book destroys it. The reader is
treated to a full array of stunning black and white and
colour photographs of the Great Gray Owl and its
world. These pictures illustrate numerous subtle and
spectacular features alike and are artistically and
technically superb. The latter quality is waived only
once (on page 132) in order to illustrate a precise
action. A particularly innovative photograph (page
76) superimposes a skull on a feathered head to (very
effectively) illustrate the relatively small body size of
this owl. Captions are excellent, adding understand-
ing (not just description) of each photograph for the
reader. They also offer interesting tid-bits (e.g. the
identification of immature birds by tail feather char-
acteristics).

Data are presented ina narrative form; the reader is
directed to upcoming scientific papers for a detailed
rationalization and analysis of Nero’s findings. This
narrative style has made the book very readable but
does not reduce its credibility. Nero’s writing is lucid,
credible and enjoyable. The subject coverage is very
complete, dealing with the Great Gray Owl’s behav-
iour, physiology, nesting habits, population trends,
research techniques, etc. These are discussed in a lar-

1981

gely smooth and natural manner although the flow
becomes a bit ragged towards the end of the book.
Preening is covered in Chapters 6 and 9 as is interspe-
cific conflict in Chapters 7 and 8 — both of which
could have been dealt with in single chapters. Chap-
ters 3(An Owl in the City) and 10(A Nesting of Owls)
though beautifully written are really expansions on
topics discussed satisfactorily in other chapters and
are therefore redundant.

For Nero this book and the studies that went into it
were obviously labours of love, a feeling he wishes to
share. The better part of Chapter |1 is a manual for
would-be Great Gray Owl nest-finders — complete
with words of caution and warning. This is a particu-
larly delightful “welcome” for the reader from the
author.

Nero estimates that there may be 50,000 of these

- _ The Warblers of America

Edited by Ludlow Griscom and Alexander Sprunt, Jr.
(Revised and updated by Edgar M. Reilly, Jr.). 1979.
Doubleday, Garden City, New York. 302 pp. US $19.95.

When I first heard that the 1957 edition of The
Warblers of America was to be revised and updated, I
was very excited because the warbler family is my
favorite group of birds. After comparing the 1979
update with the first edition, I must say that I had
hoped for more. Although there are undoubtedly
some improvements, the greatest benefit is once again
making available to the birding public a long out-of-
print popular account of warblers at a still very rea-
sonable price.

Changes include lumping of the Myrtle and
Audubon’s, and the Parula, Olive-backed, and
Socorro; some new range maps although most remain
unchanged; new chapters for Central America and the
West Indies; and a new order of presentation of spe-
cies, which now follows that found in Peters, Birds of
the World. While there is a new species account for the
Yellow-rumped (lumping of Myrtle and Audubon’s),
Reilly has retained the “old” species accounts for the
Parula and Olive-backed.

The nice 35 full-color plates by John Henry Dick
are now grouped at nine locations rather than being
distributed singly throughout the book as found in the
1957 edition. Comparing the quality of the plates in
the two editions, I found those in the older edition to
be sharper, witha richer color. Several of the plates in
the revised edition have a greenish tint. Loss of some
sharpness may be a result of the increased page size,

BOOK REVIEWS

485

magnificent owls in North America and expresses
quiet concern that trapping, forest management and
agricultural practices may be having a detrimental
effect on that population.

This book is clearly the definitive work on the Great
Gray Owl in North America. Nero’s reasoned, poetic
and at times passionate writing does justice to the
majesty of his subject. For those of us who have
waited almost ten years for this book to be published,
it was worth the wait. I’ll summarize my opinion of the
book by using Nero’s own words . . . “gawd how I
loved it all”!

DANIEL F. BRUNTON

2683 Violet Street, Ottawa, Ontario K2B 6X1

which reduced the total number of pages from 356 in
the first edition to 302 in the 1979 edition.

The seven introductory chapters remain unchanged
with a few very minor exceptions. The 58 species
breeding in the United States, Canada, and Baja are
then presented. Each species account is accompanied
by a range map and information typically found in
species write-ups. Often included in the 114-34 pages
devoted to each species is a black-and-white line draw-
ing. Species accounts are followed by | 1 chapters on
warblers of other western hemisphere regions.

In the appendix on the classification of the warbler
family Parulidae, a subspecies regional name has been
added in the new edition. An irritating change in the
1979 edition is the deletion of species page numbers in
the Table of Contents. One can no longer check the
Contents to find the page number for a particular
species. On the other hand, an improvement in the
new edition is the expansion of the index from just a
species index to a full subject index.

While this volume still provides invaluable infor-
mation on this most interesting group of birds, I was
hoping for more incorporation of information
accumulated over the past 20 years into the revised
edition. However, a price of $20 for a 300-page bird
book with 35 color plates is a steal, and it will continue
to provide interesting reading and prove to bea useful
reference whenever it is consulted.

NOEL J. CUTRIGHT

Wisconsin Electric Power Co., 231 W. Michigan, Milwau-
kee, Wisconsin 53201

486

THE CANADIAN FIELD-NATURALIST

Vol. 95

Butterflies and Moths of Newfoundland and Labrador: the Macrolepidoptera

By Ray F. Morris. 1980. Research Branch Agriculture Can-
ada Publication 1691. Supply and Services Canada, Hull.
407 pp., illus $15.00 in Canada; $18.00 elsewhere.

The completion of this excellent book has been a
life long goal for Ray Morris. It brings together a great
deal of his original research to document the biology
of the many introduced and native macrolepidoptera
from this most easterly part of Canada. Mr. Morris
emphasizes larval feeding and points out many infor-
mation gaps in life cycles thereby accentuating future
research objectives. The text has a logical and concise

format in clear bold type. This very readable, hard-
cover book with its many photos, illustrations, distri-
bution maps and 34 colour plates of adult stages is an
outstanding bargain at $15.00. I’m sure this will be a
key text for students and researchers well into the 2 Ist
century.

ROBIN T. DAY

Biology Department, University of Winnipeg, Winnipeg,
Manitoba R3B 2E9

Les salmonidés des eaux de la Plaine de Montréal: 1. Historique, 1534-1977 et 2. Biometrie,
biogéographie, 1970-1975, et registre des péches 1941-1976.

1. Par Vianney Legendre, Jean-René Mongeau, Jean Leclerc
et Jocelyne Brisebois. 1980. Ministere de Loisir, de la
Chasse et de la Péche, Service de l'aménagement et de
lexploitation de la faune, Québec. Rapport Technique N°
06-27. 280 pp., illus. et 2. par Jean-René Mongeau, Vianney
Legendre, Jean Leclerc et Jocelyne Brisebois. 1980, 2nd
edition. Ibid. Rapport Technique N° 06-28. 139 pp., illus.

This work is a delight to read because Vianney
Legendre and his team are well-versed in fishery biol-
ogy, possess a rare degree of expertise in writing, and
display a deep knowledge of history and literature that
both document and embellish their writings.

An in depth study and interpretation of the litera-
ture combined witha knowledge of the upper thermal
and lower oxygen tolerance of the Atlantic salmon,
Salmo salar, demonstrate that historically anadrom-
ous populations did not ascend the St. Lawrence
River much further upstream than Québec City and
therefore did not reach Montréal* or the Richelieu
River. Populations in Lake Champlain and Lake
Ontario must have been land-locked. Documenta-
tion, from one of the finest natural history libraries in
Canada (built up by Vianney Legendre ) is provided
from the works of Samuel Champlain, the Jésuite
Relations, Pierre-Espirit Radisson to those of modern
studies on Quaternary geology. If we do not find
reference or a footnote to Philippe Aubert de Gaspés
chapter “Le Lac Trois-Saumons” from his 1866
Mémoires, we can be confident it was not omitted
because the authors were unaware of it, but rather
through judicious selection.

The distribution of salmonids in the St. Lawrence is
also documented from the excellent and thorough fish
collections of the Ministry. Unlike those of most pro-

vinces these collections have been made according toa
systematic plan, and the results published.

From this evidence the authors are able to point out
the ecological lacunae for salmonids in rivers of the
Plaine de Montréal, voids which introduced rainbow
and brown trout have profited by, in spite of pollution
in the region. Current ministerial reorganization for
protection of the aquatic environment will doubtless
benefit the catch of salmonids and other fishes.

The second volume deals more closely with the
management and evaluation of success of fisheries
programs. A valuable long-term evaluation indicates
that plantings of some species have shown extraordi-
nary returns to the anglers while those of other species
should be discontinued.

The third chapter of the same volume explores the
role of smelt as a sport and as a forage fish. The pygmy
smelt is not mentioned. The studies of C. E. Delisle
and D. Copeman have convinced the reviewer that
Québec has two kinds of smelt, one large and wide-
spread and one small, the pygmy smelt and known in
the world only from Heney Lake, Québec and Green
Lake, Maine. The pygmy smelt, which feeds almost
entirely on plankton, would make an ideal forage
species. It would be worthwhile in the reviewer's opin-
ion to try and introduce it in lakes which lack ciscos or
whitefishes as planktivores. The introduction should
be made using eggs, not adults which might be
infected with Glugea hertwigi. The stock of pygmy
smelt and Lake Heney should be protected from
harmful environment changes.

In the smelt chapter as elsewhere, Legendre and his
team show a mastery of language and imagery that the
man on the street, the scientist, and even a linguisti-

*May occur as a very rare phenomenon as one did see recently (pers. com. V. Legendre), not as a regular migration.

1981

cally untalented reviewer can enjoy and be enriched
by. The flowing and beautiful phrases are particularly
striking in what could have been a dry explanation of
how the smelt eats plankton a thousand times smaller
than ifself and, in turn is eaten by a trout a thousand
times larger. The smelt thus acts as a converter (trans-
formateur) of energy.

The final section of volume two is devoted to con-
clusions and carefully considered recommendations,
and appendices which document the theses contained
in the text.

To these recommendations I would personally
tender one additional: that the Ministry, which has so
sagely supported the building of the library of the
Laboratoire de Recherches and the study collection of
fishes through systematic surveys and publication of
results, not only continue but increase support for
these worthy projects. The yield of the library and
collection are numerous and significant to the public,
to science in Québec, Canada, and the world. The
recently published Atlas of North American fresh
water fishes benefited greatly from the completeness

- Birds of Prey of the World

By Friedhelm Weick. 1980. Paul Parey, Hamburg and Ber-
lin, Germany (US distributor IR Unlimited, New York).
159 pp. illus. US$48.

I would not have believed that yet another general
book on birds of prey could be of any use to raptor
enthusiasts, however Friedhelm Weick manages to
write and illustrate as stated by the late Leslie Brown
in the foreword, “a much needed guide to identifica-
tion of all the world’s diurnal birds of prey, in the
order Falconiformes”. However, because the owls are
not represented, I believe the title is misleading and
would suggest a more apt one, “Falconiformes of the
World”.

This book written in German and English consists
of two sections, the first dealing with an identification
key based primarily on size, and the second being 40
colour plates comprising more than 1400 side-view
figures painted by the author. Weick has deliberately
compressed the text, emphasizing instead his consid-
erable skill as a bird artist. The index with common
and scientific names in separate lists at the back of the
book facilitates the search for a given species or sub-
species.

In my opinion, the first section is of questionable
value. The black and white ink drawings, as well as the
short descriptions of significant identifiable character-
istics, are well-done and useful, but how can one sub-
jectively discern a “very large”, a “rather large”, or a

BOOK REVIEWS

487

of surveys in southern Québec and accurately protray-
ing the distribution of fishes. The spot distribution
maps are useful to anglers, fishery managers, ecolo-
gists, biogeographers and taxonomists. Perhaps these
priceless and irrreplacable collections, together with
existing herbaria, insect and other collections, might
be gathered and properly housed as a Québec Natural
Science Museum. Only recently has society begun to
appreciate the many values of natural history collec-
tion to education, research and development.

To the authors of Les salmonideés des eaux de la
Plaine de Montréal | offer my heartiest congratula-
tions. To my fellow anglophone biologists, and natu-
ralists may I suggest they obtain copies, and sit down
beside the fireplace, for several evenings of pleasura-
ble and rewarding reading, even if it means periodic
reference to a dictionary.

DON E. MCALLISTER

Curator of Fishes, National Museum of Natural Sciences,
National Museum of Canada, Ottawa, Ontario KIA 0M8&

“large” bird from one another without being able to
measure it in the hand? On many occasions in my
experience, enthusiastic people have mistaken the
“very small” American Kestrel (Falco sparverius) for
the “medium-sized” Peregrine Falcon (Falco peregri-
nus) just after having seen the latter on television!

The second section, though, is far more useful and
certainly unique in its approach. Some minor faults
should be mentioned. Due to the large number of
subspecies, etc., some of the plates appear crowded,
creating some difficulty in attaching labels to each
bird correctly. More specifically, in selecting a species
with which I am most familiar, the eye colour in Falco
sparverius sparverius, and perhaps in many of the
other falcons, appears somewhat lighter than it should
be.

In spite of the above faults, Weick’s book will be a
source of information on my library shelf to which I
will undoubtedly refer to often. Its size obviously
restricts its use to other than in the field. I recommend
it highly to serious raptor enthusiasts.

DAVID M. BIRD

Macdonald Raptor Research Centre, Macdonald Campus
of McGill University, 21,111 Lakeshore Rd., Ste. Anne de
Bellevue, Québec H9X ICO

488

Behavioral Mechanisms in Ecology

By Douglass H. Morse. 1980. Harvard University Press,
Cambridge, Massachusetts. 371 pp., illus. US $25.

It is a common complaint of would-be reformers
that both ecology and ethology grew up in the anti-
Darwinian atmosphere of the early 20th century, and
that, despite the fact that an animal’s ecology may be
very largely considered asummation of its behaviour,
each discipline has tended to ignore both the processes
studied by the other and the consequences of its own
theories for individual fitness.

The reaction against this has been ‘modern natural
history’ or ‘sociobiology,’ here ‘behavioral ecology’,
and the hypothesis it has broadly corroborated is that
the behaviour of animals is as precisely attuned to
their ecology as their morphology is. Most of this
advance in the measurement of Darwinian variables
has been due not so much to post-Darwinian advances
in the technology of measurement, but to a new wil-
lingness to spend thousands of hours observing and
tallying selectively significant behaviours.

Behavioral Mechanisms in Ecology even-handedly
reviews this new discipline, discussing theory and
results in the study of foraging, habitat selection, pre-
dation, maintenance behaviour, reproduction, com-
petition and sociality. The 1190 references cited are a
comprehensive review of the relevant literature up to
1978. The presentation of theory is largely nonma-
thematical, but only quantitative results are consi-
dered. These are presented in an astonishing number
of graphs, mostly redrawn from previous publica-
tions. There are only 33 illustrations which are pic-
tures of animals (expressively drawn by Jaquin B,

BOTANY

THE CANADIAN FIELD-NATURALIST

Vol. 95

Schulz), but 141 figures plot one variable against
another. Many of these graphs are both very simple
and very large. A third of page 268, for example is
taken up by a scatter diagram of three points, and a
half of the page by a caption explaining the points.
There are, consequently, almost no tables of data.

It is, perhaps, a reflection of the causal importance
of natural selection that the data closest to reproduc-
tive fitness (reproductive investment and parental
roles) seem to fit simple theory most closely, while the
behaviours which are most remote from a direct
payoff in fitness (foraging strategy) seem to be the
most difficult to preduct by theories of simple optimi-
zation. Morse traces the influence of these complicat-
ing factors. The influence of the danger of predation
on behaviour is a recurrent theme, and the volume
ends with a call for more research on the hard-to-
observe events that are the foundation upon which
more visible behaviour is built: the fate of dispersers,
predator-prey interactions, and interspecific
competition.

This is material which must be understood by any
naturalist who aspires to research which goes beyond
the enumeration of data for their own sake, and
Behavioral Mechanisms in Ecology is a very fine
introduction to it.

FREDERICK W. SCHUELER

Herpetology Section, National Museum of Natural Scien-
ces, Ottawa, Ontario KIA 0M8

Guide to Forest Understory Vegetation in Saskatchewan

By W.C. Harris. 1980. Technical Bulletin No. 9, Saskatche-
wan Department of Tourism and Renewable Resources,
Regina. 106 pp. $10.00

This short, non-technical guide will be a useful,
colourful accompaniment to the average naturalist,
especially in northern Saskatchewan, but also in most
other areas of boreal forest vegetation. It is not meant
for the serious botanist. Its stated purpose is to aid
forestry oriented personnel in assessing site conditions
more fully and it will likely be a satisfactory initial
training aid. To more fully meet that aim there should
have been more emphasis on mosses and liverworts
and abbreviations of species names should have been

provided for field use. The guide includes a table with
number codes describing soil drainage classes as well
as a forest cover map of the province, a glossary, an
index and several appendices but the actual species
descriptions very rarely refer to these parts of the
guide and asa result are inefficient. In effect, the guide
may be superficially useful to government forestry
personnel but it will probably be more useful to the
common or garden naturalist.

The plant descriptions (of about 80 species) are
given one species to a page with short descriptions
under the headings of Growth Habit, Leaves, Flow-
ers, Fruit, and Habitat. This format makes for easy

1981

skimming in absence of a key. The photographs are
large and well composed: in most cases two or three
photos are given to show temporal stages in the plant
or flower’s development. Only a few of the photos are
of questionable quality, such as those of the two
strawberry species. It is unlikely that anyone could
differentiate the two just by the photos provided.
There are some omissions because of the small
number of species shown. For example there are only
two lichens shown and only a few mosses, ferns and
grasses.

This guide is of sturdy construction with plastic
binding, an unusual cover photo and pages which
don’t stick together when wet. It will fit in one’s back
pocket and overall it is pleasing to the eye and is
clutter-free. These are the main features which

BOOK REVIEWS

489

recommend it alongside several other books of similar
content. Although no acknowledgment is made, the
layout of photos and text are almost identical to those
in Wildflowers Across the Prairies (by Vance, Jowsey
and McLean) and in Saskatchewan the two books are
complementary in content. It shouldn’t take too long
for the avid naturalist to digest this book and progress
to more specialized works. As a stepping stone, it is a
worthwhile guide to most of the common understory
plants throughout most of boreal Canada.

JAMES A. ROWSELL

Beak Consultants Limited, 6870 Goreway Drive, Missis-
sauga, Ontario L4V 1L9

Flora of the Prairie Provinces: A Handbook to the Flora of the Provinces of Manitoba,
Saskatchewan ‘and Alberta; Part I. — Pteroids, Ferns, Conifers and Woody Dicopsids;
Part II. — Digitatae, Dimerae, Liberae; Part III. — Connatae; Part IV. — Monopsida.

By Bernard Boivin. Part I, Provancheria No. 2, Universite
Laval, Quebec; 1967; 202 pp. $5.00 paper (Reprinted from
Phytologia vols. 15 & 16). Part II, Provancheria No. 3;
1969; 185 pp. $5.00 paper (Reprinted from Phytologia vols
16-18). Part III, Provancheria No. 4, 1972; 224 pp. $6.00
paper (Reprinted from Phytologia vols. 22 & 23). Part IV,
Provancheria No. 5, 1979; 189 pp. $8.00 paper (Reprinted
from Phytologia vols. 42 & 43). Set $20.00, available from
the author at Herbier Louis-Marie, Universite Laval,
Quebec, GIK 7P4, Canada.

With the appearance of Part lV — The Monopsida,
Boivin’s Flora of the Prairie Provinces is now com-
plete in taxonomic coverage with the single, but
major, exception of the grass family. The latter, along
with a general index, bibliography and glossary are
now planned by the author as Part V of this Flora
series. Perhaps a review of the Flora at this time might
be perceived as either long overdue for Parts I-III, or
still premature for the whole finished work.

The introduction describes the scope of the work,
outlines some evolutionary principles, and explains
the author’s views on species concepts and key con-
struction. The text-body represents a classical flora
with simple dichotomous indented keys to major
groups, families, genera and species. For each species,
the scientific name is followed by the pertinent syn-
onyms, common names (if any), brief descriptions,
indications of phenology (general flowering time),
short habitat notes, and much abbreviated geographi-
cal ranges. Infraspecific taxa are not separated by keys
but treated within the text under each species, along
with their own synonyms, habitat, and distribution.
Short taxonomic or distributional comments are
often included.

This Flora by a foremost Canadian floristic taxon-
omist constitutes a complete, critical, concise and rel-
atively up-dated treatment of the vascular flora of the
Canadian Prairie Provinces. Preparation of the Flora
has obviously involved much extensive and painstak-
ing library and herbarium searching, compilation,
and synthesis of information by the author. Unusual
species reports have been indefatigably traced down
to verify or discount them. The resulting work is com-
prehensive, intellectually stimulating, innovative, and
often controversial, and at least indirectly it points up
numerous taxonomic difficulties in our flora needing
further study. Clearly, from a scholarly point of view,
Boivin’s Flora must be considered a landmark in
Western Canadian floristics.

Nevertheless, several shortcomings or at least dis-
tracting elements are perceived in this work. The keys
to species are composed of unusually brief leads,
mostly employing only a single character. Generally
the choice of key characters seems good, with easily
observable and decisive ones used, although other
apparently useful or even better characters are often
omitted, sometimes even from the brief descriptions.
Because of their simplicity, the keys are remarkably
efficient to use, but only if one has excellent plant
specimens on hand revealing the required characters.
Such is not necessarily nor even usually the case.

Taxonomic judgements of how narrowly or
broadly species are interpreted (i.e. — “splitting” vs.
“lumping”) must be considered privileged — not a
matter of correctness or incorrectness per se. Boivin
has explained his philosophy of species delimitation in
the introduction and has reiterated or amplified this at

490

various places elsewhere in the text. With some nota-
ble exceptions, he has followed his stated concept in
practice. In my opinion, Boivin’s species’ circumscrip-
tions for the most part tend overly conservative (i.e.
—too broadly inclusive), with the apparent result
that various taxonomic problems and the distinctions
between similar but I think often meaningful species
or variants have been obscured by sweeping them
under a carpet of synonomy. The tongue-in-cheek
accusation sometimes made that angiosperm taxon-
omists accept species differences no smaller than the
10X level of mignification, seems extended by the
author to almost the “eyeball-level”! I also think that
too many major taxonomic reinterpretations and
associated nomenclatoral changes have been made by
the author with less than satisfactory explanations.
One is thus inclined to accept any such changes only
tentatively, if at all, until substantiated by other
authorities.

This Flora seems to represent primarily a herba-
rium compilation, with field characteristics under-
stressed and habitat information sometimes lacking
precision. The taxonomic conclusions and various
habitat/ distributional notes suggest that the author
has in all likelihood acquired his knowledge of the
flora of Western Canada mainly through available
herbarium specimens rather than from extensive field
experience here.

An inconvenience to most users of Boivin’s Flora
will be his unfamiliar sequence of families, necessitat-
ing a frequent checking of the separate indices of each
booklet. Instead of accepting the conventional and
familiar Englerian System, as have most flora-
manuals, or the modern phylogenetic systems of
Cronquist or Takhtajan, as have a few recent manu-
als, Boivin has basically followed the Hutchinson Sys-
tem with its major separation of the dicotyledonous
angiosperms into primarily woody and herbaceous
families. The General Index planned for Part V will be
welcomed.

Technical botanical terminology is used through-
out. Some frequent terms are ones seldom encoun-
tered elsewhere, even in other technical works (e.g.
—isomegueth, dimegueth, tegule). A glossary is
promised in Part V. At least initially disconcerting to
some English readers may be the frequent usage of
several unfamiliar French terms, especially for topo-
graphic features (e.g. — “Boise Coteau”, for the
Cypress Hills, Wood Mountain, and adjacent pla-
teaus; “Coteau de Prairie” for the Manitoba Escarp-
ment; “La Petite Montagne de Cypres” for the Sandi-
lands Uplands in south-eastern Manitoba). Common
names are not of great concern to most professional
taxonomic botanists since these lack standardization,
but I think they do represent an important link with

THE CANADIAN FIELD-NATURALIST

Vol. 95

amateur botanists. In Boivin’s Flora this professional-
to-amateur link is much weakened by the fact that
sucha large percentage — I would estimate about half
— of the common names given by the author are not
those in common usage and familiar to Western Can-
adians.

In this day of expensive flora-manuals, the low
price of Dr. Boivin’s Flora of the Prairie Provinces
appears to be a remarkable bargain. At least some of
this cost economy, however, seems achieved at a
rather serious sacrifice in the technical quality and
visual attractiveness of the published product. The
booklets have been economically reproduced by a
reductive photographic offset method directly from
the original typescript. The work suffers from the
usual technical deficiencies of such production
methods. In addition, the original typescript appears
to have lacked careful typing, editing and
proof-reading.

The eye-disturbing, but really inconsequential,
non-capitalization of proper adjectives (e.g. — cana-
dian, europaean, linnaean) throughout the text seems
too consistent to be blamed on typographical errors.
The soft-paper covers and bindings of the booklets are
hardly of high or lasting quality, although the stapling
at least adequately maintains the booklet integrity.
Clearly the value of this publication is in its informa-
tional content rather than its form. But the quality of
the former, and its importance as a scholarly contribu-
tion to North American botany, surely deserved a
better product packaging. I find it surprising that this
important Flora would not have been subsidized or
produced as an official publication of the Canada
Department of Agriculture.

The afore-mentioned deficiencies, I fear, will likely
reduce the popular appeal and utility of this Flora for
undergraduate students, non-taxonomic botanists,
amateur plant enthusiasts, general naturalists and
other lay-persons, contrary to the stated goals in the
Preface. It does not represent a particularly practical
guide to be used singularly for plant identification, but
rather seems best used in combination with other
flora-manuals. But the possible utilitarian lack for
non-professional users does not detract at all from the
high scholarly value of this publication series as an
indispensable reference source for systematic bota-
nists studying the flora of this region and elsewhere in
North America. I would strongly recommend that no
important plant taxonomic library should be without
this Flora.

VERNON L. HARMS
The W.P. Fraser Herbarium, Departments of Plant Ecology

and Biology, University of Saskatchewan, Saskatoon, Sas-
katchewan S7N 0WO

1981

Living with plants: a guide to practical botany

By Donna N. Schumann. 1980. Mad River Press, Eureka,
California, xx + 328 pp., illus. US $14.50.

If ever you've felt the frustration of not knowing
why you are doing what to whom in the garden, Living
With Plants could solve your problem. Donna Schu-
mann has done an admirable job of bridging the gap
between pure botanical science and the strictly practi-
cal side of horticulture.

The book was written for use in the author’s course
in gardening, and thus it covers all popular aspects of
the topic. The reader is immediately introduced to a
plant, complete with reams of new vocabulary through
which to wade. It is easy to see why botany is often
referred to as the science of inventing new words.
Professor Schumann valiantly tries to make this
necessary chapter as painless as possible through the
use of profuse, clear and well-labelled drawings, some
of a rather less conventional nature. (My favourite is
the marriage of Miss Glucose and Mr. Fructose to
produce Mr. and Mrs. Sucrose as an aid to under-
standing enzymes). However, once digested, the
author moves on to more familiar ground. Armed
with this introduction, the readers are equipped to
understand why they use rooting hormone, how pest
control is really supposed to work and why poinsettias
need darkness to bloom, along with a host of other
quandries that often plague the curious gardener.

Although the book does answer a lot of horticultu-
ral whys, it functions mainly as a manual of horticul-
tural practices. Instructions for all basic gardening
chores are given clearly and are almost always accom-
panied by good illustrations. There are lots of charts
of plants the author recommends for special purposes,
as well as useful charts on comparative herbicides and
pesticides. Since Living with Plants is an American
publication, the trade names in the latter charts to not
always apply here. However, the chemical names are

BOOK REVIEWS

491

universal and a great help when deciding among the
profusion of products available on the nursery shelf. It
is interesting to note that most of the Organochlorine
pesticides, the group to which DDT belongs, have
been banned or use-restricted in the U.S., whereas
they are widely available in Canada. This section on
‘Meeting the Enemy’ is especially modern as the
author tries to integrate discussion of chemical war-
fare and organic controls to overcome garden pests.
Most gardeners are not ambitious enough to expend
the greater effort required to use organic gardening
techniques and too often fall back on the simpler but
more dangerous chemicals. Mrs. Schumann tries to
strike a medium between both approaches, discourag-
ing the indiscriminate use of chemicals.

Most of the maps used in the book generally do not
extend beyond the American border, and many of the
plants listed are only found in southern climates.
However, with care and a little imagination, the maps
and plant lists can be interpreted for use in southern
Canada.

As a whole, the book is well written and a useful
reference tool for gardeners. What makes it unique is
its ability to explain in relatively simple terms why a
procedure works so that the gardener can truly under-
stand why he or she is using a specific product, tech-
nique or tool. (Also why Grampa’s advice about prun-
ing was right!) In the long run this knowledge can be
used to help the individual intelligently adapt tech-
niques to his or her unique requirements, rather than
be left to slavishly follow mysterious recipes, making
the whole experience of gardening that much more
exciting and satisfying.

DALE Hoy

726 Pickering Beach Road, Ajax, Ontario LIS 3K8

Common and Botanical Names of Weeds in Canada/
Noms Populaires et Scientifiques des Plantes Nuisibles du Canada

By Jack F. Alex, Richard Cayouette and Gerald A. Mulli-
gan. 1980. Agriculture Canada, Publication 1397
(Revised). Supply and Services Canada, Hull. 132 pp.
$5.75 in Canada; $6.90 elsewhere.

This publication is an integration of Common and
Botanical Names of Weeds in Canada (Canada Weed
Committee 1969) and Noms des mauvaises herbes du
Québec (latest edition, Ferron and Cayoutte 1975).
With one exception, the rules for spelling English
common names, the book is completely bilingual,

with English and French parallel columns on each
page.

The bulk of the material is presented in four large
tables. In Table 1, the entries are arranged alphabeti-
cally by botanical (Latin) name. The family name, one
common name in English, one common name in
French and a four-digit code number assigned by the
Expert Committee on Weeds (formerly the Canada
Weed Committee) are given for each species. Tables 2
and 3 present the same information, except for family

492 THE CANADIAN FIELD-NATURALIST

name, arranged alphabetically by English and French
common name respectively. Table 4 gives the botani-
cal names arranged in numerical order of their code
numbers.

More than 900 species have been designated as
weeds by the authors. In addition, several species have
been divided into their most important varieties (e.g.
Rhus radicans is given as 1546 — R. radicans L. var.
negundo — poison-ivy; 1960 — var. radicans — east-
ern poison ivy; and 1961 var. rydbergii — Rydberg’s
poison-ivy). Many readers of the Canadian Field-
Naturalist may be surprised to find that species such
as Acer saccharum — sugar maple, Arisaema atro-
rubens — Jack-in-the-pulpit, and Ulmus americana
— American elm are listed as weeds. Similarly, agri-
culturalists might ask why alfalfa, orchard grass, red
clover, white clover, oats and barley are included in
the list. The authors explain that “many plants that
are regularly or occasionally cultivated for crop,
ornamental or other use may reappear at some other
time or place as unwanted plants. Such plants are
weeds in the sense that they are growing where some-
one does not want them, and have been included
ene

A few species that are considered weeds by some
people in Canada are not included. In southwestern
Ontario some farmers have dense infestations of bur
cucumber (Sicyos angulatus L.) in their fields. None
of the various species of forget-me-not (Myosotis L.)
are listed but some gardeners claim that these plants
constitute a serious weed problem. Lonicera tatarica
L. is included but L. japonica Thunb. is not; several
botanists claim that the latter is more of a weed than
the former.

Despite the above comments, this book does list the
vast majority of plants that are weedy in Canada.
Many weeds of local distribution in Canada such as
smooth pigweed (Amaranthus hybridus L.) are listed
as well as the obvious widespread species such as wild
oats (Avena fatua L).

In general, there is only one English name and one
French name recognized for each species. However, if
a second English common name is in fairly wide-
spread use throughout part of Canada it may be listed
in Table 2 with the direction “see/voir” the accepted
common name. For example, couchgrass is in Table 2
followed by “see/voir grass, quack”. Also, “‘creeping
Charlie see/ voir ground-ivy.” Only with the accepted
common name do you get the code number, botanical
name and French name. In Tables 1, 2 and 3, there is
only one French name for each species. There are a
few weed species that do not occur in French-speaking
areas of Canada and for which there was no accepta-
ble French common name (e.g. Cenchrus longispinus
(Hack.) Fern. — long-spined sandbur). These species
are left without a French name.

Vol. 95

What is the purpose of publishing this book and
who will use it? The authors point out in their intro-
duction that “the common, or vernacular, name (of a
particular Canadian weed) often differs from one
region to another.” Further, they note that there are
“variations in the form and spelling of weed names in
Canada and in foreign literature”. All of this variation
can cause confusion for farmers or gardeners who
wish to control a particular weed, but through using
the wrong name for the plant, spray it with a herbicide
to which it is resistant. Advisors and agricultural
representatives must also know exactly which weeds
are growing in a given field. There is increasing evi-
dence that many closely related species which are very
similar in appearance can differ in such critical aspects
as response to herbicides, kind and amount of dor-
mancy, pattern of seedling emergence, and ability to
perennate.

As the formulation and testing of herbicides has
become more exact the scientists involved with this
area of research have come to realize that errors can
arise because a herbicide that is recommended for the
control of one plant species has been sprayed on a
similar-appearing but quite different species. Part of
the problem obviously lies with errors in identification
but further complications arise when a single common
name is used for different species. This book will be of
great assistance in correcting the latter problem.

The main purpose of assigning code numbers to
each species or taxon is to enable the “Expert Com-
mittee on Weeds” to implement computerization for
the publication of its annual Research Reports and
Abstracts. Code numbers beginning with the numeral
1 refer to weeds per se, while those beginning with 9
refer to crops as weeds. This sytem will undoubtedly
prove useful in the future for many other purposes
such as the production of distribution maps, the list-
ing of various parasites and predators, etc. It would be
possible to extend this code number system to all
Canadian plant taxa and this could lead to many
additional uses for the system.

However, there are drawbacks to using a system of
code numbers. An obvious problem is illustrated by
the three varieties of poison-ivy mentioned earlier in
this review. When a new species is added to the list or
the name of a species is changed, the code numbers
cannot always be adjusted to accommodate this
change. Thus, within a single species such as poison-
ivy you can have two quite different code numbers. A
simple letter code using the first six letters of the
botanical name might ultimately be more useful.

The book is produced attractively. It features large
clear type and a coloured picture of a most important
toxic weed, tansy ragwort (Senecio jacobaea L.) on
the cover. At 28 X 21 cm in dimensions and with 132
pages, it is too large to be handy in the field, but

1981

undoubtedly its major use will be as a reference book
in offices across the country. It has proved to be a
well-thumbed, frequently-consulted text in our
laboratory.

ENVIRONMENT

BOOK REVIEWS

493

P.B. CAVERS

Department of Plant Sciences, University of Western Onta-
rio, London, Ontario N6A 5B7

Extinction: The Causes and Consequences of the Disappearance of Species

By Paul and Anne Ehrlich. 1981. Random-House, 304 pp.
US$21.00

Rare Birds’ Nest Stolen! Sperm Whales Protected!
Falcons Fly Again!

Recently, animals threatened with extinction are
getting a lot of press coverage. The escalating extinc-
tion rate of wildlife species, where wildlife means all

wild animals and plants, is becoming an issue of global
concern. The World Conservation Strategy, a blue-
print for global conservation of natural resources,
includes the preservation of genetic diversity as one of
the Strategy’s three principle goals. The eminent
socio-biologist Dr. E.O. Wilson considers the contin-
uing extinction of wildlife species the greatest catas-
trophe of our times because the future of mankind
becomes more constrained with each extinction.

A new book, Extinction, by Paul and Anne Ehrlich
(authors of The Population Bomb and The End of
Afluence), looks at this subject in considerable depth.
In doing so, the Ehrlich’s develop an excellent anal-
ogy: the authors see each extinct wildlife species as a
rivet popped from Spaceship Earth. Although the
ship will withstand a few missing rivets, the cumula-
tive effect will be disasterous.

As professional biologists, the Ehrlich’s have con-
siderable expertise with wildlife. They present a com-
prehensive look at the importance of wildlife, the ways
in which wildlife species have been and are being
pushed to extinction, and the potential remedies to
this “rivet popping”. Throughout, interesting insights
are made into “the extinction factors”. Of particular
interest is a section on prehistoric extinctions. It is
hard to imagine the anger of future generations
deprived through the loss of some present wildlife
species. A discussion of species deprived us by our
earliest and more recent ancestors allows us to feel
some real anger.

A second insight is into the central role of habitat
conservation in the fight to keep wildlife species thriv-
ing. The single species approach to conservation (the
tiger, the rhino, the panda), although very successful,
has to give way to a broader thrust directed at habi-
tats. This is especially true of the Tropical Rain
Forests which, although covering only 6% of the

earth’s surface, contain nearly 50% of wildlife species.
These forests are disappearing at an incredible rate
(conservatively estimated at 50 acres per minute). This
short-sighted destruction of tropical forests spawns
another analogy: trees are terrestrial whales.

The book is exhaustive, providing all the basic
information about wildlife extinctions. The book
breaks each large area into many small sections. The
notes to each chapter are extremely complete, provid-
ing tonnes of additional reading for the keeners. This
comprehensive approach leads to the stacking of
detail on detail like cord wood making the book hard
to read at times. At its worst, it reads like a pneumatic
drill sounds. Such is the danger inherent in carefully
outlining the direct economic benefits of plants to
mankind, or the nefarious ways Homo sapiens exter-
minates wildlife — directly and indirectly.

To the concerned, the book provides the vivid
background to a biological crisis. To the converted,
little of this book is new. The solutions proposed are
familiar friends: stabilizing world population, found-
ing a new economic order, replacing short-term gain
with longterm benefits. In the end, after considerable
discussion of the need for habitat preservation and a
new thrust, the Ehrlich’s conclude:

“What may be needed now is a great act of

leadership. Just as President Kennedy put the

U.S. on the road to space, another president
could put the nation — and, one would hope,
the world — onaroad to treating the Earthasa

Spaceship”.

Extinction expertly details an old but escalating
problem. What we need now is action on a grand
scale. The World Conservation Strategy suggests a
number of concrete solutions. The World Wildlife
Fund’s coming campaign on The Tropical Rain
Forests is also an encouraging sign. In short, the time
for thinking and theorizing is over. We know what to
do. Let’s do it. Now.

DAVID LOVE

World Wildlife Fund (Canada), 60 St. Clair Avenue E.,
Toronto, Ontario M4T 1N5

494 THE CANADIAN FIELD-NATURALIST

Vol. 95

Natural Resource Conservation: an ecological approach

By Oliver S. Owen. 1980. 3rd edition, MacMillan, New
York. 883 pp., illus. US$19.95

The last few decades have seen a public environ-

mental awakening. One result has been many new
university courses. It is ecological dogma that when an
unexploited resource exists, an organism will evolve
to fill that niche. The plethora of textbooks aimed at
“ecology and man” or “resource management”
courses seems consistent with this. It is inevitable that
many of these books will be less than adequate. Faced
with a continuing inflow of such books from publish-
ing houses, I have set down a list of three criteria
which, in my opinion, all such books should meet.
(1) Ecology as a science. All such textbooks should
first acquaint students with the basic concepts of ecol-
ogy. Little is to be gained by introducing complex
ecological problems until the student first under-
stands basic concepts such as energy flow, nutrient
cycles, population dynamics, ecological succession,
biomes, etc.
(2) Ecology andresource management should then be
covered. Students can learn that ecological principles
can show us how to scientifically evaluate problems
and then systematically design methods to solve them.
(3) In depth treatment. Since the foregoing would
tend to give students a superficial view of many areas,
occasional in depth treatment is necessary. In depth
treatment would enable students to examine at least a
few problems to see the different perspectives and
complex interactions that face any problem solving
institution or committee.

It is particularly important that a text on ecology
and man be well written. Many engineers, general arts
students and landscape architects, for example, may
get their one and only introduction to ecology through
sucha text. Poor understanding of basic concepts may
have severe economic, ecological or political reper-
cussions in years to come. |

Owen introduces basic ecological principles in one
chapter, 34 pages. It might be argued that a scanty 34
pages were necessitated by space limitation, but the
book is more than 800 pages long.

Resource management problems discussed include
soil erosion, pollution, forestry, wildlife, pesticides
and energy. However, the treatment of these problems
is uneven. The chapter on wildlife begins with a gen-
eral discussion of the problem of extinction. Yet, of 76
pages on wildlife, 34 are devoted solely to the produc-
tion of a few species of animals for North American
sportsmen to shoot. If there was insufficient space to
deal with the ecological factors regulating species
diversity, destruction of tropical rain forest and coral
reefs, the fate of African game reserves, etc., surely it

could have been gained by: (1) reducing emphasis on

intensive management of a few game species, or (2) by
eliminating the excessive use of photos of dead ducks,
deer, and fish.

Moreover, there is a lengthy discussion of the
infamous Kaibab deer herd, which Owen calls “classic
example of a deer irruption” and graphically describes
how... “deer now began to succumb to slow, agoniz-
ing death by starvation. Within the six-year period
from 1924-1930, 80,000 deer starved, their gaunt ema-
ciated carcasses bearing eloquent testimony. ...” Yet,
10 years ago, Caughley (1970) demonstrated that the
Kaibab example was at best careless science, at worst,
outright fiction! It is thoroughly discussed in Colin-
vaux’s (1973) text. It is inexcusable that ten years and
two printings later Owen is still repeating the Kaibab
story, complete with a redrawn figure and completely
imaginary data points neatly labelled!

This leads directly to the third criterion: illustrating
the complexity of ecological problems. Let me cite
three of many possible examples. First consider food
production. Owen starts soil management with the
title “Need for Greater Food Production by American
Farmers” and states that the United States “must
export food to help prevent massive starvation”.
There is no mention of alternate viewpoints such as
Lappé and Collins (1977) who present a convincing
case that world starvation is largely a result of inequit-
able land ownership in developing countries, and the
growing of export crops (coffee, tea, sugar cane)
instead of food. Thus, at present, starvation may be as
much a political problem as an agricultural one.
Second, consider energy. The concept of energy qual-
ity (Lovins 1977) is essential for a scientific perspective
of energy demand. For example, roughly half of all
usable energy consumed in Canada is used simply to
produce low grade heat (less than 140°C) (Croweral.,
1978). Yet, Owens does not introduce the reader to
energy quality. Neither is Lovins’ (1977) proposed
“soft energy path” presented as a distinct alternative
to the “hard energy path” we have taken the last few
decades. Third, consider fisheries. Owen adds three
paragraphs entitled “Dam versus Darter: a Classic
Confrontation” in reference to the controversy over
the Tellico Dam and the Snail Darter. This overly-
simplistic analysis ignores the fact that there were
many other economic and environmental reasons for
scrapping the Tellico Dam (Ballal, 1977).

The fact that this book is inits third edition suggests
that it has its redeeming points — and indeed it has.
For example, it provides a wealth of practical exam-
ples that are directly relevant to American students,
and many of the figures and photographs visually

1981

emphasize concepts that are presented inthe text. But
it is no longer enough to add a few new paragraphs to
an old text. It is remarkable to think how much one
additional decade of work has changed our perspec-
tives on planetary survival.

I rate the text a minimal pass on my first criterion,
and a failure on the next two. Better texts are now
available, and instructors and students would do well
to consult them.

Literature cited:

Ballal, S. K. 1977. Tellico dam furor involves more than
snail darter. Bioscience 27: 586.
Caughley.G. 1970. Eruption of ungulate populations, with

MISCELLANEOUS

BOOK REVIEWS

495

emphasis on Himalayan Thar in New Zealand. Ecology
51: 53-72.

Colinvaux, P. A. 1973. Introduction to Ecology. John
Wiley and Sons. N.Y.

Crow, R., P. Szegedy-Maszak and C. Conway. 1978.
Energy Planning in a Conserver Society. Energy Probe.
Lappé, F. M. and J. Collins. 1977. Food First: Beyond the
Myth of Scarcity. Houghton-Mifflin Company, Boston.
Lovins, A. B. 1977. Soft Energy Paths: Toward a Durable
Peace. Ballinger Publishing Company. Reprinted 1979 by

Harper and Row. N.Y.

PAUL A. KEDDY

Department of Botany and Genetics, College of Biological
Science, University of Guelph, Guelph, Ontario NIG 2W1.

Hawk Lady: the story of a woman who opened her home to care for wild birds of prey

By Stellanie Ure. 1980. Doubleday and Company, Toronto,
215 pp., illus. $14.95

The “hawk lady” of the title, Stellanie Ure, has
recorded her experiences in caring for and rehabilitat-
ing injured, sick and orphaned raptors of 26 species.
She and her family have been instrumental in return-
ing many oftheir patients to the wild, and she imparts
much information on the behavior, feeding, etc., of
the many hawks and owls she treated. She is to be

Completely Foxed

By Miles Smeeton. 1980. Van Nostrand Reinhold. Toronto.
xi+ 148 pp., illus. $9.95.

After he retired from the British army in 1946, Miles
Smeeton and his wife Beryl moved to Canada, later
sailed around the world, and finally settled down on
what has become a private game sanctuary in the
foothills of the Rocky Mountains near Calgary,
Alberta. Their aim was to do what they could to help
native endangered species; this is the story of their
efforts on behalf not only of the Swift Fox as the title
of the book suggests, but also the Trumpeter Swan,
the Wcod Bison, and various other animals, especially
the Moose.

The publisher calls this a joyous book, but such an
adjective glosses over the hard work and often heart-
ache which accompany living day by day with and for
wild animals. An example was the care of Zeus and Io,
two pinioned Trumpeter Swans which the Smeetons
obtained with great effort from the government and

commended for her efforts in assisting these threa-
tened birds. Unfortunately, the book seems overly
long and detailed, and failed to sustain my interest. I
feel it would be a real value only to those actively
involved in similar endeavours.

JO WRIGHT

118 Cedar Ave., Hudson, Quebec JOP 1HO

hoped to breed so that their progeny would return to
the wild. These birds lived happily in summer on the
sanctuary pond, but in winter when the water froze
over they were at the mercy of any passing lynx or
coyote. The Smeetons thus erected a wind and motor-
driven propeller which would theoretically continu-
ally stir up the water and keep it from freezing. This
was successful only until the weather turned really
cold. Then someone had to break up the newly-
formed ice by hand each morning, hauling it to the
edge of the pool and piling it high. Since this task took
up to two hours a day, some other solution had to be
found. The next plan was to build a geodetic dome or
wood and fabric over the water to make the swans an
indoor swimming bath. This worked well until spring,
when the 600-lb dome had to be removed before the
ice melted and it sank from sight. At first the Smee-
tons tried to raise and shift it with the help of a hot- air
balloon, a job which was beyond them and sent sev-

496

eral helpers to hospital. Finally, with friends, they
managed to carry it to shore.

The following winter the well near the swan’s pond
went dry because of drought, so the Smeetons were
forced each day to siphon into the pool about five
45-gallon drums of water brought from the house.
They only gave up their efforts on behalf of the swans
when they were informed by government that no birds
raised in captivity could be released to the wild (the
whole point of their labors) and that they must pay for
a licence for the pleasure of keeping their swans. They
reluctantly turned over Zeus and Io to the Saskatoon
Zoo.

THE CANADIAN FIELD-NATURALIST

Vol. 95

Other chapters in this book are devoted to the antics
of the Swift Foxes Emma, Napoleon, Josephine, Nel-
son, and their progeny; to the Wood Bison Martha
and Mary who likes to sneak indoors to lick the butter
on the dining room table; and to various Moose who
feed from Beryl Smeeton’s hands. Those who like to
read about such an animal wonderland will enjoy both
the text and the lovely photographs.

ANNE INNIS DAGG

Integrated Studies, University of Waterloo, Waterloo, Onta-
rio N2L 3G1

Evolution for Naturalists: the simple principles and complex reality.

By Philip. J. Darlington, Jr. 1980. Wiley, Toronto, xvi+ 262
pp., illus. US $19.95.

This book should be used by both serious amateur
naturalists and advanced professional biologists . By
reading this book serious naturalists will obtain a
“self-education” in evolution while professional biol-
ogists will obtain a fresh reorientation of evolutionary
theory. Since Darlington argues about several pro-
found aspects of evolutionary theory, serious amateur
naturalists who wish to make intelligent decisions
about evolution will have to work and think while
reading much of this text. Professional biologists may
feel the text is too verbose until one of Darlington’s
innovative, unconventional ideas causes a re-reading
of sections and careful thinking. The text is replete in
many areas with very “simple principles” that every-
body understands while in others are “complex reali-
ties” upon which professional biologists have yet to
agree. Therefore, the subtitle of this book perhaps
describes this text best.

Darlington verbally separates naturalists from pro-
fessional biologists throughout his book but he has
done an inadequate job trying to create such a dichot-
omy. Many areas of the text are clearly written for
uninformed naturalists while others are much too
involved for such persons. Steming from his objective
to write for uninformed persons by using little termi-
nology, grammatical errors and clumsy phrases
become numerous.

Darlington feels that there have been many evolu-
tionary principles uncritically accepted with bias and
therefore many theories are most likely wrong. He
provides many reassessments of evolutionary
thoughts not available in other texts. An excellent
discussion is provided on the Darwinian revolution
(chapter 1). The small print on page 5 should be

carefully read by all persons interested in evolution
because each will be awakened to the fact that a per-
fected theory of natural selection was published by a
Patrick Matthew in an obscure journal before Darwin
first boarded the Beagle. Kin and group selection are
put into contexts totally unfamiliar to both amateur
naturalists and professional biologists. Good argu-
ments are provided concerning population-wide
reciprocal altruism inhibiting kin-limited altruism.
The primacy of the chicken/egg hypothesis is nicely
discussed. Darlington sees natural selection as the
“selective elimination of individuals that walk in the
wrong directions”. He feels that competitive exclusion
is the result of convenience since the choice for or
against coexistence is facultative. Another idea is
“Genes that promote evolution . . . by reducing their
own stability actually promote their own survival.”
An informative chapter (9) on human evolution is
perhaps the best in the book. Darlington consistently
provides good reasons for deploring the oversimpli-
fied approach used in many scientific journals pur-
porting to construct mathematical models of
evolution.

Typographical errors are scarce and the text reads
with facility. A “reading and reference” list at the end
of each chapter cites literature by subject and an excel-
lent index appears. University professors should find
this text the best newer book available for evolution-
ary seminars and discussion groups held with both
undergraduate and graduate students. One’s specula-
tive power will greatly increase after reading this
book.

RICHARD M. ZAMMUTO

Kananaskis Environmental Centre, University of Calgary,
Seebe, Alberta TOL 1X0

1981

BOOK REVIEWS

497

British Natural History Books, 1495-1900: A Handlist

By R.B. Freeman. 1980. Dawson (Distributed by Shoe
String Press, Hamden, Connecticut). 437 pp. US $29.50.

In this volume the author has brought together 4206
titles of books published in a period of just over three
centuries, ending in 1900. These titles include books
about the flora and fauna of the British Islands,
including Ireland, the Channel Islands and Heligo-
land. Also included are general natural history books
written by the inhabitants of these islands, as well as
translations into English of relevant books by foreign
authors, plus a few American books which have
appeared in British editions.

The book is divided into four sections: a short
introduction which outlines the scope, problems
involved, methods of citation, inclusion of various
editions, etc.; the main alphabetical listing of titles,

NEW TITLES

Zoology

*Advances in Ephemeroptera biology. 1980. Edited by
John F. Flannagan and K. Eric Marshall. Plenum Press,
New York. 552 pp., illus. US $49.50 (US $59.50 outside
USA).

The age of birds. 1980. By Alan Feduccia. Harvard Uni-
versity Press, Cambridge. xii + 196 pp., illus. US $20.

Animal tracks of the Pacific northwest. 1981. By Karen
Pandell and Chris Stall. Mountaineers (Distributed by
Douglas and McIntyre, Vancouver). 96 pp., illus. $4.95.

Arctic animal ecology. 1980. By Hermann Remmert.
Springer-Verlag, Berlin. 250 pp., illus. US $24.80.

* Atlas of North American freshwater fishes. 1980. By D. S.
Lee, C. R. Gilbert, C. H. Hocutt, R. E. Jenkins, D. E.
McAllister, and J. R. Stauffer, Jr. North Carolina State
Museum of Natural History, Raleigh. 825 pp., illus. US
$20.

The Audubon Society book of marine wildlife. 1980. By
Les Line and George Reiger. Abrams. New York. 240 pp.,
illus US $45.

Australia’s animals discovered. 1981. By P. Stanbury and
G. Phipps. Pergamon Press, Oxford. 128 pp., illus. US
$23.40.

Basic structure and evolution of vertebrates. 1980. 1981.
By Erik Jarvik. Academic Press, London. Volume I, xvi+
576 pp. US $94.50; Volume 2, xiv + 338 pp. US $56.50.

with dates and places of publication and publisher,
where known, each entry being numbered sequen-
tially; a list of titles in date order from 1495 to 1900;
and a subject index which has been derived mainly
from information given in the titles.

It is impossible to assess the completeness of a work
suchas this, especially from this continent, but there is
no question that the author has delved deeply into the
early natural history literature of the British Isles. The
book will be of considerable use to anyone who needs
to search out an old title or who wishes to trace out the
early natural history of these islands.

WILLIAM J. CODY

Biosystematics Research Institute, Canada Agriculture,

‘Ottawa, Ontario KIA 0C6

*Behavioral mechanisms in ecology. 1980. By Douglas H.
Morse. Harvard University Press, Cambridge, Massachu-
ssetts. 371 pp., illus. US $25.

Biosocial mechanisms of population regulation. 1980.
Edited by M. N. Cohen, R. S. Malpass, and H. G. Klein.
Yale University Press, New Haven. 406 pp. US $22.50.

Birds of Africa. 1980. By J. Karmali. Viking Press, New
York. 191 pp. US $25.

Birds: readings from Scientific American. 1980. Edited by
B. W. Wilson. Freeman, San Francisco. 276 pp. Cloth US
$17.95; paper US $8.95.

Book of mammals. 1981. By The National Geographic
Society, Washington. 2 volumes. 608 pp., illus US $22.95.

+Diversity and adaptation in fish behaviour. 1979. By
Miles H. A. Keenleyside. Springer-Verlag, New York. xiv+
210 pp., illus. US $37.80.

Echinoderms: present and past. 1980. Edited by Michel
Jangoux. Proceedings of a colloquium, Brussels, Sep-
tember, 1979. Balkema (Distributed by MBS, Salem, New
Hampshire). xviii + 428 pp., illus US $45.

The ecology and conservation of large African mam-
mals. 1979. By S. K. Ehringham. University Park Press,
Baltimore x + 286 pp., Illus. US $49.95.

The evolution of air breathing invertebrates. 1981. By
David J. Randall, Warren W. Burggren, Anthony P. Far-

498 THE CANADIAN FIELD-NATURALIST

rell, and M. Stephen Haswell. Cambridge University Press,
New York. vii + 134 pp., illus. US $27.50.

*The fallacy of wildlife conservation. 1981. By JohnA. Liv-
ingston. McClelland and Stewart, Toronto. 117 pp. $14.95.

A guide to the mammals of Ohio. 1980. By J. L.
Gottschang. Ohio State University Press, Columbus. 300
pp. US $35.

How animals hunt. 1980. By J.H. Prince. Elsevier-
Nelson, New York. 128 pp., illus. US $8.95.

Insect worlds: a guide for man on making the most of the
environment. 1980. By Lorus J. Milne and Margery
Milne. Scribner’s, New York. 274 pp., illus. US $12.95.

The island waterfowl. 1980. By Milton W. Weller. Iowa
State University Press, Ames. x + 121 pp., illus. US $10.95.

The life of the hummingbird. 1980. By Alexander F.
Skutch. Crown, New York. 96 pp., illus. US $15.95.

Mammalian communication. 1980. By Roger Peters.
Brooks-Cole, Monterey, California. x + 342 pp., illus. US
$9.95.

*A manual of mammalogy: with keys to families of the
world. 1981. By Anthony F. DeBlase and Robert E. Mar-
tin. 2nd edition. Brown, Dubuque, Iowa. xii+ 436 pp., illus.
US $9.95.

+Marine bivalve molluscs of the Canadian central and east-
ern Arctic. 1980. By Irene Lubinsky. Canadian Bulletin of
Fisheries and Aquatic Sciences No. 207. Supply and Servi-
ces Canada, Hull. vit 111 pp., illus. $9.75 in Canada; $11.70
elsewhere.

Migrant birds in the neotropics: ecology, behavior, distri-
bution, and conservation. 1980. Edited by Allen Keast
and Eugene S. Morton. Smithsonian Institute Press,
Washington. 576 pp., illus. Cloth US $27.50; paper US $15.

North American elk: ecology, behavior, and management.
1979. Edited by Mark S. Boyce and Larry D. Hayden-
Wing. Papers from a symposium. Laramie, Wyoming,
April 1978. Department of Zoology, University of Wyom-
ing, Laramie. vi+ 294 pp., illus. US $8.

*Pheasants in Asia. 1979. Edited by Christopher Savage.
Proceedings of the First International Symposium on
Pheasants in Asia. Kathmandu, Nepal, November, 1979.
World Pheasant Association, London. 116 pp. £8.50.

Pleistocene mammals of North America. 1980. By Bjorn
Kurtén and Elaine Anderson. Columbia University Press,
New York. xviii + 444 pp., illus. US $42.50.

+The population ecology of cycles in small mammals. 1980.
By James Patrick Finerty. Yale University Press, New
Haven. 260 pp., illus. US $18.50.

Vol. 95

+The pronghorn antelope in Alberta. 1980. By George J.
Mitchell. Department of Biology, University of Regina,
Regina, Saskatchewan. 165 pp. Free.

*Rare birds of the west coast of North America. 1980. By
Don Robertson. Woodcock Publications, Pacific Grove,
California. 496 pp., illus. US $24.95 plus $1.50 postage.

The red fox. 1980. Edited by Erik Zimen. Proceedings of a
symposium, Saarbrucken, Germany, January, 1979. Junk,
The Hague. vi + 286 pp., illus. US $73.50.

*La répartition géographique des poissons, les ensemence-
ments, la péche sportive et commerciale, les frayéres et la
bathymétrie du fleuve Saint-Laurent dans le bassin de la
Prairie et les Rapides de Lachine. 1980. Par Jean-René
Mongeau, Jean LeClerc, et Jocelyne Brisebois. Rapport
Technique No. 06-29. Service de !aménagement et de l’ex-
ploitation de la faune, Québec. 145 pp., illus.

Returning to Eden: animal rights and human responsibili-
ty. 1980. By M. W. Fox. Viking Press, New York. xiv +
281 pp. US $13.95.

A review of the systematics and ecology of Arctic char,
Salvelinus alpinus, in the western Arctic. 1980. By P. J.
McCart. Canadian Technical Report of Fisheries and
Aquatic Sciences No. 935. Fisheries and Oceans Canada,
Ottawa. 89 pp., illus. Free.

+Revision of the snailfish genus Liparis from arctic
Canada. 1980. By K. W. Able and D. E. McAllister. Can-
adian Bulletin of Fisheries and Aquatic Sciences No. 208.
Supply and Services Canada, Hull. viii + 52 pp., illus. $3 in
Canada; $3.60 elsewhere.

Salmonid ecosystems of the north Pacific. 1980. Edited by
William J. McNeil and Daniel C. Himsworth. Oregon State
University Press, Corvallis. x + 332 pp., illus. US $15.

The soul of the wolf. 1980. By Michael W. Fox. Little,
Brown, Boston. x + 131 pp., illus. US $12.95.

Workshop on raptors and energy developments. 1980.
Edited by R. P. Howard and J. F. Gore. U.S. Fish and
Wildlife Service, Boise, Idaho. 125 pp. US $3.

Botany

All good things around us: a cookbook and guide to wild
plants and herbs. 1980. By Pamela Michael. Holt. Rine-
hart, and Winston, New York. 240 pp., illus. US $19.95.

+Biology of plants. 1981. By Peter H. Raven, Ray F. Evert,
and Helena Curtis. 3rd edition. Worth, New York. 686 pp.,
illus. US $23.95.

*The complete trees of North America; field guide and natu-
ral history. 1980. By Thomas S. Elias. Van Nostrand
Reinhold. New York. 948 pp., illus. $26.95.

1981

A dictionary of botany. 1980. By R. John Little and C.
Eugene Jones. 4th edition. Van Nostrand Reinhold, New
York. x + 400 pp., illus. US $18.50.

Flora of New Zealand: Volume 3, adventive, cyperaceous,
petalous, and spathaceous monocotyledons. 1980. By
A. S. Healy and E. Edgar. Government Printer, Welling-
ton, New Zealand. xl + 220 pp., illus. NZ $18.50.

*Guide to forest understory vegetation, Saskatche-
wan. 1980. By W. C. Harris. Technical Bulletin Number 9.
Saskatchewan Department of Tourism and Renewable
Resources, Regina. 106 pp., illus. $10.

*Guide to the orchids of the Cypress Hills (including the
most common orchids of Alberta and Saskatche-
wan). 1980. By R. M. Fisher. R. M. Fisher, Olds, Alberta.
39 pp. $4.95.

Introduction and guide to the marine bluegreen algae.
1980. By Harold J. Humm and Susanne R. Wicks. Wiley-
Interscience, New York. xii + 194 pp., illus. US $23.50.

Inventory of rare and endangered vascular plants of Cali-
fornia. 1980. By J. P. Smith, Jr., R. J. Cole, and J. O.
Sawyer, Jr. 2nd edition. California Native Plant Society,
Berkeley. viii + 116 pp. US $7.50.

Mushrooms: wild and edible; a seasonal guide to the most
easily recognized mushrooms. 1980. By Vincent Marteka.
Norton, New York. x + 290 pp., illus. US $19.95.

Petaloid monocotyledons: horticultural and botanical
research. 1980. Edited by C. D. Brickell, D. F. Cutler, and
Mary Gregory. Linnean Society Symposium No. 8. Aca-
demic Press, xii + 222 pp. US $62.

Primary productivity in the sea. 1980. Edited by Paul G.
Falkowski. Plenum, New York. x + 532 pp., illus. US
$45.90.

?+Structure and function of northern coniferous fore-
sts. 1980. Edited by T. Persson. Ecological Bulletins 32.
Swedish Natural Science Research Council, Stockholm.
609 pp., illus. 290 SK.

*Trees and shrubs of the Dominion Arboretum. 1980. By
A. R. Buckley. Agriculture Canada Publication 1697.
Supply and Services Canada, Ottawa. 252 pp., illus. $19.95
in Canada; $23.95 elsewhere.

Wildflowers of the outer banks: Kitty Hawk to Hatte-
ras. 1980. By the Dunes of Dare Garden Club. University
of North Carolina Press, Chapel Hill. xviii+ 166 pp., illus.
US $6.95.

Wild teas, coffees, and cordials: 60 natural drinks from
Pacific northwest plants. 1981. By Hilary Stewart. Dou-
glas and McIntyre, Vancouver, 128 pp., illus. $7.95.

NEW TITLES

499

Environment

+ Advances in environmental science and engineering. 1980.
Edited by James R. Pfafflin and Edward N. Ziegler.
Volume 3. Gordon and Breach, New York. 224 pp., illus.
US $59.

American environmentalism: values, tactics, priorities.
1980. By Joseph. M. Petulla. Texas A & M University
Press, College Station. xiv + 239 pp., illus. US $18.50.

*The Arctic and Antarctic: their division into geobotanical
areas. 1980. By V. D. Aleksandrova. Translated by Doris
Love. Cambridge University Press, New York. xiii + 247
pp., illus. US $34.50.

Assessing tropical forest lands: their suitability for sustai-
nable uses. 1981. Edited by Richard A. Carpenter. Natu-
ral Resources and Environment, volume 3. Tycooly, Dub-
lin. 256 pp. US $26.

Atmospheric pollutants in natural waters. 1981. Edited by
Steven J. Eisenreich. Ann Arbor Science, Ann Arbor. 400
pp., illus. US $40.

Biophysical ecology. 1980. By David M. Gates. Springer-
Verlag, New York. xxiv + 612 pp., illus. US $39.80.

Coal burning issues: a monograph reporting results of the
scoping phase of an interdisciplinary assessment of the
impact of the increased use of coal. 1980. Edited by
A. E. S. Green. University Presses of Florida, Gainesville. x
+ 390 pp., illus. US $10.

The curious naturalist. 1980. By John Mitchell and The
Massachusetts Audubon Society. Prentice-Hall, Engle-
wood Cliffs, New Jersey. Unpaged. Cloth US $15.95; paper
US $7.95.

Dynamics, exposure, and hazard assessment of toxic che-
micals. 1980. Edited by R. Haque. Ann Arbor Science,
Ann Arbor. 496 pp., illus. US $39.95.

}Ecological land survey guidelines for environmental impact
analysis. 1981. By Environmental Conservation Service
Task Force. Ecological Land Classification Series, No. 13.
Environment Canada. Ottawa. 42 pp. English + 44 pp.
French, illus. Free.

Ecology and environmental management: a geographical
perspective. 1980. By Chris C. Park. Westview Press,
Boulder, Colorado. iv + 272 pp., illus. US $30.

Ecology of fresh waters. 1980. By Brian Moss. Halsted
(Wiley), New York. xvi + 332 pp., illus. US $24.95.

Ecology of the Canadian Arctic archipelago: selected refer-
ences. 1980. By N. M. Peterson. Volume 8. Indian and
Northern Affairs Canada, Ottawa. Pages 3182-3665. Free.

Ecosystem succession: a general hypothesis and a test
model of a grassland. 1980. By L. T. Gutierrez and W. R.
Frey. MIT Press, Cambridge. 231 pp. US $25.

500

Energy and environmental balance. 1980. By Earl Finbar
Murphy. Permagon, New York. vili+ 282 pp. US $25.

Energy Conservation through land use planning: a synthe-
sis of discussions at a symposium held in Montreal 26-28
March 1980. 1980. By W. R. D. Sewell and H. D. Foster.
Working Paper No. 6. Environment Canada, Ottawa. 91
pp. Free.

Environmental awakening: the new revolution to protect
the earth. 1980. By Rice Odell. Ballinger, Cambridge,
Massachusetts. xiv + 330 pp., illus. US $12.95.

Environmental biology. 1980. By E. J. W. Barrington.
Halsted (Wiley), New York. x + 244 pp., illus. US $17.95.

Environmental biology for engineers: a guide to environ-
mental assessment. 1980. By George Camougis. McGraw-
Hill, New York. x + 214 pp., illus. US $19.95.

The environmental impacts of production and use of ener-
gy. 1981. By Essam El-Hinnawi. Natural Resource and
the Environment, Volume |. Tycooly International, Dub-
lin. 330 pp. US $33.

Environmental risk assessment. 1980. Edited by Anne V.
Whyte and Ian Burton. SCOPE 15. Wiley, New York. xxiv
+ 160 pp., illus. US $30:

Environment, ideology, and policy. 1980. By Francis
Sandbach. Allanheld, Osmun, Montclair, New Jersey. xiv+
254 pp. US $26.50.

The environment: from surplus to scarcity. 1980. By A.
Schnaiberg. Oxford University Press, New York. xiv + 464
pp. Cloth US $14.95; paper US $9.95.

+The environment in British prehistory. 1981. Edited by
Ian Simmons and Michael Tooley. Cornell University
Press, Ithaca. 334 pp., illus. US $34.50.

*Evolution for naturalists. 1980. By P.J. Darlington.
Wiley-Interscience, New York. 262 pp. US $19.95.

Fate and weathering of petroleum spills in the marine
environment: a literature review and synopsis. 1980. By
Randolph E. Jordan and James R. Payne. Ann Arbor
Science, Ann Arbor. 655 pp. US $39.95.

Forests: fresh perspectives from ecosystem analysis.
1980. Edited by Richard H. Waring. Oregon State Univer-
sity Press, Corvallis. x + 200 pp., illus. US $12.

Idle weeds: the life of a sandstone ridge. 1980. By David
Rains Wallace. Sierra Club Books (Distributed by
Scribner, New York). viii + 184 pp., illus. US $12.95.

Interactions between herbicides and the soil. 1981. Edited
by R. J. Hance. Academic Press, London. xii+ 350 pp. US
$49.50.

Marine and shoreland resources management. 1980. By

THE CANADIAN FIELD-NATURALIST

Vol. 95

Joseph M. Heikoff. Ann Arbor Science, Ann Arbor. 214
pp., illus. US $28.95

Natural radiation environment III. 1981. Edited by
Wayne Lowder and Thomas F. Gesell. U.S. Department of
Energy, Oak Ridge, Tennessee. 1784 pp., 2 volumes. US
$39.50.

The nature of Vermont: introduction and guide to a New
England environment. 1980. By C. W. Johnson. Univer-
sity Press of New England, Hanover, New Hampshire. 276
pp. Cloth US $15; paper US $7.50.

New developments in river basin management. 1981. Ed-
ited by S. J. Jenkins. Pergamon Press, Oxford, England.
320 pp., illus. US $47.

The offshore ecology investigation: effects of oil drilling
and production in a coastal environment. 1979. Edited by
C. H. Ward, M.E. Bender, and D. J. Reish. William
Marsh Rice University, Houston. 590 pp., illus. US $11.

*Protection of natural areas in Ontario. 1980. Edited by
Suzanne Barrett and John Riley. Proceedings of a confer-
ence, Toronto. 12 April, 1980. Working Paper No. 3. York
University, Toronto. 195 pp., illus. No price given.

The recovery process in damaged ecosystems. 1980. E-
dited by J. Cairns, Jr. Ann Arbor Science, Ann Arbor. 167
pp. US $29.50.

+A reference book of urban ecology. 1981. By Anne Innis
Dagg. Otter Press, Waterloo. 190 pp., illus.

Renewable natural resources and the environment. 1981.
By Kenneth Ruddle and Walther Manshard. Natural
Resources and the Environment, Volume 2. Tycooly, Dub-
lin. 404 pp. US $40.

Speaking for nature. 1980. By P. Brooks. Houghton-
Mifflin, Boston. xi+ 304 pp. US $12.95.

*Threatened and endangered species and habitats in British
Columbia andthe Yukon. 1980. Edited by Richard Stace-
Smith, Lois Johns, and Paul Joslin. Proceedings of a Sym-
posium, Richmond, British Columbia, 8-9 March, 1980.
British Columbia Ministry of Environment, Victoria. 302
pp., illus. No price given.

The waste watchers: a citizen’s handbook for conserving
energy and resources. 1980. By Arthur H. Purcell.
Anchor/ Doubleday, Garden City, New York. xxvi + 286
pp., illus. US $4.50.

+ Watershed red: the life of the Dunk River, Prince Edward
Island. 1981. By Kathy Martin. Ragweed Press, Charlot-
tetown. 155 pp., illus. $6.95 plus 50c¢ postage.

What is ecology? 1980. By D. F. Owen. 2nd edition.
Oxford University Press, New York. x + 234 pp., illus. Cloth
US $18.50; paper US $9.95.

1981 NEW TITLES 501

Window on the wild. 1980. By Jack Denton Scott. Put-
nam’s, New York. 192 pp., illus. US $8.95.

A world like our own: man and nature in Madagascar.
1980. By A. Jolly. Yale University Press, New Haven. US
$29.95.

Miscellaneous

*From arsenic to DDT: a history of entomology in western
Canada. 1980. By Paul W. Riegert. University of Toronto
Press, Toronto. 357 pp. $30.

The art of Robert Bateman. 1981. By Ramsay Derry.
Penguin, Markham, Ontario. 180 pp., illus. $40.

Bird student: an autobiography. 1980. By George Miksch
Sutton. University of Texas Press, Austin. vill + 216 pp.,
illus. US $19.95.

TtCasey A. Wood (1856-1942): ophthalmologist, bookman,
ornithologist: a biobibliography. 1981. Compiled by E. C.
Astbury. Occasional Paper 7. Graduate School of Library
Science, McGill University, Montreal. 67 pp. $5.

*A delicate arrangement: the strange case of Charles Darwin

and Alfred Russell Wallace. 1980. By Arnold C. Brack-
man. Times Books, New York. 370 pp., illus. US $14.95.

Mathematics and statistics for the bio-sciences. 1980. By
G. Eason, C. W. Coles, and G. Gettinby. Halsted (Wiley),
New York. 578 pp., illus. US $78.95.

+Mountains and mountaineering facts and feats. 1981. By
Edward Pyatt. Guinness (Distributed by Sterling, New
York ). 256 pp., illus. US $19.95.

On the road with John James Audubon. 1980. By Mary
Durant and Michael Harwood. Dodd, Mead, New York.
Xil1 + 638 pp., illus. US $19.95.

The origin: a biographical novel of Charles Dar-
win. 1980. By Irvin Stone. Doubleday, Garden City, New
York. 744 pp. US $14.95.

+The weather almanac. 1981. Edited by James A. Ruffner
and Frank E. Blair. 3rd edition. Gale Research, Detroit. 801
pp., illus. US $48.

*Assigned for review
t Available for review.

Index to Volume 95

Compiled by W. HARVEY BECK

Abies balsamea, 50

Accipiter striatus, Sharp-shinned Hawk, (Accipitriformes:
Accipitridae) on Victoria Island, Northwest Terri-
tories, 366

Acer saccharum, 53
spicatum, 50

Agelaius phoeniceus, 63, 346, 408

Agonum cupreum, 147
(complex), 147
decentis, 147
gratiosum, 147
trigeminum, 147

Agropyron caninum, 412
smithii, 412

Agrostis hyemalis, 247

Ainley, M. G., review by, 107

Alaska, Caribou (Rangifer tarandus) encounters with pipe-
lines in northern, 57

Alaska, Moose browse utilization in Mount McKinley
National Park, 85

Alberta, 69, 133, 206

Alberta, A probable Franklin’s X Ring-billed Gull pair nest-
ing in, 474

Alberta, Dynamics of Moose populations near Rochester,
1975-1978, 39

Alberta, Great Blue Heron (Ardea herodias) colony in the
Peace-Athabasca Delta, 95

Alberta, Population characteristics and movements of
Striped Skunks (Mephitis mephitis) in central, 149

Alberta, Scheduling censuses of breeding White Pelicans
(Pelecanus erythrorhynchos) in northern, 198

Alberta, Seasonal and daily use of plant communities by
Sharp-tailed Grouse ( Pedioecetes phasianellus) in the
parklands of, 287

Alberta, Terrestrial molluscs of central, 75

Alberta, Variation in frequencies of pelvic phenotypes of the
Brook Stickleback, Culaea inconstans, in Redwater
drainage, 178

Alces alces, 50, 85

Alces alces, Moose, Estimating winter defecation rates for,
208

Alnus rugosa, 247

Alvo, R. Marsh nesting of Common Loons (Gavia immer),
357

Amara cupreolata, 147

Ambloplites rupestris, 274

Ambystoma tigrinum, Viger Salamanders, and stocked
Rainbow Trout (Salmo gairdneri): potential competi-
tors for food in Manitoba prairie pothole lakes, 129

Ammodramus bairdii, 409
savannarum, 408

Ammodytes americanus, 314

Ammospiza leconteii, 409

Ampelisca macrocephala, 190

Anas clypeata, 471

crecca, 423
discors, 423, 471
platyrhynchos, 423

Anas platyrhynchos, Mallards, in Manitoba, A time-activity
budget for breeding, 266

Andropogon scoparius, 412

Anguilla rostrata, 359

Anguilla rostrata, American Eel, from the Lake Superior
drainage, An usually small, 97

Antennaria rosea, 412

Anthus spragueii, 408

Apeltes quadracus, 359

Apherusa glacialis, 190

Aprodectodea sp., 183

Aralia spp., 323

Archilochus colubris, 311

Ardea herodias, 257

Ardea herodias, Great Blue Heron, colony in the Peace-
Athabasca Delta, Alberta, 95

Ardea herodias, Great Blue Herons, Gulls robbing prey
from, 205

Arenaria interpres, 183

Artemisia frigida, 412

Asio flammeus, 423

Asplenium platyneuron, Ebony Spleenwort, (Aspleniaceae),
in the Great Lakes area, Natural history of the, 156

Aster nemoralis, 247

Athene cunicularia, 408

Avens, Three-flowered, (Geum triflorum) in Manitoba,
Northern record for, 100

Badister obtusus, 147

Ball, P. W., 89

Ball, P.W. Hill’s Oak (Quercus ellipsoidalis) in southern
Ontario, 281

Banim, Farrell E., O.M.I., 1902-1979, 102

Barbour, S. E., 359

Barkworth, M. E., review by, 117

Bartramia longicauda, 408

Bass, Largemouth, Micropterus salmoides, in an oligotro-
phic Precambrian Shield lake, Reproduction, distri-
bution, and population size of, 298

Bass, Rock, 274
Smallmouth, 274

Bat, Little Brown, 206

Beauchamp, C., 307

Beaver, R. D. and V. Lewin. Scheduling censuses of breed-
ing White Pelicans (Pelecanus erythrorhynchos) in
northern Alberta, 198

Bechard, M. J. Historical nest records for the Ferruginous
Hawk in Manitoba, 467

Bedford, R. E., 231, 403

Behavior, haulout, of radio-tagged Harbor Seals, Phoca
vitulina, Movements and, 292

502

1981

Behavior, social, of some migrant shorebirds in southern
Manitoba, Feeding and, 183

Bembidion, 147

Bergeron, J.-M. et P. Demers. Le régime alimentaire du
Coyote (Canis latrans) et du chien errant (C. familia-
ris) dans le sud du Québec, 172

Betula papyrifera, 323
spp., 53

Beyersbergen, G. W., 471

Bidens sp., 247

Bird, D. M., reviews by, 218, 487

Bittern, American, 64, 423

Bjorge, R. R., J. R. Gunson, and W. M. Samuel. Population
characteristics and movements of Striped Skunks
(Mephitis mephitis) in central Alberta, 149

Blackbird, Red-winged, 63, 346, 407
Rusty, 310, 424

Blackbirds, Brewer’s, on man-made structures and natural
sites in British Columbia, Nesting of, 476

Blarina brevicauda, 254

Blueberry, Velvet-leaf, The biological flora of Canada, 2.
Vaccinium myrtilloides Michx., 329

Boag, D. A., 35, 75

Bobolink, 408

Bog lemming, 254

Boles, B. K., 366

Bonasa umbellus, 423

Boreogadus saida, Arctic Cod, in the Bering, Chukchi, and
Beaufort seas, Distribution, growth, and foods of,
186

Bos taurus, 172

Botaurus lentiginosus, 64, 423

Bouteloua gracilis, 412

Bovin domestique, 175

Boxall, P. C. Ruby-crowned Kinglets (Regulus calendula)
feeding a Brown-headed Cowbird (Molothrus ater),
99

Branta canadensis, 423
c. hutchinsii, 276
c. interior, 276
maxima, 276

Branta canadensis, Canada Geese, in southern Manitoba
prior to development of Oak Hammock Marsh, Dis-
tribution and harvest of, 276

Brasenia schreberi, 246

Breeding, feeding, and chick growth of the Black Guillemot
(Cepphus grylle) in southern Quebec, 312

Breeding Mallards (Anas platyrhynchos) in Manitoba, A
time-activity budget for, 266

Breeding of Mountain Plovers (Charadrius montanus) in
Canada, Status and, 133

Breeding success in a North Dakota marsh, Observations on
Black-crowned Night Heron, 465

Breeding success of New Brunswick Bald Eagles, Status and,
428

Breeding White Pelicans (Pelecanus erythrorhynchos) in
Northern Alberta, Scheduling censuses of, 198

British Columbia, 99, 348, 473

British Columbia, First records of the Tundra Shrew (Sorex
tundrensis) in, 93

INDEX TO VOLUME 95

503

British Columbia, Nesting of Brewer’s Blackbirds on man-
made and natural sites in, 476

British Columbia, Nesting of Smith’s Longspurs in, 469

Britton, D. M., review by, 119

Brodo, I. M. 100 years of natural history in Canada. Intro-
duction, |

Brooks, R. J., 350

Brunton, D. F. and P. D. Pratt. Panicled Knapweed (Cen-
taurea paniculata: Compositae) new to eastern Can-
ada, 98

Brunton, D. F., reviews by, 378, 484

Buchloe dactyloides, 412

Bufo americanus hemiophrys, Canadian Toad, Attempted
avian predation by a, 346

Bullhead, Brown, 137, 274

Bunting, Lark, 407

Buteo regalis, 408, 467

Butler, R. W. Nesting of Brewer’s Blackbirds on man-made
structures and natural sites in British Columbia, 476

Butterfly, Hop Merchant, (Polygonia comma), on insular
Newfoundland, Sightings of the, 361

Cairns, D. Breeding, feeding, and chick growth of the Black
Guillemot (Cepphus grylle) in southern Quebec, 312

Calamagrostis canadensis, 244

Calamospiza melanocorys, 408

Calanus cristatus, 190
glacialis, 190
hyperboreus, 190

Calathus ingratus, 147

Calcarius lapponicus, 425
mcecownii, 409
ornatus, 409
pictus, 425, 469

Caldwell, P. J., 183

Calidris minutilla, 183, 423

Calosoma frigidum, 147

Campagnol des champs, 175

Canachites canadensis, 423

Canada in the past 100 years, Federal wildlife conservation
work in, 31

Canada, Mammalogy in: a historical overview, 6

Canada, 100 years of natural history in, |

Canada, Status and breeding of Mountain Plovers (Charaa-
rius montanus) in, 133

Canada, The biological flora of
2. Vaccinium myrtilloides Michx., Velvet-leaf Blue-
berry, 329
3. Vaccinium vitis-idaea L. var. minus Lodd. sup-
plementary account, 434

Canada, 200 years of ornithology in, 2

Canadian entomology in the last century, 18

Canis familiaris, chien errant, dans le sud du Québec, Le
régime alimentaire du Coyote (Canis latrans) et du,
172

Canis latrans, Coyote, et duchienerrant (C. familiaris) dans
le sud du Québec, Le régime alimentaire du, 172

Capelin, 212, 314

Capella gallinago, 64, 311, 423

Carabus maeander, 147

$04 THE CANADIAN FIELD-NATURALIST

Carbyn, L. review by, 113

Carex echinata, 247
eleocharis, 412
filifolia, 412
lasiocarpa, 247
michauxiana, 247
oligosperma, 247
pensylvanica, 412
rostrata, 247
sp., 323
stricta, 247
vesicaria, 247

Carex loliacea, The Sedge, in eastern North America, 89

Caribou, Barren-ground, (Rangifer tarandus), Twinning
and postpartum activity in, 354

Caribou, North American Barren-ground, ( Rangifer taran-
dus arcticus), Application of the Varri6 Snow Index
to overwintering, 363

Caribou (Rangifer tarandus) encounters with pipelines in
northern Alaska, 57

Carp, 274

Carychium exile, 77

Catharus fuscescens, 311
guttatus, 311, 423
minimus, 423
ustualtus, 423

Catoptrophorus semipalmatus, 408

Catostomus commersoni, 137, 272

Cavers, P. B., review by, 491

Centauria paniculata: Compositae, Panicled Knapweed,
new to eastern Canada, 98

Cepaea nemoralis, land snail, at London, Ontario: changes
over three decades, Polymorphism in colonies of the,
192

Cephenemyia trompe, 59

Cepphus grylle, Black Guillemot, in southern Quebec,
Breeding, feeding, and chick growth of the, 312

Cerf de Virginie, 172

Certhia familiaris, 423

Chabot Visqueux, 203

Chamaedaphne calyculata, 247

Chaoborus, 129

Charadrius montanus, Mountain Plovers, in Canada, Status
and breeding of, 133

Charadrius vociferus, 64, 408

Char, Arctic, 203, 359

Chelydra serpentina, Common Snapping Turtle, in Algon-
quin Park, Ontario, Fate of overwintered clutches of
the, 350

Chickadee, Boreal, 423

Chien errant (C. familiaris) dans le sud du Québec, Le régime
alimentaire du Coyote (Canis latrans) et du, 172

Chipmunk, Least, 253

Chlaenius pensylvanicus, 147

Chordeiles minor, 408

Chrosomus eos, 138, 274

Chub, Creek, 138, 272
Lake, 212

Cionella lubrica, 76

Vol. 95

Circus cyaneus, 408

Cisco, Bering, (Coregonus laurettae) from the Yukon River
at Dawson, Yukon Territory, First Canadian record
of, 365

Cladina spp., 325

Cladium mariscoides, 244

Cladonia spp., 325

Clethrionomys gapperi, 146, 254

Clethrionomys gapperi, Southern Red-backed Voles, in
northern Ontario, Food habits of, 325

Cochran, P. A. An unusually small American Eel (Anguilla
rostrata) from the Lake Superior drainage, 97

Cod, Arctic, (Boreogadus saida) in the Bering, Chukchi, and
Beaufort seas, Distribution, growth, and foods of,
186

Cody, W. J., reviews by, 119, 382, 497

Coker, G. A., 210

Colaptes auratus, 423

Colgan, P., review by, 221

Colorado and Missouri, Low DDT residues in plasma of
Bald Eagles (Haliaeetus leucocephalus) wintering in,
249

Columella simplex, 77

Comanara livida, 323

Conservation strategy workshop, Canadian wolf status and,
216

Conservation work, Federal wildlife, in Canada in the past
100 years, 31

Contopus virens, 311

Cook, F. R. and J. C. Cook. Attempted avian predation bya
Canadian Toad, Bufo americanus hemiophrys, 346

Cook, J. C., 346

Cooke, F. 200 years of ornithology in Canada, 2

Coregonus laurettae, Bering Cisco, from the Yukon River at
Dawson, Yukon Territory, First Canadian record of,
365

Cormorant, Double-crested, 257

Cormorants in Montana, Nest-tree sharing by herons and,
257

Corvus caurinus, 473
corax, 423

Corydalis sempervirens, 322

Corylus cornuta, 53

Cottus bairdi, 274
cognatus, 203

Couesius plumbeus, 212

Cougar, 73

Cowbird, Brown-headed, 408

Cowbird, Brown-headed (Molothrus ater), Ruby-crowned
Kinglets (Regulus calendula) feeding a, 99

Cowling, J., 85

Coyote (Canis latrans) et du chien errant (C. familiaris) dans
le sud du Québec, Le régime alimentaire du, 172

Creeper, Brown, 423

Créte, M. et P. A. Jordan. Régime alimentaire des Orginaux
du sud-ouest Québécois pour les mois d’avril a octo-
bre, 50

Cromartie, E., 249

Crossbill, White-winged, 424

1981

Crows, Northwestern, Attempted predation of juvenile Star-
lings by, 473

Cryptacanthodes maculatus, 314

Culaea inconstans, 137, 274

Culaea inconstans, in Redwater drainage, Alberta, Varia-
tions in frequencies of pelvic phenotypes of the Brook
Stickleback, 178

Curlew, Long-billed, 407

Cutright, N. J., review by, 485

Cyminidis cribicollis, 147

Cyprinus carpio, 274

Dace, Blacknose, 274
Longnose, 274
Northern Redbelly, 212
Pearl, 138, 212, 274
Redbelly, 138, 274

Dagg, A. I., reviews by, 377, 385, 495

Darter, Fantail, 273
Iowa, 212
Johnny, 212, 274

Davie, J., reviews by, 123, 219

Day, R. T., 486

DDT residues, Low, in plasma of Bald Eagles (Haliaeetus
leucocephalus) wintering in Colorado and Missouri,
249

Deer, White-tailed, 172

Deer mortality in the Pohénégamook wintering area,
Quebec, 80

deGraff, D. A. First Canadian record of Bering Cisco (Core-
gonus laurettae) from the Yukon River at Dawson,
Yukon Territory, 365

deGraff, D. A., B. K. Boles, and J. A. Lovisek. Two-lined
Salamander, Eurycea bislineata (Amphibia: Cau-
data: Plethodontidae), in Labrador, 366

Demers, P., 172

Dendroica caerulescens, 311
castanea, 424
coronata, 424
magnolia, 424
palmarum, 424
pensylvanica, 307, 311, 424
petechia, 424
striata, 424
virens, 311, 424

Dickcissel, 408

Diervilla lonicera, 53

Discus cronkhitei, 76

Distribution and harvest of Canada Geese ( Branta canaden-
sis) in southern Manitoba prior to development of
Oak Hammock Marsh, 276

Distribution, growth, and foods of Arctic Cod (Boreogadus
saida) in the Bering, Chukchi, and Beaufort seas, 186

Distribution, Reproduction, and population size of Large-
mouth Bass, Micropterus salmoides, in an oligotro-
phic Precambrian Shield lake, 298

Dixon, C. C., 276

Dolichonyx oryzivorus, 408

Dove, Mourning, 408

INDEX TO VOLUME 95

505

Dowitcher, Short-billed, 423
Drosera intermedia, 244
Dryopteris spp., 53
Duchesneau, F., 80
Dulichium arundinaceum, 244

Eagles, Bald, (Haliaeetus leucocephalus) wintering in Colo-
rado and Missouri, Low DDT residues in plasma of,
249

Eagles, Bald, Status and breeding success of New Brunswick,
428

Editor’s farewell musings, 233

Editor’s report for 1980, 215

Editor’s thanks, 232

Edwards, Y., review by, 373

Eel, American, 359

Eel, American, (Anguilla rostrata) from the Lake Superior
drainage, An unusually small, 97

Eelpout, Polar, 314

Eleocharis olivacea, 244
palustris, 247

Eley, T. J., review by, 387

Empidonax alnorum, 311, 423
minimus, 307, 423

Environmental monitoring: a symposium, 482

Epigaea repens, 323

Eremophila alpestris, 408, 423

Erethizon dorsatum, 175

Eriocaulon septangulare, 53, 244

Esox lucius, 203

Etheostoma exile, 212
flabellare, 274
nigrum, 212, 274

Eualus fabricii, 190
gaimardii, 190

Euchaeta glacialis, 190

Euconulus fulvus, 76

Euphagus carolinus, 311, 424
cyanocephalus, 476

Eurycea bislineata, Two-lined Salamander, (Amphibia:
Caudata: Plethodontidae) in Labrador, 366

Eutamias minimus, 146, 253

Falco sparverius, 423

Fancy, S. G. Daily movements of Red Squirrels, Tamiasciu-
rus hudsonicus, 348

Feeding, and chick growth of the Black Guillemot (Cepphus
grylle) in southern Quebec, Breeding, 312

Feeding and social behavior of some migrant shorebirds in
southern Manitoba, 183

Felis concolor, 73

Ferguson, M., 272

Ferguson, M.A.D., review by, 124

Festuca octoflora, 412

Field research, Support for, 480

Fishes, fossil, from Coppermine River, Northwest Territo-
ries, Canada, Postglacial, 203

Fishes, teleost, Range extensions for 15, in the Hudson Bay
lowlands, Ontario, 212

506

Flicker, Common, 423

Flora of Canada, The biological, 2. Vaccinium myrtilloides
Michx., Velvet-leaf Blueberry, 329
3. Vaccinium vitis-idaea L. var. minus Lodd. Sup-
plementary account, 434

Flounder, 205

Flycatcher, Alder, 310, 423
Least, 307, 423
Olive-sided, 423

Food habits of Deer Mice (Peromyscus maniculatus) in
northern Ontario, 319

Food habits of Southern Red-backed Voles (Clethrionomys
gapperi) in northern Ontario, 325

Food in Manitoba prairie pothole lakes, Tiger Salamanders
(Ambystoma tigrinum) and stocked Rainbow Trout
(Salmo gairdneri): potential competitors for, 129

Foods of Arctic Cod (Boreogadus saida) in the Bering,
Chukchi, and Beaufort seas, Distribution, growth,
and, 186

Foxes, Red (Vulpes vulpes) on Baccalieu Island, Newfound-
land, Predation and caching of seabirds by, 352

Fragaria virginiana, 323, 327

Franzmann, A. W., 208

Fraser, J. M. Estimates of the standing stocks of fishes in
four small Precambrian Shield lakes, 137

Fratercula arctica, 352

Freedman, B., C. Beaucamp, I. A. McLaren, and S.I. Ting-
ley. Forestry management practices and populations
of breeding birds in a hardwood forest in Nova Sco-
tia, 307

Frost, K. J., 186

Gajadhar, A., 471

Gallus gallus, 175

Gammarus, 129

Gannet, Northern, 352

Gannets, Northern, (Morus bassanus) killed by rock falls at
Great Bird Rock, Quebec, Nesting, 202

Gasterosteus aculeatus, 359

Gastrocopta tappaniana, 77

Gavia immer, Common Loons, Marsh nesting of, 357

Gee, J. H., 129

Geese, Canada, (Branta canadensis) in southern Manitoba
prior to development of Oak Hammock Marsh, Dis-
tribution and harvest of, 276

Geothlypis trichas, 307, 408, 424

Geum triflorum, Three-flowered Avens, in Manitoba,
Northern record for, 100

Ginns, J., review by, 381

Glaucomys sabrinus, 254

Glomus spp., 322, 326

Glyceria canadensis, 247

Godwit, Marbled, 407

Goldeye, 212

Goose, Canada, 423

Grackle, Common, 64

Grand Brochet, 203

Gray, P. A., reviews by, 115, 121, 219, 374, 383

Grayling, Arctic, 203

THE CANADIAN FIELD-NATURALIST

Vol. 95

Grayling, Arctic, (Thymallus arcticus) spawning in a high-
way culvert, Evidence of, 358

Greenberg, D. A., review by, 225

Greenwood, R. J. Observations on Black-crowned Night
Heron breeding success in a North Dakota marsh,
465

Griffin, C. R., 249

Grosbeak, Pine, 423
Rose-breasted, 310

Grouse, Ruffed, 423
Sharp-tailed, 408
Spruce, 423

Grouse, Sharp-tailed, ( Pedioecetes phasianellus) in the park-
lands of Alberta, Seasonal and daily use of plant
communities by, 287

Growth, and condition, Invasion of a new reservoir by fishes:
species composition, 272

Growth, and foods of Arctic Cod (Boreogadus saida) in the
Bering, Chukchi, and Beaufort seas, Distribution,
186

Growth, chick, of the Black Guillemot (Cepphus grylle) in
southern Quebec, Breeding, feeding, and, 312

Growth patterns of the American Brook Lamprey, Lethen-
teron lamottenii, in the Ottawa River, Occurrence
and, 261

Grubby, 314

Guillemot, Black, (Cepphus grylle) in southern Quebec,
Breeding, feeding, and chick growth of the, 312

Gull, Bonaparte’s, 423
Great Black-backed, 205
Ring-billed, 183

Gull pair, Franklin’s X Ring-billed, nesting in Alberta, A
probable, 474

Gulls robbing prey from Great Blue Herons (Ardea hero-
dias), 205

Gunnel, Rock, 314

Gunson, J. R., 149

Gymnocanthus tricuspis, 314

Haliaeetus leucocephalus, 428

Haliaeetus leucocephalus, Bald Eagles, wintering in Colo-
rado and Missouri, Low DDT residues in plasma of,
249

Hall, I. V., 329

Hall, I. V. and J. M. Shaw. The biological flora of Canada,
3. Vaccinium vitis-idaea L. var. minus Lodd. Sup-
plementary account, 434

Hannon, S., 469

Hanson, J. M., 261

Hanson, W.C. Caribou (Rangifer tarandus) encounters
with pipelines in northern Alaska, 57

Hare, Snowshoe, 172

Harmata, A. R:, 249

Harms, V. L., review by, 489

Harpalus fulvilabus, 147
pleuriticus, 147

Hawk, Ferruginous, 407
Marsh, 408

1981

Hawk, Ferruginous, in Manitoba, Historical nest records for
the, 467

Hawk, Sharp-shinned, Accipiter striatus (Accipitriformes:
Accipitridae) on Victoria Island, Northwest Territo-
ries, 366

Henny, C. J., C. R. Griffin, D. W. Stahlecker, A. R. Har-
mata, and E. Cromartie. Low DDT residues in
plasma of Bald Eagles (Haliaeetus leucocephalus)
wintering in Colorado and Missouri, 249

Heron, Black-crowned Night, breeding success in a North
Dakota marsh, Observations on, 465

Heron, Great Blue, 257

Heron, Great Blue, (Ardea herodias) colony in the Peace-
Athabasca Delta, Alberta, 95

Herons and cormorants in Montana, Nest-tree sharing by,
257

Herons, Great Blue, (Ardea herodias), Gulls robbing prey
from, 205

Hiodon alosoides, 212
tergisus, 212

Hirundo rustica, 63

Hogsucker, Northern, 273

Holm, E. and G. A. Coker. First Canadian records of the
Ghost Shiner (Notropis buchanani) and the Orange-
spotted Sunfish (Lepomis humilus), 210

Hop Merchant (Polygonia comma) butterfly on insular
Newfoundland, Sightings of the, 361

Hoy, D., review by, 491

Hudson, R. J., 69

Hughes, S. J., review by, 222

Hummingbird, Ruby-throated, 310

Huot, J., 80

Hybognathus hankinsoni, 274

Hypentelium nigricans, 274

Hypericum boreale, 247
ellipticum, 247

Ictalurus nebulosus, 137, 274

Tlex verticillata, 247

Incubation, Analysis of weight lost by eggs of eleven species
of birds during, 63

Iridoprocne bicolor, 311, 423

Tris versicolor, 247

Jackson, B. S. Sightings of the Hop Merchant (Polygonia
comma) butterfly on insular Newfoundland, 361
James, P. C. Attempted predation of juvenile Starlings by
Northwestern Crows, 473
Jay, Gray, 423
Johnson, D. M., 156
Jones, D. M., 93
Jordan, P. A., 50
Junco, Dark-eyed, 309
Junco hyemalis, 311
Juncus brevicaudatus, 247
canadensis, 247
filiformis, 245
gerardii, 167
militaris, 241

INDEX TO VOLUME 95

507

pelocarpus, 244
Juncus compressus (Juncaceae) in North America, Distribu-
tional history of, 167

Kantrud, H. A. Grazing intensity effects on the breeding
avifauna of North Dakota native grasslands, 404

Keddy, P. A., review by, 494

Keddy, P. A. Vegetation with Atlantic coastal plain affinities
in Axe Lake, near Georgian Bay, Ontario, 241

Keith, L. B., 39

Kennedy, A., review by, 377

Kennedy, A. J., review by, 125

Kerekes, J. J., 359

Kestrel, American, 423

Killdeer, 64, 407

Kingfisher, Belted, 423

Kinglet, Golden-crowned, 423
Ruby-crowned, 424

Kinglets, Ruby-crowned, (Regulus calendula) feeding a
Brown-headed Cowbird (Molothrus ater), 99

Kirkham, I. R. Nesting Northern Gannets (Morus bassanus)
killed by rock falls at Great Bird Rock, Quebec, 202

Kittiwake, Black-legged, 352

Knapweed, Panicled, (Centaurea paniculata: Compositae)
new to eastern Canada, 98

Koeleria pyramidata, 412

Kratt, L. F. Evidence of Arctic Grayling (Thymallus arcti-
cus) Spawning in a highway culvert, 358

Kristensen, J. Great Blue Heron (Ardea herodias) colony in
the Peace-Athabasca Delta, Alberta, 95

Kurata, P. W. Farrell E. Banim, O.M.I., 1902-1979, 102

Lamprey, American Brook, Lethenteron lamottenii, in the
Ottawa River, Occurrence and growth patterns of
the, 261

Lance, American Sand, 314

Lanteigne, J., J. M. Hanson, and S. U. Qadri. Occurrence
and growth patterns of the American Brook Lam-
prey, Lethenteron lamottenii, in the Ottawa River,
261

Lark, Horned, 407, 423

Larus argentus, 205
delawarensis, 183, 474
marinus, 205
Philadelphia, 423
pipixcan, 474

Ledingham, G. F., review by, 116

Legget, R. F. Geology, 1879-1979, 24

Lemming, Bog, 254

Lepomis gibbosus, 137, 274

Lepomis humilis, Orangespotted Sunfish, First Canadian
records of the Ghost Shiner (Notropis buchanani)
and the, 210

Lepus americanus, 172

Lethenteron lamottenii, the American Brook Lamprey, in
the Ottawa River, Occurrence and growth patterns
of, 361

Lewin, V., 198

508 THE CANADIAN FIELD-NATURALIST

Liévre d’Ameérique, 172
Limosa fedoa, 408
Liopsetta putnami, 205
Lobelia dortmanna, 244
Lobipes lobatus, 423 ~
Longspur, Chestnut-collared, 407
Lapland, 425
McCown’s, 407
Smith’s, 425
Longspurs, Smith’s, in British Columbia, Nesting of, 469
Loons, Common (Gavia immer), Marsh nesting of, 357
Loricera pilicornis, 147
Love, D., reviews by, 376, 384, 493
Lovejoy, D. A., review by, 114
Lovisek, J. A., 366
Lowry, L. F. and K. J. Frost. Distribution, growth, and
foods of Arctic Cod ( Boreogadus saida) in the Bering,
Chukchi, and Beaufort seas, 186
Loxia leucoptera, 424
sp., 424
Lycodes turneri, 314
Lycopodium inundatum, 244
Lycopus uniflorus, 244
Lysimachia terrestris, 247

Macaulay, A. L., 319

Maccarone, A. D. and W. A. Montevecchi. Predation and
caching of seabirds by Red Foxes (Vulpes vulpes) on
Baccalieu Island, Newfoundland, 352

MacCrimmon, H. R., 298

MaclInnes, K. L., review by, 380

Macrozoarces americanus, 314

Mahon, R. and M. Ferguson. Invasion of a new reservoir by
fishes: species composition, growth, and condition,
272

Maine, Seasonal pelage change of Marten (Martes a. ameri-
cana) in, 356

Mallard, 423

Mallards (Anas platyrhynchos) in Manitoba, A time-activity
budget for breeding, 266

Mallotus villosus, 212, 314

Malus pumila, 175

Manitoba, 144, 346

Manitoba, A time-activity budget for breeding Mallards
(Anas platyrhynchos) in, 266

Manitoba, Feeding and social behavior of some migrant
shorebirds in southern, 183

Manitoba, Historical nest records for the Ferruginous Hawk
in, 467

Manitoba, northeastern, Relative abundances of birds in
boreal and subarctic habitats of northwestern Onta-
rio and, 418

Manitoba, Northern record for Three-flowered Avens
(Geum triflorum) in, 100

Manitoba prairie pothole lakes, Tiger Salamanders (Ambys-
toma tigrinum) and stocked Rainbow Trout (Salmo
gairdneri): potential competitors for food in, 129

Manitoba prior to development of Oak Hammock Marsh,
Distribution and harvest of Canada Geese (Branta

Vol. 95

canadensis) in southern, 276

Manning, T. H. Analysis of weight lost by eggs of eleven
species of birds during incubation, 63

Maraldo, D. C. and H. R. MacCrimmon. Reproduction,
distribution, and population size of Largemouth
Bass, Micropterus salmoides, in an oligotrophic Pre-
cambrian Shield lake, 298

Marmota monax, 172

Marmotte commune, 172

Martell, A. M., 354

Martell, A. M. and A. L. Macaulay. Food habits of Deer
Mice (Peromyscus maniculatus) in northern Ontario,
319

Martell, A. M. Food habits of Southern Red-backed Voles
(Clethrionomys gapperi) in northern Ontario, 325

Marten (Martes a. americana) in Maine, Seasonal pelage
change of, 356

Martes a. americana, Marten, in Maine, Seasonal pelage
change of, 356

Martin, K., S. Hannon, and R. Moses. Nesting of Smith’s
Longspur in British Columbia, 469

McAllister, D. C., 292

McAllister, D. E. and Denis St-Onge. Postglacial fossil
fishes from Coppermine River, Northwest Territo-
ries, Canada, 203

McAllister, D. E., reviews by, 120, 220, 375, 486

McCorquodale, D., reviews by, 110, 111

McDonald, E. J.and A. M. Martell. Twinning and postpar-
tum activity in Barren-ground Caribou (Rangifer
tarandus), 354

McLaren, I. A., 307

McLaren, M. A. and P. L. McLaren. Relative abundances
of birds in boreal and subarctic habitats of northwest-
ern Ontario and northeastern Manitoba, 418

McLaren, P. L., 418

McNicholl, M. K. John M. Powell — Honoured, 368

McNicholl, M. K., reviews by, 218, 372

Meadowlark, Western, 407

Megaceryle alcyon, 423

Melospiza lincolnii, 425
melodia, 311, 425

Mephitis mephitis, Striped Skunks, in central Alberta, Pop-
ulation characteristics and movements of, 149

Metabletus americanus, 147

Metridia longa, 190

Mice, Deer, (Peromyscus maniculatus) in northern Ontario,
Food habits of, 319

Michigan, upper, conifer swamps, Changes in small mam-
mal populations following clear-cutting in, 253

Microgadus tomcod, 314

Micropalma himantopus, 423

Micropterus dolomieui, 274

Micropterus salmoides, Largemouth Bass, in an oligotro-
phic Precambrian Shield lake, Reproduction, distri-
bution, and population size of, 298

Microsorex hoyi, 146

Microtus pennsylvanicus, 146, 175, 253

Miller, B. N., 205

Minnesota, 97

1981

Minnow, Bluntnose, 272
Brassy, 274
Fathead, 138, 212, 274

Missouri, Low DDT residues in plasma of Bald Eagles
(Haliaeetus leucocephalus) wintering in Colorado
and, 249

Mite, bat, Spinturnix globosus and a new host, Myotis luci-
fugus, First record for Canada of the, 206

Mniotilta varia, 311, 424

Molluscs, Terrestrial, of central Alberta, 75

Molothrus ater, 408

Molothrus ater, Brown-headed Cowbird, Ruby-crowned
Kinglets (Regulus calendula) feeding a, 99

Montana, Nest-tree sharing by herons and cormorants in,
257

Montevecchi, W. A., 352

Mooneye, 212

Moose, 50

Moose, Alces alces, Estimating winter defecation rates for,
208

Moose browse utilization in Mount McKinley National
Park, Alaska, 85

Moose populations near Rochester, Alberta, 1975-1978,
Dynamics of, 39

Morgantini, L. E. and R. J. Hudson. Sex differential in use
of the physical environment by Bighorn Sheep (Ovis
canadensis), 69

Morus bassanus, 352

Morus bassanus, Northern Gannets, killed by rock falls at
Great Bird Rock, Quebec, Nesting, 202

Moses, R., 469

Mouse, Deer, 146, 253
Meadow Jumping, 254
Woodland Jumping, 146, 254

Movements of Red Squirrels, Tamiasciurus hudsonicus,
Daily, 348

Movements of Striped Skunks (Mephitis mephitis) in central
Alberta, Population characteristics and, 149

Moyles, D. L. J. Seasonal and daily use of plant communi-
ties by Sharp-tailed Grouse (Pedioecetes phasianel-
lus) in the parklands of Alberta, 287

Mudminnow, 274

Muhlenbergia uniflora, 241

Munro, D. H., 100

Munro, H. L.and D. H. Munro. Northern record for Three-
flowered Avens (Geum triflorum) in Manitoba, 100

Murre, Common, 352

Myiasis by Wohlfahrtia opaca (Coq.): a cause of mortality of
newly hatched wild ducklings, 471

Myotis, Little Brown, 206

Myotis lucifugus, First record for Canada of the bat mite
Spinturnix globosus and a new host, 206

Myoxocephalus aeneus, 314
quadricornis, 212

Myrica gale, 53, 245

Myriophyllum tenellum, 244

Mysis litoralis, 190

Mytton, W. R.and L. B. Keith. Dynamics of Moose popula-
tions near Rochester, Alberta, 1975-1978, 39

INDEX TO VOLUME 95

509

Nagorsen, D. W. and D. M. Jones. First records of the Tun-
dra Shrew (Sorex tundrensis) in British Columbia, 93

Napaeozapus insignis, 255

Nemopanthus mucronata, 247

Neomysis rayii, 190

Nest records for the Ferruginous Hawk in Manitoba, Histor-

ical, 467

Nesting in Alberta, A probable Franklin’s X Ring-billed Gull

pair, 474

Nesting Northern Gannets (Morus bassanus) killed by rock

falls at Great Bird Rock, Quebec, 202

Nesting of Brewer’s Blackbirds on man-made structures and

natural sites in British Columbia, 476

Nesting of Common Loons (Gavia immer), Marsh, 357

Nesting of Smith’s Longspurs in British Columbia, 469

New Brunswick Bald Eagles, Status and breeding success of,

428

Newfoundland, 366

Newfoundland, Freshwater fishes from northern, 359

Newfoundland, Predation and caching of seabirds by Red
Foxes (Vulpes vulpes) on Baccalieu Island, 352

Newfoundland, Sightings of the Hop Merchant (Polygonia
comma) butterfly on insular, 361

Nighthawk, Common, 408

North Dakota marsh, Observations on Black-crowned Night
Heron breeding success in a, 465

North Dakota, native grasslands, Grazing intensity effects
on the breeding avifauna of, 404

Northwest Territories, Canada, Postglacial fossil fishes from
Coppermine River, 203

Northwest Territories, Sharp-shinned Hawk, Accipiter stria-
tus (Accipitriformes: Accipitridae) on Victoria
Island, 366

Notemigonus crysoleucas, 137

Notiophilus semistriatus, 147

Notropis atherinoides, 212
cornutus, 212, 272
heterolepis, 212
hudsonius, 212

Notropis buchanani, Ghost Shiner, and the Orangespotted
Sunfish (Lepomis humilis), First Canadian records of
the, 210

Nova Scotia, Forestry management practices and popula-
tions of breeding birds in a hardwood forest in, 307

Numenius americanus, 408
Phaeopus, 423

Nuphar variegatum, 246

Nuthatch, Red-breasted, 423

Nuttallornis borealis, 423

Nycticorax nycticorax, 465

Nymphaea odorata, 246

Nymphoides cordata, 241

Oak, Hill’s, (Quercus ellipsoidalis) in southern Ontario, 281

Obbard, M.E. and R. J. Brooks. Fate of overwintered
clutches of the Common Snapping Turtle (Chelydra
serpentina) in Algonquin Park, Ontario, 350

Oceanodroma leucorhoa, 352

Odocoileus virginianus, 80, 172

510

Oedemagena tarandi, 59

Oil spills, Experimental, off Baffin Island approved, 106

Oldemeyer, J. L. and A. W. Franzmann. Estimating winter
defecation rates for Moose, Alces alces, 208

Olenick, R. J. and J. H. Gee. Tiger Salamanders (Ambys-
toma tigrinum) and stocked Rainbow Trout (Salmo
gairdneri): potential competitors for food in Manit-
oba prairie pothole lakes, 129

Omble Chevalier, 203

Ombre Arctique, 203

Ontario, 63, 89, 98, 210, 272, 298, 357

Ontario, Fate of overwintered clutches of the Common
Snapping Turtle (Chelydra serpentina) in Algonquin
Park, 350

Ontario Field Biologist, The, 217

Ontario, Food habits of Deer Mice (Peromyscus manicula-
tus) in northern Ontario, 319

Ontario, Food habits of Southern Red-backed Voles (Cle-
thrionomys gapperi) in northern, 325

Ontario, Hill’s Oak (Quercus ellipsoidalis) in southern, 281

Ontario, northwestern, and northeastern Manitoba, Rela-
tive abundances of birds in boreal and subarctic habi-
tats of, 418

Ontario, Polymorphism in colonies of the land snail Cepaea
nemoralis at London,: changes over three decades,
192

Ontario, Range extensions for 15 teleost fishes in the Hudson
Bay lowlands, 212

Ontario, Vegetation with Atlantic coastal plain affinities in
Axe Lake, near Georgian Bay, 241

Oporornis philadelphia, 311

Orignaux du sud-ouest Québécois pour les mois d’avril a
octobre, Régime alimentaire des, 50

Osmerus mordax, 359

Ottawa Field-Naturalists’ Club
Auditor’s report, 397
Balance sheet, 398
Minutes of the one hundred and first annual business
meeting, 392
Report of Council, 393
Statement of Centennial project expenditures, 400
Statement of income and expenditure, C.F.-N., 401
Statement of income and expenditure, O.F.-N.C.,
399

Ottawa Field-Naturalists’ Club, Amendment to By-laws of
The, 478

Ottawa Field-Naturalists’ Club Annual Business Meeting,
Notice of The, 370

Ottawa Field-Naturalists’ Club, Call for nominations for the
Council of The, 370

Ottawa Field-Naturalists’ Club Centennial Symposium, 100
years of natural history in Canada, The: Introduc-
tion, |
200 years of ornithology in Canada, 2
Mammalogy in Canada: a historical overview, 6
Canadian entomology of the last century, 18
Geology, 1879-1979, 24
Federal wildlife conservation work in Canada in the
past 100 years, 31

THE CANADIAN FIELD-NATURALIST

Vol. 95

Ottawa Field-Naturalists’ Club, Changes in By-laws, 478

Ottawa Field-Naturalists’ Club, New honorary members of
The, 478

Ottawa Field-Naturalists’ Club, Notice of motions to amend
the Constitution of The, 369

Ovenbird, 307

Ovis canadensis, Bighorn Sheep, Sex differential in use of the
physical environment by, 69

Owl, Burrowing, 408
Short-eared, 423

Owls, Snowy, Request for information — marked, 368

Ozoga, J. J., 253

Pacific Science Congress, X Vth, 1983, 482

Panicum lanuginosum, 247

Parathemisto libellula, 190

Parula americana, 311

Parus hudsonicus, 423

Passerculus sandwichensis, 64, 408

Passerella iliaca, 352, 424

Patrobus foveocollis, 147

Pearce, P. A., 428

Pedioecetes phasianellus, 408

Pedioecetes phasianellus, Sharp-tailed Grouse, in the park-
lands of Alberta, Seasonal and daily use of plant
communities by, 287

Pelecanus erythrorhynchos, White Pelicans, in northern
Alberta, Scheduling censuses of breeding, 198

Pelicans, White, (Pelecanus erythrorhynchos) in northern
Alberta, Scheduling censuses of breeding, 198

Perca flavescens, 212

Perch, Yellow, 212

Perisoreus canadensis, 423

Peromyscus maniculatus, 146, 175, 253

Peromyscus maniculatus, Deer Mice, in northern Ontario,
Food habits of, 319

Petrel, Leach’s Storm, 352

Pewee, Eastern Food, 310

Pfannmuller, L. A., 144

Phalacrocorax auritus, 257

Phalarope, Northern, 423
Wilson’s, 64, 408

Pheucticus ludovicianus, 311

Phipps, J. B., review by, 223

Phoca vitulina, Harbor Seals, Movements and haulout
behavior of radio-tagged, 292

Phoebe, Eastern, 63

Pholis gunnellus, 314

Phoxinus eos, 212

Picoides arcticus, 423

Pike, Northern, 203

Pimephales notatus, 272
promelas, 138, 212, 274

Pinicola enucleator, 423

Pipit, Sprague’s, 407

Pitcher, K. W. and D. C. McAllister. Movements and hau-
lout behavior of radio-tagged Harbor Seals, Phoca
vitulina, 292

Plantago patagonica, 412

1981

Plover, American Golden, 183
Black-bellied, 183

Plovers, Mountain, (Charadrius montanus) in Canada, Sta-
tus and breeding of, 133

Pluvialis dominica, 183
squatarola, 183

Poa pratensis, 412

Polygonia comma, Hop Merchant, butterfly on insular New-
foundland, Sightings of the, 361

Polymorphism in colonies of the land snail Cepaea nemora-
lis at London, Ontario: changes over three decades,
192

Pontederia cordata, 244

Pooecetes gramineus, 409

Population characteristics and movements of Striped
Skunks (Mephitis mephitis) in central Alberta, 149

Population size of Largemouth Bass, Micropterus sal-
moides, in an oligotrophic Precambrian Shield lake,
Reproduction, distribution, and, 298

Populations, Changes in small mammal, following clear-
cutting in upper Michigan conifer swamps, 253

Populations, Moose, near Rochester, Alberta, 1975-1978,
Dynamics of, 39

Populations of breeding birds in a hardwood forest in Nova
Scotia, Forestry management practices and, 307

Populus balsamifera, 86
tremuloides, 53

Porc domestique, 175

Porc-épic, 175

Porzana carolina, 63

Potamogeton confervoides, 244
epihydrus, 247
natans, 247
oakesianus, 244

Potentilla (norvegica?), 323

Potvin, F., J. Huot, and F. Duchesneau. Deer mortality in
the Pohénégamook wintering area, Quebec, 80

Poulet domestique, 175

Pout, Ocean, 314

Powell, John M. — Honoured, 368

Prairie Wildlife Interpretation Centre, 106

Pratt, P. D., 98

Predation and caching of seabirds by Red Foxes (Vulpes
vulpes) on Baccalieu Island, Newfoundland, 352

Predation, avian, by a Canadian Toad, Bufo americanus
hemiophrys, Attempted, 346

Predation of juvenile Starlings by Northwestern Crows,
Attempted, 473

Preston, W. B., reviews by, 108, 112, 113

Procyon lotor, 172

Pruitt, W. O., Jr. Application of the Varrio Snow Index to
overwintering North American Barren-ground Cari-
bou (Rangifer tarandus arcticus), 363

Prunus pensylvanica, 54
spp., 50

Pseudoma truncata, 190

Psoralea argophylla, 412

Pterostichus adstrictus, 147

Puffin, Atlantic, 352

INDEX TG VOLUME 95

511

Pumpkinseed, 137, 274
Punctum minutissimum, 77
Pungitius pungitius, 359
Pyrus arbutifolia, 247

Qadri, S. U., 261

Quebec, 50

Quebec, Breeding, feeding, and chick growth of the Black
Guillemot (Cepphus grylle) in southern, 312

Quebec, Deer mortality in the Pohénégamook wintering
area, 80

Québec, Le régime alimentaire du Coyote (Canis latrans) et
du chien errant (C. familiaris) dans le sud du, 172

Quebec, Nesting Northern Gannets (Morus bassanus) killed
by rock falls at Great Bird Rock, 202

Québécois pour le mois d’avril a octobre, Régime alimentaire
des Orignaux du sud-ouest, 50

Quercus ellipsoidalis, Hill’s Oak, in southern Ontario, 281

Quercus ellipsoidalis X velutina, 283

Quinney, K. R. S., 205

Quinney, T. E., B. N. Miller, and K. R. S. Quinney. Gulls
robbing prey from Great Blue Herons (Ardea hero-
dias), 205

Quiscalus quiscula, 64

Raccoon, 172

Range extensions for 15 teleost fishes in the Hudson Bay
lowlands, Ontario, 212

Rangifer tarandus arcticus, North American Barren-ground
Caribou, Application of the Varrid Snow Index to
overwintering, 363

Rangifer tarandus, Barren-ground Caribou, Twinning and
postpartum activity in, 354

Rangifer tarandus, Caribou, encounters with pipelines in
northern Alaska, 57

Raton laveur, 172

Raveling, D. G. and C. C. Dixon. Distribution and harvest
of Canada Geese (Branta canadensis) in southern
Manitoba prior to development of Oak Hammock
Marsh, 276

Raven, Common, 423

Redstart, American, 309

Regulus calendula, 424
satrapa, 423

Regulus calendula, Ruby-crowned Kinglets, feeding a
Brown-headed Cowbird (Molothrus ater),99 _

Reist, J. D. Variation in frequencies of pelvic phenotypes of
the Brook Stickleback, Culaea inconstans, in Red-
water drainage, Alberta, 178

Report, Editor’s, for 1980, 215

Report on new titles of books, 368

Reports of significant range extensions, 212, 366

Reproduction, distribution, and population size of Large-
mouth Bass, Micropterus salmoides, in an oligotro-
phic Precambrian Shield lake, 298

Request for information:
marked Snowy Owls, 368
Common Tern colour marking by the Canadian
Wildlife service, 481

512

Restoration and Management Notes, 369

Retinella electrina, 76

Reznicek, A. A. and P. W. Ball. The Sedge Carex /oliacea in
eastern North America, 89

Rhexia virginica, 241

Rhinichthys atratulus, 274
cataractae, 274

Rhynchospora capitellata, 241
fusca, 244

Ribes lacustre, 323

Rissa tridactyla, 352

Robertson, A., review by, 118

Robin, American, 63, 310

Rombough, P. J.,S. E. Barbour, and J. J. Kerekes. Fresh-
water fishes from northern Newfoundland, 358

Rosa acicularis, 322, 327

Ross, P. E., review by, 386

Rowsell, J. A., review by, 488

Rubus hispidus, 247
pubescens, 327
strigosus, 323

Ryder, J. P. and D. A. Boag. A Canadian paradox — pri-
vate land, public wildlife: can it be resolved? (view-
point), 35

Salamander, Two-lined, Eurycea bislineata (Amphibia:
Caudata: Plethodontidae), in Labrador, 366
Salamanders, Tiger, (Ambystoma tigrinum) and stocked

Rainbow Trout (Salmo gairdneri): potential competi-
tors for food in Manitoba prairie pothole lakes, 129
Salix arbusculoides, 86
alaxensis, 85
glauca, 86
spp., 53
Salmo gairdneri, Rainbow Trout, potential competitors for
food in Manitoba prairie pothole lakes, Tiger Sala-
manders (Ambystoma tigrinum) and stocked, 129
Salmo salar, 359
Salmon, Atlantic, 359
Sandpiper, Least, 183
Upland, 408
Salvelinus alpinus, 203, 359
fontinalis, 137, 359
Samuel, W. M., 149
Sandpiper, Least, 423
Solitary, 423
Stilt, 423
Sapsucker, Yellow-bellied, 423
Saskatchewan, 363, 471
Sayornis phoebe, 64
Schueler, F. W., review by, 488
Scirpus subterminalis, 245
torreyi, 244
Sculpin, Arctic Staghorn, 314
Four-horned, 212
Mailed, 314
Mottled, 274
Slimy, 203

THE CANADIAN FIELD-NATURALIST

Vol. 95

Seals, Harbor, Phoca vitulina, Movements and haulout
behavior of radio-tagged, 292

Sealy, S. G., 183

Sedge, Carex loliacea in eastern North America, The, 89

Seiurus aurocapillus, 307
noveboracensis, 311

Selaginella densa, 412

Semotilus atromaculatus, 138, 272
margarita, 138, 212, 274

Setophaga ruticilla, 311

Shanny, Arctic, 314

Shaw, J. M., 434

Sheep, Bighorn, (Ovis canadensis), Sex differential in use of
the physical environment by, 69

Shiner, Blacknose, 212
Common, 212, 272
Emerald, 212
Golden, 137
Rosyface, 274
Spottail, 212

Shiner, Ghost, (Notropis buchanani) and the Orangespotted
Sunfish (Lepomis humilis), First Canadian records of
the, 210

Shorebirds in southern Manitoba, Feeding and social behav-
ior of some migrant, 183

Shoveler, Northern, 471

Shrew, Arctic, 254
Dusky, 146
Masked, 146, 253
Pygmy, 146
Short-tailed, 254
Water, 255

Shrew, Tundra, (Sorex tundrensis) in British Columbia,
First records of the, 93

Shrike, Northern, 424

Singh, S. M. Polymorphism in colonies of the land snail
Cepaea nemoralis at London, Ontario: changes over
three decades, 192

Sitta canadensis, 423

Skunks, Striped, (Mephitis mephitis) in central Alberta,
Population characteristics and movements of, 149

Smelt, Rainbow, 359

Smith, D. A. Mammalogy in Canada: a historical overview,
6 ‘

Smith, H. C. First record for Canada of the bat mite Spin-
turnix globosus and a new host, Myotis lucifugus, 206

Smith, L. C. Editor’s farewell musings, 233

Smith, L. C. Editor’s report for 1980, 215

Smith, L. C. Editor’s thanks, 232

Smith, T. G. Sharp-shinned Hawk, Accipiter striatus (Acci-
pitriformes: Accipitridae) on Victoria Island,
Northwest Territories, 366

Snail, land, Cepaea nemoralis at London, Ontario: changes
over three decades, Polymorphism in colonies of the,
192

Snipe, Common, 64, 310, 423

Solmon, V. E. F. Federal wildlife conservation work in
Canada in the past 100 years, 31

1981

Sora, 63
Sorex arcticus, 254
cinereus, 146, 253
monticolus, 146
palustris, 255
Sorex tundrensis, Tundra Shrew, in British Columbia, First
records of the, 93
Souris sylvestre, 175
Soutiere, E. C. and J. D. Steventon. Seasonal pelage change
of Marten (Martes a. americana) in Maine, 356
Sparrow, Baird’s, 407
Chipping, 424
Clay-colored, 407
Fox, 352, 424
Grasshopper, 408
Harris’, 424
LeConte’s, 407
Lincoln’s, 425
Savannah, 64, 407, 424
Song, 309, 425
Tree, 424
Vesper, 409
White-throated, 307, 424
Speirs, J. M., review by, 374
Sphaeroderus lecontei, 147
Spinturnix globosus, bat mite, and a new host, Myotis luci-
fugus, First record for Canada of the, 206
Spiraea alba, 247
Spiza americana, 408
Spizella arborea, 424
pallida, 409
Passerina, 424
Spleenwort, Ebony, Asplenium platyneuron (Aspleniaceae),
in the Great Lakes area, Natural history of the, 156
Squirrel, Northern Flying, 254
Red, 254
Squirrels, Red, Tamiasciurus hudsonicus, Daily movements
of, 348
Stahlecker, D. W., 249
Starlings, juvenile, by Northwestern Crows, Attempted pre-
dation of, 473
Steganopus tricolor, 64, 408
Steventon, J. D., 356
Stichaeus punctatus, 314
Stickleback, Brook, 137, 274
Fourspine, 359
Ninespine, 359
Threespine, 359
Stickleback, Brook, Culaea inconstans, in Redwater drain-
age, Alberta, Variation in frequencies of pelvic pheno-
types of the, 178
Stipa comata, 412
viridula, 412
Stocek, R. F. and P. A. Pearce. Status and breeding success
of New Brunswick Bald Eagles, 428
St-Onge, D., 203
Storm-Petrel, Leach’s, 352
Stuckey, R. L. Distributional history of Juncus compressus
(Juncaceae) in North America, 167

INDEX TO VOLUME 95

513

Sturnella neglecta, 408

Sturnus vulgaris, 473

Succinea sp., 77

Sucker, White, 137, 272

Sugden, L. G., 471

Sunfish, Orangespotted, (Lepomis humilis), First Canadian
records of the Ghost Shiner (Notropis buchanani)
and the, 210

Sus scrofa, 172

Swallow, Barn, 63
Tree, 310, 423

Symphoricarpos occidentalis, 412

Synaptomys cooperi, 254

Synuchus impunctatus, 147

Tamiasciurus hudsonicus, 254

Tamiasciurus hudsonicus, Red Squirrels, Daily movements
of, 348

Taxus canadensis, 50

Taylor, R. Is the Impact Factor a meaningful index for the
ranking of scientific research papers? 236

Taylor, R. and R. E. Bedford. Announcement, 403

Taylor, R. and R. E. Bedford, Editorship of The Canadian
Field- Naturalist, 231

Taylor, R. J.and L. A. Pfannmuller. A test of the peninsular
effect on species diversity, 144

Teal, Blue-winged, 423, 471
Green-winged, 423

Thompson, L. S. Nest-tree sharing by herons and cormor-
ants in Montana, 257

Thrush, Gray-cheeked, 423
Hermit, 309, 423
Swainson’s, 423

Thuja occidentalis, 53

Thymallus arcticus, 203

Thymallus arcticus, Arctic Grayling, spawning in a highway
culvert, Evidence of, 358

Tingley, S. I., 307

Titman, R. D. A time-activity budget for breeding Mallards
(Anas platyrynchos) in Manitoba, 266

Toad, Canadian, Bufo americanus hemiophrys, Attempted
avian predation by a, 346

Tomcod, Atlantic, 314

Tomlin, A. D., review by, 122

Triadenum fraseri, 245

Triglops murrayi, 314

Tringa flavipes, 423
melanoleuca, 423
solitaria, 423

Troglodytes troglodytes, 423

Trout, Brook, 137, 359

Trout, Rainbow, (Salmo gairdneri): potential competitors
for food in Manitoba prairie pothole lakes, Tiger
Salamander (Ambystoma tigrinum) and, 129

Turdus migratorius, 64, 311

Turnstone, Ruddy, 183

Turtle, Common Snapping, (Chelydra serpentina) in Algon-
quin Park, Ontario, Fate of overwintered clutches of
the, 350

514

Typha spp., 53

Umbra limi, 274

Uria aalge, 352

Utricularia cornuta, 244
gibba, 244
intermedia, 247
purpurea, 241
resupinata, 244
vulgaris, 247

Vaccinium angustifolium, 321, 327
macrocarpon, 247
myrtilloides, 323
m. forma chicoccum, 331
m. vat. integrifolium, 331
oxycoccus, 247

Vaccinium myrtilloides Michx., Velvet-leaf Blueberry, The
biological flora of Canada, 2, 329

Vaccinium vitis-idaea L. var. minus Lodd., Supplementary
account, The biological flora of Canada, 3, 434

Vallonia gracilicosta, 77
Span

Van Es, J. and D. A. Boag. Terrestrial molluscs of central
Alberta, 75

Vander Kloet, S. P. and I. V. Hall. The biological flora of
Canada, 2. Vaccinium myrtilloides Michx. Velvet-
leaf Blueberry, 329

Variation in frequencies of pelvic phenotypes of the Brook
Stickleback, Culaea inconstans, in Redwater drain-
age, Alberta, 178

Veery, 310

Vegetation with Atlantic coastal plain affinities in Axe Lake,
near Georgian Bay, Ontario, 241

Verme, L. J. and J. J. Ozoga. Changes in small mammal
populations following clearcutting in upper Michigan
conifer swamps, 253

Vermivora ruficapilla, 424

Vertigo gouldi, 76
modesta, 77

Viburnum alnifolium, 53
cassinoides, 53, 247

Viola lanceolata, 244

Vireo olivaceous, 311, 424
Dhiladelphicus, 424
solitarius, 311, 424

Vireo, Philadelphia, 424
Red-eyed, 309, 424
Solitary, 309, 424

Vitrina alaskana, 76

Vockeroth, J. R. Canadian entomology of the last century,
18

Vole, Meadow, 146, 253
Southern Red-backed, 146, 254

Voles, Southern Red-backed, (Clethrionomys gapperi) in
northern Ontario, Food habits of, 325

Vulpes vulpes, Red Foxes, on Baccalieu Island, Newfound-
land, Predation and caching of seabirds by, 352

THE CANADIAN FIELD-NATURALIST

Vol. 95

Wagner, W. H., Jr., and D. M. Johnson. Natural history of
the Ebony Spleenwort, Asplenium platyneuron
(Aspleniaceae), in the Great Lakes area, 156

Wallis, C. A. and C. R. Wershler. Status and breeding of
Mountain Plovers (Charadrius montanus) in Can-
ada, 133

Warbler, Bay-breasted, 424
Black-and-white, 309, 424
Blackpoll, 424
Black-throated Blue, 310
Black-throated Green, 309, 424
Canada, 424
Chestnut-sided, 307, 424
Magnolia, 424
Mourning, 310
Nashville, 424
Northern Parula, 309
Orange-crowned, 424
Palm, 424
Tennessee, 424
Wilson’s, 424
Yellow, 424
Yellow-rumped, 424

Waterthrush, Northern, 310

Weber, W. C. review by, 111

Wein, R. W., review by, 381

Weir, J. B., 212

Wershler, C. R., 133

Weseloh, D. V. A probable Franklin’s X Ring-billed Gull
pair nesting in Alberta, 474

Weseloh, D. V. C., review by, 109

Whimbrel, 423

Wildlife area, A new, — Long Point in Lake Erie, 216

Willet, 408

Wilson Ornithological Society annual meeting, The, 217

Wilsonia canadensis, 424
pusilla, 424

Wishart, R. A., P. J. Caldwell, and S. G. Sealy. Feeding and
social behavior of some migrant shorebirds in south-
ern Manitoba, 183

Wobeser, G., A. Gajadhar, G. W. Beyersbergen, and L. G.
Sugden. Myiasis by Wohlfahrtia opaca (Coq.): a
cause of mortality of newly hatched wild ducklings,
471

Wohlfahrtia opaca (Coq.): a cause of mortality of newly
hatched wild ducklings, Myiasis by, 471

Wolf status and conservation strategy workshop, Canadian,
216

Wolff, J. O. and J. Cowling. Moose browse utilization in
Mount McKinley National Park, Alaska, 85

Wood Buffalo National Park Management Planning Pro-
gram, 481

Woodchuck, 172

Woodpecker, Black-backed Three-toed, 423

Woodwardia virginica, 241

Wren, Short-billed Marsh, 407
Winter, 423

Wright, J., review by, 495

1981

Wrymouth, 314
Xyris caroliniana, 241

Yellowlegs, Greater, 423
Lesser, 423

Yellowthroat, Common, 307, 408, 424

Yukon Territory, 354, 358

Yukon Territory, First Canadian record of Bering Cisco
(Coregonus laurettae) from the Yukon River at Daw-
son, 356

INDEX TO VOLUME 95

515

Zalewski, B. R. and J. B. Weir. Range extensions for 15
teleost fishes in the Hudson Bay lowlands, Ontario,
212 -
Zammuto, R. M., reviews by, 224, 379, 384, 386, 484, 496
Zapus hudsonius, 146, 254
Zenaida macroura, 408
Zonitoides arboreus, 77
Zonotrichia albicollis, 307, 424
leucophrys, 424
querula, 424
Zo6genetes harpa, 77

Index to Book Reviews

Botany

Alex, J. F., R. Cayouette, and G. A. Mulligan. Common
and botanical names of weeds in Canada/Noms
populaires et scientifiques des plantes nuisibles du
Canada, 491

Alexsandrova, V. D. The Arcticand Antarctic: their division
into geobotanical areas, 381

Boivin, B. Flora of the Prairie Provinces: a handbook to the
flora of the provinces of Manitoba, Saskatchewan
and Alberta; Part I. — Pteroids, ferns, conifers and

woody dicopsids; Part II. — Digitatae, Dimerae,
Liberae; Part II]. — Connatae; Part IV. — Monop-
sida, 489

Clifford, H. T. and L. Watson. Identifying grasses: data,
methods and illustrations, 117

Cronquist, A. Vascular flora of the southeastern United
States: Volume I, Asteraceae, 119

Dhir, K. K. Ferns of north-western Himalayas, 119

Dore, W. G. and J. McNeill. Grasses of Ontario, 223

Harris, W. C. Guide to forest understory vegetation in Sas-
katchewan, 488

Kendrick, B. (ed.). The whole fungus: the sexual-asexual
synthesis, 222

Maher, R. V., G. W. Argus, V. L. Harms, and J. H. Hud-
son. The rare vascular plants of Saskatchewan, 116

Orloci, L. Multivariate analysis in vegetation research, 379

Pomerleau, R. Flore des champignons au Québec et régions
limitrophes, 381

Porsild, A. E. and W. J. Cody. Vascular plants of continen-
tal Northwest Territories, Canada, 380

Ryan, A. G. Native trees and shrubs of Newfoundland and
Labrador, 118

Schumann, D.N. Living with plants: a guide to practical
botany, 491

Scoggan, H. G. The flora of Canada, 378

Environment

Baker, R. (ed.). The mystery of migration, 384

Baxter, R. M. and P. Glaude. Environmental effects of dams
and impoundments in Canada: experience and pros-
pects, 386

Cloudsley-Thompson, J. L. Terrestrial environments, 383

Ehrlich, P. and A. Extinction: the causes and consequences
of the disappearance of species, 493

Environmental Council of Alberta. The environmental
effects of forestry operations in Alberta: report and
recommendations, 121

Griffiths, D. Island forest year: Elk Island National Park,
124

Laird, M. Bibliography of the natural history of Newfound-
land and Labrador, 382

Lewis, D. B. and D. M. Gower. Biology of communication,
385

Lohmn, U. and T. Persson (eds.). Soil organisms as compo-
nents of ecosystems, 122

Owen, O. S. Natural resource conservation: an ecological
approach, 494

Pielou, E. C. Biogeography, 384

Riedl, R. Order in living organisms: a systems analysis of
evolution, 120

Shuttlesworth, D. Exploring nature with your child, 123

Simons, T. J. Circulation models of lakes and inland seas,
225

Wilson, D. S. The natural selection of populations and
communities, 224

Zoology

Allen, D. L. Wolves of Minong — their vital role ina wildlife
community, 113

Balon, E. K. (ed.). Charrs: salmonid fishes of the genus
Salvelinus, 220

Bartonek, J. C. and D. N. Nettleship (eds.). Conservation of
marine birds of northern North America, 107

Bonner, J. T. The evolution of culture in animals, 221

Burk, D. (ed.). The Black Bear in North America, 374

Campbell, R. W., H. R. Carter, C. D. Shepard, and C. J.
Guiguet. A bibliography of British Columbia orni-
thology, Volume I, 218

Cannings, R. A.and K. M. Stuart. The dragonflies of British
Columbia, 112

Dagg, A. I. Camel quest, 219

DeBlase, A. F.and R. E. Martin. A manual of mammalogy;
with keys to families of the world, 484

Erskine, A. J. The first ten years of the Co-operative Breed-
ing Bird Survey in Canada, 111

516

Goodwin, C. E. A bird-finding guide to the Toronto region,
374

Griscom, L. and A. Sprunt, Jr. (eds.). The warblers of Amer-
ica, 485

Hall, R. L.and H. S. Sharp (eds.). Wolf and man: evolution
in parallel, 376

Hamilton, W. J., Jr.and J. O. Whitaker, Jr. Mammals of the
eastern United States, 114

Hancock, L. Love affair with a cougar, 219

Harris, J. T. The Peregrine Falcon in Greenland: observing
an endangered species, 110

Hatler, D. F., R. W. Campbell, and A. Dorst. Birds of
Pacific Rim National Park, 372

Kelton, L. A. The insects and arachnids of Canada. Part 4.
The Anthocoridae of Canada and Alaska (Heterop-
tera: Anthocoridae), 108

Legendre, V., J.-R. Leclerc, et J. Brisebois. Les salmonidés
des eaux de la Plaine de Montréal: 1. Historique,
1534-1922 et 2. Biometrie, biogeographie, 1970-1975,
et registre des péches 1941-1976, 486

Lister, R. The birds and birders of Beaverhills Lake, 109

Mansell, W. North American birds of prey, 376

Martin, J. E. H. The insects and arachnids of Canada. Part
1. Collecting, preparing and preserving insects, mites
and spiders, 108

Matsuda, R. The insects and arachnids of Canada. Part 3.
The Aradidae of Canada (Hemiptera: Aradidae), 108

Miller, F. L.and A. Gunn. Responses of Peary Caribou and
Muskoxen to helicopter harassment, 115

THE CANADIAN FIELD-N ATURALIST

Vol. 95

Morris, R. F. Butterflies and moths of Newfoundland and
Labrador: the Macrolepidoptera, 486

Morrow, J. E. The freshwater fishes of Alaska, 375

Morse, D. H. Behavioral mechanisms in ecology, 488

Murie, M. (comp. and ed.). The Alaskan bird sketches of
Olaus Murie with excerpts from his field notes, 373

Nero, R. W. The Great Gray Owl: phantom of the northern
forest, 484

Newton, I. Population ecology of raptors, 218

Orians, G. H. Some adaptations of marsh-nesting black-
birds, 111

Secretariat, Entomological Society of Canada. Annotated
list of workers on systematics and faunistics of Cana-
dian insects and certain related groups, 113

Weick, F. Birds of prey of the world, 487

Woods, S. E. The squirrels of Canada, 377

Miscellaneous

Darlington, P. J., Jr. Evolution for naturalists: the simple
principles and complex reality, 496

Freeman, R. B. British natural history books, 1495-1900: a
handlist, 497

Nelson, R. K. Shadow of the Hunter — stories of Eskimo
life, 387

Patterson, F. Photography and the art of seeing, 125

Pimentel, R. A. Morphometrics: the multivariate analysis of
biological data, 386

Smeeton, M. Completely foxed, 495

Ure, S. Hawk lady: the story of a woman who opened her
home to care for wild birds of prey, 495

Instructions to Contributors

Content

The Canadian Field- Naturalist is a medium for the publi-
cation of scientific papers by amateur and professional natur-
alists 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 sub-
mit for consideration their manuscripts meeting these crite-
ria. 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 significance.

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 sub-
mitted. Also authors are expected to have complied with all
pertinent legislation regarding the study, disturbance, or
collection of animals, plants or minerals. The place where
voucher specimens have been deposited, and their catalogue
numbers, should be given.

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 biblio-
graphic 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. Under-
line only words meant to appear in italics. The names of
authors of scientific names should be omitted except in taxo-
nomic manuscripts or other papers involving nomenclatural
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.

We prefer the names of journals in the Literature Cited to
be written out in full. 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 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 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
K1A 082. 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. Webs-
ter’s New International Dictionary and le Grand Larousse
Encyclopédique are the authorities for spelling.

Illustrations— Photographs should have a glossy finish and
show sharp contrasts. 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
$50 for each page in excess of six journal pages, plus $5 for
each illustration (any size up to a full page), and up to $50 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 acommitment, especially if grant or institutional funds
are available, to pay $50 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.

Reviewing Policy of The Canadian Field-Naturalist

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

comments and constructive recommendations. Almost all
manuscripts accepted for publication have undergone revi-
sion — 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)
from OBC

Environment: Extinction: the causes and consequences of the disappearance of species — Natural
resource conservation: an ecological approach

Miscellaneous: Hawk lady: the story of awoman who opened her home to care for wild birds of prey
— Completely foxed — Evolution for naturalists: the simple principles and complex reality
— British natural history books, 1495-1900: a handlist

New Titles

Index to Volume 95 Compiled by W. HARVEY BECK

Mailing date of previous issue 15 September 1981

493

495

497

502

THE CANADIAN FIELD-NATURALIST Volume 95, Number 4 1981

Announcement 403

Articles

Grazing intensity effects on the breeding avifauna of North Dakota native grasslands
Fy H. A. KANTRUD 404

Relative abundances of birds in boreal and subarctic habitats of northwestern Ontario
and northeastern Manitoba
MARGARET A. MCLAREN and PETER L. MCLAREN 418

Status and breeding success of New Brunswick Bald Eagles
R F. STOCEK and P. A. PEARCE 428
The Biological Flora of Canada

3. Vaccinium vitis-idaea L. var. minus Lodd. Supplementary account

IVAN V. HALL and JENNIFER M. SHAY 434
Notes
Observations on Black-crowned Night Heron breeding success in a North Dakota marsh
RAYMOND J. GREENWOOD 465
Historical nest records for the Ferruginous Hawk in Manitoba MARC J. BECHARD 467
Nesting of Smith’s Longspurs in British Columbia
KATHY MARTIN, SUSAN HANNON, and RICHARD MOSES 469
Myiasis by Wohlfahrtia opaca (Coq.): a cause of mortality of newly hatched wild ducklings
GARY WOBESER, ALVIN GAJADHAR, GERRY W. BEYERSBERGEN and LAWSON G. SUGDEN 47]
Attempted predation of juvenile Starlings by Northwestern Crows PAUL C. JAMES 473
A probable Franklin’s X Ring-billed Gull pair nesting in Alberta D. V. WESELOH 474
Nesting of Brewer’s Blackbirds on man-made structures and natural sites in
British Columbia ROBERT W. BUTLER 476
News and Comment 478

Book Reviews

Zoology: A manual of mammalogy: with keys to families of the world — The Great Grey Owl: 484
phantom of the northern forest — The warblers of America — Butterflies and moths of
Newfoundland and Labrador: the Macrolepidoptera — Les salmonidés des eau de la Plaine
de Montréal: 1. Historique, 1534-1977 et 2. Biometrie, biogéographie, 1970-1975, et registre
des péches 1941-1976 — Birds of prey of the world — Behavioral mechanisms in ecology

Botany: Guide to forest understory vegetation in Saskatchewan — Flora of the Prairie 488
Provinces: a handbook to the flora of the provinces of Manitoba, Saskatchewan and
Alberta; Part I. — Pteroids, ferns, conifers and woody dicopsids; Part I]. — Digitate,
Dimerae, Liberae; Part III. — Connatae; Part IV. — Monopsida — Living with plants: a
guide to practical botany — Common and botanical names of weeds in Canada/ Noms
populaires et scientifiques des plantes nuisibles du Canada

continued on inside back cover

ISSN 0008-3550

"ute ny ¥ i

Fabel Wyk ty
Wy. a
ONT Dnt Wo

2

ans hee
Neat
Ae

eas
ica

a a

MAR 1 6

3 2044 072 176 258

aoe,
vethoee

LiSeTaaHey nes
4 4d Fen
, di

Fa Myke

Wai
Way ar ake

"
Yip a
PL ay geass
ore rk ie

Sage
Way eg
vay

RCE i ey,

Aaa
“4
Rear iy

WoT te. ae

rae

me

‘eae ny ae he
woe

erry

Carer

Woe ae